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PROCEEDINGS 
of the 


LINNEAN 
SOCIETY 


of 
NEW SOUTH WALES 


VOLUME 124 


NATURAL HISTORY IN ALL ITS BRANCHES 


THE LINNEAN SOCIETY OF 
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VOLUME 124 
January 2003 


\) SOCTETY, OF 
L1H WALES 
-ATOSTY 


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EDITORIAL 


This is the first volume to be produced entirely by electronic means and in then new A4 format. The process has 
taken a great deal more time and effort than anticipated, resulting in the publication of Volume 124 being held over 
until January 2003. Therefore no volume is dated 2002 (Volume 123 was published in Dec 2001). Volume 124 is 
covered by 2002 subscriptions and memberships. 


Volume 125 is expected in the second half of 2003 and will contain a section with papers arising from Monotreme 
If], a symposium being held 10-11 July 2003 by the Linnean Society of New South Wales and the Australian 
Mammal Society. Details of that symposium can be obtained from the Secretary of the Linnean Society. 


In an attempt to contain the increasing costs of producing this journal, the Linnean Society now uses desk-top 
publishing to provide Southwood Press with printer-ready copy. At the final stage the journal is in PDF. There- 
fore any subscriber or member who would prefer to receive the journal in this format on Compact Disc should 
contact the Secretary. Since postage costs are becoming a major financial burden to the Society, the savings 
resulting from sending a CD rather than a printed journal are important. It is very likely that in the near future the 
journal may be available on-line, but at present the files are simply too large to allow transmission of the entire 
journal volume. 


In-house production puts a considerable workload onto voluntary staff and therefore the cooperation of authors 
in submitting manuscripts in the correct format is essential. It will be necessary to return manuscripts submitted 
that are not in accordance with the ‘Instructions for Authors’ as summarized at the back of this volume and as 
available at the Society’s web site or from the Secretary. 


This volume is a first step into a new era of publication. It is certainly not perfect, and any comments or 
suggestions would be warmly welcomed by the editor. This first step has been difficult, and would have been 
impossible without the technical support of Bruce Welch from Southwood Press and the services of the Dubbo 
Secretariat in transferring author’s artwork and photographs to electronic files. 


M.L. Augee 
Editor 


Proc. Linn. Soc. N.S.W., 124, 2003 i 


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Larval Distributions of Some Commercially Valuable Fish 
Species Over the Sydney Continental Shelf 


KIMBERLEY A. SMITH 


School of Biological Science, University of New South Wales, Sydney 2052, Australia 


Present address: NSW Fisheries, Cronulla Fisheries Centre, P. O. Box 21, Cronulla, NSW 2230, Australia. 


email : smithk @fisheries.nsw.gov.au 


Smith, K.A. (2003). Larval distributions of some commercially valuable fish species over the Sydney 
continental shelf. Proceedings of the Linnean Society of New South Wales 124, 1-11. 


The cross-shelf and vertical distributions of larvae of 12 species of commercially valuable marine fish are 
described from continental shelf waters off Sydney, south-eastern Australia. Depth stratified sampling was 
conducted along a shore-normal transect on 3 and 4 consecutive nights in January and April, respectively, 
1994. Larvae of the commercially valuable species Hyperlophus vittatus, Sardinops sagax, Engraulis 
australis, Argyrosomus japonicus, Pseudocaranx dentex, Trachurus novaezelandiae, Liza argentea, Sillago 
flindersi, Acanthopagrus australis, Pagrus auratus, Rhabdosargus sarba and Gerres subfasciatus together 
represented 11947 of the 50781 total fish larvae in samples. Species distributions extended to the outer shelf 
or slope, although the majority of larvae occurred in subsurface waters of the nearshore mixed layer. The 
majority of larvae were at a preflexion stage of development. Where present, later stage larvae tended to 
exhibit a different distribution to that of earlier stage larvae, although trends were variable among species. 
Results are discussed in relation to existing information on the larval distributions and spawning times of 


each species. 


Manuscript received 8 March 2002, accepted for publication 22 May 2002. 


KEYWORDS: ichthyoplankton, fisheries, flexion. 


INTRODUCTION 


Studies of early life history can provide 
valuable information to managers of fisheries 
resources. The spatio-temporal distribution of 
planktonic larvae, at various developmental stages, can 
help to infer the timing and location of spawning, and 
may assist in the interpretation of juvenile recruitment 
variability. Such information is most useful when 
coupled with oceanographic data (e.g. Caputi et al. 
1996). 

Australian marine  ichthyoplankton 
communities are typically diverse but numerically 
dominated by species of low fisheries value (Leis 1991; 
Gray 1993; Smith and Suthers 1999). The low 
abundances of species of higher value often preclude 
their individual attention in final analyses and 
published reports, contributing to a dearth of 
information on the early life history of many of 
Australia’s commercially valuable fish species 
(hereafter “commercial species’). The limited number 
of continental shelf ichthyoplankton studies, compared 
with those conducted within estuaries, is also a 
contributing factor. 


Several ichthyoplankton studies have been 
conducted in south-eastern Australian shelf waters 
(Miskiewicz 1987; Gray et al. 1992; Gray 1993, 1996, 
1998; Dempster et al. 1997; Smith and Suthers 1999; 
Smith et al. 1999; Smith 2000). Although generally 
not targeted, larvae of numerous commercial species 
have been encountered in these studies. However, only 
some of the larval distributions and developmental 
stages of commercial species captured in shelf waters 
during previous studies have been described (Gray 
1993, 1998; Smith 2000). 

Some commercial species occurring off 
south-eastern Australia are also distributed in other 
shelf regions, suggesting inferences about early life 
history may be drawn in the absence of local 
information. However, widespread intra-specific 
differences in spawning patterns and modes of larval 
dispersal among geographic regions and 
oceanographic regimes demonstrate the importance of 
local observations (e.g. Juanes et al. 1996). The aim 
of this paper is to provide local observations on the 
cross-shelf distributions and larval developmental 
stages of 12 species of commercial fish found in shelf 
waters off Sydney during two surveys in 1994. The 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


outer shelf occurrence of larvae of these 
species has not been previously described 
off Sydney. Samples described in this paper 
constitute a valuable contribution to the 
limited body of knowledge concerning the 
early life history of commercial fish in 
Australian marine waters. 


MATERIALS AND METHODS 


Location and time of study 

Data were collected in continental 
shelf waters adjacent to Sydney, on the 
south-eastern coast of Australia (Fig. 1). 
Currents over the shelf are predominantly 
southward, due to the influence of the East 
Australian Current (EAC) and associated 
eddies (Nilsson and Cresswell 1981). 
Compared to along-shore currents, cross- 
shelf currents are small, usually <10cm/s 
(Middleton 1987). Density variability in the 
Sydney coastal ocean is primarily the result of changes 
in temperature (Griffin and Middleton 1992). During 
the summer months, shelf waters generally exhibit 
strong temperature stratification (White and Church 
1986). 

Data were collected during two 10 d cruises, 
in January and April, 1994, aboard the research vessel, 
R. V. Franklin. On both cruises, data were collected 
from 5 stations along a cross-shelf transect. The 
transect began 2.7 km offshore and ended 40 km from 
the coast. Plankton sampling stations A, B and C were 
within shelf waters (bottom depths less than 150 m). 
Station D was at the shelf break, (bottom depth 250m), 
and station E occurred over the continental slope 
(bottom depth 600 m) (Fig. 1, Table 1). Plankton was 
collected on 22, 23 and 25 January and on 5, 6, 7 and 
8 April. Some locations were not sampled on 7 April 
due to bad weather. Plankton was collected at night 


TABLE 1. 
Details of location, bathymetry and sampling depth 
intervals for ichthyoplankton sampling stations. 


Distance Bottom 
offshore depth Depth of sampling intervals (m) 


Station (km) (m) Surface Shallow Middle Deep 
A 31 67 0-1 15-30 3040 40-50 
B 83 0-1 1540 4060 60-70 
€ 16 130 0-1 15-40 40-80 80-120 
|B boat taco 2 250 0-1 15-40 40-80 80-120 
E 40 600 0-1 15-40 40-80 80-120 


Figure 1. Location of sampling transect across 
Sydney continental shelf. 200 m isobath denotes 
shelf break. 


between 2030 and 0500 hours in January, and 1900 
and 0600 hours in April. Sunrise and sunset were at 
approximately 0600 and 2000 hours in January, and 
0600 and 1745 hours in April, respectively. 


Collection and processing of samples 

Surface plankton was collected by a 75 x 75 
cm square mouth net (330 um mesh), fitted with a 
General Oceanics flow meter. Two surface hauls, each 
of 6 minutes duration, were conducted at each station 
per night. Average volume of water filtered by the 
surface net was 291 m*. 

Subsurface plankton was collected by a 
multiple, opening and closing net with a square mouth 
of 1 m? and mesh size of 330 um. The net was fitted 
with temperature, conductivity and depth sensors and 
two General Oceanics flow meters - one inside and 
one outside the net mouth. Real time data were 
communicated to an operator onboard ship who 
electronically triggered each net release. At each 
station, three subsurface depth strata were sampled. 
Strata were designed to sample above (shallow), within 
(middle) and below (deep) the thermocline where 
possible at each station. Actual sampling depths varied 
according to hydrography and water depth at each 
station (Table 1). Subsurface haul durations were 10 
minutes and each depth stratum was obliquely sampled 
once per station per night. Average volume of water 
filtered by the subsurface net was 429 m’. 

Plankton samples were immediately placed 
into seawater and 5-10% formalin. Fish were removed 
from samples between 1 and 24 months after . 


Proc. Linn. Soc. N.S.W., 124, 2003 


K.A. SMITH 


collection, counted, identified and then stored in 95% 
ethanol. Larvae were assigned a developmental stage 
of preflexion, flexion, or postflexion. Flexion is the 
stage during which the notochord tip turns upward 
(Neira et al. 1998). Fin development is largely 
completed at the end of flexion. 

Larvae of twelve commercially valuable fish 
species were identified by descriptions in Leis and 
Trnski (1989) and Neira et al. (1998). Some other 
commercially valuable species, which were present in 
samples, were excluded due to very low abundance or 
difficulty in identification to species level. A summary 
of the Australian distribution and seasonality of each 
larval species can be found in Neira et al. (1998). 

Raw abundances per sample were 
standardised to density of larvae 100 m®? of water 
filtered. Mean density of each stage of each species 
was calculated for each sampling location (i.e. depth 
within station) during each sampling period. 


AY .B C 


0 
MIB, 


40 


— 
i 


RESULTS AND DISCUSSION 


During both sampling periods, a mixed layer 
of 23 °C and 20-50 m depth, overlay the shelf. The 
mixed layer extended over the entire shelf region 
during January but, during April, was displaced from 
the nearshore region by cooler, upwelled water (Fig. 
2). There was a tendency for the distributions of some 
species to extend further from the coast in April 
compared with distributions in January. This tendency 
was most pronounced in the carangids, Pseudocaranx 
dentex (silver trevally) and Trachurus novaezelandiae 
(yellowtail scad), and coincided with the offshore 
displacement of the nearshore mixed layer (Fig. 2). 
For a full description of this coastal upwelling event 
see Smith and Suthers (1999). 

A high density (298 larvae 100 m*) of 
Hyperlophus vittatus (sandy sprat) occurred in a single 
deep sample taken at station A on 22 January (Fig. 3a, 

Table 2). Densities in other January 

samples were relatively low. Few 

D E individuals were taken in April and 
these mostly occurred at shallow 

depths over the inner shelf. Flexion 
and postflexion H. vittatus were 
rare during both sampling periods 
and no trends were apparent in the 
vertical distribution of these stages. 

The occurrence of 
Hyperlophus vittatus off Sydney in 
January is consistent with 
spawning by this species during 
late summer and early autumn off 


0 10 20 


Figure 2. Average temperature (°C) profile across 
Sydney continental shelf during sampling on a) 22- 
25 January, and b) 5-8 April, 1994. Sampling 
stations A-E shown along top of profile. 


Proc. Linn. Soc. N.S.W., 124, 2003 


30 
Distance along transect (km) 


eastern Australia (Ramm 1986; 
Miskiewicz 1987). The occurrence 
of preflexion H. vitattus off Sydney 
(this study) and to the north of 
Sydney (Miskiewicz 1987), 
suggest a large potential spawning 
area. Off Western Australia, 
spawning also occurs over a long 
length of coastline (Gaughan et al. 
1996). The extremely patchy, 
coastal distribution of H. vittatus 
larvae observed off Sydney may be 
typical for this species (Gaughan 
et al. 1996). 

Engraulis australis 
(anchovy) were less abundant in 
April than in January. Spawning 
peaks between spring and autumn 
throughout the species distribution, 
and a decline in larval density off 
Sydney between January and April 
is consistent with decreasing 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


TABLE 2. 


Range of non-zero densities (larvae 100 m™) per sample [first line] and percentage of samples 
containing larvae [second line] per month. (n/a = no larvae; * = larvae in one sample only) 


January April 
Species Pre-flexion Flexion Post-flexion Pre-flexion Flexion Post-flexion 
Hyperlophus 0.41 - 298.28 0.30-0.70 n/a 0.02-0.18 n/a 0.02* 
Vittatus 14 3 6 1 
Engraulis 0.24 - 78.33 0.42 - 16.67 0.31 - 2.00 0.01-0.83 0.01-0.02 0.01 -0.74 
australis 31 15 11 10 2 6 
Sardinops 0.41 - 40.00 0.30-2.00 n/a 0.01-0.16 n/a n/a 
sagax 25 7 8 
Gertes 0.24 - 25.00 0.61-1.33 n/a 0.02-0.09 0.02% n/a 
subfasciatus 19 3 2 1 
Liza 0.24 - 6.67 0.30* 0.02-0.89 0.02-0.35 0.12-0.33 
argentea 29 1 18 7 2 
Agyrosomus 0.32 - 5.00 n/a n/a 0.01-0.62 n/a 0.02 - 0.03 
japonicus 19 14 2 
Sillago 0.28 - 81.67 0.30 - 13.33 0.35 - 4.00 0.01-5.08  0.02-0.23 0.02 -0.33 
flindersi 42 31 14 31 16 10 
Acanthopagrus 0.35* n/a 0.32* 0.01-0.15 0.02-0.04 n/a 
australis 1 1 7 4 
Pagrus 0.35 - 0.67 0.83* n/a 0.02-0.42 0.02-0.15 n/a 
auratus 6 1 9 2 
Rhabdosargus 0.83* n/a n/a 0.02-0.17 0.02-0.16 0.02-0.31 
sarba 139 8 3 5 
Pseudocaranx 0.63 - 251.67 0.30-28.33 0.30 - 24.16 0.01-1.69 0.02-0.46 0.02 -0.37 
dentex 43 28 13 22 4 2 
Trachurus 0.31- 1001.91 0.31-32.00 0.40 - 4.00 0.01 - 20.62 0.01-6.00 0.01 - 2.62 
novaezelandiae 50 22 10 53 21 10 


spawning activity between these times. Spawning 
occurs in estuaries and shelf waters of temperate and 
sub-tropical Australia (Miskiewicz and Neira 1998). 
Low densities of larvae over the outer shelf off Sydney 
(this study, Fig 3b) and elsewhere (Hoedt and 
Dimmlich 1995) suggests that spawning is restricted 
to the inner shelf. 

Densities of preflexion and flexion stage 
Engraulis australis larvae were highest in shallow and 
middle depths, and low at the surface (Fig. 3b). The 
highest density of preflexion E. australis was 78 larvae 
100 m*°, occurring at shallow depth at station A in 
January (Table 2). Postflexion larvae occurred at 
similar densities in surface and subsurface waters in 
January, but mainly at the surface in April. Previously, 
Gray (1993, 1996) found E. australis larvae most 
abundant in subsurface waters in November, but 
equally abundant in surface and subsurface waters in 
April/May and August/September. Given the tendency 
of older larvae to dominate surface samples in the 
present study, previously noted variability may reflect 
ontogenetic changes in vertical distribution. 


Sparids were rare in shelf waters during 
January and April, 1994 (Fig. 3c). Acanthopagrus 
australis (yellowfin bream) spawns throughout the year 
along the eastern Australian coast, although there are 
regional peaks in activity, including a local peak in 
autumn. This is consistent with a slight increase in 
larval abundance between January and April. A. 
australis observed during this study were caught within 
7 km of the coast. Spawning occurs at the mouths of 
estuaries and most previous observations of A. australis 
larvae within eastern Australian shelf waters have been 
within 1 km of the coast (Gray 1993; Miskiewicz and 
Neira 1998). Additional larvae may have been present 
in coastal waters in 1994 but distributed inshore of 
station A. 

Preflexion and flexion Acanthopagrus 
australis larvae occurred in subsurface samples but 
were absent at the surface. Densities of A. australis 
were < 1 larvae 100 m®° in all samples (Table 2). A 
single postflexion larva, which appeared competent 
to settle, was taken at the surface at station A in January. 
This is consistent with observations of settlement stage 


Proc. Linn. Soc. N.S.W., 124, 2003 


K.A. SMITH 


JANUARY 
Pre-flexion Flexion 
a) Hyperlophus vittatus 


APRIL 


Pre-flexion 


Post-flexion Flexion Post-flexion 


Depth (m) 


0 


Distance from coast (km) 


Figure 3. Mean density of preflexion, flexion and postflexion stage larvae of a) Hyperlophus vittatus, 
b) Engraulis australis c) Acanthopagrus australis, and d) Rhabdosargus sarba at sampling locations in 
January and April, 1994. Circle size is proportional to density of each stage within a month. Circle 
size is not comparable among stages or months (n = total number of larvae during sampling period) 


larvae at the surface in estuarine waters (T. Trnski, 
pers. comm.). 

In January, a single preflexion stage 
Rhabdosargus sarba (tarwhine) was taken, at shallow 
depth over the inner shelf (Fig. 3d). In April, R. sarba 
was more abundant in samples. Spawning by R. sarba 
occurs during most months of the year off eastern 
Australia, but local larval abundance and recruitment 
rates peak in autumn/winter (Miskiewicz and Neira 
1998; Smith and Suthers 2000). The increase in larval 
abundance between January and April, 1994, was 
consistent with these patterns. A single larva in 
January, and the presence of early and late stage larvae 
in April, 1994, suggested that some spawning activity 
occurred prior to April. Densities of R. sarba were < 1 
larvae 100 m? in all samples (Table 2). 

In April, preflexion, flexion and postflexion 
Rhabdosargus sarba occurred mainly over the inner 
and mid shelf, which is consistent with an inner shelf 
spawning location for this species (Wallace 1975; 
Miskiewicz 1986). Some preflexion and postflexion 
larvae also occurred over the outer shelf. The 
occurrence of postflexion larvae up to 40 km offshore 
of Sydney contrasts with the estuarine distribution of 
juveniles. Postflexion larvae, with some swimming 


Proc. Linn. Soc. N.S.W., 124, 2003 


ability, are less likely to have been passively advected 
away from the inner shelf than the less developed 
preflexion larvae. The outer shelf may be within the 
typical distributional range of older larvae. 

During night-time sampling in 1994, all 
developmental stages of Rhabdosargus sarba occurred 
in subsurface samples and were absent in surface 
samples. R. sarba larvae are also in subsurface waters 
off Sydney during the day (Gray 1998), suggesting 
that larvae do not undertake daily vertical migrations. 

Pagrus auratus (snapper) occurred at low 
densities in January and April (Fig. 4a). Spawning by 
P. auratus occurs throughout the year along the south- 
eastern coast, with a local peak in spawning activity 
in autumn (Miskiewicz and Neira 1998). Relatively 
low densities of larvae off Sydney, particularly in 
January, may reflect limited spawning at this time. 
Densities of P. auratus were < | larvae 100 m° in all 
samples (Table 2). 

Preflexion and flexion Pagrus auratus mainly 
occurred in subsurface waters of the inner and mid- 
shelf, although preflexion larvae also occurred at the 
surface and at shallow depths over the outer shelf in 
April. Spawning occurs in waters of < 50 m depth 
(Kailola et al. 1993), and so the presence of preflexion 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


APRIL 
Pre-flexion 


JANUARY 
Pre-flexion 
5 a) Pagrus auratus 


N 


4 b) Gerres subfasciatus 


-* n=101 | 

<< q . 

e, ( L EEE gee 
A 


ZEEE 
2 et 


d) 
© 


0 


Distance from coast (km) 


Densities of both preflexion 
and flexion Gerres subfasciatus were 
greatest at shallow depths. Maximum 
observed density of preflexion larvae 
was 25 larvae 100 m®, at shallow depth 
at station A in January (Table 2). Larvae 
occur at the surface during the day (Gray 
et al. 1992) and at depth during the night 
(this study), suggesting diel vertical 
migration by this species. 

Sardinops sagax (pilchard) 
probably spawns off NSW during most 
months of the year although the timing 
of peak activity may vary among years 
(Miskiewicz 1987). Preflexion larvae 
n=0 were relatively abundant in January, 

1994, suggesting significant spawning 
activity in the Sydney region at this time. 
S. sagax was largely restricted to the 
inner shelf in January (Fig. 4c). Larvae 
0 were more evenly distributed across the 
shelf in April, although densities were 
very low. An inshore larval distribution 
is consistent with spawning in estuaries 


Figure 4. Mean density of preflexion and flexion stage 
larvae of a) Pagrus auratus, b) Gerres subfasciatus, c) 
Sardinops sagax and d) Argyrosomus japonicus at 
sampling locations in January and April, 1994. No 
postflexion larvae of these species in samples. Circle 
size is proportional to density of each stage within a 
month. Circle size is not comparable among stages 


and inner shelf waters throughout 
temperate Australia (Blackburn 1949; 
Jenkins 1986; Fletcher and Tregonning 
1992; Hoedt and Dimmlich 1995). A 
small number of early stage larvae were 
found 30-40 km offshore, which is more 
likely to reflect larval advection than an 


or months (n = total number of larvae during 


sampling period) 


larvae over the continental slope (i.e. 40 km offshore) 
off Sydney during April may not reflect spawning 
location. Larvae may have been advected offshore 
during coastal upwelling at this time (Smith and 
Suthers 1999). No postflexion P. auratus were taken 
in either month. 

In January, preflexion stage larvae of Gerres 
subfasciatus (silver biddy) occurred over the inner and 
mid-shelf, and flexion stage larvae over the mid-shelf 
(Fig. 4b). In April, the distribution of both stages was 
restricted to the inner shelf and densities were very 
low. No postflexion G. subfasciatus were taken in 
either month. Spawning by G. subfasciatus peaks in 
summer along the south-eastern Australian coast 
(Miskiewicz and Bruce 1998) and declines in larval 
density between January and April may reflect a 
decline in spawning activity between these times. The 
occurrence of preflexion larvae over the inner shelf 
suggests a coastal spawning location. 


offshore spawning location (Smith and 
Suthers 1999). 

In January and April, the 
densities of preflexion and flexion 
Sardinops sagax were similar at shallow, middle and 
deep depths, but low at the surface (Fig. 4c). Maximum 
density of preflexion S. sagax was 40 larvae 100 m°, 
which occurred within shallow and deep samples at 
station A in January (Table 2). Flexion stage larvae 
were absent from samples in April and post-flexion 
stage larvae were absent from all samples. 

The vertical distribution of Sardinops sagax 
is spatially and temporally variable. Off Sydney, S. 
sagax has been observed below the surface during the 
day (Gray 1996) and night (this study). However, in 
south-western Australia, S$. sagax has been observed 
at the surface during the day but dispersed throughout 
surface and subsurface waters at night (Fletcher 1999). 
Hydrology may have influenced these vertical 
distributions. The water column was stratified during 
sampling off Sydney, and most larvae occurred just 
below the thermocline. There was no thermocline 
during sampling off western Australia. 


Proc. Linn. Soc. N.S.W., 124, 2003 


K.A. SMITH 


Similar numbers of preflexion Argyrosomus 
japonicus (mulloway) were caught in January and 
April, suggesting similar levels of spawning activity 
at these times (Fig. 4d). The occurrence of larvae in 
south-eastern Australian coastal waters during January 
is one month earlier than previous observations, 
between February and May (Steffe and Neira 1998). 
However, juveniles are known to enter local estuaries 
as early as February (Gray and McDonall 1993), and 
so the presence of larvae in coastal waters during 
January is possible. In January and April, densities 
were highest over the inner and mid-shelf. Larvae of 
this species also occur in nearshore areas and estuaries 
of South Africa (Beckley 1990). An inshore larval 
distribution is consistent with spawning by this species 
along ocean beaches in temperate and subtropical 
Australia (Kailola et al. 1993). 

Preflexion Argyrosomus japonicus larvae 
occurred at all sampling depths, although surface 
densities were low. Maximum observed density of A. 
japonicus was 5 larvae 100 m®°, at shallow depth at 
station A in January (Table 2). In April, larvae occurred 
mainly in middle and deep samples. This distribution 
may reflect an increasingly demersal habit with 
increasing size (A. Miskiewicz pers. comm.). This may 
also explain the rarity of later stage larvae in samples, 
if later stages typically occurred below the deepest 
sampling strata. Flexion and postflexion stages of A. 
japonicus were absent from samples in both months. 

Liza argentea (flat-tail mullet) was more 
abundant in April than in January, 1994, suggesting 
an increase in spawning activity between these months 
(Fig. 5a). L. argentea spawn between December and 
June (Kailola et al. 1993). L. argentea larvae occurred 
over the inner and mid-shelf in January, and also 
occurred over the outer shelf in April. Maximum 
observed density of preflexion L. argentea was 7 larvae 
100 m°, at shallow depth at station A (Table 2). The 
abundance of preflexion larvae over the inner shelf 
suggested a nearshore spawning location. Larvae 
tended to occur further from shore with increasing 
stage. 

Flexion and postflexion Liza argentea were 
most abundant at the surface. Later stage L. argentea 
also occur at the surface during the day (Gray 1993), 
suggesting that older larvae do not undertake daily 
vertical migrations. 

Sillago flindersi (eastern school whiting) was 
relatively abundant in January and April, 1994, 
suggesting that the level of spawning activity was 
similar at these times. The timing of spawning by S. 
flindersi peaks in spring/summer off south-eastern 
Australia (Kailola et al. 1993). Relatively high densities 
of later stage larvae in January, 1994, suggest that 


Proc. Linn. Soc. N.S.W., 124, 2003 


spawning activity had commenced at least several 
weeks prior to sampling. Larval of all developmental 
Stages occurred mainly over the inner and mid-shelf 
regions each month (Fig. 5b). The distribution of 
preflexion larvae suggested a nearshore spawning 
location. Maximum observed density of preflexion S. 
flindersi was 82 larvae 100 m°, at shallow depth at 
station A in January (Table 2). 

Each larval stage of Sillago flindersi was 
present at the surface and at all subsurface sampling 
depths. However, the vertical distribution of larvae 
changed between months. Preflexion larvae were most 
abundant in shallow samples in January but most 
abundant at the surface in April. Postflexion larvae 
occurred in shallow samples and at the surface in 
January, but in mid and deep samples in April. 
Preflexion and flexion S. flindersi larvae have 
previously been observed mainly in subsurface waters 
during the day (Gray 1996). The shift by postflexion 
larvae from warm, surface water in January to cool, 
deep water in April suggests that vertical distribution 
is not strongly influenced by hydrography. Postflexion 
S. flindersi may be patchily distributed throughout the 
water column. 

The carangids, Pseudocaranx dentex (silver 
trevally) and Trachurus novaezelandiae (yellowtail 
scad) were considerably more abundant in January than 
in April. Larvae of these species have been collected 
off eastern Australian at most times of the year, 
although spawning probably peaks in summer (Kailola 
et al. 1993; Trnski 1998) which is consistent with very 
high densities of larvae in January. The cross-shelf 
distributions of preflexion and flexion carangid larvae 
were similar between species (Fig. 5c). Larvae 
occurred over the inner and mid-shelf in January, and 
extended to the outer shelf in April. High densities of 
preflexion larvae over the inner shelf in January 
suggested spawning by both species over the inner 
shelf. The occurrence of larvae over the outer shelf in 
April resulted from the offshore displacement of larvae 
by a coastal upwelling event (Smith and Suthers 1999). 

Preflexion and flexion carangids occurred in 
subsurface samples in January, but also occurred at 
the surface in April. Maximum observed density of 
preflexion Pseudocaranx dentex was 252 larvae 100 
m7, at shallow depth at station A in January (Table 2). 
Maximum observed density of preflexion Trachurus 
novaezelandiae was 1002 larvae 100 m>, at shallow 
depth at station A in January (Table 2). 

Larvae of both carangids displayed an 
ontogentic shift in distribution although trends differed 
between species (Fig. 5d). Larvae of Trachurus 
novaezelandiae occurred further offshore with 
increasing stage. Larvae of Pseudocaranx dentex were 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


JANUARY APRIL 
Pre-flexion Flexion Post-flexion Pre-flexion Flexion Post-flexion 
a) Liza argentea 


Of S--0-==- 2 


> 


Silla 


c) Pseudocaranx dentex 
1 e----------------- - 


Depth (m) 


40 
Distance from coast (km) 


Figure 5. Mean density of preflexion, flexion and postflexion stage larvae of a) Liza argentea, b) Sillago flindersi, c) Pseudocaranx dentex 
and d) Trachurus novaezelandiae at sampling locations in January and April, 1994. Circle size is proportional to density of each stage 
within a month. Circle size is not comparable among stages or months (n = total number of larvae during sampling period) 


Proc. Linn. Soc. N.S.W., 124, 2003 


K.A. SMITH 


more abundant at the surface with increasing stage. 
During the day, the average size of Pseudocaranx 
dentex larvae is greater in subsurface waters (Gray 
1993), which suggests diel vertical migration by this 
species. However, differences in observations between 
studies may also reflect differences in hydrography 
(i.e. lack of water column stratification during sampling 
by Gray). The shift in cross-shelf distribution between 
January and April during coastal upwelling indicates 
that carangid larvae are indeed subject to 
hydrodynamic influences. 


Conclusions 

Approximately 90% of larvae of each species 
occurred at stations A, B or C (i.e. within 17 km of the 
Sydney coast) during January and April, 1994 (Fig. 
6). The highest densities of each species generally 
occurred at ‘shallow’ or ‘middle’ sampling depths, 
which corresponded to water within the mixed layer 
or upper thermocline (Fig. 2). No species was most 
abundant at the surface. These results highlight the 
importance of the nearshore mixed layer to many 
commercially significant fish larvae off south-eastern 
Australia. 


per species (%) 


Mean (+ s.d.) larvae 


30 
Distance from coast (km) 


0 10 20 


Figure 6. Mean (+ standard deviation) percentage 
of larvae of each species at sampling stations in 
January and April, 1994. 


The vast majority of individuals of all species 
observed in this study were at a preflexion stage of 
development. This is not unexpected and is likely to 
reflect the combined effects of high larval mortality 
and increasing net avoidance with increasing larval 
size. In some species, lower catchability of older larvae 
may also result from a movement away from the 
marine, pelagic environment towards the end of the 
larval phase. For example, older Argyrosomus 
japonicus larvae are believed to adopt a benthic habit 
prior to settlement in estuaries (A. Miskiewicz pers. 
comm.). Many coastal species have a coastal or 


Proc. Linn. Soc. N.S.W., 124, 2003 


estuarine juvenile phase and marine larvae must 
eventually migrate towards the coast. Postflexion 
larvae, nearing settlement, could have been present < 
2 km from the coast, inshore of station A during this 
study. This relatively shallow region was not accessible 
to the sampling gear employed by this study. 

Limitations on sampling location imposed by 
the use of a particular gear type confound many 
plankton studies, and contribute to an incomplete 
understanding of the life history of many fish species. 
In particular, no sampling of the extreme nearshore 
zone (i.e. m’s from shore) or absolute bottom (< 1 m 
from bottom) is reported for the south-eastern 
Australian coast. Such regions have been sampled 
elsewhere by use of light-traps or diver-operated nets 
and often host settlement stage larvae (e.g. Hickford 
and Shiel 1999). Off south-eastern Australia, these 
regions may host late-stage larvae of numerous 
commercial species, especially sparids, that are highly 
abundant as juveniles within coastal waters but 
relatively infrequently observed as larvae. 

All species encountered during this study are 
coastally distributed as juveniles and adults (Kailola 
et al. 1993), and the nearshore distribution of most 

larvae suggests that the life cycle of each 
species is typically completed close to the 
coast. However, observations of some larvae 
over the outer shelf and slope during this study 
provide an insight into the extent of 
distributional variability that larvae may 
experience. Larvae of five species (Sardinops 
sagax, Hyperlophus vittatus, Pseudocaranx 
dentex, Trachurus novaezelandiae, Sillago 
flindersii) were found > 30 km offshore in 
January. Larvae of nine species (Sardinops 
sagax, Engraulis australis, Pagrus auratus, 
Rhabdosargus sarba, Pseudocaranx dentex, 
Trachurus novaezelandiae, Liza argentea, 
Agyrosomus japonicus, Sillago flindersii) were 
found > 30 km offshore in April. The increased 
incidence of larvae in offshore waters in April, 
compared with January, is unlikely to have 
arisen by chance, considering the lower 
abundances of larvae in April. Passive larval advection 
during coastal upwelling in April is the most likely 
cause of these offshore distributions. The frequency 
of such offshore excursions by coastally-spawned 
larvae is unknown. Anomalous episodes of offshore 
advection may contribute to spatial and temporal 
variability of coastal recruitment by these species. 
Similarly, downwelling events may enhance the coastal 
recruitment of surface-orientated larvae. 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


ACKNOWLEDGEMENTS 


I thank A. Miskiewicz, J. Leis, I. Suthers and the crew of the 
R. V. Franklin. Work was partly funded by an Australian 
Research Council small grant. The manuscript was improved 
by helpful comments from two anonymous referees. 


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12 


COMMERCIALLY IMPORTANT FISH LARVAE OF THE SYDNEY SHELF 


Proc. Linn. Soc. N.S.W., 124, 2003 


New Distribution and Biological Records for Native Dung 
Beetles, in the Tribe Scarabaeini, from Northern New South 
Wales 


GEOFF WILLIAMS 


Research Associate, Department of Entomology, Australian Museum, 6 College Street, Sydney, NSW 2000, 
Australia 


Williams, G. (2003). New distribution and biological records for native dung beetles, in the tribe Scarabaeini, 
from northem New South Wales. Proceedings of the Linnean Society of New South Wales 124, 13-16. 


New coastal and inland distribution records, and behavioural observations, are given for Scarabaeini dung 


beetles collected from northern New South Wales. 


Manuscript received 21 December 2001, accepted for publication 19 September 2002. 


KEYWORDS: Coleoptera, Scarabaeidae, Scarabaeini, dung beetles, perching, cave fauna, subtropical 


rainforest 


INTRODUCTION 


Northern New South Wales is a centre of 
diversity for dung beetles in the tribe Scarabaeini 
(Matthews 1974). Numerous species are restricted to 
wet forest types along the Great Dividing Range 
(Williams 2002), but the maritime fauna, and that of 
isolated forest outliers east and west of the main 
mountain range complex, has been poorly documented. 

This paper gives additional scarabaeine 
distribution and behavioural records. Unless otherwise 
stated all beetles were collected at excrement-baited 
pit-fall traps. Voucher specimens have been deposited 
in the Queensland Department of Primary Industries 
Collection, Mareeba, Lorien Wildlife Refuge field 
reference collection, Lansdowne, and the Australian 
Museum, Sydney. 


NEW RECORDS 


Amphistomus speculifer Matthews 


New western distribution record 

Cedar Brush Nature Reserve, Liverpool 
Range, NW of Scone, 12-13.11.1999, G. Williams, 
subtropical rainforest. 


Aptenocanthon hopsoni (Carter) 


New northern and eastern distribution records 
Dooragan National Park, North Brother Mtn 
(~480 m.a.s.1.), Laurieton, 30.xii.1998, G. Williams, 


subtropical rainforest; same locality except, 25- 
27.x1.1999, G. and T. Williams; Hueys Corner, Fitzroy 
Creek, Carrai State Forest, WNW of Kempsey, 8- 
911.1997, G. and B. Williams, warm temperate 
rainforest; Banda Banda Beech Reserve, NW of 
Wauchope, 10-11.1.1984, G. Williams and C. Cross, 
cool temperate rainforest; Cockerawombeeba Flora 
Reserve, NW of Wauchope, 14-15.1.1988, G. and B. 
Williams, rainforest; Wilson River Flora Reserve, NW 
of Wauchope, 5-6.xii.1988, G. Williams, subtropical 
rainforest; vicinity Mt Seaview-Oxley Highway 
turnoff, W of Wauchope, 28-29.xi1. 1988, G. Williams, 
warm temperate rainforest. 


Aulacopris maximus Matthews 


New eastern distribution and behaviour records 
Approximately 4.5 km N of Lansdowne, 
Coorabakh National Park, 19.1.1996, G. and B. 
Williams, in bat cave, at bat guano; Dooragan National 
Park, North Brother Mtn (~480 m.a.s.l.), Laurieton, 
30.xii.1998, G. Williams, subtropical rainforest; Banda 
Banda Beech Reserve, NW of Wauchope, 28.1.1985, 
G. Williams, perching at night on tree trunk (~4m 
above ground), warm temperate rainforest; “The Pines’, 
Way Way State Forest, SW of Scotts Head, 20- 
21.1.1999, G. and T. Williams, subtropical rainforest. 


Diorygopyx asciculifer Matthews 


New northern and western distribution, and habitat 
records 

Cedar Brush Nature Reserve, Liverpool 
Range, NW of Scone, 12-13.11.1999, G. Williams, dry 


DISTRIBUTION OF NATIVE DUNG BEETLES IN NEW SOUTH WALES 


sclerophyll forest, at macropod dung; Camden Head, 
25-27.xi.1999, G. and T. Williams, littoral rainforest; 
Lake Cathie, 26.11.1999, G. and T. Williams, vine 
thicket. 


Diorygopyx incomptus Matthews 


New southern distribution, and habitat records 

Glenugie Peak, Glenugie State Forest, SE of 
Grafton, 16.xii.1998, G. and B. Williams, dry rainforest 
- vine thicket complex. 


Diorygopyx incrassatus Matthews 


New northern and eastern distribution, habitat and 
behaviour records 

Dooragan National Park, North Brother Mtn 
(~480 m.a.s.1.), Laurieton, 30.x11.1998, G. Williams, 
subtropical rainforest; same locality except, 25- 
27.xi.1999, G. and T. Williams; Sea Acres Nature 
Reserve, Port Macquarie, 25-27.xi.1999, G. and T. 
Williams, subtropical rainforest; Wilson River Flora 
Reserve, NW of Wauchope, 5-6.xii.1988, G. Williams, 
subtropical rainforest, perching at night on low foliage; 
Cockerawombeeba Flora Reserve, NW of Wauchope, 
14-15.1.1988, G. and B. Williams, rainforest; same 
locality except, 25-27.x1.1999, G. and T. Williams; 
Racecourse Headland, S of Crescent Head, 20.1.1999, 
G. and T. Williams, littoral rainforest; vicinity “The 
Blowhole’ Boonanghi State Forest, 24 km W of 
Kempsey, 8.x.1993, G. Williams, riparian dry 
rainforest; “The Pines’, Way Way State Forest, SW of 
Scotts Head, 20-21.1.1999, G. and T. Williams, 
subtropical rainforest. 


Diorygopyx niger Matthews 


New southern distribution and behaviour records 

Mt Killiekrankie, New England National 
Park, 12.1.2001, G. and B. Williams, wet sclerophyll 
forest, at bird carrion; Coachwood Creek, vicinity of 
‘Kookaburra’, Carrai State Forest, WNW of Kempsey, 
8-9.i1.1997, G. and B. Williams, wet sclerophyll forest 
- warm temperate rainforest complex; vicinity “The 
Natural Arch’, Carrai State Forest, WNW of Kempsey, 
8-9.ii. 1997, G. and B. Williams, wet sclerophyll forest. 


DISCUSSION 


Species of the flightless endemic genus 
Diorygopyx exhibit restricted distribution patterns 
along the New South Wales north coast (Matthews 
1974). Diorygopyx incrassatus was previously 
recorded from the Hastings Valley and the northern 


rim of the Manning Valley (Matthews 1974, Willams 
and Williams 1983b). It is replaced to the immediate 
north by D. niger and to the south by D. asciculifer 
(Matthews 1974). In the Carrai Plateau region, west 
of Kempsey, D. niger occurs at higher elevations (e.g., 
< 1000 m.a.s.l.) and D. incrassatus is found in 
submontane forest (ie. Boonanghi State Forest). The 
maritime distribution of D. incrassatus, however, is 
skewed northwards into latitudes occupied by D. niger 
and reaches to at least Way Way State Forest, near 
Macksville. The maritime distribution of Diorygopyx 
asciculifer is similarly skewed northwards. In littoral 
rainforest south of Port Macquarie D. incrassatus is 
displaced by D. asciculifer (Williams 1979, Williams 
and Williams 1984). At Laurieton D. incrassatus 
occurs in mountainous (~480 m.a.s.l.) subtropical 
rainforest of Dooragan National Park approximately 
3 kilometres inland from the coastline, but D. 
asciculifer occurs in adjacent littoral rainforest 
remnants at Camden Head, Lake Cathie and Crowdy 
Bay National Park, to the east, north, and south 
respectively. 

Diorygopyx asciculifer is the southern-most 
member of the genus, and is widely distributed in 
rainforests of the Manning catchment (Matthews 1974, 
Williams and Williams 1983a). In addition, it 
penetrates to the isolated Liverpool Ranges, west of 
Barrington Tops, where it has been collected in dry 
sclerophyll forest. At least one further species of 
Diorygopyx, D. duplodentatus Matthews, originally 
recorded only from rainforest, also occurs in drier 
forest types at the western extremity of its known range 
(C. Reid pers. comm.). 

Diorygopyx incomptus was originally 
described from the Macpherson Ranges (Matthews 
1974) but is more widely distributed in rainforests of 
far northern New South Wales (Williams 2002). It was 
collected in large numbers in low dry rainforest and 
associated vine thickets on scree slopes at Glenugie 
Peak, southeast of Grafton. This is a small area of 
isolated rainforest occurring within an extensive 
landscape matrix of dry forest and woodland. 

Matthews (1974) cited two specimens of D. 
niger found under old wallaby bones (an association 
which he considered possibly fortuitous). Numerous 
adult D. niger were collected in and under bird carrion 
at Mt Killiekrankie in January 2001 possibly 
confirming necrophagous habits in the species. 

The genus Aptenocanthon comprises two 
species from New South Wales (A. hopsoni, A. rossi 
Matthews) and a further six species from northern 
Queensland (Storey 1984, Storey and Monteith 2000). 
Aptenocanthon rossi is known only from the Mt 
Wilson-Mt Irvine area west of Sydney (Matthews 


Proc. Linn. Soc. N.S.W., 124, 2003 


G. WILLIAMS 


1974, Williams and Williams 1982) and A. hopsoni 
was previously recorded from montane wet forests in 
Barrington Tops, Dingo Tops and the Comboyne 
Plateau (Matthews 1974, Williams and Williams 
1983a). However, the distribution of A. hopsoni 
reaches montane rainforests in the Carrai Plateau, and 
its near-maritime occurrence in submontane rainforest 
at Dooragan National Park is exceptional. 

Aulacopris maximus was recorded by Waite 
(1898) from the Yessabah bat caves, in the Macleay 
Valley, northern New South Wales. Fricke (1964) 
recorded the related southern species A. reichei White 
“densely populating” a small cave sheltering 
bandicoots in a suburban garden at Mosman, Sydney. 
Aulacopris maximus is possibly a specialist on bat 
guano (G. Monteith pers. comm.). However, no further 
records of association with bat caves have been 
published. Aulacopris maximus was collected on a 
guano heap in a small cave in Coorabakh National Park 
(formerly part Lansdowne State Forest), near Taree, 
in January 1996. This is a roost cave seasonally 
occupied by Miniopterus spp., and dissected by an 
intermittent stream. Bat guano deposits are regularly 
flushed from the cave during heavy rain, which 
presumably would limit occupation of the cave and 
utilisation of guano deposits by invertebrate fauna. 

Scarabaeine dung beetles have been recorded 
from the Americas, and tropical Australia, perching 
near the ground on plant leaves (Howden, Howden 
and Storey 1991, Howden and Nealis 1978, Young 
1982). This may be a predator avoidance strategy 
(Young 1982) or related directly to foraging (Howden 
et al. 1991). Howden et al. (1991) record the dung 
beetle genera Monoplistes, Temnoplectron 
(Scarabaeini) and Onthophagus (Onthophagini) 
perching on foliage in tropical Queensland rainforest, 
but no other Australian records are known. Two 
subtropical species, Diorygopyx incrassatus and 
Aulacopris maximus, were observed nocturnally 
perching in montane rainforests of the Upper Hastings 
Valley; several D. incrassatus on low foliage in 
subtropical rainforest, and a single A. maximus 
approximately 4 m above ground on a tree trunk in 
warm temperate rainforest. Aulacopris maximus has 
previously been collected from inside possum nest 
boxes placed on tree trunks (Williams 1993). The large 
numbers of some species recorded by Howden et al. 
(1991) from tropical Queensland suggested “perching’ 
may be a common, albiet localised, strategy. There 
are no similar abundance records for Australian 
subtropical dung beetles, and extensive spot-lighting 
in New South Wales north coast rainforests indicates 
that perching may be rare. 


Proc. Linn. Soc. N.S.W., 124, 2003 


ACKNOWLEDGEMENTS 


The Director, New South Wales National Parks 
and Wildlife Service, and the Director, New South Wales 
State Forests, are thanked for permission to undertake 
fieldwork in national parks, nature reserves and forestry 
reserves. Discussions over the years with Dr Eric Matthews 
(South Australian Museum, Adelaide), Mr Tom Weir 
(CSIRO, Canberra) and Dr Geoff Monteith (Queensland 
Museum, Brisbane) have greatly contributed to my 
understanding of the Australian dung beetle fauna. Dr Chris 
Reid (Australian Museum, Sydney) kindly commented on 
an earlier version of this manuscript. I am grateful for the 
extensive field assistance given by my family over the years, 
and the company of Dr Dan Bickel (Australian Museum) 
during many interesting field trips. 


REFERENCES 


Fricke, F.T. (1964). A note on Aulacopris reichei White 
(Col., Scarabaeidae, Coprinae). Journal of the 
Entomological Society of Australia (N.S.W.) 1: 
36. 

Howden, H.F, Howden, A.T. and Storey, R.I. (1991). 
Nocturnal perching of scarabaeine dung beetles 
(Coleoptera, Scarabaeidae) in an Australian 
tropical rain forest. Biotropica 23: 51-57. 

Howden, H.F. and Nealis, V.G. (1978). Observations on 
height of perching in some tropical dung beetles 
(Scarabaeidae). Biotropica 10: 43-46. 

Matthews, E.G. (1974). A revision of the scarabaeine dung 
beetles of Australia. Il. Tribe Scarabaeini. 
Australian Journal of Zoology, Supplementary 
Series 24: 1-211. 

Storey, R.I. (1984). A new species of Aptenocanthon 
Matthews from north Queensland (Coleoptera: 
Scarabaeidae: Scarabaeinae). Memoirs of the 
Queensland Museum 21: 387-390. 

Storey, R.I. and Monteith, G.B. (2000). Five new species of 
Aptenocanthon Matthews (Coleoptera: 
Scarabaeidae: Scarabaeinae) from tropical 
Australia, with notes on distribution. Memoirs of 
the Queensland Museum 46: 349-358. 

Waite, E.R. (1898). Notes and exhibits. Proceedings of the 
Linnean Society of New South Wales 33: 803. 

Williams, G.A. (1979). Scarabaeidae (Coleoptera) from the 
Harrington district of coastal northern New South 
Wales, with special reference to a littoral rainforest 
habitat. Australian Entomological Magazine 5: 
103-108. 

Williams, G.A. (1993). Hidden Rainforests: subtropical 
rainforests and their invertebrate biodiversity. 
New South Wales University Press, Kensington, 
in association with the Australian Museum, 
Sydney. 

Williams, G.A. (2002). A Taxonomic and Biogeographic 
Review of the Invertebrates of the Central Eastern 
Rainforest Reserves of Australia (CERRA) World 
Heritage Area, and Adjacent Regions. Technical 


15 


DISTRIBUTION OF NATIVE DUNG BEETLES IN NEW SOUTH WALES 


Reports of the Australian Museum 16: 1-208. 


Williams, G.A. and Williams, T. (1982). A survey of the 


Aphodiinae, Hybosorinae and Scarabaeinae 
(Coleoptera: Scarabaeidae) from small wet forests 
of coastal New South Wales, Part 1: Nowra to 
Newcastle. Australian Entomological Magazine 
9: 42-48. 


Williams, G.A. and Williams, T. (1983a). A survey of the 


Aphodiinae, Hybosorinae and Scarabaeinae 
(Coleoptera: Scarabaeidae) from small wet forests 
of coastal New South Wales, Part 2: Barrington 
Tops to the Comboyne Plateau. The Victorian 
Naturalist 100: 25-30. 


Williams, G.A. and Williams, T. (1983b). A survey of the 


Aphodiinae, Hybosorinae and Scarabaeinae 
(Coleoptera: Scarabaeidae) from small wet forests 


of coastal New South Wales, Part 4: Lansdowne 


State Forest. The Victorian Naturalist 100: 146- 
154. 


Williams, G.A. and Williams, T. (1984). A survey of the 


Aphodiinae, Hybosorinae and Scarabaeinae 
(Coleoptera: Scarabaeidae) from small wet forests 
of coastal New South Wales, Part 5: littoral 
rainforests from Myall Lakes to Crowdy Head 
National Park. The Victorian Naturalist 101: 127- 
135. 


Young, O. P. (1982). Perching behavior of Canthon viridis 


16 


(Coleoptera: Scarabaeidae) in Maryland. New York 
Entomological Society 90: 161-165. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Host-plant Disjunction in a New Species of Neohoodiella 
(Insecta, Thysanoptera, Phlaeothripinae),with Notes on Leaf- 
Frequenting Thrips in New South Wales Subtropical 
Rainforests 


LAURENCE Mounp ! AND GEOFF WILLIAMS 2 


' CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601 (laurence.mound @csiro.au) 
?Research Associate, Department of Entomology, Australian Museum, 6 College Street, Sydney, NSW 2000 


Mound, L. and Williams, G. (2003). Host-plant disjunction in a new species of Neohoodiella (Insecta, 
Thysanoptera, Phlaeothripinae),with notes on leaf-frequenting thrips in New South Wales subtropical 
rainforests. Proceedings of the Linnean Society of New South Wales 124, 17-28. 


Neohoodiella jennibeardae sp.n. is described breeding on the leaves of two unrelated plants in the rainforests 
of eastern Australia, the dicotyledonous tree Ficus coronata (Moraceae) and the monocotyledonous vine 
Ripogonum elseyanum (Smilacaceae). To confirm this remarkably disparate pair of host associations many 
other plants in these rainforests were examined. This new species was not found on any other plant, although 
about 40 thrips species were taken from the leaves of 40 plant species in 22 families, and these records are 
tabulated. Neohoodiella is known previously only from a single species in New Caledonia. The genus is 
characterised by the two character states: abdominal tube one third of body length; dorsal setae elongate but 
broadly capitate. The head of N. jennibeardae bears a bifurcate tubercle that is unique amongst Phlaeothripidae. 


Manuscript received 12 May 2002, accepted for publication 21 August 2002. 


KEYWORDS: Leeuweniini, Neohoodiella jennibeardae, host plant associations, subtropical rainforest. 


INTRODUCTION 


Despite the acknowledged diversity of the 
insect fauna in Australia’s eastern rainforests (Monteith 
and Davies 1991), our knowledge of many groups is 
remarkably poor. For the order Thysanoptera, over 
most of the past 100 years, taxonomic descriptive work 
far outstripped any real understanding of the fauna. 
Most of these descriptions were by A. A. Girault, who 
published more than 130 species group names for thrips 
between 1924 and 1934 (Mound 1996), based mainly 
on single, often damaged, specimens, with no 
information about biology. Recent studies have been 
directed toward recognising the species described by 
Girault, establishing their structural variation and hence 
synonymies, and discovering their host plants (Mound 
2002a). This report is part of broader project to 
understand the biology of a larger proportion of the 
Australian thrips fauna. Plant names are used as in 
Mabberley (1997). 

Thrips species are proving to exhibit a wide 
diversity of interesting biological relationships. The 
apparent lack of natural enemies in Thrips imaginis 
Bagnall, the plague thrips that occurs in such vast 
numbers during early summer in southern Australia, 


has long been commented on (Andrewartha and Birch 
1954). However, other endemic thrips species also 
produce huge and apparently unconstrained 
populations. One was reported recently as invading a 
school in vast numbers in Queensland (Mound et al. 
2002). Moreover, this species has switched from 
breeding on its native host, Araucaria, to breeding on 
introduced northern hemisphere species of Pinus. Very 
large populations are reported also for thrips species 
that pollinate certain Macrozamia cycads in Australia, 
with up to 20 000 individuals occurring on a single 
male cone (Mound and Terry 2001; Terry 2001). 
Many thrips species feed on fungi on dead 
branches or in leaf litter (Mound 2002b), whereas 
others are phytophagous either in flowers or on leaves, 
some on single plant species but with a few 
polyphagous (Mound 2002a). Thrips are increasingly 
being recognised as plant pollinators, some as 
generalists (Williams et al. 2001) but others highly 
specific (Mound and Terry 2001). Similarly, behaviour 
patterns shown by particular thrips species are 
increasingly being investigated, such as lekking by 
males as is now known in two species of Australian 
Thripidae (Gillespie et al. 2002). Domicile creation, 
with adults securing leaves together with silk or glue, 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


is described for several species of Phlaeothripidae 
(Mound and Morris 2001). This behaviour is often 
accompanied by deliberate female (but not male) de- 
alation, although the significance of such wing removal 
remains unexplored. Structural polymorphisms, within 
or between sexes, can be so great that isolated 
individuals of the same species would not be 
considered congeneric (Mound et al. 1998), but 
behaviour patterns associated with such intra-specific 
variation have been studied in few species. 

Most recent research effort on thrips has been 
directed toward the arid zone of Australia (Crespi and 
Mound 1997). In this paper, a particularly bizarre new 
species is described from the eastern rainforests and 
observations recorded of its biology, this being the 
second member of a genus known previously only 
from New Caledonia. This new species was found 
breeding on the leaves of two very distantly related 
plants. To examine this disjunct host relationship, a 
survey was made of thrips associated with the leaves 
of many different plants in eastern rainforests around 
Taree, these records being tabulated and discussed 
below. 


Neohoodiella Bournier 
Neohoodiella Bournier 1997: 143. Type-species WN. 
grandisetis Bournier. 

The only previous species in this genus was 
described from a total of eight females and two males 
collected by a canopy fogging technique from 
unidentified forest trees in the Riviére Bleu region of 
New Caledonia. The genus is a member of the tribe 
Leeuweniini, in which the adults are distinguished from 
other leaf-feeding Phlaeothripinae by their elongate 
tenth abdominal segment, the tube (Ananthakrishnan 
1970). In most Phlaeothripidae, the tube is little more 
than twice as long as the ninth abdominal segment, 
whereas in Leeuweniini the tube is usually more than 
four times as long as the preceding segment. 
Neohoodiella differs from the other genera currently 
recognised in this group in having extraordinarily long 
setae on the head and pronotum, and the tube 10 times 
as long as the ninth tergite. 


Key to species of Neohoodiella 

1. Body and legs mainly light brown; antennal segment 
III with 1 sense cone, IV with 2 sense cones; major 
setae of head and body with margins smooth; 
ocellar region not produced over bases of 
antennae; median pair of major setae on vertex 
arising anterior to postocular setae; pronotal 
anteromarginal setae minute; pronotal notopleura 
each with 2 major setae; pronotal posteroangular 
setae minute; mesonotal lateral setae minute; 


18 


abdominal tergite IX setae B2 setaceous in 
contrast to capitate setae Bl; New Caledonia 
stad. cpp argank Boe deage te peers grandisetis Bournier 
-. Body and legs mainly clear yellow, dark brown on 
metascutum, tube apex and frontal margin of head 
(Fig. 1); antennal segment III with 2 sense cones, 
IV with 3 sense cones (Fig. 4); major setae of 
head and body with margins coarsely spiculate 
(Fig. 3); ocellar region with black, V-shaped 
tubercle projecting over front ocellus and 
extending beyond apex of antennal II (Fig. 2); 
median pair of major setae on vertex arising 
posterior to postocular setae; pronotal 
anteromarginal setae elongate; pronotal 
notopleura each with one large and one minute 
seta; pronotal posteroangular setae elongate; 
mesonotal lateral setae capitate with shaft 
spiculate; abdominal tergite IX setae B1 and B2 
similar in structure but B2 shorter; eastern 
Australia ..........::cs:eceeeeeeeeeee Jenmibeardae sp.n. 


Neohoodiella jennibeardae sp.n. 

Holotype 2, New South Wales, Lorien Wildlife 
Refuge, 3km N of Lansdowne near Taree, from Ficus 
coronata leaf, 27.x1i.2000 (LAM 3991), in ANIC, 
CSIRO Canberra. 

Paratypes: 109 6¢¥ taken with larvae; 19 50" at same 
site, 11.1.2001 (G. Williams); 19km NW of Bellbrook, 
Nulla Nulla Creek, 12 10% from F. coronata leaf, 
11.i1.2001 (G. Williams); NW of Wingham, Dingo State 
Forest, 49 20% from F. coronata leaf, 16.xii.2001 (G. 
Williams); Queensland, 100km NW of Brisbane, 
Conondale N.P., from Ripogonum elseyanum leaves, 
1Q, 10.x.2000, 32 30% taken with larvae 18.iii.2001 
(Dr Jenny Beard). Paratypes will be deposited in the 
US National Museum, Washington, Natural History 
Museum, London, and the Senckenberg Museum, 
Frankfurt. 

Female macroptera. Colour: Body pale yellow; 
metanotum with dark brown area; head with ocellar 
area dark brown and bearing black forked tubercle; 
tube golden with distal quarter brown; forewings pale 
with short darker line in basal third; antennal segment 
I dark brown, II yellow, III — IV light brown, V — VIII 
yellow with apex light brown; major setae mainly 
yellow but tergites III — VII each with 2 setae pale 
brown, pronotum with midlateral and posteroangular 
setae dark brown, mesonotal lateral pair light brown, 
tergites II and VIII each with 2 setae dark brown. 
Structure: Body elongate (Fig. 1), all major setae 
unusually long with shafts spiculate and apices with 
crown-like fringe of stout spicules. Head longer than 
wide, cheeks convex; eyes slightly smaller dorsally 
than ventrally; ocellar region produced into pair of long. 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. MOUND AND G. WILLIAMS 


Figures 1-4. Neohoodiella jennibeardae. 1, Male; 2, Cephalic tubercles; 3, Meso and metanota overlayed 
by the left pronotal posteroangular seta; 4, antennal segments II — VI. 


Proc. Linn. Soc. N.S.W., 124, 2003 19 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


tubercles overlaying front ocellus and extending to 
apex of antennal segment II, these tubercles with 
margins spiculate and bearing about 12 small setae; 
two pairs of postocular setae extending beyond apex 
of antennal segment II; maxillary stylets retracted to 
postocular setae, close together medially; mouth cone 
extending across prosternum. Antennae 8-segmented; 
III with 2 sense cones, IV with 3 sense cones; VIII 
slender. Pronotum with 5 pairs of major setae, am 
shortest, pa and epim arising from pronounced 
tubercles that obscure the notopleural sutures; 
prosternal basantra not developed, ferna large, 
mesopraesternum reduced to paired lateral triangles; 
metathoracic sternopleural sutures not developed. 
Mesonotal lateral setae well developed; metanotum 
reticulate with markings internal to reticles, paired 
median setae minute (Fig. 3). Legs slender, fore tarsus 
with no tooth; all femora with one large capitate seta 
on external margin medially. Forewing slender without 
duplicated cilia; 3 sub-basal setae long and capitate. 
Pelta triangular, tergite I with one pair of major setae 
near spiracle; tergites If — VIII each with 2 pairs of 
major setae laterally arising from tubercles, II — VII 
each with 2 pairs of strongly sigmoid wing-retaining 
setae; tergite IX setae B1 and B2 capitate and spiculate, 
B3 setaceous; tube exceptionally elongate (Fig. 1). 
Measurements (holotype 9 in micrometres). Body 
length 3150. Head, length 250; width 200; midvertex 
setae 230; postocular setae 240; inner margin of ocellar 
tubercles 130. Pronotum, length 130; width 280; major 
setae — am 140, aa 200, ml 220, epim 240, pa 230. 
Forewing, length 1000; distal width 50; sub-basal setae 
100, 110, 110. Tergite II lateral setae 190, 210. Tergite 
VII lateral setae 180, 170. Tergite IX, length 100; setae 
B1 180, B2 90, B3 80. Tube, length 960; anal setae 
450. Antennal segments I — VIII length, 40, 60, 90, 
80, 70, 70, 60, 50. 

Male macroptera. Indistinguishable from female in 
colour and structure but considerably smaller; sternite 
VIII with broad transverse glandular area on posterior 
half. 

Larvae and pupae. Colour yellow, apex of tube and 
antennae light brown. All major setae unusually long 
with broadly capitate apices but shafts not spiculate; 
head with 2 pairs of setae on vertex; pronotum with 6 
pairs; meso and metanota each with 5 pairs; tergites I 
— VIII each with 2 pairs arising from tubercles; tube 
three times as long as head. 


SYSTEMATIC RELATIONSHIPS 


Members of the Leeuweniini are recorded from 
various countries between India, New Caledonia and 


20 


Australia (Ananthakrishnan 1970), but only two other 
species have been described with long setae on the 
head and pronotum. These are the Indian species, 
Kochumania excelsa Ananthakrishnan (1969), which 
has the tube little more than twice as long the ninth 
tergite, and Neohoodiella grandisetis in which the tube 
is 10 times as long as the ninth tergite. Systematic 
relationships between the genera in the Leeuweniini 
require further study. The new species is remarkable 
for the pale yellow colour of the adults as well as the 
larvae, because adults of almost all large thrips species 
are brown to black. This pale colour, in combination 
with the long dorsal setae, results in the individuals 
being well camouflaged on the leaf surface. 


OBSERVATIONS ON BIOLOGY 


In common with other members of the 
Phlaeothripidae, the life history of N. jennibeardae 
involves two larval instars and three pupal instars. All 
of these life stages, from egg to adult, have been found 
on the leaves of Ficus coronata (Moraceae), a sand- 
paper fig, at a number of rainforest sites in the region 
of Taree (NSW), and it has been taken from this plant 
at sites between northwest of Kempsey and southeast 
of Gloucester. Moreover, larvae and pupae have been’ 
found on the leaves of Ripogonum elseyanum 
(Ripogoneaceae) at Conondale National Park just north 
of Brisbane (Qld). Although unrelated, the leaves of 
these two plant species are similar in texture, with 
prominent hairy veins on the lower surfaces. 

Despite the wide separation between the two 
collection areas, the distribution of N. jennibeardae 
appears patchy and unpredictable. The population on 
one particular tree at Lorien Wildlife Refuge, 
Lansdowne, was observed regularly over a period of 
18 months. In December, 2000, the thrips could be 
found on many leaves of this tree, all life stages being 
present. However, this population progressively 
declined, until by April 2002 only a single adult could 
be found. If this fluctuation in population size is 
normal, then our failure to find the thrips on the 
majority of F. coronata trees that have been examined 
gives no information about its real distribution. The 
leaves of this tree species are particularly long-lived, 
and populations of this thrips presumably prosper only 
in years when fresh growth is abundant. 

Eggs of this thrips are deposited on the lower 
surface of leaves, but in contrast to many other leaf- 
feeding Phlaeothripinae the eggs are scattered rather 
than in groups. This is possibly an adaptation to avoid 
predation by other insects, because this thrips 
apparently overwinters primarily as these isolated eggs, | 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. MOUND AND G. WILLIAMS 


not as adults. Adults and larvae commonly position 
themselves close to prominent veins of a Ficus leaf 
and, because of the large number of setae on their 
dorsal surfaces, they blend into the hairy under-surface 
of the leaf lamina. When illuminated artificially, thrips 
move to the shaded side of a leaf, although in lower 
light intensities they remain on the hairy lower 
surfaces, even when a leaf is deliberately inverted. 
Individuals have also been observed to be active on 
the leaves of Ficus trees during the night. Pupae were 
present on leaves, but were particularly difficult to find 
beneath the curve of hairy major leaf veins. No 
evidence could be found of larvae falling to the ground 
to pupate. 

The behaviour of adults and larvae was 
observed on detached leaves of F. coronata in petri 
dishes. The thrips are noticeably sluggish in their 
behaviour, quite unlike common flower-living species 
of Phlaeothripidae. When disturbed with a brush, they 
often sat lower onto the leaf surface, usually close to a 
vein, without being stimulated to walk or run. At other 
times when molested they waved the tube from side 
to side, often quite briskly, and sometimes raised it 
over the head. No aggression was observed between 
adults and larvae, but adults clearly explore the 
possibilities of mating. When a male first encountered 
a prospective mate he sometimes arched the tube over 
the female, although during copulation the tube was 
lowered horizontally. Copulation in one pair was 
observed to take about 1.5 minutes, but the male 
continued to straddle the female for a further half 
minute after copulating. During copulation, the male 
constantly stroked the female with his antennae, and 
appeared to stroke her abdomen with his mid and hind 
legs. 

Because the two recorded host plants of this 
thrips belong to such widely unrelated plant families, 
and considering the geographical range noted above 
between Taree and Conondale, we attempted to 
discover the insect on other host plants. To this end, 
we examined the leaves of numerous tree, shrub, vine 
and fern species in subtropical rainforests at various 
sites of the mid-north coastal region of New South 
Wales. Collecting methods were either by examining 
leaf surfaces with the aid of a head-mounted magnifier, 
specimens being removed with a small artist’s brush, 
or by beating fresh foliage of individual plant species 
onto a sheet or net. This yielded a considerable number 
of foliage associated thrips species, as listed in Table 
1 (located at the end of the paper, p. 25), but produced 
no evidence for a more extensive host range for N. 
jennibeardae. In particular, this thrips was not found 
on the other common, but relatively smooth-leaved, 
species of either Ficus or Ripogonum. 


Proc. Linn. Soc. N.S.W., 124, 2003 


N. jennibeardae thus appears to be restricted 
to just two unrelated plants. The first of these, Ficus 
coronata (Moraceae), is a small tree that is distributed 
widely from the Northern Territory, through 
Queensland to Victoria (Harden 1990). The second is 
a vine, Ripogonum elseyanum (Ripogoniaceae), that 
occurs in northern NSW north from Dorrigo to 
Queensland (Harden 1993). The distribution of these 
two plants overlaps in the rainforest of northern New 
South Wales. 


NOTES ON LEAF-ASSOCIATED THRIPS 


During the survey for alternative host plants 
for N. jennibeardae, various thrips species were taken 
around Taree from numerous unrelated plant taxa. 
Thrips are generally perceived as flower-living, but a 
considerable number of species rarely, if ever, visit 
flowers. Some species feed only on fungi, whereas 
others feed only on leaves. Because some of these small 
insects disperse on the wind, determining their precise 
biology from casual observations is fraught with 
difficulties. Adult thrips can be found, sometimes in 
considerable numbers, on plants to which they have 
no biological association. Moreover, adults sometimes 
feed on a plant species on which they do not breed. 
Thus recognition of true host plant associations 
amongst Thysanoptera is particularly difficult. The 
plants listed in Table I cannot be interpreted as the 
hosts of the thrips found on their leaves without further 
field studies, but these records provide a starting point 
for future studies. Many plant species examined in the 
field did not support thrips but, as indicated above for 
Neohoodiella jennibeardae, this could equally well 
reflect seasonal or spatial patterns of presence and 
abundance rather than patterns of non-exploitation by 
thrips. 

One of the most commonly encountered species 
in this survey was the greenhouse thrips, Heliothrips 
haemorrhoidalis (Bouché), a member of the Thripidae 
sub-family Panchaetothripinae in which all species 
breed only on leaves (Mound et al. 2001). This thrips 
breeds on the leaves of a wide range of plants, 
particularly introduced species. Damage to native 
plants by introduced insects is not well documented, 
but this thrips was observed causing damage to leaves 
of Tetrastigma nitens (Vitaceae) and Palmeria 
scandens (Monimiaceae) near Taree, and large 
populations have been observed damaging the leaves 
of Doryanthes excelsa (Doryanthaceae) near Sydney. 
Adults of a related endemic species, Helionothrips 
spinosus Wilson, were found on many plant species, 
but there is currently no evidence that it breeds 


Dal 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


anywhere other than on the leaves of Smilax australis 
(Smilacaceae). A third species of Panchaetothripinae, 
Anisopilothrips venustulus (Priesner), is known only 
from isolated adult females with no reliable host data, 
taken in many tropical countries and in Australia at 
scattered localities between Taree and Cape Tribulation 
in north Queensland. Another introduced 
panchaetothripine, Parthenothrips dracaenae 
(Heeger), is well known as a pest under domestic 
situations, damaging the Parlour Palm (Chamaedorea 
elegans - Palmae), but is not commonly taken in the 
field. In contrast, Bhattithrips Mound is an endemic 
panchaetothripine genus, with three described species 
and at least two more undescribed, but with no precise 
information on the biology of any of them. 

The sub-family Dendrothripinae (Mound 
1999b) appears to be better represented in rainforest 
than in the more arid parts of Australia. In a small 
floodplain rainforest remnant at Anthoneys Brush near 
Taree (see Williams 1993), females of Ensiferothrips 
primus Bianchi were found commonly on five plant 
species in five families. Females were also taken at 
other sites on the leaves of two further plant species. 
However, females together with males and larvae have 
been taken so far only from the vine Trophis scandens 
scandens (Moraceae) Curiously, this plant is absent 
from Anthoneys Brush, so the thrips is either highly 
dispersive or polyphagous. The only other member of 
this thrips genus, FE. secundus Mound, is known only 
from Lord Howe Island, and during a recent visit to 
that island the host plant of this thrips was found to be 
the endemic sub-species, T. scandens megacarpa, 
rather than the plants mentioned with the original 
description (Mound 1999a). 

Pseudodendrothrips gillespiei Mound was 
also described from Lord Howe Island, and several 
teneral adults were taken recently on that island from 
the leaves of T. scandens megacarpa. The record of 
one female of this species given here, from subtropical 
rainforest at Lorien Wildlife Refuge near Taree, 
represents the first record from the Australian 
mainland. The species listed as Pseudodendrothrips 
sp.n. was found in large numbers breeding on the 
leaves of Ficus fraseri, and was also taken in 
considerable numbers on Ficus coronata and F. 
rubiginosa. The colour of the forewings, however, 
varies among the samples taken, from mainly dark to 
banded. One female of Dendrothrips glynn Mound was 
taken, but the true host of this species is not known as 
it was based only on three females collected near 
Cairns. Similarly, the host plant of the widespread 
Dendrothrips diaspora Mound remains unknown, 
although collecting records suggest that this thrips is 
possibly polyphagous. In contrast, the species listed 


22 


as Dendrothrips sp.n. was found breeding on the young 
leaves of the tree Scolopia braunii (Flacourtiaceae) at 
two widely separated sites. 

The third sub-family of the Thripidae, the 
Sericothripinae, includes species that breed in flowers 
as well as species that breed on leaves. The female 
listed in Table I as Neohydatothrips poeta (Girault) is 
the third known specimen of this species, and the host 
plant remains unknown. In contrast, N. haydni (Girault) 
appears to be common on the young leaves of some 
species of Indigofera (Fabaceae), and possibly also 
on some species of Swainsonia (Fabaceae). The largest 
of the four sub-families of Thripidae, the Thripinae, 
includes many flower-living species. Williams, et al. 
(2001) recorded numerous Thripinae from the flowers 
of rainforest trees and shrubs in this study area, but in 
the present study, no attempt was made to sample thrips 
from flowers. Despite this, small numbers of the 
abundant flower-living species, Thrips setipennis 
(Bagnall), were taken on the leaves of Claoxylon 
australe (Euphorbiaceae), Acradenia euodiiformis 
(Rutaceae) and Gmelina leichhardtii (Labiatae), and 
a few specimens of Anaphothrips and Bregmatothrips 
that are possibly associated with grasses were also 
taken. Of the three leaf-feeding Thripinae in Table I, 
Chaetanaphothrips orchidii (Moulton) is introduced 
from southeast Asia, and was abundant on the leaves 
of an orchard tree, Annona cherimola (Annonaceae). 
Scirtothrips dobroskyi Moulton was described from 
the Philippines but is common in northeast Australia, 
and was found in large numbers on the terminal red 
leaves of another orchard tree, Mangifera indica 
(Anacardiaceae). The Oriental genus Rhamphothrips 
has only recently been recorded from Australia 
(Mound 2002a), based on a single female taken on the 
Cobourg Peninsula (Northern Territory), but an 
undescribed species of this genus seems to be 
widespread and abundant on the youngest leaves of 
Cissus antarctica (Vitaceae) in eastern NSW. 

Amongst the Phlaeothripidae that were found, 
some host associations in the list can be dismissed; for 
example Nesothrips and Hoplandrothrips species are 
known to feed on fungi not on green leaves. However, 
the presence of large numbers of adult Herathrips 
nativus (Girault) on the leaves of Drypetes deplanchei 
(Euphorbiaceae) in dry rainforest at Kiwarrack State 
Forest south of Taree, is more difficult to understand. 
The structure of the mouthparts of this species, 
previously known only from the type series of eight 
specimens, indicates that it feeds on fungal spores. 
Single specimens of this species were also collected 
on leaves of Baloghia inophylla (Euphorbiaceae) and 
Planchonella australis (Sapotaceae) at the same site. 
It seems likely that a large population had built up 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. MOUND AND G. WILLIAMS 


locally on dead leaves or branches, possibly on the 
lichens that are abundant at this site, and the individuals 
on leaves were part of a dispersing population. 

The single specimen of Hoodiella convergens 
(Hood) from Archontophoenix cunninghamiana 
(Arecaceae) was presumably a stray, but adults and 
many larvae of this thrips were found in distorted and 
partially rolled leaves of the vine, Tetrastigma nitens 
(Vitaceae). One species, Euoplothrips bagnalli Hood, 
was taken in rolled leaf galls on several plants, 
sometimes in large numbers, but is considered more 
likely to be a kleptoparasite than a gall-inducing 
species (Marullo 2001). The rolled-leaf galls on Smilax 
are probably due to Tolmetothrips smilacis (Priesner), 
a species that is widespread northward into the tropics. 
Foliage beating produced two species of Teuchothrips, 
a genus of leaf-feeding thrips that currently includes 
20 named species in Australia and at least as many 
un-named. The one from Tetrastigma has the antennae 
largely yellow, unlike any other member of the genus, 
and the one from Citriobatus pauciflorus 
(Pittosporaceae) is unusually small with both winged 
and wingless adults. The undescribed species of 
Haplothrips from Austrosteenisia (Fabaceae) is 
particularly interesting, because it was taken in large 
numbers, although without larvae, from the terminal 
leaflets of this plant, whereas Haplothrips species are 
usually flower-living. Similar in general appearance 
to this species were two that are presumed to be 
predatory, Haplothrips bituberculatus (Girault) and 
Xylaplothrips clavipes (Karny). The first is usually 
found on dead twigs, but the second is associated with 
the galls of other thrips. 

Finally, four Phlaeothripidae are listed that 
were taken in rolled-leaf galls, three apparently 
representing new genera. The leaf galls on Drypetes 
deplanchei were unusual, involving the margin of each 
leaf folding in for a distance of about 2 mm, enclosing 
a narrow tubular space but with the actual margin 
flattened and closely adpressed to the upper surface 
of the leaf. Two very different species of thrips were 
involved; a small but abundant, micropterous species, 
similar in appearance to certain gall-inducing 
Oncothrips species, presumably induces the galls, but 
with a second and much larger species that is probably 
a kleptoparasite. The leaf rolls on Acronychia 
oblongifolia (Rutaceae) were more open and irregular, 
as iS common amongst many members of 
Teuchothrips. These galls also contained two species; 
a large but short-winged species of Teuchothrips 
presumably induced the galls; the second species is 
apparently congeneric with an undescribed genus and 
species that commonly co-exists within the rolled-leaf 
galls of Gynaikothrips australis Bagnall on Moreton 


Proc. Linn. Soc. N.S.W., 124, 2003 


Bay fig trees (Ficus macrophylla). 

These records, from a relatively small area 
but involving several undescribed taxa, indicate that 
the diversity of Thysanoptera in Australia’s eastern 
rainforests is considerably higher than published 
records suggest. 


ACKNOWLEDGMENTS 


We are grateful to Dr Jenny Beard of the University 
of Queensland, St Lucia, for the initial discovery of this new 
species, and to John Hunter (NSW National Parks and 
Wildlife Service, Coffs Harbour) for investigating the 
presence of this thrips on Ficus coronata at several sites in 
northern NSW. 


REFERENCES 


Ananthakrishnan, T.N. (1969). New gall thrips from India 
(Ins., Thysanoptera, Phlaeothripidae). Sencken- 
bergiana biologia 50, 179-194. 

Ananthakrishnan, T.N. (1970). Studies on the genus 
Leeuwenia Karny. Oriental Insects 4: 47-58. 

Andrewartha, H.G. and Birch, L.C. (1954). The Distribution 
and Abundance of Animals. University of Chicago 
Press, Chicago. 

Bournier, J.-P. (1997). Thysanoptéres des foréts primaries 
de Nouvelle-Caledonie —I. Annals de la Société 
Entomologique de France 33, 139-153. 

Crespi, B.J. and Mound, L.A. (1997). Ecology and evolution 
of social behaviour among Australian gall thrips and 
their allies. Pp 166-180. In Choe, J. and Crespi, BJ 
(eds) Evolution of Social Behaviour in Insects and 
Arachnids. Cambridge University Press. 

Gillespie, P.S., Mound, L.A. and Wang, C.-L. (2002). 
Austro-oriental genus Parabaliothrips Priesner 
(Thysanoptera, Thripidae) with a new Australian 
species forming male aggregations. Australian 
Journal of Entomology 41, 111-117. 

Harden, G.J. (ed) (1990). Flora of New South Wales. Volume 
1. New South Wales University Press, Kensington. 
601 pp. 

Harden, G.J. (ed) (1993). Flora of New South Wales. Volume 
4. New South Wales University Press, Kensington. 
775 pp. 

Mabberley, D.J. (1997). The Plant Book. 2™ edition. 
Cambridge University Press. 858 pp. 

Marullo, R. (2001). Gall thrips of the Austro-Pacific genus 
Euoplothrips Hood (Thysanoptera), with a new 
species from Australia. Insect Systematics and 
Evolution 32, 93-98. 

Monteith, G.B. and Davies, V.T. (1991). Preliminary account 
of a survey of arthropods (insects and spiders) along 
an altitudinal rainforest transect in tropical 
Queensland. Pp 345-362. In Werren, G. and 
Kershaw, P. (eds) The Rainforest Legacy. Australian 
Heritage Publication Series Number 7, 414pp. 


23 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


Mound, L.A.. (1996). Thysanoptera, pp 249-336, 397-414 
(Index). In Wells, A., Zoological Catalogue of 
Australia. Volume 26. Psocoptera, Phthiraptera, 
Thysanoptera. Melbourne. CSIRO Australia. 

Mound, L.A.. (1999a). Thysanoptera from Lord Howe 
Island. Australian Entomologist 25, 113-120. 

Mound, L.A.. (1999b). Saltatorial leaf-feeding Thysanoptera 
(Thripidae, Dendrothripinae) in Australia and New 
Caledonia, with newly recorded pests of ferns, figs 
and mulberries. Australian Journal of Entomology 
38, 257-273. 

Mound, L.A.. (2002a). Thrips and their host plants: new 
Australian records (Thysanoptera: Terebrantia). 
Australian Entomologist 29, 49-60. 

Mound, L.A.. (2002b). Zemiathrips; a new genus of fungus- 
feeding phlaeothripine Thysanoptera in Australian 
leaf-litter. Australian Journal of Entomology 41, 
209-215. 

Mound, L.A., Crespi, B.J. and Tucker, A. (1998). 
Polymorphism and kleptoparasitism in thrips 
(Thysanoptera: Phlaeothripidae) from woody galls 
on Casuarina trees. Australian Journal of 
Entomology 37, 8-16. 

Mound, L.A., Marullo, R. and Trueman, J.W.H. (2001). The 
greenhouse thrips, Heliothrips haemorrhoidalis, and 
its generic relationships within the sub-family 
Panchaetothripinae (Thysanoptera; Thripidae). 
Journal of Insect Systematics and Evolution 32, 1- 
12. 

Mound, L.A. and Morris, D.C. (2001). Domicile constructing 
phlaeothripine Thysanoptera from Acacia phyllodes 
in Australia: Dunatothrips Moulton and Sartrithrips 
gen.n., with a key to associated genera. Systematic 
Entomology 26, 401-419 

Mound, L.A., Ritchie, S. and King, J. (2002). Thrips 
(Thysanoptera) as a public nuisance: a Queensland 
case study and overview, with comments on host 
plant specificity. Australian Entomologist 29, 25- 
28. 

Mound, L.A. and Terry, I. (2001). Pollination of the central 
Australian cycad, Macrozamia macdonnellii, by a 
new species of basal clade thrips (Thysanoptera). 
International Journal of Plant Sciences 162, 147- 
154. 

Terry, I. (2001). Thrips and weevils as dual specialist 
pollinators of the Australian cycad Macrozamia 
communis (Zamiaceae). International Journal of 
Plant Science 162, 1293-1305. 

Williams, G.A. (1993). Hidden rainforests: subtropical 
rainforests and their invertebrate biodiversity. New 
South Wales University Press, Kensington. 182 pp. 

Williams, G.A., Adam, P. and Mound, L.A. (2001). Thrips 
(Thysanoptera) pollination in Australian subtropical 
rainforests, with particular reference to pollination 
of Wilkiea huegeliana (Monimiaceae). Journal of 
Natural History 35, 1-21. 


24 Proc. Linn. Soc. N.S.W., 124, 2003 


Table 1. 


L. MOUND AND G. WILLIAMS 


Thysanoptera from leaves of subtropical rainforest plants near Taree, NSW. (‘V’ vine, ‘S’ shrub, ‘T’ 
tree, ‘SF’ State Forest, ‘NR’ Nature Reserve, ‘NP’ National Park) 


Plant species 


Anacardiaceae 
Mangifera indica 


Annonaceae 
Annona cherimola 


Arecaceae 


Archontophoenix 
cunninghamiana 


Euphorbiaceae 
Baloghia inophylla 
Breynia oblongifolia 


Bridelia exaltata 


Claoxylon australe 


Drypetes deplanchei 


Eupomatiaceae 
Eupomatia laurina 


Fabaceae 
Austrosteenisia blackii 


Indigofera sp. 


Flacourtiaceae 
Scolopia braunii 


Malvaceae 
Hibiscus heterophyllus 


ny 


T/S 


S/T 


Thysanoptera species 
Scirtothrips dobroskyi 
Chaetanaphothrips 
orchidii 
Hoodiella convergens 
Liothrips sp. 
Herathrips nativus 
Anaphothrips sp. 
?Bregmatothrips sp. 
Dendrothrips diaspora 
Ensiferothrips primus 
Bhattithrips sp. n. 
Thrips setipennis 
Herathrips nativus 


Phlaeothripinae 
gen.n.2 & 3 


Heliothrips 
haemorrhoidalis 
Neohydatothrips poeta 


Haplothrips sp.n. 


Neohydatothrips haydni 
Dendrothrips glynn 


Dendrothrips sp. n. 


Dendrothrips sp. n. 


Ensiferothrips primus 


Proc. Linn. Soc. N.S.W., 124, 2003 


Location 


Lorien Wildlife Refuge, 
3 km N Lansdowne 


Lorien Wildlife Refuge, 
3 km N Lansdowne 


Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lorien Wildlife Refuge, 
3 km N Lansdowne 


Kiwarrak SF, S of Taree 
Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lorien Wildlife Refuge, 
3 km N Lansdowne 
Anthoneys Brush, 

NE of Taree 

Saltwater Reserve, 

SE Taree 

Saltwater Reserve, 

SE Taree 

Kiwarrak SF, S of Taree 


Black Head 
20km S Taree 


Saltwater Reserve, 
SE Taree 
Saltwater Reserve, 
SE Taree 


Kiwarrak SF, S of Taree 


Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 


Lorien Wildlife Refuge, 
3 km N Lansdowne 


Plant community 


wet sclerophyll forest 


wet sclerophyll forest 


riparian rainforest 
riparian rainforest 


dry rainforest 
wet sclerophyll forest 


wet sclerophyll forest 
wet sclerophyll forest 
floodplain rainforest 
littoral rainforest 
littoral rainforest 

dry rainforest 


littoral rainforest 


littoral rainforest 


littoral rainforest 


dry rainforest 


dry rainforest 
dry rainforest 


wet sclerophyll forest 


Black Head 20km S Taree littoral rainforest 


Kiwarrak SF, S of Taree 


wet sclerophyll forest 


25 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


Plant species 


Monimiaceae 
Daphnandra micrantha 


Palmeria scandens 


Moraceae 
Ficus coronata 


Ficus fraseri 


Ficus rubiginosa 


Trophis scandens 


Streblus brunonianus 


Myrtaceae 
Backhousia sciadophora 


Rhodomyrtus psidioides 


Waterhousea floribunda 


Oleaceae 
Notelaea longifolia 


26 


ml 


Thysanoptera species 


Anisopilothrips 
venustulus 

Heliothrips 
haemorrhoidalis 


Heliothrips 
haemorrhoidalis 


Pseudodendrothrips 

sp.n. 

Pseudodendrothrips 
sp. n. 

Thrips setipennis 

Ensiferothrips primus 


Ensiferothrips primus 
Pseudodendrothrips sp.n. 
Ensiferothrips primus 


Ensiferothrips primus 
Ensiferothrips primus 


Pseudodendrothrips 
gillespiei 
Ensiferothrips primus 


Heliothrips 
haemorrhoidalis 
Ensiferothrips primus 


Xylaplothrips clavipes 


Heliothrips 
haemorrhoidalis 


Heliothrips 
haemorrhoidalis 

Heliothrips 
haemorrhoidalis 

Nesothrips propinquus 


Liothrips sp. 


S/T Ensiferothrips primus 


Location 


Lorien Wildlife Refuge, 
3 km N Lansdowne 
Tapin Tops NP, 

NW Wingham 


Upsalls Ck, Kerewong 
SF, WNW of Kendall 


Lorien Wildlife Refuge, 
3 km N Lansdowne 
Red Head, SE of Taree 


Harrington 

Anthoneys Brush, 

NE of Taree 

Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 
Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lansdowne Brush, 

0.5 km SE Lansdowne 
Red Head, SE of Taree 


Lorien Wildlife Refuge, 
3 km N Lansdowne 
Anthoneys Brush, NE 
of Taree; 
Wingham Brush NR 
Kiwarrak SF, 

S of Taree 

Kiwarrak SF, S of Taree 


Kiwarrak SF, S of Taree 


Woko NP, ~24 kn NNW 
of Gloucester 

Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lansdowne Brush, 

0.5 km SE Lansdowne 
Lansdowne Brush, 

0.5 km SE Lansdowne 
Lorien Wildlife Refuge, 
3 km N Lansdowne 


Anthoneys Brush, 
NE of Taree 


Plant community 


subtropical rainforest 
mixed subtropical 
rainforest — wet 
sclerophyll forest 
riparian subtropical 
rainforest 

wet sclerophyll forest 
headland littoral 
rainforest 

littoral rainforest 
floodplain rainforest 
dry rainforest 

wet sclerophyll forest 
subtropical rainforest 
floodplain rainforest 
headland littoral 
rainforest 


subtropical rainforest 


floodplain rainforest 


dry rainforest 
dry rainforest 


dry rainforest 


dry rainforest 

mixed rainforest — wet 
sclerophyll forest 
floodplain rainforest 


floodplain rainforest 


riparian rainforest 


floodplain rainforest 


Proc. Linn. Soc. N.S.W., 124, 2003 


Plant species 


Pittosporaceae 
Citriobatus pauciflorus 


Ripogonaceae 
Ripogonum album 


Ripogonum discolor 


Ripogonum 
fawcettianum 

Rubiaceae 

Morinda jasminoides 


Rutaceae 
Acradenia euodiiformis 


Acronychia oblongifolia 


Sapindaceae 
Mischocarpus 
pyriformis 


Sapotaceae 
Planchonella australis 


Smilacaceae 
Smilax australis 


Smilax glyciphylla 


Ulmaceae 

Aphananthe 
philippinensis 

Celtis paniculata 


Za) 


L. MOUND AND G. WILLIAMS 


Thysanoptera species 


Heliothrips 
haemorrhoidalis 
Parthenothrips 
dracaenae 
Ensiferothrips primus 
Euoplothrips bagnalli 
Haplothrips 
bituberculatus 
Xylaplothrips clavipes 
Teuchothrips sp.n. 


Heliothrips 
haemorrhoidalis 

Helionothrips spinosus 

Helionothrips spinosus 


Helionothrips spinosus 


Neohydatothrips ?sp. n. 


Thrips setipennis 


Phlaeothripinae gen.n.1 
Teuchothrips sp.n. 


Heliothrips 
haemorrhoidalis 
Haplothrips sp. 


Heliothrips 
haemorrhoidalis 
Hoplandrothrips sp. 


Herathrips nativus 


Helionothrips spinosus 
Helionothrips spinosus 


Tolmetothrips smilacis 
Euoplothrips bagnalli 
Helionothrips spinosus 


Ensiferothrips primus 
Anisopilothrips 


venustulus 
Helionothrips spinosus 


Proc. Linn. Soc. N.S.W., 124, 2003 


Location 


Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 


Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 


Kiwarrak SF, S of Taree 
Kiwarrak SF, S of Taree 


Wingham Brush NR 


Wingham Brush NR 
Lorien Wildlife Refuge, 
3 km N Lansdowne 
Camden Head 


Lorien Wildlife Refuge 
3 km N Lansdowne 


Lorien Wildlife Refuge, 
3 km N Lansdowne 
Lorien Wildlife Refuge, 
3 km N Lansdowne 


Lansdowne Brush, 
0.5 km SE Lansdowne 
Lansdowne Brush, 
0.5 km SE Lansdowne 


Lansdowne Brush, 

0.5 km SE Lansdowne 
Lansdowne Brush, 

0.5 km SE Lansdowne 
Kiwarrak SF, S of Taree 


Kiwarrak SF, S of Taree 
Red Head, SE of Taree 


Plant community 


dry rainforest 
dry rainforest 
dry rainforest 
dry rainforest 


dry rainforest 


dry rainforest 
dry rainforest 


floodplain rainforest 


floodplain rainforest 
subtropical rainforest 


headland littoral 
rainforest 


subtropical rainforest 


subtropical rainforest 


Wet sclerophyll forest 


floodplain rainforest 


floodplain rainforest 


floodplain rainforest 
floodplain rainforest 
dry rainforest 
dry rainforest 


headland littoral 
rainforest 


Black Head 20km S Taree littoral rainforest 
Black Head 20km S Taree littoral rainforest 


Saltwater Reserve, 
SE Taree 


Anthoneys Brush, 
NE Taree 
Camden Head 


Camden Head 


littoral rainforest 


floodplain rainforest 


headland littoral 
rainforest 
headland littoral 
rainforest 


27 


LEAF-FREQUENTING THRIPS IN SUBTROPICAL RAINFORESTS 


Plant species Thysanoptera species Location Plant community 
Verbenaceae 
Gmelina leichhardtii T Thrips setipennis Saltwater Reserve, littoral rainforest 
SE Taree 
Vitaceae 
Cissus antarctica VY -Rhamphothrips sp.n. Lorien Wildlife Refuge, wet sclerophyll forest 
3 km N Lansdowne 
Tetrastigma nitens Vs -Heliothrips Woko NP, ~24 km dry rainforest 
haemorrhoidalis NNW of Gloucester 
Heliothrips Kiwarrak SF, dry rainforest 
haemorrhoidalis S of Taree 


Hoodiella convergens _—_ Kiwarrak SF, S of Taree dry rainforest 
Euoplothrips bagnalli | Kiwarrak SF, S of Taree dry rainforest 
Teuchothrips sp.n. Kiwarrak SF, S of Taree dry rainforest 


28 Proc. Linn. Soc. N.S.W., 124, 2003 


Late Ordovician Allochthonous Limestones in Late Silurian Barnby 
Hills Shale, Central Western New South Wales 


Y.Y. Zuen", I.G. PerctvAv”” AND J.R. FARRELL? 


! Division of Earth and Environmental Sciences, The Australian Museum, 6 College Street, Sydney, N.S.W. 

2010 (yongyi@austmus.gov.au); * Specialist Geological Services, Geological Survey of New South Wales, 

P.O. Box 76, Lidcombe, N.S.W. 2141 (percivai@minerals.nsw.gov.au); > School of Education, Australian 
Centre for Educational Studies, Macquarie University, N.S.W. 2109 (jfarrell@ Aces 1.aces.mq.edu.au); 


“Honorary Research Associate, Centre for Ecostratigraphy & Palaeobiology, Macquarie University, N.S.W. 


Zhen, Y.Y., Percival, I.G. and Farrell, J.R. (2003). Late Ordovician allochthonous limestones in Late Silurian 
Barnby Hills Shale, central western New South Wales. Proceedings of the Linnean Society of New South 
Wales 124, 29-51. 


Allochthonous limestone blocks exposed in the Eurimbla area, west of the Mitchell Highway between 
Molong and Wellington, are substantially older than the enclosing Barnby Hills Shale of Late Silurian age. 
Nine of the blocks yielded a diverse Late Ordovician conodont fauna, dominated by Panderodus gracilis, 
Belodina confluens, Periodon grandis, Paroistodus? nowlani and Yaoxianognathus? tunguskaensis. 
Occurrence of Taoqupognathus blandus in seven sampled blocks indicates an early Eastonian (Ea2) age, 
although rare Taoqupognathus tumidus in one suggests an extension into the late Eastonian (Ea3). These 
age determinations are confirmed by the presence of a silicified brachiopod fauna with typical elements 
(predominantly Mabella halis and Doleroides mixticius) of the previously defined fauna B of Eastonian 2 
age. The conodont and articulate brachiopod faunas from the Eurimbla blocks are comparable with those 
from autochthonous limestones of Eastonian age elsewhere in the Molong Volcanic Belt, in particular the 
Bowan Park Group, except for occurrence of the conodont Webbygnathus munusculum and brachiopod 
Sowerbyella billabongensis which, in the Lachlan Orogen, are otherwise known only from the Junee- 
Narromine Volcanic Belt to the west. The allochthonous blocks may have been subject to one or more 


episodes of erosion and redeposition, prior to final emplacement in the Barnby Hills Shale. 


Manuscript received 19 June 2002, accepted for publication 27 November 2002. 


KEYWORDS: allochthonous limestones, Barnby Hills Shale, brachiopods, conodonts, Late Ordovician, 


Late Silurian. 


INTRODUCTION 


Regional mapping recently conducted by the 
Geological Survey of New South Wales in the central 
Lachlan Orogen has revealed that allochthonous 
limestones within the Late Silurian Barnby Hills Shale 
were derived from two major sources with substantial 
age differences. Morgan (1999: 92) determined that 
the Narragal Limestone, of early to middle Ludlow 
age (Percival 1998), provided the source of the Late 
Silurian limestone blocks and calcareous debris within 
the Barnby Hills Shale, based on faunal similarities 
and regional contact relationships. However, this is 
only the case within the belt of Barnby Hills Shale 
situated east of the Mitchell Highway (Fig. 1), for 
which Meakin and Morgan (1999) provided an 
extensive list of fossils from both the Silurian limestone 
blocks and the enclosing siliceous sediments. 


Other allochthonous limestones emplaced 
within the Barnby Hills Shale in the Eurimbla district, 
west of the Mitchell Highway between Molong and 
Wellington, were first recognised as Late Ordovician 
in age by Webby (1969) who identified in them the 
stromatoporoids Ecclimadictyon amzassensis and E. 
nestori, indicative of his early Eastonian coral- 
stromatoporoid Fauna II. Locations of the larger of 
these blocks were shown on a regional map by Byrnes 
(in Pickett 1982; reprinted in Lishmund et al. 1986), 
but only relatively recently (Farrell in Talent 1995) 
was detailed mapping of the area undertaken. Percival, 
Engelbretsen and Brock (1999) noted the occurrence 
and Eastonian age of a diverse lingulate brachiopod 
fauna (12 species) from one of the blocks; systematic 
description of this fauna is underway. Documentation 
herein of the remainder of the fauna, including 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Ups 
Zz 


S 


——. Geological boundary §7 Dip and strike 
=--o* Inferred geological 

boundary Ku» Quarry 
—<« Fault KiILS # Sample locality 
——— Thrust [FT} Measured section 
—?== Inferred fault line GL2%* Graptolite locality 
easaaae oe | Catombal Group 


a Dgg Garra Limestone 


Early i 
V Deg Cuga Burga Volcanics - 
Devonian 9 volcanilitharenite and volcanilithrudite. 


—_ hes Dmt Camelford Limestone 
Barnby Hills Shale - shale, chert, 
Silurian Smb meta-dolerite, rhyolite and 
Ordovician limestone blocks (L) 
teens Smq_ Narragal Limestone 


Ordovician Fairbridge Volcanics 


Figure 1. Locality maps. A. New South Wales showing location of figure 1B; B. simplified geological map of 
the area between Wellington and Cumnock, central New South Wales, showing position of figure 1C; C. 
geological map of the area to the east of Eurimbla, showing locations of sampled limestone blocks within the 
Barnby Hills Shale. Position of Figure 2, ‘KIL’ blocks, is also indicated. 


30 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


conodonts and articulate brachiopods, provides 
confirmation of the Eastonian age of this suite of 
allochthonous blocks, and further enables informed 
speculation as to their origin. 


GEOLOGICAL SETTING 


The Barnby Hills Shale was initially 
proposed by Strusz (1960) for shales, siltstones and a 
few small limestone lenses comprising the upper 
member of the Mumbil Formation. He also identified 
Monograptus bohemicus in a siliceous siltstone bed 
within the unit, supporting a mid to late Ludlow age. 
The unit was subsequently raised to formation status 
by Vandyke and Byrnes (1976). Byrnes (in Pickett 
1982: 146) regarded the section near the old “Mumbil” 
homestead, which was originally investigated by Strusz 
(1960), as the type section. However, due to faulting 
at the top and some internal folding at this locality, 
Morgan (1999: 90, photo 19 and fig. 21) more recently 
nominated the section exposed along a railway cutting 
on “Narrellen” property near Dripstone as the type 
section. At this locality, the formation is estimated to 
be 290 m thick, but it reaches a maximum thickness 
(over 700 m) in the Eurimbla area. 

Strusz (1960) stated that the Barnby Hills Shale 
was in conformable contact with the Early Devonian 
Cuga Burga Volcanics, but Chatterton et al. (1979) 
showed that the Camelford Limestone was intercalated 
between these two units, and that it had a conformable 
contact with both the Cuga Burga Volcanics and the 
Barnby Hills Shale. At Neurea, the Barnby Hills Shale 
conformably overlies the Narragal Limestone, the 
upper layers of which have been dated, using conodont 
data, as mid Ludlow siluricus Zone (Percival 1998). 
Graptolites from the Barnby Hills Shale at Neurea 
include Bohemograptus bohemicus tenuis Boucek, 
Pristiograptus dubius cf. frequens Jaekel and 
Egregiograptus egregius byrnesianum Rickards and 
Wright, indicative of a late Ludlow age (Rickards and 
Wright 1997). Hence in this eastern belt of outcrop of 
the Barnby Hills Shale, the age of allochthonous 
limestone blocks derived from the Narragal Limestone 
(Morgan 1997, 1999) is only slightly older than that 
of the clastic sediments into which they were 
redeposited. 

This is not the case, however, in the belt of 
Barnby Hills Shale situated west of the Mitchell 
Highway, and east of Eurimbla (Fig. 1). In the mapped 
area (Fig. 1B), the Barnby Hills Shale is fault-bounded, 
having over-ridden the Late Devonian Catombal 
Group to the east along the Curra Creek Thrust, and 
being bounded by the Cuga Burga Volcanics along 
the Eurimbla Fault to the west. Graptolites recovered 


Proc. Linn. Soc. N.S.W., 124, 2003 


from locality “GL2’ (GR 672700E 6352100N, 
Cumnock 1:50,000 8632-S), within the upper horizons 
of the siltstone sequence, have been identified as 
Monograptus ludensis (Murchison), indicating a latest 
Wenlock age (R.B. Rickards pers. comm.). From near 
the ‘KIL’ section at GR 673850E 6359110N, Sherwin 
(1997) reported the occurrence of Monograptus 
(Saetograptus) colonus, indicative of an early Ludlow 
age (Neodiversograptus nilssoni to early Lobograptus 
scanicus zones). North-east of ‘The Gap’ (Fig. 1), 
several graptolite species including Bohemograptus 
praecornutus Urbanek (praecornutus Biozone, ie 
middle Ludlow) were discovered in the uppermost 
horizons of the Barnby Hills Shale (R.B. Rickards pers. 
comm.). Given the evidence in this area of a latest 


“Wenlock to mid Ludlow age for the upper part of the 


Barnby Hills Shale, that name is retained despite 
arguments (Talent and Mawson 1999; Cockle 1999) 
for its suppression in favour of the more restricted, in 
both location and depositional time frame, late Ludlow 
(and younger) Wallace Shale (Sherwin and Rickards 
2002) — these two formations being demonstrably non- 
contemporaneous. 


DISTRIBUTION OF ALLOCHTHONOUS 
LIMESTONES 


Eighteen allochthonous limestone 
bodies of various sizes, ranging from less than a few 
metres to nearly 200 m in length, are emplaced at or 
near the faulted base of the Barnby Hills Shale, along 
the western side of the Curra Creek Thrust, and extend 
laterally for approximately 7 km on a NS trend (Fig. 
2). The limestone outcrops occur in three major 
groupings. One group is situated in close proximity to 
the “KIL’ locality; another incorporates the outcrops 
at ‘KILN’, ‘WKILN’ and ‘FT’ localities, while the 
third is grouped around a large outcrop at ‘EURO’. In 
addition, two very small isolated outcrops occur at 
‘“CWNN’ and ‘KILS’. 

The limestones are predominantly fine-grained 
and light to dark grey in colour with some isolated 
light fawn beds. Muddy intervals and rubbly beds are 
confined to a single block at the ‘KIL’ locality. Calcite 
veining is extensive at locality ‘FT’ and sets this 
outcrop at variance to all the others. Apart from a large 
syncline towards the top of block ‘KIL-C’, no other 
folding or faulting is apparent within the allochthonous 
blocks. Bedding is clearly evident in all blocks and 
varies from a few centimetres to 1 m in thickness. Apart 
from the silicified section in block “KIL-B’ (which 
yielded the abundant brachiopod fauna documented 
herein) and a coarsely silicified interval around sample 
number 78 in the same block, macro-fossils are 


Sil 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


BARNBY 
HILLS 


SHALE 


pill 
containing 
inarticulate 
brachiopods 


section 


ao] 
@ 
3S 
7) 


Silicified 
section 


metres 


Soil cover 


Late Devonian 
Catombal Group 


Silurian 
Barnby Hills Shale 


Late Ordovician limestone 


x= Barren samples 


Figure 2. Detailed outcrop map of, and stratigraphic sections measured through, three limestone blocks ‘KIL- 
A’, ‘KIL-B’ and ‘KIL-C’; samples producing conodonts are listed to the left of the columns. 


32 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


extremely rare. Only a few tabulate corals, some 
stromatoporoids, several brachiopods and one 
gastropod were recorded from other horizons in the 
allochthonous blocks. 


COMPOSITION AND AGE SIGNIFICANCE 
OF CONODONT FAUNA 


Conodonts recovered from nine individual 
limestone blocks are represented by 28 species (Fig. 
3), and show that these blocks are more or less similar 
in age (early Eastonian). Twenty-six of these species 
were previously recorded from the Cliefden Caves 
Limestone Group (Zhen and Webby 1995), Bowan 
Park Group (Zhen et al. 1999), Reedy Creek Limestone 
(Percival, Morgan and Scott 1999), and various other 
Eastonian successions in central New South Wales 
(Percival 1999; Packham et al. 1999; Pickett and 
Furey-Greig 2000; Pickett and Percival 2001), the New 
England Fold Belt (Furey-Greig 1999, 2000a, 2000b), 
and eastern Queensland (Palmieri 1978; Simpson 
1997). Coelocerodontus trigonius Ethington, 1959 and 
Panderodus serratus Rexroad, 1967 are recorded from 
the Ordovician of eastern Australia for the first time; 
both are relatively rare, but had a long 
biostratigraphical range, and are widely distributed. 

The six most common species in the fauna - 
Panderodus gracilis (37% of elements), Belodina 
confluens (14%), Periodon grandis (12%), 
Panderodus sp. (7%), Paroistodus? nowlani (6%), and 
Yaoxianognathus? tunguskaensis (5%) - are all 
cosmopolitan or widely distributed geographically. 
Biostratigraphically significant species present include 
Y.? tunguskaensis, which seems to be confined to 
Eastonian strata of central New South Wales (Trotter 
and Webby 1995, Zhen and Webby 1995, Zhen et al. 
1999), or time equivalents in North China (Wang and 
Luo 1984; An and Zheng 1990), northwestern China 
(Wang and Qi 2001), Siberian Platform (Moskalenko 
1973), and Canada (McCracken 2000). Chirognathus 
cliefdenensis has previously been reported only from 
Eastonian rocks of eastern Australia, apart from a 
recent record from the early Eastonian equivalent 
confluens Zone of southern Baffin Island, Canada 
(McCracken 2000). Typical species of the 
Aphelognathus webbyi biofacies, which characterises 
the early Eastonian Fossil Hill Limestone of the 
Cliefden Caves area, are extremely rare in the present 
collection, with only two specimens of the Pa element 
referrable to A. webbyi Savage, 1990. Webbygnathus 
munusculum was probably endemic to eastern 
Australia, with an age range from Ea2 in central New 
South Wales to Ea3 in the New England Fold Belt 
(Pickett and Furey-Greig 2000). 


Proc. Linn. Soc. N.S.W., 124, 2003 


Of particular note is the recovery of three 
species of the Eastonian genus Taoqupognathus (T. 
philipi, T. blandus and T. tumidus) from the ‘KIL’ 
locality, where three outcrops were mapped (Fig. 2) 
and sampled for conodonts (Figs 2 and 3). Zhen and 
Webby (1995) proposed a lineage for this genus from 
T. philipi to T. blandus to T. tumidus, which are now 
recognised as three succeeding conodont zones in the 
Eastonian (Zhen 2001). Taogupognathus blandus is 
much more common than the other two species, and 
indicates an early Eastonian age for these 
allochthonous limestone blocks in the Barnby Hills 
Shale. Taoqupognathus tumidus is very rare and is only 
represented by one specimen referable to the P element, 
which was recovered from the basal part of limestone 
block ‘KIL-B’. However, geopetal structures observed 
in thin sections prepared from samples taken from the 
silicified section (Fig. 2) indicate that this outcrop is 
overturned. Therefore, stratigraphically this late 
Eastonian (Ea3) species occurs in the highest level of 
this limestone block. Specimens referrable to T. philipi 
have been recovered from the lower part of limestone 
block ‘KIL-C’, and two more specimens doubtfully 
referred to T. philipi are recognised in the two samples 
of limestone block “KIL-B’. Occurrence of T. blandus 
and W. munusculum in other samples from the same 
stratigraphic horizon or below suggest that T. philipi 
might well extend upwards into the 7. blandus Zone 
or even T. tumidus Zone as a relic species (Zhen 2001). 
Therefore, despite the occurrence of T. philipi in some 
samples, a mid Eastonian (Ea2) age (T. blandus Zone) 
is postulated for the majority of the allochthonous 
limestone blocks. 


BRACHIOPOD FAUNA OF THE 
ALLOCHTHONOUS BLOCKS 


Brachiopods recovered from the silicified 
portion of the “KIL-B’ section (Fig. 2) include: Mabella 
halis, Doleroides mixticius, Rhynchotrema oepiki, 
Australispira disticha, Sowerbyella billabongensis, 
with rare Sowerbyites isotes, Zygospira carinata, 
Protozyga definitiva, Skenidioides quondongensis? and 
Chaganella speciosa, together with an indeterminate 
form provisionally identified as a craniid. All named 
species recorded here had previously been described 
(Percival 1991) from in situ Eastonian limestones in 
the Molong Volcanic Belt, such as the Cliefden Caves 
Limestone Subgroup and the Bowan Park Group, and 
the Billabong Creek Limestone in the Junee-Narromine 
Volcanic Belt to the west. Closest correlation is with 
Brachiopod Fauna B of Ea2 age (Percival 1992), as 
indicated by the presence of D. mixticius and 
Sowerbyella billabongensis which make their first 


33 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


appearances at this level. 

Mabella halis and D. mixticius overwhelmingly 
dominate the brachiopod fauna. While M. halis is 
ubiquitous throughout the Eastonian of the central 
Lachlan Orogen, D. mixticius is common only in the 
Billabong Creek Limestone. Another significant 
species, previously believed to be restricted to this latter 
formation, is S. billabongensis. On the other hand, two 
of the rare species in the allochthonous block fauna — 
Skenidioides quondongensis? and C. speciosa 
(represented by a solitary fragmentary specimen) — are 
otherwise known only from the Molong Volcanic Belt. 
However, given the location of the Eurimbla area in 
this Belt, this is not unexpected. The occurrence of S. 
billabongensis in the Eurimbla allochthonous blocks 
is unusual; seen in the context of the presence of the 
conodont Webbygnathus munusculum, which also is 
unknown in any other Molong Volcanic Belt 
limestone, this suggests the possibility of a linkage 
between the Junee-Narromine Volcanic Belt and the 
western flank of the northern Molong Volcanic Belt. 
However, affinities of the majority of the conodont 
fauna, discussed below, indicate likely derivation of 
the limestone blocks from a more proximal source area. 


STRUCTURAL AND DEPOSITIONAL 
SETTINGS 


To determine the most likely potential source 
of the allochthonous blocks, an analysis was 
undertaken of contemporaneous conodont faunas 
known from Eastonian limestones of the Molong 
Volcanic Belt. Many species are common to these in 
situ limestones and the allochthonous blocks, but it is 
the presence or absence of certain restricted species 
which is critical to revealing closest affinities. The 
nearest known in situ carbonate of Late Ordovician 
age, which might have provided a potential source area 
for the Eurimbla blocks, is the Reedy Creek Limestone, 
exposed near Molong about 20 kms to the south 
(Percival, Morgan and Scott 1999). The Reedy Creek 
conodont fauna is almost an exact duplicate of the more 
fully documented fauna from the Cliefden Caves 
Limestone Group, described by Zhen and Webby 
(1995). The latter shares 18 of the 28 species in the 
allochthonous assemblages, but significantly lacks 
Paroistodus? nowlani and Protopanderodus liripipus. 
Both of these species are present in allochthonous 
limestones near the base of the overlying Malongulli 
Formation (Trotter and Webby 1995), which is of 
slightly younger Ea3 age. However, the basal 
Malongulli Formation fauna is apparently devoid of 
Chirognathus cliefdenensis, Taoqupognathus blandus, 
T. philipi and Yaoxianognathus wrighti, which are 


34 


regarded as biostratigraphically important species. The 
Bowan Park Group fauna (Zhen et al. 1999), 
particularly that of the Quondong Limestone 
(Eastonian 2 age), has more species in common with 
the Eurimbla allochthonous blocks than do the 
previously mentioned faunas. Thus the affinities of the 
Eurimbla faunas are closer to in situ carbonates on the 
western flank of the Molong Volcanic Belt, rather than 
with the Reedy Creek Limestone, contrary to what 
might have been expected from the latter’s 
geographical proximity. 

The presence of Webbygnathus munusculum, 
although extremely rare in the Eurimbla allochthonous 
blocks (a single specimen recovered from “KIL-C’), 
is of some importance in being the first record of the 
taxon from the Molong Volcanic Belt. Pickett and 
Furey-Greig (2000), who described this species from 
Eastonian 2 horizons in the Billabong Creek Limestone 
of the Junee-Narromine Volcanic Belt, and Ea3 strata 
in the New England Orogen, commented that 
“curiously” their new monotypic genus had not been 
reported from any of the extensive assemblages 
(discussed above) described from the Molong Volcanic 
Belt. Hence it is worthwhile investigating other 
affinities between the conodont faunas from 
allochthonous limestones at Eurimbla with those from 
the upper Billabong Creek Limestone (itself in part an 
allochthonous horizon, although this was deposited 
penecontemporaneously with the surrounding 
sediments). A number of significant species found in 
the Eurimbla allochthonous blocks were not recorded 
from the Gunningbland area by Pickett and Percival 
(2001), particularly Paroistodus? nowlani, 
Protopanderodus liripipus, Pseudobelodina dispansa, 
Taoqupognathus philipi and Yaoxianognathus wrighti. 
Thus we conclude that the Billabong Creek Limestone 
is perhaps not as strong a contender as is the Bowan 
Park area for a potential source of the Eurimbla 
allochthonous blocks, and the presence of 
Webbygnathus munusculum in the latter remains 
enigmatic. 

Other erosional remnants in the nearby region 
are represented by Late Ordovician limestones in the 
Sourges Shale (Percival, Morgan and Scott 1999), 
especially the limestone containing conodont sample 
C1547 that is interpreted as having a middle Eastonian 
(Ea2-3) age. This outcrop is located about 3 km 
northeast of Cumnock, and is approximately 6 km west 
of the Eurimbla allochthonous blocks. Almost due 
north of the Eurimbla area, along regional strike, 
further evidence of Late Ordovician allochthonous 
limestones is found at “Narrawa” in the Wellington 
district (Percival, Morgan and Scott 1999). This 
particular occurrence yields corals of coral- | 
stromatoporoid Fauna III age, together with an 


Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


undescribed inarticulate (lingulate and acrotretid) 
brachiopod fauna containing several elements in 
common with the fauna found at “KIL-C’ at Eurimbla. 
The associated conodont fauna at “Narrawa’ is sparse, 
but includes Periodon grandis and Protopanderodus 
liripipus, both of which are represented at Eurimbla. 
These allochthonous blocks appear to have been 
emplaced into the Oakdale Formation, here of similar 
Late Ordovician (late Eastonian to early Bolindian) 
age. 

Allochthonous limestones of Late Ordovician 
age have also been reported from the Apsley and 
Bodangora areas, approximately 5 kms SSE and 15 
kms NNE, respectively, from Wellington. The Apsley 
block, located about 400 m E of the railway crossing, 
is apparently surrounded by Early Devonian Cuga 
Burga Volcanics. Conodonts recovered from this 
limestone (collection of the late G.C.O. Bischoff) 
included Yaoxianognathus, Periodon grandis, 
Panderodus, and Taoqupognathus blandus, indicative 
of an Eastonian 2 age — identical to that interpreted for 
the Eurimbla blocks. The Bodangora occurrence is 
shown on the most recent mapping by the Geological 
Survey of N.S.W. as an elongate limestone block 
within the Oakdale Formation at GR 685800E 
6409500N (Dubbo 1:100,000 mapsheet). Geological 
Survey microfossil sample C 061 yielded a small 
assemblage including the following conodonts: 
Belodina_ confluens, Panderodus_ gracilis, 
Protopanderodus liripipus and Taoqupognathus sp.., 
which can be dated no more accurately than Eastonian. 

Having established the age of the Eurimbla 
allochthonous blocks as early Eastonian (Ea2), and 
their most likely source as the Bowan Park Group, 
based on overall similarities in conodont faunas, the 
mechanism of their emplacement in the Barnby Hills 
Shale remains to be determined. Clumping of the 
limestone blocks in three separate groups, two to three 
kilometres apart, may reflect the presence of discrete 
channels or submarine valleys. The blocks are also 
emplaced at various stratigraphic levels within the 
Barnby Hills Shale, indicating that erosion and 
redeposition of material was not confined to a single 
episode. Timing of this series of events is constrained 
only by the age of the enclosing sediments, ie middle? 
to late Wenlock to mid Ludlow. 

In one possible scenario, the former Molong 
Volcanic Belt (in which the Bowan Park Group was 
deposited in the Late Ordovician) subsided in the Early 
Silurian, becoming the site for further shallow water 
sedimentation along the Molong High. Uplift of this 
area, concurrent with deposition of the Barnby Hills 
Shale in late Wenlock to mid Ludlow time, would have 
led to erosion of the Molong High succession and 
emplacement of allochthonous blocks in the deeper 


Proc. Linn. Soc. N.S.W., 124, 2003 


water sediments flanking that tectonic feature. The lack 
of carbonate debris forming breccia deposits within 
the Barnby Hills Shale in the study area suggests that 
the limestone blocks either slid down slope 
individually, or else were associated with a mass flow 
deposit but, due to their momentum, travelled further 
into deeper water after the bulk of finer-grained debris 
had settled. This model pre-supposes that only 
Eastonian limestone was available at the source site 
and that any carbonate material aged between 
Eastonian and the onset of deposition of the Barnby 
Hills Shale was either not present or had previously 
been eroded away. 

However, elsewhere in central New South 
Wales, Sherwin (1971) reported allochthonous blocks 
of Late Ordovician Malongulli Formation and Reedy 
Creek Limestone redeposited in the Wallace Shale of 
Late Silurian age, at “Mirrabooka” near Molong. In 
this instance, blocks with intervening ages are known 
to have been reworked into the succession, with detritus 
(including boulder beds) eroded from progressively 
older deposits. Thus clasts derived from the early 
Ludlow Molong Limestone appear lower in the 
Wallace Shale, to be succeeded by the Late Ordovician 
olistoliths as the Mirrabooka submarine valley (Byrnes 
1976; Byrnes in Pickett 1982: 159, figs 19, 20) 
excavated through the western shelf edge of the 
northern Molong Rise. 

Along the eastern margin of the Molong Rise, 
allochthonous block deposition in Late Silurian to 
Early Devonian fill of the Hill End Trough (Talent 
and Mawson 1999) derived from erosion of rocks 
forming the platform margin to the Mumbil Shelf. This 
was exposed during the Late Silurian, allowing 
limestone blocks of various sizes to detach and 
redeposit in the mud and silt matrix of a lower slope to 
basinal setting. Again, in this well-documented 
example, a considerable variety of ages of redeposited 
blocks, from late Wenlock to Emsian, are evident 
(Talent and Mawson 1999: text-fig. 7), which is at 
variance with the Eurimbla situation. 

An alternative model to account for the lack of 
any allochthonous Early Silurian carbonate material, 
involves tectonic uplift with multiple episodes of 
redeposition — the first concurrent with limestone 
breccias emplaced into the basal Malongulli Formation 
in the Cliefden Caves area (Rigby and Webby 1988). 
The source of these limestones is interpreted as the 
Ballingoole Limestone (Eastonian 3 age) in the upper 
Bowan Park Group. In the northern Molong Volcanic 
Belt, the Eurimbla blocks appear to have been derived 
from the slightly older Quondong Limestone, with 
initial emplacement in the deeper water Oakdale 
Formation flanking the volcanic belt. Subsequent 
tectonic uplift of this unit would lead to a second 


35 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


erosional episode in which only the more competent 
limestones were redeposited as recognisable clasts into 
the Barnby Hills Shale. This may explain removal of 
associated finer-grained carbonate debris to leave only 
the larger blocks in the final depositional episode. 

Large-scale faulting on the Curra Creek Thrust 
and related structures to the west (Scott and Glen 1999; 
Glen 1999: fig. 95) provides a possible mechanism to 
explain the occurrence of allochthonous blocks in the 
Barnby Hills Shale, by bringing Late Ordovician 
sediments to sufficiently shallow depths to expose them 
directly to subaerial or submarine erosion without 
having to wear through Early Silurian cover. In the 
extensional tectonic regime prevailing in the region 
during the Silurian, it is probable that some rotational 
component was involved in such faulting (R.A. Glen, 
pers. comm.). Further south on the Bathurst 1:250 000 
sheet, the Columbine Mountain Fault (Glen 1998: 302) 
defines the present-day crest of the preserved Molong 
Volcanic Belt, separating its western side (where the 
Bowan Park Group was deposited) from the eastern 
flank, site of the Cliefden Caves Limestone Subgroup 
and Reedy Creek Limestone. Webby (1992: 56) 
invoked early movement on the Columbine Mountain 
Fault as responsible for subsidence of the eastern flank 
while shallow water carbonate deposition continued 
in the Bowan Park area, the latter shedding debris into 
the deep water Malongulli Formation. Timing of this 
tectonic activity coincides with emplacement of 
shallow water carbonate blocks into the Oakdale 
Formation in the Wellington region, and quite feasibly 
caused displacement of Quondong Limestone 
equivalents into deeper water sediments. 


MATERIAL AND METHODS 


Five larger limestone bodies were measured and 
sampled, and a further four limestone bodies were also 
spot-sampled (Figs 1 and 2). The majority of conodont 
samples came from two measured sections through 
limestone blocks “KIL-B’ (16 samples) and “KIL-C’ 
(28 samples) (Figs 2 and 3). These samples, each 
weighing approximately 6 kg, were collected at regular 
intervals along the measured sections. Conodonts were 
extracted by completely dissolving the samples in 
dilute (10%) acetic acid; the residues were separated 
using the Sodium polytungstate technique outlined in 
Anderson et al. (1995). Sixty-five samples yielded a 
total of 1884 conodont elements (Fig. 3), which are 
relatively well preserved with a CAI of 4. Photographs 
of the conodonts are SEM photomicrographs captured 
digitally. Figured conodont specimens bearing the 
prefix AMF are deposited in the Australian Museum, 


36 


Sydney. The majority of the conodont species 
identified are documented by illustration only, as 
comparable material has been adequately described 
in recent publications on Late Ordovician conodont 
faunas of central-western NSW. Only those species 
providing new or comparative taxonomic information 
are discussed in detail in the following section. 

Brachiopods were obtained by acid dissolution 
of silicified horizons from limestone block ‘KIL’. 
Illustrated brachiopods are housed in the 
Palaeontological Collections of the Geological Survey 
of New South Wales at Lidcombe, and have the prefix 
MME. Silicified specimens were not whitened prior 
to being photographed digitally. As all taxa recognised, 
with the exception of a possible craniid, were 
comprehensively described by Percival (1991), only 
brief remarks are made on significant species. 

Grid references of the sampled localities (all 
on Cumnock 8632-S 1:50,000 sheet, using AMG66 
co-ordinates) are as follows: ‘KIL-A’: 674000E 
6358850N, ‘KIL-B’: 673950E 6358800N, ‘KIL-C’: 
673900E 6358950N, ‘KILS’: 673850E 6357900N, 
“‘CWNN’: 673600E 6357400N, ‘KILN’: 674350E 
6361700N, “‘WKILN’ (top): 674350E 6362000N, 
‘FT’: 674500E 6362100N, and ‘EURO’: 674350E 
6364650N. 


SYSTEMATIC PALAEONTOLOGY 
[Conodont taxonomy by Zhen; brachiopod 
taxonomy by Percival] 


Phylum CHORDATA Bateson, 1886 
Class CONODONTATA Pander, 1856 
Genus CHIROGNATHUS Branson and Mehl, 1933 


Type species 
Chirognathus duodactylus Branson and Mehl, 1933. 


Chirognathus? cliefdenensis Zhen 
and Webby, 1995 
Fig. SA-C 

Synonymy 

?Oulodus cf. oregonia (Branson, Mehl and 
Branson): Trotter and Webby, 1995, p. 483, 
pl. 4, figs 16-17. 

Chirognathus cliefdenensis Zhen and Webby, 1995, 
p. 281, partim, pl. 2, figs 13-22, pl. 3, figs 
2-4; non pl. 3, fig. 1; Zhen et al., 1999, p. 
86, Fig. 6.13-17. 

Yaoxianognathus? tunguskaensis (Moskalenko): 
McCracken, 2000, partim, only pl. 3, figs 
226, 28-30. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


ss sl Bes Ee = fe (2 bees lo sel ones 


fevro(basey| | of tet | tt tt tet tt TE 

fe eo Soo seee eee eeeaees 

SROs BESS SSSSs5e5 BEE mS 
ea ee OA | [Se 


abe 
wi7aa__[=| BORE 
eicteaa [|_| lamas (off 


iKictesa__| [=| 


Kuis26 | SISOS 

ec UC ee | IN Vl] 0 | a es see SP a ee ace af a ze | 
cece |_ Baal ese ee ae ee ae Se eae ee ae Sees 
Gigs JBoss bees Se Ree eRe saa eese 
UTS |e | | See ee esa pase | 
eit s a  a | OP |  f= ete e  ee eed Phe  az 
A RT aera Pa Pg ce | a eee fe a | 
er RS NT | | fa | | a a i ee es ez 


KIL1125 
KIL110.7 
KIL106.3 
KIL102 
KIL100.1 
KIL97.5 
KIL95.7 
KILOS 
KIL91 
KIL89 
KiL86.5 
KiL78.4 
KIL76.4 
KIL73 
KIL7Q 
KIL66.8 
KIL64.3 
KIL62 
KIL60 


KIL-C 
x 
r 
iil 
LI 
ie 
L 
ie 
ul 
a 
2| 
| 
| 
| 
5) 
| 
[| 
[| 
[| 
J] 
Ey 
| 
it) 
| 
| | 
[| 
a 
| 
a 
a 
be 
' 


a a sf ee ese opt ae eer) 
eS ee ae ee eS ae 


ESuUcuo eb JS ooe sense eoe eae 
acre | A a ee a eee za | 
eae 22 ee a | Fe | ap eae Pee de sp 3] 
ASRS saa Sea See aaa eee 
ae isa a | | aT | Sap eae eee Pt ee 
i a | a ee epee eae [ex] a 

ft So Pea eee Se Beas BES 
ese a | ce oa FY aa eal ee eas 
DLR OS 6 aoe noose Soe 
SLA SAS eae sea aA eae ae ees 
BEE EEE EE EERE EEE EEE 


KIL-B 


ra 
LE 
MERA ABEOEOO COCO Sao 
HEBEREROE IEC COCO ae 


KILS2.4 
KIL49.3 
KIL48 
KIL42.5 
KIL38 
KIL33.6 
KILS.4 
KIL8.4 


Soong eosr aSseLoes 


KIL-A 


cf. venustus 


inathus webbyi 
Belodina baiyanhuaensis 
Belodina confluens 


limestone bodies 


gnathus? cliefdenensis |_| 


Coelocerodontus tngonius 
gmodus undatus 


Conodont taxa 
Ansella sp. 

phelo: 
Besselodus sp. 
Drepanoistodus suberectus 
"Oistodus" sp. 
Panderodus serratus 
Panderodus sp. 
Paroistodus? nowlani 
Pseudobelodina dispansa 
Pseudobelodina sp. 
Pseudooneotodus mitratus 


Proc. Linn. Soc. N.S.W., 124, 2003 


Figure 3. Distribution chart of conodont species from nine individually sampled allochthonous limestone blocks. 


3H 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Figure 4. SEM photographs of Late Ordovician conodonts from allochthonous limestones within the Barnby 
Hills Shale. A, B, Ansella sp., asymmetrical nondenticulated element; A, AMF121400, “KIL’ 9.4; B, AMF121401, 
‘KIL’ 95. C, Aphelognathus webbyi Savage, 1990, Pa element, AMF121402, ‘KILS’. D, Belodina 
baiyanhuanensis Qiu in Lin et al., 1984, compressiform element, AMF121403, ‘KILN’ 52. E-G, Belodina 
confluens Sweet, 1979, E, eobelodiniform element, AMF121404, ‘KIL’ 97.5; F, grandiform element, 
AMF121405, ‘KIL’ 70; G, compressiform element, AMF121406, ‘KIL’ 173.1. H-K, Belodina sp., grandiform 
element; H, AMF121407, ‘KIL’ 56; I, J, AMF121408, ‘KIL’ 48; K, enlargement of surficial ornament, 
AMF121409, ‘KIL’ 49.3. L-O, Besselodus sp., short-based distacodiform element; L, M, AMF121410, “KIL’ 
161.4; N, AMF121411, ‘KILN’ 52; O, AMF121412, ‘KIL’ 161.4. P-X, Coelocerodontus trigonius Ethington, 
1959; P, Q, symmetrical trigoniform element, P, AMF121413, ‘KIL’ 173.1; Q, AMF121414, ‘KIL’ 9.4; R, S, 
asymmetrical trigoniform element, AMF121415, ‘KIL’ 173.1; T, U, asymmetrical tetragoniform element, 
AMF 121416, ‘KIL’ 173.1; V-X, symmetrical tetragoniform element (W: basal view), AMF121417, ‘KIL’ 
173.1; scale bars 100 um, except as indicated for K. 


38 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


Material 
Fifteen specimens (6 Pa, 4 Pb, 5 S) from limestone 
blocks ‘KIL-B’, “KIL-C’, ‘FT’ and ‘KILN’. 


Discussion 

All elements, except for the symmetrical Sa, have been 
recovered from various limestone blocks within the 
Barnby Hills Shale, and they are identical with the type 
material from the Cliefden Caves Limestone Group 
(Zhen and Webby 1995). 

McCracken (2000) proposed a septimembrate 
apparatus for Yaoxianognathus? tunguskaensis 
(Moskalenko). This reconstruction is rather different 
from that recognised by Zhen et al. (1999) on the basis 
of homologic characters, such as widely spaced, robust 
denticles on all elements of Y.? tunguskaensis. 
McCracken’s illustrated Sc and M elements of Y.? 
tunguskaensis are typical for Yaoxianognathus, 
especially the Sc (McCracken 2000: pl. 3, fig. 25) 
which is referable to Y. tunguskaensis. However, P (Pa, 
Pb, Pc) and Sa elements figured by McCracken (2000: 
pl. 3, figs 26, 28-30) are at least congeneric if not 
conspecific with C. cliefdenensis. 


Genus COELOCERODONTUS Ethington, 1959 


Type species 
Coelocerodontus trigonius Ethington, 1959. 


Coelocerodontus trigonius Ethington, 1959 
Fig.4P-X 

Synonymy 

Coelocerodontus trigonius Ethington, 1959, p. 273, 
pl. 39, fig. 14; Webers, 1966, p. 25, pl. 2, 
figs 12-14; ?Orchard, 1980, p. 19, pl. 2, 
figs 17, 22, 23, 29; Nowlan et al., 1988, p. 
14, pl. 3, figs 1-5, 8-10 (cum. syn.); 
McCracken and Nowlan, 1989, p. 1888, pl. 
2, fig. 18; Nowlan et al., 1997, pl. 1, fig. 4; 
Zhang, 1998, p. 56, pl. 5, figs 1-4. 

Coelocerodontus tetragonius Ethington, 1959, p. 
ZIBe plesOmtiesiS: 

Coelocerodontus digonius Sweet and Bergstrém, 
1962, p. 1224, pl. 168, fig. 1, Text-fig. 1F. 


Material 

Eight specimens (6 trigoniform, 2 tetragoniform) 
from limestone blocks “KIL-A’, ‘KIL-B’ and ‘KIL- 
(Or 


Discussion 

Webers’ (1966) initial species concept of a bimembrate 
(trigoniform and tetragoniform) apparatus was revised 
by Nowlan et al. (1988) as a trimembrate apparatus 
including a symmetrical trigoniform, a slightly 


Proc. Linn. Soc. N.S.W., 124, 2003 


asymmetrical trigoniform and a nearly symmetrical 
tetragoniform element. All three elements have been 
recognised in the central New South Wales material, 
together with an additional slightly asymmetrical 
tetragoniform specimen. All these elements are 
characterised by having thin walls and a very deep 
basal cavity, with the apex nearly reaching the tip of 
the cusp. The trigoniform elements, with a broad 
anterior face, a sharp costa on each antero-lateral 
corner, and a costa along the posterior margin, are 
either symmetrical (Fig. 4P, Q) or slightly 
asymmetrical (Fig. 4R, S), with a triangular opening 
of the basal cavity. The latter is identical with the 
holotype of the form species C. trigonius, except for 
its more antero-posteriorly compressed cusp. The 
tetragoniform element is quadrate in cross section with 
four prominent costae, situated on the antero-lateral 
and postero-lateral corners of each side. Our specimens 
are identical with the holotype of the form species C. 
tetragonius Ethington, 1959, except that the latter 
(Ethington 1959: pl. 39, fig. 15) is more laterally 
compressed. 

Zhang (1998) illustrated (but neither defined 
nor described) laterally compressed, non-costate P 
elements, and costate Sb, Sc and Sd elements from the 
Middle Ordovician of South China. Of the two 
specimens (Zhang 1998: pl. 5, figs 1, 2) referred to as 
P elements, the more slender is identical with the 
holotype of the form species C. digonius Sweet and 
Bergstrom, 1962. The other specimen (Zhang 1998: 
pl. 5, fig. 1) is a wider conical unit with a more 
posteriorly extended base. These elements may be 
differentiated as Pa and Pb, respectively. The specimen 
designated as the Sd element (Zhang 1998: pl. 5, fig. 
4) seems identical with one illustrated as 
Coelocerodontus? sp. from the Middle Ordovician 
(Darriwilian) of northern Sweden (Lofgren 1978: pl. 
1, fig. 40). Neither the P elements, nor the Sd element 
with a mid-costa on each side, have been recognised 
in the central New South Wales material. 

Coelocerodontus trigonius ranged through 
the Middle and Upper Ordovician. The type specimen 
(Ethington 1959: pl. 39, fig. 14) from the upper Galena 
Formation (confluens Zone) of Iowa is associated with 
a rich conodont fauna including Belodina confluens, 
Phragmodus undatus, Periodon grandis, and 
Drepanoistodus suberectus. Zhang (1998) reported this 
species from the Middle Ordovician Guniutan 
Formation (Darriwilian) of South China. It was also 
identified from the Upper Ordovician of North 
America (Winder 1966; Webers 1966; Nowlan et al. 
1988, 1997; McCracken and Nowlan 1989), 
Scandinavia (Hamer 1964) and north England 
(Orchard 1980). This is the first record of this species 
in the Ordovician of eastern Australia. 


39) 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Figure 5. SEM photographs of Late Ordovician conodonts from allochthonous limestones within the Barnby 
Hills Shale. A-C, Chirognathus? cliefdenensis Zhen and Webby, 1995; A, Pa element, AMF121418, ‘CWNN’; 
B, Sd element, AMF121419, ‘KIL’ 78.4; C, Sc element, AMF121420, ‘KIL’ 78.4. D-I, Drepanoistodus suberectus 
(Branson and Mehl, 1933); D, M element, AMF121421, ‘KIL’ 86.5; E, Sb element, AMF121422, ‘KIL’ 62; F, 
Sa element, AMF121423, ‘KIL’ 157.8; G, P element, AMF121424, ‘KIL’ 155.2: H, Sc element, AMF121425, 
‘KIL’ 157.8; I, Sa element, AMF121426, ‘KIL’ 95.7. J, “Oistodus” sp. cf. venustus Stauffer, 1935 s.f., M 
element, AMF121427, ‘KIL’ 110.7. K-M, Panderodus gracilis (Branson and Mehl, 1933); K, falciform element, 
AMF121428, ‘KIL’ 175.6; L, M, asymmetrical graciliform element, L, AMF121429, ‘KIL’ 175.6, M, 
AMF121430, ‘CWNN’. N, O, Panderodus panderi (Stauffer, 1940); N, short-based element, AMF121431, 
‘KIL’ 175.6; O, long-based element, AMF121432, ‘KIL’ 175.6. P-T, Panderodus serratus Rexroad, 1967; P, 
S, T, serrated arcuatiform element, P, AMF121433, ‘KIL’ 173.1, S, AMF121435, ‘KIL’ 175.6, T, AMF121436, 
‘KIL’ 173.1; Q, R, non-serrated falciform element, AMF121434, ‘KIL’ 173.1. U-W, Panderodus sp.; falciform 
element, U, AMF121437, ‘KIL’ 175.6; V, AMF121438, ‘KIL’ 175.6; W, AMF121439, ‘CWNN’; scale bars 
100 um. 


Genus DREPANOISTODUS Lindstrém, 1971 Synonymy 
Oistodus suberectus Branson and Mehl, 1933, p. 
Type species 111, pl. 35, figs 22-27. 
Oistodus forceps Lindstrém, 1955. Drepanoistodus suberectus (Branson and Mehl); 
Nowlan and McCracken in Nowlan et al., 
Drepanoistodus suberectus (Branson and Mehl, 1988, p. 16, pl. 3, figs 19-22 (cum syn.); 
1933) Dzik, 1994, p. 78, pl. 17, figs 2-6, text-fig. 
Fig. 5D-I 12b; Zhen and Webby, 1995, p. 282, pl. 3, 


40 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


figs 8-10 (cum syn.); Nowlan et al., 1997, 
pl. 1, figs 7-9; Zhen et al., 1999, p. 88, Fig. 
6.1-7; Furey-Greig, 1999, p. 310, pl. 2, 
figs1-3; Furey-Greig, 2000b, p. 137, Fig. 
5.8; McCracken, 2000, pl. 1, fig. 12, pl. 2, 
figs 20, 21; Leslie, 2000, Fig. 5.16-19; 
Sweet, 2000, Fig. 9.23-25. 


Material 

Seventy-six specimens (9 P, 6 M, 5 Sa, 12 Sb, 10 Sc 
and 34 undifferentiated elements) from limestone 
blocks ‘KIL-A’, ‘KIL-B’, ‘KIL-C’, ‘FT’, ‘CWNN’, 
“WKILN’ and ‘EURO’. 


Discussion 

Elements forming the quinquimembrate apparatus of 
this species are laterally compressed, with sharp 
posterior and anterior margins and smooth lateral faces. 
The P element is weakly asymmetrical with a convex 
outer lateral face and an inner-laterally curved cusp, 
and is characterised by having a triangular, anticusp- 
like extension at the antero-basal corner. The M 
element is geniculate with a robust cusp. The nearly 
symmetrical Sa element has a sub-erect cusp, an antero- 
posteriorly extended base, and a shallow, but open 
inflated basal cavity. The Sb element is asymmetrical, 
with a suberect to slightly reclined cusp, a strongly 
curved basal margin, and the base extended only 
posteriorly. The Sc element somewhat resembles the 
P element, but is strongly asymmetrical with a 
posteriorly reclined cusp, and apparently lacks the 
prominent antero-basal extension of the latter. The 
present material is identical with that from the Bowan 
Park Group and basal Malachi’s Hill Beds (Zhen et 
al., 1999), except that the P element from the 
limestones within Barnby Hill Shale shows a more 
strongly extended antero-basal corner. 


Genus PANDERODUS Ethington, 1959 


Type species 
Paltodus unicostatus Branson and Mehl, 1933. 


Panderodus serratus Rexroad, 1967 
Fig. 5P-T 

Synonymy 

Panderodus unicostatus serratus Rexroad, 1967, p. 
47, pl. 4, figs 3, 4. 

Panderodus serratus Rexroad; Cooper, 1975, p. 
993, pl. 1, figs 3-5, 7-9, 13, 14, 23; Nowlan 
et al., 1988, p. 23, pl. 8, figs 5-7; Miller, 
1995, pl. 1, figs 15, 16; Jeppsson, 1997, 
p.107, Fig. 7.4. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Material 

Six specimens (4 serrated arcuatiform, 2 falciform 
elements) from limestone blocks ‘KIL-B’ and ‘KIL- 
cr. 


Discussion 

Panderodus serratus ranges from the Upper 
Ordovician (Ethington and Schumacher 1969; 
McCracken and Barnes 1981; Nowlan and Barnes 
1981; Nowlan et al. 1988) where it is relatively rare, 
through the Lower and Middle Silurian (Miller 1995; 
Jeppsson 1997), where it is widely distributed although 
by no means common. The distribution patterns of 
Panderodus serratus and its abundant ubiquitous 
associate P. unicostatus in the Silurian suggested to 
Jeppsson (1997) that they represented two distinct 
species (rather than morphotypes of a single species), 
distinguishable on presence or absence of the serrated 
element. In our collections, the serrated element is only 
represented by four specimens, which are comparable 
with those recorded from the Upper Ordovician of 
Canada (Nowlan et al. 1988), except for their smaller 
denticles and more prominently inner laterally curved 
cusp. Two additional specimens, which are recognised 
as the nonserrated falciform element of this species, 
are laterally compressed with sharp anterior and 
posterior margins. 


Genus PERIODON Hadding, 1913 


Type species 
Periodon aculeatus Hadding, 1913. 


Periodon grandis (Ethington, 1959) 


Fig. 6D-L 
Synonymy 
Loxognathus grandis Ethington, 1959, p. 281, pl. 
40, fig. 6. 


Periodon grandis (Ethington); Bergstrom and 
Sweet, 1966, p. 363-5, pl. 30, figs 1-8 (cum 
syn.); Lindstrom in Ziegler, 1981, p. 243- 
244, Periodon-pl. 1, figs 13-18; Zhang and 
Chen, 1992, pl. 1, figs 13-16; Ding et al., in 
Wang, 1993, p. 190, pl. 35, figs 18-21; 
Zhen and Webby, 1995, p. 284, pl. 4, figs 
3, 4 (cum syn.); Zhen et al., 1999, p. 90, fig. 
8.19-8.21; Furey-Greig, 1999, p. 310, pl. 2, 
figs 21, 22, pl. 3, figs 1, 2. 


Material 

Two hundred and thirty specimens (36 Pa, 8 Pb, 123 
M, 63 S elements) from limestone blocks ‘KIL-C’ and 
“WKILN’. 


4] 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Figure 6. SEM photographs of Late Ordovician conodonts from allochthonous limestones within the Barnby 
Hills Shale. A-C, Paroistodus? nowlani Zhen et al., 1999; distacodiform (b) element, A, AMF121440, ‘KIL’ 
73; B, AMF121441, ‘KIL’ 145; C, AMF121442, ‘KIL’ 149.1. D-L, Periodon grandis (Ethington, 1959); D, M 
element, AMF121443, ‘KIL’ 152.6; E, Sc element, AMF121444, ‘KIL’ 166.4; F, Pa element AMF121445, 
‘KIL’ 155.2; G, Sb element, AMF121446, ‘KIL’ 157.8; H, Pb element, AMF121447, ‘KIL’ 173.1; I, Sc element, 
AMF 121448, ‘KIL’ 166.4; J, Pa element, AMF121449, ‘KIL’ 157.8; K, ?Sd element, AMF121450, ‘KIL’ 
173.1; L, Sa element, AMF121451, ‘KIL’ 173.1. M, Pseudobelodina dispansa (Glenister, 1957); AMF121452, 
‘KIL’ 70. N, O, Phragmodus undatus Branson and Mehl, 1933; N, Pa element, AMF121453, ‘KIL’ 70; O, Sc 
element, AMF121454, ‘KIL’ 60; P, Yaoxiangnathus wrighti Savage, 1990; Pa element, AMF121455, “KIL’ 


48. Q, Pseudooneotodus mitratus (Moskalenko, 1973); AMF121456, ‘KIL’ 173.1; scale bars 100 um. 


Discussion 

Bergstrom and Sweet (1966) reconstructed the species 
as having a seximembrate apparatus, and this concept 
has been accepted since then by most conodont 
workers. The holotype of the species is a ramiform 
specimen referred to as the Sb element (previously the 
form species Loxognathus grandis Ethington, 1959). 
The Sa position is taken by the form species 
Trichonodella insolita Ethington, 1959, the Sc element 
is the form species Eoligonodina magna Ethington, 
1959 (see Bergstrom and Sweet 1966; and Sweet 
1988), while the form species Prioniodina araea 
Webers, 1966 and Ligonodina tortilis Sweet and 
Bergstro6m 1962 were assigned to the Pa and Pb 
positions respectively. As admitted by Bergstroém and 


42 


Sweet (1966: 364), P. grandis and P. aculeatus 
Hadding, 1913, the likely direct ancestor of the former, 
are “similar in overall shape and in most morphologic 
features.” These authors suggested that the geniculate 
M element of P. grandis, as the most characteristic 
form, could be differentiated from the same element 
of P. aculeatus by having a large, subtriangular base 
with essentially straight basal margin, and by having 
denticles on the anterior margin closely appressed to 
it rather than developed into an anterior process. 
Lindstrém (in Ziegler 1981) suggested distinguishing 
these two species by the denticles along the anterior 
margin of the M element reaching higher towards the 
tip of the cusp, and by the greater number (about 6 or 
more) of denticles between the cusp and the biggest 


Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


denticle on the posterior process of the S elements in 
P. grandis. However, the present central NSW material 
exhibit a rather centrally, downwardly arched basal 
margin, and typically more than six smaller denticles 
between the cusp and the largest denticles on the 
posterior process. Specimens referred as P. grandis 
showing similar arched basal margin of the M element 
were also recorded previously from North America 
(Webers 1966). 


Genus TAOQUPOGNATHUS An in An et al., 1985 


Type species 
Taoqupognathus blandus An in An et al., 1985. 


Taoqupognathus blandus An in An et al., 1985 
Fig. 7E-H 
Synonymy 

1985 Taoqupognathus blandus An in An et al., p. 
104, pl. 2, figs 18, 19; An, 1987, p. 192, pl. 
30, fig. 20; An and Zheng, 1990, pl. 7, figs 
5, 6, 20; Zhen and Webby, 1995, p. 287, pl. 
6, figs 1-13; Zhen et al., 1999, p.94-96, Fig. 
14.10-16; Wang and Qi, 2001, pl. 2, fig. 15. 


Material 

Forty-one specimens (11 M, 3 Sb2, 1 Sc2, 6 Sc3, 16 
Sc5, 4 undifferentiated S elements) from limestone 
blocks “KIL-A’, “KIL-B’, “KIL-C’, ‘FT’, ‘KILN’, 
“WKILN’, and ‘EURO’. 


Discussion 

In the lower part of limestone block KIL-C, T. philipi 
and T. blandus co-occur. One specimen, from sample 
‘KIL’ 95.7 (Fig. 7E), shows features transitional 
between typical Sc5 elements of 7. philipi and T. 
blandus. Considering its rather prominent posterior 
bulging, we regard this element as an early 
representative of the Sc5 element of T. blandus. A 
specimen of the Sb2 element of T. blandus is also 
recovered from a slightly lower level (Fig. 7F). 
Specimens from the upper part of limestone block 
‘KIL-C’ with more prominent and stronger posterior 
bulging (Fig. 7G), are identical with those illustrated 
from the upper Belubula Limestone and Vandon 
Limestone (Zhen and Webby 1995). 


Taoqupognathus philipi Savage, 1990 
Fig. 71, J 
Synonymy 
Taoqupognathus philipi Savage, 1990, p. 828, fig. 
8.1-8.12; Zhen and Webby, 1995, p. 287, 
pl., 5, figs 7-22; non McCracken, 2000, p. 
194, pl. 1, fig. 28. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Taoqupognathus tumidus Trotter and Webby, 1995; 
Furey-Greig, 1999, p. 312, partim, only pl. 
4, figs 2, 9. 


Material 

Twenty-two specimens (5 M1, 8 M2, 1 Sc3, 8 Sc5 
elements) from the lower part of the limestone block 
*KIL-C’, and two doubtful specimens from limestone 
block ‘KIL-B’. 


Discussion 

Taoqupognathus philipi, the oldest species of the 
genus, is characterized by having slender, elongated 
elements, with only weakly developed posterior 
bulging on the S elements. It has been recorded from 


’ the Fossil Hill Limestone of the Cliefden Caves 


Limestone Group (Savage 1990; Zhen and Webby 
1995), and the lower Reedy Creek Limestone 
(Percival, Morgan and Scott 1999) of central New 
South Wales. Several Sc5 specimens showing the 
characteristic features of the species (Fig. 7H, I, J) were 
recovered from block ‘KIL-C’. In comparison with 
the holotype of T. philipi from the Fossil Hill 
Limestone (Savage 1990: fig. 8.11, 8.12), these 
specimens exhibit a weaker development of posterior 
bulging and a gently curved anterior margin. No P 
elements have been recovered from any of our samples. 
So far Taoqupognathus has only been 
recorded from Australia and China (Zhen 2001). 
McCracken (2000) reported the occurrence of T. philipi 
from the Frobisher Bay and Amadjuak formations of 
southern Baffin Island. McCracken’ s identification was 
based on two specimens, one from each formation, 
but only one supposed Sc5 element was illustrated 
(McCracken 2000: pl. 1, fig. 28). In our opinion, this 
specimen cannot even be assigned with any certainty 
to the Panderodontidae. It lacks any distinctive 
characters of Taoqupognathus, except for a 
superficially similar outline, especially the tip of the 
cusp. Similar shape outlines are also seen in some Early 
Ordovician taxa, like Macerodus Fahraeus and 
Nowlan, 1978 (also see Ji and Barnes, 1994). 


Taoqupognathus tumidus Trotter and Webby, 1995 


Fig. 7K 
Synonymy 
Drepanodus? altipes? Palmieri, 1978, pl. 2, figs 24, 
2d) 


gen. unident. Pickett, 1978, fig. 4. 

Belodina cf. B. blandus (An); Duan, 1990, p. 31, pl. 
Do 1D Te 

Taoqupognathus tumidus Trotter and Webby, 1995, 
p. 487, pl. 7, figs 10-24; Zhen et al., 1999, 


43 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Figure 7. SEM photographs of Late Ordovician conodonts from allochthonous limestones within the Barnby 
Hills Shale. A, Pseudobelodina sp. Zhen et al., 1999; AMF121457, ‘WKILN-C’. B-D, Protopanderodus liripipus 
Kennedy, Barnes and Uyeno, 1979; B, symmetrical element, AMF121458, ‘KIL’ 119.5; C, asymmetrical element, 
AMF121459, ‘KIL’ 173.1; D, weakly asymmetrical element, AMF121460, ‘KIL’ 173.1. E-H, Taoqupognathus 
blandus An in An et al., 1985; E, ?Sc5 element, AMF121765, ‘KIL’ 95.7; F, Sb2 element, AMF121766, ‘KIL’ 
91; G, Sc5 element, AMF121463, ‘WKILN-C’; H, Sc5 element, AMF121464, “‘WKILN-B’. I, J, Taoqupognathus 
philipi Savage, 1990; Sc5 element, I, AMF121465, ‘KIL’ 97.5, J, AMF121466, ‘KIL’ 100.1. K, Taoqupognathus 
tumidus Trotter and Webby, 1995; P element, AMF121467, ‘KIL’ 38. L-N, Webbygnathus munusculum Pickett 
and Furey-Greig, 2000; Pa element, AMF121468, ‘KIL’ 95. O, P, Yaoxianognathus sp.; O, Pa element, 
AMF121470, ‘KIL ‘100.1; P, Pb element, AMF121469, ‘KIL’ 48; Q, R, Yaoxianognathus? tunguskaensis 
(Moskalenko, 1973); Q, Sc element, AMF121461, ‘KIL’ 95; R, Sd element, AMF121462, ‘KIL’ 89; scale bars 
100 um. 


: Material 
p. 96, fig. 14.1-14.9; Percival, 1999, fig. ; : 
3.1, 3.2, 3.5; Packham et al., 1999, fig. = ee oe from the lower part of limestone 
3.14-3.16; Furey-Greig, 1999, p. 312, ee ary 
partim only, pl. 4, figs 1, 3-8. 


s Discussion 
T. th W d Zhou, 1998, p. 190, a" ' 2 ap) Dor 
PodeP bie fig. 4 ae on a P This species seems much wider in distribution than 


the other two stratigraphically older species in eastern 


44 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


Australia and China. After the submission of a review 
paper on the genus (Zhen 2001), YYZ had the 
opportunity to examine the late Professor An’s 
conodont collection from the Taoqupo section near 
Yaoxian (An and Zheng 1990: 81-87), housed at the 
Department of Geology, Beijing University. Four 
specimens referable to P, Sb1, and Sb2 elements of T. 
tumidus were recognised in the sample TP34Y-5, 
which was taken about 30 m below the occurrence of 
Favisitina sp. at the top of the Taoqupo Formation. 
T. ani Wang and Zhou, 1998 from the Upper 
Ordovician of the Tarim Basin is a form species based 
only on a single element, which is identical to the P 
element of T. tumidus from eastern Australia. 


Phylum BRACHIOPODA Duméril, 1806 
?Subphylum CRANIIFORMEA Popov et al., 1993 
?Class CRANIATA Williams et al., 1996 
?Order CRANIIDA Waagen, 1885 
?Family CRANIIDAE Menke, 1828 


unnamed ?crantid 
Fig. 8 ff-kk 


Material. 
Five ventral? valves from ‘KIL’ section at 53.5, 58, 
and 59 m. 


Discussion. 

Classification of these specimens is problematic, as — 
if in fact they are craniids — they are atypical of this 
group. The valves are assumed to be ventral in position 
because of the apparent and consistent presence of a 
pedicle foramen. If so, this would be the first 
recognition of this feature in the remarkably long 
history of the craniids, which have displayed 
morphological conservatism from their appearance in 
the Ordovician to the present day. Most representatives 
of the group (including all living forms) are cemented 
to the substrate by their ventral valves. Two genera 
(Orthisocrania and Pseudocrania) interpreted as free- 
living are known from the Ordovician, but neither 
possesses a foramen. 

The specimens from the Eurimbla block are 
subconical in profile, and circular to ovate in outline 
(although incompletely preserved, so that orientation 
is uncertain). There is a suggestion that growth may 
have been mixoperipheral, as there appears to be a 
straight hingeline in the largest valve (Fig. 8 kk). 
Ornament comprises coarse radial costae of irregular 
size; that extending from the pedicle foramen is often 
(but not always) the most pronounced. Internal 
musculature is not preserved, and the dorsal valve is 


Proc. Linn. Soc. N.S.W., 124, 2003 


unknown. For these reasons, the taxon is not formally 
named. No other craniids have been described from 
Lachlan Orogen strata of Late Ordovician age. 
However, their recognition may have been impeded 
by the fact that fragmentary remains would appear 
indistinguishable from other silicified debris retrieved 
from acid dissolution of limestones; it is only when 
the valves are reasonably intact that their affinities can 
be determined. 


Subphylum RHYNCHONELLIFORMEA Williams 
et al., 1996 
Class STROPHOMENATA Williams et al., 1996 
Order STROPHOMENIDA Opik, 1934 
Superfamily PLECTAMBONITOIDEA Jones, 1928 
Family LEPTELLINIDAE Ulrich and Cooper, 1936 
Subfamily LEPTELLININAE Ulrich and Cooper, 
1936 
Genus MABELLA Klenina in Klenina, Nikitin and 
Popov, 1984 


Mabella halis (Percival, 1991) 
Fig. 8 0-t 


Wiradjuriella halis Percival, 1991, p. 138-141, fig. 12: 
1-38; Percival 1992, fig. 4: 10, fig. 5: 37-38. 

Mabella halis (Percival, 1991), Cocks and Rong Jia- 
yu 2000, p. 322-323, fig. 208: 2f-i. 


Material. 
Six valves figured from ‘KIL’ section, plus several 
hundred additional specimens. 


Discussion. 

These specimens conform in all regards to the 
description given by Percival (1991). Mabella sp. is 
also known from younger (late Eastonian) strata at 
Gunningbland in the Junee-Narromine Volcanic Belt, 
where it was first described as aff. Leptellina sp. by 
Percival (1979) and subsequently illustrated as 
Wiradjuriella sp. in Percival (1992: fig. 6, 31). A 
potential ancestor of M. halis may be Leptellina? sp. 
from the Wahringa Limestone Member of the 
Fairbridge Volcanics, which is exposed at Bakers 
Swamp, less than 15 kms north of Eurimbla; this 
occurrence is of earliest Gisbornian (basal Late 
Ordovician) age (Percival et al. 2001). Other species 
referred to Mabella (according to Percival et al. 2001: 
228) include M. solida (Nikitin and Popov, 1984) and 
M. namasensis (Klenina, 1984), as well as M. conferta 
(Popov, 1985); all three are from Kazakhstan. 


45 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


ff 


Figure 8. Silicified brachiopods of Late Ordovician age from allochthonous limestone block “KIL’ within the Barnby Hills 
Shale; all photographs taken by digital camera of unwhitened specimens. All specimens from horizon “KIL’ 53.5 unless 
otherwise stated. Magnifications all x2 unless otherwise stated. a, Chaganella speciosa (Percival, 1991), ventral valve 
interior, MMF36922. b-h, Doleroides mixticius Percival, 1991; b, ventral valve interior, MMF36923; c, ventral valve 
interior, MMF36924; d, ventral valve exterior, MMEF36925; e, ventral valve exterior, MMF36926; f, dorsal valve interior, 
MMF36927; g, dorsal valve interior, MMF36928; h, dorsal valve exterior, MMF36929. i-l, Sowerbyella billabongensis 
Percival, 1991; i, dorsal valve interior, MMF36930; j, ventral valve interior, MMF36931; k, dorsal valve exterior, MMF 

36932; 1., dorsal valve interior, MMF36933. m, n, Skenidioides quondongensis? Percival, 1991; exterior and interior views 


46 Proc. Linn. Soc. N.S.W., 124, 2003 


Y.Y. ZHEN, I.G. PERCIVAL AND J.R. FARRELL 


(Figure 8 continued) (both x3) of incomplete ventral valve, MMF36934, from ‘KIL’ 59. o-t, Mabella halis (Percival, 
1991); 0, ventral valve exterior, MMF36935; p, ventral valve interior, MMF36936; q, ventral valve interior, MMF36937; 
r, dorsal valve exterior, MMF36938; s, dorsal valve interior, MMF36939; t, dorsal valve interior, MMF36940. u-y, 
Rhynchotrema oepiki Percival, 1991; u, ventral valve interior, MMF36941; v, x, dorsal valve interior and exterior, MMF36942; 
w, dorsal valve interior with interlocked posterior fragment of ventral valve; y, enlargement (x3) of posterior fragment of 
dorsal valve interior, showing crura, MMF36943. z, aa, Australispira disticha Percival, 1991; z, incomplete dorsal valve 
exterior, MMF36944; aa, dorsal view (x3) of juvenile conjoined valves, MMF36945. bb, cc, Zygospira carinata Percival, 
1991; dorsal and ventral views of conjoined valves, MMF36946, x4. dd, ee, Protozyga definitiva Percival, 1991; ventral 
valve exterior, and oblique lateral view of conjoined valves, dorsal valve to right, MMF36947, x4. ff-kk, unnamed craniid?, 
all x3; ff, gg, exterior and interior views of presumed ventral valve, MMF36948, from ‘KIL’ 58; hh, exterior view of 
presumed ventral valve, MMF36949, from ‘KIL’ 59; ii, exterior view of presumed ventral valve, MMF36950, from ‘KIL’ 
58; jj, exterior view of presumed ventral valve, MMF36951, from ‘KIL’ 59; kk, interior view of ?ventral valve, MMF36952. 


Family HESPEROMENIDAE Cooper, 1956 
Genus CHAGANELLA Nikitin, 1974 


Chaganella speciosa (Percival, 1991) 
Fig. 8a 


Tylambonites speciosa Percival 1991, p. 143-144, fig. 
13: 14-35; Percival 1992, fig 4: 11-3, fig. 5: 
46-47. 

Chaganella speciosa (Percival, 1991) Cocks and Rong 
Jia-yu 2000, p. 339, fig. 222: 2e-h. 


Material. 
One specimen, a fragmentary ventral valve from ‘KIL’ 
section at 53.5 m. 


Discussion. 

Percival (1991: 143) noted substantial similarities 
between TJylambonites and Chaganella, but 
differentiated these genera on lack of a pedicle callist 
in Chaganella and chilidial plates being partially fused 
in the latter rather than discrete as in Tylambonites. 
Such distinctions are now regarded as of specific rather 
than generic significance. 


Class RHYNCHONELLATA Williams et al., 1996 
Order ORTHIDA Schuchert and Cooper, 1932 
Superfamily PLECTORTHOIDEA Schuchert and 
LeVene, 1929 
Family PLECTORTHIDAE Schuchert and LeVene, 
1929 
Genus DOLEROIDES Cooper, 1930 


Doleroides mixticius Percival, 1991 
Fig. 8b-h 


Doleroides mixticius Percival 1991, p. 127, 129, fig. 
8: 17-39; Percival 1992, fig. 5: 19-21. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Material. 
Abundant valves from ‘KIL’ section at 53.5 m. 


Discussion. 

The characteristic Mimella-like appearance of the 
ventral muscle field, previously noted by Percival 
(1991) when establishing the species, is well in 
evidence in some specimens (eg, Fig. 8b) from the 
Eurimbla allochthonous block, where D. mixticius is 
one of the most dominant forms. In all other 
morphological aspects, this species conforms to the 
generic concept of Doleroides as summarised in the 
recently revised brachiopod Treatise (Williams and 
Harper 2000: 759). 


ACKNOWLEDGEMENTS 


Specimens documented in this paper were collected and 
processed in the course of Ph.D. thesis research by John 
Farrell, carried out through the Macquarie University Centre 
for Ecostratigraphy and Palaeobiology (MUCEP). Michael 
Engelbretsen (MUCEP) is thanked for his assistance in 
processing some of the samples. Zhen’s study of the 
conodont fauna was undertaken with the support of a research 
fellowship provided by Sydney Grammar School. Scanning 
electron microscopy was carried out at the Electron 
Microscope Unit of the Australian Museum. John Farrell is 
grateful to Elisabeth Arundell (nee Morgan) for field 
discussions on stratigraphy and allochthonous blocks and 
to Bill Ebzery who was indispensable in the field. Farrell is 
very appreciative to the landowners in the district — the Lee 
family, the Weston family, Ian Henry and Roy Barrett — 
who allowed ready access to their properties. The manuscript 
was improved by the careful reviews of Lawrence Sherwin 
and John Pickett. Ian Percival publishes with the permission 
of the Director General, NSW Department of Mineral 
Resources. This is a contribution to IGCP Project no.410: 
the Great Ordovician Biodiversification Event. 


47 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


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52 


ORDOVICIAN LIMESTONES IN SILURIAN SHALE, CENTRAL NSW 


Proc. Linn. Soc. N.S.W., 124, 2003 


The Middle Triassic Megafossil Flora of the Basin Creek 
Formation, Nymboida Coal Measures, New South Wales, 


Australia. Part 3. Fern-like Foliage. 


W.B.KeitH HoLMEs 


“‘Noonee Nyrang’, Gulgong Road, Wellington, NSW, 2820, Australia. (Hon. Research Fellow, Geology 


Department, University of New England, Armidale, NSW, 2351, Australia). 


Holmes, W.B.K. (2003). The Middle Triassic megafossil flora of the Basin Creek Formation, Nymboida 
Coal Measures, New South Wales, Australia. Part 3. Fern-like foliage. Proceedings of the Linnean Society 
of New South Wales 124, 53-108. 


Two quarries in the Basin Creek Formation of the Middle Triassic Nymboida Coal Measures have yielded 
numerous examples of fern-like foliage. No affiliated fertile material is available to place the fronds in a 
natural classification. Twenty three species in twelve genera are described as morpho-taxa in Order and 
Family Incertae Sedis. Plants described in this paper are:- Cladophlebis conferta sp. nov., C. octonerva sp. 
nov., C. paucinervasp. nov., C. retallackiisp. nov., C. sinuatasp. nov., C. fenuipinnulasp. nov., Dictyonymba 
sparnosa gen. et sp. nov., Gouldianum alethopreroides gen. et sp. nov., Leconama stachyophylla gen. et sp. 
nov., Micronymbopteris repens gen. et sp. nov., Vymbiella lacerata gen. et sp. nov., Nymboidiantum 
elossophyllum (Tenison-Woods) gen. et comb. nov., V. mu/tilobatum gen. et sp. nov., VV. elegans gen. et sp. 
nov., WV. fraciiflexum gen. et sp. nov., V. robustum gen. et sp. nov., Vymbophlebis polymorpha gen. et sp- 
nov., Vymbopteron dejerseyi (Retallack) gen. et comb. nov., VV. fo/ey/ gen. et sp. nov., VV. uncinatum gen. et 
sp. nov., Vymborhipteris radiata gen. et sp. nov., Plotonymba curvinervia gen. et sp. nov. and Sphenopreris 
speciosa sp. nov. The diversity of this new material demonstrates the remarkable recovery of Gondwana 


vegetation following the end-Permian extinction event. 


Manuscript received 9 May 2002, accepted for publication 27 October 2002. 


KEYWORDS: Gondwana Middle Triassic, megafossil fern-like foliage, Nymboida Coal Measures, 


palaeobotany. 


INTRODUCTION 


This paper is the third in a series describing the 
rich and diverse megafossil flora from two quarries 
near the village of Nymboida in north eastern New 
South Wales. A locality map and details of the geology 
of the Nymboida area were provided in Part 1 of this 
series (Holmes 2000) which dealt with the Thallophyta 
and Sphenophyta. Part 2 (Holmes 2001b) included 
descriptions of 14 taxa of the Filicophyta representing 
true ferns preserved in a fertile state or remains of 
sterile material with known fern relationships. 

This paper describes fern-like foliage of 
uncertain systematic position due to the lack of fertile 
material. It is acknowledged that some at least of these 
fossil plants are not true ferns. They may be new forms 
of pteridosperms or even belong to plant groups that 
are presently unknown. Twenty four species are placed 
in twelve genera. The names published below are 
defined as morpho-taxa under the provisions of the 
International Code of Botanical Nomenclature (ICBN 
2000). 


The diversity of plants with fern-like foliage 
that are described below is a remarkable demonstration 
of the recovery of the world’s vegetation following 
the end-Permian extinction event. That catastrophic 
and devastating event caused the disappearance of the 
Gondwana G/ossopreris Flora and up to 90% of the 
world’s living organisms. It brought about the cessation 
of all coal formation throughout the world. The “Coal 
Gap” (Retallack 1996) persisted until early Middle 
Triassic time. Some of the earliest coal seams following 
the “Coal Gap” are preserved in the Basin Creek 
Formation. 

Included in this Middle Triassic coal flora are 
some specimens which in gross morphology are 
closely similar to ferns. Although some are common 
and widespread, none have been found with associated 
or identifiable fertile remains. Most of the morpho- 
taxa are rare and in some cases are based on a single 
specimen or just a few individuals. Even though 
specimens may be rare and fragmentary, provided there 
are significant diagnostic features for differentiation, 
they are illustrated and described in order to make 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


known their presence in the Nymboida Flora. The 
minimum criteria for formal naming are that the 
specimens demonstrate the attachment of pinnae to a 
main rachis and preferably with pinnules showing a 
reasonable state of preservation of the venation. 
Lebedev (1974) believed that through the use of good 
descriptions, accurate drawings and adequate 
photographs, sterile fern fronds could be confidently 
identified and even placed in natural groupings. In this 
paper I have attempted as far as possible to follow the 
guidelines of Lebedev. However, the novelty of most 
specimens, the lack of associated fertile material and 
the state of preservation precludes the placement of 
this material in a natural classification. 

A tectonic heating event during the Cretaceous 
(Russell 1994) has destroyed the cuticle of otherwise 
often exquisitely preserved leaves and fruits of the 
entire flora at the two Nymboida localities. There is 
great scope for further investigations from localities 
where cuticle and fine cell structure may be preserved. 

As in extant floras, the assemblages of fossil 
plants are directly related to their habitats. Several 
species of one genus may co-exist in a limited area 
but each within its own habitat. The presence of several 
species in the Cladophlebis, Nymbopreron and 
Nymboidiantum genera may relate to the taxa being 
derived from differing facies. 

The sediments included in the 20 metres depth 
exposed in the working faces of the Nymboida quarries 
range from coarse conglomerates to fine shales and 
coal bands. Each horizon represents a flood event and 
a particular facies that existed through a short period 
of geological time (Holmes 2000) on an alluvial 
floodplain. Fossil soils formed during pauses in the 
deposition of sediments are also present The 
reconstruction of the Middle Triassic Nymboida 
floodplain by Retallack (1977) demonstrates the range 
of habitats that existed simultaneously. The fossil 
assemblages, supported by facies evidence, range from 
in-situ accumulations and fossil soils (autochthonous) 
to partially dispersed (semi-authochthonous) to long 
distance transport dispersal and fragmentation 
(allochthonous). 

The origins and relationships of the majority 
of plants in the Early to Middle Triassic Gondwana 
floras are problematical. Due to the virtual spatial and 
climatic isolation of Gondwana and especially eastern 
Australia from the Northern Hemisphere following the 
end-Permian extinction event, I believe there is dubious 
value in attempting to determine close relationships 
between plants from those two macro-regions based 
only on gross morphology. This problem, relevant to 
sphenophytes, was discussed by Holmes (2001a). In 
this paper, with a few exceptions, I have compared 
the Nymboida fern-like foliage only with previously 


54 


described Gondwana material. Where there is no 
published description matching my material I have 
erected new species and in some cases, new genera. 
These new taxa will provide a reference for 
comparisons with other Gondwana assemblages and, 
hopefully, will be the foundation for future studies 
based on better preserved and more complete material 
that will allow the fossil plants to be placed in a natural 
classification system. At that stage, more meaningful 
comparisons may be made with fossils from Triassic 
northern floras. 

Most specimens are illustrated at both natural 
size and enlarged. The size of reproduction is indicated 
in the Figure legends and by a bar measure representing 
one centimetre on each photograph. 

All the described, illustrated and mentioned 
specimens in this paper have been lodged with the 
Australian Museum, Sydney and have been catalogued 
with AMF numbers. 


SYSTEMATIC PALAEOBOTANY 


Order Incertae sedis 
Family Incertae sedis 
Genus C/adophlebis Brongniart 1849 
Type species Cladophlebis albertsii (Dunker) 
Brongniart 1849 


The genus C/adophlebis was erected by Brongniart 
(1849) for sterile fern fronds from both the Late 
Palaeozoic and Mesozoic. It was generally regarded 
as a form genus. Fertile material has been placed 
variously in a number of natural genera. In an attempt 
to define C/adoph/ebis in a strict sense as a natural 
genus, Frenguelli (1947) carried out an extensive 
review of over 150 species and forms that had been 
attributed to Cladoph/ebis. On his restricted definition 
he recognised 28 Mesozoic species worldwide, 
including 14 species from the Triassic to Cretaceous 
in Argentina. Later Herbst (1971, 1978) revised both 
the Argentinean and Australasian species of 
Cladoph/ebis. For Australasia he retained as valid 
species the variable and long-ranging C/adoph/lebis 
australis with a synonymy list of 26 entries, and C 
mendozaensis. C. patagonicaand C. gondwanicawere 
regarded as doubtful. However, the Nymboida Flora 
is rich in ferns and fern-like foliage and contains at 
least six species described below which are referable 
to this genus. In Part 2 of this series, fertile ferns were 
placed in Asterotheca, Rhinipteris, Todites, 
Herbstopteris, Osmundopsis and Nymbofelicia 
(Holmes 2001b). In most cases the associated sterile 
fronds would have fitted broadly in Cladophilebis. 

Sterile fronds with no associated fertile material 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


are here placed in the morpho-genus C/adophlebis 
which includes bipinnate fronds, pinnules separated 
to the base, broadly attached to pinna rachis and 
variously decurrent, base occasionally slightly lobed 
either basiscopically or acroscopically, entire to 
slightly lobed or serrate, parallel-sided to slightly 
tapering, straight or variously falcate; midvein 
prominent and usually persistent almost to the apex, 
lateral veins alternate, forking once, twice or rarely 
three times, often simple distally. It must be understood 
that this is an artificial classification based merely on 
gross form and the individual morpho-taxa may belong 
in a range of natural genera. 


Cladophlebis conferta Holmes sp. nov. 
Figures 1A—C 


Diagnosis 

Medium sized bipinnate frond; pinnae closely 
spaced at high angle to main rachis; pinnules opposite 
to subopposite, broadly falcate, midvein fine, four pairs 
of lateral veins, proximal two pairs twice forked, 
second pair once forked, distal pair unforked or once 
forked close to margin. 


Description 

This taxon is based on two specimens each 
showing a midportion of bipinnate fronds that may 
have possibly reached one metre long in life. 
AMF 121014 shows a main rachis tapering from 8 mm 
to 6 mm over a length of 300 mm. The holotype frond 
fragment (Fig. 1A) has a smooth main rachis 5 mm in 
width with seven pairs of overlapping opposite pinnae 
attached at c. 80° to main rachis at intervals of c. 10 
mm. Pinnae linear, tapering in distal half to acute apex, 
to 100 mm long, 15 mm wide. Pinnules opposite to 
alternate, closely spaced to overlapping, to 8 mm long, 
5 mm wide; length to width ratio of c. 1.6:1; attached 
at c. 45° at midpinna and at a higher angle closer to 
the main rachis; basiscopic margin decurrent, slightly 
contracted above base, margin broadly convex, 
acroscopic margin straight to slightly convex, apex 
obtuse; midvein fine, once forked before the pinnule 
apex; with four pairs of alternate lateral veins at c. 35° 
to midvein; first two pairs twice forked, second pair 
once forked and distal pair usually simple; forking very 
acute with c. 20 vein endings around pinnule margin 
(Fig. 1C). 


Holotype 
AMF 120987, and portion of the counterpart as 
isotype AMF121015. Australian Museum, Sydney. 


Type locality 


Proc. Linn. Soc. N.S.W., 124, 2003 


Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF121014, Reserve Quarry, Nymboida. 


Name derivation 
conferta — confertus, (Lat.), dense, crowded, 
referring to the closely spaced pinnae and pinnules. 


Discussion 

Although known from only two specimens, the 
preservation is sufficient to provide diagnostic 
characters. C. conferfapinnules have a similar number 
of lateral veins as C. octonervia (described below), 
but differ by the shorter broader pinnules, the more 
acute and radiating form of the forking veins and by 
the fewer vein endings around the pinnule margin. By 
the closely spaced, long, opposite pinnae and by the 
form of the sparse and fine venation, C’ con/er/a differs 
from all other Gondwana C/adoph/ebis spp. 


Cladophlebis octonerva Holmes sp. nov. 
Figures 2 A—C 
1921 — Cladophlebis mesozoica vat typica Kuriz pl. 
30, Fig. 6. 


Diagnosis 

Medium sized bipinnate frond, pinnae opposite 
to alternate, pinnules thick-textured, alternate, oblong 
to triangular, entire or occasionally lobed, apex obtuse, 
median vein weak, continuing to apex, four pairs of 
lateral veins once forked close to midvein then 
diverging to margin. 


Description 

Complete fronds not available. The holotype 
(Fig. 2A), a 180 mm long midportion of a narrow 
elliptic bipinnate frond, suggests a total frond length 
of c. 400 mm. The main and pinna rachises are strongly 
ribbed and grooved. Pinnae opposite to alternate, 
attached basally at right angles or obtuse, in the middle 
of the frond at c. 80° to 90°, distally more acute; linear, 
longest in midportion of frond, to 60 mm long and 14 
mm wide, tapering from midway to the acute apex. 
Pinnules alternate, oblong to triangular or straight to 
slightly falcate, attached at 90° to 60° to pinna rachis, 
basiscopic margin slightly contracted, acroscopic base 
slightly decurrent upwards, margin more convex on 
proximal side, entire to slightly lobed, apex obtuse, 
length to width ratio, c. 2—2.5:1. Midvein continuing 
to the apex, four pairs of lateral veins each once broadly 
forked close to the midvein; venules continuing to 


>) 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


diverge straight or slightly arching to the margin; 
number of lateral veins decreasing on smaller pinnules 
distally and apically. First lateral veins occasionally 
forking again near the margin (Fig. 2C). Number of 
vein endings around margin c. 16. 


Holotype 
AMF 113484, Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF113485-86. Coal Mine Quarry, 
Nymboida. 


Name derivation 

octonerva — octo, (Lat.), eight; nerva, (Lat.) 
veins, referring to the usually four pairs of arching 
lateral veins in each pinnule. 


Discussion 

C. octonerva differs from other Nymboida 
Cladoph/ebis spp. by the shorter broader pinnules with 
four pairs of once-forking lateral veins in the pinnules. 
A frond fragment illustrated in Kurtz (1921, pl. 30, 
Fig. 6) from the Triassic of Argentina, as C’ mesozoica 
var. Aypica has pinnule form and once-forking lateral 
venation closely similar to C: ocfonerva. Stipanicic et 
al. (1995), in a revision of the material figured in Kurtz 
(1921), followed both Frenguelli (1947, pl. 7, Figs 2,5) 
and Herbst (1971, Figs 17, 19), who defined C. 
mesozoica as having twice forked lateral veins. The 
C mesozoicavat typica specimen was not accepted as 
C. mesozoica and was merely identified as C. sp. By 
restricting C: mesozoica to leaves with twice forked 
lateral veins, forms such as C. ocfonervaand C. sinuata 
(described below), which are only partially twice 
forked, are excluded from that taxon. 


Cladophlebis paucinerva Holmes sp. nov. 
Figures 3A-G 


Diagnosis 

A small bipinnate frond with short narrow- 
ovate, opposite to alternate pinnae; pinnules 
alternate, broad-ovate, apex obtuse, length to width 
ratio of c. 2:1; midvein straight, dividing well before 
the apex, with two pairs of alternate lateral veins 
once-forking halfway to margin. 


Description 


This taxon is based on a number of small 
midportions of incomplete fronds to 35 mm long, 


56 


which bear to five pairs of sub-opposite short pinnae; 
estimated total length of frond in life c. 50 mm. Rachis 
1 mm wide, strongly ribbed and grooved; pinnae well- 
spaced, attached at c. 45°, narrow ovate to 15 mm long 
and 6 mm wide, tapering from halfway to the acute 
apex with c. 6 pairs of alternate pinnules. Pinnules 
broadly attached at c. 45°, decurrent basiscopically, 
slightly contracted acroscopically, broad-ovate, 3-4 
mm long, 1.5—2 mm wide, becoming more triangular 
in midportion of pinna then coalescing apically; length 
to width ratio of c. 2:1; pinnule margin entire or slightly 
serrate; midvein straight or faintly undulate, forking 
well before the apex; two pairs of alternate lateral veins 
attached at a very acute angle, first pair once-forked 
halfway or closer to the margin, distal lateral veins 
simple (Fig.3A,D); six to eight vein endings around 
margin. 


Holotype 
AMF 120979; isotype AMF 120980, Australian 
Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF120981-120984. 


Name derivation 
Daucinerva—paucus, (Lat.), jew, nervus (Lat.), 
nerve, referring to the small number of lateral veins. 


Discussion 

Portions of two fronds are preserved on one 
small slab (Fig. 3C) but their appearance does not 
suggest that they are detached pinnae of a tripinnate 
frond. A pinna fragment from Argentina which was 
illustrated by Kurtz (1921, pl. 32, Fig. 11) as Asplenium 
whitbyense has small pinnules and venation similar to 
Cladophlebis paucinerva. Stipanicic et al. (1995) have 
reclassified that specimen as Cladophlebis (Todites?) 
ugartei Herbst 1964. As fertile fronds of C. paucinerva 
are not known, it is difficult to make a closer 
comparison with C. ugartei. C. paucinerva differs from 
all other known Gondwana Triassic Cladophlebis spp 
by its small size and the few lateral veins. The small 
creeping fern Af%icronymbopreris repens described 
below differs by the elongated pinnae and much 
smaller pinnules with no obvious venation. C. parva 
(Fontaine) Bell (1956), a Cretaceous fern from North 
America, has pinnules similar in size to C. paucinerva 
but differs by the tripinnate form and by the sometimes 
trilobed first basiscopic pinnules. 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Cladophlebis retallackii Holmes sp. nov. 
Figures 4A,B; 5A,B 
1977 — Cladophlebis gondwanica non Frenguelli, 
Retallack in Retallack et al. p.86, Figs D-F. 


Diagnosis 

Medium to large bipinnate frond with robust 
primary rachis and opposite pinnae rachises. Pinnules 
opposite, straight to slightly falcate, slightly constricted 
above the basiscopically and acroscopically decurrent 
base, margins entire, parallel for c. half length then 
basiscopic margin curves apically to form acute to 
obtuse rounded apex. Median vein strongly decurrent, 
basal lateral veins attached to the primary rachis at the 
base of the midvein and thrice forked. Distally, lateral 
veins arch away from midvein, forking up to three 
times proximally, thence twice in mid-region and once 
apically. 


Description 

A robust bipinnate lanceolate frond with stout 
rounded primary rachis and pinnae rachises. In life 
possibly to one metre in length. Figure 4A is a portion 
of a large frond, primary rachis to c. 8 mm wide near 
broken lower section, base missing; pinnae opposite 
on lower portion of the frond, upwards becoming 
subopposite and then alternate apically, linear to 100 
mm long and 20 mm wide, basal pinnae attached at 
right angles, becoming more acute, to 45° apically. 
Pinnules opposite to sub-opposite (Figs 4B, 5B), 
overlapping to well-separated, mostly oblong, straight 
to slightly falcate, slightly contracted in width above 
the decurrent base, attached at right angles close to 
primary rachis, becoming more acute distally, to 14 
mm long and 5 mm wide, with length to width ratio of 
c. 2.5 (2—3.4):1. Midvein strongly decurrent on pinna 
rachis then decurving into pinnule and continuing 
almost to pinnule apex where it divides into two short 
veinlets. First basiscopic and acroscopic lateral veins 
attached at or just below the base of the median vein, 
basiscopic vein arching and forking three times to meet 
margin at a high angle, first acroscopic vein also 
branching three times with the adaxial vein running 
parallel to the pinna rachis. The subsequent five to 
seven pairs of lateral veins attached at c. 30° to midvein 
and arching to meet the pinnule margin at c. 45° to 
60°, mostly twice forked then once forked near the 
pinnule apex, with c. 40-50 vein endings around the 
pinnule margin. 


Holotype 
AMF 120959, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 


Proc. Linn. Soc. N.S.W., 124, 2003 


Formation, Nymboida Coal Measures. Middle Tniassic. 


Other material 

UNEF133363-4, AMF120954-120959, 
120993 and 121167, Coal Mine Quarry. The material 
from the Cloughers Creek Formation of the Nymboida 
Sub-Basin referred by Retallack (in Retallack et al. 
1977), (see below), to Cladophlebis gondwanica and 
included here in Cladophlebis retallackitis also housed 
in the Australian Museum collections. 


Name derivation 

retallackii — for Dr G.J. Retallack who carried 
out research on the Nymboida Flora; pioneer 
investigator of fossil soils. 


Discussion 

Retallack (in Retallack et al. 1977) illustrated 
and briefly described fronds from the Cloughers Creek 
Formation of the Nymboida Coal Measures which he 
assigned to Cladophlebis gondwanica Frenguelli. 
Herbst (1978), with reservations, also included this 
material in C. gonawanica. Retallack noted that similar 
fronds also occurred in the Basin Creek Formation. 
This is confirmed by my collections, some of which 
are here illustrated and form the basis for the new taxon 
C. retallackiz. The material selected by Frenguelli 
(1947) as the type for C. gondwanica was based on 
material from Tonkin in Vietnam that had earlier been 
identified by Zeiller (1903) with the European species 
C. roessertii(Presl) Krystofovich. The Tonkin material 
illustrated by Frenguelli (1947, Fig.19) has a similar 
length to width ratio as C. rera//ackii but differs by the 
more slender, canaliculated primary and pinnae 
rachises, by the alternate pinnules with straight lateral 
veins and fewer vein endings around the pinnule 
margins. Frenguelli included in C. gondwanicaa frond 
from the Carnian Molteno Formation of South Africa 
which had been briefly described by Seward (1908, 
p.98, pl.8) and assigned to C. (Todites) roessertii. 
Seward’s description and one illustration are lacking 
in details and I have not examined the specimen. From 
the illustration, the length to width ratio is lower and 
the pinnae are more acutely attached to the main rachis, 
so its affinity with C. refa//acki7is doubtful. With the 
exception of the material described by Retallack (in 
Retallack et al. 1977), there are no other records of 
fronds similar to C. reta//ackii in Triassic Gondwana 
floras. 


Cladophlebis sinuata Holmes sp. nov. 
Figures 6A-C 
1921 — Cladophlebis mesozoica forma typica Kurtz 
pl. 32, Fig. 13 
1921 — Cladophlebis mesozoica forma crenulata 


ad 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Kurtz pl. 32, Fig. 14 
?1982- Cladophlebis mesozoica Holmes p. 5, Fig. 
3A 


Diagnosis 

Medium sized bipinnate frond; pinnae opposite, 
well-spaced, attached at a high angle. Pinnules 
alternate, elongate-triangular, closely spaced, margin 
irregularly dentate or lobed. Midvein fine, sinuate, 
forking near the apex, c. five pairs of lateral veins 
attached to median vein at c. 45°. First two pairs of 
veins twice forked, following veins once forked. 


Description 

Cladophlebis sinuata is based on seven 
specimens. The holotype (Fig. 6A) is a portion of a 
bipinnate frond showing three pairs of pinnae attached 
opposite at c. 90° to a conspicuously ribbed and 
grooved primary rachis 4.5 mm wide. The complete 
frond is estimated to have been c. 400 mm long. The 
pinnae are well separated, 45 mm and 35 mm apart, 
with a length perhaps of 150 mm. Pinnules sub- 
opposite to alternate, closely spaced to overlapping, 
broadly attached at c. 80° to ribbed pinna rachis, 
basiscopic margin contracted and acroscopic base 
expanded, elongate-triangular to slightly falcate, to 15 
mm long and to 10 mm wide just above the base; length 
to width ratio of c. 1.6:1, becoming smaller and more 
acutely attached towards the pinna apex, margin 
broadly lobed or dentate, apex broadly acute to obtuse. 
Midvein fine, slightly undulate to sinuate, forking close 
to apex; four to five pairs of alternate secondary veins 
leave midvein at c. 45°; the proximal two pairs of veins 
fork twice, the distal two pairs fork once with the last 
one or two veins unforked (Fig. 6C); 18-24 vein 
endings around pinnule margin. 


Holotype 
AMF113512, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF113513-113518, AMF113541, Coalmine 


Quarry. 


Name derivation 
sinuata — sinuatus, (Lat.), wavy — referring to 
the course of the fine median vein. 


Discussion 


Cladophlebis sinuata is close to C. wielandii 
Jain and Delevoryas (1967) from the Carnian Cacheuta 


58 


Formation of Argentina, but differs by the typically 
twice forked lateral veins in the basal portion of the 
pinnule lamina. Material from the Carnian Ipswich 
Basin, referred to C’ concinnaby Jones and de Jersey 
(1947, text Figs 4 and 5) has closely spaced pinnules 
with sinuous midveins. Their text fig. 4 differs from 
C. sinuata by the lateral veins which all fork twice, 
and text fig. 5 differs by the lateral veins which are all 
once forked in a manner similar to those referred to C. 
concinna by DuToit (1927) from the Carnian Molteno 
Formation of South Africa. C. s/nuara is similar to 
some forms of the variable C. mendozaensis (Geinitz) 
Frenguelli 1947, Herbst (1971, Figs 13, 14 and 21) 
from the Upper Triassic of South America, but differs 
by the finer sinuous midrib, by the more closely spaced 
to overlapping pinnules and by the generally lower 
length to width ratio of the pinnules. The Upper 
Triassic Queensland material which was identified by 
Herbst as C. mendozaensis (Herbst 1978, pl 1, Figs 4 
and 5, pl. 3, Figs 16-18) has alternate pinnae and a 
significantly higher length to width ratio. Specimens 
illustrated in Kurtz (1921, pl. 32, Figs 13, 14) as C. 
mesozoica forma typica and C. mesozoica forma 
crenulata have a sinuous midvein and similar lateral 
venation to C. s‘nuata. These two Carnian Cacheuta 
Formation specimens of Kurtz were placed by 
Stipanicic et al. (1995) in C’ mesozoica, a taxon defined 
by its twice forking lateral veins, as discussed under 
C. octonerva. Frond fragments from the Middle 
Triassic Benolong Flora of central-western New South 
Wales which were placed in C. mesozoica by Holmes 
(1982) are similar in venation pattern and length to 
width ratio, but are of smaller size and have less vein 
endings around the pinnule margin. Another sterile 
pinna fragment from the same assemblage attributed 
to Zodites pattinsoniorum Holmes (1982, Fig. 2D) also 
has similar venation to C. s‘zwara, thus suggesting that 
the latter could be the foliage of an osmundaceous fern. 


Cladophlebis tenuipinnula Holmes sp. nov. 
Figures 7A, 8 A-C 


Diagnosis 

Large tripinnate frond, tertiary pinnae alternate, 
linear; pinnules alternate, small, straight, closely 
spaced; lateral veins well-spaced and once forked. 


Description 

Large tripinnate frond (Fig.8A), primary rachis 
35 mm wide near base which suggests a total frond 
length of from two to three metres. Secondary rachises 
(Fig. 7A) well separated, to 5 mm wide, straight, to 
200 mm long. Secondary rachises in basal region 
obtuse, changing to right angles in mid-frond then 
slightly acute apically. Tertiary rachises at high angle 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


to secondary rachis, c. 10 mm apart, linear to 50 mm 
long, c. 10 mm wide. Pinnules alternate, attached by 
whole base, linear, straight to slightly curved, 4-5 mm 
long, 1-1.5 mm wide, length to width ratio of c. 4:1, 
closely spaced, free to base, margin entire, apex obtuse. 
Pinnule lamina apparently thick textured which 
obscures the venation. On some pinnules on 
AMF 113526, which are preserved at an angle to the 
bedding plane (Fig. 8B), the venation is faintly evident 
and shows a fine midvein with c. 8 pairs of lateral veins 
attached at a high angle and once-forked close to the 
midvein; c. 30 vein endings around the margin. 
Fragmentary fertile pinnules appear to bear five or six 
pairs of As/erorheca-like synangia but the preservation 
is too poor to determine the structure. 


Holotype 
AMF113523; isotypes AMF113524-27, 
AMF113406, Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF121016-121018, Reserve Quarry. 


Name derivation 
tenuipinnula— tenuis, (Lat.), Jong and slender, 
referring to the attenuated pinnules. 


Discussion 

This is one of the largest fern fronds in the 
Nymboida Flora, but is known only from dispersed 
fronds on a single horizon. Some very fragmentary 
fertile material is available and while the preservation 
is very poor, the distribution and size of the sori on the 
pinnules appear to differ from all Asterotheca fronds 
previously described from Nymboida (Holmes 2001b). 
By the large frond size, by the very stout secondary 
rachises and by the very small densely spaced 
attenuated pecopteroid-like pinnules, C. texuipinnula 
differs from all other Gondwana Triassic ferns. 


? Cladophlebis sp. A 
Figures 9 A-C 


Description 

Several frond fragments with characteristic 
narrow-elliptic pinnae with the first pinnules well 
detached from the main rachis, are here placed 
doubtfully in C/adoph/ebis due to lack of preserved 
venation. The available specimens indicate living 
fronds were from 80 mm to 150 mm long. The pinnae 
are opposite to sub-opposite, attached in lower portion 


Proc. Linn. Soc. N.S.W., 124, 2003 


of frond at c. 90° and becoming more acute (to 60°) 
apically. Pinnae elliptic to 35 mm long, with four to 
twelve opposite triangular to rhombic pinnules attached 
at c. 45° to 80°; pinnules separated proximally but soon 
coalescing and decreasing in size towards the acute 
apex; venation obscured by the obviously thick texture 
of the pinnules. 


Material 
AMF113508-113510, Coal Mine Quarry, 
Nymboida. 


Discussion 

The three specimens illustrated (Figs 9 A-C) 
show a range of variation but appear to represent 
intergrading forms. There are some similarities with 
foliage fragments of the seed ferns Dicroidium and 
Lepidopteris. However there is no indication that the 
fronds may have been forked as in Dicroidium, and 
the lack of pinnules on the main rachis precludes 
affinities with Zeprdopreris. 


?Cladophlebis sp. B 
Figures 9 D, E 


Description 

An apical portion of a tiny bipinnate fern, 30 
mm long, pinnae opposite, pinnules broadly attached 
and well-spaced, rounded, c. 2 mm in diameter. The 
venation is poorly preserved. 


Material 
AMF 120978, Coal Mine Quarry, Nymboida. 


Discussion 

Only a single specimen of this tiny sterile fern 
frond has been collected. It is illustrated to draw 
attention to its presence. From the matching size and 
form, this could be a sterile frond affiliated with fertile 
Todites parvum described previously from Nymboida 
in Holmes (2001b). 


? Cladophlebis sp. C 
21883 Alethopieris currani Tenison-Woods 1883 p. 
77, pl. 6, Fig. 4 
Figures 10A, B 


Description 

A small fragment of a pinnate (? bipinnate) 
frond shows eight pairs of well-spaced opposite 
decurrent elongate, slightly falcate pinnules with 
serrate margins, attached at c. 60° to a slender ribbed 
rachis; midvein decurrent, with four pairs of once- 
forked lateral veins attached at an acute angle to the 
midvein. Length to width ratio of c. 4:1. 


59 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Material 
AMF 120994, Coal Mine Quarry, Nymboida. 


Discussion 

? Cladophlebis sp. C has (?pinnae) pinnules 
somewhat similar in shape and venation to those of 
Alethopteris currani Tenison-Woods (1883) from the 
Middle Triassic Napperby Formation at Ballimore near 
Dubbo in central-western New South Wales, but differs 
by the wider spacing of the pinnules. The Ballimore 
and Nymboida specimens are too incomplete to 
warrant formal naming. 


Dictyonymba Holmes gen. nov. 
Dictyonymba sparnosa Holmes gen. et sp. nov. 
Figures 10 C, D 


Combined diagnosis 

Bipinnate frond; pinnules broad ovate, 
conjoining, tapering to obtuse apex; lateral veins 
forking and occasionally joining to form coarse 
irregular reticulations in the conjoined area between 
the pinnules and towards the distal margin of each 
pinnule. 


Description 

D. sparnosa is described from a single 
fragmentary specimen showing portions of two parallel 
pinnae which bear partially conjoined pinnules with 
well-preserved venation (Fig. 10C). The size and form 
of the complete frond is not known. The larger pinna 
fragment is c. 39 mm long, bearing nine pairs of 
opposite pinnules attached at c. 60° to the pinna rachis, 
which tapers from 1 mm wide at the broken base. 
Pinnules broadly ovate but somewhat variable in shape, 
with convex entire margins, apices rounded, 5—6.5 mm 
long, 3.3-4.6 mm wide (Fig. 10D). Adjacent pinnules 
are conjoined for one third to one half of their length. 
Three to four pairs of once to three times forked lateral 
veins are attached at c. 45° to 60° to a slightly sinuous 
midvein. A separate vein enters the basiscopic portion 
of each pinnule lamina directly from the pinna rachis, 
forks three times and then meets in the conjoined region 
with some of the lateral veins from the adjacent pinnule 
to form an irregular mesh. In the distal free region of 
each pinnule where the lateral veins fork once or twice 
and sometimes anastomose, there are c. 18 vein endings 
around the margin. 


Holotype 
AMF113507. Australian Museum, Sydney. 


Type locality 


Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


60 


Name derivation 

Dictvonymba - dictyos, (Gt.) net, referring to 
the net venation and the Nymboida locality. 

sparnos - (Gt.) scarce, rare, recognising that 
one fragment only has been collected. 


Discussion 

Dictvonymbais a monotypic genus represented 
only by the type species D. sparnosa. 

Dunedoonia reticulata Holmes (1977) is a 
pinnate fern-like frond with reticulate venation from 
the Late Permian of eastern Australia. Dunedoonia 
reticulata differs from Dictyvonymba sparnosa by the 
basally contracted and much larger, broader pinnae 
and by the form of the anastomoses. 

Lonchopreris and Lonchopteridium spp from 
the Carboniferous of Europe (Boureau 1975) and 
Emplectopreris Halle (1927) from the Permian of 
China are fern-like bipinnate fronds with varying forms 
of anastomosing venation. The form of the pinnules 
and venation pattern of Dictvonymba sparnosa differs 
in detail from any species described in the above 
genera. The differences in geographical distribution 
and time also strongly suggest that this Nymboida form 
is at least generically distinct. Dictvonymba sparnosa 
is unique in Gondwana Triassic floras. 


Gouldiopteris Holmes gen. nov. 
Gouldiopteris alethopteroides Holmes gen. et sp. 
nov. 

Figures 11A-D 


Combined diagnosis 

An alethopteroid pinnatifid frond; pinna lobes 
with a distinct midrib, opposite, broad-linear, margin 
entire or slightly undulate, well-spaced, conjoined by 
a broad wing along primary rachis. Secondary veins 
closely spaced, arising directly from the main rachis 
and from the midvein in each pinna lobe at an acute 
angle (c. 45°), forking once, arching slightly then 
running parallel to wing and pinna lobe margins. 


Description 

This is a rare element in the Nymboida Flora. 
Only three specimens and their counterparts have been 
collected. The form and dimensions of complete fronds 
are not known. The holotype (Figs 11A, B) is a mid- 
section 55 mm long of a pinnatifid frond. Four pairs 
of opposite lobes are attached to the 3 mm wide primary 
rachis at c. 70° to 85°. The lobes are well-separated, c. 
15 mm apart, broad-linear, 45 mm long and 10 mm 
wide, bases strongly decurrent to form a conjoining 
wing c. 2-3 mm wide along the main rachis, lobe 
margins straight or slightly undulate, apex broadly. 
acute. A broad tapering midvein runs the length of 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


each lobe and forks close to the apex. Lateral veins 
from both the main rachis and midvein are attached at 
c. 45°, forking once, usually near the base, then arching 
slightly and running parallel to each other, reaching 
the wing or pinna margin at c. 45° to 80°. The density 
of veins along the pinna lobe margin is c. 20 per 10 
mm. 

AMF113563 (Fig.11D) is a 100 mm long 
section of a frond which in life may have reached 200 
mm in length. The lobes, to 35 mm long and 6 to 8 
mm wide, are attached at c. 40° to 50° to the primary 
rachis at a spacing closer than those of the holotype. 


Holotype 
AMF113561, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF113562, AMF112563, Coal Mine Quarry. 


Name derivation 

Gouldiopteris — Gould, for Dr R.E.Gould, 
former palaeobotanist at the University of New 
England, who encouraged my collecting and research 
of the Nymboida Flora; preris, (Lat.), 7erz. 

alethopteroides — Alethopteris, a Palaeozoic 
morpho-genus of fern-like fronds similar in gross 
morphology to the Nymboida material. 


Discussion 

Gouldiopteris, a monotypic genus with G: 
alethopreroides as the type species, is erected for 
foliage fragments that conform with the diagnosis of 
the Palaeozoic form genus A4/erhopreris Sternberg 
(Wagner 1968). A/erhopreris is not a natural genus and 
includes both ferns and pteridosperms. Plants placed 
in the genus are common and widespread in the 
Carboniferous of the Northern Hemisphere, the Permo- 
Carboniferous of China and the Permian of Thailand. 
Boureau (1975) listed 96 species, varieties and 
synonyms. Because of the vast time and geographical 
differences between Northern Hemisphere plants that 
have been placed in A/erhopreris and the Nymboida 
material, it is highly improbable that there is any close 
relationship. I have erected the new morpho-genus 
Gouldiopteris to avoid suggestions that A/erhopreris 
had Mesozoic representatives and a cosmopolitan 
distribution. 

Most early records of A/eopreris species in 
Australia have subsequently been revised or were 
based on unidentifiable material. 4/e*hopreris australis 
(Morris) Johnston (1888), Feistmantel (1890) and 


Proc. Linn. Soc. N.S.W., 124, 2003 


Alethopteris serratifolia Johnston (1887) from 
Tasmania in which the pinnules are separated to the 
base, were placed in Cladophlebis australis (Morris) 
by Walkom (1926). Specimens from the Upper Triassic 
of Queensland, originally placed in A/erthopreris 
lindleyana (Etheridge Jnr 1892) and ?A/erhopteris 
lindleyana (Shirley 1898), were removed by Walkom 
(1917) to Cladophlebis royatlei, a species which was 
described from the Permian of India (Arber 1905). All 
the above specimens previously placed in 4/erhopreris 
are bipinnate and do not conform with the diagnoses 
of Alethopteris or Gouldiopteris. 

Dejerseyia lobata (Jones and de Jersey) Herbst 
1977 has simple to pinnatifid leaves with elongate 
lobes as in Gouldiopteris alethopteroides (see also 
Dejerseyia lobata ‘forma D’ of Anderson and 
Anderson 1983), but differs from G. alethopteroides 
by the lateral venation of well spaced decurrent 
secondary veins which arch and divide several times. 
Dejersevia lJobata is now considered to be a 
gymnosperm (Anderson and Anderson, in press). The 
form of Gouwldiopteris alethopteroides is unique in 
Gondwana Triassic floras. 


Leconama Holmes gen. nov. 
Leconama stachyphylla Holmes gen. et sp. nov. 
Figures 12A-C 


1975 Cladophlebis lobifolia non (Phillips) Seward, 
Flint and Gould, pl.1, Fig. 6 only 

1977 Lobifolia dejerseyi Retallack (in Retallack et 
al.) pp 88-89 


Combined diagnosis 

Medium sized bipinnate frond; pinnae opposite, 
pinnules broad falcate, basiscopic margin sub-circular, 
acroscopic margin straight to slightly concave; midvein 
decurrent, weak, soon dissolving into several radiating 
and forking venules. 


Description 

This taxon is based on single pinnae (Fig. 12B, 
C and Flint and Gould 1975, pl.1, fig 6) and a frond 
fragment showing linear opposite pinnae attached 
acutely to a slender main rachis (Fig. 12A). Pinnae 
opposite, linear to 100 mm long; pinnules opposite, 
closely spaced to overlapping, free to the base, attached 
at c. 60°, broad falcate, entire, c. 6 mm long and wide; 
basiscopic margin sub-circular convex through c. 90°; 
acroscopic margin decurrent upwards at base then 
straight or slightly concave at an obtuse angle to the 
rachis; apex rounded-acute. A decurrent weak midvein 
arches into each pinnule from near the basiscopic 
margin, giving off two pairs of closely spaced lateral 
veins then dissolving into forking and radiating 


61 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


venules. First one or two basiscopic lateral veins 
forking twice to three times; first acroscopic lateral 
vein forking three times with the near venule running 
parallel to the pinna rachis; following lateral veins once 
forked or simple, radiating, c. 24 to 28 vein endings 
around margin. 


Holotype 
AMF121183, Australiam Museum, Sydney. 
(Formerly UNEF14124). 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF121160 formerly UNEF 
AMF 113544 and its counterpart. 


14104, 


Name derivation 

Leconama — lekos, (Gt.), basin, nama, (Gt.), 
stream, referring to the type locality in the Basin Creek 
Formation. 

stachyophylla — stachys, (Gt.), ear of grain, 
Dhylton, (Gr.), leaf, referring to the leaf outline 
resembling an ear of wheat. 


Discussion 

Leconama is a monotypic genus with Z. 
stachyophylla as the type species. A pinna fragment 
of Z. stachyophyla, previously illustrated by Flint and 
Gould 1975, pl. 1, fig. 6, and referred to Cladoph/ebis 
lobifolia, was included by Retallack (in Retallack et 
al. 1977) in Lobifolia dejerseyi, now Nymbopteron 
dejerseyi Holmes (see below). Leconama 
stachyophylla differs from Mymbopteron species by 
the absence of the conjoining of the first acroscopic 
pinnule to the main rachis. The pinnule shape and 
venation pattern distinguishes Z. stachyophylla from 
all other described Gondwana Triassic fronds. 


Micronymbopreris Holmes gen. nov. 
Micronymbopteris repens Holmes gen. et sp. nov. 
Figures 13, 14A-D 


Combined diagnosis 

A small procumbent or climbing plant with 
elliptic to lanceolate bipinnate fronds to 70 mm long, 
irregularly spaced along a curved tapering rhizotomous 
stem. Pinnae alternate, linear, bearing oblong, thick- 
textured pecopteroid pinnules. 


Description 


The holotype (Figs 13A, 14A) shows portions 
of five elliptic bipinnate fronds attached spirally and 


62 


irregularly to an elongated tapering stem or rhizome. 
At the broken base the width is 5 mm and tapers 
through a length of 85 mm to 2 mm wide. Another 
specimen (Fig. 14B) was retrieved from the same 
fractured block. I believe that it is the lower portion of 
the holotype but the section of the stem that would 
join the two is missing. The lower stem fragment is 12 
mm wide with one frond attached at right angles then 
decurving vertically. The holotype shows an almost 
complete frond and another four acutely attached 
incomplete fronds as well as detached fragments 
nearby. The complete frond is acutely decurrent, 
bipinnate, elliptic to 70 mm long, c. 35 mm wide; main 
rachis robust, tapering from 25 mm wide at base; 
pinnae c. 4 mm apart, with thick rachises, opposite, 
becoming subopposite to alternate apically, to 30 mm 
long and 3 mm wide, mostly linear but tapering close 
to the base and towards apex; pinnules alternate, 
closely spaced, free to the base, attached at c. 80° to 
60°, oblong with rounded apex, c. 1—1.5 mm long, c. 
0.5—0.75 mm wide, length to width ratio of c. 2:1; 
venation obscured by the apparent very thick texture 
of the pinnule lamina. 


Holotype 
AMF 120962, isotype AMF 120963, Australian 
Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF120964—120966. 


Name derivation 

Micronymbopteris — contrived, for sma// 
Nymboida fern. 

repens — (Lat.), creeping, for the inferred 
growth form. 


Discussion 

Micronymbopteris is a monotypic genus with 
the type species 1Z repens. Most ferns and fern-like 
taxa in the fossil record are known only from detached 
fronds, many of which are incomplete or even 
fragmentary. The holotype of AZ repens is a rare and 
important find as it demonstrates the elongated stem 
and probable creeping or climbing growth form of the 
once living plant. 

M. repens is unlike any other described fern- 
like material and its fronds are perhaps the smallest in 
the Gondwana Triassic fossil record. 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Nymbiella Holmes gen. nov. 
Nymbiella lacerata Holmes gen. et sp. nov. 
Figures 15A, 16A, B 


Combined diagnosis 

Medium to large bipinnate-bipinnatifid frond; 
pinnae opposite to subopposite, broad linear; pinnules 
irregular in width and shape, separated to base or 
conjoined; apices obtuse, lobed or irregularly lacerated. 
First basiscopic pinnules sometimes attached to the 
main rachis. Venation odontopteroid, from one to three 
or more veins arching from pinna rachis into each 
pinnule, dividing once, occasionally twice and running 
parallel to each other to the distal margin. 


Description 

Medium to large bipinnate-bipinnatifid fronds 
estimated to reach 600 mm in length and 220 mm in 
width. No fronds are complete and no bases are present. 
The holotype (Fig. 15A) is an apical portion of a frond 
220 mm long with another frond adjacent and slightly 
diverging. There is no evidence that the two fragments 
may be pinnae of a tripinnate fern. The main rachis in 
the base of this specimen is 3 mm wide and tapers 
gradually apically. In the length preserved there are 
13 pairs of opposite to subopposite pinnae attached at 
12-15 mm apart. The pinnae have a decurrent base 
then continue straight for their whole length at an angle 
of 60° to 75° to the main rachis. Pinnae broad-linear to 
120 mm long and 20 mm wide, tapering in distal 
quarter to an acute apex. Pinnules of irregular shape 
and width, 8-10 mm long, attached from 60° to 80°, 
usually parallel-sided, sometimes contracted at the base 
(Fig.16A), sometimes several pinnules conjoined (Fig. 
16B), apices obtuse, flattened or variously lobed or 
lacerated. Basal basiscopic pinnules attached around 
the junction of the pinna rachis with the main rachis 
and sometimes directly to the main rachis. The pinnule 
lamina is apparently thin with the venation clearly 
defined. Venation odontopteroid; one to three or more 
veins arching into each pinnule, forking once or 
occasionally twice and running straight and parallel 
to each other to the distal margin. 


Holotype 
AMF 113530, Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 

AMF113531, 113536, 121174 from Reserve 
Quarry. AMF113532, 113535, 113537, 121173 from 
Coal Mine Quarry. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Name derivation 

Nymbiella — contrived from Vymbo/da, the 
source of the material. 

lacerata — (Lat.) lacerated, irregularly torn; 
referring to the apical margin of many pinnules. 


Discussion 

Nymbiella is a monotypic genus with V. 
lacerataas the type species. Over forty specimens are 
in the collections, all are incomplete and none show 
the basal region. The extreme variability of pinnule 
shape and form of V. /acera/ais unique in Gondwana 
fern-like fronds. The two frond fragments on the 
holotype slab appear to be arising from a common base, 
and suggest a growth form perhaps similar to that of 
Osmundopsis scalaris (Holmes 2001b). 

Odontopteris? (Callipteris) laceratifolia ftom 

the Upper Palaeozoic of China, illustrated by Halle 
(1927 pl. 32, figs 1, 2) has bipinnatifid leaves with 
lacerated margins and odontopteroid venation. 
However Halle’s material differs from Vymbiella 
/acerataby the less divided pinnae and by the veins at 
a more acute angle. Because of the geographic and 
time differences I believe that V /acerafa should be 
regarded as generically distinct. 


Nymboidiantum Holmes gen. nov. 
Type species Sphenopteris (? )glossophylla Tenison- 
Woods 1883 p. 58, pl. 4, fig. 4 


Diagnosis 

Fern-like foliage; fronds bipinnate or rarely 
pinnate; pinnules attached at c. 45°, contracted at the 
base, sessile, lamina elliptic, margin entire, lobed or 
divided into segments. Three veins enter pinnule base, 
each forking one or more times as they pass into the 
lobes or distal portion of the entire lamina. 


Discussion 

From the gross morphology of pinnules with 
contracted bases and radiating venation, 
Nymboidiantum resembles the sterile fronds of some 
extant species of the fern genus Adiantum but no close 
relationship is inferred. Fossil leaves of Palaeozoic age 
with somewhat similar morphology have been placed 
variously in the genera 77~phy/opreris (Boureau 1975; 
Morris 1975), Adiantites (Boureau 1975), Genselia 
(Knaus and Gillespie 2001), Pa/maropreris (Boureau 
1975) and Archaeopreris (Feistmantel 1890; Boureau 
1970). Others of Late Jurassic to Tertiary age from 
the Northern Hemisphere have been placed in 
Adiantopteris (Boureau 1975). Because of the vast 
differences in time and distance, the new genus 
Nymboidiantum is erected to separate the Nymboida 


63 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


material from the genera listed above. Undescribed 
Nymboidiantum foliage from the Carnian Molteno 
Formation of South Africa has been figured by 
Anderson and Anderson (1983, pl. 9; Figs la, 1b) as 
‘Incertae sedis gen. A, sp. A’. All the specimens from 
Nymboida here placed in VWymboidiantum are known 
only from sterile fronds, so their natural affinities are 
not known. With the exception of Vymboidiantum 
glossophyllum which is known from c. 40 specimens, 
the other taxa are very rare. I have distinguished five 
species mainly on the size, form and placement of the 
pinnules, and there appear to be no intergrading forms. 
The different fossil species are each preserved in 
sediments representing specific facies and it may be 
reasonable to assume that in life they grew in different 
habitats. 


Nymboidiantum glossophyllum (Tenison-Woods 
1883) Holmes gen. et comb. nov. 
Figures 17A-E, 18C, D 


1883 Sphenopteris (7) glossophyla Tennison- 
Woods p.58, pl.4, Fig.4 

1983 Incertae sedis foliage gen. A, sp. A Anderson 
and Anderson, pl.9, Figs la and 1b. 


Diagnosis emended 

Medium sized bipinnate frond; pinnules well- 
spaced, alternate, elliptic with contracted decurrent 
base; proximal pinnules sometimes tri-lobed to deeply 
incised; venation sparse, forking and radiating to apical 
margin. 


Description 

The type specimen of Tenison-Woods (1883) 
is a small fragment showing three pairs of incomplete 
pinnae attached alternately to a main rachis that is 
mostly missing in the coarse siltstone matrix (refigured 
here, Fig. 17A). The elliptic pinnules are closely similar 
to those on many specimens in the Nymboida 
collections. However the Nymboida material exhibits 
a range of variation amongst specimens that I include 
in this species, especially in the lobing of the proximal 
pinnules. AMF120946 is a siltstone slab with some 
almost complete ovate fronds to c. 250 mm long (Fig. 
17C), with alternate pinnae bearing to eight pairs of 
sub-opposite to alternate elliptic pinnules. The pinnules 
are from 5-10 mm long and 3-5 mm wide, attached at 
c. 45°-60° to the pinna rachis. Basally the shorter 
pinnae are at right angles to the main rachis becoming 
acute apically. Proximal pinnules are sometimes 
partially divided into two or three lobes (Figs 17B, D, 
E). On some specimens (Figs 18C, D) the pinnules are 
divided to the base into three linear lobes. The texture 
of the pinnule laminae appears to be thick and the 


64 


venation is rarely observed, except in some rare types 
of preservation where sparse radiating veins may be 
faintly visible. 


Lectotype 

AMF68449, Australian Museum, Sydney. 
Formerly catalogued as SUF35 in the fossil collections 
of Sydney University. 


Type locality 

“Talbragar Mines” of Tenison-Woods (1883). 
Probably north-east of the present day village of 
Ballimore on the Talbragar River east of Dubbo, in 
the Napperby Formation, Middle Triassic. 


Other material 
AMF121158—-121149, 121152, Coal Mine 
Quarry, AMF121150, 121153, Reserve Quarry. 


Discussion 

I have examined the specimen of Sphenopreris 
(7) glossophyla Tenison-Woods which is now housed 
in the Australian Museum, Sydney. It is a small 
fragment of a pinna rachis with ?alternate ovate 
pinnules with contracted bases and no visible venation. 
It agrees well with much of the Nymboida material, 
which, however, demonstrates a wide range of 
variation. The emended diagnosis reflects the diversity 
present in the Nymbcida specimens which I now 
include in this taxon. The type locality, ‘Talbragar 
Mines’ near Ballimore on the Talbragar River, is ina 
very poorly collected area of the Middle Triassic 
Napperby Formation (previously included in the 
Wallangarra Formation, Cameron et al. 1999), which 
outcrops along the south-eastern margin of the Great 
Artesian Basin in central-western New South Wales. 
An assemblage of fossil plants from the same formation 
at Benolong, south west of Dubbo, has been described 
by Holmes (1982, 2001a). 

In gross morphology V. g/ossophyllum is 
closely similar to some forms of 777p/y//opreris known 
from the Carboniferous of Peru and the Northern 
Hemisphere (Boureau 1975, Figs 557, 558), but these 
forms are regarded as distinct due to the geographic 
and time differences. 

Comparisons between MWymboidiantum 
glossophyllum and those species described below are 
given under the respective species. 


Nymboidiantum multilobatum Holmes gen. et sp. 
nov. 
Figures 18A, B 
Diagnosis 
Medium sized bipinnate frond, pinnae sub- 
opposite; pinnules broad-flabellate, divided into 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


several more or less deeply incised lobes. 


Description 

Frond bipinnate, estimated length to c. 250 mm; 
main rachis 4 mm wide near base, tapering gradually, 
apical portion of frond not preserved. Pinnae sub- 
opposite, 30 mm wide, length not known. Pinnules 
alternate attached by contracted base at c. 45° to pinna 
rachis, rhombic to sub-circular, 15 mm long and wide 
near main rachis, decreasing in size distally. Pinnules 
deeply divided into three to five segments, each 
segment with two or three elongated, obtuse lobes. A 
single vein enters each pinnule, soon forking into each 
segment and again into each lobe. 


Holotype 
AMF113558; isotypes AMF113559, 113560, 
Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
F ormation, Nymboida Coal Measures, Middle Triassic. 


Name derivation 
multi, (Lat.) many; lobata, (Lat.) lobed, 
referring to the much dissected pinnules 


Discussion 

This morphotaxon is known only from a single 
incomplete specimen that is associated with large 
Sphenobaiera \eaves. It differs from all other species 
of Mymboidiantum by the larger flabellate pinnules 
which are divided into several segments, each segment 
with apical lobes. 


Nymboidiantum elegans Holmes gen. et sp. nov. 
Figures 19A, B 


Diagnosis 

Pinnae alternate, linear; pinnules well-spaced, 
opposite, elliptic, not lobed. First basiscopic pinnule 
attached in angle between the main and pinna rachis 
or directly on the main rachis. 


Description 

The holotype (Fig.19A) is an apical fragment, 
130 mm long, of a frond which in life was probably 
twice that length. Width of the main rachis at the base 
of the portion preserved is 2.5 mm, tapering gradually 
to the apex. Main rachis longitudinally striated but not 
conspicuously ribbed or grooved. Pinnae alternate, 
decurrently attached at c. 45° to the main rachis, linear, 
15 mm wide, to 65 mm long, tapering in the distal 
portion to an acute apex. Pinnules opposite, evenly 


Proc. Linn. Soc. N.S.W., 124, 2003 


spaced, symmetrically elliptical, entire, apex acute, c. 
8 mm long and 3 mm wide, well separated, decreasing 
in size distally and apically, 12 pinnules in a pinna 
length of 60 mm. The first basiscopic pinnule on each 
pinna is attached at the base of the pinna rachis or 
directly to the main rachis. Towards the frond apex 
the pinnules coalesce to form lobed or entire pinnae. 
Venation not preserved. 


Holotype 
AMF 113504; isotype AMF113505. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF 113506, Coal Mine Quarry. 


Name derivation 
elegans — (Lat.), e/egant, referring to the neat 
and even spacing of the pinnules. 


Discussion 

N. elegans differs from V. glossophyllum, N. 
robustum and NV. multilobatumby the opposite, elliptic, 
well-spaced pinnules and by the absence of lobing of 
the proximal pinnules. 


Nymboidiantum fractiflexum Holmes gen. et sp. nov. 
Figures 19C, D 


Diagnosis 

Planate, pinnate frond; rachis flattened, 
grooved, changing direction slightly at each pair of 
opposite pinnae. Pinnae sessile, rhombic, venation fine, 
forking and radiating from a short midvein. 


Description 

This taxon is based on a single specimen which 
consists of five pairs of opposite pinnae on a slightly 
zigzag rachis in which the rachis and pinnae appear to 
be flattened in one plane (Fig. 19C). The rachis as 
preserved is 50 mm long, with a conspicuous medial 
groove and longitudinal striations, 2 mm in width near 
base and not visibly decreasing to the tip, where it 
forms a short broken projection beyond the last pair 
of pinnae. Pinnae well-spaced c. 12 mm apart, 
decurrent on rachis, sessile, attached at c. 45°, c. 35 
mm long and 10 mm wide, obtrullate to rhombic, entire 
with acute apex. Fine lateral veins diverge from a 
midvein or groove in the proximal one third of the 
pinna and fork several times distally. In the proximal 
half of the pinna, the radiating veins run parallel to the 


65 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


margin. In the distal half, the veins terminate along 
the margin with a density of c. 12 per 10 mm (Fig. 
19D). 


Holotype 
AMF 113502, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Name derivation 
Jractiflexus — (Lat.) — deviating from side to 
side, zigzag, as in the rachis of this taxon. 


Discussion 

By its pinnule shape and venation this leaf 
fragment is placed in the morpho-genus 
Nymboidiantum, but by the flattened zigzag rachis and 
pinnate form it differs from all other described species. 
The type specimen of V. /racizflexus may be a pinna 
of a large bipinnate frond as in other Vymboidiantum 
species. 


Nymboidiantum robustum Holmes gen. et sp. nov. 
Figures 20A-C 


Diagnosis 

Pinnules broad, ovate, opposite, well-separated 
proximally, coalescing distally; first acroscopic pinnule 
broadly but shallowly lobed. 


Description 

Medium to large bipinnate frond; midportions 
only of fronds preserved. Complete fronds estimated 
to exceed 300 mm long. The main rachis of the 
holotype (Fig. 20B) decreases in width from 2.5 mm 
to 2 mm over a length of 125 mm. Pinnae opposite to 
subopposite, broad-linear, 15 mm wide, 40-50 mm 
long. Pinnules opposite, attached at c. 45°, base slightly 
contracted, strongly decurrent, well separated 
proximally, coalescing distally, 4-8 mm wide, 5-12 
mim long, margins entire, apex obtuse. First acroscopic 
pinnule shallowly lobed. The venation is not clear but 
three veins appear to enter base with each vein forking 
once. 


Holotype 
AMF 113496, Australian Museum, Sydney. 


Type locality 


Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


66 


Other material 
AMF113495, 113497, 121164, 121166, all 
Reserve Quarry. 


Name derivation 
robusta — (Lat.), robust, referring to the broad 
compact pinnules. 


Discussion 

NV. robustum differs from all other 
Nymboidiantum species by the larger pinnules with 
broadly decurrent bases. 


Nymbophlebis Holmes gen. nov. 


Nymbophlebis polymorpha Holmes gen. et sp. nov. 
Figures 21A, 22A-C, 23A-C 


Combined diagnosis 

Large polymorphic frond with long opposite 
linear, slightly arching primary pinnae distally bearing 
cladophieboid pinnules; in basal and proximal portions 
of fronds, primary pinnae tripinnate with fine pinnules; 
distally pinnae becoming bipinnate to pinnate as 
ultimate segments coalesce and conjoin to form large 
linear entire pinnules. 


Description 

This taxon is based on large bi-tri-quadripinnate 
sterile fronds in which the polymorphic characters may 
occur on a single primary rachis. Figure 20 is a basal 
portion of a frond estimated to have reached | m long. 
Secondary rachises opposite, slightly arching or 
straight, to 150 mm long, 20-30 mm wide with 20 to 
30 alternate pinnae-pinnules; proximal portions of the 
lower rachises often tripinnate to bipinnate but pinnules 
distally conjoining to progressively coalesce until the 
primary rachis is simply pinnate (Figs 22. A, B; Figs 
23A-C). The cladophleboid distal and apical pinnules 
are attached by a wide base, slightly contracted 
basiscopically and enlarged or decurrent upwards on 
the acroscopic side; broad-linear or slightly tapering, 
margin entire or slightly lobed close to the tripinnate 
segments of the frond, 12—18 mm long, 3-5 mm wide, 
with length to width ratio of c. 3.5—4:1; midvein 
decurrent, strong, persisting almost to pinnule apex; 
8-10 pairs of lateral veins; first basiscopic lateral vein 
attached in angle between the pinna rachis and midvein, 
forking twice with proximal venule forking again; 
other lateral veins decurrent, all twice forked; first 
dichotomy close to the midvein then again at 1/3 the 
distance to the margin; venules travelling straight, close 
and parallel to meet margin at c. 45° to 70°. 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Holotype 
AMF120995, Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Tmiassic. 


Other material 
AMF120988, 120996-121007, 121009- 
121013, Reserve Quarry; AMF121008, Coal Mine 


Quarry. 


Name derivation 

Nymbophlebis contrived, for Cladophlebis-like 
plant from Nymboida. 

Polymorpha — (Gt.) many forms, referring to 
the diversity of pinnule shapes within a single frond. 


Discussion 

Nymbophiebis is a monotypic genus with WV. 

polymorpha as the type species. Almost complete 

fronds of Vymbophlebis polymorpha occur at the 
Reserve Quarry in a bed of olive-grey mudstone. 
Reliable identification can only be achieved from 
substantially complete fronds. Retallack (1977) listed 
frond fragments from the Coal Mine Quarry at 
Nymboida which agreed well with the the bipinnate 
portions of Vymbophlebis polymorphaas *C: australis 
sensu stricta Morris with twice forked lateral veins’. 

The interpretation of Pecopreris australis 
Morris (1845) [= Cladophlebis australis (Motris) 
Walkom (1917)] from the Triassic Newtown Beds of 
Tasmania has had a long and confused history. The 
pinnules in the bipinnate portions of Vymbophlebis 
Dolymorpha are similar in shape to those of Morris’s 
fronds and the lateral veins are twice forking. However, 
NV. polymorpha differs in that the primary and 
secondary forking of the lateral veins occurs close to 
the midvein and the veins then proceed close and 
parallel to each other to the margin, in contrast to those 
of Morris’s illustration which fork midway to the 
margin and then widely diverge. 

The polymorphic nature of the complete fronds 
of Nymbophlebis polymorpha argues for their 
placement in a genus separate to C/adophi/ebis. Isolated 
bipinnate fragments may be best placed in 
Cladophlebis sp. indet. 

The diverse form of the pinnules of WV. 
Dolymorpha is reflected in some forms of the extant 
genera Preris and Preridium, but no relationship is 
inferred. 


Nymbopteron Holmes gen. nov. 


Type species Lobijolia dejerseyi Retallack (in 
Retallack et al. 1977) 


Proc. Linn. Soc. N.S.W., 124, 2003 


Diagnosis 

Small to large bipinnate-bipinnatifid fronds, the 
first acroscopic pinnule of each pinna always confluent 
between the pinna and main rachises to form a 
triangular wing. First basiscopic pinnule sometimes 
enlarged, triangular, rectangular, rounded or variously 
lobed, often attached between pinna and main rachis 
or directly to the main rachis; subsequent 
cladophleboid pinnules of even size and shape for most 
of the pinna length but decreasing in size and 
conjoining distally and apically. 


Name derivation 

Nymbopteron —nymbo, referring to Nymboida, 
the source of the material; preror, (Gr.), wing; referring 
to the winged shape of the first acroscopic pinnules. 


Discussion 

The genus Vymbopreron is erected to include 
Australian material formerly placed in Cladophlebis 
/obifolia (Flint and Gould 1975), Lobijolia dejerseyi 
Retallack (in Retallack et al. 1977) and possibly 
Cladophlebis lobifolia (Walkom 1924, 1928). 
Cladophlebis lobifolia sensu stricta is a Northern 
Hemisphere species, having been recorded from the 
Middle Jurassic to Early Cretaceous of Europe, Russia 
and China. Fertile specimens are placed in Z4oracea 
lobifolia Thomas (Harris 1961). Zod7o/ra, a genus with 
a new type, Lobifolia novopokrovskii was erected by 
Lebedev and Rasskazova (1968) who also included 
the combination Lobijolia Jobifolia, contrary to ICBN 
tules (as discussed by Rigby 1977). Retallack (in 
Retallack et al. 1977) described the new species 
Lobifolia dejerseyi and illustrated a small fragment 
from the Cloughers Creek Formation (Retallack et al. 
1977, Fig.5A). He selected as the holotype a specimen 
from the Basin Creek Formation that had been assigned 
to Cladophlebis lobifolia by Flint and Gould (1975 
Pl.1, fig.4). That specimen is reillustrated here as Fig. 
25A. Apart from the geographical and time differences, 
the Nymboida material described by Retallack and the 
additional new species described below differ 
significantly from both the Northern Hemisphere 
Lobifolia species and Cladoph/ebis sensu Frenguelli 
(1947) by the placement and form of the first basiscopic 
and acroscopic pinnules. It is essential when 
identifying fossils as Vymbopteron that the material 
includes the portions of the pinnae attached to the main 
rachis. Isolated distal pinna fragments may be confused 
with Cladophlebis or Dicroidium species. 

Four species of Vymbopreron are described 
below. They are distinguished by the frond size, shape 
and venation of the pinnules and particularly by the 
shape and position of the first basiscopic pinnules. 


67 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


While the collected material of some of these species 
is limited, each species occurs in a sediment type 
representing a different facies similarly to that noted 
for Vymboidiantum spp. above, and it is most probable 
that each species grew in a different vegetation type. 
In the present collection there are no intergrading forms 
between the species. 


Nymbopteron dejerseyi (Retallack, in Retallack et al. 
1977) Holmes gen. et comb. nov. 
Figures 24A—E and 25A 


1975 Cladophlebis lobifolia non (Phillips) Seward, 
Flint and Gould, pl.1, Fig.4 only 

1977 Lobifolia dejerseyi Retallack in Retallack et 
al., p.88 


Emended diagnosis 

Medium sized ovate fronds; first basiscopic 
pinnule enlarged and of irregular shape, attached in 
angle between pinna rachis and main rachis or directly 
on main rachis; venation radiating and forking; 
following pinnules with midvein persisting almost to 
apex, first basiscopic vein attached at base of midvein 
or directly to pinna rachis, lateral veins twice or once 
broadly forked. 


Description 

Retallack’s holotype, reillustrated here as Fig. 
25A, is a midportion of a broad ovate frond estimated 
to have been up to c. 200 mm long. No complete fronds 
are available. The main rachis is deeply grooved and 
ridged (Figs 24A-D), 2—3 mm wide near base. Pinnae 
well separated, sub-opposite to alternate, basal pinnae 
short and obtuse with conjoined pinnules (Fig. 24B); 
in mid-frond, pinnae attached at a right angles and 
pinnules separated to the base, apically the pinnae 
becoming moderately acute and pinnules again 
coalescing (Fig. 24C). First basiscopic pinnule 
enlarged and of irregular shape, attached in angle 
between pinna and main rachis or directly to main 
rachis (Figs 24D, E) in the manner of zwischenfiedern 
as in Lepidopreris species (Anderson and Anderson 
1989); with strong midvein at centre base of pinnule, 
forking into three major lateral veins with each vein 
again forking once to three times, 8 to 28 vein endings 
around the margin. The venation in the triangular first 
acroscopic pinnule bifurcates three times, distal veins 
almost parallel. The succeding pinnules opposite to 
alternate, free to the base but closely spaced, ovate to 
broad-falcate, 4-10 mm long and 24 mm wide, length 
to width ratio of c. 1.6—2:1; midvein slightly sinuous, 
persistent almost to apex, three to four pairs of alternate 
lateral veins broadly forking once or twice, first lateral 
vein sometimes arising directly from the pinna rachis. 


68 


Holotype 
AMF121158 (formerly UNEF14102), 
Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 

AMF113543-113557, 120972, 120974, 
120975, Coal Mine Quarry; AMF120973, 120977, 
Reserve Quarry. 


Discussion 

The holotype of this species was placed by 
Retallack (in Retallack et al. 1977) in the genus 
Lobifolia, which is essentially a Northern Hemisphere 
genus of Jurassic to Cretaceous age (Lebedev and 
Rasskazova 1968; Boureau 1975) as discussed above. 
The conjoining of the first acroscopic pinnules to form 
a decurrent triangular membrane between the base of 
the pinna rachis and the main rachis does not occur in 
Lobifolia but is diagnostic for Mywbopreron. 

N. dejerseyi differs from the other Vymzbopreron 
species by the broadly enlarged basal basiscopic 
pinnules with radiating venation and by the persistent 
midvein with broadly forking lateral veins in the 
following pinnules. 


Nymbopteron foleyi Holmes gen. et sp. nov. 
Figure 25B, C; 26A 


Diagnosis 

Small to medium sized fronds, first basiscopic 
pinnules rounded, attached along base of pinna rachis; 
following pinnules triangular-falcate; midvein weak, 
forking four or five times into fine radiating venules. 


Description 

Several almost complete broad elliptic to 
ovate fronds on AMF113538 (Fig. 26A) are up to 150 
mm long and 65 mm wide; main rachis 2 mm wide 
near base. Pinnae opposite to alternate, basal pinnae 
short, with decurved attachment, at high angles in mid 
frond and becoming more acute apically. First 
basiscopic pinnules attached near the base of the 
pinnae, semicircular; first acroscopic pinnule enlarged, 
triangular and conjoined in the angle between the main 
rachis and the pinna rachis and reaching almost to the 
next pinna rachis; following pinnules opposite, closely 
spaced to overlapping (Figs 25B, C), short, broad, 
rhombic to slightly falcate, apex obtuse; length to width 
ratio of c. 1.5:1. Venation usually obscure; midvein 
arising close to the basiscopic margin at c. 45°, then 
dividing four to five times into venules which radiate 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


evenly at an acute angle to each other to the pinnule 
margin; c. 16-22 vein endings around margin. 


Holotype 
AMF113538, part and counterpart, Australian 
Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF113522, 113539, 113540, 120972, 
120985, all from Coal Mine Quarry. 


Name derivation 

foleyi — for Mr Brian Foley, Nymboida Quarries 
operator, in recognition of his valuable on-site 
assistance over a period exceeding 30 years. 


Discussion 

N. foleyi differs from the other Nymbopteron 
species by the first basiscopic pinnules which are 
mostly rounded and attached only to the pinna rachis, 
and by details of the venation pattern in the pinnules. 


Nymbopteron rhomboidale Holmes gen. et sp. nov. 
Figures 27A-D 


Diagnosis 

Medium sized bipinnate frond, pinnae 
subopposite, closely spaced, first basiscopic pinnules 
large, square to rounded, attached mostly to the main 
rachis from which a single vein enters the pinnule, 
forking four times, ultimate veinlets meeting margin 
at 90° to main rachis; first acroscopic pinnules forming 
equilateral triangles between the upper side of the pinna 
rachis and main rachis, proceeding pinnules 
thomboidal, overlapping; midvein short, with several 
radiating and forking lateral veins. 


Description 

The only specimen (Fig. 27A) is an upper mid 
portion of a frond with the base and apex missing; 
estimated length of c. 150 mm; main rachis at broken 
base 2.5 mm wide, tapering apically to 1.5 mm wide, 
longitudinally grooved, with six pairs of subopposite 
broad linear pinnae attached at c. 45° and c. 11 mm 
apart. First basiscopic pinnules (Fig. 27B) c. 5 mm 
wide and 5 mm long, attached to basal 3 mm of pinna 
rachis and 5 mm along the main rachis, distal margin 
square to rounded, a single vein enters the base of the 
pinnule from the main rachis, forking four times, the 
ultimate venules are parallel and pass into the pinnule 
margin at 90° to the main rachis. First acroscopic 


Proc. Linn. Soc. N.S.W., 124, 2003 


pinnules equilateral triangular in shape with proximal 
margin conjoined to the main rachis; a single vein 
enters the pinnule from the junction of the pinna rachis 
with the main rachis, forking three times, with ultimate 
veinlets parallel to each other and at right angles to 
the sloping pinnule margin. Succeeding pinnules along 
the pinnae (Fig. 27D) alternate, overlapping, c. 5 mm 
wide and 5 mm long hence a length to width ratio of 
1:1, rhomboidal, proximal margin curving through 90°, 
distal margin at right angles to pinna rachis, straight 
to slightly convex, apex acute, a single vein enters the 
pinnule at a very acute angle, forking four to six times, 
with adaxial branches running parallel to the pinna 
rachis and the other veins radiating evenly to the 
pinnule margin where there are 16—22 vein endings, 
distal pinnules decreasing in size and becoming 
rounded. 


Holotype 
AMF113556, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Name derivation 

rhomboidale — rhombus, (Lat.), an equilateral 
parallelogram with unequal pairs of opposite angles; 
referring to the shape of the pinnules. 


Discussion 

N. rhomboidale differs from all other 
Nymbopteron species by the closely spaced pinnae, 
by the shorter broader overlapping pinnules and by 
the distinctive radiating venation. The lateral venation 
in the proximal portions of the broad overlapping 
pinnules may be confused as conjoining with the veins 
of adjacent pinnules as described for Merianopteris 
major Feistmantel (Boureau 1975). 


Nymbopteron uncinatum Holmes gen. et sp. nov. 
Figure 28A 


Diagnosis 
A large bipinnate frond with broad uncinate 
basal basiscopic pinnules. 


Description 

N. uncinatum is based on two incomplete 
specimens of large bipinnate fronds, both over 200 
mm long and with bases and apices missing. On the 
portion preserved of the holotype (Fig. 28A) the main 
rachis is smooth, 4 mm wide at the broken base, 
tapering gradually upwards, with 15 pairs of opposite, 
straight pinnae to 80 mm long attached at c. 45°. The 


69 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


specimen AMF120969 has 10 pairs of opposite pinnae 
to 60 mm long attached at c. 60° to 80°. First basiscopic 
pinnules attached in the angle between the main and 
pinna rachises, triangular to 10 mm long, with 
recurving tip which forms a broad hook-like outline; 
first acroscopic pinnule vertically elongate and 
conjoined to the main rachis. Succeeding pinnules 
opposite to subopposite, separated to the base except 
at the distal extremity of the pinnae where they 
coalesce, broadly triangular to slightly falcate with 
proximal margin broadly convex, the distal margin 
straight or slightly convex, c. 8 mm long and c. 4 mm 
wide with a length to width ratio of 2:1. Venation not 
preserved. 


Holotype 
AMF120968, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF120969, Coal Mine Quarry. 


Name derivation 
uncinatum — (Lat.), hooked, referring to the 
hook-shaped outline of the first basiscopic pinnules. 


Discussion 

N. uncinatum differs from all other 
Nymbopteron species by its larger and more robust 
frond size and by the hook-shaped outline of the first 
basiscopic pinnules. The main and pinna rachises of 
N. uncinatum appear to be rounded and smooth in 
contrast to the ribbed and grooved rachises of N. 
dejerseyi and N. foleyi. This feature may be real or 
perhaps it is an artifact dependent upon the orientation 
or deterioration of the frond in the sediment at the time 
of fossilisation (see comments on Osmundopsis 
scalaris in Holmes 2001b). 


Nymborhipteris Holmes gen. nov. 


Nymborhipteris radiata Holmes gen. et sp. nov. 
Figures 29A, B 


Combined diagnosis 

Medium sized bipinnate frond, stout main 
rachis, pinnae alternate, bearing opposite sub-circular 
pinnules with radiating and forking venation. 


Description 


A fragment of a mid-portion of a bipinnate 
frond; main rachis on portion preserved longitudinally 


70 


wrinkled, 7 mm wide, tapering to 6 mm over length of 
90 mm; total length of frond estimated to reach 300 
mm. Pinnae sub-opposite to alternate, 1 1-15 mm apart, 
attached at c. 60°, straight, length unknown. Pinnules 
opposite, sub-circular, base broadly attached but 
slightly constricted basiscopically, margin entire, apex 
rounded; a single decurrent vein enters the base, soon 
dividing four times into finer veins which again fork 
two or three times to radiate throughout the lamina 
with c. 24 vein endings around the margin. 


Holotype 
AMF1 13489, Australian Museum, Sydney. 


Type locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Name derivation 

nymborhipteris — contrived; from Nymboida, 
the source of the material; rhipis, (Gr.) fan; pteris, (Gr.) 
fern; referring to the fan-shaped appearance of the 
pinnules. 

radiata (Lat.), radiating, referring to the 
venation pattern in the pinnules. 


Discussion 

N. radiata, the type species of the monotypic 
genus Nymborhipteris, is based on a single fragmentary 
specimen (Fig. 29A). The stout striated main rachis 
and the venation pattern distinguishes N. radiata from 
the forked bipinnate fronds of the corystosperm 
Dicroidium zuberii (Retallack 1977; Anderson and 
Anderson 1983). In gross morphology this frond 
differs from all known Gondwana material. 


Ptilotonymba Holmes gen. nov. 


Ptilotonymba curvinervia Holmes gen. et sp. nov. 
Figures 30A-C, 31 
1977 Arctopteris sp? Retallack in Retallack et al. p. 
86, Fig. 5C 
1977 Cladophlebis sp. cf C. oblonga, Bourke et al. 
Fig. 3.2 


Combined diagnosis 

Medium to large bipinnate frond; pinnae 
broad-linear; pinnules strongly decurrent, rhombic; 
venation asymmetrical; two lateral veins attached to 
pinna rachis; 4-6 basiscopic lateral veins once forked 
or simple, slightly recurved; 3-4 acroscopic lateral 
veins mainly unforked, arching to follow parallel to 
pinnule margin. 


Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Description 

This morpho-taxon is based on five fragments 
of bipinnate fronds. The holotype (Figs 30B, C) shows 
incomplete pinnae to 80 mm long in two parallel lots 
of four apparently attached to separate main rachises. 
Fig. 31 is a portion of a large frond with the main rachis 
18 mm wide near base, decreasing to 12 mm over the 
300 mm length preserved, which suggests a total length 
in excess of 1 metre. Pinnae opposite, to >80 mm long, 
20-30 mm apart. Pinnules rhomboidal to rectangular, 
alternate overlapping, c. 7-10 mm long and 5-6 mm 
wide, base broad, decurrent on basiscopic side, 
contracted on acroscopic side, length to width ratio of 
proximal pinnules c. 1.6:1, margin entire, apex acute 
to obtuse. Pinnule midvein entering pinnule at c. 30°, 
arching to c. 45°, at 2/3 of way through lamina to fork 
and continue as two parallel veins to the apex. Four 
pairs of asymmetrical lateral veins attached at c. 45°. 
First one or two lateral veins on basiscopic side of the 
pinnule fork twice, subsequent veins forking once or 
unforked near apex, running straight or slightly 
recurved and parallel to each other to the pinnule 
margin. On the acroscopic side of the midvein four 
lateral veins are attached at c. 30°, once forked, the 
proximal one or two recurving strongly to follow 
parallel to the pinnule margin for half its length; 
subsequent veins leaving the midvein at c. 30° and 
running parallel to each other round distal portion of 
pinnule margin. Two veins enter the decurrent portion 
of each pinnule at 45° directly from the main rachis, 
first vein simple, second vein once forked; 14-18 or 
more vein endings around pinnule margin with twice 
as many endings on the basiscopic margin as on the 
acroscopic margin. Distally the pinnules coalesce and 
the pinna becomes lobed to entire. 


Holotype 
AMF113479, Australian Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF11380-83, Coal Mine Quarry. 


Name derivation 
Ptilotonymba — ptilotos, (Gr.) feathered, 
referring to the feather-like appearance of the pinnules; 
nymba, for Nymboida, the source of the material. 
curvinervia — contrived, for the strongly arching 
lateral veins. 


Discussion 
Ptilotonymba is a monotypic genus with P. 


Proc. Linn. Soc. N.S.W., 124, 2003 


curvinervia as the type species. Arctopteris sp? of 
Retallack (in Retallack et al. 1977) and Cladophlebis 
sp. cf C. oblonga (Bourke et al. 1977) are frond 
fragments that agree with P. curvinervia. Arctopteris 
spp from the Lower Cretaceous of Siberia (Samalyna 
1964; Boureau 1975) have similar curving lateral 
venation and coalescing pinnules, but differ by the 
unforked lateral veins and by the presence of pinnules 
decurrent on, or attached directly to, the main rachis 
between the pinnae. Pecopteris arcuata Halle (1927 
pl. 19, Figs 1-7; pl.20, Figs 1-13) from the Palaeozoic 
of China has closely spaced to coalescing pinnules with 
curving venation similar to P. curvinervia, but differs 
by the opposite arrangement of the pinnules and by 
the unforked lateral veins. 


Genus Sphenopteris (Brongniart) Sternberg, 1825 
Type species Sphenopteris elegans (Brongn.) 
Sternberg 1825, see Boureau 1975 pp 427-429 


Sphenopteris speciosa Holmes sp. nov. 
Figures 32A-D 


Diagnosis 

Medium sized bipinnate frond. Pinnae opposite. 
Pinnules elongate triangular with deeply incised lobes 
on proximal pinnae, decreasing in size, number of lobes 
and degree of lobation distally and apically. Midvein 
straight and strong in proximal half of pinnule, 
decreasing in width and forking in the apical lobe. 
Lateral veins arching and forking into each lobe. 


Description 

This leaf form is known only from incomplete 
fronds; the largest being a midportion 150 mm long 
suggesting a total length of c. 300 mm. Pinnae opposite, 
recurving, slightly overlapping close to the main rachis; 
attached at c. 60° to 70°, to c. 100 mm long, 45 mm 
wide near the base, with up to 10 pairs of pinnules. 
Pinnules alternate, base contracted, to 25 mm long x 
15 mm wide across proximal portion of pinnule, 
elongate triangular, with to five pairs of deeply incised 
semicircular lobes, becoming less lobate to entire 
distally and apically. A single straight midvein 
traverses each pinnule, decreasing in width and forking 
in terminal lobe. Lateral veins arch into each lobe, 
forking to three times into the proximal lobes then 
decreasingly forked to simple distally. 


Holotype 
AMF113520 and counterpart, Australian 
Museum, Sydney. 


Type locality 
Coal Mine Quarry, Nymboida. Basin Creek 


71 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Formation, Nymboida Coal Measures, Middle Triassic. 


Other material 
AMEF113521, UNEF13557, 13558, Coal Mine 


Quarry. 


Name derivation 
speciosa - (Lat.) showy, beautiful, for a fossil 
that I regard as aesthetically pleasing. 


Discussion 

A number of Australian Triassic fronds have 
been placed in the form genus Sphenopteris (Tenison- 
Woods 1883; Shirley 1898; Walkom 1917, 1928; Jones 
and de Jersey 1947). However they all differ 
significantly from S. speciosa, which appears to be 
unique in Gondwana Triassic floras. 


Fern bases with radiating fronds 
Figures 33A-C, 34A-C 


Description 

Eight specimens in the collection are of 
rhizomes or stems with closely spaced, spirally 
attached fronds. Figure 33A shows a number of short, 
straight and parallel-sided rachises radiating from a 
common rhizome or stem. One rachis (Fig. 33B) bears 
four pairs of opposite rounded pinnules 12 mm apart 
arranged pinnately and apparently attached in a plane 
at right angles to the main rachis. The size of the three 
lower pairs of pinnules (pinnae?) is not known as they 
are compressed at an angle to the bedding plane. The 
uppermost preserved pinnule is 9 mm long and 7 mm 
wide. It is ovoid in outline with a contracted base, 
margin slightly lobed, apex obtuse. The venation is 
not clear but appears to be an indistinct short median 
vein that forks repeatedly to form fine lateral veins 
radiating throughout the lamina. The specimens 
illustrated in Figs 34A, B have c. 12 frond rachises 
radiating from a central axis c. 10 mm in diameter. 
They are all incomplete. From an expanded base, the 
rachises are smooth, straight, to 4 mm in width and to 
an incomplete length of 80 mm without pinnae or 
pinnules. Figure 33C is a vertical section of a rhizome 
with persistent frond rachises radiating three 
dimensionally into the matrix. 


Material 
AMF113498-113503, Coal Mine Quarry, 
Nymboida. 


Discussion 


The growth form and foliage pattern of the 
specimen with the pinnate rachis can not be related to 


1P2 


any other fern or fern-like plant at Nymboida or other 
Gondwana localities. All these specimens were at first 
mistaken for isoetalean plants until AMF1 13498 (Figs 
33A, B) demonstrated their fern-like nature. However 
the poor preservation of the present material does not 
warrant the erection of a new taxon. 


Circinate frond 
Figure 33D 


Despite the abundance of fern fossils at the 
Nymboida quarries, the specimen illustrated 
(AMF113519) is the only example in the collections 
of a circinate frond. As the specimen is detached and 
incomplete it cannot be affiliated with any known 
taxon. 


ACKNOWLEDGEMENTS 


I wish to thank my family for co-operation and 
assistance over many years; Mrs Adele Romanowski of NBI, 
Pretoria for providing many of the photographic prints; the 
curators at the Australian Museum and the Geology 
Department, University of New England for access to their 
fossil collections; Dr R. Herbst and Dr Maria Kataeva for 
assistance in obtaining South American and Russian 
references respectively; Dr H.A. Anderson for valuable 
support and advice; the Director of the National Botanical 
Institute, Pretoria, South Africa for providing research 
facilities and the two anonymous referees for their valuable 
comments. 


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Jones, O.A. and de Jersey, N.J. (1947). The flora of 
the Ipswich Coal Measures — morphology and 
floral succession. Department of Geology, 
University of Queensland Paper n.s. 3, 1-88. 

Knaus, M.J. and Gillespie, W.H. (2001). Genselia 
compacta (Jongmans et al.) Knaus et Gillespie 
comb. nov.: New insights into possible develop- 
mental pathways of early photosynthetic units. 
Palaeontologica B 256, 39-87. 

Kurtz, F. (1921). Atlas de los plantas fosiles de la 
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Lebedev, Y.L. (1974). Methods for the study of 
sterile leaves of Mesozoic ferns. Palaeontologi- 
cal Journal 2, 237-245. 

Lebedev, Y.L. and Rasskazova, E.S. (1968). 
Lobifolia, a new genus of Mesozoic ferns. Akad. 
Nauk. SSSR Geol. Inst. Tr. 191, 56-69. 

Morris, J. (1845). Fossil flora. In De Strezelecki, 
P.E., ‘Physical description of New South Wales 
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Green: London). 

Morris, N. (1975). The Rhacopteris Flora in New 
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‘Gondwana Geology’. (Australian National 


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University Press: Canberra). 

Retallack, G.J. (1977). Reconstructing Triassic 
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Alcheringa 1, 247-277. 

Retallack, G.J. (1996). Global coal gap between 
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Retallack, G.J., Gould, R.E. and Runnegar, B. 
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Rigby, J.F. (1977). New collections of Triassic 
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Russell, N.J. (1994). A palaeothermal study of the 
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van de Rijks Geologische Dienst Series C 3, 1- 
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Walkom, A.B. (1917). Mesozoic floras of Queens- 
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Walkom, A.B. (1926). Notes on some Tasmanian 
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nia, Papers and Proceedings (1925), 63-74. 

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Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 1. A-C. Cladophlebis conferta Holmes sp. nov. A. AMF120987, holotype X2; B. AMF121015; C. 
AMF120987, X4. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 75 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 2. A-C. Cladophlebis octonerva Holmes sp. nov. A. AMF113484, holotype; B. AMF113485; C. 
AMF113485, X3. Scale bar = 1 cm. 


76 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 3. A-G. Cladophlebis paucinerva Holmes sp. nov. A. AMF120980, X3; B. AMF120979 and AMF 120980 


part and counterpart, holotype and isotype; C. AMF120981; D. AMF120979, X3; E. AMF120984, X2; F. 
AMF 120982; G. AMF120983, X2. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 Wil 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 4. A, B. Cladophlebis retallackii Holmes sp. nov. A. AMF120954; B. AMF120958, X2. Scale bar = 1 
cm. 


78 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 5. A, B. Cladophlebis retallackti Holmes sp. nov. A. AMF120959, holotype; B. AMF120958, X2. Scale 
bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 79 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 6. A-C. Cladophlebis sinuata Holmes sp. nov. A. AMF113512, holotype; B. AMF113518; C. 
AMF113518, X2. Scale bar = 1 cm. 


80 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 7. A. Cladophlebis tenuipinnula Holmes sp. nov. AMF113523, holotype. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 


81 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 8. A-C. Cladophlebis tenuipinnula Holmes sp. nov. A. AMF113406; B. AMF1 13526, X3; C. AMF113524. 
Scale bar = 1 cm. 


82 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


E. ?Cladophlebis 


C. AMF113509. D, 


° 


B. AMF113510; 


?Cladophlebis sp.A. A. AMF113508; 


Figure 9. A-C. 


1 cm. 


X3. Scale bar = 


> 


E. AMF120978 


sp.B. D. AMF120978; 


83 


Proc. Linn. Soc. N.S.W., 124, 2003 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 10. A, B. ?Cladophlebis sp.C. A. AMF120994; B. AMF120994, X2.5; C. D. Dictyonymba sparnosa 
Holmes gen. et sp. nov.; C. AMF113507, holotype; D. AMF113507, X3. Scale bar = 1 cm. 


84 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 11. A—D. Gouldiopteris alethopteroides Holmes gen. et sp. nov. A. AMF113561, holotype; B. 
AMF113561, X2; C. AMF113562, X2; D. AMF113563. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 85 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 12. A-C. Leconama stachyophylla Holmes gen. et sp. nov. A. AMF121183, holotype; B. AMF121160; 
C. AMF113544, X2. Scale bar = 1 cm. 


86 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 13. A. Micronymba repens Holmes gen. et sp. nov. AMF120962, holotype, X2. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 87 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 14. A-D Micronymba repens Holmes gen. et sp. nov. A. AMF120962, holotype; B. AMF120963, isotype; 
C. AMF120962, X3; D. AMF120965. Scale bar = 1 cm. 


88 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 15. A. Nymbiella lacerata Holmes gen. et sp. nov. AMF113530, holotype. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 


89 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


i ye 
i. 

Pa, 
in: 


4 WF tim 
qt ‘s Was / 


bas ae ee ; 


Figure 16. A, B. Nymbiella lacerata Holmes gen. et sp. nov. A. AMF113532; B. AMF113533. Scale bar = 1 cm. 


90 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 17. A-E. Nymboidiantum glossophyllum (Tenison-Woods) Holmes gen. et comb. nov. A. AMF68449, 
lectotype; B. AMF120945; C. AMF120946; D. AMF120949; E. AMF120948. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 91 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 18. A, B. Nymboidiantum multilobatum Holmes gen. et sp. nov. A, B, AMF113558, holotype; B. X2 
C, D. Nymboidiantum glossophyllum (Tenison-Woods) Holmes gen. et comb. nov.; C. AMF120947; D. 
AMF12950. Scale bar = 1 cm. 


92 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 19. A, B. Nymboidiantum elegans Holmes gen. et sp. nov. A. AMF113504, holotype; B. AMF113506 


C, D. Nymboidiantum fractiflexus Holmes gen. et sp. nov. C. AMF113502, holotype; D. AMF113502, X3. 
Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 93 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 20. A-C. Nymboidiantum robustum Holmes gen. et sp. nov. A. AMF113495; B. AMF113497; C. 
AMF113496, holotype. Scale bar = 1 cm. 


94 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 21. A. Nymbophlebis polymorpha Holmes gen. et sp. nov. AMF121010, X0.75. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 95 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 22. A-C. Nymbophlebis polymorpha Holmes gen. et sp. nov. A. AMF120996; B. AMF121000, X2; C. 
AMF121001, X2. Scale bar = 1 cm. 


96 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 23. A-C. Nymbophlebis polymorpha Holmes gen. et sp. nov. A. AMF120998; B. AMF120999; C. 
AMF120997, X2. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 97 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 24. A-E. Nymbopteron dejerseyi (Retallack) Holmes gen. et comb. nov. A. AMF113549; B. AMF113548; 
C. AMF113549, X2; D. AMF113545, X2; E. AMF113547, X2. Scale bar = 1 cm. 


98 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 25. A. Nymbopteron dejerseyi (Retallack) Holmes gen. et comb. nov. AMF121158, holotype. 
B, C. Nymbopteron foleyi Holmes gen. et sp. nov. B. AMF113540; C. AMF113522, X2. Scale bar = 1 cm 


Proc. Linn. Soc. N.S.W., 124, 2003 99 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 26. A. Nymbopteron foleyi Holmes gen. et sp. nov. AMF113538, holotype. Scale bar = 1 cm. 


100 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


B. X2; 


A. holotype, X1; 


> 


nov. All AMF113556; 


Figure 27. A-D. Nymbopteron rhomboidale Holmes gen. et sp 


C. X1; D. X2. Scale bar 


1 cm. 


101 


Proc. Linn. Soc. N.S.W., 124, 2003 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 28. A. Nymbopteron uncinatum Holmes gen. et sp. nov. AMF120968, holotype. Scale bar = 1 cm. 


102 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


B. AMF113489, 


holotype; 


b) 


Figure 29. A, B. Nymborhipteris radiata Holmes gen. et sp. nov. A. AMF113489 


X4. Scale bar = 1 cm. 


103 


Proc. Linn. Soc. N.S.W., 124, 2003 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 30. A—C. Ptilotonymba curvinervia Holmes gen. et sp. nov. A. AMF113480, holotype, X2; B. 
AMF113479; C. AMF113479, X2. Scale bar = 1 cm. 


104 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


Figure 31. A. Ptilotonymba curvinervia Holmes gen. et sp. nov. AMF113483. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 105 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 32. A-D. Sphenopteris speciosa Holmes sp. nov. A. AMF113520, holotype; B. AMF113520, X2; C. 
UNEF13557; D. AMF113521. Scale bar = 1 cm. 


106 Proc. Linn. Soc. N.S.W., 124, 2003 


W.B.K. HOLMES 


. ae : 
Paes 


Figure 33. A-C. Fern bases. A. AMF113498; B. X2, pinnae arrowed; C. AMF113501, longitudinal section. 
Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 107 


TRIASSIC FLORA FROM NYMBOIDA - FERN-LIKE FOLIAGE 


Figure 34. A-C. Fern bases. A. AMF113499; B. AMF113503; C. AMF113500; D. Circinate frond. AMF1 13519. 
Scale bar = 1 cm. 


108 Proc. Linn. Soc. N.S.W., 124, 2003 


First Australian Records of Three Species and Two Genera of 
Aquatic Oligochaetes (Clitellata: Annelida) 


A.M. PINDER 


Wildlife Research Centre, Science Division, Department of Conservation and Land Management, P.O. Box 
51, Wanneroo, 6946, Western Australia. 


Pinder, A.M. (2003). First Australian records of three species and two genera of aquatic Oligochaetes 
(Clitellata: Annelida). Proceedings of the Linnean Society of New South Wales 124, 109-114. 


Recent collections of aquatic oligochaetes in New South Wales and South Australia included three 
species not previously reported from Australia. These are Nais barbata Miiller 1774 and Haemonais 
waldvogeli Bretscher, 1900 (Naididae) and Monopylephorus limosus (Hatai, 1898) (Tubificidae). The last 
two of these represent the first Australian records of their genera. Brief descriptions and locality details of 


Australian specimens are provided. 


Manuscript received 17 September 2002, accepted for publication 19 November 2002. 


KEYWORDS: Naididae, Annelida, Monopylephorus limosus, Haemonais waldvogeli, Nais barbata, 


Australia. 


INTRODUCTION 


Although knowledge of the Australian aquatic 
oligochaete fauna has improved significantly in recent 
years, there are still large areas of the country 
(including much of south-eastern Australia) for which 
few records exist. This paper presents the first 
Australian records of three species from New South 
Wales and South Australia. These are the tubificid 
Monopylephorus limosus (Hatai, 1898) and the naidids 
Haemonais waldvogeli Bretscher, 1900 and Nais 
barbata Miiller, 1774. Neither the genus 
Monopylephorus, which is represented by several 
Species in the northern hemisphere, nor the 
monospecific Haemonais have been reported from 
Australia until now. The occurrence of Nais barbata 
brings the number of species of this genus in Australia 
to six. Almost all of the naidids that occur in Australia 
are cosmopolitan or at least circumtropical (Pinder 
2001) and cosmopolitan species constitute about a third 
of the Australian tubificid fauna (Pinder and Brinkhurst 
2000). The species reported below add to this non- 
endemic component of these families in Australia. 


MATERIALS AND METHODS 


Serially sectioned specimens were cut at 6 


uum, stained with haematoxylin and eosin and mounted 
in DePeX. Dissected specimens were stained in 
Grenacher’s Borax Carmine and mounted in Permount. 
Body measurements are of preserved and slide 
mounted specimens. Specimens are held by the author 
(AP colln), returned to the New South Wales 
Department of Land and Water Conservation (DLWC) 
in Sydney or the Australian Water Quality Centre 
(AWQC), Salisbury, South Australia or deposited with 
the Australian Museum in Sydney (AMS). Collection 
localities are in New South Wales (NSW) or South 
Australia (SA). 


DESCRIPTIONS AND RECORDS 


Tubificidae 
Monopylephorus limosus (Hatai) 
(Figs la, 2) 
Synonymy 
Vermiculus limosus Hatai, 1898, 103-111, Figs 
1-5. 
Rhizodrilus limosus (Hatai): Michaelsen, 1900, 
41; Yamaguchi, 1953, 297. 
Monopylephorus limosus (Hatai): Nomura, 
1915, 1, Figs 1-30; Brinkhurst, 1971, 558, Fig. 
8.34H. 


FIRST AUSTRALIAN RECORDS OF SOME AQUATIC OLIGOCHAETES 


and rarely those of other segments simple- 
pointed, most chaetae bifid with 
rudimentary upper tooth (Fig. 1a). Chaetae 
present but not modified on X, absent on 
XI. 

Clitellum indistinct due to pale 
nature of body, but well developed on 
entire circumference of X and XI, starting 
after the spermathecal pores ventrally. 
Male and spermathecal ducts (Fig. 2a, b) 
paired but terminating in unpaired 
spermathecal and male bursae. Testes 
antero-ventral on IX. Ovaries antero- 
ventral on X. Male funnels ventro-lateral 
on 10/11, asymmetrical, usually folded, 
feeding long ciliated vasa deferentia. Vasa 
deferentia with 3 parts: a short naked 
section entally, a much longer and broader 
(up to 350 um long and 100 um wide) 
middle section covered with diffuse 
prostate tissue and then a narrow ectal 
portion without prostates. Vasa deferentia 
enter protrusible pseudopenes apically. 


iS) Pseudopenes broad medially, narrower at 


either end, with thick muscle layers 
(longitudinal over circular), thin lining 


Figure 1. Chaetae of Monopylephorus limosus and 
Haemonais waldvogeli. a, bifid chaeta of M. limosus. 
b-f, H. waldvogeli: b, two anterior ventral chaetae; 
c, posterior ventral chaeta; d. penial chaetae; e. a 
pair of dorsal chaetae (one hair and one bifid 


tissue and with a broad lumen with cilia 
on ental two-fifths. Pseudopenes 
terminating on papillae on lateral walls of 
voluminous left or right lobes of common 
median bursa. Bursa muscular, with thin 


chaeta). Scales 10 pm. 


Material examined. 

Two serially sectioned (AMS W28529 and 
W28530 ), 4 dissected (W28528 and AP collection), 
sludge in pipe from sewage treatment plant to 
abandoned ocean outfall between North Head (33°48’S 
151°18’E) and Malabar (33°57’S 151°15’E) sewage 
treatment plants, NSW, 1 May 1995. Collection by staff 
of Sydney Water. 


Description of preserved Australian material. 
Worms off-white when preserved. Length of 
body 14 — 26 mm, up to 125 segments. Prostomium 
rounded conical, anal end bluntly tapered. 
Coelomocytes abundant throughout body, circular 
(most 8 — 10 um) to elongate oval (up to 5 x 15 pm) 
with large nucleus. Ventral and dorsal chaetae similar 
in size (125 - 165 pum) and form (all crotchets, hairs 
absent), starting on II, 4 - 5/bundle in pre-clitellar 
segments, mostly 3/bundle after clitellum and 2 - 3/ 
bundle in posterior-most part of body. Chaetae of II 


‘110 


uneven lining tissue, opening to the 

exterior as a small medial pore on XI half- 

way between 10/11 and 11/12 beneath 

ventral nerve cord. Each lobe of bursa with 
a blind sac entad of the pseudopenis/bursa union. 
Spermathecal ducts each connected apically to bi-lobed 
common median bursa. Latter with lobed lining tissue 
and thick muscle layer leading to a single pore behind 
9/10. Spermathecal ducts consisting of an ectal section 
with narrow lining and thick muscle tissue, a middle 
section, with thinner muscle tissue and broad, deeply 
lobed lining tissue and an ental section which has a 
broad lumen (sperm filled in mated specimens) and 
thin lining and muscle layers. 


Remarks 

The terminology used for the male genitalia is 
based on the revelation (Gustavsson and Erséus 1999) 
that the ciliated ducts partially covered by prostate 
tissue are modified vasa deferentia in Monopylephorus, 
rather than atria as previously assumed. The Australian 
specimens mostly match the descriptions of M. limosus 
provided by Nomura (1915) and Erséus and Paoletti 


Proc. Linn. Soc. N.S.W., 124, 2003 


A.M. PINDER 


Figure 2. Genital anatomy of Monopylephorus limosus. a, spermathecal ducts entering corresponding 
lobe of median bursa. b, male ducts with pseudopenis shown passing behind (dotted lines) the 
corresponding lobe of median bursa and entering it laterally. Abbreviations: mb, male bursa; p, 
prostate; pp, pseudopenis; pp-b, union of pseudopenis with male bursa; sa, spermathecal ampulla; 
sb, spermathecal bursa; vd, vas deferens. Scale 100 um. 


(1986), which expand on the account of the original 
Japanese specimens by Hatai (1898). The new 
specimens differ from previous descriptions in that they 
have pseudopenes which are ciliated on their ental two- 
fifths. However, for the present, the Australian 
specimens are seen as being variants of M. limosus. 
Other Monopylephorus species, which are mostly 
marine, have less complex male bursae (where present 
at all), and some have eversible rather than protrusible 
pseudopenes (Baker and Brinkhurst 1981; Brinkhurst 
and Marchese 1987; Rodriguez 1999). 
Monopylephorus limosus is the only oligochaete 
known from Australian inland waters that has almost 
all chaetae bifid with minute upper teeth and with 
unpaired spermathecal and male pores. 

The tolerance of M. limosus to a range of 
salinities and to organic pollution (Chen 1940; Nomura 
1915) as well as its apparent introduction to Europe 
from Asia, suggests this is an opportunistic species 
(Erséus and Paoletti 1986). It is not clear whether the 
Australian records are part of a natural distribution, or 
whether they represent an introduction. In either case, 
M. limosus can be expected to occur at other localities 
in Australia, at least near coastal population centres 
but possibly further inland where suitable conditions 
occur, as in China (Wang Hongzhu pers. comm.). 
Embolocephalus yamaguchii (Brinkhurst 1971), 


Proc. Linn. Soc. N.S.W., 124, 2003 


known from Japan and Australia (Pinder and McEvoy 
2002), is the only other non-marine tubificid to have 
an Australasian distribution. Other freshwater species 
are either cosmopolitan or are known only from 
Australia (Pinder and Brinkhurst 2000). 


Naididae 
Haemonais waldvogeli Bretscher 
(Figs 1b-e) 
Synonymy 
Haemonais waldvogeli Brestcher, 1900, 16, Pl. 
I, Figs 11-14. 
Haemonais waldvogeli Brestcher: Sperber, 
1948, 154, Fig. 18C; Brinkhurst, 1971, 356, Fig. 
7.11M-P. 
Haemonais laurentii Stephenson, 1915, 769, 
Figs 1-5, Pl. LXXIX. 


Material examined 

Material identified by A. Pinder (on which 
the following description is based): 1 immature in 
alcohol (in 2 parts) (AMS W28531), Namoi River at 
Duncans Junction, 30°18’ 16"S 149°05’58"E, 30 Mar 
2000; 2 immature in alcohol and 2 mature mounted 
whole on slides (AMS W28532, W28533 and 
W28534), Macquarie Marsh (North) Third Crossing 


111 


FIRST AUSTRALIAN RECORDS OF SOME AQUATIC OLIGOCHAETES 


Lagoon, 30°44’33"S 147°34’27"E, 12 Apr 2000; 
Gingham Channel (Gwydir catchment) at Crinolyn, 
29°12’37"S 149°08'49"E, 10 Jan 2001; Murrumbidgee 
River at McKennas Lagoon, 34°25’53"S 145°30’32"E, 
8 Feb 2001; 1 immature, Murrumbidgee River at 
Ganmain Station 1 Storage, 35°00’40"S 147°01’58"E, 
22 Nov 2000; Murrumbidgee River at Iris Park Swamp, 
35°05’ 16"S 147°13’30"E, 20 Nov 2000. 

Additional material identified by WSL 
Consultants: Gingham Channel at Rookery, 
29°14’51"S 149°19’52"E, 11 Jan 2001; Murrumbidgee 
River at Coldene Lagoon Storage Gauge, 35°04’26"S 
147°45’32"E, 18 Dec 2000;Murrumbidgee River at 
Ganmain Station | Storage, 35°00’40"S 147°01°58"E, 
17 Jan 2001; Murrumbidgee River at McKennas 
Lagoon, 34°25’53"S 145°30’32"E, 29 Nov 2000; 
Murrumbidgee River at Sunshower Lagoon, 
34°36’28"S 146°01’06"E, 27 Nov _ 2000; 
Murrumbidgee River at Yarradda Lagoon, 34°35’ 10"S 
145°49°21"E, 28 Nov 2000 and 23 Jan 2001. 
Collection by Chris Burton, Sean Grimes, Lorraine 
Hardwick, David Hohnberg, James Maguire, Warwick 
Mawhinney and Sue Powell (DLWC). 


Description of Australian Material 

Most specimens incomplete, 2 complete 
specimens 6.5 mm long, width at VI up to 0.55 mm, 
number of segments up to 55. Gut containing a variety 
of diatoms and other algae. Dorsal chaetae (Fig. le) 
starting in XVI to XXI, each bundle with 1 - 2 short 
(130-150 um long by 2.5 um wide), curved hairs with 
a blunt tip and an equal number of crotchet chaetae. 
The latter are 100 — 120 um long by 4.5 — 5 um wide 
at the distal (1/3 - 2/5 from the distal end ) nodulus 
with long slightly curved teeth, the upper tooth 1.5 — 2 
times longer than the lower. Ventral chaetae all bifid 
(Fig. 1b-d), normally 3 per bundle (rarely 2), those of 
first 15 - 19 segments longer, thinner and straighter 
(105-120 um long by 3-3.5 um wide at the nodulus) 
than those of posterior segments, and with upper teeth 
slightly longer than the lower. Posterior ventral chaetae 
85 — 100 um long by 4-4.5 um wide at the nodulus and 
with upper tooth shorter than the lower. Ventral chaetae 
with nodulus slightly ental in first few segments, 
thereafter becoming medial and then slightly ectal. 
Ventral chaetae of VI (penial chaetae) (Fig. 1d) possibly 
shorter (laying at awkward angle for measurement) 
but definitely broader (5-6 ym at the nodulus) than 
adjacent somatic chaetae with more curved tips and 
shorter teeth than other ventral chaetae. 

Mature specimens too flattened under 
coverslips to make out much detail of the genitalia, 
but spermathecae in V and atria in VI, both small and 
globose with short ducts. 


112 


Remarks 

The combination of similarly sized dorsal and 
ventral crotchet chaetae and the position and form of 
the dorsal chaetae are unique to Haemonais and the 
Australian specimens conform to descriptions of the 
type and only species, Haemonais waldvogeli, given 
in the literature (e.g. Sperber 1948; Harman and Harrel 
1975 and Ohtaka and Nishino 1999). Dorsal chaetae 
are known to be present in newly developed anterior 
segments (Sperber 1948) but are gradually shed, so 
may be found more anteriorly than XVI in some 
specimens. Outside of Australia, H. waldvogeli is 
widespread, with reports from Europe, Africa, Asia and 
North and South America. A variant of this species, 
not recorded from Australia but known from India and 
North America, lacks teeth on the dorsal crotchet 
chaetae. Haemonais waldvogeli is known from a wide 
variety of lentic and lotic habitats overseas, particularly 
where submerged plants are present. Australian 
specimens have been collected from a variety of 
wetland habitats, including snags, detritus and from 
samples taken amongst a wide variety of aquatic plants, 
including Azolla, water ribbon, water hyacinth, Typha, 
water couch and lignum. This species is currently 
known only from the Murray-Darling catchment of 
New South Wales, but is probably more widespread, 
at least in south-eastern Australia. 


Nais barbata Miiller 
(Figs 3a-d) 
Synonymy 
Nais barbata Miiller, 1774, 23. 
Nais barbata Miiller: Sperber, 1948, 116, Pl. 
VII, Fig. 4; Brinkhurst, 1971, 338, Fig. 7.7F- 
I. 
(?) Opsonais obtusa Gervais, 1838, 17. 
Nais obtusa (Gervais): Michaelsen, 1900, 25. 


Material examined 

1 immature on slide (AMS W28535), Nattai 
River at The Crags, NSW, 34°23’21.7"S 
150°25’31.6”E, 10 Dec 2001; 2 immature on slide 
(AMS W28536), Wingecarribee River at Berrima, 
NSW, 34°29’26.3”S 150°19’57.7”E, 10 Dec 2001; 1 
immature in alcohol, Mongarlowe River at Monga 
Bridge, NSW, 35°32734.6”S 149°55’47.5”E, 5 Dec 
2001; 1 immature in alcohol, Mongarlowe River at 
Charleyong Bridge, NSW, 35°15’02.8”S 
149°55’13.5”E, 5 Dec 2001. 2 immature in alcohol 
(AP colln), 2 immature in alcohol (AMS W28537) and 
several returned to AWQC, Murray River at Craignook 
Landing, SA, 34°53’S 139°39’E, 28 May 2002; 3 
immature in alcohol (AMS W28538) and several 
returned to AWQC, Murray River downstream of Lock 


Proc. Linn. Soc. N.S.W., 124, 2003 


A.M. PINDER 


yi 


d 


_ Figure 3. Chaetae of Nais barbata. a, ventral chaeta 
of II; b, ventral chaetae of posterior segment; c, 
dorsal needle chaetae; d, dorsal needle and hair 


chaetae. Scales 10 pm. 


3, 5km upstream of Overland Corner, 34°11’S 
140°21’E, 11 Jun 2002. Collection by Kim Clarke 
(Ecowise Environmental, Melbourne) or by Vlad 
Tsymbal, Chris Madden and D. Hicks (AWQC). 


Description of Australian Material 

Length 1.0 - 2.9 mm long and width at VI 0.15 
— 0.2 mm. Gut contents fine detritus and various 
diatoms. Dorsal chaetae (Fig. 3c, d) from VI, each 
bundle with 1 - 3 hair chaetae (150 - 200 um long by 2 
uum wide at level of body wall) with an equivalent 
number of simple-pointed needle chaetae. The latter 
58 - 82 um long by 2 um wide at the slightly distal 
nodulus, bent slightly at the nodulus, with a parallel 
sided shaft ental to the nodulus and an ectal end that 
tapers evenly to a fine point. Ventral chaetae all bifid 
(Fig. 3a, b), 3 - 4 per bundle, anteriorly with ental 
nodulus and with upper teeth longer than and slightly 
thinner than the lower, posteriorly with slightly distal 
nodulus and teeth about equal in length but the upper 
slightly thinner. Ventral chaetae of anterior bundles 
longer and thinner (69 - 78 um long and 2.5 um wide 
at the nodulus) than those of posterior bundles (53 - 
74 um long by 4 um wide). 


Remarks 


These specimens match descriptions of N. 
barbata (the type species of Nais) provided in the 


Proc. Linn. Soc. N.S.W., 124, 2003 


literature (e.g. Sperber 1948). Although these 
accounts usually state 1-5 hairs and 1-5 
crotchet chaetae per dorsal bundle, this refers 
to ranges for the species and individuals with 
a maximum of 2 or 3 hairs per bundle and 
an equivalent number of crotchets (like the 
Australian specimens) are not uncommon 
elsewhere (Tarmo Timm, Estonia, pers. 
comm.). Nais pseudobtusa Piguet, 1906 is 
the only other Nais with simple-pointed 
needles known from Australia (Naidu and 
Naidu 1980), but in that species the upper 
teeth of the posterior ventral chaetae are 
longer than the lower and the nodulus on the 
needle is located closer to the distal end than 
in N. barbata. 

Nais barbata has previously been 
collected from North America, Europe, the 
near-east, north-east Asia (Japan and 
Kamchatka) and northern India (Brinkhurst 
1971; Brinkhurst et al. 1990; Liang 1964; 
Ohtaka and Nishino 1995; Stephenson 1923; 
Timm 1999). Thus, while other Australian 
non-endemic naidids are part of more 
continuous cosmopolitan or circum-tropical 
distributions, the Australian records of Nais 
barbata are the first for the southern 
hemisphere and represent more of an outlier. The sites 
listed above are all from the Murray Darling Basin, 
ranging from upper tributaries to the main channel of 
the lower Murray, but the species is likely to be more 
widespread, at least in the south-east. Specimens from 
the upper tributaries were collected from fresh, neutral 
to moderately alkaline waters in riffles and stream 
edge/backwater samples with sediments dominated by 
coarse material (gravel to cobble) and often with 
significant algal growth but no macrophytes. 
Specimens from the lower Murray were collected from 
reaches with macrophytes and filamentous algae and 
fresh moderately alkaline water with high turbidity. 
Learner et al. (1978) found this species to be most 
abundant amongst filamentous algae in British streams 
and noted its abundance in organically polluted 
European rivers. 


ACKNOWLEDGEMENTS 


Specimens of Monopylephorus limosus were 
collected by Sydney Water and provided to the authors by 
Paul McEvoy (then at Australian Water Technologies). 
Haemonais waldvogeli specimens were collected as part of 
the NSW DLWC’s Integrated Monitoring of Environmental 
Flows program. These were provided to the author via Kylie 
Swingler (WSL Consultants) and Paul McEvoy (AWQC) 


113 


FIRST AUSTRALIAN RECORDS OF SOME AQUATIC OLIGOCHAETES 


and habitat data were provided by Sarah Rish (DLWC). 
Specimens of Nais barbata and habitat data were collected 
by Ecowise Environmental for a project managed by the 
Sydney Catchment Authority and were provided to the author 
via Phil Mitchell (Water ECOscience, Melbourne), or were 
collected by AWQC as part of the Murray Darling Basin 
Commission’s Sustainable Rivers Audit Pilot Study, funded 
through the South Australian Department of Water, Land 
and Biodiversity Conservation. Gordon Thomson (Murdoch 
University) sectioned specimens of M. limosus. Thanks to 
Christer Erséus and Wang Hongzhu for up to date information 
on the distribution of M. limosus and to Tarmo Timm and 
Reinmar Grimm for information on the morphology and 
distribution of Nais barbata. 


REFERENCES 


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genus Monopylephorus and redefinition of the 
subfamilies Rhyacodrilinae and Branchiurinae 
(Tubificidae: Oligochaeta). Canadian Journal of 
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Bretscher, K. (1900). Mitteilungen uber die 
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Brinkhurst, R.O. (1971). Naididae. In “Aquatic Oligochaeta 
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Brinkhurst, R.O. and Marchese, M. (1987). A contribution 
to the taxonomy of the aquatic Oligochaeta 
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America. Canadian Journal of Zoology 65, 3154- 
3165. 

Brinkhurst, R. O., Qi, S. and Liang, Y. (1990). The aquatic 
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Chen, Y. (1940). Taxonomy and faunal relations of the 
limnitic Oligochaeta of China. Contributions from 
the Biological Society of China, Zoological Series 
14, 1-131. 

Erséus, C. and Paoletti, A. (1986). An Italian record of the 
aquatic oligochaete Monopylephorus limosus 
(Tubificidae), previously known only from Japan and 
China. Bollettino di Zooligia. 53, 115-118. 

Gervais, P. (1838). Note sur la disposition systematique des 
annelides chetopodes de la familie des Nais. Bulletin 
de l’Academie Royale de Belgique Classe des 
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Gustavsson, L.M. and Erseus, C. (1999). Development of 
the genital ducts and spermathecae in the 
Rhyacodrilines Rhyacodrilus coccineus and 
Monopylephorus rubroniveus (Oligochaeta, 
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Harman, W.J. and Harrel, R.C. (1975). Haemonais 
waldvogeli (Naididae: Oligochaeata) now established 
in North America. The Texas Journal of Science 26, 
621-623. 


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Hatai, S. (1898.) On Vermiculus limosus, a new species of 
aquatic oligochaeta. Annotationes Zoologicae 
Japonenses 2, 103-111. 

Learner M.A., Lochhead G. and Hughes B.D. (1978). A 
review of the biology of British Naididae 
(Oligochaeta) with emphasis on the lotic 
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Liang, Y. (1964). Studies on the aquatic Oligochaeta of China 
II. On some species of Naididae from Siankiang with 
description of a new species, Allodero prosetosa. 
Acta Zoologica Sinica 16, 643-652. 

Michaelsen, W. (1900). Oligochaeta. Das Tierreich 10: 1- 
565. 

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Havniae et Lipsiae 1773-74. 

Naidu, K. V. and K. A. Naidu (1980). Nais pseudobtusa 
Piguet, 1906 (Oligochaeta: Naididae) new to 
Australia. Hydrobiologia 68, 91-92. 

Nomura, E. (1915). On the aquatic oligochaete 
Monopylephorus limosus (Hatai). Journal of the 
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1-46. 

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oligochaete fauna in Lake Biwa, Central Japan. I. 
Checklist with taxonomic remarks. The Japanese 
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Ohtaka, A. and Nishino, M. (1999). Studies on the aquatic 
oligochaete fauna in Lake Biwa, central Japan. II. 
Records and taxonomic remarks of nine species. 
Hydrobiologia 406, 33-47. 

Pinder, A. M. (2001). Notes on the diversity and distribution 
of Australian Naididae and Phreodrilidae 
(Oligochaeta: Annelida). Hydrobiologia 463: 49-64. 

Pinder, A.M. and Brinkhurst, R.O. (2000). A review of the 
Tubificidae (Annelida: Oligochaeta) from Australian 
inland waters. Memoirs of the Museum of Victoria 
58, 39-75. 

Pinder, A.M. and McEvoy, P.K. (2002). Embolocephalus 
yamaguchii Brinkhurst, 1971 (Clitellata: Tubificidae) 
from South Australian streams. Records of the South 
Australian Museum 35, 139-145. 

Rodriguez, P. (1999). Monopylephorus camachoi nov. sp., 
a new rhyacodriline worm (Tubificidae, Clitellata) 
from the Coiba Island, on the east Pacific Coast of 
Panama. Hydrobiologia 406, 49-55. 

Sperber, C. (1948). A taxonomical study of the Naididae. 
Zoologiska Bidrag Fran Uppsala 28, 1-296. 
Stephenson, J. (1915). On Haemonais laurenti n. sp., a 
representative of a little known genus of Naididae. 
Transactions of the Royal Society of Edinburgh 50: 

769. 

Stephenson, J. (1923). Oligochaeta. In “The Fauna of British 
India, Including Ceylon and Burma’ (Ed. Shipley, 
A.E.) pp. 95-518. (Taylor and Francis: London). 

Timm, T. (1999). Distribution of freshwater oligochaetes in 
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Ocean. Hydrobiologia 406, 67-81. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Behavioural Responses of Two Native Australian Fish Species 


(Melanotaenia duboulayi and Pseudomugil signifer) to 
Introduced Poeciliids (Gambusia holbrooki and Xiphophorus 
helleri) in Controlled Conditions 


KEVIN WARBURTON AND CHRISTINE MADDEN 


Department of Zoology and Entomology, University of Queensland, St. Lucia, Queensland 4072, Australia 


Warburton, K. and Madden, C. (2003). Behavioural responses of two native Australian fish species 
(Melanotaenia duboulayi and Pseudomugil signifer) to introduced Poeciliids (Gambusia holbrooki and 
Xiphophorus helleri) in controlled conditions. Proceedings of the Linnean Society of New South Wales 124, 
115-123. 


Experimental treatments to compare behavioural responses included native fish species only, natives 
plus one exotic species and natives plus both exotic species. The mosquitofish, Gambusia holbrooki frequently 
attacked both native species, but tended to nip Melanotaenia duboulayi (especially small individuals) and 
chase Pseudomugil signifer. The frequency of attacks by G. holbrooki on M. duboulayi rose when all four 
fish species were present. When food was added, all four species showed a strong increase in aggression, 
especially in the four-species treatment, where there were significant increases in the frequency of attacks 
by the swordtail Xiphophorus helleri on M. duboulay and by M. duboulayi on G. holbrooki, and of conspecific 
attacks by M. duboulayi. Increased attack frequency was associated with aggregation closer to the water’s 
surface, regardless of the presence of food. The results support the hypothesis that introduced poeciliids can 
have deleterious competitive effects on native species. However, while juvenile M. duboulayi were particularly 
vulnerable to the secondary effects of fin-nipping, P. signifer appeared to be more susceptible to physical 


displacement and reduced food capture success. 


Manuscript received 16 April 2002, accepted for publication 21 December 2002. 


KEYWORDS: rainbowfish, blue-eye, poeciliids, mosquitofish, aggression, displacement 


INTRODUCTION 


Prominent among the species of freshwater 
fish that have been introduced to Australia are the 
poeciliids Gambusia holbrooki (mosquitofish) and 
Xiphophorus helleri (swordtail). Gambusia holbrooki 
is widespread and prolific throughout much of 
Australia (Arthington 1991). Xiphophorus helleri is 
locally abundant in south-eastern Queensland 
(Arthington et al. 1983, McKay 1989, McDowall 
1996), and there is a strong chance of further spread 
of this species in tropical and subtropical Australia 
(Arthington 1991). 

Expanding ranges of introduced fishes and 
widespread declines of native populations have 
heightened interest in interactions between non- 
indigenous and resident species. Although the 
underlying mechanisms have been poorly documented, 
it is likely that Gambusia species exert diverse effects 


on other fish through interference competition, 
resource competition and predation (Lloyd 1990). 
Gambusia holbrooki are aggressive towards other 
species, and damage to fins and scales resulting from 
nipping by G. holbrooki (Lloyd 1987; Howe et al. 
1997), coupled with secondary infection, can prove 
fatal. Such aggressive behaviour by Gambusia may 
cause physiological stress, and in topminnows 
(Poeciliopsis occidentalis) it leads to reduced rates of 
feeding, fecundity and survival (Schoenherr 1981). 
Gambusia holbrooki may also consume the larvae of 
Pseudomugil signifer, melanotaeniids (rainbowfishes) 
and other Australian native fish species (Aarn & 
Ivantsoff 2001). 

The ecological impacts of X. helleri are poorly 
understood (Arthington 1991). Xiphophorus helleri 
is primarily herbivorous (Arthington et al. 1983; 
Arthington 1989) and its diet is likely to overlap more 
with small omnivores (e.g. Melanotaenia duboulayi, 


RESPONSE OF NATIVE AUSTRALIAN FISH TO INTRODUCED POECILIIDS 


Arthington 1992) than insectivorous carnivores (e.g. 
P. signifer). Xiphophorus helleri can occur at very 
high densities in shallow Australian creeks (pers. obs.), 
and it is possible that this may lead to enhanced 
interspecific resource competition through the local 
depletion of invertebrate prey. Xiphophorus helleri 
and G. holbrooki often co-occur (Arthington et al. 
1983), and the pressure of high numbers of these two 
species together could subject native species to 
significant stress (McKay 1978, cited by Arthington 
1991; Howe et al. 1997). 

Several groups of Australian native fish tend 
to occur in much lower numbers in the presence of G. 
holbrooki (Arthington et al. 1983; Lloyd 1990), but 
further work is necessary to disentangle the impacts 
of introduced species from the effects of habitat 
modification (Arthington 1991). To this end, there is 
a clear need for behavioural analyses of interspecific 
interactions (Lloyd 1990), which are likely to vary 
with the species involved (Arthington & Lloyd 1989). 
In particular, because the impact of such interactions 
tends to be more severe in confined or still water 
environments (Pen and Potter 1991) and when the 
native fish are relatively small, it is important to gather 
comparative information on the effects of density and 
body size (Howe et al. 1997). 

The present study investigated the 
behavioural responses of Melanotaenia duboulayi 
(Family Melanotaentidae) and Pseudomugil signifer 
(Family Pseudomugilidae) to Gambusia holbrooki and 
Xiphophorus helleri (Family Poeciliidae). 
Melanotaeniids and pseudomugilids are speciose and 
abundant in tropical inland waters in the Australia / 
Papua New Guinea region (McDowall 1996). 
However, although M. duboulayi and P. signifer are 
the commonest representatives of these groups in 
south-eastern Queensland streams, they have been 
driven to very low densities in disturbed habitats that 
support large populations of introduced poeciliids 
(Arthington et al. 1983). To assess the relative 
competitive potential of M. duboulayi and P. signifer 
in the presence of G. holbrooki and/or X. helleri, we 
tested the prediction that the frequency of chasing, fin 
nipping and displacement of the native species would 
increase in the presence of the exotics - especially when 
food was available, when both exotic species were 
present, and when the body size of the native 
individuals was relatively small. 


MATERIALS AND METHODS 
On several occasions in winter (May- 


September) 1998, fish were collected from a site in 
Moggill Creek, Brisbane, using bait traps and a seine 


116 


net. Individuals of these four species were transferred 
to a laboratory at the University of Queensland, where 
they were maintained in filtered, single-species aquaria 
(tank volume 0.05 m?; c. 50 fish per tank) for 1-3 weeks 
prior to experimentation. During this time they were 
fed on a commercial fish flake diet, which was the 
food used in the experiments. The tanks were filled 
with aged tapwater and maintained under artificial light 
(L:D 12:12) at a constant temperature of 23 °C 


Experiment 1. : Effects of species composition on 
behavioural interactions. 

Experimental trials were conducted in May 
and September 1998 in a 90 litre tank (1.5 x 0.5 x 
0.35m) maintained under the same conditions as the 
holding tanks. Grid markings on the tank allowed the 
vertical height in the water column of the fish to be 
estimated. The tank was screened by black plastic 
sheeting to avoid external visual bias and an eye slit 
was cut into the sheet to allow observations. 

In these trials, mixed species groups totalling 
20 fish were used and individuals were used only once. 
The ratio of mean catch rates from repeated sampling 
at the field site (7:3:6:4 for M. duboulayi, P. signifer, 
G. holbrooki and X. helleri respectively) was used as a 
guide when determining the relative numbers of fish 
in the four treatments (Table 1). The treatments were 
as follows: Treatment 1, native species; Treatment 2, 
native species plus X. helleri; Treatment 3, native 
species plus G. holbrooki; Treatment 4, native species 
plus both exotic species. 

Attempts were also made to equalise sex ratios 
and to ensure that the fish used in each trial were of a 
representative range of sizes. Fish biomass was not 
measured directly but on average the aggregate total 
length of the fish used (all species combined) was 55.6 
cm, 58.4 cm, 50.0 cm and 53.4 cm for Treatments | to 
4, respectively. The order of trials was randomised 
with respect to treatment, and ten replicate trials were 
performed for each treatment. 

In each trial, the fish were placed in the tank 
and left to settle for 15 minutes. This was followed by 
a period of focal sampling (Martin and Bateson 1990) 
to record the behaviour of individual fish: a randomly 
selected fish of each species was watched for two 
minutes, during which time its depth was recorded 
every 20 seconds by reference to the markings on the 
tank. This was followed by a five-minute sampling 
period in which the occurrence and direction of all 
agonistic interactions (chases, nips) involving a 
randomly chosen focal individual were recorded. A 
single fish flake was then placed on the water’s surface 
in the central, intermediate depth section of the tank 
and the above procedure, involving both periods of 
focal sampling, was repeated. Pilot studies indicated 


Proc. Linn. Soc. N.S.W., 124, 2003 


K. WARBURTON AND C. MADDEN 


Table 1. Summary of experiments and treatments with mean (+SE) total length (excluding tail sword 


of X.helleri) and numbers of individuals. 


Species /size group Mean length SE 


Treatment 
Expt 1 2 3 4 5 6 


(cm) 


X. helleri 

Mixed 3.39 0.25 
Small 2.50 0.13 
Large 4.50 0.11 

M. duboulayi 
Mixed 3.07 0.15 
Small 2.61 0.11 
Large 3.89 0.20 
P. signifer 2.10 0.07 
2.10 0.07 
G. holbrooki 2.00 0.05 
2.00 0.05 


that a single food source was sufficient to elicit 
behavioural responses in all fish, while promoting 
competition. Each trial therefore yielded data on 
aggressive encounters and the depth of the species 
concerned, both in the absence and presence of food. 
Treatment differences in mean chasing frequency, 
mean nipping frequency and mean depth were 
examined using analysis of variance and multiple range 
(Tukey) tests, separate analyses being carried out on 
the “food absent” and “food present” data. Primary 
data (numbers of nips and chases) were standardised 
by dividing the number of attacks received by focal 
fish per trial by the number of potential aggressors. 
For interspecific attacks the number of aggressors was 
n, where n = group size for the attacking species. For 
conspecific attacks the number of aggressors was n-1. 
Data were log-transformed prior to analysis since 
means and standard deviations were linearly related. 


Experiment 2. Effects of fish size on interspecific 
interactions. 

This experiment was designed to investigate 
the impact of relative body size on interspecific 
interactions. Melanotaenia duboulayi and X. helleri 
were each divided on the basis of body size into two 
groups (large and small) (Table 1). Because P. signifer 
and G. holbrooki are relatively small species, only one 
size class (adult fish, which were similar in size to small 
M. duboulayi and X. helleri) were used. 

There were six mixed-species treatments, as 
follows: Treatment 1, large M. duboulayi, small X. 
heller; Treatment 2, small M. duboulayi, large X. 
helleri; Treatment 3, large M. duboulayi, G. holbrooki; 


Proc. Linn. Soc. N.S.W., 124, 2003 


1 6 4 
z 8 12 
Z, 8 12 
l CA: peat 9®° we 2 0 
2 12 11 
z 12 11 
1 6 SA 3 
8 68 


— 
~ 
ON 


Treatment 4, small M. duboulayi, G. holbrooki; 
Treatment 5, P.signifer, large X. helleri; Treatment 6, 
P. signifer, small X. helleri. Table 1 gives numbers of 
fish used in the different treatments. 

There were seven replicate trials for each 
treatment. The focal sampling and data analysis 
procedures were as described for Experiment 1. 
Separate analyses were carried out to compare the M. 
duboulayi treatments (1-4) and the P. signifer 
treatments (5-6). 


RESULTS 


Experiment 1 


Aggressive encounters 
There were several statistically significant 


between-treatment differences in attack frequency 
(Fig.1; Table 2). Two main trends were apparent, 
namely: 

(a) Attack levels were significantly higher with 
the combined treatment (Treatment 4) than 
the other treatments. A notable exception to 
this trend occurred in the case of chasing of 
G. holbrooki by P. signifer, where the mean 
frequency for Treatment 3 was marginally 
significantly higher than that for Treatment 4 
(F, g= 3-99; p = 0.061). 

Significant treatment differences occurred in 
the presence of food. The only exception here 
involved attacks by G. holbrooki on M. 
duboulayi, where although the difference 


(b) 


117 


RESPONSE OF NATIVE AUSTRALIAN FISH TO INTRODUCED POECILIIDS 


between __ treatment 0.2 
means for nipping was 
only marginally 
significant in the 
presence of food (F, ,, 
=" 4,12; p=0.057), 
treatment means for 
chasing varied 
significantly in the 
absence of food (Table 


3) Gh-Md (n+) 


Table 3 summarises total 15 
attack frequency, the frequency 
of nips relative to chases, and 
total attack frequency in the 
presence of food relative to that 
in the absence of food. The 
following trends were clear: 0 

(a) On a standardised per 
capita basis, M. 
duboulayi received 
more attacks than the 0.4 
other species. Mostof 93 
these came from G. 
holbrooki pute eos 
conspecific attacks by 0.1 
M. duboulayi were also 0 
relatively common. In 
contrast, attacks on P 
signifer were less 
frequent and were 
mainly conspecific in 
nature. 

(b) Attacks by G. holbrooki 
on M. duboulayi mainly 
involved nipping. 
Other interspecific and 
conspecific attacks 
were dominated by 
chasing. 

(c) The addition of food 
increased attack 
frequency. Food addition had a greater 
relative impact on attacks by X. helleri than 
on attacks by the other species, but absolute 
levels of aggression by X.helleri were very 
low. Observations indicated that X. helleri 
were territorial and defended localised areas 
of the tank. Attacks by M. duboulayi on 
conspecifics and on P.signifer also increased 
markedly in relative terms. 


Attacks 


Depth preferences 
The four species tended to swim at different 


depths, with P. signifer nearest the surface, followed 


118 


Xh-Md (c+) 


Natives only 


Gh-Md (c-) 


Natives + X.h. 
+G.h. 


Natives +X.h. Natives + G.h. 


Treatment 


Figure la. Standardised (per capita) attack frequencies per trial. Bars 
represent means and standard errors. c, chases; n, nips; - no food; + 
food added. Species names are abbreviated (see caption for Table 2) ; 
the first-named of each pair is the attacking species. Attacks by exotic 
and native species are shown in Fig. 1(a), above, and Fig. 1(b), facing 
page, respectively. 


by G. holbrooki, M. duboulayi and X. helleri in order 
of increasing depth (Fig. 2). However, in the presence 
of food, P. signifer moved deeper while G. holbrooki 
moved closest to the surface. The addition of food 
had little impact on the preferred depth of M. 
duboulayi, but it affected X. helleri differently 
depending on treatment: when X. helleri was alone with 
the native species, its mean depth increased, but when 
G. holbrooki was also present, the mean depth of X. 
helleri decreased (Fig. 2). 

For P. signifer, G. holbrooki and M. duboulayi 
there was a consistent effect of treatment, individuals 
of all these species moving closer to the surface in 


Proc. Linn. Soc. N.S.W., 124, 2003 


K. WARBURTON AND C. MADDEN 


Attacks 


Natives only Natives + Xh. 


Treatment 
Figure 1b. 


Treatment 4 (i.e., when both G. holbrooki and X. helleri 
were present; Fig. 2). However, between-trial variation 
was high and these trends were only marginally 
significant (F, , = 4.51, p = 0.048 for M. duboulayi; 


F_.= 4.11, p = 0.058 for P. signifer; comparisons of 


1,18 
Treatments | and 4 in the absence of food). 


Feeding success 
Individual M. duboulayi tended to take food 


before individuals of the other species (Table 4). 
Overall, X. helleri had a lower per capita feeding 
success than the other species. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Md-Md (c+) 
2 
15 
1 
05 
0 
0.4 
Md-Md (n+) 
0.3 
0.2 
0.1 
0 
a Md-Gh (c+) 
0.25 
0.2 
0.15 
0.1 
0.05 
0 
07>  Ps-Gh (c+) 
06 
05 
04 
03 
02 
0.1 
0 


Natives + G.h. 


Experiment 2 


Larger individuals of M. 
duboulayi were significantly less 
susceptible to attack by G. 
holbrooki. Yn the presence of food, 
G. holbrooki chased small M. 
duboulayi approximately six times 
more often than large M. duboulayi, 
and G. holbrooki-small M. 
duboulayi chases were significantly 
more frequent than those for the 
other three combinations of M. 
duboulayi and exotic species (F, ,= 
6.74; p = 0.0019). Similarly, there 
were significant differences 
between treatments in terms of the 
frequency of nips on M. duboulayi 
(F,, ,4= 34.75, p=0.0001), with small 
M. duboulayi being nipped by G. 
holbrooki about eight times more 
frequently than were large M. 
duboulayi. Attacks by X. helleri on 
P. signifer and M. duboulayi were 
infrequent and not significantly 
affected by the relative size of X. 
helleri. 

When offered food and 
mixed with G. holbrooki, large M. 
duboulayi spent significantly more 
time in the deep section of the tank 
than did smaller individuals (en = 
4.54, p = 0.012). There were no 
significant differences in tank usage 
with respect to depth between the 
treatments with natives plus X. 
helleri. 

Natives + Xh. + 
G.h. 
DISCUSSION 


The results supported the 

prediction that the addition of food 

and the number of exotic species present would 
positively influence the frequency of attacks on the 
native species. In most cases, the highest attack rates 
were recorded when all four species were present, and 
most significant treatment differences occurred in the 
presence of food. Food has been shown to increase 
rates of aggression in other fish species (Syarifuddin 
& Kramer 1996). It is noteworthy that, although G. 
holbrooki was responsible for most interspecific 
attacks, chasing and nipping of M. duboulayi by X. 
helleri also increased to relatively high levels in the 
four-species treatment. The species enhancement 


119 


RESPONSE OF NATIVE AUSTRALIAN FISH TO INTRODUCED POECILIIDS 


Table 2. Main between-treatment differences in attack frequency. Species names have been abbreviated 
(see key); the first-named of each pair is the attacking species. Underlined treatments are not significantly 
different. Key to abbreviations: Gh = Gambusia holbrooki; Md = Melanotaenia duboulayi; Ps = Pseudomugil 


signifer; Xh = Xiphophorus helleri 


Interaction Type of Food / no F df p Treatment 
attack food differences 

Gh - Md chases No food 7.46 1,18 0.014 T4>T3 

Gh - Md nips Food 4.12 l, 0.057 T4>T3 

Xh - Md chases Food 5.10 1,18 0.037 T4>T2 

Md - Md chases Food 11.57 3,36 0.0001 T4>T3>I1>T2 

Md - Md nips Food 2.90 3,36 0.048 T4>1T2>T3>T1 

Md - Gh chases Food 5.26 1,18 0.034 T4>T3 

Ps - Gh chases Food 3.99 1,18 0.061 T3 >T4 


effect was associated with aggregation involving 
vertical movement toward the surface. When all four 
species were present in low numbers there was a 
general tendency (for all species except X. helleri in 
the absence of food) to move closer to the surface. By 
increasing the local density of fish, such aggregation 


Table 3. Frequency of attacks (Summary table). ‘Total 
attacks’ refers to nips plus chases. For each species 
combination, the ratio of nips to chases and the ratio of 
total attacks during all food trials to total attacks during all 
non-food trials are also shown. (+) indicates an 
increase in the presence of food, but where a ratio can- 
not be calculated due to a zero “no food” value. Species 
names have been abbreviated (see caption for Table 2). 


Interaction Total Nips : Food : 
attacks/ chases no food 
trial 
Gh — Md 1.57 PTB) 6.45 
Gh — Ps 0.60 0.86 4.41 
Xh — Md 0.26 0.82 24.50 
Xh — Ps 0.08 1.00 (+) 
Md — Md 1.06 0.24 9.34 
Md — Ps 0.08 0.23 9.67 
Ps — Md 0.14 0.46 (+) 
Ps —> Ps 0.79 0.16 3.59 
Md — Gh 0.21 0.52 (+) 
Md — Xh 0.05 1.00 1.33 
Ps — Gh 0.49 0.56 3.41 
Ps — Xh 0.04 (+) (+) 


120 


appeared to promote elevated levels of activity and 
more aggressive interactions. The broader diversity 
of species-specific behaviours and salient stimuli may 
also have encouraged heightened levels of activity. In 
a study of conspecific and interspecific interactions 
between brook trout, Salvelinus fontinalis, and rainbow 
trout, Salmo gairdneri (=Oncorhynchus mykiss), 
Newman (1956) postulated that the presence of 
food increased feeding activity, which in turn 
increased aggressive activity as the focus of 
attacks was displaced from food to fellow fish of 
both species. He noted that feeding fish displayed 
some movements that are associated with 
aggression, such as body undulations, swift 
darting and biting, and suggested that such 
movements constituted sign stimuli eliciting 
attacks from other fish. The increased excitement 
associated with the four-species treatment 
(Treatment 4) could not be explained in terms of 
the total number of fish, which remained constant 
across treatments. Treatment 4 had the lowest 
numbers of individuals of each of the four species 
studied. Working with gouramis (Trichogaster 
trichopterus), Syarifuddin & Kramer (1996) 
found that fish were more aggressive in smaller 
groups and attributed this to greater costs of 
contest competition with increasing group size. 
In the present study it is possible that fish were 
responding more to the numbers of individuals 
belonging to each species than to the size of the 
group as a whole, but testing this hypothesis 
would require more work. 

The four species exhibited considerable 
variation in the extent and type of aggression 
displayed. Although M. duboulayi were often 
attacked by G. holbrooki, they concentrated 


Proc. Linn. Soc. N.S.W., 124, 2003 


K. WARBURTON AND C. MADDEN 


14 mainly on conspecific rather than 
a) No food interspecific exchanges. In 
Experiment 1, attacks by G. 
holbrooki on M. duboulayi were 
over 2.5 times more frequent than 
attacks by G. holbrooki on P. 
signifer. Further, G. holbrooki 
tended to nip M. duboulayi but 
chase P. signifer. However, these 
differences were not due simply to 
the difference in body size between 
the two native species, since in 
Experiment 2 G. holbrooki attacked 
small M. duboulayi more readily 
than large M. duboulayi. Juvenile 
M. duboulayi therefore appear to be 
14 particularly susceptible to nipping. 

The depths occupied by 


Mean depth (cm) 


b) Food present the four species examined were a 
function of (a) species- and size- 
specific differences in mean depth 
preference, (b) an increased 
tendency to aggregate near the 
surface as species diversity 
increased, and (c) species-specific 
changes in depth in response to the 
addition of food. In the absence of 
food, P. signifer occurred closest to 
the surface. However, when food 
was present, G. holbrooki moved 
Natives only Natives+Xh. Natives+G.h. Natives+Xh. closest to the surface while P. 

+ G.h. signifer retreated toward the deeper 
regions of the tank. These results 
suggest that P. signifer is arelatively 
poor competitor for food at the 
surface, as does the fact that for P 
signifer (unlike M. duboulayi, X. 
helleri and G. holbrooki) feeding 
success was lowest when all four 
species were present. 


Treatment 


Figure 2. Mean (+ SE) swimming depths (in cm) by fish species, 
treatment and the absence or presence of food.. M. duboulayi, oblique 
bars; P. signifer, black bars; X. helleri, crossed oblique bars; G. 
holbrooki, crossed vertical/horizontal bars. Exotic species names are 
abbreviated (see caption for Table 2). 


Table 4. Per capita feeding success of the four species. In this table the percentage of trials when any 
member of a given species took the offered food before members of any other species has been divided by 
the number of individuals of the focal species. Species names have been abbreviated (see caption for 
Table 2). 


Treatment 
1 2 B 4 
Species Natives only Natives+ X.h. Natives+Gh. Natives + X.h.+Gh. 
Md 6.6 8.9 6.4 8.7 
Ps 12 40 45 1.0 
Xh - 0 - 2.8 
Gh - - 3.4 4.2 


Proc. Linn. Soc. N.S.W., 124, 2003 121 


RESPONSE OF NATIVE AUSTRALIAN FISH TO INTRODUCED POECILIIDS 


In Experiment 2, larger M. duboulayi were 
less attracted to the shallow part of the tank than were 
smaller M. duboulayi. In the wild, adult M. duboulayi 
tend to occur at greater depths than juveniles and are 
less inclined to feed at the surface (Hattori and 
Warburton, in press). These observations may reflect 
the general tendency for small fish species, and small 
individuals within species, to occur closer to the surface 
(Helfman et al. 1997). 

There are substantial overlaps among the four 
focal species in terms of habitat use and diets 
(Arthington et al. 1983; Arthington 1992), so that the 
potential exists for interspecific competition. Further, 
in disturbed urban streams invaded by exotic 
semiaquatic grasses the extent of open flowing water 
habitats preferred by M. duboulayi and P.signifer is 
typically reduced (Arthington et al. 1983), and the 
success of these species will depend more heavily on 
their ability to utilise the low velocity grassy edge 
habitats favoured by poeciliids. Similarly, the 
prerequisites for competition exist when mixed 
populations of native and exotic species are trapped in 
shrinking ponds during droughts (Howe et al. 1997). 
Although the present laboratory-based findings should 
not be applied in a precise predictive way to wild 
populations, they do illustrate behavioural mechanisms 
by which exotic fish species may negatively impact 
on natives under confined conditions in streams and 
ponds. Where exotic species occur at high density, 
and especially where G. holbrooki and X. helleri 
coexist, associated increases in activity and aggression 
are likely to lead to elevated stress levels, increased 
energy expenditure, reduced attentiveness to foraging 
and anti-predator vigilance, and reduced per capita 
feeding success in the native species. More research 
is required on how dynamic behavioural interactions 
between shoaling fish species affect the composition 
of local stream assemblages. There is also a need for 
allied work on the effects of variation in abundance 
ratios, food availability, temperature and cover on 
behaviour. 

In summary, in laboratory experiments the 
behaviour of G. holbrooki had clear, species-specific 
impacts on M. duboulayi and P. signifer. Gambusia 
holbrooki was responsible for both direct aggression 
and displacement, but it also encouraged increased 
activity and aggression by other species, including X. 
helleri. The presence of multiple exotic species may 
therefore exacerbate the negative impact of high 
poeciliid densities on native species such as M. 
duboulayi and P. signifer. 


122 


ACKNOWLEDGMENTS 


We are very grateful to Louise Weaver for 
collecting data for Experiment 2, to Culum Brown for his 
help in the design and preparation of this study, and to Angela 
Arthington and anonymous reviewers for commenting on 
the manuscript. 


REFERENCES 


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by Gambusia holbrooki in the Orara River, New 
South Wales. Fishes of Sahul 15, 726-732. 

Arthington, A.H. (1989). Diet of Gambusia affinis holbrooki, 
Xiphophous helleri, X. maculatus and Poecilia 
reticulata (Pisces: Poeciliidae) in streams of 
southeastern Queensland, Australia. Asian 
Fisheries Science 2, 92-212. 

Arthington, A.H. (1991). Ecological and genetic impacts of 
introduced and translocated freshwater fishes in 
Australia. Canadian Journal of Fisheries and 
Aquatic Science 48 (Suppl. 1), 33-43. 

Arthington, A.H. (1992). Diets and trophic guild structure 
of freshwater fishes in Brisbane streams. 
Proceedings of the Royal Society of Queensland 
102, 31-48. 

Arthington, A.H. and Lloyd, L.N. (1989). Introduced 
Poeciliidae in Australia and New Zealand. In 
‘Evolution and Ecology of Livebearing Fishes 
(Poeciliidae)’ (Eds. Meffe, G.K. and Snelson, F-F.) 
pp. 333-348. Prentice-Hall, New York, N.Y. 

Arthington, A.H., Milton, D.A. and McKay, R.J. (1983). 
Effects of urban development and habitat 
alterations on the distribution and abundance of 
native and exotic freshwater fish in the Brisbane 
region, Queensland. Australian Journal of 
Ecology 8, 87-101. 

Hattori, A. and Warburton, K. (in press). Microhabitat use 
by the rainbowfish Melanotania duboulayi in a 
subtropical Australian stream. Journal of 
Ethology. 

Helfman, G.S., Collette, B.B. and Facey, D.E. (1997). “The 
Diversity of Fishes’. Blackwell, Mass. 

Howe, E., Howe, C., Lim, R. and Burchett, M. (1997). 
Impact of the introduced poeciliid Gambusia 
holbrooki (Girard, 1859) on the growth and 
reproduction of Pseudomugil signifer (Kner, 1865) 
in Australia. Marine & Freshwater Research 48, 
425-34. 

Lloyd, L. N. (1987). ‘Ecology and distribution of the small 
native fish of the lower River Murray, South 
Australia and their interactions with the exotic 
mosquitofish, Gambusia affinis holbrooki 
(Girard)’. M.Sc. thesis, University of Adelaide. 


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290 pp. 

Lloyd, L.N. (1990). Ecological interactions of Gambusia 
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‘Introduced and Translocated Fishes and their 
Ecological Effects’ (Ed. Pollard, D.A.) pp. 94-97. 
Proceedings of Australian Society for Fish Biology 
Workshop, Magnetic Island 24-25 August 1989. 
Bureau of Rural Resources Proceedings No. 8. 
Australian Government Publishing Service, 
Canberra. 

Martin, P. and Bateson, P. (1990). ‘Measuring Behaviour’. 
Cambridge University Press, Cambridge. 

McDowall, R.M. (1996). “Freshwater Fishes of Southeastern 
Australia’. Reed, Sydney. 247 pp. 

McKay, R.J. (1978). “The Exotic Freshwater Fishes of 
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McKay, R.J. (1989). Exotic and translocated freshwater 
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Proceedings of the Workshop on Introduction of 
Exotic Aquatic Organisms in Asia. Asian Fisheries 
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Fisheries Society, Manila, Philippines. 

Newman, M.A. (1956). Social behavior and interspecific 
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Ecology 29, 64-81. 

Pen, L.J. and Potter, I.C. (1991). Reproduction, growth and 
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Schoenherr, A.A. (1981). The role of competition in the 
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Naiman, R.J. and Stolz, D.L.) pp. 173-203. Wiley 
Interscience, New York. 

Syarifuddin, S. & Kramer, D.L. (1996). The effect of group 
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Proc. Linn. Soc. N.S.W., 124, 2003 


123 


RESPONSE OF NATIVE AUSTRALIAN FISH TO INTRODUCED POECILIIDS 


124 Proc. Linn. Soc. N.S.W., 124, 2003 


Kchinoids of the Kairuku Formation (Lower Pliocene), 
Yule Island, Papua New Guinea: Clypeasteroida 


I1.D. LINDLEY 


Department of Geology, Australian National University, Canberra, A.C.T. 0200 


Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea: 
Clypeasteroida. Proceedings of the Linnean Society of New South Wales 124, 125-136. 


The Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea, contains a rich and diverse 
echinoid fauna. Clypeasteroid (sand dollar) echinoids are an important component of this fauna and seven 
taxa are recognised. A seagrass community included the clypeasterids Clypeaster reticulatus (Linné), 
Clypeaster latissimus (Lamarck) and Clypeaster humilis (Leske) and the laganid Laganum depressum Lesson 
in L. Agassiz, 1841. A current-swept, shallow water, sand-dwelling community included Laganum decagonale 
(de Blainville, 1827), Laganum depressum delicatum Mazzetti, 1894 and Laganum depressum sinaiticum 
Fraas, 1867. Bathymetric ranges of extant forms of these clypeasteroids suggest water depths from littoral 
to about 40 m. This diverse fauna has much in common with modern communities of the tropical Indo- 
Pacific, as well as fossil Plio-Pleistocene faunas of the Indonesian archipelago and the western Indian 


Ocean region. 


Manuscript received 18 June 2002, accepted for publication 21 August 2002. 


KEYWORDS: Echinoidea, Clypeasteroida, Laganum, Clypeaster, Tertiary, Lower Pliocene, Faunal 


Affinities, Papua New Guinea. 


INTRODUCTION 


The Lower Pliocene Kairuku Formation, 
Yule Island, Papua New Guinea (PNG) contains a rich 
and diverse assemblage of regular and irregular 
echinoids typical of faunas from the tropical Indo- 
Pacific. The fauna remains poorly understood, with 
only Tenison-Woods (1878) providing an early note 
on two species, and Lindley (2001) describing several 
species. Several faunal lists have also been compiled 
(F. Chapman in Mayo et al. 1930; F. Chapman and I. 
Crespin in Montgomery 1930). The fauna is 
geographically remote from other Tertiary echinoid 
faunas described in the literature. These include faunas 
from the Indonesian archipelago (Jeannet and R. 
Martin 1937) and at Barrow Island off the NW coast 
of Australia (McNamara and Kendrick 1994). 

The present paper is the second in a series 
describing the echinoid faunas of the Kairuku 
Formation. It is based on collections made by the writer 
during fieldwork completed in January 2002. 
Collection details are provided in the Appendix. 
Echinoid terminology used herein follows that of 
Durham (1966). For a map of the Indo-Pacific realm 
as used herein, the reader is referred to Hemminga and 
Duarte (2000: fig. 1.1). Specimens have been 
temporarily allocated Department of Geology, 


Australian National University (ANU) repository 
numbers, pending their repatriation to Papua New 
Guinea at the conclusion of studies, where they will 
be housed in the Department of Geology, University 
of Papua New Guinea, Port Moresby. 


KAIRUKU FORMATION AND ITS ECHINOID 
FAUNA 


Stratigraphy 

The Kairuku Formation was defined by 
Francis et al. (1982) as a 250 m thick sequence of 
gently easterly dipping limestone outcropping ina NW 
trending belt along the eastern coastline of Yule Island 
(Fig. 1). The formation is dominated by massive, 
variably lithified biosparite with interbedded cream- 
coloured micrite and biomicrite. The sequence can be 
traced SE from Yule Island, across Hall Sound to 
Poukama and Delena on the mainland. The formation 
contains a diverse assemblage of marine invertebrates 
including corals, gastropods, echinoids, larger 
foraminifera and pelecypods with rare vertebrate 
remains (shark teeth). Prominent bright yellow-brown 
coloured bluffs, consisting of massive, poorly lithified, 
medium-coarse grained biosparite, typically forming 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


PAPUA NEW 
GUINEA 


PORT MORESBY 


CORAL SEA 


YON 
<7) 8280-2 
2 Tete ne‘ina Beach 
' 


Catholic 
Mission 


Figure 1. Yule Island, Central Province, Papua New Guinea 
showing distribution of Kairuku Formation (after Francis et al. 
1982) and collection localities. Base from Kairuku 1: 50 000 Sheet 


8280-III (Edition 1). 


recessive sections of the eastern coastline of the island, 
were found to be richly fossiliferous for echinoids. The 
sequences examined during fieldwork represent the 
lower to mid-levels of the formation. Foraminifera 
from the unit indicate a Lower Pliocene age — zones 
N18-N19/20 (Haig et al. 1993). 

The Kairuku Formation was deposited during 
a period of rapid shallowing from mid-neritic in the 
lower part of the formation to innermost neritic higher 
in the formation (Francis et al. 1982; Haig et al. 1993). 
Haig et al. (1993) interpreted the carbonate sand facies, 
with coral-rich beds and Marginopora-rich sands, as 
a deposit of seagrass meadows. However, Lindley 
(2001) noted that the echinoid faunas of the formation 
also included robust, highly turbulent, shallow-water 
dwelling forms such as the temnopleurid Temnotrema 
macleayana (Tenison-Woods). The limestone- 
dominated Kairuku Formation, is thus interpreted to 
have accumulated on a rimmed tropical carbonate shelf 
with actively growing seaward patch reefs, seagrass 
covered banks, carbonate sand shoals and elongate 


126 


2 
BISMARCK GEA as 
I 


HALL SOUND 


depressions. The entire area 
was subject to wholesale 
disruption during storm surges 
or as a result of wind action. 
These turbulent events helped 
spread skeletal sands and coral 
rubble derived from patch 
reefs. 


SOLOMON SEA 


200 km 
ae) 


Echinoid faunas 

The Kairuku Formation 
contains contrasting echinoid 
faunas: an infaunal seagrass 
meadow community 
dominated by clypeasteroids 
with locally abundant 
burrowing spatangoids 
confined to coarse-grained 
carbonate sands in shallow 
water, and an epifaunal 
community from a highly 
turbulent niche of reef and 
creviced reef rock, with 
pockets of current-swept sand, 
consisting of temnopleuroids 
and other regular echinoids 
(Lindley 2001). The seagrass 
community included the 
clypeasterids Clypeaster 
reticulatus (Linné), C. 
latissimus (Lamarck) and C. 
humilis (Leske) and the 
laganid Laganum depressum 
Lesson in L. Agassiz, 1841. A 
current-swept, shallow water community was 
dominated by the laganids L. decagonale (de 
Blainville, 1827), L. depressum delicatum Mazzetti, 
1894 and L. depressum sinaiticum Fraas, 1867. 
Bathymetric ranges of extant forms of these 
clypeasteroids suggest water depths from littoral to 
about 40 m. 

Clypeasteroid-dominant seagrass meadow 
dwelling echinoid faunas predominate in the lower 
Kairuku Formation, south of Tete ne’ina Beach. 
Modern species of Clypeaster are shallow burrowers 
confined to medium-coarse sand, with their elevated 
respiratory petaloids serving as ‘snorkel’ structures 
(Seilacher 1979). The laganids are also shallow 
burrowers, only sieving the uppermost sand layer 
(Seilacher 1979). Burrowing heart urchins are very 
common and/or dominant in the mid-levels, north of 
Aru’re village. Of the regular echinoids, the compact, 
robust Temnotrema macleayana (Tenison-Woods), an 
uncommon echinoid observed only in the lower levels 
of the fomation, was a highly turbulent, shallow-water 


Poukama 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


dwelling form, confined to patch reef habitats. A 
toxopneustid echinoid, a common component of the 
fauna at locality 8280-3, about mid-level in the 
formation, inhabited a current-swept, shallow water 
habitat. 

Despite their fine preservation, some of the 
echinoid faunas of the Kairuku Formation represent 
death assemblages. At locality 8280-4, north of Aru’re 
village, high bluffs of massive yellow-brown biosparite 
are crowded with an infauna of burrowing spatangoids, 
whose tests are preserved in a range of orientations, 
and numerous small laganid tests, many resting on their 
edges or inverted (Fig. 1). This fauna is indicative of 
the effects of the wholesale disruption and 
redistribution of sediment resulting from a succession 
of large storm events. A thin interbed of coarse, chaotic 
limestone breccia, containing a fauna dominated by 
Pecten sp. and the large clam Tridacna sp. with broken 
tests of heart urchins and laganids, is probably the 
shallow water deposit derived from a storm related 
redistribution of coral and skeletal debris from seaward 
patch reefs. 


Faunal affinities 

The tropical Indo-Pacific affinity of the 
Tertiary echinoid faunas of PNG was noted by Lindley 
(2001). The diverse clypeasteroid fauna of the Kairuku 
Formation has much in common with modern 
communities and Plio-Pleistocene faunas. The 
formation contains three Clypeaster species, all found 
in waters of the Indo-Pacific. Only C. humilis is found 
as fossil, in Java (Miocene), Taiwan (Pliocene) and 
the Red Sea (Pleistocene). The diversity of the Yule 
Island clypeasterids contrasts with the impoverished 
faunas in Australia where just two species are known. 
Clypeaster gippslandicus McCoy from the Middle to 
Upper Miocene of southern Australia is a member of 
an echinoid fauna distinct from those of the tropical 
Indo-Pacific (McNamara and Kendrick 1994). 
Clypeaster butleri McNamara and Kendrick, 1994 is 
a Middle Miocene species from the Barrow Island 
fauna, regarded as having stronger affinities with 
tropical northern, rather than southern, faunas 
(McNamara and Kendrick 1994). 

Four laganids are present in the Kairuku 
Formation and reflect not only the strong Indo-West 
Pacific affinity, but interestingly, close relationships 
with Plio-Pleistocene faunas of the western Indian 
Ocean region (Red Sea, Persian Gulf and Zanzibar). 
Only Laganum depressum sinaiticum is not present in 
modern communities. Laganum decagonale and L. 
depressum are common Indo-Pacific species, known 
from fossils in Java (Mio-Pliocene), and Fiji 
(?Miocene) and Java (Pliocene), respectively. Fossil 
Laganum depressum delicatum is found in the Pliocene 


Proc. Linn. Soc. N.S.W., 124, 2003 


of Zanzibar and var. sinaiticum is known only from 
fossils in the Persian Gulf region and the Pleistocene 
of the Red Sea. 


SYSTEMATIC PALAEONTOLOGY 


Class ECHINOIDEA Leske, 1778 

Subclass EUECHINOIDEA Bronn, 1860 
Superorder GNATHOSTOMATA Zittel, 1879 
Order CLYPEASTEROIDA A. Agassiz, 1872 
Suborder CLYPEASTERINA A. Agassiz, 1872 
Family CLYPEASTERIDAE L. Agassiz, 1835 
Genus CLYPEASTER Lamarck, 1801 


Type species 
Clypeaster rosaceus (Linné), by subsequent 
designation of Desmoulins, 1835; Recent, 
Caribbean. 


Remarks 

The literature contains a great number of 
nominal species (more than 400) of Clypeaster 
(Durham 1966). Mortensen (1948a) in his monograph 
on the Clypeasteroida added to the earlier attempts of 
A. Pomel, J. Lambert and other workers in identifying 
groupings using sections or subgenera and, at the same 
time, recognised difficulties and limitations because 
of the great variation in most species of Clypeaster. 
Mortensen’s (1948a) classification emphasised internal 
test structure, ahead of test shape, nature of petals, 
nature of peristome, situation of periproct, genital 
pores, the spines etc. Although Durham (1966: U463) 
argued there is no systematic basis for the recognition 
of subgeneric groupings, Mortensen’s (1948a) key to 
the clypeasterids, based on Recent species, is useful 
for the description of the Yule Island fossils. 

F. Chapman in 1920 (in Mayo et al. 1930) 
first noted the occurrence of Clypeaster sp. in the 
Kairuku Formation. However, his subsequent detailed 
echinoid faunal list (F. Chapman and I. Crespin in 
Montgomery 1930) did not include the genus. 


Clypeaster reticulatus (Linné) 
Figs 2a-c 


Synonymy 

Echinus reticulatus Linné 1758, p. 666. 

Clypeaster reticulatus (Linné), Jeannet and R. 
Martin 1937, p. 244; A.M. Clark and Rowe 
1971, p. 144, 160. 

T. Mortensen (1948a), A Monograph of the 
Echinoidea 4(2), Clypeasteroida, p. 71-73, 
lists the previous synonymies. 


127 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


Figure 2. Clypeasterid echinoids. Clypeaster reticulatus (Linné). Lower Pliocene, Yule Island, Central 
Province. 2a-c, ANU 60557, aboral, oral and lateral views. Bar scale = 1.0 cm. Clypeaster latissimus 
(Lamarck). Lower Pliocene, Yule Island, Central Province. 2d, ANU 60558, aboral view; 2e, ANU 60571, 
sectional view showing marginal internal skeleton of isolated pillars. Bar scale = 1.0 cm. Clypeaster humilis 
(Leske). Lower Pliocene, Yule Island, Central Province. 2f-g, ANU 60561, aboral and posterolateral views. 
Bar scale = 1.0 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Description 

Test of medium size 41 mm x 32 mm, low 
with elongate pentagonal outline, with more or less 
concave sides and rounded corners. Margin of test is 
rounded. Aboral side is slightly raised towards the 
centre with a shallow depression between the petaloid 
region and the edge. The oral side is distinctly concave, 
with the distal part gently rounded and only the 
proximal part sinking steeply towards a deeply sunken 
peristome. The test is fairly strong. 

Petals are broad and petaloid area about 1/2 
to 2/3 test length. The anterior petal is the longest, with 
the antero-lateral petals slightly shorter than the other 
petals. With the exception of the anterior petal, all 
petals are closed distally. Petals are broadest distally, 
because of the greater width of pore zone in this region. 
Pore zones along petals are about 2/3 width of 
interporiferous zones; pores are small and circular. The 
ridges between pore-pairs carry 1-3 primary tubercles, 
amongst miliary tubercles. The interporifeous zone is 
distinctly raised with numerous scattered primary and 
miliary tubercles. Ambulacral furrows on oral side are 
weak and very little sunken. 

The interambulacral areas on the aboral side 
of juvenile specimens of this species have a distinctive 
sculpturing of sutural pits, recalling that of the 
temnopleurids (Mortensen 1948a: 74). This sculpturing 
disappears totally with age, with no trace in adult 
specimens, as is the case with ANU 60557. 
Interambulacral areas carry scattered primary tubercles 
set amongst a close covering of miliary tubercles. On 
the oral side there is a belt of closely spaced primary 
tubercles and miliary tubercles on the gently rounded 
distal region. Primary tubercles are weakly scattered 
on the steeply dipping region towards the peristome. 

Apical system is very small, 2 mm in 
diameter. Genital pores are small. The periproct is close 
to the posterior edge of the test; it is subrounded and 
distinctly smaller than the peristome. The peristome 
appears to be subpentagonal, about 2 mm diameter, 
and is surrounded by numerous primary tubercles. 


Remarks 

Clypeaster reticulatus (Linné), a 
characteristically small species with great variation in 
form, is type of the Rhaphidoclypus Section 
(Mortensen 1948a). This section is characterised by 
the complete absence of marginal internal laminae and 
a conspicuously concave oral side. The species was 
previously known only from extant forms found in 
the tropical waters of the Indo-Pacific, from the Red 
Sea through to the Indonesian Archipelago, to the 
Hawaiian Islands (Mortensen 1948a). The species is 
also noted from Barrow Island, NW Australia 
(McNamara and Kendrick 1994). Mortensen (1948a) 


Proc. Linn. Soc. N.S.W., 124, 2003 


recognised a subspecies, Clypeaster reticulatus 
sundaicus Mortensen 1948, distinguished by its 
possession of thin edged test. The subspecies is widely 
distributed throughout the tropical Indo-West Pacific, 
and is also known as fossil from the Lower Miocene 
of Java and the Plio-Pleistocene of East Africa and the 
New Hebrides (Jeannet and R. Martin 1937; Mortensen 
1948a). 


Material 

ANU 60557, a complete test from locality 
8280-4, northwest of Aru’re village, east coast of Yule 
Island, Central Province, PNG. Kairuku Formation, 
Lower Pliocene. 


Clypeaster latissimus (Lamarck) 
Figs 2d-e 


Synonymy 
Scutella latissima Lamarck 1816, p. 286. 
T. Mortensen (1948a), A Monograph of the 
Echinoidea 4(2), Clypeasteroida, p. 63, lists 
the previous synonymies. 


Description 

Test of small size 86 mm x 48 mm by 
comparison with measurements provided by 
Mortensen (1948a) for this largest of the Clypeasters. 
Test very low, very gradually rising towards the centre, 
with an ovoid-subpentagonal outline, distinctly 
truncated at posterior end. Edge of test is very thin. 
Oral side flattened. 

Petals are elongate-ovoid shape, broadest in 
the middle. Anterior petal slightly longer than others. 
Petals closed distally. Petals area slightly more than 
half test-length. Pore zones are wide, 2-2.5 mm, with 
the interporiferous area just over three times the width 
of a pore zone. Inner pores are small, circular; outer 
pores may be elongate as in ANU 60559. There are 2- 
5 primary tubercles on ridges between pore pairs. 
Mortensen (1948a) noted for young specimens there 
are only 3-5 tubercles on the ridge and for mid-sized 
specimens, c. 100 mm length, 8-10 primary tubercles. 
The interporiferous zone is flush with the test. 
Ambulacral furrows shallow. 

Interambulacra are in contact with the apical 
system. Tuberculation on aboral and oral surfaces is 
uniformly fine and dense, in excess of 300 per cm2. 

The apical system is very small, only c. 2 mm 
diameter. Details of genital pores, peristome and 
periproct unknown. 


Remarks 


The large flat, thin edged test of ANU 60558, 
with broad, ovoid and closed petals indicates 


129 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


placement in the Coronanthus Section (Lambert 1910; 
Mortensen 1948a). Mortensen (1948a) described this 
section as including forms with a large flat test and 
thin edge. The largest specimen noted by Mortensen 
(1948a) has a length x width of 235 mm x 190 mm; 
the smallest 33 mm x 31 mm, with many greater than 
102 mm x 87 mm. The oral side is flat and periproct 
near the edge, circular about the size of the peristome. 
Petals are broad, ovoid and nearly closed. In addition 
to C. latissimus (Lamarck), the section also includes 
C. amplificatus Koehler, 1922, C. pateriformis 
Mortensen (1948b) and C. telurus H.L. Clark, 1914. 
Clypeaster latissimus is a Recent species known from 
the tropical waters of the Indonesian Archipelago 
(Sunda Strait, Celebes and Java Sea) and the coast of 
Indo-China (Mortensen 1948a). The Sunda Strait 
specimens were collected at water depths of 25-45 m 
(Mortensen 1948a). 


Material 

ANU 60558, a complete test; ANU 60559, 
60560 and 60571 fragmentary tests from locality 8280- 
1, immediately south of Tete ne’ina Beach, east coast 
of Yule Island, Central Province, PNG. Kairuku 
Formation, Lower Pliocene. 


Clypeaster humilis (Leske) 
Figs 2f-g 


Synonymy 

Echinanthus humilis Leske 1778, p. 185. 

Clypeaster rosaceus (Linné), Gerth 1922, p. 
504: Miocene, Java. 

Clypeaster humilis Jeannet and R. Martin 1937, 
p. 242: Upper Miocene, Java; H.L. Clark 
1946, p. 337; A.M. Clark and Rowe 1971, p. 
144, 161. 

T. Mortensen (1948a), A Monograph of the 
Echinoidea 4(2), Clypeasteroida, p. 88-90, 
lists the previous synonymies. 


Description 

Test of moderate size, distinctly longer than 
broad; estimated at 113 mm x 90 mm, at the upper end 
of test sizes noted by Mortensen (1948a). Greatest 
breadth at the antero-lateral petals. Edge is concave in 
the lateral ambulacra, while the posterior end is 
rounded. The edge is rather thin, the distal part of the 
test gently rising from the ambitus to the petaloid 
region, from where it steepens towards the apex. 
Marginal internal skeleton consists of isolated pillars. 

Petaloid region is a little more than 1/2 of 
test length. The posterior and antero-lateral petals are 
closed distally, details of anterior petal not known. 
Petals are of narrow, elongate ovoid shape, distinctly 


130 


broadest at the distal end. The interporiferous zone is 
slightly elevated in this large specimen. Pore zones 
along petals are about 1/2 width of interporiferous 
zone; inner pores are small and circular, outer pores 
elongate. The ridges between pore pairs carry a regular 
series of 7-9 primary tubercles. Oral ambulacral 
furrows distinct and extend nearly to edge of test. 

Interambulacral areas are in contact with the 
apical system because of narrow petals. Tuberculation 
on both aboral and oral surface is uniformly dense, 
with a close covering of primary tubercles set amongst 
miliary tubercles. 

Apical system is poorly preserved, c. 3 mm 
diameter; details of genital pores unknown. The 
periproct small, situated about its own diameter from 
the edge of the test. Details of peristome unknown. 


Remarks 

The Indo-Pacific Clypeaster humilis (Leske) 
and the closely allied C. subdepressus (Gray) from the 
West Indies, belong to the Stolonoclypus Section. 
These clypeasterids possess a test with flattened margin 
and raised central part; a flat or concave oral side; petals 
more or less distinctly closed; and a well developed 
marginal internal skeleton (Mortensen 1948a). 
Clypeaster humilis and C. subdepressus are 
distinguished using petaloid characters, including the 
degree of elevation of the interporiferous zone and the 
relative number of tubercles on ridges between pore 
pairs (Mortensen 1948a). Specimen ANU 60561 is 
readily referred to C. humilis using these characters. 
The species exhibits considerable variation in test 
outline, profile of the oral surface and size of the 
petaloid region (Mortensen 1948a). Although the Yule 
Island specimen appears to possess a distinctly 
elongated pentagonal test outline, with a gently sloping 
aboral margin, this variation falls within the limits 
described for the species by Mortensen (1948a). 

Extant forms of C. humilis are distributed 
throughout tropical waters of the Indo-Pacific, with a 
water depth range from littoral to 216 m. The species 
is well represented in the fossil record. Gerth (1922) 
and Jeannet and R. Martin (1937) described the species 
from the Miocene of Java and fossils have also been 
recorded from the Pleistocene of the Red Sea and a 
form with affinities to the species, is noted from the 
Lower Pliocene of Taiwan (Mortensen 1948a). 


Material 

ANU 60561, an incomplete test from locality 
8280-4, northwest of Aru’re village, east coast of Yule 
Island, Central Province, PNG. Kairuku Formation, 
Lower Pliocene. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Suborder LAGANINA Mortensen, 1948 
Family LAGANIDAE A. Agassiz, 1873 


Remarks 

Mortensen (1948a) and Durham (1966) 
disagree on the systematic position of several laganids 
which are represented in the Yule Island fauna. This 
confusion arises from the great variation within species 
and particularly applies to the relationships between 
Peronella Gray, 1855, Hupea Pomel, 1883 and 
Laganum Link, 1807, and Jacksonaster Lambert and 
Thiéry, 1914 and Laganum, all recognised as valid taxa 
by Durham (1966). 

Both workers disagree on the status of Hupea. 
The number of genital pores is fundamental in 
separating Laganum (five genital pores) from 
Peronella (four genital pores) and accordingly 
Mortensen (1948a) regarded Hupea as a synonym of 
Peronella. However, Durham (1966) reinstated Hupea 
as a taxon with five genital pores. The test of Hupea, a 
Pliocene-Recent form from the tropical Indo-Pacific, 
is described and figured by Durham (1966), and 
especially resembles that of Laganum decagonale (de 
Blainville, 1827). Both L. decagonale and the type 
species Hupea decagonale (Lesson) have small petals 
about 0.5 length of radius, raised apical area, a 
distinctly polygonal test outline, with submarginal 
periproct. 

Durham’s (1966) diagnoses for Laganum and 
Jacksonaster provide very little distinction between 
these taxa. Only the shape of the periproct, elongate 
in the case of Laganum, round or transversely elliptical 
for Jacksonaster, separates them. They are identical 
in many respects, including petaloid area, genital pores 
and test size and shape. That this single character is 
used for generic distinction is, in the words of 
Mortensen (1948a: 302) ‘rather ridiculous’. Mortensen 
(1948a) regarded Jacksonaster as a synonym of 
Laganum. The writer follows Mortensen’s (1948a) 
scheme. 

The faunal list of Chapman and Crespin (in 
Montgomery 1930) included four laganids: Laganum 
sp., L. bonani Klein, 1734, L. depressum Lesson in L. 
Agassiz, 1841 and Peronella sp. Laganum laganum 
(Leske) (= L. bonani) is noteworthy for its oblong 
periproct, situated midway between the mouth and the 
edge of the test (Mortensen 1948a), and was not 
observed during the present fieldwork. Similarly, 
Peronella sp., characterised by an apical system with 
four genital pores, is not recorded. 


Genus LAGANUM Klein, 1734 


Type species 
Laganum laganum (Leske), by subsequent 


Proc. Linn. Soc. N.S.W., 124, 2003 


designation. 


Remarks 

The laganid Laganum depressum Lesson in 
L. Agassiz, 1841, present in the Kairuku Formation 
on Yule Island, was described by Lindley (2001). 


Laganum decagonale (de Blainville, 1827) 
Figs 3a-b 


Synonymy 

Scutella decagonalis de Blainville 1827, p. 229. 

Scutella decagona Herklots 1854, p. 9: 
Miocene, Java. 

Peronella decagonalis Lesson in A. Agassiz, 
1872-74; Tenison-Woods 1878, p. 126; 
Etheridge 1889, p. 173, 178; Etheridge 1892, 
p. 209, 215; Jack and Etheridge 1892, p. 692; 
Tate 1894, p. 213, 214; Carne 1913, p. 17. 

Peronella sp., F. Chapman and I. Crespin in 
Montgomery 1930, p. 57. 

Echinodiscus lesueuri Jeannet and R. Martin 
1937, p. 254 [non Peronella lesueuri 
Valenciennes in L. Agassiz, 1841]. 

Jacksonaster decagonus (de Blainville), 
Jeannet and R. Martin 1937, p. 206: Pliocene, 
Java 

Laganum decagonale (de Blainville, 1827), 
A.M. Clark and Rowe 1971, p. 144, 162; 
Gibbs, A.M. Clark and C.M. Clark 1976, p. 
133. 

Fibulariid (?), Lindley 2001, p. 130. 

T. Mortensen (1948a), A Monograph of the 
Echinoidea 4(2), Clypeasteroida, p. 331, 332, 
lists previous synonymies. 


Description 

Test markedly flattened, with rounded 
decagonal outline; interambulacral edges are distinctly 
convex and broader than the linear ambulacral edges. 
In young specimens the test outline is elliptical. ANU 
60562, the largest specimen has a test length x width 
of 52 mm x 45 mm; ANU 60564, the smallest is 28 
mm x 22.5 mm. Test is thin with a weakly inflated 
edge forming a narrow margin. Oral surface shallowly 
concave. 

Petaloid area is small, about 1/2 test length 
and slightly anterior. Petals are relatively narrow and 
closed distally; the anterior petal is distinctly longer 
than the others. Plates of petals apparently simple, 
running across half the width of the petal. Pores small, 
about equal sized, conjugate. Interporiferous area is 
fairly narrow, covered by scattered primary tubercles 
and numerous miliary tubercles. Ambulacral furrows 
shallow and rapidly disappearing distally. 


131 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


Figure 3. Laganid echinoids. Laganum decagonale (de Blainville, 1827). Lower Pliocene, Yule Island, 
Central Province. 3a-b, ANU 60562, aboral and oral views. Bar scale = 1.0 cm. Laganum depressum 
sinaiticum Fraas, 1867. Lower Pliocene, Yule Island, Central Province. 3c-d, ANU 60565, aboral and oral 
views. Bar scale = 1.0 cm. Laganum depressum delicatum Mazzetti, 1894. Lower Pliocene, Yule Island, 
Central Province. 3e-f, ANU 60567, aboral and oral views; 3g-h, ANU 60566, aboral and oral views. Bar 
scale = 1.0 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Interambulacra covered by scattered coarse 
primary tubercles and fine miliary tubercles; densely 
spaced primary tubercles are present around edge of 
test on the weakly inflated margin. Tuberculation on 
the oral surface is similar to that of the aboral surface. 

Apical system raised; composed of single 
large madreporite plate; stellate with apices opposite 
interambulacra. Genital pores five, tightly grouped and 
located in adapical interambulacral position. UPNG 
F1183 presents an anomaly with genital pores located 
midway along sides of central plate, at adapical end of 
ambulacra (Lindley 2001: fig. 7c). Peristome is round. 
Periproct transversely elongate, near the posterior end, 
about its own diameter from the edge of the test. It is 
small, c. 3 mm transverse diameter in ANU 60562, 
about the same size as peristome. 


Remarks 

A single small specimen of this species, 
lacking well-defined food grooves, was tentatively 
assigned to Family Fibulariidae by Lindley (2001). 
However, the collection of additional material has 
permitted referral to Laganum decagonale (de 
Blainville, 1827). Tenison-Woods (1878) originally 
assigned the Yule Island species to Peronella 
decagonalis Lesson in A. Agassiz, 1872-74. Peronella 
and Laganum are similar in many respects, but are 
separated by the former having only four, rather than 
five, genital pores (Mortensen 1948a; Durham 1966). 

Laganum decagonale is noteworthy for its 
ten-sided and very thin test. It is a very common extant 
species in the tropical seas of the Indo-Pacific, 
particularly the Java and Philippine Seas (Mortensen 
1948a). The species has also been recorded from the 
Admiralty Islands, PNG, and the northern Great Barrier 
Reef (H.L. Clark 1925; Gibbs et al. 1976). The species 
is found in water depths ranging from c. 5 to 275 m 
(Mortensen 1948a). Fossil species synonymous with 
L. decagonale (Scutella decagonus Herklots 1854 and 
Jacksonaster decagonus Jeannet and R. Martin 1937) 
have been described from the Miocene and Pliocene 
of Java (Mortensen 1948a). 

The Yule Island species, with its densely 
spaced tubercles around the edge, bearing frill spines, 
was interpreted by Lindley (2001) to have inhabited a 
current-swept sandy substrate. The frill spines were 
used for burrowing and, by bending down, to reduce 
dislocation by currents. 


Material 

Four complete tests: ANU 60562 from 
locality 8280-4, northwest of Aru’re village; ANU 
60563 from locality 8280-1, south of Tete ne’ina 
Beach; ANU 60564 from locality 8280-2, immediately 


Proc. Linn. Soc. N.S.W., 124, 2003 


north of Tete ne’ina Beach; and UPNG F1183 collected 
by R. Perembo from locality 24 of Francis et al. (1982) 
= locality 8280-3, east coast of Yule Island, Central 
Province, PNG. Kairuku Formation, Lower Pliocene. 


Laganum depressum sinaiticum Fraas, 1867 
Figs 3c-d 


Synonymy 
Laganum attenuatum L. Agassiz and Desor 
1847, p. 132: Red Sea and Persian Gulf. 
Laganum sinaiticum Fraas 1867, p. 333: 
Pleistocene, Red Sea region. 
Laganum tumidum Duncan and Sladen 1886, 
[ds oS) 


Description 

Test of medium size, 37 mm x 33 mm, with 
subrounded outline. Edge of test is thick and rounded. 
Test margin is broadly inflated, the width of margin 
extending about 1/3 of distance to apex. Inside this 
inflated margin the test is moderately sunken, width 
of sunken region about 1/3 of distance from ambitus 
to apex. Test then rises gradually to apical system, with 
a height just above that of inflated margin. Oral surface 
is Shallowly concave. Test is fairly strong. 

Petaloid region is about 0.7 of test length. 
Petals are relatively narrow, distinctly broadest 
adapically, extend to the inflated margin and are open 
distally. Pore zones are narrow, about 1/3 width of 
interporiferous zone, with small circular pores. 
Interporiferous zone is flush with the test. 
Tuberculation of interporiferous zones is distinct with 
a covering of numerous primary and miliary tubercles, 
separated by a narrow, central corridor with only 
miliary tubercles. Ambulacral furrows are shallowly 
sunken and extend 1/2 distance to edge of test. 

Interambulacra in contact with apical system; 
covered by a uniformly scattered covering of primary 
and miliary tubercles to the mid-region of the inflated 
margin. The remainder of the inflated margin, the 
ambitus and the margin of the oral surface has a dense 
covering of primary tubercles with scattered miliary 
tubercles. The anterior section of this densely 
tuberculated margin, level with the anterolateral petals 
on both the aboral and oral surfaces, contains scattered, 
very coarse noncrenulate tubercles. 

Apical system low, stellate with apices 
opposite interambulacra. Genital pores five, tightly 
grouped and located in apical disc. Details of peristome 
unknown. Periproct is transversely elongate, located 
closer to posterior edge, about 2/3 of distance from 
mouth to ambitus. Periproct is surrounded by a narrow 
rim of primary tubercles. 


133 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


Remarks 

Laganum depressum sinaiticum Fraas, 1867 
is a subrounded, thick-edged, heavy form of L. 
depressum found in abundance in the raised 
Pleistocene deposits of the coastal plains of the Red 
Sea and also the Persian Gulf (Mortensen 1948a). The 
writer has not read Fraas’ (1867) description of var. 
sinaiticum. However, Mortensen (1948a: Plate LII, 
figs 14, 26-27) figured three fossil specimens collected 
from these beds. The figured specimens range in length 
x width from the smallest 28 mm x 19 mm to the largest 
56 mm x 48 mm. Test size and shape, petaloid area 
and shape, genital pores, ambulacral furrows and 
periproct details indicate ANU 60565 is a closely allied 
form, reinforcing the strong Indo-Pacific affinities of 
the Yule Island fauna. There are no Recent occurrences 
of var. sinaiticum Fraas. 

The distinctive marginal tuberculation, 
indicative of the possession of robust frill spines, and 
the thick, strong test of ANU 60565 suggests this 
species inhabited a sandy substrate constantly swept 
by currents. Robust frill spines borne by scattered, large 
antero-marginal tubercles, by bending down, served 
to reduce shifting and maintain test orientation. 


Material 

ANU 60565, a complete test from locality 
8280-1, south of Tete ne’ina Beach, east coast of Yule 
Island, Central Province, PNG. Kairuku Formation, 
Lower Pliocene. 


Laganum depressum delicatum Mazzetti, 1894 
Figs 3e-h 


Synonymy 

Laganum fragile Mazzetti 1894, p. 217. 

Laganum delicatum Mazzetti 1894, p. 241. 

Laganum depressum Lesson in L. Agassiz 
1841, Stockley 1927, p. 115: Pliocene, 
Zanzibar. 

Laganum depressum, var. delicatum Mortensen 
1948a, p. 318. 


Description 

Test very small, length x width ranging from 
11.5 mm x 11.0 mm in ANU 60567 to 19.5 x 17.0 mm 
in ANU 60568. Outline distinctly more rounded than 
that typical of L. depressum and even quite circular in 
the tests of ANU 60566 and 60567. Test is low with 
thickened edges; oral side is distinctly concave. Test 
is fairly strong. 

Petaloid region about 0.7 of test length. Petals 
are relatively narrow, broadest adapically, open 


134 


distally. Interporiferous zone is flush with test, with 
even covering of fine tubercles. Ambulacral furrows 
are shallow and extend 2/3 distance to edge of test. 

Interambulacra in contact with apical system; 
covered by an even scattering of primary and miliary 
tubercles. Rounded ambital region with dense covering 
of primary tubercles. Density of primary tubercles on 
marginal oral surface rapidly diminishes towards the 
peristome, with primary tubercles confined to 
interambulacral regions adjacent to the peristome. 
Remainder of oral surface is covered by fine miliary 
tubercles. 

Apical system typically low, although it may 
be distinctly raised in some specimens (ANU 60568, 
60569). Genital pores five, visible in the smallest 
specimen, ANU 60567, with a test length of 11.5 mm. 
Mortensen (1948a) made a similar observation, noting 
their presence in tests of 9-10 mm length. Peristome 
round. Periproct is transversely elliptical to rounded, 
located closer to posterior edge, varying from two to 
four times its own diameter from the edge. 


Remarks 

Laganum depressum delicatum Mazzetti, 
1894 is a smaller form than the typical L. depressum, 
with none exceeding 30 mm test length (Mortensen 
1948a). As such, Mortensen (1948a) was of the opinion 
that it should not simply be included with L. depressum, 
although he was undecided whether it represented a 
distinct species or a variety of L. depressum. The 
distinctive development of marginal frill spines on the 
test of var. delicatum from Yule Island also serves to 
differentiate it from local specimens of L. depressum. 
Fossils figured by Stockley (1927) from the Pliocene 
of Zanzibar are, in the view of Mortensen (1948a: 319), 
likely to be var. delicatum. Extant var. delicatum is 
only known from the Red Sea (Mortensen 1948a). 

The presence of well developed outer 
marginal frill spines on var. delicatum were probably 
used to sieve sand, burrow and to act as a steering 
device. It is likely that the laganid was a sand dweller, 
possibly a seagrass meadow constantly swept by 
currents. 


Material 

Ten complete tests including ANU 60566 and 
60567 from locality 8280-1, south of Tete ne’ina 
Beach; ANU 60568 and 60569 from locality 8280-2, 
immediately north of Tete ne’ina Beach; and ANU 
60570 from locality 8280-4, northwest of Aru’re 
village, east coast of Yule Island, Central Province, 
PNG. Kairuku Formation, Lower Pliocene. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


ACKNOWLEDGMENTS 


Fieldwork was completed during a visit to Yule 
Island in January 2002. Travel and accommodation were 
kindly arranged by Sr. Elizabeth of the Bishop’s Office, 
Diocese of Bereina, Port Moresby. The OLSH Sisters at the 
Yule Island Mission are especially thanked for their 
hospitality. Alphonse Aisi and Daniel Salamas provided 
assistance and Nahau and Ben Roama dinghy hire, during 
the course of fieldwork. Photography of specimens was 
completed by Dr. Richard Barwick, Department of Geology, 
ANU. The helpful comments of an anonymous reviewer is 
acknowledged. This work was completed while the writer 
was a Visiting Fellow in the Department of Geology, ANU, 
and Dr. Patrick De Deckker, Head of Department, is thanked 
for the provision of departmental facilities. 


REFERENCES 


Carne, J.E. (1913). Notes on the occurrence of coal, 
petroleum, and copper in Papua. Bulletin of the 
Territory of Papua 1, 116p. 

Clark, A.M. and Rowe, F.W.E. (1971). Monograph of 
shallow-water Indo-Pacific echinoderms. British 
Museum (Natural History) Publication 690, 1-238. 

Clark, H.L. (1925). A catalogue of Recent sea-urchins in 
the British Museum. 1-278. (London). 

Clark, H.L. (1946). The Echinoderm Fauna of Australia: Its 
Composition and Its Origin. Carnegie Institution 
of Washington, Publication 566, 1-567. 

Durham, J.W. (1966). Clypeasteroids. In “Treatise on 
Invertebrate Paleontology, Part U, Echinodermata 
3’ (Ed. R.C. Moore) pp. U450-U491. (Geological 
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Etheridge, R. Jr. (1889). Our present knowledge of the 
palaeontology of New Guinea. Records of the 
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179. 

Etheridge, R. Jr. (1892). Our present knowledge of the 
palaeontology of New Guinea. In ‘British New 
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Philip and Son: London). 

Francis, G., Rogerson, R., Haig, D.W. and Sari, J. (1982). 
Neogene stratigraphy and structure of the Yule 
Island-Delena region. Geological Survey of Papua 
New Guinea Report 82/9, 45p. 

Fraas, A. (1867). Geologisches aus dem Orient. 
Jahresbericht des Vereins fur Vaterlandische 
Naturkunde zu Wiirttemberg 23. 

Gerth, H. (1922). Echinoidea. In “Die fossilien von Java’ 
(K. Martin). Sammlungen des Geologischen 
Reichs-Museums in Leiden 2, 497-520. 

Gibbs, P.E., Clark, A.M. and Clark, C.M. (1976). 
Echinoderms from the northern region of the Great 
Barrier Reef, Australia. Bulletin of the British 
Museum Natural History (Zoology) 30(4), 103- 
144. 

Haig, D.W., Perembo, R.C.B., Lynch, D.A., Milner, G. and 


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Zammit, M. (1993). Marine stratigraphic units in 
Central Province, Papua New Guinea: Age and 
depositional environments. In ‘Petroleum 
Exploration and Development in Papua New 
Guinea: Proceedings of the Second PNG 
Petroleum Convention, Port Moresby’ (Eds. G.J. 
Carman and Z. Carman) pp. 47-60. (PNG Chamber 
of Mines and Petroleum: Port Moresby). 

Hemminga, M.A. and Duarte, C.M. (2000). “Seagrass 
Ecology’. (Cambridge University Press: 
Cambridge). 

Jack, R.L. and Etheridge, R. (1892). ‘The Geology and 
Palaeontology of Queensland and New Guinea’. 
(Government Printer: Brisbane). 

Jeannet, A. and Martin, R. (1937). Ueber Neozoische 
Echinoidea aus dem Niederlaendisch-Indischen 
Archipel. Leidsche geologische mededeelingen 
8(2), 215-308. 

Lambert, J. (1910). Descriptions des Echinides néogénes du 
Bassin du Rhone: Mémoires de la Société 
Paléontologique Suisse 37, 1-48. 

Lindley, I.D. (2001). Tertiary Echinoids from Papua New 
Guinea. Proceedings of the Linnean Society of New 
South Wales 123, 119-139. 

Mayo, H.T., Montgomery, J.N. and De Verteuil, J.P. (1930). 
Part I - Reports of the First Geological Expedition, 
1920-1923: Yule Island-Delena-Bokama Area. In 
“The oil exploration work in Papua and New 
Guinea, conducted by the Anglo-Persian Oil 
Company on behalf of the Government of the 
Commonwealth of Australia, 1920-1929, Volume 
1’, pp. 17-20. (Harrison and Sons: London). 

McNamara, K.J. and Kendrick, G.W. (1994). Cenozoic 
Molluscs and Echinoids of Barrow Island, Western 
Australia. Records of the Western Australian 
Museum Supplement No. 51, 5Opp. 

Montgomery, J.N. (1930). Part V - A contribution to the 
Tertiary geology of Papua. In “The oil exploration 
work in Papua and New Guinea, conducted by the 
Anglo-Persian Oil Company on behalf of the 
Government of the Commonwealth of Australia, 
1920-1929, Volume 4’, pp. 3-85. (Harrison and 
Sons: London). 

Mortensen, T. (1948a). A Monograph of the Echinoidea 4(2), 
Clypeasteroida. C.A. Reitzel, Copenhagen. 471p. 

Mortensen, T. (1948b). Report on the Echinoidea collected 
by the United States Fisheries Steamer ‘Albatross’ 
during the Philippine Expedition, 1907-1910. Part 
3. United States National Museum Bulletin 100, 
93-140. 

Seilacher, A. (1979). Constructional morphology of sand 
dollars. Paleobiology 5(3), 191-221. 

Stockley, B. (1927). Neogene Echinoidea from the Zanzibar 
Protectorate. Report on Paleontology of the 
Zanzibar Protectorate. 

Tate, R. (1894). Note on the Tertiary fossils from Hall Sound, 
New Guinea. Proceedings of the Linnean Society 
of New South Wales 19, 213-214. 

Tenison-Woods, J.E. (1878). On a Tertiary Formation at New 
Guinea. Proceedings of the Linnean Society of New 
South Wales 2, 125-128. 


135 


LOWER PLIOCENE ECHINOIDS (CLYPEASTEROIDA) FROM PAPUA NEW GUINEA 


APPENDIX 


Collection details 


All collection sites are located along the 
eastern shoreline of Yule (Rabao) Island, Central 
Province, PNG (Fig. 1). Grid references are from the 
Kairuku 1: 50 000 Sheet 8280-III (Edition 1). 

8280-1: Collection from a 400 m interval 
along low coastal cliffs north from D’ Albertis Point, 
near old Kairuku Government Station, to Tete ne’ina 
Beach (GR 502240 to GR 502244). Interval collected 
represents the lower levels of the Kairuku Formation. 

8280-2: Collection from an approximately 
250 m interval of coastline immediately north of Tete 
ne’ina Beach. Centered on GR 501246. Interval 
collected is in the mid-levels of the Kairuku Formation, 
slightly higher level than for locality 8280-1. 

8280-3: Collection interval is a 100 m section 
of shoreline cliffs, extending south from the mouth of 
a small stream at GR 492260. This sequence probably 
represents the highest level of the Kairuku Formation 
collected. Same locality as microfossil locality 24 of 
Francis et al. (1982). Foraminifera from this site were 
assigned a N18/19 (Lower Pliocene) age. 

8280-4: Lower section of high coastal cliffs 
behind and to the immediate south of a small beach 
approximately 500 m north of Aru’re village at GR 
495259. Stratigraphically equivalent to locality 8280- 
oF 


136 


Proc. Linn. Soc. N.S.W., 124, 2003 


Echinoids of the Kairuku Formation (Lower Pliocene), 
Yule Island, Papua New Guinea: Regularia 


I.D. LINDLEY 


Department of Geology, Australian National University, Canberra, A.C.T. 0200 


Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea: 
Regularia. Proceedings of the Linnean Society of New South Wales 124, 137-151. 


Regular sea urchins are an important component of the rich and diverse echinoid fauna of the Lower Pliocene 
Kairuku Formation, Yule Island, Papua New Guinea. Seven taxa are recognised, including the cidaroids 
Phyllacanthus imperialis var. javana K. Martin, 1885, Phyllacanthus sp. and Prionocidaris verticillata 
(Lamarck, 1816), the toxopneustids Cyrtechinus verruculatus (Liitken) and Schizechinus cf. tuberculatus 
(Pomel), a temnopleurid Temnotrema macleayana (Tenison-Woods) and a parasaleniid Parasalenia poehli 
Pfeffer, 1887. The cidaroids, parasaleniid and temnopleurid occupied shallow-water reef habitats. The 
toxopneustids were dominant herbivores in adjacent seagrass meadows. The strong affinities evident between 
the seagrass meadow- and shallow-water sand-dwelling echinoid faunas of Yule Island and fossil and extant 
faunas of the Red Sea region, parallel the geographic patterns of species diversity of Indo-Pacific seagrasses, 


corals and mangroves. 


Manuscript received 30 September 2002, accepted for publication 19 November 2002. 


KEYWORDS: Cyrtechinus, Echinoidea, Palaeoecology, Papua New Guinea, Parasalenia, Phyllacanthus, 
Pliocene, Prionocidaris, Regularia, Schizechinus, Temnotrema 


INTRODUCTION 


The Tertiary faunas of Yule Island, and along 
the south coast of eastern New Guinea (Montgomery 
1930), represent the nearest marine fossil occurrences 
to the Great Barrier Reef (Fig. 1). The Great Barrier 
Reef’s rich and diverse shallow-water echinoderm 
fauna, in particular, has been intensively documented, 
including (from north to south) the Murray Islands in 
Torres Strait (H.L. Clark 1921), Cairns-Pipon Island- 
Raine Island region (Gibbs et al. 1976), Low Isles (H.L. 
Clark 1932; Endean 1956), Swain Reefs (A.M. Clark 
1975) and the Capricorn-Bunker Groups (Endean 
1953). Although it is widely accepted that a significant 
component of the echinoderm faunas of tropical 
Australia migrated southward from the East Indies via 
the Torres Strait or the Arafura Sea (A.H. Clark 1911; 
H.L. Clark 1946; Endean 1957), with no marine 
Tertiary record in Queensland, palaeontology has not 
been of use in determining migrations to this region 
(Endean 1957). Torres Strait prior to the late 
Quaternary was emergent (Australasian Petroleum 
Company 1961; Struckmeyer et al. 1993) and the 
eastern coast of New Guinea was in closest faunistic 
contact with the Great Barrier Reef, the coastline at 
the time (H.L. Clark 1946; Ekman 1953). Accordingly, 


many of the Lower Pliocene echinoids from Yule Island 
may be ancestral to those now living on the Great 
Barrier Reef and elsewhere along tropical northern 
Australian coasts. 

This paper is the third in a series describing the 
echinoid fauna of the Kairuku Formation, Yule Island, — 
Central Province, Papua New Guinea (Fig. 2). 
Elements of the Kairuku fauna have been described 
by Lindley (2001, 2003a). The present descriptions 
are based on collections made by the writer in January 
2002, and the reader is referred to Lindley (2003a) for 
collection details. Specimens have been temporarily 
allocated Department of Geology, Australian National 
University repository numbers, pending their 
repatriation to Papua New Guinea (PNG) at the 
conclusion of studies, where they will be housed in 
the Department of Geology, University of Papua New 
Guinea, Port Moresby. The classification used herein 
follows that of Fell and Pawson (1966). 


PALAEOECOLOGY 


Foraminifera have been used by Haig et al. 
(1993: Fig. 5) to demonstrate rapid shallowing during 
the deposition of the Kairuku Formation, from mid- 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


= 0 \iet eae 


(<) 

A 
(iter 
a ee 
PO is 
fa * a 
5 aig: 
pg 
wee: 
rap ote 
Bote 2 

eae 
mend be 
oe 
ae 


Ee 


8 Swain Reefs 


a | Capricorn- 
"A. > Bunker Groups 


Figure 1. Locality map showing proximity of Yule Island to northern Great Barrier Reef. 


neritic in the lower part to innermost neritic carbonate 
sand facies higher in the formation. The carbonate sand 
facies, predominant in the sequence collected during 
the present work, includes coral-rich beds and 
Marginopora-tich sands and has been interpreted by 
Haig et al. (1993) to have accumulated in seagrass 
meadows. However, Lindley (2001, 2003a) using the 
echinoid fauna of the Kairuku Formation, concluded 
that although some elements of the fauna, including 
the clypeasterids and laganids, were undoubtedly 
members of an infaunal seagrass community confined 
to shallow-water coarse sands, others occupied 
turbulent-water reef habitats and shallow-water sandy 
substrates constantly swept by currents. Bathymetric 
ranges of extant species of Yule Island echinoids 
indicate water depths during the deposition of the 
Kairuku Formation from littoral to about 40 m (Lindley 
2003a), closely coinciding with Hemminga and 
Duarte’s (2000) observation that the majority of 
modern seagrasses are confined to depths of less than 
20 m. 

Tropical echinoids are typically generalist 
feeders, consuming algae, seagrasses and invertebrates, 
and considerable data is available on the impacts of 
their grazing patterns on other coral reef organisms 
(Hatcher 1983). Echinoids are the dominant 


138 


invertebrate herbivore in tropical and sub-tropical 
seagrass communities (McPherson 1965, 1968; Ogden 
et al. 1973; Hemminga and Duarte 2000). High 
population densities of the toxopneustid Lytechinus 
A. Agassiz, 1863 commonly leads to overgrazing 
events in the seagrass beds of the northern Gulf of 
Mexico (Valentine and Heck 1991). The toxopneustid 
Tripneustes L. Agassiz, 1841 is a dominant consumer 
of live seagrass leaves in PNG and Philippine seagrass 
beds (Nojima and Mukai 1985; Klumpp et al. 1993). 
Therefore, it is likely that Cyrtechinus verruculatus 
(LYtken) and Schizechinus sp., close relatives of 
Lytechinus and Tripnesutes, were important herbivores 
in the seagrass meadow community at the time of 
deposition of the Kairuku Formation. The relative 
abundance of C. verruculatus as a fossil in the 
formation is probably related to the enhanced 
preservation potential associated with this habitat. 
Turbulent, shallow-water habitats were 
developed in and around fringing reef, located seaward 
of the apparently extensive seagrass meadows. 
Parasalenia poehli Pfeffer, 1887 occupied well 
concealed habitats, among branches of corals or hidden 
in crevices beneath coral rock. The small cidaroid 
Prionocidaris verticillata (Lamarck, 1816) was 
restricted to the reef or lived on adjacent coral sands. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


HALL SOUND 


7 >». 8280-3 
DX, 8280-4 


Tsiria 


4 Kairuku Formation 


8280-4 Collection locality 


y Aru're 


8280-2 
Tete ne’ina Beach 
1 


8280-1 
~ D'ALBERTIS POINT 


Kairuku 


Catholic 
Mission 


HALL SOUND 


Poukama 


Figure 2. Yule Island, Central Province, Papua New Guinea, showing distribution of Kairuku 
Formation and collection localities. Base from Kairuku 1:50 000 Sheet 8280-III (Edition 1). 


The large cidaroids Phyllacanthus imperialis var. 
jJavana K. Martin, 1885, and Phyllacanthus sp. may 
have favoured the outer side of the reef below low- 
tide mark, similar to living P. imperialis on the nearby 
Murray Islands (H.L. Clark 1946). The small 
temnopleuroid Temnotrema macleayana (Tenison- 
Woods) was also an element of the highly turbulent- 


Proc. Linn. Soc. N.S.W., 124, 2003 


water habitat (Lindley 2001). 

Echinoids play a major role in reef destruction and 
may be growth limiting (Davies 1983). Glynn et al. 
(1979) have shown that the cidaroid Eucidaris Pomel, 
1883, an inhabitant of intertidal reefs, grazes heavily 
on live corals, estimating an annual rate of bio-erosion 
of marine limestone attributable to this echinoid in the 


139 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


order of 1.7 kg/m2. The high rate of marine erosion 
caused by Eucidaris is important in the generation of 
major reef-flanking sediment accumulations composed 
of coral and coralline particles excreted by the echinoid 
(Glynn et al. 1979). 

The strong affinities of elements of the Yule 
Island fauna with both Indo-Pacific and Red Sea 
echinoid communities have been highlighted in 
Lindley (2003a) and the present work. These affinities 
are strongest amongst the species-rich infaunal and 
epifaunal echinoid communities of the seagrass 
meadow and the current-swept shallow-water sands. 
Similar geographic patterns of species richness are 
evident in corals and mangroves, and parallel seagrass 
species richness (Heck and McCoy 1979). Seagrass 
meadows with the richest species diversity are found 
in the Indo-Pacific and the Red Sea region and the 
parallelism present between the Yule Island echinoids 
and seagrasses suggests, in the words of Hemminga 
and Duarte (2000), that the ‘constraints and processes 
responsible for the development and maintenance of 
species diversity of these taxa have been linked 
throughout their evolutionary history’. 


SYSTEMATIC PALAEONTOLOGY 


Class ECHINOIDEA Leske, 1778 

Subclass PERISCHOECHINOIDEA M’Coy, 1849 
Order CIDAROIDA Claus, 1880 

Family CIDARIDAE Gray, 1825 

Subfamily RHABDOCIDARINAE Lambert, 1900, 
emended Fell 1966 

Genus PHYLLACANTHUS Brandt, 1835 


Synonymy 
Leiocidaris Desor, 1885, p. 48. 


Type species 
Cidarites (Phyllacanthus) dubia Brandt, 1835, 
p. 67, by original designation. 


Remarks 

Phyllacanthus Brandt, 1835 is a strictly Indo- 
Pacific and Australasian genus, generally restricted by 
most workers (Mortensen 1928; Chapman and 
Cudmore 1934; H.L. Clark 1946; Philip 1963; A.M. 
Clark and Rowe 1971) to forms having thick, smooth 
and cylindrical spines. It is on this basis that the Yule 
Island specimens can clearly be assigned to 
Phyllacanthus. Mortensen (1928) and Philip (1963) 
provided reviews of extant and fossil species of this 
genus. Seven living species have been described by 
Mortensen (1928, 1936), with 4 being confined to the 
Australian coast (Mortensen 1928; H.L. Clark 1946; 


140 


Philip 1963). While fossil test fragments are rare, spines 
of Phyllacanthus are well documented in the Tertiary 
sequences of the Indo-Pacific region and include 
Phyllacanthus javana K. Martin, 1885 and 
Phyllacanthus imperalis var. javana K. Martin from 
the Miocene of Java (Jeannet and R. Martin 1937); 
Phyllacanthus imperialis (Lamarck) from Madagascar, 
Middle Pliocene of Java, Upper Miocene of Vanuatu, 
Lower Miocene to Pleistocene of Fiji and the 
Quaternary of the Suez region (Mortensen 1928; 
Jeannet and R. Martin 1937; Philip 1963); 
Phyllacanthus dubius Brandt, from the Middle 
Pliocene of Java (Jeannet and R. Martin 1937); 
Phyllacanthus dubius var. sundaica (R. Martin) (non 
Phyllacanthus sundaica K. Martin, 1885) from the 
Lower Miocene of Java (Jeannet and R. Martin 1937); 
and Phyllacanthus sp. from the Pliocene of Kenya 
(Philip 1963). 

Phyllacanthus is a littoral genus that exhibits a 
marked preference for seas in which the surface 
temperature does not fall below the winter isotherm 
of 15°C (Fell 1966). Like other cidaroids it is a shallow- 
water dweller, with a preference for hard bottom, such 
as reefs (Mortensen 1928; Fell 1966). Cidaroids feed 
upon bottom animals, including molluscs, annelids, 
bryozoans, foraminifera and sponges (Fell 1966). 

F. Chapman and I. Crespin (in Montgomery 
1930) recorded the presence of spines of Phyllacanthus 
sundaica K. Martin, 1885 from ‘the limestones of the 
upper part of e stage’ at Delena, across Hall Sound on 
the mainland (= Ou Ou Limestone Member of the 
Middle to Late Miocene Lavao Group, Yule Island: 
Francis et al. 1982) (Fig. 2). This species is now 
considered a synonymn of Chondrocidaris gigantea 
(A. Agassiz) (Mortensen (1928: 492). The primary 
spines of this large species carry very coarse thorns in 
a random arrangement. Spines of this type were not 
collected from the Kairuku Formation during the 
present fieldwork. 


Phyllacanthus imperialis var. javana K. Martin, 
1885 
Figs 3a-b 


Synonymy 

Phyllacanthus javana K. Martin, 1885, p. 289: 
Mortensen 1928, p. 503; Philip 1963, p. 202; 
Upper Miocene, Java. 

Phyllacanthus cf. imperialis, cf. dubia Duncan 
and Sladen, 1885, p. 284; Miocene. 

Phyllacanthus javanus: Gerth 1922, p. 517; F. 
Chapman and I. Crespin in Montgomery 
1930, p. 57, 58; Chapman and Cudmore 
1934, p. 131; Lindley 2001, p. 119; Upper. 
Miocene, Java; Lower Pliocene, Yule Island. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Figure 3. Cidaroid echinoids. Phyllacanthus imperialis var. javana K. Martin, 1885. Lower Pliocene, Yule 
Island, Central Province. 3a, ANU 60592, lateral view of spine. Bar scale = 10 mm; 3b, ANU 60598, detail 
of distal spine end. Bar scale = 2.5 mm. Phyllacanthus sp. Lower Pliocene, Yule Island, Central Province. 
3c, ANU 60590, lateral view of spine. Bar scale = 10 mm. Prionocidaris verticillata (Lamarck). Lower 
Pliocene, Yule Island, Central Province. 3d, ANU 60589, lateral view of incomplete spine. Bar scale = 2.5 


mm. 


Leiocidaris imperialis: Jeannet 1928, p. 465. 

Leiocidaris (Phyllacanthus) imperialis: Jeannet 
1934, p. 13; Miocene, Java. 

Phyllacanthus imperialis var. javana K. Martin, 
Jeannet and R. Martin 1937, p. 222, 223; 
Philip 1963, p. 202; Upper Miocene, Java. 


Description 

No test fragments which belong to this species 
have been identified. Primary spines are moderately 
thick, cylindrical, slightly fusiform. Spine length 
ranges from 36-80 mm, with a maximum diameter 
(measured on largest spine) of 9 mm, occurring at 1/3 
distance from proximal end; spine gently tapers 


Proc. Linn. Soc. N.S.W., 124, 2003 


towards apex. Collar is finely longitudinally striated; 
collar length between 3.5-5 mm, with an obvious 
swelling at about 7 mm above collar on spine ANU 
60592 of 80 mm length. Surface of shaft with a smooth 
appearance, but with low magnification a fine ornament 
is evident, consisting of low, rounded granules forming 
numerous (> 50) longitudinal ridges. On the distal 1/3 
of spine the numerous, occasionally sinuate ridges 
merge, and interspaces become narrower, to form fewer 
(15-20), somewhat higher ridges, passing to a fluted 
spine end. 


141 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


Remarks 

Jeannet and R. Martin (1937) noted that the 
spines of Phyllacanthus imperialis var. javana K. 
Martin, 1885 from the Upper Miocene of Java, are 
generally slimmer and are often more pointed than 
those of Phyllacanthus imperialis (Lamarck). They 
also noted that the spine collar length of var. javana, 
ranging from 2.5-4 mm, is distinctly shorter than the 6 
mm of P. imperialis. The spines of P. imperialis, and 
by association those of P. javana, are relatively short, 
usually equal to test diameter (Mortensen 1928). 
Mortensen (1928) considered var. javana a close 
relative to P. imperialis. 

Phyllacanthus imperialis is the only species of 
Phyllacanthus that is widespread throughout the 
tropical Indo-Pacific (Mortensen 1928; H.L. Clark 
1946; A.M. Clark and Rowe 1971). Phyllacanthus 
imperialis from the Murray Islands, Torres Strait, has 
a distinctive habitat, favouring the outer side of reef 
below the low-tide mark (H.L. Clark 1946). 


Material 

Eight isolated primary spines: ANU 60593, 
ANU 60596-98 from locality 8280-1; ANU 60592 
from locality 8280-3; ANU 60594-95, ANU 60599 
from locality 8280-4. All localities are from the east 
coast of Yule Island, Central Province, PNG. Kairuku 
Formation, Lower Pliocene. 


Phyllacanthus sp. 
Fig. 3c 


Description 

No test fragments which belong to this species 
have been identified. Primary spines are moderately 
thick, cylindrical, fusiform, with length ranging from 
37.5-67.5 mm, with a maximum diameter of 9.5 mm 
measured at 1/3 distance from proximal end on largest 
spine. Spine gently tapers to apex. Collar length 3 mm 
in ANU 60590; details of collar ornament unknown. 
Surface of shaft is finely and uniformly granulated (not 
visible to the naked eye), the granules arranged in 
regular longitudinal series along length of spine. The 
distal 1/5 length of spine carries nine high ridges 
passing to a fluted spine end. 


Remarks 

The primary spines of Phyllacanthus sp. in 
many respects resemble those of var. javana, but the 
presence of notably fewer high ridges on the distal part 
of the spine is in contrast to the many ridges (15-20) 
present on spines of the latter form. The Yule Island 
spine probably represents another variety of P. 
imperialis. 


142 


Material 

Two isolated primary spines: ANU 60590-91 
from locality 8280-1, immediately south of Tete ne’ ina 
Beach, east coast of Yule Island, Central Province, 
PNG. Kairuku Formation, Lower Pliocene. 


Genus PRIONOCIDARIS A. Agassiz, 1863 


Synonymy 
Stephanocidaris A. Agassiz, 1872 (non 1863). 
Plococidaris Mortensen, 1909, p. 51, 51. 


Type species 
Cidarites pistillaris Lamarck, 1816, p. 55. 


Remarks 

The status of the name Plococidaris Mortensen, 
1909, with Cidarites bispinosa Lamarck, 1816 
designated as type species, has been the subject of 
discussion following Mortensen’s (1928) referral of 
bispinosa to Prionocidaris A. Agassiz, 1863 (H.L. 
Clark 1946: 287; A.M. Clark and Rowe 1971: 151). 
These workers considered Mortensen’s (1928) 
retention of the name Plococidaris, with Cidarites 
verticillata Lamarck, 1816 redesignated as type 
species, to be contrary to the Rules of Nomenclature, 
and it is regarded as a synonym of Prionocidaris. 


Prionocidaris verticillata (Lamarck, 1816) 
Fig. 3d 


Synonymy 

Cidarites verticillata Lamarck, 1816, p. 56. 

Phyllacanthus cf. verticillata Duncan and 
Sladen, 1885, p. 284; Miocene, India. 

Cidaris verticillatus: Lemoine, 1906, p. 256; 
Miocene, Madagascar. 

Phyllacanthus verticillatus Cottreau, 1908, p. 38. 

Plococidaris verticillata Mortensen, 1909, p. 51, 
53; Mortensen 1928, p. 428; Jeannet and R. 
Martin 1937, p. 220. 

Prionocidaris verticillata Déderlein, 1911, p. 
242; H.L. Clark 1921, p. 145; H.L. Clark 
1932, p. 211; H.L. Clark 1946, p. 287; A.M. 
Clark and Rowe 1971, p. 151. 

Leiocidaris (Plococidaris) verticillata: Jeannet 
1934, p. 11; Pliocene, Ceram. 

T. Mortensen (1928), A Monograph of the 
Echinoidea 1, Cidaroidea, p. 428, details the 
previous synonymies. 


Description 

No test fragments which belong to this species 
have been identified. Primary spine small, slim, 
cylindrical with an incomplete length of 15 mm and a 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


diameter of 1.5 mm. Collar length ca. 0.7 mm; details 
of collar ornamentation unknown. ANU 60589 with 
characteristic prominent whorls of blunt thorns, placed 
one above the other; two are preserved in specimen 
with three-four usually present in complete primary 
spines (Mortensen 1928; A.M. Clark and Rowe 1971). 
Smaller blunt thorns arranged in longitudinal series 
occur between successive whorls. 


Remarks 
Prionocidaris verticillata (Lamarck) is a small 

(30-40 mm diameter) extant species, widely distributed 
throughout the Indo-Pacific (Mortensen 1928; H.L. 
Clark 1946; A.M. Clark and Rowe 1971). In particular, 
the species has been recorded from the Torres Strait 
by H.L. Clark (1921), with a single specimen collected 
from Low Isles reef on the Great Barrier Reef, making 
it “one of the rarest of Australian echini’ (H.L. Clark 
1946). In spite of the species’ widespread distribution, 
specimens are not common (H.L. Clark 1946). The 
species is restricted to coral reefs and many have been 
found by dredging on coral sand (Mortensen 1928). 

P. verticillata, with the ‘very peculiar character 
of primary spines...may well claim to be the easiest 
recognizable of all species of Cidarids’ (Mortensen 
1928). The species, with its distinct spines, stands alone 
in genus Prionocidaris (see key of A.M. Clark and 
Rowe 1971), so much so that Mortensen (1928: 428) 
was adamant that it should represent the type of a 
separate genus. Prionocidaris verticillata, or forms 
closely related to the species, have been found as fossil 
from the Miocene of Madagascar and India, Pliocene 
of Ceram (Indonesia), and the Quaternary of East 
Africa (Mortensen 1928). 


Material 

One isolated primary spine ANU 60589 from 
locality 8280-4, northwest of Aru’re village, east coast 
of Yule Island, Central Province, PNG. Kairuku 
Formation, Lower Pliocene. 


Subclass EUECHINOIDEA Bronn, 1860 
Superorder ECHINACEA Claus, 1876 
Order TEMNOPLEUROIDA Mortensen, 1942 


Remarks 

The temnopleuroid Temnotrema macleayana 
(Tenison-Woods) is present in the Kairuku Formation 
on Yule Island and was redescribed by Lindley (2001). 


Family TOXOPNEUSTIDAE Troschel, 1872 
Genus CYRTECHINUS Mortensen, 1943 


Proc. Linn. Soc. N.S.W., 124, 2003 


Type species 
Psammechinus verruculatus Liitken, 1864, p. 
98, by original designation. 


Remarks 

Cyrtechinus Mortensen, 1943 and the closely 
related Nudechinus H.L. Clark, 1912 and Gymnechinus 
Mortensen, 1903, are distinguished by the number of 
plates in the buccal membrane. Accordingly, it is very 
difficult or even impossible to distinguish between 
naked tests of species of these taxa. All are small forms 
found in the tropical Indo-Pacific and western Indian 
Ocean (A.M. Clark and Rowe 1971: 142, 143) and, as 
with the temnopleurids, appear to be well adapted to 
life in shallow-water tropical regions, especially in the 
case of Nudechinus which has developed into a 
considerable number of species (Mortensen 1943). All 
three taxa have been noted from the Torres Strait and 
surrounding waters (Mortensen 1943; H.L. Clark 1946; 
A.M. Clark and Rowe 1971). The Yule Island 
specimens are assigned to Cyrtechinus because of the 
striking similarity of both ambulacral and 
interambulacral plating diagrams with those of 
Hawaiian specimens of the only species, Cyrtechinus 
verruculatus (Liitken) figured by Mortensen (1943: 
Figs 245a-b). 


Cyrtechinus verruculatus (Liitken) 
Figs 4a-g, 5a-b 


Synonymy 

Echinus (Psammechinus) verruculatus Litken, 
1864, p. 98. 

Echinus verruculatus Liitken, 1864: Sluiter 
1899, p. 110. 

Lytechinus verruculatus H.L. Clark, 1912: H.L. 
Clark 1921, p. 147; H.L. Clark 1946, p. 321. 

Cyrtechinus verruculatus (Liitken): Mortensen 
1943, p. 393; A.H. Clark 1952, p. 267; A.M. 
Clark and Rowe 1971, p. 142, 143. 

Cyrtacanthus verruculatus (Liitken): A.M. Clark 
and Rowe 1971, p. 156. 

Stomechinid (?), Lindley 2001, p. 126. 

T. Mortensen (1943), A Monograph of the 
Echinoidea 3(2), Camarodonta 1, p. 393, and 
A.M. Clark and F.W.E. Rowe (1971), 
Monograph of Shallow-Water Indo-West 
Pacific Echinoderms, p. 142, 143, details the 
previous synonymies. 


Description 

Test small, very regularly hemispherical; 
distinctly subpentagonal circumference in the largest 
specimens. Ambitus relatively low; distinctly sunken 
towards the peristome. It appears not to exceed a size 


143 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


Figure 4. Cyrtechinus verruculatus (Liitken). Lower Pliocene, Yule Island, Central Province. 4a-b, ANU 
60576, aboral view. Bar scale = 2.5 mm; portion of apical disc, showing periproct and small tubercles 
adapical to genital pores. Bar scale = 0.5 mm. 4c-f, UPNG F1184, oral, lateral views. Bar scale = 5 mm; 
apical disc with madreporite top centre, posterior to lower right. Bar scale = 1 mm; ambulacral plating at 
ambitus (refer to Fig. 5b for plating diagram). Bar scale = 0.5 mm. 4g, ANU 60572, interambulacral 
plating at ambitus (refer to Fig. 5a for plating diagram). Bar scale = 1 mm. 


144 Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Figure 5. Cyrtechinus verruculatus (Liitken). Lower Pliocene, Yule Island, Central Province. 5a-b, plating 
diagrams at ambitus for interambulacrum, ambulacrum. 


of c. 24 mm horizontal diameter (Table 1). Dimensions 
are comparable with those presented by Mortensen 
(1943) for the species, ranging from 10.5 to 24 mm 
horizontal diameter, with a single specimen of 27 mm 
considered by him to be very old. Both ambulacra and 
interambulacra densely covered with tubercles. 

Apical system small with a central periproct. 
In smaller specimens (ANU 60576) Ocular I is insert; 
in larger specimens (UPNG F1184) both Oculars I and 
V are narrowly insert. The same relationships were 
described by Mortensen (1943) in Recent specimens 
of the species from the central Pacific and East African 
coast. Tubercles on apical plates increase in size with 
age, with older specimens having a single large tubercle 
located adapically to genital pore; young specimens 
(ANU 60576) have three small, equi-sized tubercles 
adapical to the genital pore. 

Ambulacra at ambitus are about 2/3 width of 
interambulacra. Ambulacral plates compound, 
trigeminate, pore pairs in arcs of three. A prominent 
elevated primary tubercle is present in the middle of 
each plate, forming a vertical series. Tubercles 


imperforate, noncrenulate, as large as interambulacral 
primaries. Large secondary tubercles present 
admesially to the primary ambulacral tubercles, but 
do not form prominent vertical series. The number and 
arrangement of tubercles generally remains the same 
in various sized specimens. Interambulacral plates are 
about equal in height to opposite ambulacral plate. 
Each possesses a sub-central prominent imperforate, 
noncrenulate primary tubercle; each primary tubercle 
is surrounded by a semi-circular series of variously 
sized secondaries. In larger specimens (ANU 60572) 
there is a tendency for the development of weak ridging 
radiating from the primary tubercle towards 
secondaries, forming a sculpturing reminiscent of the 
temnopleurids. 

Peristome large, about 1/3-1/2 horizontal 
diameter of test; twice size of apical system in larger 
specimens (Table 1). Gill slits are distinct and small. 
Tubercles of the oral surface, particularly surrounding 
the peristome, with a distinctly enlarged, elevated boss 
compared with those of the apical surface. 


TABLE 1. Cyrtechinus verruculatus (Liitken): Test dimensions in millimetres. 


Specimen Diameter 
F1184 24 
60578 eS 
60579 ZA, 
60577 19 
60572 19 
60582 18.5 
60580 18 
60583 17 
60581 IOS 
60576 16 
60573 14.5 
60575 14 
60574 11 


Proc. Linn. Soc. N.S.W., 124, 2003 


Peristome 
10 


Apical system 


145 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


Remarks 

Cyrtechinus verruculatus (Liitken) is a Recent 
species from the tropical Indo-West Pacific (Mortensen 
1943; A.M. Clark and Rowe 1971). In particular, it 
has been recorded from the Java Sea, Sulu Sea and 
Torres Strait (Mortensen 1943). The fossil record of 
C. verruculatus is problematic given the uncertainty 
in distinguishing naked tests from those of the closely 
allied Nudechinus H.L. Clark, 1912, and Gymnechinus 
Mortensen, 1903 (Mortensen 1943: 398, 399; A.M. 
Clark and Rowe 1971). Fossil specimens from the 
Pleistocene deposits of Egypt have been referred to 
Nudechinus scotiopremnus H. L. Clark, 1912, a species 
common in the Red Sea (Mortensen 1943). Specimens 
possibly referrable to C. verruculatus have been 
described from the East African coast, from the 


Pliocene of Mombasa Island and from Mozambique 
(Mortensen 1943). However, it is suffice to say that 
forms identical with C. verruculatus or an ancestor of 
the species were present in the western Indian Ocean 
during the Plio-Pleistocene. 

Cyrtechinus is a small form typically restricted 
to the littoral zone. The low hemispherical test with a 
low down ambitus, was interpreted by Lindley (2001) 
as an adaptive strategy, giving stability in currents on 
either rocky or sandy substrates. Closely allied forms 
such as Lytechinus and Tripneustes are dominant 
consumers of live seagrass leaves in tropical seagrass 
communities (Nojima and Mukai 1985; Klumpp et al. 
1993). Cyrtechinus verruculatus is interpreted as a 
seagrass grazer in meadows constantly swept by 
currents. 


Figure 6. Schizechinus cf. tuberculatus (Pomel). Lower Pliocene, Yule Island, Central Province. 6a-c, ANU 
60600, lateral view with peristome to top. Bar scale = 5 mm; ambulacral plating at ambitus (refer to Fig. 
7a for plating diagram). Bar scale = 1 mm; portion of peristome showing deep gill slits. Bar scale = 2 mm. 


146 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Figure 7. Schizechinus cf. tuberculatus (Pomel). Lower Pliocene, Yule Island, Central Province. 7a-b, plating 
diagrams at ambitus for ambulacrum, interambulacrum. 


Material 

Fourteen complete tests: ANU 60572-73, ANU 
60575-83 from locality 8280-3; ANU 60574, ANU 
60587 from locality 8280-4; and UPNG F1184 
collected by R. Perembo from locality 24 of Francis et 
al. (1982) = locality 8280-3. All localities are northwest 
of Aru’re village, east coast of Yule Island, Central 
Province, PNG. Kairuku Formation, Lower Pliocene. 


Genus SCHIZECHINUS Pomel, 1869 


Synonymy 
Toxophyma Lambert and Thiéry, 1925, p. 280. 


Type species 
Anapesus tuberculatus Pomel, 1887, p. 298, by 
original designation. 


Remarks 

The referral of the Yule Island form to Echinus 
cf. stracheyi by F. Chapman and I. Crepsin (in 
Montgomery 1930) is doubtful. The presence of a 
primary tubercle on every ambulacral plate clearly 
distinguishes the Yule Island species from the echinid 
Echinus Linné, 1758, in which primaries are present 
only on every alternate (or every third) ambulacral plate 
(Fell and Pawson 1966). 

The Yule Island species can be confidently 
assigned to Schizechinus Pomel, 1869 using 


Proc. Linn. Soc. N.S.W., 124, 2003 


Mortensen’s (1943) key for the Family 
Toxopneustidae. Schizechinus is the only large 
toxopneustid with trigeminate ambulacral plates each 
bearing primary and secondary tubercles arranged in 
regular parallel series (Mortensen 1943; Fell and 
Pawson 1966). Lytechinus A. Agassiz, 1863 is another 
toxopneustid with a similar test, but differs from 
Schizechinus in that the secondary tubercles do not 
form a regular series. Mortensen (1943) regards 
Schizechinus a near relation of Lytechinus. 
Schizechinus is known only from fossils, from the Mio- 
Pliocene of North Africa (Malta and Algeria) and 
Europe. 


Schizechinus cf. tuberculatus (Pomel) 
Figs 6a-c, 7a-b 


Synonymy 
Schizechinus tuberculatus Pomel, 1869, p. XLII. 
Echinus cf. stracheyi: F. Chapman and I. Crespin 
in Montgomery 1930, p. 57; Lower Pliocene, 
Yule Island. 


Description 

No complete test is available. However, the 
portion of test preserved in ANU 60600 suggests a 
large test, c. 60 mm diameter, comparable with forms 
figured by Mortensen (1943). Test apparently of high 
hemispherical shape, inferred from shape of ambital 
region preserved in ANU 60601. Ambital outline 
inferred to be circular, relatively low; oral surface 


147 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


Figure 8. Parasalenia poehli Pfeffer, 1887. Lower Pliocene, Yule Island, Central Province. 8a-d, ANU 
60584, aboral, lateral views. Bar scale = 5 mm; ambulacral plating at ambitus (refer to Fig. 9a for plating 
diagram). Bar scale = 2 mm; apical disc, with posterior to lower left. Bar scale = 2 mm. 


flattened and distinctly sunken towards peristome 
(ANU 60600). Details of apical system unknown. 
Ambulacra at ambitus are about 1/2 width of 
interambulacra. Ambulacral plates compound, 
trigeminate pore pairs in arcs of three. A prominent 
elevated imperforate, noncrenulate primary tubercle, 
about the same size as interambulacral ones, is present 
in the middle of each plate, forming a vertical series. 
Large secondary tubercles are also present on each 
plate, forming a regular perradial series parallel with 
the primary series. A varying number of randomly 
placed small tubercles cover the plates. 
Interambulacral plates at ambitus are about 
equal in height and twice the width of opposite 
ambulacral plate. Each possesses a sub-central 
imperforate, noncrenulate primary tubercle, forming 
a regular vertical series. Each primary is flanked by 
two secondaries, also forming regular series parallel 
with the primary series. Varying numbers of randomly 
placed small tubercles cover the remainder to each 


148 


plate. 
Peristome sunken, c. 10 mm diameter in ANU 
60600; gill slits are distinct, deep. 


Remarks 

This large regular echinoid is represented only 
by fragmentary material, indicative of the poor 
preservation potential of such fragile forms (Kier 
1977). However, the Yule Island material is tentatively 
referred to Schizechinus tuberculatus (Pomel), figured 
by Mortensen (1943: Fig. 291a,b) and Fell and Pawson 
(1966: Fig. 320, 3b,c), on the basis of similarity of (a) 
ambulacral and interambulacral plating in the ambital 
region, and (b) inferred test shape. Schizechinus 
tuberculatus (Pomel) is a Miocene echinoid from 
Algeria. The Yule Island occurrence is the first record 
of the subtropical Schizechinus in the southern 
hemisphere. 

Schizechinus also includes another 12 species, 
predominantly from the Miocene and Pliocene of the 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


oO 


b 


Figure 9. Parasalenia poehli Pfeffer, 1887. Lower Pliocene, Yule Island, Central Province. 9a-b, plating 
diagrams at ambitus for ambulacrum, interambulacrum. 


Mediterranean region and Europe (Mortensen 1943). 
Mortensen (1943) figured two of these species, viz: 
Schizechinus angulosus (Pomel) from the Miocene of 
Algeria and Schizechinus duciei (Wright) from the 
Miocene of Malta. Test shape is an obvious difference 
between figured species. Little information on the 
remaining ten species is presently available to the writer 
and the Yule Island form may well represent a new 
species. 


Material 

Three fragmentary tests: ANU 60600 is a 
preserved portion of the oral surface, with the margin 
of the peristome and gill slits; ANU 60601 is a portion 
of an interambulacrum at the ambitus, with some of 
the oral surface; ANU 60602 is a portion of the ambital 
surface. ANU 60600-01 were collected from locality 
8280-1; ANU 60602 from locality 8280-3. All localities 
are from the east coast of Yule Island, Central Province, 
PNG. Kairuku Formation, Lower Pliocene. 


Order ECHINOIDA Claus, 1876 
Family PARASALENIIDAE Mortensen, 1903 
Genus PARASALENIA A. Agassiz, 1863 


Synonymy 
Cladosalenia A. Agassiz, 1872, p. 148. 


Type species 
Parasalenia gratiosa A. Agassiz, 1863, p. 22. 


Parasalenia poehli Pfeffer, 1887 
Figs 8a-d, 9a-b 


Synonymy 
Parasalenia pohlii Pfeffer, 1887, p. 110: H.L. 
Clark, 1912, p. 369; Lambert and Thiéry, 
1914, p. 269; H.L. Clark, 1922, p. 142; HLL. 
Clark, 1925; H.L. Clark, 1928, p. 469; 
Mortensen, 1940, p. 49; Mortensen, 1943, p. 


Proc. Linn. Soc. N.S.W., 124, 2003 


272; H.L. Clark, 1946, p. 331. 
Parasalenia gratiosa de Meijere, 1904, p. 98. 
Parasalenia poehli: A.M. Clark and Spencer 
Davies, 1966, p. 599, 603; A.M. Clark and 
Rowe, 1971, p. 142, 157. 


Description 

Test small, distinctly elongate and low, ANU 
60585 measuring 8 x 7.5 x 4.5 mm and ANU 60584 
20 x 18 x 9 mm. Dimensions are comparable with those 
of Mortensen (1943), who noted a range in test length 
x height from 8 x 3 mm to 17 x 6 mm, and HL. Clark 
(1928, 1946), who noted a single specimen from the 
northern Great Barrier Reef 16 x 13 x 7 mm. Details 
of oral surface unknown. Mortensen (1943) noted that 
the test is not very strong, and is easily broken. 

Apical system is dicyclic, 8 mm x 6.5 mm, 
elongate in the plane of long axis of test. Diameter of 
periproct is about 1/4 of the long diameter of the apical 
system. The madreporite is limited in size, with pores 
only occupying about 1/3 of plate. Genital plates are 
smooth and are without tubercles. Oculars are widely 
exsert and small. 

Ambulacra at ambitus match the width of 
interambulacra. Ambulacral plates compound, 
trigeminate, pore pairs in arcs of three. Pore zones very 
narrow. A large primary tubercle occupies much of 
plate, leaving little space for development of secondary 
tubercles and granules. Primary tubercles are 
imperforate and non-crenulate; the mamelon is 
relatively large and areole indistinct. Secondary 
tubercles and granules are either absent or number at 
most one or two per plate. Primary tubercles on the 2- 
3 uppermost ambulacral plates are very much 
diminished in size. Interambulacral plates are about 
equal in height to opposite ambulacral plate. Primary 
interambulacral tubercles very large, leaving only a 
restricted adradial space for 2-3 secondary tubercles. 
Secondary tubercles do not form regular horizontal or 
vertical series. 


149 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


Details of peristome unknown. 


Remarks 

Parasalenia A. Agassiz, 1863 includes two 
Recent species, Parasalenia gratiosa A. Agassiz, 1863 
and Parasalenia poehli Pfeffer, 1887, with an Indo- 
Pacific and Red Sea distribution, and a single fossil 
species, Parasalenia fontannesi Cotteau, 1913, from 
the Lower Miocene of France (Mortensen 1943; Fell 
and Pawson 1966). H.L. Clark (1928, 1946) noted a 
single specimen of P. poehli from the northern Great 
Barrier Reef, representing the only documented 
occurrence of the species in Australian waters. 

There has been considerable discussion 
regarding the validity of poehli as a species (Mortensen 
1943; H.L. Clark 1946). Parasalenia poehli is a much 
smaller form, with the largest recorded being 17 mm 
long, half that of gratiosa (Mortensen 1943). Both 
Mortensen (1943) and H.L. Clark (1946) agree that 
periproct size is important in distinguishing the two 
forms. The periproct of poehli is relatively smaller, 
about 1/4 of the long diameter of the apical system; 
this compares with 1/3-1/2 that length in gratiosa. H.L. 
Clark (1946) regarded the presence or absence of 
tubercles on genital plates as an important diagnostic 
character, but Mortensen (1943) believed this character 
to be ‘quite unreliable’. Both Mortensen (1943) and 
H.L. Clark (1946) observed that the genital plates of 
gratiosa carry tubercles, but not in poehili. In addition 
to periproct size, Mortensen (1943) emphasised the 
size of primary tubercles, with poehli having smaller 
primary tubercles, with more space for secondary 
tubercles, than gratiosa. A.M. Clark and Rowe (1971) 
recognised both poehli and gratiosa as valid species, 
but using differing characters to those of Mortensen 
(1943) and H.L. Clark (1946). They distinguished 
species using, amongst other characters, the length of 
primary spines relative to test length. In the case of 
gratiosa spines are about equal to test length, whereas 
for poehli, primary spines are only about half as long 
as test length. Spine length in the parasaleniids is 
expected to be proportionate to primary tubercle size, 
and this latter character, as used by Mortensen (1943), 
is therefore clearly useful in distinguishing naked tests. 

Parasalenia occupies generally well concealed 
habitats, among the branches of corals or hidden in 
crevices and cavities beneath coral rock (H.L. Clark 
1946). Mortensen (1943) noted a bathymetric range 
extending from littoral to c. 70 m. 
Material 

Two tests: ANU 60584 from locality 8280-3; 
ANU 60585 from locality 8280-4. Both localities are 
northwest of Aru’re village, east coast of Yule Island, 
Central Province, PNG. Kairuku Formation, Lower 
Pliocene. 


150 


ACKNOWLEDGMENTS 


The specimens described in this paper were collected 
by the writer during fieldwork on Yule Island in January 
2002. For the efficient execution of this fieldwork, the writer 
gratefully acknowledges the assistance and hospitality of Sr. 
Elizabeth of the Bishop’s Office, Diocese of Bereina, Port 
Moresby and the OLSH Sisters at the Yule Island Mission. 
Alphonese Aisi and Ben and Nahau Roama of Yule Island, 
with Daniel Salamas of Port Moresby, all helped during 
fieldwork. Photography of specimens was completed by Dr. 
Roger Heady of the Scanning Electron Microscopy Unit, 
ANU, and Dr. R.E. Barwick. Parts of the manuscript were 
read by Prof. K.S.W. Campbell. Dr. Ulrike Troitzsch is kindly 
acknowledged for her translation from German of parts of 
Jeannet and R. Martin (1937). This work was completed 
while the writer was a Visiting Fellow in the Department of 
Geology, ANU, and Dr. Patrick De Deckker, Head of 
Department, is thanked for the provision of departmental 
facilities. 


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Department of Zoology, University of Queensland 


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Department of Zoology, University of Queensland 
1, 53-60. 

Endean, R. (1956). Queensland faunistic records - 4. Further 
records of Echinodermata (excluding Crinoidea). 
Papers of Department of Zoology, University of 
Queensland 1, 123-140. 

Endean, R. (1957). The biogeography of Queensland’s 
shallow-water echinoderm fauna (excluding 
Crinoidea), with a rearrangement of the faunistic 
provinces of tropical Australia. Australian Journal 
of Marine and Freshwater Research 8, 233-273. 

Fell, H.B. (1966). Cidaroids. In “Treatise on Invertebrate 
Paleontology, Part U, Echinodermata 3’ (Ed. R.C. 
Moore) pp. U312-U340. (Geological Society of 
America and University of Kansas Press: 
Lawrence). 

Fell, H.B. and Pawson, D.L. (1966). Echinacea. In “Treatise 
on Invertebrate Paleontology, Part U, 
Echinodermata 3’ (Ed. R.C. Moore) pp. U367- 
U440. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Gibbs, P.E., Clark, A.M. and Clark, C.M. (1976). 
Echinoderms from the northern region of the Great 
Barrier Reef, Australia. Bulletin of the British 
Museum Natural History (Zoology) 30(4), 103- 
144. 

Glynn, P.W., Wellington, G.M. and Birkeland, C. (1979). 
Coral reef growth in the Galapagos: Limitation by 
sea urchins. Science 203, 47-49. 

Hatcher, B.G. (1983). Grazing in coral reef ecosystems. In 
‘Perspectives on Coral Reefs’ (Ed. D.J. Barnes) 
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Heck, K.L. and McCoy, E.D. (1979). Biogeography of 
seagrasses: evidence from associated organisms. 
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128. 

Kier, P.M. (1977). The poor fossil record of the regular 
echinoid. Palaeobiology 3, 168-174. 

Klumpp, D.W., Salita-Espinosa, J.T. and Fortes, M.D. (1993). 
Feeding ecology and trophic role of sea urchins in 
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McPherson, B.F. (1965). Contributions to the biology of the 
sea urchin Tripneustes ventricosus. Bulletin of 
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McPherson, B.F. (1968). Contributions to the biology of the 
urchin Eucidaris tribuloides (Lamarck). Bulletin 
of Marine Science 18, 400-433. 

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Mortensen, T. (1936). Phyllacanthus forcipulatus, sp. nov., 
a new cidarid from the Indian Ocean. Records of 
the Indian Museum, Calcutta 38, 307-309. 

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151 


LOWER PLIOCENE ECHINOIDS (REGULARIA) FROM PAPUA NEW GUINEA 


152 Proc. Linn. Soc. N.S.W., 124, 2003 


Echinoids of the Kairuku Formation (Lower Pliocene), 
Yule Island, Papua New Guinea: Spatangoida 


J.D. LINDLEY 


Department of Geology, Australian National University, Canberra, A.C.T. 0200 


Lindley, 1.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea: 
Spatangoida. Proceedings of the Linnean Society of New South Wales 124, 153-162. 


Spatangoid echinoids are well represented in the rich and diverse echinoid fauna of the Lower Plicoene 
Kairuku Formation, Yule Island, Papua New Guinea. Five taxa are recognised, including the schizasterid 
Schizaster (Schizaster) alphonsei sp. nov., the palaeostomatid Palaeostoma kairukuensis sp. nov., the brissid 
Eupatagus (Eupatagus) pulchellus (Herklots) and the spatangids Maretia planulata (Lamarck) and Maretia 
cordata Mortensen, 1948. The spatangoids, by comparison with the clypeasteroids and regularia, exhibit a 


high degree of endemism. 


Manuscript received 18 November 2002, accepted for publication 11 December 2002. 


KEYWORDS: Echinoidea, Spatangoida, Schizaster, Palaeostoma, Eupatagus, Maretia, Lower Pliocene, 


Papua New Guinea. 


INTRODUCTION 


This paper, describing spatangoid or burrowing 
echinoids, is the fourth in a series (Lindley 2001, 
2003a, 2003b) on the rich and diverse echinoid fauna 
of the Kairuku Formation, Yule Island, Central 
Province, Papua New Guinea (PNG). Two new species 
are described and the spatangoids, by comparison with 
the clypeasteroids (Lindley 2003a) and regularia 
(Lindley 2003b), exhibit a high degree of endemism. 
The present descriptions are based on collections made 
by the writer in January 2002, and the reader is referred 
to Lindley (2003a) for locality details. Some specimens 
have been temporarily allocated Department of 
Geology, Australian National University repository 
numbers, pending their repatriation to PNG at the 
conclusion of studies, where they will be housed in 
the Department of Geology, University of Papua New 
Guinea, Port Moresby. The classification used herein 
follows that of Fischer (1966) and A.M. Clark and 
Rowe (1971). 


SYSTEMATIC PALAEONTOLOGY 


Class ECHINOIDEA Leske, 1778 
Subclass EUECHINOIDEA Bronn, 1860 
Superorder ATELOSTOMATA Zittel, 1879 
Order SPATANGOIDA Claus, 1876 


Suborder HEMIASTERINA Fischer, 1966 
Family SCHIZASTERIDAE Lambert, 1905 
Genus SCHIZASTER L. Agassiz, 1836 


Type species 
Schizaster studeri Agassiz, 1836, by subse- 
quent designation of ICZN, 1948. 


Diagnosis 

Test small to large with anterior ambulacrum 
and frontal notch shallow to deep. Apical system 
located posterior of centre, with 2-4 genital pores. 
Anterior ambulacrum shallow to deep with pore pairs 
oblique or transverse and arranged in single or 
irregular, double rows. Anterior petals long and almost 
straight although at times distally flexed; at least twice 
the length of the posterior petals. Both peripetalous 
and lateroanal fascioles present and complete 
(McNamara and Philip 1980a, 1980b). 


Subgenus SCHIZASTER L. Agassiz, 1836 


Type species 
See above. 


Diagnosis 

Moderate to large species of Schizaster with 
deep anterior notch. Anterior ambulacrum deep and 
long; pore pairs almost transversely oriented and in 


LOWER PLIOCENE ECHINOIDS (SPATANGOIDA) FROM PAPUA NEW GUINEA 


single rows. Anterior petals long, deep and flexed; 
diverging at a low angle usually less than 80° 
(McNamara and Philip 1980a). 


Schizaster (Schizaster) alphonsei sp. nov. 
Figs la-c 


Synonymy 
Brisaster latifrons (A. Agassiz, 1898), Lindley 
2001, p. 135. 


Diagnosis 

A small species of Schizaster with a relatively 
low test; apical system is 54 percent of test length from 
anterior. Anterior ambulacrum broad posteriorly, 
narrowing anteriorly, very deep, partially overhung by 
its sides, with the adjoining interambulacra forming 
narrow vertical keels; anterolateral ambulacra also 
incised, with sides partially overhung; moderately deep 
frontal sinus. 


Etymology 
Named for Alphonse Aisi of Yule Island, Central 
Province, PNG. 


Description 

Test of small size, elongate oval, only specimen 
with length x width x height measuring c. 36.5 x 30 x 
17 mm. Test rather low, vertex located 54 percent of 
test length from anterior; the test slopes very gradually 
towards the anterior end; details of posterior end 
unknown. Oral surface gently convex. 

Apical system located at vertex, slightly 
posterior of centre; number of genital pores unknown. 
Frontal ambulacrum broad posteriorly, but becomes 
noticeably narrower anteriorly; very deeply sunken, 
with sides partially overhung. Adjoining 
interambulacra forming narrow vertical keels. Frontal 
ambulacrum passes to a moderately deep frontal notch. 
Pore pairs arranged in a single row. Anterior petals 
sunken with sides partially overhung; petals are gently 
curved. Posterior petals short, closed and sunken. 
Posterior interambulacrum forming a prominent keel. 
Details of periproct unknown. 

Peristome close to anterior end of test, shallowly 
sunken, with the sunken peristomal region continuing 
directly to frontal notch. Peripetalous fasciole well 
developed; a lateral fasciole passes posteriorly to meet 
with an anal fasciole that is partially preserved in F 
1179. Both oral and aboral surfaces with a uniform 
covering of tubercles. 


Remarks 


Schizaster (Schizaster) alphonsei sp. nov. can 
be distingusihed from Schizaster excavatus Jeannet and 


154 


R. Martin, 1937, Middle Pliocene, Java, by its shorter 
frontal ambulacrum and anteriorly positioned apical 
system (54 percent of test length from anterior 
compared with 65 percent, respectively). Jeannet and 
R. Martin (1937) described two specimens of S. 
excavatus, with length x width x height measuring 47 
x 36 x 24 mm and 56 x 45 x 30 mm. This species and 
S. (S.) alphonsei both have low tests and a very deep 
frontal ambulacrum, wide at its posterior end, narrower 
anteriorly. Schizaster (Schizaster) alphonsei is readily 
distinguished from Schizaster (Schizaster) aff. 
compactus Koehler, 1914, described from the Middle 
Miocene of Barrow Island, northwestern Australia, by 
McNamara and Kendrick (1984), by the latter’s 
possession of a smaller (length of 30 mm), globose 
test and shallow frontal notch. Schizaster (Schizaster) 
sphenoides Hall, 1907, redescribed by McNamara and 
Philip (1980a) from the Middle Miocene of Victoria, 
is similar to S. (S.) alphonsei, with a very deep frontal 
ambulacrum with overhung sides. However, this 
temperate water species can be distinguished from S. 
(S.) alphonsei by its large, subcircular test. 

McNamara and Philip (1980a) described the 
progressive morphological changes in schizasterid 
echinoids from the Palaeocene to Recent, reflecting 
adaptation to the occupation of new ecological niches, 
and noted the following general trends: 

(a) the posterior migration of the apex and apical 
system, allowing more water to flow over the anterior 
edge of the test toward the peristome; 

(b) an increase in declination of the anterior 
slope, also increasing the flow of water to the anterior; 

(c) lengthening of the test, allowing a further 
posterior migration of the apex and apical system; and 

(d) lengthening and deepening of the frontal 
ambulacrum and the anterior notch, assisting in 
channelling water to the peristome. 

McNamara and Philip (1980a) believed that 
these morphological adaptations in Schizaster were 
related to the need to enhance the current flow over 
the aboral surface of the test in a sediment of low 
permeability. These changes were considered by 
McNamara and Philip (1980a) to reflect adaptation to 
deeper burrowing and the occupation of finer 
sediments and were at their ‘extreme’ with the 
Schizaster morphotype during the Miocene. 

The Lower Pliocene S.(S.) alphonsei is an 
intermediate form transitional between McNamara and 
Philip’s (1980a) Paraster form, a sand-dwelling 
echinoid, and the Schizaster form, a mud-dwelling 
form. The echinoid’s small test size, with an apical 
system slightly posterior of centre, short frontal 
ambulacrum and gentle anterior slope from the apex 
are typical of Paraster Pomel, 1869. A very deep frontal . 
ambulacrum with a deep frontal notch is typical of the 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Figure 1. Schizaster (Schizaster) alphonsei sp. nov. Lower Pliocene, Yule Island, Central Province. 1a-c, 
UPNG F1179, aboral, lateral and oral views. Bar scale = 1.0 cm. Palaeostoma kairukuensis sp. nov. 
Lower Pliocene, Yule Island, Central Province. 1d-f, UPNG F1186, aboral, lateral and posterior views. 
Bar scale = 0.5 cm; 1g, UPNG F1186, detail of apical system with elevated processes separating pore 
pairs, visible along the margins of the frontal ambulacrum (refer to Fig. 2 for an interpretation of 
frontal ambulacrum). Bar scale = 0.25 cm; 1h, UPNG F1186, view of pentangular peristome. Bar scale 
= 0.5 cm; 1i, ANU 60635, view of pentangular peristome. Bar scale = 0.5 cm. 


Proc. Linn. Soc. N.S.W., 124, 2003 155 


LOWER PLIOCENE ECHINOIDS (SPATANGOIDA) FROM PAPUA NEW GUINEA 


Schizaster morphotype. 


Material 

Holotype UPNG F1179, a near complete test 
collected by R. Perembo from locality 24 of Francis et 
al. (1982) = locality 8280-3. Northwest of Aru’re 
village, east coast of Yule Island, Central Province, 
PNG. Kairuku Formation, Lower Pliocene. 


Family PALAEOSTOMATIDAE Lovén, 1867 


Remarks 

Family Palaeostomatidae differs from the 
Hemiasteridae in possessing a pentangular, rather than 
labiate, peristome (Mortensen 1950; Fischer 1966). 


Genus PALAEOSTOMA Loven in A. Agassiz, 1872 


Synonymy 
Leskia Gray, 1851, p. 134. 
Skouraster Lambert, 1937, p. 89. 


Type species 
Leskia mirabilis Gray, 1851, p. 184. 


Emended diagnosis 

Test small, ovoid, inflated. Apical system 
central, or nearly so, 2 genital pores; paired ambulacra 
broadly petaloid, closed distally; frontal ambulacrum 
non-petaloid with pores arranged in a single radial row. 


Remarks 

The emended diagnosis for Palaeostoma is 
broadened from those presented by Mortensen (1950) 
and Fischer (1966), to include the Lower Pliocene 
species from Yule Island, strikingly similar in all 
characters except pore shape and placement in the 
frontal ambulacrum. 


Palaeostoma kairukuensis sp. nov. 
Figs 1d-i, 2 


Synonymy 
Hemiaster sp., F. Chapman in Mayo et al. 
1930; F. Chapman and I. Crespin in 
Montgomery 1930, p. 57; Lower Pliocene, 
Yule Island. 
Ditremaster sp., Lindley 2001, p. 133; Lower 
Pliocene, Yule Island. 


Diagnosis 

A relatively large species of Palaeostoma with 
a little marked frontal ambulacrum; pores positioned 
adapically, arranged in a single radial row, 6-7 pore 


156 


pairs per row; pores small, circular, each pore pair 
separated by an elevated process of rectilinear shape. 


Etymology 

Named after Kairuku, the former Government 
Station and present village, southeast coast of Yule 
Island, Central Province, PNG. 


Description 

Test small, ovoid outline and high vaulted; the 
largest F1185, a slightly deformed test, with a length 
x width x height measuring 25 x 20 x c. 11 mm and 
the smallest ANU 60635 c. 14 x 11 x 8 mm. Vertex 
subcentral, about 52-60 percent of test length from 
anterior; test slopes both posteriorly and anteriorly 
from vertex, with an abrupt steepening near margins. 
Oral surface is convex. 

Apical system located at vertex, ethmophract 
with madreporite not separating posterior oculars; two 
genital pores on top of conical elevations. Frontal 
ambulacrum non-petaloid, sunken posteriorly but flush 
and little marked anteriorly; frontal sinus very shallow. 
Pores of frontal ambulacrum are intra-fasciolar, 
arranged in a single radial row, 6-7 pore pairs per row; 
pores small, circular. Each pore pair oriented obliquely 
to line of radial row, pointing perradially distally; 
separated by an elevated process of rectilinear shape 
(Fig. 2). Ornament of frontal ambulacrum consists of 
sparse, irregular granules confined to margins of 
ambulacrum. Paired ambulacra straight, broad, closed 
distally, and sunken; posterior petals about 1/3 size of 
anterior ones. Pores are slit-like, equi-sized; adjacent 
pore-pairs are separated by 4-5 secondary tubercles. 
Interporiferous zone, narrow, naked. 

Interambulacral areas within the peripetalous 
fasciole with a covering of fine primary tubercles with 
prominent bosses, set in a dense covering of granules; 
a particularly dense clustering of primary tubercles is 
present in interambulacral areas flanking the pore zone 
of the frontal ambulacrum. Otherwise, plates of aboral 
and oral surface with a scattered covering of primary 
tubercles with prominent bosses and granules, 
increasing in density towards anterior end of test. 

Peristome distinctly pentangular, clearly evident 
in F1186 and ANU 60635. Periproct circular, located 
at upper end of curved posterior. Peripetalous fasciole 
very well marked on F1186; no other fascioles. 


Remarks 

Palaeostoma Lovén in A. Agassiz, 1872 is a 
locally common extant genus of the Indo-Pacific 
(Mortensen 1950; A.M. Clark and Rowe 1971). 
Palaeostoma mirabile (Gray), regarded as a primitive 
spatangoid, is particularly common in the Java Sea and: 
is known to occur in the Philippine islands, south Japan, 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


distal 


adapical 


Figure 2. Palaeostoma kairukuensis sp. nov. Lower 
Pliocene, Yule Island, Central Province. Anterior 
ambulacrum (Amb III) showing pores arranged in 
a single radial row and pore pairs separated by an 
elevated process. Plate boundaries are 
interpretative. 


the Indian Ocean, South Pacific islands and the Red 
Sea (Mortensen 1950; A.M. Clark and Rowe 1971). 
The maximum test size recorded by Mortensen (1950) 
was 18 mm length. Palaeostoma mirabile lives in mud 
and has a bathymetric distribution of c. 20-110 m 
(Mortensen 1950). Only two fossil species have been 
recorded, Palaeostoma zitteli de Loriol, 1881 and 
Palaeostoma rochi Lambert, 1937, from the Eocene 
of Egypt and Morocco, respectively (Mortensen 1950). 
Both Eocene forms have distinctive frontal depressions 
and are clearly not related to the Lower Pliocene 
species from Yule Island. 

Eocene species of Palaeostoma and P. mirabile 
are distinguished by their possession of a frontal 


Proc. Linn. Soc. N.S.W., 124, 2003 


ambulacrum with pores arranged in a single radial row, 
the distal one of each pair comma-shaped. In the frontal 
ambulacrum of P. kairukuensis sp. nov. (F1186, 
measuring 20 mm length), each radial row possesses 
6-7 pore pairs, each pore being small and circular. 
Although Mortensen (1950) and Fischer (1966) did 
not state the number of pore pairs in the frontal 
ambulacrum of P. mirabile, an illustrated specimen in 
Mortensen (1950: Plate V, Fig. 4; reproduced in Fischer 
1966: Fig. 451) indicates 8-9 pore pairs in each radial 
row. This specimen measures 15 mm length. The 
frontal ambulacrum of P. kairukuensis is flush at the 
ambitus, indicating that the entire structure was 
unlikely to be involved in food gathering and 
transportation (Smith 1984). The pores of the frontal 


-ambulacrum are all intra-fasciolar, located adapically 


in a sunken region of the ambulacrum high on the 
dorsal side of the test. A dense cluster of primary 
tubercles is present in the flanking interambulacra. 

The presence of pore pairs in the adapical region 
of the frontal ambulacrum, as opposed to the ambital 
region, indicate that funnel-building tube feet were 
present (Nichols 1959; Smith 1984). Normally, each 
pore of a pair gives passage to a tube foot connected 
internally to its ampulla, and the more perradial pore 
is grooved to house a branch of the radial nerve (Fischer 
1966; Smith 1984). A rimmed area surrounding each 
pore is an attachment area for the stem retractor muscle, 
and the width of this area gives a measure of the 
thickness of the stem retractor muscle and hence an 
idea of the strength of the tube foot (Smith 1984: 
41).These tube feet, typical of the dorsal region of 
burrowing spatangoids, were primarily engaged in 
building and maintaining a long open respiratory 
funnel to the sediment/water interface (Nichols 1959; 
Durham 1966). The prominent elevated process 
separating pore pairs in the frontal ambulacrum of P 
kairukuensis (Fig. 2) may have been an attachment 
structure for the stem retractor muscle, its small surface 
area interpreted to indicate tube feet of limited length. 
Nichols (1959: Figs 41 and 45) figured and described 
a similar process between pore pairs of ambulacra I 
and II of Micraster coranguinum Leske, 1778, 
interpreting it as a muscle attachment feature. 

Palaeostoma kairukuensis, with its poorly 
developed frontal ambularcral pore pairs, was probably 
a shallower burrower than P. mirabile. The relatively 
dense covering of aboral tubercles and the presence of 
a well developed peripetalous fasciole on the test of 
the Yule Island species also indicate not only an 
infaunal mode of life, but also burrowing in fine sands 
and muds (Smith 1984). 


Material 
Holotype UPNG F1186, a near complete test; 


157 


LOWER PLIOCENE ECHINOIDS (SPATANGOIDA) FROM PAPUA NEW GUINEA 


and paratypes UPNG F1185, a deformed test, collected 
by R. Perembo from locality 24 of Francis et al. (1982) 
= locality 8280-3; and ANU 60635-36, two worn 
specimens, from locality 8280-3. Locality 8280-3 is 
northwest of Aru’re village, east coast of Yule Island, 
Central Province, PNG. Kairuku Formation, Lower 
Pliocene. 


Suborder MICRASTERINA Fischer, 1966 
Family BRISSIDAE Gray, 1855 
Genus EUPATAGUS L. Agassiz, 1847 


Synonymy 
Pseudopatagus Pomel, 1885, p. 18. 
Melitia Fourtau, 1913, p. 68. 
Heterospatangus Fourtau, 1905, p. 606. 
Euspatangus Cotteau, 1869, p. 257. 
Perispatangus Fourtau, 1905, p. 605. 
Koilospatangus Lambert, 1906, p. 185. 
Zanolettiaster Sanchez Roig, 1952, p. 14. 


Megapatagus Sanchez Roig, 1952, p. 58. 


Type species 
Eupatagus valenciennesi L. Agassiz, 1847, by 
subsequent designation of Pomel, 1883, p. 


28; Recent, Australia. 


Diagnosis 

Test ovoid in outline, low, oral side flat; apical 
system anterior, ethmolytic, with 4 genital pores; paired 
ambulacra with closed petals; frontal ambulacrum 
nonpetaloid, pores in single series; primary tubercles 
on aboral side only within peripetalous fasciole 
(Fischer 1966). 


Subgenus EUPATAGUS L. Agassiz, 1847 


Type species 
See above 


Diagnosis 
Ambitus rounded, frontal sinus weak or 
absent (Fischer 1966). 


Eupatagus (Eupatagus) pulchellus (Herklots) 
Figs 3a-e 


Synonymy 
Spatangus pulchellus Herklots, 1854, p. 12; 
Miocene, Java. 
Hemipatagus pulchellus, Desor, 1858, p. 418. 
Maretia? pulchella, K. Martin 1880, p. 5; 
Gerth 1922, p. 512; Miocene, Java. 
Brissoides (Brissoides) pulchellus, Lambert 


158 


and Thiéry 1924, p. 451. 

Maretia pulchella, Lambert and Thiéry 1924, 
p.451 

Eupatagus pulchella, F. Chapman and I. 
Crespin in Montgomery 1930, p. 57; Lower 
Pliocene, Yule Island. 

Eupatagus (Brissoides) pulchellus (Herklots), 
Jeannet and R. Martin 1937, p. 273; 
Miocene, Java. 

Eupatagus pulchellus (Herklots), Mortensen 


1951, p. 472, 473; Miocene, Java. 


Description 

Test of small size, elongate oval, the largest 
ANU 60603 measuring 40 x 34.5 x 18 mm, and the 
smallest, ANU 60616, 27 x 25 x 14 mm. Test low, 
vertex located centrally; test slopes very gently towards 
anterior end, with an abrupt steepening near margin; 
test slopes gently towards posteriorly, to an abrupt 
vertical termination. Oral surface is gently convex. 

Apical system located anteriorly of vertex, at 
about 37 percent of test length from anterior; four 
genital pores. Frontal ambulacrum non-petaloid, 
narrow, not sunken, with a very shallow frontal sinus. 
Paired ambulacra closed petaloid, weakly sunken. 
Outer and inner pores of pore pairs are equi-sized; 
consecutive pore pairs are separated by a rounded 
transverse ridge and, adapically, a shallow transverse 
depression. Interporiferous zone covered by randomly 
arranged secondary tubercles and numerous miliary 
tubercles. 

Posterior interambulacrum V is developed into 
a prominent keel with irregularly arranged secondary 
tubercles and numerous miliary tubercles. The paired 
interambulacra have 4-7 large, perforate, crenulate 
tubercles within the peripetalous fasciole. Otherwise, 
the plates of the aboral side with dense covering of 
secondary and miliary tubercles. 

Peristome is large, transversely elliptical, 
without a prominent lip. Periproct is pear-shaped, 
situated high up on the vertical posterior end of test, 
with the point of it just visible in aboral view. Plastron 
is distinctly inflated particularly in juvenile specimens 
(ANU 60616), which bear a high, medial keel. 

Peripetalous fasciole is difficult to recognise in 
many specimens; however, in ANU 60608 fasciole is 
distinct, passing around adoral extremity of petals of 
paired ambulacra, before disappearing towards frontal 
ambulacrum. Subanal fasciole present. 


Remarks 

Eupatagus (Eupatagus) pulchellus (Herklots) 
is locally very common in the lower to middle Kairuku 
Kormation. The species appears to be identical in all 
respects to the specimen figured by Jeannet and R. 


Proc. Linn. Soc. N.S.W., 124, 2003 


I.D. LINDLEY 


Figure 3. Eupatagus (Eupatagus) pulchellus (Herklots). Lower Pliocene, Yule Island, Central Province. 
3a-d, ANU 60603, aboral, oral, lateral and posterior views. Bar scale = 1.0 cm; 3e, ANU 60607, oral view. 


Bar scale = 1.0 cm. 


Martin (1937: Figs 50a, b) from the Miocene of Java. 
The largest of the four specimens described by Jeannet 
and R. Martin (1937) has dimensions of 40.9 x 35.3 x 
23.0 mm and the smallest 33.5 x 29.0 x >20 mm. The 
difficulty in proving the presence of a peripetalous 
fasciole in the specimens described and figured by 
Herklots (1854) led to uncertainty in referral to 
Eupatagus L. Agassiz, 1847 (Mortensen 1951). 
However, Jeannet and R. Martin (1937) proved the 
existence of a peripetalous fasciole and Mortensen 
(1951) believed there was little doubt that the East 
Indies form is an Eupatagus. 

Mortensen (1951: 472) considered E. (E.) 
pulchellus to be a near relation of the extant Eupatagus 
(Eupatagus) rubellus Mortensen, 1948. The only 
known specimen of this species was collected near 
Tinakta Island of the Tawi Tawi group (5° 12’N; 119° 
55’E), Sulu Archipelago, Philippines, at a depth of 24 
m (Mortensen 1948b). Many examples of E. (E.) 
pulchellus, preserved in a range of orientations, were 
collected from the middle Kairuku Formation at 


Proc. Linn. Soc. N.S.W., 124, 2003 


locality 8280-4, north of Aru’re village on the east coast 
of Yule Island. This shallow-water sequence was 
interpreted by Lindley (2003a) to have suffered 
wholesale disruption and redistribution of sediment 
resulting from a succession of large storm events. 


Material 

Twenty-nine complete tests including: ANU 
60608, 60611 from locality 8280-2; ANU 60612 from 
locality 8280-3; and ANU 60603, 60607, 60609-10, 
60613-34 from locality 8280-4. All localities are on 
the east coast of Yule Island, Central Province, PNG. 
Kairuku Formation, Lower Pliocene. 


Family SPATANGIDAE Gray, 1855 
Genus MARETIA Gray, 1855 


Synonymy 


Hemipatagus Desor, 1858, p. 416. 
Tuberaster Peron and Gauthier, 1885, p. 46. 


159 


LOWER PLIOCENE ECHINOIDS (SPATANGOIDA) FROM PAPUA NEW GUINEA 


Thrichoproctus A. Agassiz (M.S. nom. nud.). 
Plagiopatagus Liitken (in litteris, nom. nud.). 


Type species 
Spatangus planulatus Lamarck, 1816, p. 326, 


by original designation; Recent. 


Maretia planulata (Lamarck) 
Fig. 4d 


Synonymy 

Spatangus planulatus Lamarck, 1816, p. 326. 

Spatangus praelongus Herklots, 1854, p. 11; 
Miocene, Java. 

Spatangus affinis Herklots, 1854, p. 10; 
Miocene, Java. 

Maretia planulata Gray, 1855, p. 48; Tenison- 
Woods 1878. p. 173: Recent, Australia; 
Tenison-Woods 1881, p. 204: Recent, 
Australia; Gerth 1922, p. 512: Pliocene, 
Java; Jeannet and R. Martin 1937, p. 277: 
Miocene, Java; Mortensen 1951, p. 21; 
A.M. Clark and Rowe 1971, p. 146: 
Recent, northern Australia; Gibbs et al. 
1976, p. 135: Recent, Low Isles, northern 
Great Barrier Reef; De Ridder 1986, p. 48: 
Recent, New Caledonia. 

Maretia ?planulata, K. Martin 1885, p. 286: 
Miocene, Java. 

Maretia ovata, H.L. Clark 1932, p. 277; H.L. 
Clark 1946, p. 380: Recent, northern Great 
Barrier Reef. 

T. Mortensen (1951), A Monograph of the 
Echinoidea 5(2), Spatangoida II, p. 27-29, 
lists the previous synonymies. 


Description 

Test of moderate size, very low, only specimen 
measuring 44 x c. 50 x c. 10 mm, length:height in the 
range of 3-4:1 of A.M. Clark and Rowe (1971); low 
arched above, with moderately sharp edges; outline 
elongate oval. Anterior notch apparently lacking, 
although test is broken in this region. Oral side is gently 
concave about a medial line passing posterially through 
frontal ambulacrum. 

Details of apical system unknown. Frontal 
ambulacrum flush and indistinct. Pore pairs are 
indistinct. Anterior petals flush, straight and closed 
distally; interporifeous zone is broad, conspicously 
raised, densly covered with fine miliary tubercles and 
occasional irregularly placed secondary tubercles. 
Details of posterior petals unknown. 

Details of periproct and peristome unknown. 
Details of subanal fasciole unknown. The aboral 


160 


surface between paired petals and frontal ambulacrum 
with large primary tubercles arranged in horizontal 
series. Tubercles are perforate, crenulate, with 
moderately sunken aureoles. Anterolateral margins of 
test with dense covering of secondary tubercles. Oral 
surface with uniform covering of distinctively shaped 
tubercles, best described (and figured) by Mortensen 
(1951; 33 and Fig. 14), with ‘the boss forming a screw, 
and the aureole unequally deepened, the whole 
structure almost resembling an ear’. 


Remarks 

ANU 60606, an incomplete test lacking details 
of the posterior, peristome and apical system, is 
assigned to Maretia planulata (Lamarck) on the basis 
of test shape, tubercle arrangement on the aboral 
surface and the distinctive shape of tubercles on the 
oral surface. As fossil, M. planulata has been recorded 
from the Pliocene of the Red Sea region and Zanzibar, 
and the Mio-Pliocene of Java (Mortensen 1951; Jeannet 
and R. Martin 1937). Extant forms of the species are 
widely distributed throughout the tropical-subtropical 
Indo-West Pacific, from East Africa (Mozambique, 
Madagascar), northern Australia, China and southern 
Japan, and Fiji and the Gilbert Islands (Mortensen 
1951; A.M. Clark and Rowe 1971). On the Australian 
coast the species occurs from Cooktown southward to 
Port Jackson, but not from within, or to the west of, 
the Torres Strait region (H.L. Clark 1946). The 
spatangoid lives buried in muddy sand within the ebb 
zone and often comes to the sand surface during 
exposure on day-time spring tides (Mortensen 1951; 
Gibbs et al. 1976). Mortensen (1951) records the 
species to a depth of c. 60m. 


Material 

ANU 60606, a fragmentary test including left 
anterior petal and frontal ambulacrum, from locality 
8280-1, south of Tete ne’ina Beach, east coast of Yule 
Island, Central Province, PNG. Kairuku Formation, 
Lower Pliocene. 


Maretia cordata Mortensen, 1948 
Figs 4a-c 


Synonymy 
Maretia cordata Mortensen, 1948, p. 132; 


Mortensen 1951, p. 41: Recent, East Indies. 


Description 

Test small, distinctly heart-shaped, aboral side 
low arched, not flattened. ANU 60604, slightly 
deformed at its right-anterior end, has a length x width 
x height of 27 x 26 x 10 mm, similar to the largest of 
four specimens described by Mortensen (1951), 


Proc. Linn. Soc. N.S.W., 124, 2003 


LD. LINDLEY 


Figure 4. Maretia cordata Mortensen, 1948. Lower Pliocene, Yule Island, Central Province. 
4a-b, ANU 60604, aboral view and detail of apical system showing anterior series of the 
right anterior petal (Amb II) with upper 4-5 plates with rudimentary pores or none. Bar 
scales = 1.0 and 0.5 cm respectively; 4c, ANU 60605, aboral view of incomplete specimen, 
with left anterior petal (Amb IV) visible. Bar scale = 1.0 cm. Maretia planulata (Lamarck). 
Lower Pliocene, Yule Island, Central Province. 4d, ANU 60606, aboral view of incomplete 
test, with left anterior petal (Amb IV) visible. Bar scale = 1.0 cm. 


measuring 28 x 28 x 10 mm; ANU 60605, an 
incomplete specimen, is slightly larger than 
Mortensen’s (1951) specimens with a width x height 
of 29 x 14 mm. Test slopes very gently both anteriorly 
and posteriorly from apical system, with an abrupt 
steepening near margins. Details of oral surface 
unknown. 

Apical system located at about 44 percent of 
test length from anterior; number of genital pores 
unknown. Frontal ambulacrum distinctly sunken, pores 


Proc. Linn. Soc. N.S.W., 124, 2003 


pairs indistinct, passing to a conspicuous frontal notch. 
Petals flush, closed distally. Anterior series of the 
anterior petals with upper 4-5 plates with rudimentary 
pores or none. Interporiferous zone is flat, with a sparse 
covering of miliary and some secondary tubercles. 
Details of periproct and subanal fasciole unknown. 
Posterior interambulacrum V is raised forming 
a low keel that slightly overhangs the periproct; 
ornamentation consists of a sparse covering of fine 
miliary tubercles and some secondary tubercles. The 


161 


LOWER PLIOCENE ECHINOIDS (SPATANGOIDA) FROM PAPUA NEW GUINEA 


paired interambulacra each with 8-10 perforate, 
crenulate primary tubercles set amongst a sparse 
covering of miliary tubercles and occasional secondary 
tubercles; placement of primary tubercles is random 
with a tendency toward arrangement in horizontal 
series near the ambitus. 

Details of peristome unknown. 


Remarks 

Maretia cordata Mortensen, 1948 was erected 
by Mortensen (1948b) to distinguish tests with a 
distinctly different shape, deeper frontal depression and 
broader petals than M. planulata. He recorded the 
species from Palawan Island, Philippines, and the Bali 
Sea and Flores Sea, Indonesia, at bathymetric ranges 
of 50-150 m. ANU 60604 and 60605 from the Pliocene 
of Yule Island, are referable to VM. cordata on the basis 
of test size and shape, the presence of a distinct frontal 
depression and sinus, paired petals distinctly broader 
than M. planulata, and the nature of the anterior series 
of the anterior petals. Mortensen (1951) noted that large 
specimens of M. cordata carry some large tubercles in 
the posterior ambulacrum, but the Yule Island specimen 
ANU 60604, equal in size to the largest described by 
him, does not have primary tubercles in the posterior 
ambulacrum. 

Mortensen (1951: 44) noted that Hemipatagus 
bandaensis R. Martin (in Jeannet and R. Martin 1937) 
from the (?) Pliocene of Banda, Indonesia, is a probable 
ancestor of M. cordata. The Pliocene record of M. 
cordata from Yule Island makes this unlikely. 
Hemipatagus bandaensis does not possess a raised keel 
in the posterior interambulacrum, and the posterior 
aboral surface curves gradually from the vertex towards 
margin, contrasting with the flattened-gently sloping 
surface of M. cordata. 


Material 

Two tests: ANU 60605, an incomplete test 
lacking posterior details from locality 8280-1, south 
of Tete ne’ina Beach, and ANU 60604, a complete, 
though slightly deformed test, from locality 8280-3, 
northwest of Aru’re village, east coast of Yule Island, 
Central Province, PNG. Kairuku Formation, Lower 
Pliocene. 


ACKNOWLEDGMENTS 


Most of the specimens described in this paper were collected 
by the writer during fieldwork on Yule Island in January 
2002. For the efficient execution of this fieldwork, the writer 
gratefully acknowledges the assistance and hospitality of Sr. 
Elizabeth of the Bishop’s Office, Diocese of Bereina, Port 
Moresby and the OLSH Sisters at the Yule Island Mission. 
Alphonse Aisi and Ben and Nahau Roama of Yule Island, 


162 


with Daniel Salamas of Port Moresby, all helped during 
fieldwork. Photography of specimens was completed by Dr. 
R.E. Barwick. Some species determinations were discussed 
with Prof. K.S.W. Campbell. This work was completed while 
the writer was a Visiting Fellow in the Department of 
Geology, ANU, and Dr. P. De Deckker, Head of Department, 
is thanked for the provision of departmental facilities. 


REFERENCES 
[Additional to those listed in Lindley (2003a, 
2003b)] 


De Ridder, C. (1986). Les échinides. In ‘Guide des étoiles 
de mer, oursins et autres échinodermes du lagon de 
Nouvelle-Calédonie’ (Eds A. Guille, P. Laboute and 
J.-L. Menou) pp. 22-53. (Institut Francais de 
Recherche Scientifique pour le Développement en 
Coopération. Collection Faune Tropiclae 25). 

Durham, J.W. (1966). Ecology and palaeoecology. In 
“Treatise on Invertebrate Paleontology, Part U, 
Echinodermata 3’ (Ed. R.C. Moore) pp. U257-U265. 
(Geological Society of America and University of 
Kansas Press: Lawrence). 

Fischer, A.G. (1966). Spatangoids. In “Treatise on 
Invertebrate Paleontology, Part U, Echinodermata 3’ 
(Ed. R.C. Moore) pp. U543-U628. (Geological 
Society of America and University of Kansas Press: 
Lawrence). 

Herklots, J.A. (1854). “Fossiles de Java’. Echinodermes. 

Lindley, I.D. (2003a). Echinoids of the Kairuku Formation 
(Lower Pliocene), Yule Island, Papua New Guinea: 
Clypeasteroida. Proceedings of the Linnean Society 
of New South Wales, 124, 125-136. 

Lindley, I.D. (2003b). Echinoids of the Kairuku Formation 
(Lower Pliocene), Yule Island, Papua New Guinea: 
Regularia. Proceedings of the Linnean Society of New 
South Wales, 124, 137-152. 

McNamara, K.J. and Philip, G.M. (1980a). Australian 
Tertiary schizasterid echinoids. Alcheringa 4, 47-65. 

McNamara, K.J. and Philip, G.M. (1980b). Living Australian 
schizasterid echinoids. Proceedings of the Linnean 
Society of New South Wales, 104, 127-146. 

Mortensen, T. (1950). A Monograph of the Echinoidea 5(1), 
Spatangoida I. C.A. Reitzel, Copenhagen. 422p. 

Mortensen, T. (1951). A Monograph of the Echinoidea 5(2), 
Spatangoida II. C.A. Reitzel, Copenhagen. 593p. 

Nichols, D. (1959). Changes in the chalk heart-urchin 
Micraster interpreted in relation to living forms. 
Philosophical Transactions of the Royal Society of 
London, Series B, 242, 347-437. 

Smith, A. (1984). ‘Echinoid Palaeobiology’. (George Allen 
and Unwin: London). 190p. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Holocene Foraminifera from Tuross Estuary and Coila Lake, 
South Coast, New South Wales: A Preliminary Study 


LUKE STROTZ 


Centre for Ecostratigraphy and Palaeobiology, Macquarie University, Sydney, NSW, 2109 


Strotz, L. (2003). Holocene Foraminifera from Tuross Estuary and Coila Lake, south coast, New South 
Wales: A preliminary study. Proceedings of the Linnean Society of New South Wales 124, 163-182. 


Two estuaries on the New South Wales south coast, Tuross Estuary and Coila Lake, were sampled for 
Foraminifera. Thirty-seven taxa were identified from surface samples but only those requiring extensive 
taxonomic revision are discussed. The species composition of the total assemblage at each of the sample 
sites was analysed and the reasons for species distribution explored. A new species, Fissurina breviductus 


sp. noy., is described. 


Manuscript received 13 October 2002, accepted for publication 11 December 2002. 


KEYWORDS: Coastal; Estuary; Foraminifera; Holocene; New South Wales 


INTRODUCTION 


Tuross Estuary and Coila Lake are adjacent 
estuaries situated on the New South Wales south coast 
(Fig. 1). Despite their close proximity to each other, 
the two estuaries differ markedly in geomorphological 
character and depositional history. Tuross Estuary is a 
convoluted, complex estuary, characterised by 
numerous sandbars, variable depth and is open to the 
Pacific Ocean via a narrow channel. Coila Lake, in 
contrast, is a relatively simple, shallow estuary, 
periodically closed off from the ocean by a large barrier 
beach and is therefore more correctly defined as a saline 
coastal lake. Both are drowned river valleys, filled 
with sediments of Holocene age (Roy and Peat, 1976). 

A study of the modern foraminiferal faunas 
present in the two estuaries was undertaken in an 
attempt to define the environmental parameters that 
control the distribution of the assemblages that inhabit 
the two estuaries and to note differences in assemblage 
composition in a marine influenced estuarine 
environment (Tuross Estuary) in comparison to a 
recently closed lagoonal environment (Coila Lake). 
Tuross Estuary and Colia Lake were chosen for this 
study because of their close proximity to each other 
and the limited amount of anthropogenic activity in 
the surrounding area. 


PHYSICAL CHARACTERISTICS OF STUDY 
AREA 


Tuross Estuary 

Tuross Estuary is the larger and more complex 
of the two estuaries with a total water area of 12.95 
km? (Ozestuaries database, 2002). It is classified as a 
barrier estuary system (Roy, 1984) and a narrow 
channel at the north end of a large sand spit is the only 
link between the estuary and the open ocean. The 
estuary is considered to be one of the least modified 
estuaries on the New South Wales coastline (Brierley 
et al., 1995), due mainly to the limited amount of 
anthropogenic activity in the area. 

The sedimentological environments that 
dominate the estuary have been comprehensively 
documented by Roy and Peat (1976) and Brierley et 
al. (1995). According to these two studies, Tuross 
Estuary can be divided into four major depositional 
environments with the differences in each of these 
environments due to differences in the source of the 
sediment and the energy of the sedimentation processes 
present in each environment. Deposition in the estuary 
is attributed to a variety of mechanisms, including tide, 
wave and fluvial processes (Brierley et al., 1995; Fig. 
1). 


Coila Lake 

Coila Lake is classified as a saline coastal 
lake, since it is usually cut off from the ocean by a 
barrier beach (Roy and Peat, 1976). Total water area 
of the lake is 6.85 km? (Ozestuaries database, 2002), 
making it one of the largest coastal lakes on the New 
South Wales coastline (Roy, 1984). Only one major 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


Figure 1. Map of Tuross Estuary and Coila Lake (modified from Tibby, 1996). 


tributary, Coila Creek, enters the lake. Freshwater 
discharge from the creek is low and does not normally 
possess enough energy to force a channel through the 
barrier beach located along its southern margin (Roy 
and Peat, 1976). The lake has however been open to 
the sea on a number of occasions in recent years. In 
the period between 1975 and 1999 the barrier beach 
has been breached four times due to natural events 
(such as major flooding) and on thirteen occasions by 
mechanical means in order to lower water levels in the 
lake (Coila Lake Estuary Management Committee, 
2001). 

Bathymetrically, Coila Lake is flat bottomed 
with moderately steep sides. Maximum depth is 
approximately four metres (Roy and Peat, 1976). Much 
of the bottom sediment consists of dark coloured sandy 
muds, deposited by low energy tidal and wind induced 
currents (Roy and Peat, 1976). 


METHODS 


Material was described from a total of ten 


164 


sample sites (see Fig. 1; TL1, TL3, TL4, TL5, TL7 
and TL8 in Tuross Estuary and CL1, CL2, CL3, and 
CL4 in Coila Lake). Each site was examined for 
foraminifera. The ten sites were chosen because they 
form a transect across the two estuaries and best 
exemplify the full gamut of sedimentary environments 
present in the two estuaries, based upon previous 
studies of the estuaries by Roy and Peat (1976) and 
Brierley et al. (1995). Surface sediment was collected 
at each of the sample sites and was immediately treated 
with ethanol to perserve any live specimens for later 
staining. Upon return to the laboratory, each sample 
was treated with rose Bengal, using the method 
described by Bell (1996). Samples were then washed 
through a 1 mm and 63 mm sieve. dried at room 
temperature and split into 10 gram aliquots for picking. 

Water chemistry analysis was undertaken 
with the use of Hydrolab 4 datasonde. Only the results 
obtained for conductivity, directly analgous to salinity, 
obtained at each locality will be discussed herein as 
the greater portion of this work will form the basis of 
a later paper (Hostetler et al. in prep). 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


DISTRIBUTION OF MODERN 
FORAMINIFERAL FAUNAS 


Thirty-seven species of foraminifera were 
identified from surface samples collected in the two 
estuaries. In terms of species composition, the 
foraminiferal fauna present in Tuross Estuary and Coila 
Lake is similar to other estuaries on the New South 
Wales south coast (Yassini and Jones, 1989; Yassini 
and Jones, 1995; Cotter, 1996). Results of staining with 
rose Bengal revealed only minor variation between live 
and dead assemblages. Because of this, all analysis is 
based upon total assemblages, rather than only the live 
assemblage. Of the 37 species identified, thirty four 
are benthic species and the remaining three planktonic. 
All three planktonic species, Globigerina bulloides 
d’Orbigny, Neogloboquadrina pachyderma 
(Ehrenberg) and Pulleniatina obliquiloculata (Parker 
and Jones), are confined to sample site TL1, suggesting 
that marine influence over sedimentation and water 
movement in Tuross Estuary does not extend beyond 
Tuross Lake (Fig. 1). 

Three main foraminiferal assemblages were 
identified in the two estuaries. These are the Lower 
Estuary Assemblage, the Upper Estuary Assemblage 
and the Coastal Lake Assemblage. Assemblages were 
differentiated based upon the faunal composition of 
the assemblage and the relative abundances of the 
various taxa that make up the assemblage. Besides a 
few notable exceptions, dicussed more extensively 
below, the distribution of these assemblages is 
unsurprising, with the composition of the foraminiferal 
faunas at each locality conforming to what would be 
expected. 

The most diverse fauna is the Lower Estuary 
Assemblage, found at localities TL1 and CL2. The 
fauna is dominated by rotaliine forms; taxa such as 
Rosalina australis (Parr), Ammonia aoteana (Finlay), 
E. crispum crispum (Linne), E. advenum advenum 
(Cushman) and Parrellina papillosa (Cushman) having 
the highest relative abundances. An assemblage of this 
type is indicative of open estuary conditions where 
normal marine salinities dominate (Hayward et al., 
1999). This assemblage is expected at locality TL1 
considering its location (Fig. 1) and the conductivity 
results obtained for the site, approximately 57 mS/cm. 
Fully marine conditions return conductivity values 
around 60 mS/cm. Its presence at CL2 however is 
surprising as the locality is not currently subject to open 
estuary conditions, cut off from the open ocean by a 
barrier beach, and conductivity levels obtained for the 
site were brackish in nature, around 43 mS/cm. The 
site has been exposed to fully marine conditions in the 
past when the barrier beach has been opened by either 
natural or unnatural causes (Coila Lake estuary 


Proc. Linn. Soc. N.S.W., 124, 2003 


management committee, 2001). It is therefore proposed 
that the assemblage found at this site is a relict fauna 
associated with a time when the lake was open to the 
ocean. This assertion is supported by the lack of “live” 
specimens obtained at the site, determined using rose 
Bengal stain on the collected samples, suggesting that 
a living foraminiferal fauna does not currently inhabit 
the locality and the noticeably abraded nature of the 
material, suggesting deposition occurred some time 
ago. 

The remaining sample localities in Tuross 
Estuary, excluding TL4, possess a fauna that is herein 
designated the Upper Estuary assemblage. This 
assemblage is characterised by a mixture of 
agglutinated and calcareous taxa; including 


_ Scherochorella barwonensis (Collins), Ammobaculites 


exiguus Cushman and Bronniman, Quinqueloculina 
oblonga (Montagu), Elphidium advenum advenum 
(Cushman), Elphidium excavatum clavatum Cushman 
and Elphidium lene Cushman and McCulloch. This 
fauna is indicative of shallow sub-tidal middle estuary 
conditions, with salinity levels slightly below normal 
marine conditions (Hayward et al., 1999). This 
assemblage is found at localities TL3, TL5, TL7 and 
TL8, sites characterised by either mangroves or reed 
beds. The conductivity values obtained for these sites 
were between 53-56 mS/cm, only slightly below 
normal marine values. The presence of this fauna at 
locality TL8 indicates that even the extremities of 
Tuross Estuary are subject to moderately saline 
conditions (Fig. 1). 

The Coastal Lake assemblage dominates at 
the three remaining sample sites in Coila Lake, CL1, 
CL3 and CL4 and is also present at locality TL4. This 
assemblage consists of predominantly agglutinated 
taxa, in most cases dominated by the species 
Ammobaculites exiguus Cushman and Bronnimann and 
Scherochorella barwonensis (Collins). In areas where 
high amounts of aqueous vegetation are present, such 
as marsh settings, Trochammina inflata (Montagu) is 
also a major constituent of the fauna. This fauna is 
indicative of a shallow sub-tidal situation where 
brackish conditions dominate (Collins, 1974; Hayward 
et al., 1999). Conductivity levels for these four sites 
do not exceed 45 mS/cm. The presence of this 
assemblage in Coila Lake is unsurprising, considering 
the lake is cut off from the open ocean but its presence 
at locality TL4 is unexpected. Located is the eastern 
part of Tuross Estuary, close to the estuary mouth, the 
foraminiferal fauna at the site should be comparable 
to the Lower Estuary Assemblage found nearby at TL1. 
This is not the case, with the fauna made up mainly of 
two taxa, Ammobaculites exiguus Cushman and 
Bronnimann and Scherochorella barwonensis 
(Collins), which make up to 91% of the total 


165 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


abundance; the remaining 9% is composed of minor 
agglutinated forms and rare calcareous species. The 
difference in the faunal assemblage found at locality 
TL4 to that found in other parts of Tuross Estuary is 
related to the presence of a large sandbar, which isolates 
the site from the main part of the estuary. This isolation 
has created an environment, which, in terms of both 
water chemistry and sedimentary environment, is more 
comparable to a slightly brackish lagoonal situation, 
and a faunal assemblage that reflects these conditions. 


DISCUSSION 


Salinity is the main controlling factor on the 
distribution of foraminiferal species in Tuross Estuary 
and Coila Lake. Sites that are subject to, or have been 
subject to, fully marine conditions, such as TL1 and 
CL2 , possess a fauna that is relatively high in diversity 
and characterised by numerous calcareous taxa. 
Hayward et al. (1999) noted that localities of this kind, 
where normal marine conditions are prevalent and 
there is little tidal exposure, are generally the most 
diverse localities in any particular estuary. Conversely, 
sites in the upper, or northern, part of Coila Lake, where 
brackish conditions dominate, are characterised by a 
low diversity fauna that is composed of agglutinated 
taxa. Those areas in Tuross Estuary where salinity is 
variable, in the middle and upper parts of the Tuross 
Estuary, possess a fauna which lies somewhere 
between the previous two and is a mixture of calcareous 
and agglutinated taxa. 


SYSTEMATIC DESCRIPTIONS 


Unless otherwise stated, the higher level 
classification follows Loeblich and Tappan (1987). 
Although revised classification schemes have been 
published (e.g. Loeblich and Tappan, 1992; Sen Gupta, 
1999), Loeblich and Tappan’s (1987) scheme is 
considered less controversial and since it is in 
widespread use, allows comparisons to be made 
between this study and others. The only departure that 
has been made from Loeblich and Tappan (1987) is 
the use of -OIDEA rather than -ACEA as the ending 
of all superfamily names, following recommendation 
29A of the ICZN 4" edition. 

No attempt has been made to describe those 
taxa that have been comprehensively monographed in 
other systematic studies (Albani, 1968a; Albani 1979; 
Hayward et al., 1997; Hayward et al., 1999). Only new 
taxa or those requiring substantial revision are 
described in detail. A full list of species recovered from 


166 


the two estuaries can be found in Table 1. 


Order FORAMINIFERIDA Eichwald, 1830 
Suborder TEXTULARIINA Delage and Herouard, 
1896 
Superfamily HORMOSINOIDEA Haeckel, 1894 
Family HORMOSINIDAE Haeckel, 1894 
Subfamily REOPHACINAE Cushman, 1910 


Scherochorella Loeblich and Tappan, 1984 


Type species: 
Reophax minuta Tappan, 1940 


Scherochorella barwonensis (Collins), 1974 
(Plate 1, Figs. 1-3) 


1974 Reophax barwonensis Collins, p. 8; Pl. 1, fig 1. 

1980 Reophax barwonensis Apthorpe, Pl. 29, Fig. 7. 

1989 Protoschista findens Yassini and Jones, Figs. 
10.10, 10.11. 

1992 Reophax barwonensis Bell and Drury, p. 12; 
Fig. 4.5. 

1995 Reophax barwonensis Bell, p. 229; Fig. 2.1. 

1995 Protoschista findens Yassini and Jones, p. 69; 
Figs. 39, 43. 

1996 Reophax barwonensis Bell, p. 5; Pl. la. 


Description: 
see Collins (1974), p. 8. 


Remarks: 

The designation of this species has been highly 
contentious. Much of the confusion associated with 
this taxon has arisen due to its variable morphology. 
Specimens found in Tuross Estuary and Coila Lake 
display variability in the total number of chambers 
present, the coarseness of the test and the size of the 
proloculus relative to the subsequent chambers (Plate 
1, Figs. 1-2). This variation could be explained by the 
presence of both microspheric and megalospheric 
forms (Bell, 2002 pers. comm), although this 
possibility was not mentioned by Collins (1974) in his 
original description of the species. 

I have followed Loeblich and Tappan (1984) 
in assigning this species to the genus Scherochorella 
based upon its subglobular proloculus, appressed 
chambers and depressed sutures. This assignment has 
been previously rejected by Bell and Drury (1992) who 
state that the specimens illustrated as part of their study 
(Bell and Drury 1992, Fig 4.5) do not exhibit a flattened 
test, nor could they be considered tiny; both diagnostic 
features of the genus Scherochorella (Loeblich and 
Tappan, 1984). However, as test size cannot be 
considered a good generic character in foraminifera, 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


Table 1. Species recovered from Tuross Estuary and Coila Lake. X indicates the localities at which each 


species occurs. 


ep) 
xe) 
© 
Q. 
© 
” 
Fa 
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= 
0) 


Reophax barwonensis 
eptohalysis collinsi 
Ammobaculites barownensis 


[id 


as often it is environmentally controlled, and the 
flattened nature of the type species could be due to 
burial distortion (Loeblich and Tappan, 1987), Bell and 
Drury’s (1992) assertion is herein rejected. 
Hayward and Hollis (1994), in their 
assessment of New Zealand brackish water 
foraminifera, asserted that S. barwonensis Collins is a 
junior synonym of Reophax moniliforme Siddall. Bell 


Proc. Linn. Soc. N.S.W., 124, 2003 


Ammobaculites barownensis 

Ammobaculites exiguus | |s(X(XX[X| XI XT | [XTX] | |X| x] x] x, 
Miliamminafusca | | SX OTT XY XT TX xixt TT 
Portatrochamminasorosa |X| X|X|X[X[X|x{x1 | [ [x] [TT TT TT | 
Trochamminainfiata | S| SIXTX] | OX XT TT xt TT tT x] x] 
Eggerellasubconica | «| SIX] | | | | | | XT | TT tT Ix] 
Comuspirainvolvens | SXIXE | TT TT TT TE TT TT 
Spiroloculinacarinata = IX] | | TdT TdT Td TdT TdT ET TT TT tT 
Quinqueloculina oblonga |XX |X| X |X] [XI X[xIxt | | | xix? | 
Quingueloculinapoeyana | (|X| | | | IX] | | | dT dT TE TT TT 
Quinqueloculina sp.ct.Q.dispariis | |X| | | | | | | | | | | | | | Tt TT 
Triloculina giadius sp.nov. |X| | | | dT dT dT dT dT dT TUT ET ET 
Lenina SEER 
lLagenablomaeformis | | | dT TCT TCT TTT Xt TT TT tT 
Fissurina breviductus sp.nov. __—|_“‘{X| {| | {| | | | | | | TT TT | tt I 
Guttulinairreguiaris XIX] | | TdT TT TT TE TT TE ET 
Eoiynetsiiati ees SSUES eae a 
Enis. a ae ae ee ee eee eee 
Trochulinadimidiatus XIX} | | TT TT TT TT ET TET 
fiposainaausel same Lie XL ae 
Pileolina austalensis, IX} | | | dT | dT TdT TP TT TE TE TT TT 
Cibicides dispars XTX] | OT | | TT | TT | TT xt 
Ammonia aoteana IX XIXIXIXIXIX] TT [Xt | TT | TT 
Elphidium advenum advenum___—[X|X{_| | {| | | | | | tt | xix] | TT 
Elphidium advenum macelliforme |X| _| | | | | | | | [x] | | | | | tt | 
Elphidium crispum crispum —|X|X] | | | | TT | TT | | xt 
Elphidium crispumssp. ss IX|X| | | | | | | dT dT dT dT TT Tt 
Elphidium excavatum clavatum |_| |X{x{ | |x{x] | [x} | { | {| | tt Tt 
Eoin hee eae 
Elphidium macetum XIX] | | TdT dT TT TT TT TE ET TT 
Parrellinapapilosa XIX] | | | CC sso Gs XAT a 
Parrelinaverriculata XIX] | | TdT TT TT TE TT ETT tT 
Pulleniatina obliquilaculata ss [X|X| | | | | [| | | TT TT TT Tt 


Neogloboquadrinapachyderma__|X|x| | | | | | [| | | | | | | Ty tT 
Globigerina bulloides IXIXT TT TT TTT TT TT TT PT tT 


(1996) indicated this synonymy cannot be justified 
since barwonensis has a more robust test than 
moniliforme, and both taxa have differing habitats: R. 
moniliforme Siddall is associated with inner shelf 
environments and S. barwonensis Collins is found in 
shallow-water intertidal environments. 

This species has only previously been 
described from Victoria and Tasmania. This may be 


167 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


due, in part, to misidentification of the species as 
Protoschista findens Parker, particularly in studies 
conducted on modern material in New South Wales 
(Yassini and Jones, 1989; Yassini and Jones, 1995). 
Protoschista is characterised by branching at the 
proloculus and the formation of two or three uniserial 
series, a feature that none of the specimens illustrated 
from New South Wales possess (e.g. Yassini and Jones, 
1989 Figs 10.10-10.11; Yassini and Jones, 1995 Figs. 
39,43). The range of the species can thus be extended 
to the south coast of New South Wales (Yassini and 
Jones, 1995; this study). 

Specimens of S. barwonensis Collins in 
Tuross Estuary and Coila Lake are generally found in 
low energy environments, such as localities TL3, TL8 
and CL3, where the substrate is composed of 
predominantly muddy material and where aqueous 
vegetation, such as reed beds, are absent. The species 
tolerates a wide range of salinity conditions, from fully 
marine through to conductivity values below 34 mS/ 
cm. 


Superfamily LITUOLOIDEA de Blainville, 1827 
Family LITUOLIDAE de Blainville, 1827 
Subfamily AMMOMARGINULININAE Podobina, 
1978 


Ammobaculites Cushman, 1910 


Type species: 
Spirolina agglutinans d’Orbigny, 1846 


Ammobaculites barwonensis Collins, 1974 
(Plate 1, Figs. 6-7) 


1974 Ammobaculites? barwonensis Collins, p. 9; Pl. 
1, Figs. 3a-b. 

1980 Ammobaculites barwonensis Apthorpe, p. 225; 
Pl. 28, figs. 4, 5, 10-13. 

1989 Ammobaculites foliaceus Yassini and Jones, 
Fig. 10.4. 

1992 Ammobaculites barwonensis Bell and Drury, p. 
13; Figs. 4.7-4.9. 


1995 Ammobaculites barwonensis Bell, p. 229; Fig. 
Dale 
1995 Ammobaculites foliaceus Yassini and Jones, p. 


71; Figs. 51-53. 


Description: 
see Collins (1974), p. 9; Apthorpe (1980), p. 
Was). 


Remarks: 

In his original description of Ammobaculites 
barwonensis, Collins (1974) was doubtful of the 
generic placement of this species because of the 
absence of a definite terminal aperture and suggested 
that “it is possible that minute interspaces between 
grains on the distal face function as such” (Collins, 
1974, p. 9). An emended description by Apthorpe 
(1980) showed that the aperture is terminal and is 
generally an ellipse or elongate slit. 

As noted by Apthorpe (1980) the morphology 
of this species is variable, particularly the overall shape 
and degree of compression of the test. Typical 
specimens, as illustrated by Collins (1974, Pl. 1 Figs. 
3a-b) and Apthorpe (1980, Pl. 28 Figs. 4, 5, 10-13), 
are moderately compressed and rectangular in outline 
but a highly compressed, flabelliform variant does exist 
(Apthorpe, 1980, Pl. 28, Fig. 11; Bell and Drury, 1992, 
Fig. 4.7). There is a morphological continuum between 
the two forms (Apthorpe, 1980), suggesting that both 
morphotypes probably belong to A. barwonensis 
Collins. Both variants, as well a number of intermediate 
forms, were recovered from both surface and core 
samples collected in Tuross Estuary and Coila Lake, 
but the majority of specimens tend to be highly 
compressed and flabelliform in shape. It is unknown 
whether the morphology of the species is affected by 
environmental conditions. 

Like Reophax barwonensis Collins, this 
species has previously only been described from 
Victoria. Also like R. barwonensis Collins, this is 
probably due to misidentification of specimens as 
Ammobaculites foliaceus (Brady) (Yassini and Jones, 
1989; Yassini and Jones, 1995). Ammobaculites 


Plate 1 Facing page: (Unless otherwise specified all scale bars = 100 um) - 1. Scherochorella barwonensis 
(Collins), TL4 July; 2-3. Scherochorella barwonensis (Collins), TL4 515-535; 4. Leptohalysis collinsi Bell, 
TL4 140-160; 5. Ammobaculites exiguus Cushman and Bronnimann, CL3 July; 6-7. Ammobaculites 
barwonensis Collins, TLA July; 8-9. Portatrochammina sorosa (Parr), TL4 490-510; 10. Eggerella subconica 


Parr, CL3 690-710; 11. Miliammina fusca (Brady), TL3 July; 12. Trochammina inflata (Montagu), TL3 
110-130; 13. Trochammina inflata (Montagu), TL3 70-90; 14-15. Quinqueloculina oblonga (Montagu), 
TL4 490-510 (Scale bar for Fig. 15 = 10 um); 16-17. Quinqueloculina seminula (Linne), TL1 560-580; 18- 
19. Triloculina tricarinata d’Orbigny, TL1 July; 20-21. Spiroloculina carinata Fornasini, TL1 300-320 


Proc. Linn. Soc. N.S.W., 124, 2003 


169 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


foliaceus (Brady) differs from A. barwonensis Collins 
by its extremely thin, almost transparent wall, very 
compressed test and smooth exterior (Brady, 1881; 
Brady, 1884). The range of A. barwonensis Collins 
can therefore be extended to the south coast of New 
South Wales (Yassini and Jones, 1989; Yassini and 
Jones, 1995; This study). 

The species is found in both Tuross Estuary 
and Coila Lake, with highest abundances occurring at 
shallow intertidal localities, such as CL4 and TL8 (Fig. 
1). The species has not been recovered from the eastern, 
more marine, part of Tuross Estuary, suggesting this 
species is confined to brackish water conditions. 


Superfamily TROCHAMMINOIDEA Schwager, 
1877 
Family TROCHAMMINIDAE Schwager, 1877 
Subfamily TROCHAMMININAE Schwager, 1877 


Portatrochammina Echols, 1971 


Type species: 
Portatrochammina eltaninae Echols, 1971 


Portatrochammina sorosa (Parr), 1950 
(Plate 1, Figs. 8-9) 


1950 Trochammina sorosa Parr, p. 278; Pl. 5, Figs. 
15-17. 

1967 Trochammina sorosa Hedley et al., p. 23; Pl. 6, 
Figs. 4a-c, Text Figs. 11-15. 

1992 Trochammina sorosa Bell and Drury, p. 13; 
Fig. 4.12. 

1996 Trochammina sorosa Bell, Pl. 11. 

1996 Tritaxis conica Cotter, Figs. 4.10-4.11. 

1999 Trochammina sorosa Bell and Neil, p. 221; 
Fig. 3E. 

1999 Portatrochammina sorosa Hayward et al., p. 
87; Pl. 2, Figs. 4-5. 


Description: 
see Hayward et al. (1999), p. 87. 
Remarks: 
Hayward et al. (1999) placed this species in the 


genus Portatrochammina based on the presence of an 
umbilical flap, discovered upon re-examination of the 
topotype material by Hedley et al. (1967). This feature 
was neither described or illustrated in Parr’s (1950) 
original description of the species but is present on 
specimens found in Tuross Estuary and Coila Lake. P. 
sorosa (Parr) has a restricted distribution and is 
currently only known from the south-eastern coastline 
of Australia and New Zealand (Hayward et al., 1999). 
Specimens of P. sorosa (Parr) found in Tuross Estuary 
and Coila Lake tend to be variable in the number of 
whorls and are generally more trochospiral than the 
specimens illustrated by Parr (1950, Pl. 5 Figs. 15-17) 
from off the east coast of Tasmania. In the modern 
setting, the distribution of P. sorosa (Parr) is widespread 
in Tuross Estuary but no specimens were recovered 
from surface samples collected in Coila Lake. This 
would suggest that P. sorosa (Parr) prefers normal 
marine salinities. 


Suborder MILIOLINA Delage and Herouard, 1896 
Superfamily CORNUSPIROIDEA Schultze, 1854 
Family CORNUSPIRIDAE Schultze, 1854 
Subfamily CORNUSPIRINAE Schultze, 1854 


Cornuspira Schultze, 1854 


Type species: 
Orbis foliaceus Philippi, 1844 


Cornuspira involvens (Reuss), 1850 
(Plate 2, Fig. 15) 


1850 Operculina involvens Reuss, p. 370; Pl. 46, 
Figs. 20a-b. 

1884 Cornuspira involvens Brady, p. 200; Pl. 11, 
Figs. 1-3. 

1967 Cyclogyra involvens Hedley et al., p. 24-25; 
Text Fig. 16. 

1999 Cornuspira involvens Hayward et al., p. 94; PI. 
3, Fig. 16. 


Description: 
see Hayward et al. (1999). 


Plate 2 Facing page: (Unless otherwise specified all scale bars = 100 um) - 1-2. Quinqueloculina poeyana 
d’Orbigny, TLS July; 3. Quinqueloculina sp. cf. Q. disparilis d Orbigny, TL1 January; 4. Bolivina striatula 
Cushman, TL4 515-535; 5-7. Fissurina breviductus sp. nov., TL1 January; 8-9. Lagena blomaeformis 
Yassini and Jones, TL1 410-430; 10-11. Guttulina irregularis (d’ Orbigny), TL1 July; 10-11. Bulimina sp., 


TLS July; 12. Lenticulina sp., TL1 July; 13. Cornuspira involvens (Reuss), TL1 January; 14. Rosalina 
australis (Parr), TL1 January; 15. Rosalina australis (Parr), TL1 300-320; 16-18. Pileolina australensis 


(Heron-Allen and Earland), TL1 630-650 


170 


Proc. Linn. Soc. N.S.W., 124, 2003 


GY 9 fa 7 
Qi isAyss 


“< 


E 
a4 
al 
n 
=] 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


Remarks: 

This species has a cosmopolitan distribution 
(Culver and Buzas, 1986; Loeblich and Tappan, 1994) 
but it has rarely been described from Australia and 
never from the south-eastern coastline of New South 
Wales. It is generally found in fully marine conditions, 
with those specimens found in marsh environments 
probably carried there by tidal currents (Hayward et 
al., 1999). 

Specimens of C. involvens (Reuss) are 
extremely rare in the study area and are only found in 
Tuross Estuary, at TL1 and TL3. Although C. involvens 
(Reuss) is generally found inhabiting inner shelf 
environments (Hayward et al., 1999), one of the tests 
recovered from TL1 during January collection did stain 
with rose Bengal. This, along with the pristine nature 
of the test, suggests that live specimens of this species 
do inhabit the estuary. 


Suborder LAGENINA Delage and Herouard, 1896 
Superfamily NODOSARIOIDEA Ehrenberg, 1838 
Family LAGENIDAE Reuss, 1862 


Fissurina Reuss, 1850 


Type species: 
Fissurina laevigata Reuss, 1850 


Fissurina breviductus sp. nov. 
(Plate 2, Figs. 5-7) 


Diagnosis: 

Differs from other species of Fissurina in its 
small size, short entosolenian tube, perforate flanks 
with smooth central area and its distinctively depressed 
simple aperture. 


Description: 

Test unilocular, tiny, approximately 0.1 mm in 
length and 0.05 mm in width. Relatively ovate in 
outline but gently tapering towards apertural end. Test 
is laterally compressed. Aperture simple, ovate in 
shape, with area immediately around aperture slightly 
depressed. Wall glassy, perforate on flanks with central 
area smooth. Possesses a short entosolenian tube that 
is free, straight and central. 


Type Material: 

Holotype: MU59414; Paratype A: MU5S9379; 
Paratype B; MU59415; All type specimens collected 
from locality TL1 in Tuross Estuary. 


Etymology: 


Latin for “short tube”; in reference to its 
short entosolenian tube. 


172 


Remarks: 

Based upon its ovate outline, smooth surface, 
ovate terminal aperture and entosolenian tube, this 
species is assigned to genus Fissurina. It does not 
however, accord with any of the described species 
within this genus, nor does it resemble any species 
previously described from shallow water environments 
along the south-eastern Australian coastline. This may 
in part be due to its small size, generally less than 100 
tum in length. 

This species does resemble a variant of 
Lagena globosa Montagu, documented by Sidebottom 
(1912, Pl. 14 Figs. 13-15) from the outer shelf and 
abyssal plain of the south-west Pacific, Lagena globosa 
var. emaciata Reuss. Both the specimens illustrated 
by Sidebottom (1912) and those found as part of this 
study are ovoid in shape, smooth walled, possess a 
free, centralised entosolenian tube, and have a single 
terminal aperture. It is unclear whether the specimens 
shown by Sidebottom (1912) are depressed around the 
aperture. 

Sidebottom’s (1912) material was collected 
from depths of below 710 fathoms, whereas the 
material from the Tuross Estuary is from shallow water. 
Such extreme bathymetric differentiation make it 
unlikely that the specimens illustrated by Sidebottom 
(1912) and the specimens collected from the study area 
are conspecific. Specimens of F: breviductus sp. nov. 
from the study area are confined to surface and core 
samples from locality TL1 and all specimens found in 
surface samples had taken up the rose Bengal stain, 
suggesting the tests were in-situ and their natural 
habitat is a shallow water open estuary environment. 


Family POLYMORPHINIDAE d’Orbigny, 1839 
Subfamily POLYMORPHININAE d’Orbigny, 1839 


Guttulina d’Orbigny, 1839 
Type species: 
Polymorphina (les Guttulines) communis 
d’Orbigny, 1826 


Guttulina irregularis (d’Orbigny), 1846 
(Plate 2, Figs. 10-11) 


1846 Globulina irregularis d’ Orbigny, p. 226; Pl. 
13, Figs. 9-10. 

1930 Guttulina irregularis Cushman and Ozawa, p. 
25; Pl. 3, Figs. 3,4; Pl. 7, Figs. 1,2. 

1937 Guttulina irregularis Parr and Collins, p. 192; 
Pl. XII, Fig. 2. 

1995 Nevellina coronota Yassini and Jones, p. 89; 
Fig. 244. 

1999 Guttulina irregularis Hayward et al., p. 117; 


Pl. 7, Figs. 10-11. 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


Description: 
see Hayward et al. (1999), p. 117. 


Remarks: 

This species can be distinguished from other 
species of Guttulina by its pyriform test, rounded 
periphery and non-depressed sutures (Hayward et al., 
1999). It has been recorded from a number of localities 
throughout the west Pacific (Parr and Collins, 1937; 
Nomura, 1981; Hayward et al., 1999). 

In the study area, this taxon was recovered 
from surface samples as locality TL1. The abraded 
nature of the tests suggests they have been transported 
to the site by onshore oceanic currents, since G. 
irregularis (d’ Orbigny) is generally found at inner- and 
mid-shelf depths (Hayward et al., 1999) 


Suborder ROTALIINA Delage and Herouard, 1896 
Superfamily BOLIVINOIDEA Glaessner, 1937 
Family BOLIVINIDAE Glaessner, 1937 


Bolivina d’Orbigny, 1839 


Type species: 
Bolivina plicata d’Orbigny, 1839 


Bolivina striatula Cushman, 1922 
(Plate 2, Fig. 4) 


1922 Bolivina striatula Cushman, p. 27; Pl. 3, Fig. 
10. 

1937 Bolivina striatula Cushman, p. 154; Pl. 18, 
Figs. 30, 31. 

1950 Bolivina striatula Parr, p. 239. 

1974 Brizalina striatula Collins, p. 30. 

1979 Brizalina striatula Albani, p. 33; Fig. 56-6. 

1980 Bolivina striatula Apthorpe, Pl. 27, Fig. 2. 

1989 Brizalina striatula Yassini and Jones, Fig. 13.3. 

1995 Brizalina striatula Yassini and Jones, p. 132; 
Figs. 526-529, 543-544, 655. 

1996 Bolivina striatula Bell, Pl. 5d. 

2001 Brizalina striatula Albani et al. 


Description: 
see Hayward et al. (1999), p. 127. 


Remarks: 

The amendments made by Sgarrella (1992) to 
the genus Bolivina, with Brizalina now a junior 
synonym of Bolivina, are adopted herein. B. striatula 


Cushman is easily distinguished from other species of 
Bolivina by its parallel fine ribs on the lower half of 


the test. This species is generally found in sheltered, 
slightly brackish environments (Hayward et al., 1999) 
and its geographic distribution is extensive (Murray, 


Proc. Linn. Soc. N.S.W., 124, 2003 


1991). This species is extremely rare in Tuross Estuary 
with only small numbers recovered from localities 
TL1 and TL3. The species was not recovered from 
Coila Lake. 


Superfamily DISCORBOIDE Ehrenberg, 1838 
Family DISCORBIDAE Ehrenberg, 1838 


Lamellodiscorbis Bermudez, 1952 


Type species: 
Discorbina dimidiata Jones and Parker in 
Carpenter et al., 1862 


Lamellodiscorbis dimidiatus (Jones and Parker), 
1862 
(Plate 3, Figs. 3-4). 


1862 Discorbina dimidiata Jones and Parker in 
Carpenter et al., p. 201; Text Fig. 32b. 

1945 Discorbis dimidiatus Parr, p. 208. 

1967 Discorbina dimidiatus Hedley et al., p. 33; 
Text-Figs. 28-43. 

1974 Discorbis dimidiatus Collins, p. 34. 

1989 Lamellodiscorbis dimidiatus Yassini and Jones, 
Figs. 17.9-17.11. 

1992 Lammellodiscorbis dimidiatus Hansen and 
Revets, p. 176; Pl. 4, Figs. 1-3, 7-8 

1995 Trochulina dimidiata Yassini and Jones, p. 158; 
Figs. 916-917. 

1996 Lamellodiscorbis dimidiatus Bell, P1. 5b. 

1999 Trochulina dimidiatus Hayward et al., p. 139; 


Pl. 10, Figs. 9-11. 


Description: 

see Hayward et al. (1999), p. 139. 
Remarks: 

A number of authors (Loeblich and Tappan 
1987, Yassini and Jones 1995, Hayward et al. 1999) 
have assigned this species to the genus Trochulina, 
suggesting that both Discorbina and Lamellodiscorbis 
are junior synonyms of Trochulina. However, Hansen 
and Revets (1992) clearly illustrate the validity of both 
Discorbina and Lamellodiscorbis and so herein 
dimidiatus has been assigned to Lamellodiscorbis. 

This species is distinguished by its 

characteristic umbilical side, which has short, 
thickened umbilical plates bordered by deep clefts 
along the sutures (Hayward et al., 1999). Collins (1974) 
noted that specimens are sometimes strongly biconvex 
with secondary thickening on both faces. The 
geographic distribution of L. dimidiatus (Jones and 
Parker) is confined to coastal waters of Australia and 
the Pacific (Heldey et al., 1967; Yassini and Jones, 
1995; Hayward et al, 1999) and it is generally found 


173 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


in stenohaline environments (Cann et al., 2000) with 
greatest abundance occurring in exposed, shallow, high 
energy environments. 

Specimens from the study area were only 
recovered from samples collected at locality TL1. The 
broken and abraded nature of the tests, as well as the 
lack of any “live” specimens of L. dimidiatus (Parker 
and Jones) in the surface samples from locality TL1, 
suggests that living examples of this species do not 
inhabit the estuary but rather are transported in via 
currents. 


Superfamily GLABRATELLIOIDEA Loeblich and 
Tappan, 1964 
Family GLABRATELLIDAE Loeblich and Tappan, 
1964 


Pileolina Bermudez, 1952 


Type species: 
Valvulina pileolus d’Orbigny, 1839 


Remarks: 

Pileolina was recorded as a genus of uncertain 
status by Loeblich and Tappan (1988), however I 
follow Hayward et al. (1999). in placing the genus in 
the family Glabratellidae. 


Pileolina australensis (Heron-Allen and 
Earland), 1932 
(Plate 2, Figs. 18-20) 


1932 Discorbis australensis Heron-Allen and 
Earland, p. 416. 
1995 Glabratella australensis Yassini and Jones, p. 
160; Figs. 731-734. 
Description: 
see Heron-Allen and Earland (1932), p. 416. 


Remarks: 

With the validity of the genus Pileolina 
established by Hayward et al. (1999), the species 
australensis Heron-Allen and Earland is herein re- 


assigned to the genus. The genus Glabratella, where 
australensis Heron-Allen and Earland was most 
recently assigned by Yassini and Jones (1995), is 
characterised by the presence of globular chambers 
and a rounded periphery (Loeblich and Tappan, 1988; 
Hayward et al., 1999) whereas specimens of 
australensis described by Heron-Allen and Earland 
(1932) and illustrated by Yassini and Jones (1995, Figs. 
731-734), as well as those found as part of this study, 
do not have globular chambers and have an acute 
periphery. These features, along with the flat, involute 
umbilical site with radiating striae and papillae clearly 
place the species in Pileolina. 

Pileolina australensis (Heron-Allen and 
Earland) is easily distinguished from most other species 
of Pileolina by the ornament present on its umbilical 
side, which consists of strong tubercules located 
centrally and numerous prominent striae around the 
outer edge (Plate 2, fig. 20). P australensis (Heron- 
Allen and Earland) does resemble P. zealandica Vella, 
which has a similar ornament on the umbilical side, 
but can be discriminated by the nature of the chambers 
on the spiral side, which in P. australensis (Heron- 
Allen and Earland) are much longer than in P. 
zealandica Vella (Hayward et al. 1999). 

Pileolina australensis (Heron-Allen and 
Earland) is endemic to Australia and its distribution is 
confined to a variety of marine dominated 
environments (Yassini and Jones, 1995). Specimens 
found in the study area conform to these ecological 
parameters and tests were only found at locality TL1, 
where marine conditions dominate. 


Superfamily PLANORBULINOIDEA Schwager, 
1877 
Family CIBICIDIDAE Cushman, 1927 
Subfamily CIBICIDINAE Cushman, 1927 


Cibicides de Montfort, 1808 


Type species: 
Cibicides refulgens de Montfort, 1808 


Plate 3 Facing page: (Unless otherwise specified all scale bars = 100 um) - 1-2. Cibicides dispars (d’ Orbigny), 
TLI July; 3. Lamellodiscorbis dimidiatus (Jones and Parker) TL1 300-320; 4. Lamellodiscorbis dimidiatus 
(Jones and Parker), TL1 July; 5-6. Ammonia aoteana (Finlay), TL4 515-535; 7-8. Ammonia aoteana (Finlay) 
TL4 140-160; 9. Elphidium lene Cushman and McCulloch, TL7 July; 10. Elphidium lene Cushman and 
McCulloch, TL7 July (Scale bar for figure 5 = 10um); 11-12 Elphidium excavatum clavatum Cushman, 


TL4 515-535; 13-14. Elphidium advenum advenum (Cushman), TL1 410-430; 15-16. Elphidium crispum 
crispum (Linne), TL1 180-200; 17-18. Elphidium crispum (Linne) ssp. TL1 300-320; 19-20. Elphidium 
advenum macelliforme McCulloch, TL1 515-535 


174 


Proc. Linn. Soc. N.S.W., 124, 2003 


Proc. Linn. Soc. N.S.W., 124, 2003 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


Cibicides dispars (d’Orbigny), 1839 
(Pl. 3, Figs. 1-2) 


1839 Truncatulina dispars @’ Orbigny, p. 38; Pl. 5, 
Figs. 25-27. 

1980 Cibicides dispars Boltovskoy et al., p. 24; PI. 
8, Figs.12-16. 

1995 Cibicidoides floridanus Yassini and Jones, p. 
169, Figs. 889-896. 

1999 Cibicides dispars Hayward et al., p. 154; Pl. 
14, Figs. 22-24. 


Description: 
see Hayward et al. (1999), p. 154. 


Remarks: 

There has been a great deal of confusion 
concerning the taxonomic assignment of planoconvex 
species of Cibicides (Hayward et al., 1999). Because 
of this, the broad view taken by Hayward et al. (1999) 
with respect to C. dispars (d’Orbigny) is followed 
herein. Whilst the material found in the study area does 
not exactly match the specimens of C. dispars 
(d’Orbigny) illustrated by Boltovskoy et al. (1980, PI. 
8 Figs. 12-16) and Hayward et al. (1999, Pl. 14 Figs. 
22-24), mainly due to the lack of perforations on much 
of the involute side in the Tuross Estuary specimens, 
they match the original description given by d’ Orbigny 
(1839) and have an overall morphology that is 
otherwise strikingly similar. Therefore specimens from 
the study area are referred to C. dispars (d’ Orbigny). 
Also of note is that the specimens recovered from the 
study area are clearly conspecific with the material 
identified by Yassini and Jones (1995) as Cibicidoides 
floridanus (Cushman), a designation that was later 
recognised as a junior synonym of C. dispars 
(d’Orbigny) by Hayward et al. (1999). 

Specimens of C. dispars are present in Tuross 
Estuary and Coila Lake but are confined to localities 
where marine influences dominate (TL1, CL2). This 
is not unexpected since the species thrives in fully 
marine inner- to mid- shelf environments. Some 
specimens found in surface samples from TL1 did take 
up the rose Bengal stain, indicating that C. dispars 
(d’Orbigny) can survive in estuarine conditions and is 
not necessarily washed in from shelf environments. 


Superfamily ROTALIOIDEA Ehrenberg, 1839 
Family ROTALIIDAE Ehrenberg, 1839 
Subfamily AMMONIINAE Saidova, 1981 


Ammonia Briinnich, 1772 


Type species: 
Nautilus becarii Linne, 1758 


176 


Ammonia aoteana (Finlay), 1940 
(Plate 3, Figs. 5-8) 


1940 Strebulus aoteanus Finlay, p. 461. 

1967 Ammonia aoteanus Hedley et al., p. 47; Pl. 11, 
Figs. 4a-c, Text Figs. 56-60. 

1974 Ammonia aoteanus Collins, p. 40; Pl. 3, Figs. 
30a-c. 

1968a Ammonia beccarii Albani, p. 30; Fig. 129 

1968b Ammonia beccarii Albani, p.110; Pl. 9, Figs. 
7, 9-10 

1979 Ammonia beccarii Albani, p. 40; Fig. 88-1 

1980 Ammonia aoteanus Apthorpe, p. 225; Pl. 27, 
Figs. 5-6; Pl. 29, Figs. 1-2. 

1994 Ammonia beccarii (Linne) forma aoteanus 
Hayward and Hollis, p. 213; Pl. 4, Figs. 1-3. 

1996 Ammonia aoteanus Bell, p. 6. 


1996 Ammonia beccarii List, p.19; Pl. 1 Figs. G-H 


Description: 
see Finlay (1940), p. 461; Hayward and 
Hollis (1994), p. 213. 


Remarks: 

The problems surrounding the species, 
subspecies and formae within the genus Ammonia have 
been discussed extensively in the literature (see Bell, 
1996 for summary). Due to the large deal of 
morphological overlap within the genus and subjective 
recognition of key features by workers (Murray, 1979), 
it appears that genetic studies will be required to sort 
out the relationships between the numerous 
morphological variants. 

Because of the uncertainty surrounding the 
genus, two main approaches have traditionally been 
used in descriptions of Ammonia in estuarine faunas 
in south-eastern Australia. In New South Wales, 
specimens falling within the genus Ammonia have been 
referred to as Ammonia beccarii (Linne) (Albani, 
1968a; Albani, 1978; Albani, 1979; Yassini and Jones, 
1989; Yassini and Jones, 1995; Cotter, 1996) whereas 
in Victoria, specimens have been referred to Ammonia 
aoteana (Finlay) (Collins, 1974; Apthorpe, 1980; Bell, 
1996), a species of Ammonia described by Finlay 
(1940) from the south-west Pacific. The lack of 
strongly beaded sutures on the umbilical side of 
specimens from Tuross Estuary and Coila Lake, a 
feature typical of A. beccarii (Linne) sensu stricto 
(Hayward and Hollis, 1994, Pl. 4 Figs. 1-3), indicates 
that the specimens of Ammonia from the study area 
can be assigned to A. aoteana (Finlay). 

The material from the study area displays a wide 
variation in morphology that is characteristic of the: 
species and widely distributed throughout the two 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


estuaries. The only discernible trend in morphology is 
a tendency for specimens recovered from localities 
where marine conditions dominated, such as TL1, to 
have an umbilical boss. Specimens from slightly 
brackish localities such as TL3 and TL8, tend to lack 
this feature. The correlation between the presence or 
absence of an umbilical boss with prevailing salinity 
is also a feature that has been noted in variants of 
Ammonia beccarii (Linne) (Murray, 1979). 


Family ELPHIDIIDAE Galloway, 1933 


Remarks: 

The family Elphidiidae, and in particular the 
genus Elphidium, has undergone a great deal of 
reassessment since it was first described by Galloway 
(1933), with a number of genera of doubtful validity 
erected within the family. In an attempt to gain 
standardisation in identification of the members of this 
family, particularly in Australia, all Elphidiidae in this 
study were identified following the concept of the 
family presented by Hayward et al. (1997), except 
where otherwise noted. 


Subfamily ELPHIDIINAE Galloway, 1933 


Elphidium de Montfort, 1808 


Synonymy: 
see Loeblich and Tappan (1988), p. 674. 


Type species. 
Nautilus macellus var.? Fichtel and Moll, 
1798 


Elphidium advenum (Cushman), 1922 


1922 Polystomella advena Cushman, p. 56; Pl. 9, 
Figs. 11-12. 


1997 Elphidium advenum Hayward et al., p. 64. 


Description: 
see Hayward et al. (1997), p. 64. 


Elphidium advenum advenum (Cushman), 1922 
(Plate 3, Figs. 13-14) 


Synonymy: 
see Hayward et al. (1997), p. 65. 


Description: 
see Hayward et al. (1997), p. 65. 


Remarks: 
E. advenum advenum (Cushman) is easily 


Proc. Linn. Soc. N.S.W., 124, 2003 


distinguished from other species of E. advenum by its 
distinctive umbilical boss, which fills the entire 
umbilical area. The original material illustrated by 
Cushman (1922, Pl. 9 Figs. 11-12) did not clearly 
display the presence of an umbilical boss, however it 
is described in the text as a feature of the species. In a 
later publication by Cushman (1939), the boss is clearly 
illustrated. This species has been recorded from a 
variety of environments, from brackish water intertidal 
conditions to exposed inner shelf environments 
(Hayward et al., 1999). E. advenum advenum 
(Cushman) is found throughout the south-west Pacific 
(Hayward et al., 1999) as well as the coastline of 
Australia (Albani and Yassini, 1993). 

Specimens of E. a. advenum (Cushman) from 
the study area differ from the material illustrated by 
Hayward et al., (1997), in that they have a much weaker 
keel and are slightly less laterally compressed. Tests 
of this subspecies were recovered from localities TL1, 
TL8 and CL2 in both surface and core material. The 
sites represent a range of environmental conditions, 
particularly TL8 in comparison to TL1 and CL2, 
supporting the assertion that this species tolerates a 
wide range of conditions. 


Elphidium advenum macelliforme McCulloch, 
1981 
(Plate 3, figs. 19-20) 


1981 Elphidium macelliforme McCulloch, p. 119; 
Pl. 40, Fig. 1 

1993 Elphidium macelliforme Albani and Yassini, p. 
28; Figs. 65-66 

1997 Elphidium advenum macelliforme Hayward et 
al., p. 68; Pl. 5, Figs. 6-12 

Description: 

see Hayward et al. (1999), p. 68 


Remarks: 

This species is distinguished from most other 
sub-species of E. advenum_Cushman by its less inflated 
chambers and narrower, less incised sutures. It is 
distinguished from E. advenum limbatum (Chapman) 
by its distinct umbonal boss. This species is found 
along the coastline of Australia (Albani and Yassini, 
1993) as well as in the eastern part of the Pacific Ocean 
(McCulloch, 1981). In the study area, this species 
appears to be only tolerant of marine conditions as 
specimens were confined to site TL1. 


Elphidium crispum (Linne), 1758 


Synonymy: 
see Hayward et al. (1997), p. 74. 


177 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


Description: 
see Hayward et al. (1997), p. 74. 


Elphidium crispum crispum (Linne), 1758 
(Plate 3, Figs. 15-16) 


Synonymy: 
see Hayward et al. (1997), p. 74. 


Description: 
see Hayward et al. (1997), p. 74. 


Remarks: 

This subspecies is distinguished from other 
similar species of Elphidium, in particular E. macellum 
(Fichtel and Moll) and E. craticulatum (Fichtel and 
Moll), by its distinctive sparsely pitted boss. This 
subspecies is extremely common along the east coast 
of Australia and is abundant in shallow subtidal 
situations where normal marine salinities are present. 
Specimens of E. c. crispum (Linne) are found in 
material collected at localities TL1 and CL2, the two 
localities where marine conditions dominate. 


Elphidium excavatum (Terquem), 1875 


Description: 
see Hayward et al. (1997) 


Remarks: 

As stated by Hayward et al. (1997), this species 
is distinguished by its broadly rounded unkeeled 
periphery, low number of chambers in the final whorl 
(less than 12) and the presence of papillae of the sides 
of sutural pits, the umbilical area and the base of the 
apertural face. Its distribution is extensive, recorded 
from a variety of environments worldwide (Miller et 
al., 1982). 


Elphidium excavatum clavatum Cushman, 1930 
(Plate 3, Figs. 11-12) 


Synonymy: 
see Hayward et al. (1997), p. 76 


Description: 
see Hayward et al. (1997), p. 76. 


Remarks: 

E. excavatum clavatum Cushman differs from 
other subspecies of E. excavatum (Terquem) because 
of the intermediate length of its septal bridges, it 
umbilical collar and the presence of a small umbonal 
boss (Hayward et al., 1997). 


178 


In the study area, E. e. clavatum Cushman is 
distinguished from specimens of E. lene Cushman and 
McCulloch, also found in Tuross Estuary, by its less 
numerous septal bridges and more numerous papillae. 
The highest abundance of this species was recorded 
from locality TL3, indicating it favours middle estuary, 
intertidal environments. The species is possibly tolerant 
of a wide range of conditions however, as rare 
specimens were also recorded from localities TL1 and 
CL2. 


Elphidium lene Cushman and McCulloch, 1940 
(Plate 3, Figs. 9-10) 


1940 Elphidium incertum (Williamson) var. lene 
Cushman and McCulloch, p. 170; P1.19, Figs. 2, 
4. 

1968a Elphidium poeyanum Albani, p. 34; Fig. 158. 

1979 Cribroelphidium poeyanum Albani, p. 47; Fig. 
110-1. 

1989 Elphidium depressulum Yassini and Jones, p. 
263; Figs. 16.1-16.3. 

1992 Elphidium poeyanum Bell and Drury, p. 15; 
Fig. 4.20. 

1993 Cribroelphidium poeyanum Albani and Yassini, 
p. 17; Figs. 10-15. 

1995 Cribroelphidium poeyanum Yassini and Jones, 
p. 178; Figs. 1074-1075. 

1997 Elphidium lene Hayward et al., 1997, p. 84; Pl. 
13, Figs. 1-8. 


Description: 
see Hayward et al. (1997), p. 84 


Remarks: 

This species resembles E. excavatum (Terquem) 
in overall appearance, but differs in its more 
compressed test, higher apertural face, more numerous 
septal bridges, and less papillose ornament (Hayward 
et al., 1997). Elphidium lene Cushman and McCulloch 
is found along the east coast of Australia, but has 
generally been referred to either Cribroelphidium 
poeyanum (d’Orbigny) or E. poeyanum (d’ Orbigny) 
(Albani, 1968; Bell, 1992; Albani and Yassini, 1993; 
Yassini and Jones, 1995). E. poeyanum (d’ Orbigny) is 
characterised by a coarsely perforate surface (Hayward 
et al., 1999), a feature not evident on specimens found 
in Tuross Estuary and Coila Lake, or from other parts 
of eastern Australia. The specimens are thus referred 
to E. lene Cushman and McCulloch. 

In the study area, E. lene Cushman and 
McCulloch is confined to Tuross Estuary. The highest 
abundance was recorded at locality TL7, a shallow salt 
marsh environment but it is also recorded from locality 


Proc. Linn. Soc. N.S.W., 124, 2003 


L. STROTZ 


TL1, where fully marine conditions dominate, 
suggesting that this species tolerates a wide range of 
conditions. 


Elphidium macellum (Fichtell and Moll), 1798 
(Plate 4, Figs. 1-2) 


Synonymy: 
See Hayward et al., 1997, p. 84 


Description: 
See Hayward et al., 1997, p. 84 


Remarks: 

This species is distinguished from other species 
of Elphidium by its compressed profile, narrow radial 
ribs that extend to the peripheral keel and its depressed 
umbilical area with few irregular papillae (Hayward 
et al., 1997). Most specimens found in the study area 
have between 8-10 septal bridges extending across 
each chamber but rare specimens have less than seven. 
This species is common of shallow subtidal 
foraminiferal associations along the eastern Australian 
coastline. In the study area, specimens are confined to 
sites were normal marine conditions dominate (TL1, 
Cl): 


Plate 4 (Unless otherwise specified all scale bars = 100 um) - 1-2. Elphidium macellum (Fichtell and Moll), TL1 
300-320; 3. Parrellina papillosa (Cushman), TL1 July 4; 4. Parrellina papillosa (Cushman), TL1 July; 5-6. 
Parrellina verriculata (Brady), TL1 July; 7-8. Pulleniatina obliquilculata (Parker and Jones), TL1 July; 9-10. 
Neogloboquarina pachyderma (Ehrenberg), TL1 515-535; 11-12. Globigerina bulloides d’ Orbigny, TL1 515- 
535 


Proc. Linn. Soc. N.S.W., 124, 2003 


179 


HOLOCENE FORAMINIFERA FROM THE SOUTH COAST, NSW 


ACKNOWLEDGEMENTS 


I am grateful to Dr Glenn Brock, who provided 
never-ending assistance throughout the duration of this 
project and helped with editing of this manuscript; to Dave 
and Helen Mathieson; to Dave for all his assistance with 
field work and to Dave and Helen collectively for keeping 
me well fed in the field; The other members of MUCEP, too 
numerous to name here, for their general advice and 
assistance throughout the duration of this project; Team 
Hostetler (Blair Hostetler, James Ray, Grieve Brown, Peter 
Weiland) who assisted with chemical analysis and to Blair 
and James, in particular, for fruitful discussions on the 
implications of the resulting data; Ken Bell for acting as a 
sounding board for my taxonomic interpretations and 
provided advice on processing techniques; David Haig and 
two anonymous referees who reviewed this manuscript; 
Russel Field, Nigel Wilson and Dan Keating for help with 
obtaining equipment; Peter Evans, the caretaker at Horse 
Island, who allowed access to the property for collection of 
samples. The Betty Mayne Fund of the Linnean Society of 
New South Wales which provided funds towards this project. 


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Proc. Linn. Soc. N.S.W., 124, 2003 


Radio-tracking Studies of Common Ringtail Possums, 
Pseudocheirus peregrinus, in Manly Dam Reserve, Sydney 


BARBARA Suoru, MICHAEL AuGEE2 AND STEFAN Rose? 


Biological Science, University of NSW, Sydney 2052 
Current address: 185 Morgans Road, Mount White NSW 2250; 2Current address: Wellington Caves Fossil 
Studies Center, 89 Caves Road, Wellington NSW 2820; 3Current address: Ecotone Ecological Consultants 
Pty Ltd, 39 Platt Street, Waratah NSW 2298. 


(This paper edited by J-C. Herremans) 


Smith, B., Augee, M. and Rose, S. (2003). Radio-tracking studies of Common Ringtail Possums, 
Pseudocheirus peregrinus, in Manly Dam Reserve, Sydney. Proceedings of the Linnean Society of New 


South Wales124, 183-194. 


In radio-tracking studies in Manly Dam War Memorial Reserve, Common Ringtail Possums were found to 
survive an average of 319 days, with 80% of known deaths being due to predation by foxes and cats. The 
study area contained few large trees with hollows and 88% of the nest sites used were dreys. Any drey might 
be used by several possums, although rarely simultaneously. Ringtails were found to be sedentary, usually 
occupying dreys and foraging within a Banksia ericifolia thicket. Only 37% of the radio-tracked possums 
moved more than 50 m from their point of capture, and such movements often resulted in the establishment 
of a new foraging range. Males were more likely to make such shifts than females. 


Manuscript received 10 Oct 2002, accepted for publication 6 January 2003. 


KEYWORDS: Common Ringtail Possum, drey, home range, Pseudocheirus peregrinus, radio-tracking, 


INTRODUCTION 


This paper reports a long-term study (1994- 
1999) of Common Ringtail Possums, Pseudocheirus 
peregrinus, in the Manly Dam War Memorial Reserve. 
It complements and extends an earlier study (Augee 
et al. 1996) carried out in Ku-ring-gai Chase National 
Park. The latter study was terminated by fires which 
burnt the study area in January 1994. 

The Ku-ring-gai Chase study found mean 
survival of Common Ringtail Possums to be 101 days 
after the commencement of radio-tracking. It provided 
the first quantitative evidence of high levels of 
predation on possums by foxes and cats (Rose et al. 
1994, Augee et al. 1996), perhaps related to the 
unexpected finding that up to 10% of the nest sites 
were on the ground (Augee et al. 1996). Unlike other 
possums, ringtails are not dependent on tree hollows 
for nesting sites. They use tree hollows but also 
construct free-standing nests, known as “dreys”, that 
are often mistaken for birds’ nests. Ringtail dreys are 
larger and more spherical than those of birds except 
for some babblers, none of which occurred in either 
study area. Nests on the ground were usually 
constructed in grass clumps and varied in structure 


from simple depressions to fully lined shelters similar 
to arboreal dreys. 

Most of the possums radio-tracked in the Ku- 
ring-gai study were introduced into the area and it is 
possible that their behaviour was atypical compared 
to lifetime residents. The Manly Dam Reserve study 
was based on resident, “wild” ringtails and was 
designed to determine if the high levels of predation 
and patterns of nest usage reported by Augee et al. 
(1996) are a widespread occurrence or simply the result 
of unusual conditions prevailing in the Ku-ring-gai 
Chase study. 

The Manly Dam Reserve study site was selected 
because it seemed likely to be less susceptible to 
bushfire than Ku-ring-gai Chase. That assumption 
turned out to be incorrect as part of Many Dam Reserve 
did burn, but only after this study was completed. Both 
studies areas are Sydney sandstone woodland, although 
Manly Dam Reserve contains relatively few large trees 
with hollows compared to Ku-ring-gai Chase. 

The Manly Dam Reserve study was designed 
not only to provide data on survival and nest usage of 
wild Common Ringtail Possums, but also to run long 
enough to gather data on long term movements and 
dispersal of individuals and intraspecific relationships. 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


In the course of radio-tracking incidental data were 
obtained, particularly in regard to reproduction. 


MATERIALS AND METHODS 


The study area 
The study area comprised 8 ha in the north- 


west corner of the Manly Dam War Memorial Reserve 
as shown in Fig. 1. Its topography was undulating 
and consisted of moderately sloping hillsides around 
the headwaters of Curl Curl Creek and tributaries. The 
vegetation was predominantly dry sclerophyll 


Manly 
Warringah 


shrubland and heathland on Hawkesbury Sandstone. 

Five vegetation communities were identified within the 

Manly Dam Reserve study area: 

Banksia ericifolia Thicket: Scrub to 4 metres height 
and 60-70% cover. Predominantly Banksia 
ericifolia, with Kunzea ambigua and Hakea 
teretifolia. 

Low Open Woodland: Low eucalypts to 8 metres at 3 
- 4 metre intervals. Dominated by Red Bloodwood 
Corymbia gummifera and stringybarks. Sparse 
shrub layer and dense sclerophyllous ground layer. 

Heathland/Low Open Woodland: Dense low shrubs 
and grassland to 2 metres, with areas of low, scat- 


Study 
Area 


War 
Memorial 
Reserve 


1 km . 
— a 
\ Manly 
Middle N 
Harbour 
x 
9 
30 
¢ 
7 N 
SNane? 
Sydney 
CBD 
5 km 


Figure 1. The study area in Manly Dam War Memorial Reserve. 


184 


Proc. Linn. Soc. N.S.W., 124, 2003 


B. SMITH, M. AUGEE AND S. ROSE 


tered eucalypts to 8 metres. 

Open Forest: Trees to 20 metres and 30 - 40% cover. 
Dominant species include Sydney Peppermint Eu- 
calyptus piperita, Smooth-barked Apple Angophora 
costata, Red Bloodwood Corymbia gummifera, Sil- 
ver-top Ash E. sieberi and stringybarks. 

Riparian Open Forest: Trees to 20 metres height along 
the bank and floodplain of the creekline. Dominated 
by Black Wattle Callicoma serratifolia and euca- 
lypts with a shrub layer of Banksia ericifolia and a 
ground layer with sedges such as Gahnia spp. and 
Coral Fern Gleichenia spp. 


Animals 

Seventy-nine ringtails were caught in the 
study area and fitted with radio-collars. They were 
designated with numbers 242-320. Sex is indicated by 
the prefix F for females and M for males. Individual 
details are given in Appendix I. Individuals weighing 
less than 600 g were classified as juveniles. 

For ringtails that had been followed since they 
were pouch young, their mother was of course known. 
We were able to identify probable fathers in cases 
where nest sharing with the mother had been observed 
at about the right time for conception and where nest 
sharing between an adult male and the juvenile was 
observed. 


Radio-tracking system 
Ringtails were caught by shaking them from 


low lying trees or bushes, fitted with radio-transmitters 
built into collars and subsequently located by tracking 
the radio signal as described in Augee et al. (1996). 
The process of capture and collaring usually 
resulted in the animal moving to a nest other than the 
one in which it had been captured, but on only one 
occasion was that nest site out of the foraging range. 
One individual (F245), after it took several attempts 
to recapture her and replace a faulty transmitter, 
dispersed immediately by 110 m to a new foraging 
range in which she remained for more than two years. 


Data collection 

The position of each Ringtail was usually 
determined weekly, although more frequent 
determinations were often made in the first few weeks 
after a collar was fitted. The nest site (drey, tree hollow, 
ground nest or other) was recorded. 

When radio collars were replaced in order to 
change batteries, pouches of females were examined. 
The presence and estimated weight of any joeys was 
noted. Approximate date of birth was extrapolated from 
the growth curve published by Smith (1995, p. 37). 

Probable cause of death was determined from 
corpses using the criteria set out in Augee et al. (1996). 


Proc. Linn. Soc. N.S.W., 124, 2003 


Briefly, intact transmitter collars with little distortion 
found in association with scattered fur but no body 
parts were scored as fox kills. Cached collars and 
collars located a considerable distance from the last 
recorded nest site were also scored as fox kills. 
However mangled corpses or collars found with body 
parts (usually heads, paws, intestines and often the 
caecum) were scored as cat kills. 


Foraging and home ranges 
In a separate study of ringtails carried out by 


Newton (1997) at Manly Dam, the areas used by 
individual ringtails foraging around their nesting sites 
were found to average 0.020 ha for females and 0.034 
ha for males. These foraging ranges had a maximum 
diameter of about 50 m. In the present, long-term study 
therefore we considered movements less than 50 m as 
foraging movements and movements over 50 m as 
exploratory. These longer movements may or may not 
have resulted in a shift of foraging range. When 
ringtails were found to establish a new foraging range, 
new nest sites more than 50 m from their previous site 
were considered to be dispersal. We use the term “home 
range’ to refer to the sum of all foraging areas used by 
an individual throughout the course of the study. Home 
ranges are illustrated in figures by the smallest convex 
polygon that can be drawn to connect the outermost 
recorded nest sites. Only data for those animals (62 
out of 79) that were tracked for more than a month 
were used in determination of home ranges. 


Survival statistics 

Where mean survival is given it has been 
calculated only for individuals whose date of death 
could be determined. 

Survival functions for various data subsets 
were estimated using software provided by K.H. 
Pollock based on his (Pollock et al. 1989) modification 
for staggered entry of animals of the Kaplan-Meier 
product limit estimator (Kaplan and Meier 1958) as 
detailed in Augee et al. (1996). The survival functions 
in this study and the previous Ku-ring-gai Chase study 
(Augee et al. 1996) were based on weekly observations. 


RESULTS 


Survival 

The mean survival of all ringtails was 319 
days after commencement of radio-tracking (n = 60, 
SD = 336, median = 172 days). Mean survival of adults 
was 465 days (n = 33, SD = 374, median = 371 days). 
Mean survival of juveniles was 140 days (n = 27, SD 
= 156, median = 90 days). 

The mean survival of all resident ringtails was 


185, 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


Survival 


—O- SURVIVAL Ku-ring-gai Wild 
—O— SURVIVAL Manly Dam Wild 


Week 


Figure 2. Comparison of Kaplan-Meier survival functions (modified for staggered entry) for all wild 
ringtails radio-tracked at Manly Dam Reserve (n=78) and Ku-ring-gai Chase (n=41). Chi-squared = 10.227; 


P<0.01. 


much greater at Manly Dam Reserve (319 days) than 
at Ku-ring-gai Chase (182 days). The Kaplan-Meier 
survival functions, modified for staggered entry of 
animals, for all wild possums tracked at Manly Dam 
Reserve and at Ku-ring-gai Chase are shown in Fig. 2. 
The survival function for the Manly Dam Reserve 
population is significantly lower (Chi-squared = 
10.227, P<0.01). 


0.6 5 


Survival 


0.4 + 


0.2 + 


Figure 3 compares the survival functions for 
juvenile and wild ringtails at Many Dam Reserve. The 
relatively low survival function for juveniles is highly 
significant ( Chi-squared = 13.069, P<0.001). 


Predation 
The fates of all 79 resident ringtails radio- 
tracked are shown in Table 1. Predation by introduced 


—*— SURVIVAL Adult 
—®— SURVIVAL Juvenile 


150 200 250 
Week 


Figure 3. Comparison of Kaplan-Meier survival functions (modified for staggered entry) for adult ringtails 
(top curve, n=44) and juveniles (n=34) at Manly Dam Reserve. Chi-squared = 13.069, P<0.001. 


186 


Proc. Linn. Soc. N.S.W., 124, 2003 


B. SMITH, M. AUGEE AND S. ROSE 


Table 1. Fate of Common Ringtail Possums tracked in Manly Dam Reserve 1994-1999 and Ku- 


ring-gai Chase National Park 1990-1994 


FATE 


Killed by fox 

Killed by cat 

Radio-signal lost, reason undetermined 
Killed on road 

Still alive at end of study (collar removed) 
Transmitter failed, seen but not recaught 
Killed by python 

Killed by goanna 

Killed by raptor 

Killed by unknown predator 

Killed in bush fire 

Total 


carnivores (foxes and cats) was heavy; 80% of known 
causes of death (Table 1). 


Nest sites 

During this study a total of 2,907 daytime 
positions was determined by radio tracking. Of these, 
88% were in dreys, 6.1% were on the ground and 3.6% 
were in tree hollows. 

Dreys were constructed in tree/shrub species 
listed in Table 2 with the majority in Banksia ericifolia. 
On average, individual ringtails at Manly Dam Reserve 
used seven different dreys during the period they were 
radio-tracked. The actual number ranged from one to 
21 depending mainly on the length of time any one 
possum was tracked. 

The figure for usage of ground positions 
(6.1%) is skewed by one individual (M275) that was 
located 70 times on the ground after having dispersed 
to an area of heathland where there were no shrubs, 
bushes or small trees sturdy enough to support a drey. 


Manly Dam_ Ku-ring-gai 


Ku-ring-gai Introduced 


Wild 
28 18 39 
20 4 28 
12 8 27 
9 1 
4 
3 
1 3 5 
l 6 
1 2 
1 4 
4 1 
79 38 113 


If this animal is excluded, only 4% of the locations 
were on the ground. 


Sharing of nest sites 
Dreys were used and kept in repair by more 


than one individual ringtail. Table 3 lists single and 
multiple occupancies for all known nesting sites in one 
thicket. The thicket illustrated had the greatest use of 
any in the study area. Table 3 contains an example of 
nest sharing within a family, with each parent (F266 
and M295) sharing with each other and on separate 
occasions with offspring (M307 and M308). F266 also 
shared with her daughter F292, the father of which is 
unknown. 


Competition 
There is some evidence for exclusion as a 


result of competition. In two cases ringtails moved into 
an area immediately after an occupying ringtail died 
(M261 replaced M244, and F242 replaced F243). In 


Table 2. Characteristics of four thickets in the Many Dam study area. Common Ringtail Possums 
listed as occupants did not necessarily overlap in time. Area and floristic data from Newton (1997). 


OCCUPANTS THICKET 
AREA (ha 

M263, F266, M295, 0.165 

M307, M308 

F242, M254, M279, 0.12 

M300 

M290, M297, F298, 0.18 

F311 

F136, M246, F248, 0.275 

F249, M282 

F253, M264, M284, 0.16 

F287, M310 


Proc. Linn. Soc. N.S.W., 124, 2003 


SPECIES IN WHICH NESTS OCCURRED 

Banksia ericifolia, Kunzea ambigua and Corymbia gummifera 
B. ericifolia, K. ambigua, Hakea teretifolia, and E. punctata 
B. ericifolia, K. ambigua, H. teretifolia, and E. haemastoma 
B. ericifolia, K. ambigua, B. serrata, H. teretifolia, and E. 


haemastoma 


B. ericifolia, K. ambigua, Callicoma serratifolia, E. punctata 
and Leptospermum trinervium 


187 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


Table 3. Occupancy of dreys in a single thicket by Radio-collared ringtails 
during the period 1994-1999. Cases where occupancy was simultaneous 
are indicated by / and bold. *M307 and M308 are siblings; M295 is their 
father; F266 is their mother, ** F266 is the mother of F292 


Table 5 shows that some 
ringtails remain for extended 
periods, in some cases their 
entire life, in the same foraging 
area (thicket) in which they 


Drey designation Occupiers were born. 
Cla M259/F 132 5 aca “tear? 
Cl F242, M263, M261, M289 Distribution of nesting sites and 
C M259/F266, M263, F285 home_ranges 
€3 F266/M259 Fig. 4 is a plot of all 
C4 M295/F266, M308, M307 nesting sites on a map of the 
C5 M259, M263 study area. All nesting sites for 
C6 F285, M259, M263 any given individual are 
Gi M259 enclosed in the smallest 
C8 M263, M259 polygon that can be formed by 
@ M259, F266/M295, M263, M244, M261, F249, joining outer sites for that 
M279/F292, M295/M307/M308* animal to form an estimate of 
a oe M259, M263, M244 home range. The distribution is 
95 patchy across the study area 
oe ra | and the overlapping 
concentrations of home ranges 
ae aes WOES a Th correspond to the distribution 
C15 M263/F277 of thickets (Fig. 4). The thickets 
C16 F277 were composed primarily of 
C17 F266, M295, M263, M307/M308* Banksia ericifolia and Kunzea 
C18 F289 ambigua. Details of the 
C19 M259, F266/F292**, F289 floristics (from Newton 1997) 
C20 F266, M295 of four thickets in the study area 
C21 M295 are given in Table 2. Nest sites 
C22 M295/F266, M307 rarely occurred in large trees. 
C23 F266, M307/M308* Over the entire period of this 
C24 M295 study only five dreys were 
C25 F266/M307, M295 found in Eucalyptus or 
C26 M295 


another case, M275 dispersed from the foraging area 
he had occupied for four months after another male, 
M264, incorporated that area into his own. M275 
thereafter remained in his new foraging range. 


Movements 

Table 4 sets out all instances for ten 
individuals in which exploratory movements from one 
nest site resulted in the occupation of a new nest site 
more than 50 m away. The number of these movements 
for an individual varied from one (F245) to 13 (F296) 
(Table 4). It is important to note that 39 of the 62 
animals tracked for more than one month did not make 
any exploratory movements and remained within 50 
m of the point at which they were initially captured 
and collared. 

Forty-four percent of males tracked made 
exploratory movements while only 28% of the females 
did so. 


188 


Angophora spp. 


Reproduction 
When the pouches of females at the time of 


radio-collar replacement were examined, almost all 
were found to contain joeys. Although Pahl (1987) 
found numerous single births in southern Victoria, in 
this study all mothers with pouch young were found 
to be carrying twins; not one instance of single birth 
was found. Pouch young were attached to the posterior 
pair of nipples (ringtails have two pairs of nipples in 
the pouch). The annual distribution is shown in Fig. 5, 
from which it appears that breeding occurs throughout 
the year, with a peak of births in May and November, 
and a trough in January-March. 


DISCUSSION 


Mean survival of wild possums at Manly Dam 
Reserve (319 days) was greater than determined for 


Proc. Linn. Soc. N.S.W., 124, 2003 


B. SMITH, M. AUGEE AND S. ROSE 


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0 Ty KEY: 


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ee eS a ao Walking con Wocdlan 


Track 


xy Banksia v ~]| Heathland/ Riparian 
UN ericsfolia Low Open Uy, Open Forest 
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Figure 4. Home ranges (as defined in the text) of all ringtails radio-tracked at Manly Dam Reserve 1994- 
1999 superimposed on a vegetation map of the study area. Dots = nest sites. Vegetation communities are 
defined in “Methods” above. 


Proc. Linn. Soc. N.S.W., 124, 2003 189 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


Table 4. Examples of long distance movements by Common Ringtail 
Possums in Manly Dam Reserve. For each ringtail listed all known 
movements over 50 m are given. 


Animal Distance (m) Duration (days) _ Terminal event 


F242 55 (from a to b) 18 

55 (return to a) 14 

55 (return to b) 1140 killed by fox 
F245 110 741 killed by goanna 
M254 122 7 

105 14 road kill 
M261 50 36 

68 21 

53 14 

65 21 

64 539 

56 21 killed by fox 
M264 75 28 

110 764 killed by cat 
M275 122 370 

88 7 

65 78 transmitter failed 
M279 65 (a to b) " 

63 (return to a) Fi 

63 (return to b) 266 

94 72 | 

104 14 

89 1 roadkill 
F289 86 14 

61 7 

61 27 

74 39 

51 17 

61 42 

121 43 

83 293 killed by fox 
F296 110 14 

121 28 

135 (a to b) 7 

135(return to a) 34 

131 7) 

131 21 

145 U 

136 28 

159 21 

232 14 

293 7 

94 30 

373 1 killed by cat 
M317 88 28 

84 (a to b) 10 

84 (return to a) 4 killed by fox 


the Ku-ring-gai study. The 
major cause of death is 
predation by foxes and cats; 
80% at Manly Dam Reserve and 
76% at Ku-ring-gai Chase. 
Usage of ground nest sites is 
also similar in both studies; 6% 
at Manly Dam Reserve and 7% 
at Ku-ring-gai Chase. The most 
likely explanation for the 
apparently greater life 
expectancy at manly Dam 
Reserve is lower numbers of 
introduced predators, but there 
is no way to test this hypothesis 
with available data. 

Juveniles are clearly at 
greater risk than adults, having 
a significantly lower survival 
function (Fig. 3). 

The vegetation within the 
Manly Dam Reserve study area 
consisted mostly of low heath, 
with taller mature eucalypts and 
riparian vegetation along a 
creek line and Banksia/Kunzea 
thickets separated by low 
scrubby heath. Ringtails 
preferred the _ thickets, 
constructing dreys primarily in 
Banksia ericifolia and to a lesser 
extent in the low trees and 
sturdy bushes listed in Table 2. 
Some ringtails utilized hollows 
in the trees near the creek line 
and one individual utilized the 
heath, making several nests on 
the ground. 

In Ku-ring-gai Chase the 
figure for usage of tree hollows 
was about 33% (Augee et al. 
1996). In Manly Dam Reserve 
there are few large, mature trees 
and only 3.6% of daytime 
positions were in hollows. One 
animal also nested briefly in an 
arboreal termite mound about 5 
m up a eucalypt tree. 

At any given time, a 
ringtail would use several nest 


wild possums in the previous study at Ku-ring-gai _ sites, most often within a single thicket. Around these 
Chase (182 days; Augee et al. 1996). Likewise the sites the animal would forage over an area usually less 
survival functions for these two data sets (compared than 0.2 ha with a maximum diameter of 50 m. This 
in Fig. 2) indicate a significantly (p<0.01, Chi-squared foraging range was consistent regardless of the habitat,. 


= 10.227) lower survival function for wild possums in 


190 Proc. Linn. Soc. N.S.W., 124, 2003 


B. SMITH, M. AUGEE AND S. ROSE 


Table 5. Residency time of progeny that never left their natal sites. All were 
juveniles (<600 g) at time of collaring. Days are measured from time of collaring. 


Female Parent | Probable Male 


Subject 


M259 F249 

F274 F248 

F278* F242 

M279* F242 

M282 F248 

M284 F245 

F291 F245 

F296 F287 

M297 F298 

M307* F266 M295 
M308* F266 M295 
F311 F298 M290 
M318* Uncollared Uncollared 
F319* Uncollared Uncollared 


being observed for animals living along the creekline 
amongst large trees as well as in thickets. 

Like many arboreal mammals (e.g. squirrels, 
McDonald 1984), ringtails are sedentary. At Manly 
Dam Reserve 39 of the 62 animals tracked remained 
within the foraging range where they were first caught. 
One female (F273) remained in the same foraging 
range for 582 days until being killed by a fox. 

The sedentary nature of the species is further 
evidenced by data obtained from 14 individuals for 
whom their female parent and place of birth were 
known (Table 5). Of the seven ringtails that were killed 
within the natal area (i.e. within 50 m of the site of 
their birth), four had lived there for extended periods 
from the time of being collared (293, 519, 638 and 
967 days). 

Some ringtails did move out of their foraging 


Natal Area 


range, occasionally 
establishing a new nest 
site more than 50 m from 
the previous nest site. In 
a few instances the 
animal quickly returned 
(F242, M279, F296 and 
M317 in Table 4), 
however most such 
movements resulted in 
the establishment of a 
new foraging range. For 
example F312 remained 
in the foraging area 
where she was first 
caught for 90 days, then 
moved about 800 m to a 
new foraging range, 
remaining there 174 
days. At that time her collar was removed since this 
unusually large dispersal movement had taken her out 
of the study area. 

Dispersal movements, resulting in the 
establishment of a new foraging range, were made by 
both sexes, although more males (15) did so than 
females (8). However, the animal making the greatest 
number and longest dispersal moves was a female 
(F296, Table 4). The animal making the second greatest 
number of moves was also a female (F289, Table 4). 
The reasons for such shifts in foraging range are 
unclear. 

In many mammalian species there is a pattern 
of dispersal by juveniles (McDonald 1984). Some of 
the ringtail dispersals observed in this study were by 
juveniles (e.g. M264, M279, F296 and M317 in Table 
4). However dispersal of juveniles was not a consistent 
pattern as only 10 out of 26 
(38%) of juveniles tracked for 
more than a month made 


Killed in natal 
area or moved 


ASSSANASSASAARSAA 


dispersal movements. Of the 
juveniles with known parents 


(Table 5), three (M282, M279 


and M259) remained within the 


parental foraging range after 


BIRTHS 


the death of the parents. As can 
be seen from Table 3, M307 


and M308 remained in 


association with both parents. 


O-NWHAAHON @W O 


Figure 5. Annual distribution of births of Common Ringtail Possums at 


Manly Dam Reserve. 


Proc. Linn. Soc. N.S.W., 124, 2003 


One female of known parents 
(F296) made frequent moves 
greater than 50 m from the 
parental foraging area, usually 
returning, until killed by a cat 
280 m from the parental 
foraging area (at which time 


191 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


both parents were still alive). 

On the other hand, most ringtails in this study 
were mutually tolerant with considerable overlap in 
home range (Fig. 4) and foraging range (Table 2). 
Individual dreys were used by as many as 11 different 
individuals (Table 3). Simultaneous occupancy 
occurred (Table 3), usually by adult males with adult 
females and rarely two males. We did not observe 
simultaneous occupation of a drey by two adult 
females, although on one occasion the same drey was 
used by two adult females (F242 and F243) on different 
nights over a period of 3 weeks . We observed many 
instances of females sharing with joeys that were too 
small to radio-collar. In the only instance where we 
were able to track parents and their offspring 
simultaneously (F266 and M295, parents of M307 and 
M308), they were found to frequently share nest sites 
(see Table 3). The degree to which this familial 
tolerance continues as the juveniles reach maturity is 
unknown. 

Although there was no evidence that any 
movements made by the ringtails were related to 
predation, the majority of ringtails in this study, as in 
the study carried out earlier in Ku-ring-gai Chase 
(Augee at al. 1996), were killed by predators, usually 
foxes or cats (Table 1). While it is possible that deaths 
due to “unknown predators” in the Ku-ring-gai Chase 
study might have been due to dogs, we feel it unlikely 
that any deaths were due to dogs but misidentified at 
Manly Dam Reserve. Dogs are not allowed in the 
reserve unless on a lead and this rule is actively policed 
and well respected by local residents. During the entire 
course of the study we saw only one dog off the leash. 


CONCLUSION 


Ringtail possums are usually sedentary, 
remaining within a foraging range of approximately 
50 m diameter in the Manly Dam Reserve study area. 
They may on occasion move beyond this range, 
although the reasons for such long distance movements 
are unknown. They are probably all exploratory, but 
most result in the establishment of a new foraging range 
and can be considered dispersal movements. 
Presumably the new foraging range provides improved 
feeding or reproductive resources or less competition 
with conspecifics. Predation by foxes and cats was 
severe throughout the study area. 


REFERENCES 


Augee, M.L., Smith, B. and Rose, S. (1996). Survival of 
wild and hand-reared ringtail possums 


192 


(Pseudocheirus peregrinus) in bushland near 
Sydney. Wildlife Research 23, 99-108. 

Kaplan, E.L. and Meier, P. (1958). Nonparametric estimation 
from incomplete observations. Journal of the 
American Statistical Association 53, 457-481. 

McDonald, D. (ed.). (1984). The encyclopedia of mammals. 
Unwin Hyman, London 

Newton, T. (1997). Foraging range in the common Ringtail 
Possum (Pseudocheirus peregrinus). Honours 
Thesis, University of NSW (held by the School of 
Biological, Earth and Environmental Sciences, 
UNSW, Sydney 2052). 

Pahl, L.I. (1987). Survival, age determination and population 
age structure of the Common Ringtail Possum, 
Pseudocheirus peregrinus, in a Eucalyptus 
woodland and a Leptospermum thicket in southern 
Victoria. Australian Journal of Zoology 35, 625- 
639. 

Pollock, K.H., Winterstein, S.R., Bunck, C.M. and Curtis, 
P.D. (1989). Survival analysis in telemetry studies: 
the staggered entry design. Journal of Wildlife 
Management 53, 7-15. 

Rose, S., Augee, M.L. and Smith, B. (1994). Predation by 
introduced foxes on native mammals: a study of 
fox scats from Ku-ring-gai Chase National Park, 
Sydney. In “Proceedings of the 1994 Conference 
of the Australian Association of Veterinary 
Conservation Biologists” (Eds A.W. English and 
G.R. Phelps). pp. 111-116. Australian Association 
of Veterinary Conservation Biologists: Sydney. 

Smith, B. (1994). Caring for possums. Kangaroo Press, 
Sydney. 


Proc. Linn. Soc. N.S.W., 124, 2003 


B. SMITH, M. AUGEE AND S. ROSE 


APPENDIX I 
Details for all ringtails in the Manly Dam Reserve study 


No. Sex Weight Collar Collar Survival Fate 
attached retrieved (days) 

242 F 710g 7/9/94 3/12/97 1181 FOX 

243 F 995g 7/9/94 4/5/95 238 CAT 

244 M 7852 7/9/94 27/4/95 231 CAT 

245 F 800g 7/9/94 17/12/97 1196 GOANNA 

246 M 900g 14/9/94 26/9/96 742 FOX 

247 F 940g 14/9/94 26/7/95 314 FOX 

248 F 750g 21/9/94 29/8/96 708 CAT 

249 F 880g 21/9/94 8/8/96 687 FOX 

250 F 920g 21/9/94 13/9/95 356 CAT 

251 F 950g 21/9/94 27/9/95 370 CAT 

Dy) M 770g 12/10/94 6/4/95 175 CAT 

USS) F 780g 12/10/94 6/4/95 175 FOX 

254 M 810g 7/11/94 10/8/95 275 ROADKILL 

255) M 870g 7/11/94 23/3/95 135 TX.Failure 

256 F 400g 7/11/94 30/3/95 142 ROADKILL 

Dil, F 930g 7/11/94 30/11/94 22 CAT 

258 M 420g 16/11/94 8/2/95 83 CAT 

259 M 550g 11/1/95 10/10/96 638 CAT 

260 M 800g 11/1/95 22/2/95 4] ROADKILL 

261 M 700g 11/1/95 13/11/96 672 FOX 

262 M 700g 15/2/95 20/5/95 93 ROADKILL 

263 M 675g 15/2/95 26/9/96 588 CAT 

264 M 620g 13/4/95 23/4/98 1106 CAT 

265 M 940g 4/5/95 29/6/95 39 SIGNAL LOST 

266 F 700g 11/5/95 27/5/98 1112 CAT 

267 M 890g 11/5/95 5/6/96 389 CAT 

268 M 720g 12/7/95 2/8/95 20 RAPTOR 

269 F 940g 12/7/95 2/8/95 20 FOX 

270 M 930g 12/7/95 21/5/98 1005 FOX 

271 M 995g 19/7/95 11/10/95 83 FOX 

OD) F 550g 19/7/95 9/8/95 20 CAT 

DIB) F 760g 9/8/95 20/3/97 588 FOX 

274 F 595g 11/10/95 15/11/95 35 FOX 

275 M 830g 8/11/95 8/10/97 700 TxEXPIRE 

276 F 675g 8/11/95 15/5/96 199 SIGNAL LOST 

DUT F 675g 15/11/95 13/12/95 28 FOX 

278 F 550g 3/1/96 31/1/96 28 ROADKILL 

279 M 580g 17/1/96 19/6/97 519 ROADKILL 

280 M 970g 17/1/96 26/6/96 159 FOX 

281 M 920g 17/1/96 28/2/96 42 SIGNAL LOST 

282 M 625g 13/3/96 6/11/98 967 CAT 

283 M 460g 26/6/96 22/8/96 i FOX 

284 M 510g 26/6/96 10/3/97 258 FOX 

285 F 570g 22/8/96 21/9/96 30 FOX 

286 F 860g* 29/8/96 13/2/97 168 PYTHON 

287 F 720g 5/9/96 19/2/98 532 SIGNAL LOST 


Proc. Linn. Soc. N.S.W., 124, 2003 193 


MOVEMENTS OF COMMON RINGTAIL POSSUMS 


No. Sex Weight Collar Collar Survival Fate 

attached retrieved (days) 
continued 
288 M 560g 12/9/96 7/11/96 56 FOX 
289 F 850g 26/9/96 28/1/98 489 FOX 
290 M 900g 17/10/96 14/10/98 726 SIGNAL LOST 
291 F 455g 29/12/96 5/2/97 ai SIGNAL LOST 
292 F 460g 8/1/97 13/2/97 35 SIGNAL LOST 
293 M 440g 15/1/97 19/6/97 155 FOX 
294 F 490g 26/2/97 26/6/97 120 CAT 
295 M 580g 20/3/97 8/12/98 509 SIGNAL LOST 
296 F 420g 27/3/97 22/1/98 300 CAT 
297 M 300g 24/4/97 12/3/98 352 FOX 
298 F 890g 1/5/97 29/12/99 971 RELEASED 
299 M 520g 29/5/97 19/6/97 20 FOX 
300 F 375g 13/8/97 27/8/97 13 ROADKILL 
301 F 725g 20/8/97 3/11/99 804 FOX 
302 F 700g 20/8/97 16/9/98 385 CAT 
303 F 625g 20/8/97 29/12/99 860 RELEASED 
304 M 650g 20/8/97 1/1/98 133 Tx.FAILURE 
305 M 850g 27/8/97 10/12/97 118 SIGNAL LOST 
306 F 355g 15/10/97 26/10/97 10 FOX 
307 M 265g 10/12/97 30/9/98 293 CAT 
308 M 255g 10/12/97 27/5/98 167 ROADKILL 
309 M 450g 21/1/98 18/11/98 300 SIGNAL LOST 
310 M 475g 11/2/98 6/11/98 119 SIGNAL LOST 
311 F 340g 30/9/98 30/9/98 90 ROADKILL 
312 F 320g 6/11/98 12/2/98 173 RELEASED 
313 F 450g 9/2/98 30/9/98 Dil SIGNAL LOST 
314 F 350g 14/10/98 11/11/98 Di FOX 
315 F 245g 13/1/99 28/4/99 104 FOX 
316 M 300g 27/1/99 5/12/99 104 FOX 
Sil 7/ M 400g 24/3/99 30/6/99 97 FOX 
318 M 310g 4/7/99 29/12/99 265 RELEASED 
319 F 300g 4/7/99 30/6/99 83 FOX 
320 M 250g 6/2/99 23/6/99 20 CAT 


194 Proc. Linn. Soc. N.S.W., 124, 2003 


BOOK REVIEW 


BIRDS OF AUSTRALIA’S TOP END 


Denise Lawungkurr Goodfellow (2001) 


Scrubfowl Press, Parap N.T. 
RRP $A29.50 


I wondered at first if a review of yet another 
bird guide was appropriate for the Proceedings, 
however it seems to me that all natural scientists have 
some interest in birds. Even on a geology field trip it 
is hard to ignore the most obvious and varied daytime 
fauna. A field guide is a useful part of anyone’s field 
gear. The big three (Pizzey and Knight, Simpson and 
Day, and Slater and Slater) are of course Australia wide, 
but there are also a number of local guides and lists of 
varying quality to supplement them. In some localities 
the landscape and the avifauna are so unique that a 
local guide is pretty near essential. The Northern 
Territory, or more specifically the Top End, is one such 
place. 

This book is primarily aimed at visitors to the 
top end who come specifically to see birds. Australian 
and overseas twitchers looking to add to their bird lists 
will find this book invaluable. Unlike the big three, it 
provides specific instructions on where to find 
particular species and the best time and means. 

The author has a great interest in birdwatching 
tourism, which is discussed in the opening pages, and 
this book does much to encourage and facilitate this. 
There is a section on safety, which includes some hints 
for driving safely, most of which are usually ignored 
by Top Enders themselves. 

For the locals there is a section on how to attract 
birds to your garden. 

For the beginner or the student looking for a 
general reference on bird biology there are sections 
on bird evolution, physiology, behaviour and sex. 

For everyone there is a concise discussion of 
the problems of conservation in the Top End. 

The above features are found in the 
introduction, but of course the heart of the book is the 
species descriptions, backed up by watercolour 
illustrations by the author. For those few species which 
I have observed myself in the Top End, the colours 
appear correct. However no bird guide ever written 
provides colours acceptable to everyone and the user 
must always be aware that variation is the theme of 
life. 


Proc. Linn. Soc. N.S.W., 124, 2003 


Besides describing the appearance, of both 
sexes where necessary, species accounts usually 
include details of flight, calls, similar species, breeding, 
habitat, range where found, alternative common names 
and birdwatching hints. In many cases there is also 
the Kunwinjku name. This is an Aboriginal language 
understood widely in the Top End. This of course has 
nothing to do with species identification but has 
everything to do with Denise’s strong respect for 
Aboriginal culture and knowledge of the Australian 
environment. She has included some of this in notes 
which follow many species accounts. These notes, as 
well as frequent footnotes and “author’s notes” make 
this book great reading as well as useful. The “author’s 
notes” usually relate Denise’s own experiences with 
the bird in question. 

Of great use to the occasional bird watcher is 
the inclusion of description of higher taxa such as 
families. This level is usually left out of field guides 
and may not be essential for identifying species, but it 
is of great use in understanding relationships and 
overall characteristics. 

With each entry there are references to the 
matching entries in both Pizzey and Knight (“Field 
Guide to the Birds of Australia”) and Simpson and Day 
(also titled “Field Guide to the Birds of Australia’); 
not only a generous feature but a very useful one except 
for those like me who are loyal to “The Slater Field 
Guide to Australian Birds”. 

This is a unique field guide and represents an 
incredible amount of work by the author, Denise 
Lawungkurr Goodfellow, who still finds time and 
enthusiasm to show visitors the avifauna of the Top 
End. A trip to the Leanyer Sewage Ponds with Denise 
is an experience, like the book, not to be missed. 


M.L. Augee 


Sydney 
November 2002 


195 


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198 Proc. Linn. Soc. N.S.W., 124, 2003 


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PROCEEDINGS OF THE LINNEAN SOCIETY OF NSW 
al ‘wii 


1205 6 


ITUTIO 


NUN 


Issued 31 January 2003 
CONTENTS 


1 Kimberley A. Smith 
Larval distributions of some commercially valuable fish species over the Sydney continental shelf 
13 Geoff Williams 


New distribution and biological records for native dung beetles, in the tribe Scarabaeini, from 
northern New South Wales 

Wve Laurence Mound and Geoff Williams 
Host-plant disjunction in a new species of Neohoodiella (Insecta, Thysanoptera, 
Phlaeothripinae),with notes on leaf-frequenting thrips in New South Wales subtropical rainforests 

29 Y.Y. Zhen, |.G. Percival and J.R. Farrell 
Late Ordovician allochthonous limestones in Late Silurian Barnby Hills Shale, central western 
New South Wales 

53 W.B. Keith Holmes 
The Middle Triassic megafossil flora of the Basin Creek Formation, Nymboida Coal Measures, 
New South Wales, Australia. Part 3. Fern-like foliage 

109 A.M. Pinder 
First Australian records of three species and two genera of aquatic Oligochaetes (Clitellata: 
Annelida) 

115 ~ Kevin Warburton and Christine Madden 
Behavioural responses of two native Australian fish species (Melanotaenia duboulayi and 
Pseudomugil signifer) to introduced Poeciliids (Gambusia holbrooki and Xiphophorus helleri) in 
controlled conditions 

125 |.D. Lindley 
Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea: 
Clypeasteroida : 


17 1.D. Lindley 
Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New Guinea: Regularia 
153 1.D. Lindley 
Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Rapua New Guinea: 
Spatangoida 


163 Luke Strotz 
Holocene Foraminifera from Tuross Estuary and Coila Lake, south coast, New South Wales: A 
preliminary study 

183 Barbara Smith, Michael Augee and Stefan Rose 
Radio-tracking studies of Common Ringtail Possums, Pseudocheirus peregrinus, in Manly Dam 
Reserve, Sydney 

195 Book review: Birds of Australia's Top End. 

197 Instructions for authors 


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February 2004 


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EDITORIAL 


This volume consists of three parts. The first part contains general contributions. The second part contains 
research papers and review papers arising from the symposium “Monotreme III” held by the Linnean Society of 
NSW and the Australian Mammal Society at the University of Sydney in July 2003. The third section contains 
book reviews and an obituary to Merv Griffiths, the pre-eminent monotreme biologist of all time. 


The publication of this volume has been delayed by the preparation of the papers from “Monotreme III” and is 
covered by subscriptions and membership fees for 2003. 


Intending authors should read the summary of “Instructions for Authors” at the back of this volume carefully. 
More details are available in the full version available at the Society’s web site or from the Secretary. The 
preparation of this volume has been prolonged and made difficult by the failure of some authors to provide 
figures and tables in the format required. In order to keep the costs at a level which allows our Society to 
continue publication, we set the journal completely ourselves. Therefore we do not have the flexibility of large 
commercial publishers. We can only deal with figures as photographs, original line drawings or .TIF files. Jpeg 
files for example are useless in our system. Auto-formatting and track changes are a disaster, as are tables and/ 
or figures that have been put inside the text. To date we have taken the time to re-set and sometimes re-scan 
figures, however in future we will apply the policy that final copy not prepared in accordance with the instructions 
will simply be returned and held over if necessary until the next issue. 


M.L. Augee 
Editor 


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a 


Review of Australian Cave Guano Ecosystems with a Checklist 
of Guano Invertebrates 


TimotTHy Moutps 


Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, University 
of Adelaide, North Terrace, Adelaide 5005. 
timothy.moulds @ adelaide.edu.au 


Moulds, T. (2004). Review of Australian cave guano ecosystems with a checklist of guano invertebrates. 
Proceedings of the Linnean Society of New South Wales 125, 1-42. 


This work provides a check-list of all invertebrate species known, or believed to be, associated 
with cave guano in Australia. A total of 240 species in 121 families, representing 25 orders is listed. These 
species inhabit 60 karst areas in all mainland states of Australia and Christmas Island (Indian Ocean). 
Comprehensive assessment of all available records (published and in collections) show that the distribution 
of several species is more extensive than previously believed. It is unknown whether this is because of 
inadequate identification of specimens, poorly defined taxonomy or unrecognised intra-species variation 
due to a lack of specimens. Twenty species from five orders show restricted distributions and guano 
dependence, although endemic status can not yet be assigned. Amongst these species, eight pseudoscorpions 
and eight Coleoptera are distributed across several mainland states and Christmas Island. 


Manuscript received 2 July 2003, accepted for publication 22 October 2003. 


Keywords: Arthropoda, Australia, biospeleology, cave, checklist, ecosystem, guano, invertebrate. 


INTRODUCTION 


Australian cave guano ecosystems are poorly 
known with only a few communities studied in detail 
(e.g. Richards 1971; Harris 1973; Bellati et al. 2003). 
Previous studies concerned with the terrestrial 
cavernicolous fauna of Australia have mentioned 
species associated with guano, but have provided little 
in the way of detail with regard to the ecology of 
specialised guano communities and species. This paper 
seeks to synthesise the knowledge of guano ecosystems 
and communities in Australian caves. A review of 
guano ecosystems and habitats precedes a checklist of 
all species known to be associated with Australian cave 
guano deposits. 

Populations of cavernicolous animals are 
usually small because of limited food supplies. 
However, caves containing guano differ fundamentally 
because there is a virtually unlimited food supply, 
commonly resulting in large animal populations. 
Guano in caves is deposited by bats, birds, orthopterans 
(crickets and grasshoppers), and small mammals, with 
each type of guano sustaining a unique assemblage of 
taxa. Guano deposits are extremely variable, unlike 
other cave habitats, and consist of numerous micro- 


habitats differentiated by fluctuating temperature, 
moisture, and pH. Guano ecosystems contain obligate 
guano-dwelling organisms (guanobites), opportunistic 
guano-dwelling animals (guanophiles), and transient 
guano-using animals (guanoxenes) (Gnaspini and 
Trajano 2000). The basis for many guano ecosystems 
is the numerous species of fungi and bacteria that can 
grow on guano, even in complete darkness. 


Cave food sources 

Cavernicolous populations are dependant for 
their survival upon energy inputs into cave systems. 
These inputs can vary widely, with availability of food 
usually being the primary limiting factor (Peck 1976). 
Inflowing streams and periodic floodwaters introduce 
significant amounts of zooplankton, accidentals, and 
organic debris that, for many cave ecosystems, 
represent the main energy inputs (Peck and 
Christiansen 1990; Humphreys 1991). Tree roots 
penetrating the roofs and walls are another important 
energy source found commonly in tropical caves and 
lava tubes (Hoch 1988; Hoch and Howarth 1999). 
Dead animals can be a source of food for scavengers 
near cave entrances (Richards 1971). Accidentals 
wandering in from cave entrances also provide a food 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


source, although this is generally periodic in nature 
and inconsistent in quantity. 

For the most part cave environments are 
generally depaurporate in food and consequently are 
sparsely populated with cavernicolous animals. 
However, caves containing guano deposits differ 
substantially because they have a virtually unlimited 
food supply. When present, guano from bats, birds, 
and Orthoptera (crickets and grasshoppers) generally 
forms the major energy source (Park and Barr 1961; 
Poulson 1972; Martin 1977), with large, varied and 
unique ecosystems often revolving around such 
deposits. 


SOURCES AND DIVERSITY OF CAVE GUANO 


Cave guano deposits from specific sources 
can each possess a unique assemblage of taxa (Horst 
1972; Poulson 1972). Throughout the world’s 
biogeographic provinces different taxa are responsible 
for being the most important guano producers. The 
most widespread and common guano is that produced 
by bats and these deposits are generally the largest in 
volume. The spatial and temporal deposition of bat 
guano differs from tropical to temperate caves. Cave- 
dwelling bats in temperate regions show an annual 
cycle of occupancy over summer months when pups 
are born, before colonies disperse to cooler, wintering 
caves where they enter torpor. This annual cycle results 
in large amounts of guano deposited over summer 
months and then a cessation of guano input for 
approximately seven months. In contrast, tropical caves 
generally show constant bat occupancy rather than an 
annual cycle and less congregation of individuals due 
to warmer ambient temperatures. Gnaspini and Trajano 
(2000) note that many bat populations in tropical Brazil 
are commonly nomadic, resulting in roaming colonies 
varying their location in an irregular and non-seasonal 
fashion. This results in non-continuous deposition. The 
diet of bats (either haematophagous, insectivorous, 
frugivorous, or nectarivorous) also influences the 
composition of guano piles and hence the associated 
guanophilic communities (Gnaspini 1992; Ferreira and 
Martins 1998, 1999). Large populations of the common 
vampire bat (Desmodus rotundus Geoffroy) 
predominate in Brazilian karst near inhabited areas, 
due to large numbers of domestic livestock resulting 
in haematophagous guano deposits. Guano from non- 
haematophagous bats is absent, or greatly reduced as 
vampire bats exclude other bat species, thus changing 
the guanophilic communities present. 

Birds are common guano producers in the 
northern parts of South America, the Caribbean and 


tropical caves of south-east Asia. Cave-dwelling birds 
nest in the dark zone, providing an important energy 
resource for many cavernicolous animals. Cave- 
dwelling birds in South American and Caribbean caves 
include guacharos (Steatornis caripensis Humboldt) 
(Snow 1975; Gnaspini and Trajano 2000). This bird 
discards palm seeds, sometimes with flesh still 
attached, and deposit droppings in caves, thus 
providing a wide range of organic matter for 
cavernicolous animals. Because of the presence of 
discarded seeds, some taxa associated with seeds and 
detritus, such as lygaeid bugs are found only in guano 
of this type. Swiftlets (Aerodramus spp) nest in the 
entrance and dark zones of tropical caves in south- 
east Asia, northern Australia and the Pacific and are 
insectivorous (Medway 1962). These birds also support 
a range of guanophilic taxa in the caves of Christmas 
Island (Humphreys and Eberhard 2001). Richards 
(1971) reported that droppings from several species 
of birds nesting in the entrance zone of Nullarbor Plain 
caves support a wide variety of cavernicolous animals. 

Rhaphidophorid crickets are often important 
producers of guano in temperate caves such as those 
of the Nullarbor Plain (Richards 1971). The sometimes 
large populations of these crickets can accumulate 
sizeable guano deposits in caves. These deposits are 
important as few other food sources exist in areas such 
as the Nullarbor Plain because the low mean rainfall 
limits organic flood debris and bat populations are 
generally small. Rhaphidophorid guano is also utilised 
in Mammoth Cave, Kentucky, where it is widely 
dispersed through the cave system (Howarth 1983). 

Small mammals are often significant guano 
producers in temperate zones of North America. The 
guano of porcupines (Erethizon dorsatum L.) is 
reported by Calder (1965) to support a community of 
collembolans and mites active throughout the year in 
Frenchman’s Cave (Hants County, Nova Scotia, 
Canada). Cave rats (Neotoma spp), navigate using 
urine trails (Howarth 1983). Although common in the 
caves of the Canadian Rockies and Vancouver Island, 
their faeces are mostly unusable as a food source due 
to the high ammonia content from systematic urination 
at these sites (Trapani 1997). 


GUANO ECOSYSTEMS AND FOOD WEBS 


Guanobites are animals that require the 
presence of guano for survival. They will only feed 
on guano and will not use other food sources within 
caves. Although guanobitic species are occasionally 
found on other substrates in caves as they move 
between discontinuous guano deposits, they do not feed 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


or reproduce on these substrates (Gnaspini and Trajano 
2000). When guano deposition is seasonal (e.g. bat 
maternity caves), guanobites will commonly become 
quiescent until bats return and restore fresh guano 
input. Other guanobite populations crash when guano 
input ceases and then quickly reproduce when guano 
input recommences. 

Guanophiles use guano resources 
opportunistically and are able to complete their entire 
life cycle using the guano substrate. Guanophiles will 
however utilise other cave food resources when 
available and do not have to rely upon guano to feed 
or reproduce. Abundance of guanophilic animals will 
decrease if fresh guano is not available, simply due to 
food limitation, but individuals will attempt to exploit 
other food resources to survive until fresh guano is 
available. Troglobites and troglophiles that have a 
generalist role in epigean ecosystems are classified as 
guanophiles if they utilise guano when available, even 
though they are capable of surviving subterranean 
habitats without this resource. 

Guanoxenes will exploit a guano resource for 
feeding or reproduction but require other substrates 
within a cave to complete their life cycle (Gnaspini 
and Trajano 2000). Guanoxenes can be either 
troglobites, troglophiles or trogloxenes (Gnaspini and 
Trajano 2000). 

The cyclical nature of many guano deposits 
resulting from the annual breeding cycle of bats, leads 
to a similar cycle in arthropod abundances. Low 
population numbers of many species reflect changes 
in micro-habitat conditions resulting from the cessation 
of fresh guano deposition and lower air and guano 
temperatures. Guano communities decrease in numbers 
as many species stop breeding until the food supply 
(i.e. fresh guano) is restored. This has been observed 
in the mite Uroobovella coprophila Womersley, which 
is quiescent during winter months in Carrai Bat Cave, 
northern New South Wales (Harris 1971). 

Arthropods in guano communities feed either 
directly on guano or fungus growing on guano deposits 
and these in turn support a number of predators 
scavengers and omnivores (Gillieson 1997). 
Generalised guano food webs have a guano source 
directly supporting a range of guanivores including 
Phoridae (Diptera), Anobiidae (Coleoptera), Tineidae 
(Lepidoptera), Collembola and mesostigmatid mites 
(Acarina). Predators that prey upon these consumers 
include spiders, pseudoscorpions, beetles and 
opiliones. Specialised parasites and parasitoids are also 
active in many guano ecosystems. Braconid wasps 
(Hymenoptera) are found in many Australian guano 
caves and parasitise the larvae of Monopis spp 
(Lepidoptera: Tineidae). The larvae of the guanobite 


Proc. Linn. Soc. N.S.W., 125, 2004 


Derolathrus sp. (Coleoptera: Jacobsoniidae) are 
parasitised by small myrmarid wasps (Hymenoptera). 
Parasitic relationships in guano ecosystems are 
generally poorly understood and further research will 
undoubtedly reveal many more examples. Some of the 
most numerous taxa associated with guano deposits 
are mites (Acarina), particularly from the families 
Gamasidae, Actinedidae, Oribatidae and Armadillidae 
(Womersley 1963a, b; Gnaspini and Trajano 2000). 
Extremely high numbers (>33 million/m?) have been 
recorded on fresh guano (Harris 1973; Bellati 2001). 
Guanivores from all biogeographic regions are 
taxonomically similar, usually belonging to the same 
families. Differences, however, are found among the 
predators of guanivore communities and are often 
represented by taxa from different families depending 
on the biogeographical region (Gnaspini and Trajano 
2000). 


Bat guano micro-habitat variation 

Guano environments are extremely variable, 
consisting of numerous micro-habitats when compared 
with the majority of subterranean habitats (Harris 
1970). Bat guano deposits have been found to exhibit 
variable temperature of both the ambient air above 
deposits and within deposits (Harris 1970). In addition, 
the relative humidity,CO concentration, and ammonia 
concentration also change when bats occupy a cave 
due to their breathing and urine (Decu 1986). 
Variations in pH can be extreme, resulting in strong 
differentiation between fresh and old guano deposits. 
The annual cycle of bat roosting adds a temporal 
component to many guano deposits and also serves to 
alter air temperature in roosting chambers. Bat maternal 
chambers are especially variable when extremely large 
numbers of bats enter a chamber on an annual basis to 
birth young (Harris 1970). 

Large numbers of bats can raise the air 
temperature in a chamber by up to 10°C. This effect is 
most prevalent in high-domed chambers where heated 
air is trapped, but Harris (1970) also noted small 
increases in air temperature close to guano piles of up 
to 1.4°C due to heat released from guano breakdown. 
Increased air temperature of up to 12°C has also been 
noted in Cuban caves where large numbers of the leaf- 
nosed bat, Phyllonycteris poeyi Gundlach, roost (Decu 
1986). This temperature increase can act as a barrier 
for colonisation by generalist cavernicolous 
invertebrate species, but allows guanophilic and 
guanobitic populations to reach large numbers. 

Temperature within a guano pile can increase 
significantly with depth. Temperatures 5 cm below the 
surface of guano piles in Carrai Bat Cave, New South 
Wales are 1.7°C higher compared with surface 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


temperatures, and 15 cm below the surface 
temperatures are 3.0°C higher (Harris 1970). Surface 
guano temperatures have also been reported to increase 
by 9.3°C, and these increases in both surface and 
subsurface temperatures were attributed by Harris 
(1970) to the increase in the metabolic rate of the 
organisms inhabiting the guano pile. The initiation of 
growth and reproduction of mites in guano may be 
linked to the increase in temperature associated with 
bat occupation of a chamber (Harris 1971). 

Varying water content of guano due to 
desiccation with increasing age, results in noticeable 
micro-habitat differentiation. Fresh guano collected 
from the tops of piles in Bat Cave (U2), Naracoorte, 
South Australia, has been measured at up to 85% water 
by weight (Moulds 2003). Guano from the base of piles 
is a lighter grey colour due to desiccation and can 
contain as little as 6% water by weight (Moulds 2003). 
Guano moisture content increases with the birth of pups 
as their faecal matter is predominately liquid prior to 


being weened (approximately 6-8 weeks after birth 
for the large bent-wing bat Miniopterus schreibersii 
bassanii Cardinal and Christidis) (T. Moulds 
unpublished data). The surface of guano deposits 
commonly exhibit a patchwork appearance of dark 
moist areas and light grey drier areas. Different species 
within guano ecosystems prefer different micro- 
habitats. Richards (1971) noted the majority of 
guanophilic arthropods in Nullarbor Plain caves were 
only found in completely or partially dry guano. 
Guano shows a marked difference in pH 
between fresh and old deposits. Fresh guano is 
commonly basic, with the pH varying according to 
the volume of urine deposited with faeces. Fresh guano 
commonly has a pH of 8.5-9.0 that rapidly becomes 
acidic (5.0-5.5) with age and depth, although the centre 
of guano piles has a stable pH of around 4 (Harris 
1971). In bat maternity caves the pH of piles will 
gradually decrease over winter as no fresh guano is 
deposited. Data from Bat Cave (U2) (Naracoorte, 


Protochelifer cavernarum 
(Pseudoscorpionida) 


Ptinus exulans 
(Coleoptera) 


Shawella douglasi 
(Blattodea) 


Uroobovella coprophila 
(Acarina) 


Figure 1. Different distribution patterns of guano associated species across Australia. 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Table 1. Possibly endemic, guano dependent species in Australia. 


State Order Genus and Species Dependance Cave 
Cave Guano 

QLD Pseudoscorpionida Sathrochthonius webbi Tb Gp Holy Jump Lava Cave (BM1) 
QLD Coleoptera Choleva australis Tp Gp Royal Arch Cave (CH9) 
QLD Coleoptera Dermestes uter Tp Gp Royal Arch Cave (CH9) 
QLD Coleoptera Alphitobius diaperinus Tp? Gp? Bat Cleft (E6) 
QLD Coleoptera Omorgus costatus Tp Gp? Johannsens Cave (J1-2) 
QLD Coleoptera Anomotarus subterraneus Tp Gp Riverton Main Cave (RN1) 
NSW Pseudoscorpionida Oratemnus cavernicola Tp Gp? Jump Up Cave, Gray Range 
NSW Pseudoscorpionida Sundochernes guanophilus Tp2 Gb Fig Tree Cave (W148) 
NSW Pseudoscorpionida Tyrannochthonius cavicola Tp2 Gb Grill Cave (B44) 
NSW Acarina Neotrombidium gracilipes Tp2 Gb Fig Tree Cave (W148) 
NSW Acarina Hypoaspis annectans Tp Gp Carrai Bat Cave (SC5) 
Nullarbor Pseudoscorpionida Cryptocheiridium australicum Tp2 Gp Murra-E]-Elevyn Cave (N47) 
Nullarbor Isopoda Abedaioscia troglodytes Tb Gp? Pannikin Plain Cave (N49) 
Nullarbor Coleoptera Quedius luridipennis Tp? Gp Abrakurrie Cave (N3) 
VIC Pseudoscorpionida Pseudotyrannochthonius Tp2 Gp Mount Widderin Cave (H1) 

hamiltonsmithi 
VIC Coleoptera Achosia lanigera Tp? Gp Wilson Cave (EB4) 
SA Pseudoscorpionida Austrochthonicus cavicola Tp2 Gp Cathedral Cave (U12) 
SA Pseudoscorpionida Protochelifer naracoortensis Tp2 Gp Bat Cave (U2) 
WA Blattodea Paratemnopteryx atra Tb Gp Mines nr Marble Bar 
Christmas I Coleoptera Alphitobius laevigatus Unknown Gp Upper Daniel Roux Cave (C156) 


South Australia), show that late in spring, before guano 
deposition recommences, tops of guano piles can 
become acidic, occasionally as low as pH 5.0 (Moulds 
2003). The ever changing pH of guano piles due to 
age and urine content creates marked micro-habitats 
used by differing species. 

Micro-habitat variation of bat chambers is 
further complicated by the movement of bat roosts in 
a chamber within a breeding season. These movements 
are a response to avoiding unfavourable conditions 
caused by ammonia concentrations and high local 
temperatures (Poulson 1972). 


DISTRIBUTION, BIOGEOGRAPHY AND 
ENDEMISM 


This is the first checklist for Australian guano- 
associated invertebrates. The full geographic range of 
many guanobitic and guanophilic species can now 
easily be appreciated. Many species have been shown 
to have unexpectedly wide distributions, sometimes 
spanning several climatic regions. Several possible 
explanations exist for these patterns. The lack of 
systematic searching and collation of published 
records, and collections has resulted in a poor 


Proc. Linn. Soc. N.S.W., 125, 2004 


understanding of many species distribution and degree 
of endemism. This is commonly combined with a lack 
of accurate identification by taxonomic experts leading 
to the lumping of several similar species into one. 
Inadequate species definitions from groups requiring 
systematic revision will also result in species being 
artificially lumped or split (eg Diptera: Phoridae, David 
McAlpine, pers. comm. 2002). A lack of collections 
from most karst areas, both above and below ground, 
is the greatest problem, resulting in large gaps in 
distributions and a poor knowledge of variation within 
species. The paucity of records among some taxa also 
provides a focal point for future collecting priorities. 

The collation of this checklist has revealed 
associations of species across wide geographic regions. 
Figure la shows the extensive range of Protochelifer 
cavernarum Beier (Pseudoscorpionida) from Jurien 
Bay, Western Australia, across southern Australia and 
north to Undara Lava Tubes in northern Queensland. 
The distribution of Shawella douglasi Princis 
(Blattodea: Blattellidae) (Fig. 1b) is disjunct with 
records from northern New South Wales and Jurien 
Bay, Western Australia. This may be the result of 
misidentification, poor taxonomic description or a 
paucity of collecting between these localities, 
especially throughout northern Australia. Despite a 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


number of invertebrate collections from the Nullarbor 
karst no individuals have been recorded, possibly due 
to extremely small populations of troglobitic species 
and the extremely large size of the karst area concerned. 
Several species including Ptinus exulans Erichson 
(Coleoptera: Anobiidae) show very wide distributions 
from mid-north New South Wales across the Nullarbor 
Plain to the west coast of Western Australia (Fig. Ic). 
The distribution of U. coprophila (Acarina: 
Urodinychidae) (Fig. 1d) is directly linked to the 
distribution of maternal sites for the large bent-wing 
bat M. schreibersii. The single record of this species 
from Undara (north Queensland) may be spurious, a 
misidentification or an individual transported via 
phoresy, especially as no records exist between 
southern and northern Queensland despite large bat 
maternity caves around Rockhampton. These data raise 
further questions regarding the colonisation of guano 
deposits by invertebrates and the boundaries of 
possibly ill-defined species concepts. 

Endemic status of guano species, has, in the 
past been assigned without a full understanding of the 
distribution of Australian guano fauna. This is apparent 
for the maternal chamber of Bat Cave (U2), 
Naracoorte, where previous studies (Hamilton-Smith 
2000), identified ‘several endemic species’ to the 
maternal chamber or Bat Cave as a whole. This 
checklist has shown that Bat Cave contains only a 
single endemic species, Protochelifer naracoortensis 
Beier, and this pseudoscorpion may possibly be found 
in other caves in the continuous karst of the Otway 
Basin. Bat Cave does, however, form the most diverse 
guanophilic arthropod community in Australia. This 
highlights the amount of assumed knowledge 
concerning guano invertebrates in Australia and their 
distribution. The number of endemic species to specific 
bat caves is currently unknown but is almost certainly 
significantly lower than previously believed. Several 
species have been identified as possessing restricted 
distributions and guano dependence, although none 
can yet be positively identified as endemic (Table 1). 
The restricted distribution status of all species listed 
in Table | is tentative and more extensive collecting, 
both above and below ground, must be undertaken 
before distribution can be confirmed. This is especially 
true for troglophilic species as epigean occurrence of 
these species will effect their endemic status. The 
degree of a species’ guano dependence will also affect 
its endemic status and more ecological knowledge is 
required to confirm species habits. Species confined 
to single caves or isolated areas are more likely to be 
endemic when combined with guano dependence. 
Only Fig Tree Cave (W148) (Wombeyan, NSW) and 
Royal Arch Cave (CH9) (Chillagoe, QLD) are found 


to contain two species showing both restricted 
distribution and guano dependence (Table 1). 

The presence of nematodes is almost a 
certainty in guano caves as they are almost ubiquitous 
in every other habitat both above and below ground. 
Despite this the records of nematodes from guano are 
extremely limited primarily because the majority of 
caves and karst areas remain completely unsampled 
for these invertebrates. Nematodes play a potentially 
important role in the micro-habitat of guano piles and 
have been recorded in large numbers from overseas 
caves (Decu 1986). Nematodes are also believed to be 
one of the first colonisers of new bat caves, being 
deposited by in urine and faeces (Decu 1986). Further 
sampling of Australian cave guano will almost 
certainly reveal a greater diversity of species. Currently 
no free living nematodes have been recorded by the 
author from Bat Cave, Naracoorte despite several 
collection events. 

Currently no guano invertebrates are recorded 
from Tasmania, primarily due to the absence of cave- 
dwelling bats. The possibility remains however, that 
guano communities occur in orthopteran guano or 
other invertebrate guano deposits or even bird guano. 
The guanophilic mite Macrocheles tenuirostris Krantz 
and Filipponi was first recorded from mutton bird nests 
in Tasmania and has since been collected from bat 
guano in Victorian and New South Wales caves. 
Further field observations within Tasmanian caves may 
yet reveal these communities. 

Opportunities for future research in this field 
are vast with only limited knowledge existing for most 
karst areas. The ecological classification for many 
species is poorly known and this will only be achieved 
through increased observations in situ. The 
microbiology of guano deposits also remain very 
poorly known in Australian, as well as in overseas 
caves. Many karst areas remain completely unstudied 
biologically, especially with regard to the diversity of 
invertebrate guano communities. 


SYSTEMATIC CHECK LIST OF AUSTRALIAN 
GUANO INVERTEBRATES 


This checklist includes all Australian 
cavernicolous species found in association with guano 
from both caves and mines. Records have been 
compiled from the speleological literature (both 
scientific and amateur), unpublished records, and 
personal observations. Parasites of cave-dwelling 
mammals (bats) have been included as they are often 
found in guano, although their potential roles in guano 
ecosystems is currently unknown. Taxa are arranged 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


systematically by Phylum, Class and Order then 
alphabetically by Family. Undetermined taxa have 
been placed at the end of their respective order or 
family. Due to changes in taxonomy and higher 
systematics of many taxa the names and position of 
species can be uncertain. This checklist has adopted 
the most recent higher classifications attainable and 
many old names have been updated to reflect changes 
in the literature. Many groups in this checklist are in 
need of revision and so some species concepts may be 
altered in the future resulting in the splitting of some 
species and the lumping of others. This will obviously 
affect the distribution of species as presented in this 
work. 

Cave names and numbers following the 
Australian Karst Index (Mathews 1985), and are listed 
for all species’ records along with appropriate 
references. Records from caves in the Nullarbor Plain, 
southern Australia, have not been divided along state 
boundaries in order to reflect the extremely large and 
continuous nature of this karst area. Taxa previously 
considered to be obvious accidentals to cave 
environments have been excluded from this checklist. 

The following ecological classification is 
modified from Hamilton-Smith (1967), and Gnaspini 
and Trajano (2000), and is based on the degree of cave 
and guano dependence of taxa. Abbreviations are those 
used in the checklist. 


Trogloxene (Tx): an organism that regularly uses the 
cave environment for part of its lifecycle or as shelter 
but must leave the cave to feed and or breed. 

1* order Troglophile (Tp1): an organism that can 
complete its entire lifecycle within a cave but possess 


no specific adaptations to the cave environment and 
recorded in both epigean and hypogean habitats. 

2™¢ order Troglophile (Tp2): an organism that can 
complete its entire lifecycle within a cave but possess 
no specific adaptations to the cave environment and 
recorded only from hypogean habitats. 

Troglobite (Tb): obligate cavernicolous organisms 
that possess specific adaptations to the cave 
environment. 

Guanoxene (Gx): an organism that may use guano 
for reproduction and/or feeding but requires other 
substrates to complete its life cycle. 

Guanophile (Gp): an organism that inhabits and 
reproduces both in guano piles as well as other 
substrates within a cave. 

Guanobite (Gb): an organism that requires guano 
deposits to complete its entire life cycle. 

Bat Parasite (P): an animal that is an obligate bat 
parasite requiring bats to complete its lifecycle. 


Ecological classifications have been assigned 
to taxa wherever possible. These designations were 
made using available knowledge concerning 
behaviour, life history, and distribution within caves. 
However, information regarding species’ ecology was 
found to be lacking or minimal in most cases. Because 
of such constraints some taxa have not been assigned 
a guano classification. Further, information on other 
taxa was insufficient to confirm their association with 
guano ecosystems. Thus, taxa previously recorded only 
from guano caves, but without a confirmed association 
with guano, have been included for completeness even 
though some of these species may be unassociated with 
guano. 


Phylum Platyhelminthes 


Class Tubellaria 


Order undetermined 


Undetermined genus and species, Tx, Gx?. VICTORIA: Dickson Cave (M30), Murrindal (Yen and 


Milledge 1990). 


Phylum Nemathelminthes 


Class Nematoda 


Order Strongyloidea 
Trichostrongylidae 


Nycteridostrongylus unicollis Baylis, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 


(Hamilton-Smith unpublished data). 


Molinostrongylus dollfusi Mawson, Tx, Gx, P. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 


(Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Order Undetermined 

?Rhabditida 
Undetermined genus and species, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds 
unpublished data), bacterial feeder (K. Davies pers. comm. 2003). 


Undetermined Family 
Undetermined genus and species, Gp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard 
Creek (Harris 1970). 


Undetermined genus and species, Gp. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 


Phylum Mollusca 


Class Gastropoda 
Order Stylommatophora 
Charopidae 
Elsothera funera Cox, Gx?. NEW SOUTH WALES: Grill Cave (B44), Bungonia (Hamilton-Smith 
unpublished data); VICTORIA: Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades 
of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen 
and Milledge 1990). 


Undetermined Family 
Undetermined genus and species, Tx, Gx?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), 
Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Carn Dum (E15), Mount Etna 
(Hamilton-Smith unpublished data). 


Phylum Annelida 


Class Oligochaeta 
Order Haplotaxida 
Lumbricidae 
Undetermined genus and species, Tp, Gx?. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990). 


Order Undetermined 
Undetermined genus and species, Tp?, Gx?. QUEENSLAND: Four Mile Cave (C14), Camooweal 
(Hamilton-Smith unpublished data). 


Phylum Arthropoda 


Class Arachnida 
Order Scorpionida 
Undetermined Family 
Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990). 


Order Araneae 

Agelenidae 
Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990). 


Amaurobiidae 
Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (B1), Buchan (Yen and 


8 Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), 
East Buchan (Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). 


Ctenizidae 
Misgolas sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). 


Cyatholipidae 
Undetermined genus and species, Gx?. VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and 
Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), 
Murrindal (Yen and Milledge 1990). 


Cycloctenidae 
Cyclotenus abyssinus Urquhart, Tp. VICTORIA: Shades of Death Cave (M3), Murrindal (Hamilton- 
Smith unpublished data). 


Toxopsioides sp., Tp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Gray 
1973b); Yessabah Bat Cave (YE1), Yessabah (Gray 1973b). 


Undetermined genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 
1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave (M3), 
Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); 
Dickson Cave (M30), Murrindal (Yen and Milledge 1990). 


Desidae 
Badumna socialis Rainbow, Tp, Gx?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia 
(Hamilton-Smith unpublished data). 


Colcarteria carrai Gray, Tp?. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1992). 


Colcarteria yessabah Gray, Tp. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1992). 


Dictynidae 
Undescribed genus and species, Gx?. VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 
1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). 


Filistatidae 
Undescribed genus and species, Tp2. WESTERN AUSTRALIA: Cape Range peninsula (Gray 
1994). 


Gradungulidae 
Progradungula carraiensis Forster and Gray, Tp1, Gp . NEW SOUTH WALES: Carrai Bat Cave 
(SC5), Stockyard Creek (Forster et al. 1987). 


Linyphiidae 
Laetesia weburdi Urquhart, Gx?. NEW SOUTH WALES: Jenolan Caves (Hamilton-Smith 
unpublished data). 


Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990). 


Lycosidae 


Lycosa speciosa Koch, Tp1. NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek 
(Gray 1973b). 


Proc. Linn. Soc. N.S.W., 125, 2004 9 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Mimetidae 
Australomimetus maculosus Rainbow, Tp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), 
Yessabah (Gray 1973b); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); 
Jenolan Caves (Hamilton-Smith unpublished data). 


Undetermined genus and species, Gx?. VICTORIA: Spring Creek Cave (B1), Buchan (Yen and 
Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990). 


Pholcidae 
Physocyclus sp., NEW SOUTH WALES: Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b); 
Colong Main Cave (CG3), Colong (Gray 1973b). 


Psilochorus sp., NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah (Gray 1973b) . 


Pisauridae 
Undetermined genus and species, NEW SOUTH WALES: Comboyne C4 Cave, Comboyne (Gray 
1973b); Carrai Bat Cave (SC5), Stockyard Creek (Gray 1973b). 


Salticidae 
Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990). 


Segestriidae 
Undetermined genus and species, Gx?. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990). 


Stiphidiidae 
Stiphidon sp., Gx?. NEW SOUTH WALES: Colong Cave (CG1), Colong (Hamilton-Smith 
unpublished data). 


Theridiidae 
Theridon sp., Tp, Gp. NEW SOUTH WALES: Colong Cave (CG1), Colong (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Steatoda sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Theridiosomatinae 
Undetermined genus and species, Gp?. NEW SOUTH WALES: Colong Cave (CG1), Colong 
(Hamilton-Smith unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished 
data). 


Uloboridae 
Philoponella patherinus Keyserling, Tp. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data). 


Undetermined Family 
Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith unpublished data); 
Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Cave (CG1), Colong 
(Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton-Smith unpublished 
data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave 
(T1), Tuglow (Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton- 


10 Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Smith unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data); Basin Cave (W4), Wombeyan (Hamilton-Smith unpublished data); Fig Tree 
Cave (W148), Wombeyan (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta 
Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Abrakurrie 
Cave (N3) (Hamilton-Smith unpublished data); Madura Cave (Madura 6 Mile Cave) (N62) 
(Hamilton-Smith unpublished data); QUEENSLAND: Four Mile Cave (C14), Camooweal 
(Hamilton-Smith unpublished data); Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data); Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished 
data); Barker’s Cave (U34), Undara (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), 
Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), 
Mt Etna (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith 
unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet 
Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data); Ilium Cave (E31), Mount 
Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Snowflake Cave (L1), Glenelg River (Hamilton-Smith 
unpublished data); Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan 
(Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Trogdip 
Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), 
Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 1990); 
Anticline Cave (M11), Murrindal (Yen and Milledge 1990); SSS Cave (M44), Murrindal 
(Hamilton-Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith 
unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Mt Widderin Cave 
(H1), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mount Napier (Hamilton- 
Smith unpublished data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
Grassmere Cave (W6), Warrnambool (Hamilton-Smith unpublished data). 


Order Opilionida 

Triaenoncychidae 
Holonuncia cavernicola Forster, Tp2. NEW SOUTH WALES: Basin Cave (W4), Wombeyan 
(Hamilton-Smith 1967); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data). 


Holonuncia seriata Roewer, Tp1, Gx. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 


Undetermined genus and species, Tp, Gp. VICTORIA: Moon Cave (B2), Buchan (Yen and 
Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); Shades of Death Cave 
(M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal (Yen and Milledge 
1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave (M30), Murrindal 
(Yen and Milledge 1990). 


Undetermined Family 
Undetermined genus and species, Tp, Gp? NEW SOUTH WALES: The Drum Cave (B13), 
Bungonia (Hamilton-Smith unpublished data); Chalk Cave (B26), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main 
Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Gable Cave (CL7), Cliefden (Hamilton- 
Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith unpublished data); 
Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith unpublished data); 
Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith unpublished data); Glen 
Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), 
Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 
unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Lynch Cave (N60) (Hamilton-Smith unpublished data); QUEENSLAND: 
Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); 


Proc. Linn. Soc. N.S.W., 125, 2004 11 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Unnamed 
Cave (NG1), New Guinea Ridge (Hamilton-Smith unpublished data). 


Order Pseudoscorpionida 
Atemnidae 


Oratemnus cavernicola Beier, Tp, Gp?. NEW SOUTH WALES: Jump Up Cave, Gray Range (Beier 
1976). 


Cheiridiidae 


Cryptocheiridium australicum Beier, Tp2, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave 
(N47) (Richards 1971). 


Cheliferidae 


Protochelifer naracoortensis Beier, Tp2, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Beier 1968; Bellati et al. 2003). 


Protochelifer cavernarum Beier, Tp2, Gb. NEW SOUTH WALES: Murder Cave (CL2), Cliefden 
(Beier 1967, 1968); Island Cave (CL6), Cliefden (Hamilton-Smith unpublished data); Belfry Cave 
(TR2), Timor (Beier 1967); Ashford Caves, Ashford (Beier 1968); NULLARBOR PLAIN: Warbla 
Cave (N1) (Richards 1971); Weebuddie [Weebubbie, sic] Cave (N2) (Beier 1975); Abrakurrie Cave 
(N3) (Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Mullamullang Cave (N37) 
(Richards 1971); Lynch Cave (N60) (Richards 1971); QUEENSLAND: Taylor Cave (4U4), Undara 
(Howarth 1988); Collins Cave No.1, Undara (Howarth 1988); VICTORIA: Clogg’s Cave (EB2), 
East Buchan (Beier 1968); WESTERN AUSTRALIA: Gooseberry Cave (J1), Jurien Bay (Beier 
1968); Eneabba Caves (E1-3), Eneabba (Lowry 1996); Arramall Cave (E22), Eneabba (Lowry 
1996); River Cave (E23), Eneabba (Lowry 1996); Weelawadji Cave (E24) Eneabba (Lowry 1996); 
Super Cave (SH1), South Hill River (Hamilton-Smith unpublished data). 


Protochelifer sp. Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Bellati et al. 
2003). 


Chernetidae 


Sundochernes guanophilus Beier, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Beier 1967). 


Troglochernes imitans Beier, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elvyn Cave (N47) (Beier 
1975); Cocklebiddy Cave (N48) (Beier 1975); Pannikin Plain Cave (N49) (Beier 1975); Dingo 
Cave (Dingo-Donga) (N160) (Richards 1971). 


Chthoniidae 


12 


Austrochthonius cavicola Beier, Tp, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte 
(Beier 1968). 


Paraliochthonius cavicolus Beier, Tp2, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 


Pseudotyrannochthonius hamiltonsmithi Beier, Tp2, Gp. VICTORIA: Mt Widderin Cave (H1), 
Skipton (Beier 1968). 


Sathrochthonius tuena Chamberlin, Tp2, Gp. NEW SOUTH WALES: Basin Cave (W4), 
Wombeyan (Beier 1967), Deua Cave (DE1), Deua (Eberhard and Spate 1995); Punchbowl Cave 
(WJ8), Wee Jasper (Beier 1968); Imperial Cave (J4), Jenolan (Hamilton-Smith 1967; Gibian et al. 
1988); Southern Limestone, Jenolan (Hamilton-Smith 1967; Beier 1968; Gibian et al. 1988); 
Paradox Cave (J48), Jenolan (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Sathrochthonius webbi Muchmore, Tb, Gp. QUEENSLAND: Holy Jump Lava Cave (BM1), 
Bauer’s Mountain southern Queensland (Muchmore 1982). 


Tyrannochthonius cavicola Beier, Tp2, Gb. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Beier 1967; Harvey 1989). 


Undetermined Family 
Undetermined genus and species, Tp, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Royal Arch Tower, 
Chillagoe (Matts 1987). 


Undetermined genus and species, Tp?, Gx. VICTORIA: Anticline Cave (M11), Murrindal (Yen and 
Milledge 1990). 


Mites 
The mites have been arranged according to the higher classification used by Halliday (1998). Many changes 
to nomenclature have occurred since previous checklists of cavernicolous fauna have been published so the 
family placement of some species has been updated to reflect this. Previous family placements have not been 
recorded but where synonymy has occured the old name (either family or genus) has been included in 
brackets. Previous generic placements have been recorded in brackets with the prefix “=”. 


Order Acariformes 

Suborder Astigmata 

Histiostomatidae 
Histiostoma sp. NULLARBOR PLAIN: Mullamullang Cave (N37) (Hamilton-Smith unpublished 
data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Rosensteiniidae 
Nycteriglyphus (Coproglyphus) dewae Zakhvatkin, Tp2, Gb. NEW SOUTH WALES: Basin Cave 
(W4), Wombeyan (Womersley 1963a; Richards 1967b); Fig Tree Cave (W148), Wombeyan 
(Womersley 1963a; Richards 1967b); Railway tunnel, North Sydney (Womersley 1963a); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Womersley 1963a). 


Nycteriglyphus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); 
Dingo Cave (160) (Richards 1971). 


Glycyphagus sp., Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 1971); 
Dingo Cave (N160) (Richards 1971). 


Suborder Prostigmata 

Labidostomidae 
Undetermined genus and species. NEW SOUTH WALES: Island Cave (CL6), Cliefden (Hamilton- 
Smith unpublished data). 


Neotrombidiidae 
Neotrombidium gracilare Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Womersley 1963a); Murder Cave (CL2), Cliefden (Womersley 1963a); Punchbowl 
Cave (WJ8), Wee Jasper (Womersley 1963a); VICTORIA: O’Rourke’s Cave (B12), Buchan 
(Hamuilton-Smith 1967); Wilson Cave (EB4), East Buchan (Hamilton-Smith 1967). 


Neotrombidium gracilipes Womersley, Tp2, Gb. NEW SOUTH WALES: Fig Tree Cave (W148), 
Wombeyan (Hamilton-Smith 1967). 


Proc. Linn. Soc. N.S.W., 125, 2004 13 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Neotrombidium neptunium Southcott, VICTORIA: Clogg’s Cave (EB2), East Buchan (Hamilton- 
Smith unpublished data). 


Neotrombidium sp., Tp, Gb. NULLARBOR PLAIN: Firestick Cave (N70) (Richards 1971); Dingo 
Cave (N160) (Richards 1971). 


Trombiculidae 


Rudnicula barbarae Domrow (= Trombicula), Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor 
Mine, Pine Creek (Hamilton-Smith unpublished data). 


Trombicula thomsoni Womersley, Tx, Gx, P. NEW SOUTH WALES: Bonalbo Colliery (Hamilton- 
Smith unpublished data); Riverton (Hamilton-Smith unpublished data); NORTHERN 
TERRITORY: Kuhinoor Mine, Pine Creek (Hamilton-Smith unpublished data). 


Trombicula dewae Domrow, Tx, Gx, P. NORTHERN TERRITORY: Kuhinoor Mine, Pine Creek 
(Hamilton-Smith unpublished data). 


Order Parasitiformes 
Suborder Ixodida 
Argasidae 


Ixodidae 


Argas sp., Tx, Gx, P. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data). 


Amblyomma moreliae Koch, Gx, P. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 


Ixodes simplex simplex Neumann, Gx, P. Bat parasite in eastern Australia (Hamilton-Smith 1966b; 
Eberhard 1998); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished 
data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Spring 
Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); Slocombe’s Cave (BA1), The Basin 
(Hamilton-Smith unpublished data); Anticline Cave (M11), Murrindal (Hamilton-Smith 
unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data); Starlight 
Cave (W5), Warrnambool (Hamilton-Smith unpublished data); Grassmere Cave (W6), 
Warrnambool (Hamilton-Smith unpublished data). 


Undetermined genus and species, Gx, P. QUEENSLAND: Clam Cavern (CH26), Walkunder 
Tower, Chillagoe (Matts 1987); Spatial Cavern (CH41), Walkunder Tower, Chillagoe (Matts 1987); 
Royal Arch Cave (CH9), Royal Arch Tower, Chillagoe (Matts 1987); VICTORIA: Nargun’s Cave 
(NN1), Nowa Nowa (Hamilton-Smith unpublished data) 


Suborder Mesostigmata 
Ameroseiidae 


Ameroseius plumosus Oudemans, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et 
al. 2003). 


Laelapidae 


14 


Cosmolaelaps sp., Tp2, Gb. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper (Hamilton- 
Smith 1967); QUEENSLAND: Railway tunnel, Samford (Hamilton-Smith 1967). 


Hypoaspis (Gaeolaelaps) annectans Womersley, Tp, Gp. NEW SOUTH WALES: Carrai Bat Cave 
(SC5), Stockyard Creek (Harris 1971). 


Hypoaspis (Gaeolaelaps) sp.1, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Hypoaspis (Gaeolaelaps) sp.2, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 


Hypoaspis (Gaeolaelaps) sp., Tp2, Gb. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith 1967). 


Ichoronyssus (Pleisiolaelaps) miniopteri (Zumpt and Patterson 1952) (= Neospinolaelaps, 
Spinolaelaps), Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 1998); 
Bonalbo Colliery (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), 
Naracoorte (Hamilton-Smith unpublished data). 


Ichoronyssus (Pleisiolaelaps) aristippe Domrow, NEW SOUTH WALES: Cheitmore Cave, 
Cheitmore (Hamilton-Smith unpublished data); Wombeyan Caves (Hamilton-Smith unpublished 
data); Bonalbo Colliery (Hamilton-Smith unpublished data). 


Macrochelidae 
Macrocheles spatei Halliday, Tp1, Gp. NEW SOUTH WALES: Deua Cave (DE1), Deua National 
Park (Halliday 2000). 


Macrocheles tenuirostris Krantz and Filipponi, Tp1, Gp. NEW SOUTH WALES: Paradox Cave 
(J48), Jenolan Caves (Halliday 2000); Cleatmore Cave, Deua National Park (Halliday 2000); 
Colong Cave, Woof’s Cavern (CG1), Colong (Halliday 2000); Church Cave (WJ31), Wee Jasper 
(Halliday 2000); TASMANIA: Fisher Island, in nests and burrows of muttonbird (Krantz and 
Fillipponi 1964); VICTORIA: Panmure Cave (H5), Warrnambool (Hamilton-Smith 1967). 


Macronyssidae 
Macronyssus aristippe Domrow (= Ichoronyssus), Tp, Gx, P. NEW SOUTH WALES: Bungonia 
various caves (Eberhard 1998). 


Trichonyssus australicus Womersley, Tx, Gx, P. NULLARBOR PLAIN: Warbla Cave (N1) 
(Hamilton-Smith unpublished data). 


Parantennulidae 
Micromegistus gourlayi Womersley. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 


Parasitidae 
?Eugamasus sp., Tp, Gp. NULLARBOR PLAIN: Dingo Cave (N160) (Richards 1971). 


Sejidae (Ichthyostomatogastridae) 
Asternolaelaps australis Womersley and Domrow, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2) 
Naracoorte (Womersley and Domrow 1959; Hamilton-Smith 1967); VICTORIA: O’Rourkes Cave 
(B12), Buchan (Hamilton-Smith 1967). 


Spinturnicidae 
Spinturnix psi Kolenati, Tp, Gx, P. NEW SOUTH WALES: Bungonia various caves (Eberhard 
1998). 


Undetermined genus and species, Tp, Gx, P. NEW SOUTH WALES: Colong Main Cave (CG1), 
Colong (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) 
(Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished 
data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); 
Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton (Hamilton-Smith 


Proc. Linn. Soc. N.S.W., 125, 2004 15 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished 
data); WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished 
data). 


Urodinychidae 
Uroobovella (Austruropoda) coprophila Womersley (= Cilliba), Tp2, Gp. NEW SOUTH WALES: 
Cave C4, Comboyne (Smith 1982b); Carrai Bat Cave (SC5), Stockyard Creek (Harris 1973); 
Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith unpublished data); Church Cave (WJ31), 
Wee Jasper (Hamilton-Smith 1966b, 1967); Fig Tree Cave (W148), Wombeyan (Hamilton-Smith 
1966b, 1967); Cheitmore Cave, Cheitmore (Hamilton-Smith unpublished data); QUEENSLAND: 
Arch Cave (U22), Undara (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), 
Riverton (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Bellati et al. 2003); VICTORIA: Anticline Cave (M11), Murrindal (Hamilton-Smith 1967). 


Genus and species undetermined, NEW SOUTH WALES: Deua Cave (DE1), Deua (Eberhard and 
Spate 1995). 


Undetermined Family 
Undetermined sp. 1, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 
1971). 


Undetermined sp. 2, Tp, Gp. NULLARBOR PLAIN: Murra-El-Elevyn Cave (N47) (Richards 
1971). 


Undetermined Acarina 

Undetermined Family 
Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Grimes Cave (CI53) 
(Humphreys and Eberhard 2001). 


Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Gable Cave (CL7), Cliefden 
(Hamilton-Smith unpublished data); NORTHERN TERRITORY: Kintore Cave (K2), Katherine 
(Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- 
Smith unpublished data); Murra-El-Elevyn Cave (N47) (Hamilton-Smith unpublished data); 
QUEENSLAND: Flogged Horse Cave (Cammoo Cave) (J83), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Asbestos mine near Arkaba, Flinders 
Ranges (Hamilton-Smith unpublished data); Drop Drop Cave (L29), Lower south east (Hamilton- 
Smith unpublished data); Joanna Bat Cave (U38), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); O’Rourkes Cave (B12), 
Buchan (Hamilton-Smith unpublished data); Mabel Cave (EB1), East Buchan (Yen and Milledge 
1990); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave 
(EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal (Yen 
and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson Cave 
(M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith 
unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); Grassmere Cave 
(W6), Warrnambool (Hamilton-Smith unpublished data). 


Class Crustacea 
Order Isopoda 
Armadillidae 
Merulana sp. nov., Tp. NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan (Dennis 1986). 


Oniscidae 


Plymophiloscia sp. Vandel, Tp, Gp. NULLARBOR PLAIN: Pannikin Plain Cave (N49) (Richards — 
1971; Gray 1973a). 


16 Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Undetermined genus and species, Tp, Gp. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone 
Ridge, Rockhampton (Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna 
(Hamilton-Smith unpublished data); Carn Dum (E15), Mount Etna (Hamilton-Smith unpublished 
data); VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data). 


Philosciidae 
Abebaioscia troglodytes Vandel, Tb, Gp?. NULLARBOR PLAIN: Pannikin Plain Cave (N49) 
(Vandel 1973). 


Eurygastor montanus troglophilus Vandel, Tp?. VICTORIA: Anticline Cave (M11), Murrindal 
(Vandel 1973). 


Laevophiloscia dongarrensis Wahrberg, Tp, Gx?. WESTERN AUSTRALIA: Yanchep Cave 
(YN16), Yanchep (Vandel 1973); Minnie’s Grotto (YN28), Yanchep (Vandel 1973); Gooseberry 
Cave (J1), Jurien Bay (Vandel 1973). 


Laevophiloscia hamiltoni Vandel, Tp, Gx. WESTERN AUSTRALIA: Weelawadji Cave (E24), 
Eneabba (Vandel 1973); Labyrinth Cave (AU16), Augusta (Vandel 1973) 


Laevophiloscia michaelseni Vandel, Tp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Vandel 
1973). 


Porcellionidae 
Porcellio scaber Latreille, Tp1. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 


Undetermined Family 
Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Cliefden Main 
Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Cave C4, Comboyne (Hamilton-Smith 
unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith 
unpublished data); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Hamilton-Smith 
unpublished data); Main Cave (Ballroom Cave) (TR1), Timor (Hamilton-Smith unpublished data); 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave 
(T1), Tuglow (Hamilton-Smith unpublished data); Piano Cave (Long Cave) (WA12), Walli 
(Hamilton-Smith unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith 
unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data); Yessabah Bat Cave (YE1), Yessabah (Hamilton-Smith unpublished data); 
NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith unpublished data); Cocklebiddy 
Cave (N48) (Hamilton-Smith unpublished data); QUEENSLAND: Barker’s Cave (U34), Undara 
(Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished 
data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith unpublished data); Cathedral Cave 
(U12), Naracoorte (Hamilton-Smith unpublished data); Fox Cave (U22), Naracoorte (Hamilton- 
Smith unpublished data); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East 
Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and Milledge 1990); 
Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Anticline Cave (M11), Murrindal 
(Yen and Milledge 1990); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith unpublished data); 
Stockyard Cave (E3), Eneabba (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 U7 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Order Amphipoda 
Undetermined Family 


Undetermined genus and species, Tp, Gx. VICTORIA: Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990). 


Class Myriapoda 


Order Diplopoda 
Undetermined Family 


Undetermined genus and species, Tp2, Gx. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 


Undetermined genus and species, Tp?, Gx. NEW SOUTH WALES: Island Cave (CL6), Cliefden 
(Hamilton-Smith unpublished data); The Drum Cave (B13), Bungonia (Hamilton-Smith 
unpublished data); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Paradox Cave 
(J48), Jenolan (Hamilton-Smith unpublished data); Moparabah Cave (Temagog Cave) (MP1), 
Moparabah (Hamilton-Smith unpublished data); Carrai Bat Cave (SC5), Stockyard Creek 
(Hamilton-Smith unpublished data); Belfry Cave (TR2), Timor (Hamilton-Smith unpublished data); 
Tuglow Cave (T1), Tuglow (Hamilton-Smith unpublished data); Fig Tree Cave (W148), Wombeyan 
(Hamilton-Smith unpublished data); Punchbowl Cave (WJ8), Wee Jasper (Hamilton-Smith 
unpublished data); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Willi 
Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); Yessabah Bat 
Cave (YE1), Yessabah (Hamilton-Smith unpublished data); QUEENSLAND: Barker’s Cave (U34), 
Undara (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished 
data); Piglet Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data); Jolly Roger 
Cave (E29), Mountt Etna (Hamilton-Smith unpublished data); Glen Lyon River Cave (GL1), Glen 
Lyon (Hamilton-Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith 
unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and Milledge 1990); Mabel 
Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson Cave (EB4), East Buchan (Yen and 
Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), 
Nowa Nowa (Hamilton-Smith unpublished data). 


Order Chilopoda 
Scolopendromorpha 


Undetermined genus and species. NULLARBOR PLAIN: Mullamullang Cave (N37) (Richards 
1971). 


Undetermined Family 


18 


Undetermined genus and species, Gp?. NEW SOUTH WALES: Cave C4, Comboyne (Hamilton- 
Smith unpublished data); Youndales Cave (Hut Cave) (KB1), Kunderang Brook (Hamilton-Smith 
unpublished data); Carrai Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); 
Moparabah Cave (MP1), Moparabah (Hamilton-Smith unpublished data); Belfry Cave (TR2), 
Timor (Hamilton-Smith unpublished data); NORTHERN TERRITORY: Cutta Cutta Cave (K1), 
Katherine (Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith 
unpublished data); NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Hamilton-Smith unpublished 
data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith unpublished data); 
Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Superclass Hexapoda 
Class Insecta 


Order Collembola 

Armadillidae 
Buddelundia albomarginata Wahrberg, Tp, Gx?. NULLARBOR PLAIN: Murrawyinee [sic] No.1 
Cave (N7) (Vandel 1973); Cocklebiddy Cave (N48) (Vandel 1973); Lynch Cave (N60) (Vandel 
1973); Madura Cave (N62) (Vandel 1973); Old Homestead Cave (N83) (Vandel 1973); Unnamed 
cave (N140) (Vandel 1973). 


Entomobryidae 
Lepidocyrtus sp., Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Lepidosira australica Schott, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 


Undetermined genus and species, Gp? NEW SOUTH WALES: Belfry Cave (TR2), Timor (James 
et al. 1976); Chalk Cave (B26), Bungonia (Hamilton-Smith unpublished data). 


Hypogastruridae 
Hypogastrura sp., NEW SOUTH WALES: Grill Cave (B44), Bungonia (Wellings 1977); SOUTH 
AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Isotomidae 
Folsomia candida Willem, Tp. NEW SOUTH WALES: Paradox Cave (J48), Jenolan (Eberhard 
1993); Imperial Cave (J4), Jenolan (Eberhard and Spate 1995); Tuglow Main Cave (T1), Tuglow 
(Eberhard 1993); Jillebean Cave (Y22), Yarrangobilly (Eberhard 1993). 


Paronellidae 
Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Eberhard and Spate 1995). 


Undetermined Family 
Undetermined genus and species, Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) 
(Richards 1971); Lynch Cave (N60) (Richards 1971); Dingo Cave (N160) (Richards 1971); 
VICTORIA: SSS Cave (M44), Murrindal (Hamilton-Smith unpublished data); Mt Widderin Cave 
(H1), Skipton (Hamilton-Smith unpublished data). 


Undetermined genus and species, Tp?, Gp?. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith 
unpublished data); Glen Dhu Cave (Allston Cave) (TR15), Timor (Hamilton-Smith unpublished 
data); NORTHERN TERRITORY: 16 Mile Cave, Katherine (Hamilton-Smith unpublished data); 
QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton-Smith unpublished data); SOUTH 
AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); VICTORIA: 
Moon Cave (B2), Buchan (Yen and Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and 
Milledge 1990); Wilson’s Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip 
Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Lilly Pilly Cave (M8), Murrindal 
(Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Panmure 
Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 


Order Diplura 

Undetermined family 
Undetermined genus and species, SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 
2003). 


Proc. Linn. Soc. N.S.W., 125, 2004 19 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Order Blattodea 
Blattellidae 


Blattidae 


20 


Neotemnopteryx australis Saussure, Tp, Gp. NEW SOUTH WALES: Moparabah Cave (Temagog 
Cave) (MP1), Moparabah (Hamilton-Smith 1967); Cave C4, Comboyne (Hamilton-Smith 1967); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Neotemnopteryx fulva Saussure (= Gislenia australica Brunner), Tp, Gb. NEW SOUTH WALES: 
Glen Dhu Cave (Allston Cave) (TR15), Timor (Richards 1967a); Murder Cave (CL2), Cliefden 
(Richards 1967a); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Richards 1967a); Haystall 
Cave (U23), Naracoorte (Richards 1967a); VICTORIA: Mabel Cave (EB1), East Buchan (Richards 
1967a). 


Neotemnopteryx sp., Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith 
unpublished data); Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith unpublished data); 
Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Viator Main Cave (VR1), 
Viator Hill (Hamilton-Smith unpublished data). 


?Neotemnopteryx (?Gislenia sp.), Tp, Gp?. NEW SOUTH WALES: Ashford Main Cave (AS1), 
Ashford (Richards 1967a); Cave 4, Comboyne (Richards 1967a); Hill Cave (TR7), Timor (Richards 
1967a); Moparabah Cave (Temagog Cave) (MP1), Moparabah (Richards 1967a); Swallow Cave 
(CU1), Cudgegong (Richards 1967a); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Richards 1967a); Riverton Main Cave (RN1), Riverton, southern Queensland (Richards 1967a); 
Viator Cave (VR4), Viator Hill, southern Queensland (Richards 1967a); Johannsen’s Cave (J1), 
Limestone Ridge, Rockhampton (Hamilton-Smith 1967); Winding Stairway Cave (4E2), Mt Etna 
(Hamilton-Smith 1967); SOUTH AUSTRALIA: Alexandra Cave (5U3), Naracoorte (Richards 
1967a); Bat Cave (U2), Naracoorte (Richards 1967a). 


Paratemnopteryx atra Princis, Tb, Gp. WESTERN AUSTRALIA: Mines near Marble Bar (Princis 
1963; Richards 1967a; Moore et al. 2001). 


Paratemnopteryx rufa Tepper, Gb?. NULLARBOR PLAIN: Murrawijinie No.3 Cave (N9) 
(Richards 1971); Abrakurrie Cave (N3) (Richards 1971). 


Paratemnopteryx sp., Tp, Gb?. QUEENSLAND: Pinwill Cave (4U17), Undara (Howarth 1988). 


Shawella douglasi Princis, Tp, Gb?. NEW SOUTH WALES: River Cave (SC1), Stockyard Creek 
(Hamilton-Smith 1967); WESTERN AUSTRALIA: Drovers Cave (J2), Jurien Bay (Hamilton-Smith 
unpublished data); Jurien Bay caves (Princis 1963; Richards 1967a); Eneabba Caves (E1-3), 
Eneabba (Lowry 1996); Weelawadji Cave (E24), Eneabba (Lowry 1996). 


Trogloblattella nullarborensis Mackerras, Tb, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) 
(Mackerras 1967; Richards 1971); Koonalda Cave (N4) (Mackerras 1967); Mullamullang Cave 
(N37) (Mackerras 1967); Roaches Rest Cave (N58) (Mackerras 1967); Arubiddy Cave (N81) 
(Mackerras 1967). 


Polyzosteria mitchelli Angas, Tp. NULLARBOR PLAIN: Warbla Cave (N1), (Mackerras 1965); 
Kestrel Cavern (N40) (Mackerras 1965; Richards 1967a). 


Polyzosteria pubescens Tepper, Tp. NULLARBOR PLAIN: Weebubbie Cave (N2) (Hamilton- 
Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Zonioploca medilinea Tepper, Tp?. NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1967a). 


Order Orthoptera 

Rhaphidophoridae 
Australotettix carraiensis Richards, Tp, Gx. NEW SOUTH WALES: Barnett’s Cave (SC6), 
Stockyard Creek (Richards 1964); Carrai Bat Cave (SC5), Stockyard Creek (Richards 1964); Col’s 
Cave, Stockyard Creek (Richards 1964); Lot’s Mansion, Stockyard Creek (Richards 1964); River 
Cave (SC1) Stockyard Creek (Richards 1964). 


Cavernotettix buchanensis Richards, Tx, Gx. VICTORIA: Wilson Cave (EB4), East Buchan 
(Richards 1966; Yen and Milledge 1990); Trogdip Cave (EB10), East Buchan (Hamilton-Smith 
unpublished data); Spring Creek Cave (B1), Buchan (Richards 1966; Yen and Milledge 1990); 
Shades of Death Cave (M3), Murrindal (Yen and Milledge 1990); Lilly Pilly Cave (M8), Murrindal 
(Yen and Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990); Dickson 
Cave (M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave (NN1), Nowa Nowa Caves 
(Richards 1966; Yen and Milledge 1990); Weta Cave (NN2), Nowa Nowa Caves (Richards 1966; 
Yen and Milledge 1990). 


Cavernotettix montanus Richards, Tx, Gx. NEW SOUTH WALES: small cave nr Glory Cave, 
Yarrangobilly (Richards 1966); Jersey Cave (Y23), Yarrangobilly (Richards 1966); Restoration 
Cave (Y50), Yarrangobilly (Richards 1966); Unnamed cave, Yarrangobilly (Richards 1966); 
Cooleman Cave (CP1), Cooleman Plains (Richards 1966); Unnamed cave opp. Blue Waterhole, 
Cooleman Plains (Richards 1966); Unnamed cave nr Murray Cave, Cooleman Plains (Richards 
1966). 


Cavernotettix wyanbenensis Richards, Tx, Gx. NEW SOUTH WALES: Wyanbene Cave (WY1), 
Wyanbene (Richards 1966); Bat Cave, Cheitmore (Richards 1966). 


Pallidotettix nullarborensis Richards, Tx, Gx. NULLARBOR PLAIN: Warbla Cave (N1) (Richards 
1971); Weebubbie Cave (N2) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); 
Cocklebiddy Cave (N48) (Richards 1971); Pannikin Plain Cave (N49) (Richards 1971); Tommy 
Grahams Cave (N56) (Richards 1971). 


Undetermined genus and species, Tx, Gx. NEW SOUTH WALES: Grill Cave (B44), Bungonia 
(Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong (Hamilton-Smith 
unpublished data); QUEENSLAND: Danes Four Cave (C4), Camooweal (Hamilton-Smith 
unpublished data); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), Camooweal 
(Hamilton-Smith unpublished data); Haunted Cave (CH1), Chillagoe (Hamilton-Smith unpublished 
data); VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds unpublished data). 


Order Psocoptera 

Liposcelidae 
Liposcelis corrodens Broadhead, Tp1, Gp. WESTERN AUSTRALIA: Arranmall [sic] Cave (E22), 
Eneabba (Smithers 1975); undetermined caves (Smithers 1975). 


Psyllipsocidae 
?Psyllipsocus ramburi Selys-Longcamp, Tp1, Gp. NEW SOUTH WALES: Murder Cave (CL2), 
Cliefden (Hamilton-Smith 1967); Island Cave (CL6), Cliefden (Smithers 1964); Hill Cave (TR7), 
Timor (James et al. 1976); Basin Cave (W4), Wombeyan (Smithers 1964); Fig Tree Cave (W148), 
Wombeyan (Smithers 1975); Punchbowl Cave (WJ8), Wee Jasper (Smithers 1964); Church Cave 
(WJ31), Wee Jasper (Smithers 1964); NULLARBOR PLAIN: Weebubbie Cave (N2) (Richards 
1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); Koonalda Cave (N4) 
(Richards 1971); Madura Cave (N62), (Richards 1971); QUEENSLAND: Riverton Main Cave 
(RN1), Riverton, southern Queensland (Hamilton-Smith 1967); SOUTH AUSTRALIA: Bat Cave 


Proc. Linn. Soc. N.S.W., 125, 2004 21 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


(U2), Naracoorte (Smithers 1964; Bellati et al. 2003); Blackberry Cave, Naracoorte (Smithers 
1964); VICTORIA: Clogg’s Cave (EB2), East Buchan (Smithers 1964); O’Rourkes Cave (B12), 
Buchan (Smithers 1964); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith 1967). 


Trogiidae 
Lepinotus inquilinus Heyden, Tp1, Gp. WESTERN AUSTRALIA: Arranmall (sic) Cave (E22), 
Eneabba (Smithers 1975). 


?Lepinotus reticulatus Enderlein, Tp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Smithers 
1964; Bellati et al. 2003) 


Undetermined genus and species, NEW SOUTH WALES: Fig Tree Cave (W148), Wombeyan 
(Dennis and Mayhew 1986). 


Undetermined Family 
Undetermined genus and species, Tp, Gx. NEW SOUTH WALES: Gable Cave (CL7), Cliefden 
(Hamilton-Smith unpublished data); QUEENSLAND: Viator Main Cave (VR1), Viator Hill 
(Hamilton-Smith unpublished data); VICTORIA: Lilly Pilly Cave (M8), Murrindal (Yen and 
Milledge 1990). 


Order Hemiptera 

Cixiidae 
Undetermined genus and species, Tp. QUEENSLAND: Mount Etna Main Cave (E1), Mount Etna 
(Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith unpublished data). 


Lygaeoidea 
Undetermined family and genus, Gp?. VICTORIA: Starlight Cave (W5), Warrnambool (T. Moulds 
unpublished data). 


Reduviidae 
Armstrongula sp. Tp, Gp. SOUTH AUSTRALIA: McKinley’s Daughter’s Cave (F175), Flinders 
Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla Gorge, Gammon Ranges (T. 
Moulds unpublished data). 


Centrogonus sp. Tp, Gp. NORTHERN TERRITORY: Kintore Cave (K2), Katherine (Hamilton- 
Smith unpublished data). 


Undetermined Emesinae genus and species, Tp, Gp. QUEENSLAND: Crazy Cracks Cave, Jacks 
Gorge, Broken River (T. Moulds unpublished data); Not Another Frig Tree Crave, Jacks Gorge, 
Broken River (T. Moulds unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), Riverton 
(Hamilton-Smith unpublished data). 


Undetermined genus and species, Tp, Gp. NORTHERN TERRITORY: Cutta Cutta Cave (K1), 
Katherine (Hamilton-Smith unpublished data); QUEENSLAND: Queenslander Cave (CH15), 
Queenslander Tower (CH5246) Chillagoe (T. Moulds unpublished data); Trezkinn Cave (CH14), 
Chillagoe (T. Moulds unpublished data); Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data). 


Undetermined genus and species, Tp, Gp?. QUEENSLAND: Elephant Hole (E8), Mount Etna 
(Hamilton-Smith unpublished data). 


22 Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Undetermined Family 
Undetermined genus and species, QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- 
Smith unpublished data). 


Order Neuroptera 

Myrmeleontidae 
Aeropteryx sp., Tp, Gp. SOUTH AUSTRALIA: McKinley’s Daughter’s Cave (F175), Flinders 
Ranges (T. Moulds unpublished data); Moro Bat Cave (F47), Flinders Ranges (T. Moulds 
unpublished data); Unnamed cave, Brachina Gorge, Flinders Ranges (T. Moulds unpublished data); 
Unnamed bat cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed 
cave, Chambers Gorge, Flinders Ranges (T. Moulds unpublished data); Unnamed mine, Weetootla 
Gorge, Gammon Ranges (T. Moulds unpublished data). 


Myrmeleontinae sp., Tp?. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data). 


Undetermined Family 
Undetermined genus and species, QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s 
Mountain (Hamilton-Smith unpublished data). 


Order Coleoptera 

Anobiidae (Ptinidae) 
Ptinus exulans Erichson, Tp1, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith 1967); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Island 
Cave (CL6), Cliefden (Hamilton-Smith 1967); Jenolan Caves (Hamilton-Smith 1967); Willi Willi 
Bat Cave (WW1), Willi Willi (Hamilton-Smith 1967); Bungonia various caves (Eberhard 1998); 
Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Colong Main Cave (CG1), Colong 
(Hamilton-Smith unpublished data); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); 
Murrawijinie No. 1 Cave (N7) (Richards 1971); Murra-El-Elevyn Cave (N47) (Hamilton-Smith 
1967; Richards 1971); Firestick Cave (N70) (Richards 1971); Dingo Cave (N160) (Richards 1971); 
SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 1967; Bellati et al. 2003); 
Blanche Cave (U4), Naracoorte (Hamilton-Smith 1967); VICTORIA: Starlight Cave (WS), 
Warmambool (Hamilton-Smith 1967); Clogg’s Cave (EB2), East Buchan (Hamilton-Smith 1967); 
WESTERN AUSTRALIA: Goosebury Cave (J1), Jurien Bay (Hamilton-Smith 1967). 


Carabidae 
Anomotarus subterraneus Moore, Tp, Gp. QUEENSLAND: Riverton Main Cave (RN1), Riverton, 
southern Queensland (Moore 1967). 


Cratogaster melus Laporte, Tp?. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data). 


Darodilia sp., Tp?. QUEENSLAND: Winding Stairway Cave (E2), Mt Etna (Hamilton-Smith 
unpublished data). 


Gnathaphanus pulcher Dejean, Tp?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine 
(Hamilton-Smith unpublished data); Kintore Cave (K2), Katherine (Hamilton-Smith unpublished 
data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton (Hamilton-Smith 
unpublished data). 


Lecanomerus sp., Gp?. NEW SOUTH WALES: Youndales Cave (Hut Cave) (KB1), Kunderang 
Brook (Hamilton-Smith unpublished data). 


Mecyclothorax ambiguus Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 23 


24 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Meonis sp., Tp, Gp. QUEENSLAND: Main Mount Etna Cave (E1), Mount Etna (Hamilton-Smith 
unpublished data). 


Mystropomus subcostatus Chaudoir, Tp?. QUEENSLAND: Johannsen’s Cave (J1-2), Limestone 
Ridge, Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); Speaking Tube (E7), Mount Etna (Hamilton-Smith 
unpublished data); Elephant Hole (E8), Mount Etna (Hamilton-Smith unpublished data); Piglet 
Help! Help! Cave (E17), Mount Etna (Hamilton-Smith unpublished data). 


Notonomus angustibasis Sloane, Tp?, Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 


Notospeophonus castaneus castaneus Moore, Tp2. SOUTH AUSTRALIA: Blanche Cave (U4), 
Naracoorte (Hamilton-Smith 1967); Blackberry Cave (U8), Naracoorte (Hamilton-Smith 1967); 
Stick Cave (U11), Naracoorte (Moore 1964); Cathedral Cave (U12), Naracoorte (Moore 1964); Fox 
Cave (U22), Naracoorte (Hamilton-Smith 1967); Haystall Cave (U23), Naracoorte (Hamilton-Smith 
1967); Cave Park Cave (U37), Naracoorte (Hamilton-Smith unpublished data); Tantanoola Caves 
(Hamilton-Smith 1967); VICTORIA: Bat Cave (P6), Portland (Moore 1962); Byaduk Caves, 
Byaduk (Moore 1962); Panmure Cave (H5), Mount Napier (Moore 1964); Mt Widderin Cave (H1), 
Skipton (Hamilton-Smith 1967); Snowflake Cave (L1), Glenelg River (Hamilton-Smith 1967); 
Curran’s Creek Cave (G4), Glenelg River (Hamilton-Smith 1967). 


Notospeophonus castaneus consobrinus Moore, Tp, Gp. VICTORIA: Spring Creek Cave (B1), 
Buchan (Hamilton-Smith unpublished data); Moon Cave (B2), Buchan (Hamilton-Smith 
unpublished data); Mabel Cave (EB1), East Buchan (Hamilton-Smith unpublished data); Wilson’s 
Cave (EB4), East Buchan (Hamilton-Smith unpublished data); Trogdip Cave (EB10), East Buchan 
(Hamilton-Smith unpublished data); Slocombe’s Cave (BA1), The Basin (Hamilton-Smith 
unpublished data); Shades of Death Cave (M3), Murrindal (Hamilton-Smith unpublished data); 
Anticline Cave (M11), Murrindal (Hamilton-Smith unpublished data); SSS Cave (M44), Murrindal 
(Hamilton-Smith unpublished data). 


Notospeophonus jasperensis jasperensis Moore, Tp2, Gp. NEW SOUTH WALES: Punchbowl Cave 
(WJ8), Wee Jasper (Moore 1964); Pylon 58 Cave (WJ99), Wee Jasper (Moore 1964); Basin Cave 
(W4), Wombeyan (Hamilton-Smith unpublished data). 


Notospeophonus jasperensis vicinus Moore, Tp2, Gp. NEW SOUTH WALES: Bungonia various 
caves (Eberhard 1998). 


Notospeophonus pallidus Moore, Tp2, Gp?. NEW SOUTH WALES: Childrens Cave (CL12), 
Cliefden (Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Myponga (Moore 1964); 
NULLARBOR PLAIN: Warbla Cave (N1) (Hamilton-Smith 1967; Richards 1971); Weebubbie 
Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Hamilton-Smith 1967; Richards 1971); 
Koonalda Cave (N4) (Hamilton-Smith 1967; Richards 1971); Koomooloobooka Cave (N6) 
(Richards 1971); Murrawijinie No.3 Cave (N9) (Richards 1971); Knowles Cave (N22) (Hamilton- 
Smith 1967; Richards 1971); Mullamullang Cave (N37) (Richards 1971); Joe’s Cave (N39) 
(Hamilton-Smith 1967; Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Madura Cave 
(Madura 6 Mile South Cave) (N62) (Richards 1971); Lynch Cave (N60) (Richards 1971). 


Notospeophonus sp., Tp, Gp?. QUEENSLAND: Viator Main Cave (VR1), Viator Hill (Hamilton- 
Smith unpublished data). 


Phloeocarabus sp. Tp?, Gp?. QUEENSLAND: Haunted Cave (CH1), Chillagoe (Hamilton-Smith 
unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Pogonoglossus sp., Tp, Gp?. NORTHERN TERRITORY: Cutta Cutta Cave (K1), Katherine 
(Hamilton-Smith unpublished data). 


Pseudoceneus sp. Tp, Gp?. WESTERN AUSTRALIA: Stockyard Cave (E3), Eneabba (Hamilton- 
Smith unpublished data). 


Speotarus lucifugus Moore, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Moore 
1964; Bellati et al. 2003); NULLARBOR PLAIN: Warbla Cave (N1) (Richards 1971); Weebubbie 
Cave (N2) (Richards 1971); Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 
1971); Winbirra Cave (N45) (Richards 1971); Murra-El-Elevyn Cave (N47) (Richards 1971); 
Cocklebiddy Cave (N48) (Richards 1971); Moonera Tank Cave (N53) (Richards 1971); Lynch 
Cave (N60) (Richards 1971); Unnamed cave (N139) (Richards 1971). 


Speotarus princeps Moore, Tp2, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Moore 1964); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished 
data). 


Speotarus sp., Tp, Gp. NULLARBOR PLAIN: Warbla Cave (N1) (Hamilton-Smith unpublished 
data); Weebubbie Cave (N2) (Hamilton-Smith unpublished data); Murra-El-Elevyn Cave (N47) 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Mount Sims Cave (F7), Walpunda 
Creek, Flinders Ranges (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry 
Cave (J1), Jurien Bay (Hamilton-Smith unpublished data). 


Thenarotes speluncarius Moore, Tp, Gp. NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 
1971); Koonalda Cave (N4) (Richards 1971); New Cave (N11) (Richards 1971); Lynch Cave (N60) 
(Richards 1971); Decoration Cave (N84) (Richards 1971); SOUTH AUSTRALIA: Cave No. 1, 
Buckalowie, Flinders Ranges (Hamilton-Smith unpublished data). 


Trechimorphus diemenensis Bates, Tp1, Gx. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); Grill Cave (B44), Bungonia (Hamilton-Smith unpublished data); Jenolan Caves 
(Moore 1964); VICTORIA: Dalley’s Sinkhole (M35), Murrindal (Hamilton-Smith 1967). 


Trichosternus vigorsi Gory, Tp? Gx. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 


Undetermined genus and species, NEW SOUTH WALES: Grill Cave (B44), Bungonia (Eberhard 
and Spate 1995); Belfry Cave (TR2), Timor (James et al. 1976); Glen Dhu Cave (Allston Cave) 
(TR15), Timor (Hamilton-Smith unpublished data); Tuglow Cave (T1), Tuglow (Hamilton-Smith 
unpublished data); QUEENSLAND: Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum Cave), 
Camooweal (Hamilton-Smith unpublished data); Mount Etna Main Cave (E1), Mount Etna 
(Hamilton-Smith unpublished data); Cave with the thing that went thump! (E5), Mount Etna 
(Hamilton-Smith unpublished data). 


Undetermined genus and species, Tp, Gp. QUEENSLAND: Barker’s Cave (U34), Undara 
(Hamilton-Smith unpublished data); VICTORIA: Spring Creek Cave (B1), Buchan (Yen and 
Milledge 1990); Mabel Cave (EB1), East Buchan (Yen and Milledge 1990); Wilson’s Cave (EB4), 
East Buchan (Hamilton-Smith unpublished data); Shades of Death Cave (M3), Murrindal (Yen and 
Milledge 1990); Anticline Cave (M11), Murrindal (Yen and Milledge 1990). 


Cryptophagidae 
Anchicera sp., Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Atomaria sp., Gp. Southern Australia (Hamilton-Smith 1968). 


Proc. Linn. Soc. N.S.W., 125, 2004 25 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Fox Cave (U22), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Wilson’s Cave (EB4), East Buchan (Hamilton- 
Smith unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data). 


Curculionidae 


Mandalotus sp. Gp?. NEW SOUTH WALES: Chalk Cave (B26), Bungonia (Hamilton-Smith 
unpublished data). 


Talaurinus sp. Gp?. QUEENSLAND: Johannsen’s Cave (J1-2), Mount Etna (Hamilton-Smith 
unpublished data). 


Dermestidae 


Dermestes ater DeGeer, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton- 
Smith unpublished data). 


Undetermined genus and species, Tp, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Bellati et al. 2003); QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s Mountain (Hamilton- 
Smith unpublished data); Unidentified cave in southern Queensland (Hamilton-Smith 1967); 
VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 


Endomychidae 


Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith unpublished data). 


Histeridae 


26 


Carcinops sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) 
(Humphreys and Eberhard 2001). 


Saprinus sp., Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford (Hamilton-Smith 
unpublished data); QUEENSLAND: Riverton Main Cave (RN1), Riverton (Hamilton-Smith 
unpublished data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Hamilton-Smith 
unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished 
data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Clogg’s Cave 
(EB2), East Buchan (Hamilton-Smith unpublished data). 


Tomogenius ?ripicola Marseul, Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati 
et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data); 
NULLARBOR PLAIN: Lynch Cave (N60) (Richards 1971); Thylacine Hole (N63) (Richards 
1971); Dingo Cave (N160) (Richards 1971). 


Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998); Ashford Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); Carrai 
Bat Cave (SC5), Stockyard Creek (Hamilton-Smith unpublished data); Willi Willi Bat Cave (Main 
Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); QUEENSLAND: Holy Jump Lava 
Cave (BM1), Bauer’s Mountain (Hamilton-Smith unpublished data); Riverton Main Cave (RN1), 
Riverton (Hamilton-Smith unpublished data); Johannsen’s Cave (J1-2), Limestone Ridge, 
Rockhampton (Hamilton-Smith unpublished data); Winding Stairway Cave (E2), Mt Etna 
(Hamilton-Smith unpublished data); SOUTH AUSTRALIA: Sand Cave (Joanna) (U16), Naracoorte 
(Hamilton-Smith unpublished data); VICTORIA: Chimney Cave (BR1), Bat Ridges, Portland 
(Hamilton-Smith unpublished data); Clogg’s Cave (EB2), East Buchan (Hamilton-Smith 
unpublished data); Nargun’s Cave (NN1), Nowa Nowa (Hamilton-Smith unpublished data); Bat 
Cave (P6), Portland (Hamilton-Smith unpublished data); WESTERN AUSTRALIA: Gooseberry 
Cave (J1), Jurien Bay (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Jacobsoniidae 
Derolathrus sp., Tp, Gb. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); 
Various caves in southern Australia (Hamilton-Smith 1967). 


Undetermined genus and species, Tp, Gb. VICTORIA: Bat Cave (P6), Portland (Hamilton-Smith 
unpublished data); Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished data). 


Lathridiidae 
Corticaria sp., Gp. Southern Australia (Hamilton-Smith 1968); NEW SOUTH WALES: Ashford 
Main Cave (AS1), Ashford (Hamilton-Smith unpublished data); NULLARBOR PLAIN: Weebubbie 
Cave (N2) (Hamilton-Smith unpublished data); Abrakurrie Cave (N3) (Hamilton-Smith unpublished 
data): VICTORIA: Skipton Cave (Mount Widderin Cave) (H1), Mount Napier (Hamilton-Smith 
unpublished data). 


Leiodidae 
Choleva australis, Tp, Gp. QUEENSLAND: Royal Arch Cave (CH9), Chillagoe (Hamilton-Smith 
unpublished data). 


Choleva sp., Tp, Gp. NULLARBOR PLAIN: Cocklebiddy Cave (N48) (Richards 1971); Lynch 
Cave (N60) (Richards 1971). 


Nargomorphus minusculus Blackburn, Tp1, Gp. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte 
(Hamilton-Smith 1967; Bellati et al. 2003); VICTORIA: Anticline Cave (M11), Murrindal 
(Hamilton-Smith 1967). 


Pseudonemadus adelaidae Blackburn, Tp, Gp. NEW SOUTH WALES: Glen Dhu Cave (Allston 
Cave) (TR15), Timor (Hamilton-Smith unpublished data); QUEENSLAND: Riverton Main Cave 
(RN1), Riverton (Hamilton-Smith unpublished data). 


Pseudonemadus australis Erichson, Gp. VICTORIA: Chimney Cave (BR1), Bat Ridge, Portland 
(Hamilton-Smith unpublished data); Bat Cave (P6), Portland (Hamilton-Smith unpublished data); 
Panmure Cave (H5), Mt Napier (Hamilton-Smith unpublished data). 


Pseudonemadus integer Portevin, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data); QUEENSLAND: Speaking Tube (E7), Mount Etna (Hamilton- 
Smith unpublished data); Viator Main Cave (VR1), Viator Hill (Hamilton-Smith unpublished data); 
SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte (Hamilton-Smith unpublished data); 
VICTORIA: Trogdip Cave (EB10), East Buchan (Hamilton-Smith unpublished data); Mt Widderin 
Cave (H1), Skipton (Hamilton-Smith unpublished data); Panmure Cave (H5), Mt Napier (Hamilton- 
Smith unpublished data). 


Pseudonemadus sp., Gp. Southern Australia (Hamilton-Smith 1968). 


?Leiodidae 
Undetermined genus and species, NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Smith 
1982a). 


Melyridae 
Heteromastix sp. Tx?, Gx?. NEW SOUTH WALES: Colong Main Cave (CG1), Colong (Hamilton- 
Smith unpublished data). 


Merophysiidae 


Undetermined genus and species, Gp. NEW SOUTH WALES: Ashford Main Cave (AS1), Ashford 
(Hamilton-Smith 1967). 


Proc. Linn. Soc. N.S.W., 125, 2004 Da 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Pselaphidae 


Ptilidae 


Rybaxis? sp., Tp, Gp. NEW SOUTH WALES: Basin Cave (W4), Wombeyan (Hamilton-Smith 
1966a); Bungonia various caves (Eberhard 1998). 


Tyromorphus speciosus King, Tp1. NEW SOUTH WALES: Unidentified cave, Southern 
Limestone, Jenolan (Hamilton-Smith 1966a); Paradox Cave (J48), Jenolan (Hamilton-Smith 
unpublished data); QUEENSLAND: Johannsen’s Cave (J1-2), Limestone Ridge, Rockhampton 
(Hamilton-Smith 1966a); VICTORIA: Anticline Cave (M11), Murrindal (Hamilton-Smith 1966a). 


Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave, Mingella (Weinstein 
and Slaney 1995). 


Undetermined genus and species, Tp, Gp. VICTORIA: Wilson’s Cave (EB4), East Buchan 
(Hamilton-Smith unpublished data). 


Achosia lanigera Deane, Tp?, Gp. VICTORIA: Wilsons Cave (EB4), East Buchan (Hamilton-Smith 
unpublished data). 


Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Comboyne C4 Cave, Comboyne 
(Hamilton-Smith unpublished data). 


Rhizophagidae 


Undetermined genus and species, Gp. QUEENSLAND: Rope Ladder Cave (FR2), Mingella, 
Fanning River (Weinstein and Slaney 1995). 


Scarabaeidae 


Aulacopris maximus Matthews, Tp1, Gb. NEW SOUTH WALES: Yessabah Bat Cave (YE1), 
Yessabah (Waite 1898); Unknown cave in Coorabakh National Park (formerly part Lansdowne 
State Forest), Taree (Williams 2003). 


Aulacopris reichei White, Tp1, Gp. NEW SOUTH WALES: Yessabah Bat Cave (YE1), Yessabah 
(Lea 1923); Unknown cave, Mosman (Fricke 1964). 


Amphistomus accidatus Matthews, Tx, Gp. QUEENSLAND: Elephant Hole (E8), Mount Etna 
(Hamilton-Smith unpublished data). 


Saprosites mendax Blackburn, Gp. SOUTH AUSTRALIA: Cathedral Cave (U12), Naracoorte 
(Hamilton-Smith unpublished data). 


Undetermined genus and species, Gp. NEW SOUTH WALES: Willi Willi Bat Cave (Main Cave) 
(WW1), Willi Willi (Hamilton-Smith unpublished data). 


Silphidae 


Ptomaphila lachrymosa Schreibers, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 


Staphylinidae 


28 


Myotyphlus jansoni Matthews, Tp1, Gp. NEW SOUTH WALES: Unidentified cave, Southern 
Limestone, Jenolan (Hamilton-Smith and Adams 1966); Paradox Cave (J48), Jenolan (Hamilton- 
Smith unpublished data); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith and 
Adams 1966); Bat Cave (P6), Portland (Hamilton-Smith 1967). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


Tineidae 
Lindera tessellatella Blanchard, Gb?. NEW SOUTH WALES: Humicrib Cave (WJ34), Wee Jasper 
(Eberhard and Spate 1995). 


Monopis crocicapitella Clemens, Tp, Gb. NEW SOUTH WALES: Drum Cave (B13), Bungonia 
(Eberhard 1998); Grill Cave (B44), Bungonia (Eberhard 1998); SOUTH AUSTRALIA: Bat Cave 
(U2), Naracoorte (Bellati et al. 2003); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton- 
Smith unpublished data). 


Monopis sp., Gb. NEW SOUTH WALES: Gable Cave (CL7), Cliefden (Eberhard and Spate 1995); 
Colong Main Cave (CG3), Colong (Eberhard and Spate 1995); Jenolan undetermined cave (Gibian 
et al. 1988); Basin Cave (W4), Wombeyan (Smith 1982b); Undetermined caves, Wombeyan (Dew 
1963); Signature Cave (WJ7), Wee Jasper (Hamilton-Smith unpublished data); Punchbowl Cave 
(WJ8), Wee Jasper (Hamilton-Smith unpublished data); Dogleg Cave (WJ10), Wee Jasper 
(Eberhard 1993); Church Cave (WJ31), Wee Jasper (Hamilton-Smith unpublished data); Humicrib 
Cave (WJ34), (Eberhard 1993); Carey’s Cave (WJ100), Wee Jasper (Eberhard 1993); 
NULLARBOR PLAIN: Abrakurrie Cave (N3) (Richards 1971); Koonalda Cave (N4) (Richards 
1971); Mullamullang Cave (N37) (Richards 1971); Cocklebiddy Cave (N48) (Richards 1971); 
Moonera Tank Cave (N53) (Richards 1971); Thylacine Hole (N63) (Richards 1971); Old 
Homestead Cave (N83) (Richards 1971); Dingo Cave (N160) (Richards 1971). 


Undetermined genus and species, Gb. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) 
(Humphreys and Eberhard 2001); Upper Daniel Roux Cave (CI56) (Humphreys and Eberhard 
2001); NEW SOUTH WALES: Carrai Bat Cave (SCS), Stockyard Creek (Hamilton-Smith 
unpublished data); Cliefden Main Cave (CL1), Cliefden (Hamilton-Smith unpublished data); Willi 
Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith unpublished data); 
QUEENSLAND: Rope Ladder Cave (FR2), Mingella, Fanning River (Weinstein and Slaney 1995); 
Queenslander Tower (CH5246), Chillagoe (Matts 1987); Spring Tower (CH5223-5), Chillagoe 
(Matts 1987); Donna Tower (CH5155), Chillagoe (Matts 1987); Royal Arch Tower (CH5158-9), 
Chillagoe (Matts 1987); Tea Tree Tower (CH5137), Chillagoe (Matts 1987); Ryan Imperial Tower 
(CH5239), Chillagoe (Matts 1987); Wallaroo Tower (CH5201), Chillagoe (Matts 1987); Tower of 
London Cave (CH5) Chillagoe (Matts 1987); Kaiser Creek Cave (C12) (Two Mile Cave, Tar Drum 
Cave), Camooweal (Hamilton-Smith unpublished data); Holy Jump Lava Cave (BM1), Bauer’s 
Mountain (Hamilton-Smith unpublished data); VICTORIA: Anticline Cave (M11), Murrindal (Yen 
and Milledge 1990); Dickson Cave (M30), Murrindal (Yen and Milledge 1990); Nargun’s Cave 
(NN1), Nowa Nowa (Hamilton-Smith unpublished data); Grassmere Cave (W6), Warrnambool 
(Hamilton-Smith unpublished data). 


Undetermined Family 
Undetermined genus and species, Gp. CHRISTMAS ISLAND (Indian Ocean): Smiths Cave (CI9) 
(Humphreys and Eberhard 2001); Swiflet Cave (CI30) (Humphreys and Eberhard 2001); Managers 
Alcove (CI50) (Humphreys and Eberhard 2001); Grimes Cave (CI53) (Humphreys and Eberhard 
2001); Upper Daniel Roux Cave (CIS6) (Humphreys and Eberhard 2001). 


Undetermined genus and species, NULLARBOR PLAIN: Abrakurrie Cave (N3) (Hamilton-Smith 
unpublished data). 


Order Hymenoptera 

Braconidae 
Apanteles ?carpatus Say, Tp1, Gp. NEW SOUTH WALES: Humidicrib Cave (WJ34), Wee Jasper 
(Eberhard and Spate 1995); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003); 
VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 37 


CAVE GUANO ECOSYSTEMS AND INVERTEBRATE CHECKLIST 


Apanteles sp., Tp, Gp. NEW SOUTH WALES: Church Cave (W31), Wee Jasper (Hamilton-Smith 
unpublished data); Willi Willi Bat Cave (Main Cave) (WW1), Willi Willi (Hamilton-Smith 
unpublished data). 


Undetermined genus and species. Tp?. QUEENSLAND: Holy Jump Lava Cave (BM1), Bauer’s 
Mountain (Hamilton-Smith unpublished data). 


Formicidae 


Amblyopone australis Erichson, VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data). 


Iridomyrmex purpureus Smith, Tx, Gx. SOUTH AUSTRALIA: Eregunda Mine near Blinman, 
Flinders Ranges (T. Moulds unpublished data). 


Oligomyrmex sp., Tp?, Gx?. QUEENSLAND: Crazy Cracks Cave, Jacks Gorge, Broken River (T. 
Moulds unpublished data). 


Pachycondyla sp., Gp. CHRISTMAS ISLAND (Indian Ocean): Upper Daniel Roux Cave (CI56) 
(Humphreys and Eberhard 2001). 


Undetermined genus and species, NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper 
(Hamilton-Smith unpublished data); QUEENSLAND: Royal Arch Cave (CH9), Chillagoe 
(Hamilton-Smith unpublished data); Spring Cave, Mount Surprise (Hamilton-Smith unpublished 
data); SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Ichneumonidae 


Undetermined Cryptinae genus and species, Gp?. NEW SOUTH WALES: undetermined caves 
(Hamilton-Smith 1967); VICTORIA: Starlight Cave (W5), Warrnambool (Hamilton-Smith 
unpublished data); Spring Creek Cave (B1), Buchan (Hamilton-Smith unpublished data); Wilson’s 
Cave (EB4), East Buchan (Hamilton-Smith unpublished data). 


Myrmaridae 


Gonatocerinae sp., Gp?. SOUTH AUSTRALIA: Bat Cave (U2), Naracoorte (Bellati et al. 2003). 


Undetermined Family 


38 


Undetermined genus and species, Tp, Gp. NEW SOUTH WALES: Bungonia various caves 
(Eberhard 1998). 


Undetermined genus and species, Gp?. NEW SOUTH WALES: Church Cave (WJ31), Wee Jasper 
(Hamilton-Smith unpublished data); Willi Willi Bat Cave (WW1), Willi Willi (Hamilton-Smith 
unpublished data); VICTORIA: Panmure Cave (H5), Mount Napier (Hamilton-Smith unpublished 
data); Starlight Cave (W5), Warrnambool (Hamilton-Smith unpublished data). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T. MOULDS 


ACKNOWLEDGEMENTS 


This work was only possible due to the financial support 
of the Department of Environment and Heritage, South 
Australia, and the University of Adelaide. Thanks to Stefan 
Eberhard and Sue White for providing the stimulus for 
writing this paper. Many thanks to Elery Hamilton-Smith 
whose critical comments and unlimited access to his personal 
unpublished records greatly improved this paper. Thank you 
to Mike Gray, Courtenay Smithers and Judy Bellati who 
brought additional references to my attention. I also wish to 
thank John Jennings and Andy Austin for editorial comments. 


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42 Proc. Linn. Soc. N.S.W., 125, 2004 


A Devonian Brachythoracid Arthrodire Skull (Placoderm Fish) 


from the Broken River Area, Queensland 


Gavin C. YOUNG 


Department of Earth and Marine Sciences, Australian National University, Canberra ACT 0200 
gyoung @ geology.anu.edu.au 


Young, G.C. (2004). A Devonian brachythoracid arthrodire skull (placoderm fish) from the Broken River 
area, Queensland. Proceedings of the Linnean Society of New South Wales 125, 43-56. 


An incomplete brachythoracid arthrodire skull acid-prepared from the Devonian limestones of the 
Broken River area of Queensland is described as Doseyosteus talenti gen. et sp. nov. It supposedly comes 
from strata dated by conodonts as late Early Devonian in age (Emsian stage), but shows several derived 
features of the skull, typical of Middle-Late Devonian brachythoracids, and not seen in any arthrodire from 
the Emsian limestones of the Burrinjuck area of NSW. The alignment with conodont zones of stratigraphic 
subdivisions of the Burrinjuck sequence is revised. Published information on the provenance and age of all 
previously described placoderm taxa from Broken River is reviewed and amended. The new taxon may be 
most closely related to Late Devonian (Frasnian) brachythoracids from Iran and the Gogo Formation of 


Western Australia. 


Manuscript received 27 May 2003, accepted for publication 22 Oct 2003. 


KEYWORDS: Placoderm fishes, Arthrodira, Brachythoraci, Broken River, Devonian, new genus 


Doseyosteus, Queensland. 


INTRODUCTION 


Devonian sedimentary rocks, including many 
marine limestones, are well exposed in the Broken 
River area of Queensland (Fig. 1). Conodonts form 
the basis for dating the sedimentary sequence (Mawson 
and Talent 1989; Sloan et al. 1995). Vertebrate remains 
reported from this sequence include microfossils from 
many horizons (De Pomeroy 1996; Turner, Basden 
and Burrow 2000), and less well known vertebrate 
macro-remains. The latter include two genera of 
antiarch placoderms described by Young (1990), a 
ptyctodont toothplate ascribed to ?Ptyctodus sp. by 
Turner and Cook (1997), a new species of the 
brachythoracid arthrodire Atlantidosteus Leliévre 1984 
described by Young (2003a), an isolated suborbital 
plate of another arthrodire illustrated by Turner et al. 
(2000, fig. 8.7), and jaw remains of an onychodontid 
(Turner et al. 2000, fig. 5). Undescribed vertebrate 
macro-remains include various placoderm bones, most 
of which belong to brachythoracid arthrodires. The 
Arthrodira is the most diverse order within the class 
Placodermi, and its major subgroup, the Brachythoraci, 
comprises nearly 60% of about 170 genera within the 
Arthrodira (Carr 1995). The brachythoracid arthrodires 
were one of the most successful groups of early 


gnathostome fishes (e.g. Young 1986; Janvier 1996). 
In marine environments of the Late Devonian they 
included probably the largest predators of their time. 
The major radiation of brachythoracid subgroups had 
apparently already occurred by the Middle Devonian, 
and primitive representatives were already widespread 
in shallow marine environments of the Early Devonian 
(e.g. Young et al. 2001; Mark-Kurik and Young 2003), 
and are important in considering the origins and 
interrelationships of major brachythoracid subgroups 
(e.g. Leliévre 1995). 

The stratigraphic occurrence of various 
placoderm remains in the Broken River sequence were 
reviewed by Young (1993, 1995, 1996), De Pomeroy 
(1995, 1996), and Turner et al. (2000), and they have 
been mentioned in relation to conodont studies by 
Sloan et al. (1995). There has been conflicting 
information published about the provenance of some 
of the described placoderm taxa. These were collected 
from the Broken River area many years ago by 
Professor John Jell, University of Queensland, and sent 
to Canberra for acid preparation and study. In this paper 
I describe a new arthrodire skull from this collection, 
and review the locality information and age 
determinations for previously described placoderm 
taxa. 


Wurungulepis denisoni Young 1990 


specimen, it came from University of Queensland 


44 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


{vy +y CAINOZOIC [incl. basalt] 
ia L. DEVONIAN- 
La CARBONIFEROUS 


MIDDLE DEVONIAN 
= [Broken River Group] 
vw disconformity 


Feo EARLY DEVONIAN 
~ [72 ST SILURIAN 
nconformity 

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A FOSSIL FISH LOCALITIES 


unconformi 
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Figure 1. (A) location of the Broken River area in Queensland, Australia; (B) geological 
map of the collecting area (modified from Turner, Basden and Burrow 2000, fig. 2), 
showing localities for previously described placoderm taxa, and the specimen described 
in this paper (ANU V1026). 


LOCALITY AND AGE OF DESCRIBED 


PLACODERM TAXA FROM THE BROKEN locality L4399 (not L4339, given in error by Young 
RIVER AREA 1990: 45), on the north bank of the Broken River, Grid 


Reference 640 460 on the Burges 1:100 000 sheet, 
and was assigned a Middle Devonian (?Eifelian) age 
within the Broken River Formation (J.S. Jell, letter of 


According to information provided with this . ; 
17 April 1980). Judging by the map of the area 


Proc. Linn. Soc. N.S.W., 125, 2004 


published by Sloan et al. (1995: fig. 2), the 
locality lies within outcrop referred to as 
‘undifferentiated Broken River Group’. 

A ‘Wurungulepis-Atlantidosteus 
fauna’, of assumed Eifelian age, was listed in 
the macrovertebrate zonation of Young (1993, 
1995, 1996). However De Pomeroy (1995: 480) 
assigned Wurungulepis to the late Emsian 
serotinus Conodont Zone (CZ), citing a personal 
communication of J.A. Talent. This information 
was repeated by Turner et al. (2000: 498). Later 
(pers. comm. 28/8/95) J.A. Talent had advised 
A. Basden that this specimen was collected from 
the grid reference cited above, situated on a bend 
of the Broken River in an anticline, in strata 
which were pre-Dosey Limestone in the 
sequence, and equivalent to the Bracteata 
Formation and Lomandra Limestone (spanning 
the Emsian-Eifelian boundary; Sloan et al. 1995: 
fig. 3). 

No conodont data were obtained from 
the specimen, so its precise position relative to 
the standard conodont zonation is uncertain. 
Wurungulepis is an early representative of the 
asterolepidoid antiarchs, with a high short trunk 
armour (Young 1990), and was placed within 
the asterolepidoid clade adjacent to 
Sherbonaspis, and as sister group to Stegolepis, 
Asterolepis, Remigolepis and Pambulaspis, by 
Zhu (1996: fig. 29). As earlier discussed (Young 
1990: 48) the initially suggested Eifelian age 
was consistent with the oldest asterolepid 
(pterichthyodid) occurrence in Europe, cited as 
Gerdalepis from the Eifelian of Germany by 
Denison (1978), although this occurrence is 
slightly younger (early Givetian) according to 
Otto (1998: 118). However Gardiner (1994) 
cited Young (1974) for an older record (Emsian) 
of the asterolepid antiarchs, but the ‘cf. 
Pterichthyodes’ mentioned by Young (1974) 
was based on an erroneous attribution by Hills 
(1958: 88) to the Early Devonian limestone 
sequence of an ‘Antiarchan from Taemas’. In 
fact, the specimen in question came from the 
overlying Hatchery Creek Formation, of 
presumed Eifelian age (Fig. 2). This specimen 
was assigned to the new genus Sherbonaspis by 
Young and Gorter (1981). Previously, the 
suggested Emsian age of a pterichthyodid 
antiarch from the Georgina Basin (Young 
1984a) was noted as possibly the oldest 
occurrence of this group anywhere recorded. 


G.C. YOUNG 


costatus 


partitus 


patulus 


serotinus 


inversus- 
laticostatus 


perbonus- 
gronbergi 


dehiscens 


pireneae 


kindlei 


z 
< 
z 
O 
> 
tf 
Q 
a 
~ 
< 
ui 


Figure 2. Proposed alignment with conodont zones 
of subdivisions of the Early Devonian limestone 
sequence (Murrumbidgee Group) around 
Burrinjuck Dam, N.S.W., revised from Basden et 
al. (2000: fig. 2). Abbreviations for stratigraphic 
subdivisions are: B - Bloomfield Limestone 
Member; CB - Cavan Formation; CR - Crinoidal 
Limestone Member; CU - Currajong Limestone 
Member; HC - Hatchery Creek Formation; M - 
Majurgong Formation; R - Receptaculites 
Limestone Member; SY - Spirifer yassensis 
Limestone Member; W - Warroo Limestone 
Member; 1-6 - units of Upper Reef Formation. 
V1370 — horizon for highest known arthrodire in 
the sequence. 


in press), with the limestone occurrence yielding the 
antiarch probably younger than the diverse 


New evidence now indicates that two assemblages may 
have been mixed in this region (Burrow and Young, 


Proc. Linn. Soc. N.S.W., 125, 2004 45 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


Wuttagoonaspis fauna from underlying sandstones 
(Young and Goujet 2003). 

The antiarchs are a major subgroup of the 
class Placodermi, ranging in age from Early Silurian 
to latest Devonian. In recent years there has been a 
significant expansion in our knowledge of the group. 
A cladistic analysis of their distribution in relation to 
phylogeny by Young (1984b) involved 22 taxa and 
40 characters. In a recent review of antiarch phylogeny, 
Zhu (1996) noted some 45 genera and 154 species, 
and his data matrix used 66 characters for 40 genera. 
The original age assessment of Eifelian for 
Wurungulepis from Broken River is most consistent 
with our current knowledge of this large and diverse 


group. 


Nawagiaspis wadeae Young 1990 

This specimen is recorded from locality 
BRJ68D (University of Queensland locality L4428; 
‘small limestone outcrop on eastern side of gully 1 
km upstream from Six Mile yard’), Grid Reference 
596 442 on the Burges 1:100 000 sheet, which was 
assigned a Middle Devonian (?Givetian) age within 
the Broken River Formation (J.S. Jell, letter of 17 April 
1980). Apparently this specimen was found by Dr 
Mary Wade. 

Again, De Pomeroy (1995: 480) referred this 
taxon to the significantly older (late Emsian) serotinus 
CZ, based on its assigned position within the Bracteata 
Formation in section Br4 of Sloan et al. (1995, fig. 6). 
This information was repeated by Turner et al. (2000: 
498, 506). However Prof. J.A. Talent’s previous advice 
to the author (pers. comm. 5/8/92), was that this 
specimen was considerably younger (ensensis — varcus 
Zones; late Eifelian - Givetian). Clearly, there was 
some confusion about which fish specimen was being 
referred to. Subsequent advice given to A. Basden 
(pers. comm. 28/8/95), was that NV. wadeae came from 
the bank of Dosey Creek (Grid Reference 616 437), 
the location of section Br2 within outcrop of the 
Bracteata Formation (Sloan et al. 1995: fig. 2). The 
different, and presumably correct, locality information 
provided with the specimen, as cited above, 
corresponds to the vicinity of the boundary between 
the Papilio and Mytton Formations on the map of Sloan 
et al. (1995: fig. 2). This is consistent with the Givetian 
age first suggested by J.S. Jell. 

Nawagiaspis wadeae is another antiarch, 
originally interpreted as possibly a primitive 
bothriolepidoid (Young 1990), although in Zhu’s 
(1996) phylogeny it comes out as a basal 
asterolepidoid. Apart from primitive Chinese antiarchs, 
and the erroneous Emsian pterichthyodid occurrence 
discussed above, the stratigraphic record of this group 


46 


is Middle-Late Devonian (Gardiner 1994, fig. 32.1). 
The bothriolepidoid clade had an earlier history in Asia, 
and apparently expanded its range to most regions of 
the world in the Givetian (Young 2003b). 

The confusion about the provenance of this 
specimen may have resulted from the misconception 
that it was a recognisable ‘skull’ when collected. 
Turner et al. (2000) used this term to refer to the type, 
but the specimen as collected was a largely complete 
trunk armour, and the incomplete skull, missing its 
central portion, formed a minor part of the specimen. 
The whole specimen may have appeared to a non- 
vertebrate worker to represent a ‘skull’. Such fish 
remains, when collected in the field, are generally not 
determinable until after acid preparation (e.g. the type 
specimen of Atlantidosteus pacifica Young 2003a, 
before preparation, was assumed to be a ventral plate 
of the trunk armour, rather than a large suborbital bone 
from the cheek). 

A summary list of prepared fish remains from 
the original J.S. Jell collection was provided to J.A. 
Talent in 1995 to check on age and locality data. This 
list mentioned only one skull, the brachythoracid 
specimen described below, of which locality data 
provided by J.S. Jell are almost the same as stated by 
Sloan et al. (1995) for N. wadeae. Thus it seems that 
the specimen described below, previously listed as a 
‘skull’, has been confused with the type of N. wadeae, 
leading to erroneous locality and age information being 
given in De Pomeroy (1995), Sloan et al. (1995), and 
Turner et al. (2000). In the context of the global 
distribution in time and space of this major placoderm 
subgroup (see above), it is almost certain that 
Nawagiaspis is Middle Devonian in age, and a Givetian 
age, as first suggested by J.S. Jell, is most consistent 
with other information about the stratigraphic 
distribution of the more derived antiarchs. 


Atlantidosteus pacifica Young 2003a 

This specimen came from locality BRJ 67B 
(University of Queensland locality L 4472), Grid 
Reference 675 485 on the Burges 1:100 000 sheet, 
described as “Top of ridge to three-quarters way down 
western slope, west of road between Six Mile Dam 
and Diggers Creek’ (J.S. Jell, letter of 17 April 1980). 
This is the locality (with a slightly different grid 
reference) referred to as ‘Fish Hill’ by Turner et al. 
(2000: 507). They assigned it a middle Eifelian age 
(costatus - australis conodont zone), but noted that 
Sloan et al. (1995) gave a slightly longer partitus - 
early kockelianus zonal range for the Fish Hill section. 
This is consistent with the original assignment of a 
Middle Devonian (?Eifelian) age within the Broken 
River Formation by Prof. J.S. Jell. This occurrence is 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.C. YOUNG 


part of the evidence for proposing an Eifelian 
‘Wurungulepis-Atlantidosteus fauna’ in the 
macrovertebrate zonation of Young (1993, 1995, 
1996). 


Doseyosteus talenti gen. et sp. nov. 

This specimen, described below, was the only 
one in the J.S. Jell collection lacking a sample number 
at the time of preparation. It is highly probable that it 
was a sample collected the year before the other 
material, and was taken to Canberra separately by Dr 
P. Jell (J.S. Jell, letter of 17 April 1980). The following 
locality details, provided by Prof. J.S. Jell (letter of 17 
April 1980), indicate that it is the specimen collected 
from the alternative erroneous locality for Nawagiaspis 
just discussed: “‘BRJ34 = L 4054. Grid Reference 616 
438 Burges 1:100,000 sheet. Western bank of Dosey 
Creek, 750 m upstream from its junction with Broken 
River. Base of thick limestone lens in Broken River 
Formation, Middle Devonian. ? Eifelian’. 

In a published listing of University of 
Queensland locality numbers (Turner et al. 2000: 506), 
UQL4054 is assigned to ‘basal part of limestone, 
Lomandra/Dosey Limestone, Broken River Group’, 
with a slightly different grid reference (615 438), but 
the same locality description as above. However, it is 
assigned to the Emsian serotinus CZ, citing Sloan et 
al. (1995). 

Again, no conodonts were obtained from the 
sample, and section Br4 through the Bracteata 
Formation at this locality did not produce identifiable 
conodonts (Sloan et al. 1995: caption to fig. 6). 
Nevertheless, these authors (p.5) considered the entire 
formation to belong to the serotinus CZ, making it 
equivalent to the upper part of the Burrinjuck (NSW) 
limestone sequence, which extends from the top of 
the pirenae CZ (latest Pragian) into the serotinus CZ 
(the second youngest zone of the late Emsian). It is 
therefore relevant to make comparisons with the 
stratigraphic distribution of the diverse arthrodire 
assemblage described from the Burrinjuck limestone 
sequence. 

The described arthrodire fauna from the 
Burrinjuck sequence (White 1952, 1978; White and 
Toombs 1972; Young 1979, 1981, in press a, b; Young 
et al. 2001; Mark-Kurik and Young 2003) includes 10 
genera of brachythoracids, amongst which the most 
derived taxa (Cathlesichthys and Dhanguura) come 
from the upper part of the Wee Jasper limestone 
sequence. Basden et al. (2000, fig. 2) showed the 
youngest arthrodire skull from the Wee Jasper section 
(ANU V1370; the holotype of Dhanguura) to come 
from the uppermost unit 6 of the ‘Upper Reef 
Formation’ of Young (1969). This specimen is more 


Proc. Linn. Soc. N.S.W., 125, 2004 


advanced than other arthrodires known from the 
Burrinjuck sequence in possessing several derived 
characters of the skull, the most obvious being the T- 
shaped rostral plate, a feature of more derived 
eubrachythoracids (character 5 of Carr 1991; character 
4 of Leliévre 1995). Eubrachythoracids were the most 
diverse fish group of the Middle and Late Devonian, 
and the new Broken River brachythoracid described 
below clearly belongs to this group, with a skull which 
is more advanced in several respects than any of the 
known Burrinjuck arthrodires (see below). Gardiner 
(1994) lists the first occurrence of this grouping (his 
family Coccosteidae) as Coccosteus Miller 1841 from 
the Middle Devonian (Eifelian) of Scotland, for which 
a late Eifelian age is indicated by spores of the 
devonicus-naumovae zone (V.T. Young 1995). The 
same species (Coccosteus cuspidatus) is recorded from 
the Kernave Member of the Narva Formation in the 
Baltic sequence, although a related brachythoracid 
‘Protitanichthys’ occurs a little earlier, and in 
equivalent strata (costatus CZ) in the Rhenish sequence 
(Mark-Kurik 2000). However Otto (1997: 115) 
suggested that remains of early eubrachythoracids 
(coccosteids) first occur in the early Eifelian of 
Scotland, Germany, and the Baltic sequence. 
Dhanguura johnstoni Young (in press a) 
comes from a horizon about 420 m stratigraphically 
above the boundary equivalent of the Bloomfield and 
Receptaculites Members of the Taemas Limestone. A 
similar horizon high in the limestone sequence has 
produced the large lungfish Dipnorhynchus cathlesae 
Campbell and Barwick 1999. The lungfish locality is 
close to localities L537 and L538 of Pedder et al. 
(1970) which yielded tetracorals Vepresiphyllum 
dumosum, Sulcorphyllum pavimentum, 
Chalcidophyllum vesper and C. gigas. This represents 
the uppermost ‘tetracoral teilzone’ of the 
Murrumbidgee Group (Pedder et al. 1970: fig. 4), and 
is Coral Fauna F in the scheme of Garratt and Wright 
(1989). These authors considered the succeeding G 
and H Coral Faunas to overlap, and belong to the late 
Emsian, rather than Eifelian as previously assessed. 
Garratt and Wright (1989) also aligned Coral Fauna F 
from Wee Jasper (and the Sulcor Limestone of northern 
NSW) with the mid-Emsian inversus CZ (see column 
13 of Young 1995, 1996). However Basden et al. 
(2000: fig. 2) showed the uppermost beds of the 
limestone sequence at Wee Jasper (containing Coral 
Fauna F) extending well into the next youngest 
serotinus CZ. Evidence supporting this (summarised 
by Basden 2001, table 2.1) derives from reassignment 
of some of the conodonts from the highest productive 
sample (C62) in Pedder et al.’s (1970) section 2, 
referred by them to Polygnathus linguiformis 


47 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


linguiformis, but reassigned to Polygnathus inversus 
by Klapper and Johnson (1975), and to Polygnathus 
serotinus (delta morphotype) by Mawson (1987). On 
the other hand, the age in terms of conodont zone 
alignment of several constituent members of the 
Taemas Limestone, as indicated by Basden et al. (2000, 
fig. 2), seem to be too young, and should be revised 
downwards on the following evidence. Lindley (2002a: 
275) noted that the occurrence of the index species of 
Coral Fauna D (Chalcidophyllum recessum) in the 
Currajong Limestone Member indicates that it should 
be aligned with the dehiscens rather than the perbonus 
CZ. The overlying Bloomfield Limestone Member 
may also have lower beds of dehiscens rather than the 
perbonus CZ age (Basden 2001: table 2.1). The Warroo 
Limestone Member contains perbonus CZ elements 
(Nicoll, in Lindley 2002b), and the uppermost 
Crinoidal Limestone Member in the Taemas sequence 
may align with both the inversus and the serotinus CZ 
(Basden 2001: table 2.1). 

These revised alignments are summarised in 
Fig. 2. Correlation with the upper part of the Wee 
Jasper sequence is unclear, because the constituent 
members of the Taemas Limestone are difficult to 
recognise in the thicker upper part of the sequence, 
represented by units 1-6 of Young (1969). If the new 
arthrodire skull described below from Broken River 
is of serotinus CZ age, as proposed by Sloan et al. 
(1995), it is still considerably more derived (see below) 
than any arthrodire from the Burrinjuck sequence. If 
correctly dated, this would indicate that derived 
features characterising the Middle-Late Devonian 
eubrachythoracid arthrodires had originated at least 
by late Emsian time. 

To summarise, it is emphasised that there is 
no overlap in the arthrodire skull characters just 
discussed between the Burrinjuck and Broken River 
limestone sequences, even though the youngest 
occurrences in the former sequence are also the most 
derived taxa within the better-documented Burrinjuck 
arthrodire fauna. For the new taxon described below, 
this evidence would support either a latest Emsian age 
(but younger than the Burrinjuck sequence), or an 
Eifelian age as originally suggested by Prof. J.S. Jell. 


ABBREVIATIONS 


The specimen described below (prefix ANU 
V) is housed in the Earth and Marine Sciences 
Department, Australian National University, Canberra 
(GCY Vertebrate Collection). Standard abbreviations 
for placoderm dermal bones are used in the text and 
figures, and together with other morphological 
abbreviations are listed as follows: 


48 


anth, anterior nuchal thickening; 

Ce, central plate; 

cf.Ce, area overlapping Ce plate; 

cf.M, area overlapping marginal plate; 

cf.PM, area overlapping postmarginal plate; 
cf.PtO, area overlapping postorbital plate; 
cr.im, inframarginal crista; 

csc, central sensory line canal; 

d.end, openings of dermal tube for endolymphatic 
duct; 

dep, depression; 

gr.M, groove on Ce plate which received the edge of 
the marginal plate; 

ifc.ot, otic branch of infraorbital sensory groove; 
if.r, infranuchal ridge; 

if.pt, infranuchal pit; 

kb, knob-like thickening of inframarginal crista; 
Icp, lateral consolidated part of skull roof; 

llc, main lateral line sensory canal; 

M, marginal plate; 

mp, middle pitline; 

mppr, posterior median process of nuchal plate; 
Nu, nuchal plate; 

oa.Ce, area overlapped by Ce plate; 

oa.M, area overlapped by M plate; 

oa.Nu, area overlapped by Nu plate; 

orb, orbital notch; 

Pi, pineal plate; 

plpr, posterolateral process or lobe on Ce plate; 
PM, postmarginal plate; 

pmc, postmarginal sensory groove; 

pnp, postnuchal process of paranuchal plate; 
PNu, paranuchal plate; 

pp, posterior pitline; 

PrO, preorbital plate; 

PtO, postorbital plate; 

R, rostral plate; 

soa, subobstantic area; 

soc, supraorbital sensory canal; 

th.end, endolymphatic thickening; 

th.pre, pre-endolymphatic thickening; 

tnth, transverse nuchal ridge or thickening; 

vg, vascular grooves. 


SYSTEMATIC PALAEONTOLOGY 


Class PLACODERMI McCoy, 1848 
Order ARTHRODIRA Woodward, 1891 
Suborder BRACHYTHORACTI Gross, 1932 


Doseyosteus talenti gen. et sp. nov. 
Name 

From Dosey Creek, the type locality, and the 
Greek osteus (bone). The species name recognises 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.C. YOUNG 


Professor John A. Talent, Macquarie University, who 
has had a long and distinguished career in Devonian 
research, including extensive work in the Broken River 
area of Queensland. 


Diagnosis 

A eubrachythoracid arthrodire in which the 
skull shows an embayed anterior margin of the nuchal 
plate resulting from overlap by the central plates, the 
central plates have strong posterolateral lobes 
separating the nuchal and paranuchal, and a mesial 
process of the marginal plate extends to the anterior 
angle of the paranuchal. Subobstantic area of skull 
extending onto marginal plate. Dermal bones smooth, 
or ornamented with fine tubercles. 


Remarks 

Since only the skull is known, and it is 
incomplete, several features characterising the derived 
subgroup “Eubrachythoraci’ are for the present inferred 
for this new taxon. Definition of the eubrachythoracid 
arthrodires is discussed by Carr (1991: 379-381) and 
Long (1995: 55). Thus Doseyosteus talenti gen. et sp. 
nov. is assumed to have had a T-shaped rostral plate, a 
posteriorly placed pineal plate separating the 
preorbitals, a dermal process of the preorbital plate 
forming the anterodorsal margin of the orbit, and 
trilobate central plates. The holotype shows a strongly 
developed posterior thickening of the skull roof, which 
in the midline is represented by the anterior nuchal 
thickening. This is much more prominent than the 
transverse ridge on the posterior margin of the nuchal 
plate, and is a derived feature seen in coccosteomorph 
and pachyosteomorph brachythoracids, but generally 
lacking in Early Devonian taxa, for example the genus 
Cathlesichthys from Burrinjuck, NSW (Young, in press 
a). The embayed anterior margin and inferred 
proportions of the nuchal plate, and the strong 
posterolateral lobe of the Ce plate, are resemblances 
to the Late Devonian taxa Eastmanosteus and 
Golshanichthys, but the former differs in having the 
posterior pitline well developed on the posterolateral 
lobe of the central plate, and both forms lack the mesial 
process of the marginal plate inferred for this new 
taxon. 


Material 
ANU V1026 (holotype), an incomplete skull 
preserved as two unconnected portions. 


Locality and Horizon 

Locality BRJ34 (University of Queensland 
locality L4054), Grid Reference 616 438, Burges 1:100 
000 sheet; western bank of Dosey Creek, 750 m 


Proc. Linn. Soc. N.S.W., 125, 2004 


upstream from its junction with Broken River (J.S. Jell, 
letter of 17 April 1980; see discussion above). Horizon 
was described as the ‘base of thick limestone lens in 
Broken River Formation’, assigned to the Bracteata 
Formation (Sloan et al. 1995) or the ‘Lomandra/Dosey 
Limestone, Broken River Group (Turner et al. 2000). 
Age: ?late Emsian - Eifelian (see discussion above). 


Description 

ANU V1026 represents a large part of the 
posterolateral region of a brachythoracid skull roof, 
preserved as two separate portions. The larger portion 
(Fig. 3A,D) includes parts of the Nu, PNu and Ce plates 
(Fig. 4A,B), and the right postmarginal corner of the 
skull is preserved as a separate portion (Figs. 3B,C, 
4C,D). The specimen was extracted from the rock in 
six pieces, but they are well preserved, suggesting that 
it was broken up before incorporation in the sediment. 
The nuchal (Nu) plate is represented by most of its 
right half, including the midline, so its overall shape 
can be estimated. Midline length of the Nu is about 70 
mm. It has an embayed posterior margin, with a 
prominent posterior median process (mppr, Fig. 4). 
Except for the posterior lateral corner the right lateral 
margin of the Nu plate is fairly well displayed on the 
external surface. The bone is fractured in its middle 
region, and shows anteriorly that it was both 
overlapped and underlapped by the central (Ce) plate, 
a condition also reported in Holonema (Miles 1971). 
Along the anterior margin of the plate a thin 
overlapping lamina of the Ce plate has broken away 
to reveal an extensive overlap area (0a.Ce, Fig. 4B). 
In unbroken condition the anterior margin of the Nu 
plate would have been deeply embayed (Fig. 5). On 
its visceral surface extensive contact faces for the 
central plates are developed in the normal manner 
(cf.Ce, Fig. 4A). Other features shown are the 
prominent infranuchal pits (if.pt) and ridge (if.r) and 
the transverse nuchal thickening or ridge (tnth). 

Noteworthy is the strong development of the 
anterior nuchal thickening (anth), which is relevant to 
the question of the age of this specimen (see discussion 
above). This is a derived feature of brachythoracids, 
and in ANU V1026 is more pronounced than in any 
Emsian brachythoracid from the Burrinjuck fauna. 
These have Nu plates which are fairly flat in front of 
the infranuchal pits. This is the case even in a form 
like Cathlesichthys, which is derived in having a very 
strong transverse nuchal ridge (Young in press a). In 
posterior view ANU V1026 shows that the anterior 
nuchal thickening is more pronounced than the 
transverse nuchal ridge, the reverse of the condition 
in Cathlesichthys. This advanced character is also seen 
in most Middle-Late Devonian brachythoracids, such 


49 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


Figure 3. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). Larger (A,D) and smaller (B,C) 
skull portions in external (B,D) and internal (A,C) views. 


as Golshanichthys, Tafilalichthys, and various Gogo 
forms (e.g. Leliévre et al. 1981; Leli¢évre 1991; Miles 
and Dennis 1979; Long 1988, 1995; Dennis-Bryan 
1987). These taxa all resemble the giant Famennian 
form Dunkleosteus, where the ‘posterior consolidated 
arch’ of the skull roof (‘PCA’ of Heintz 1932: fig. 13) 
is a broad thickening running in front of the infranuchal 
pits, as the main transverse thickening of the skull. In 
contrast, in the Early Devonian form Cathlesichthys 
from Burrinjuck, the transverse nuchal ridge located 
behind the infranuchal pits forms the main thickening 
supporting the posterior skull margin. 

The right paranuchal (PNu) plate of 
Doseyosteus gen. nov. is represented externally by an 
elongate portion including the mesial margin forming 
sutures with the Nu and Ce plates (PNu, Fig. 4B). There 
is also a small broken part of the postnuchal process 
(pnp). The PNu and Ce plates were also connected by 
a complex interlocking suture; a broken part around 


50 


the anterior end of the PNu exposes an overlap area 
(oa.Ce, Fig. 4B), and the edge of a more extensive 
contact face is shown on the visceral surface (cf.Ce, 
Fig. 4A). The endolymphatic thickening forms a broad 
thickened area mesially (th.end), combining with the 
thickened portion of the Nu plate (anth). This thickened 
part of the skull is much more prominent than in 
primitive brachythoracids like Buchanosteus or 
Taemasosteus (White 1978; Young 1979). Along the 
broken edge of the specimen, maximum bone thickness 
(in the part enclosing the endolymphatic duct) is almost 
15 mm, which is three times the bone thickness at the 
anterior preserved extremity of the Nu. The exoskeletal 
division of the right endolymphatic duct opens on the 
visceral skull roof surface at the anterior edge of the 
area of thickened bone (th.end), and is also visible on 
the broken margin of the specimen (d.end, 
Fig.4A).This is also an advanced character of the 
brachythoracid skull — in large Emsian brachythoracids 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.C. YOUNG 


Figure 4. Doseyosteus talenti gen. et sp. nov. Holotype (ANU V1026). A,B. Larger portion of skull in 
internal (A) and external (B) views. C,D. Smaller skull portion in internal (C) and external (D) views. 


from Burrinjuck the endolymphatic duct is not within 
the bone, but anteriorly forms a bony tube attached to 
or projecting from the inner surface of a much thinner 
PNu plate (Young in press a: figs. 3, 4, 7A, 9B). A 
similar condition occurs in Holonema from Gogo (J.A. 
Long, pers. comm.; Miles 1971: fig. 53). 

The preserved part of the right Ce plate is 
crossed by a prominent sensory groove (csc), which 


Proc. Linn. Soc. N.S.W., 125, 2004 


must be the central sensory canal rather than the 
supraorbital sensory canal, because of its oblique 
orientation to the midline. Middle and posterior pitlines 
are represented by faint markings in the region of the 
ossification centre (mp, pp). Anterolateral and 
posterolateral margins of the preserved part of the Ce 
plate are somewhat fractured, but appear to 
approximate natural margins. The former is bevelled 


51 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


Ses fas ie 


~—--— - 


I le 


= 


/ Y 
| , 
Oe ; 
| / 
H DD, Fy 
] / t 
/ / : b 
\ Fi Pro ee 
s IN 
wet \ / fate 
7 I} : : Pt * 
\ N 
<4 I = Vag | 
| ! 
| H 
| t 
Ce | 
| \ 
\ \ 


Figure 5. Doseyosteus talenti gen. et sp. nov. Attempted skull roof reconstruction, preserved 


portions shaded. 


externally, and internally shows a contact face for the 
postorbital plate (cf.PtO), showing that it overlapped 
the PtO extensively, as in most other brachythoracids 
(e.g. Miles and Westoll 1968: fig. 2; Young 1979: fig. 
1; 1981: fig. 5). Holonema is an exception in this 
respect (Miles 1971: fig. 12). Subdivisions of the 
posterior part of this contact face suggest that it also 
overlapped the marginal (M) plate (cf.M, Fig. 4A). 
The posterolateral margin of the Ce plate is 
somewhat thicker, and carries a deep groove (gr.M) 
for an interlocking suture, the Ce plate providing 
external and internal laminae to enclose the margin of 
the contiguous bone. The nature of the preserved 
margins suggests that they approximate the suture 


52 


position. Since the anterior end of the PNu is well 
shown on the specimen, and is most unlikely to have 
extended to this margin of the Ce plate, it seems that 
the intervening space must have been occupied by a 
mesial projection of the M plate (M, Fig. 5). This 
arrangement has not previously been recorded in 
brachythoracids. A similar but smaller process of the 
M intrudes the Ce plate of Buchanosteus, but this is 
some distance in front of the PNu (Young 1979: fig. 
1). 

There is a long posterolateral projection of 
the Ce plate partly separating the Nu and PNu plates 
(plpr, Fig. 4B), a feature seen in various other 
brachythoracids. An early example with this 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.C. YOUNG 


morphology is Ulrichosteus Leliévre, 1982a from the 
Givetian of Germany, but this form has the Nu plate 
extending anteriorly in front of the PNu, whereas in 
Doseyosteus the PNu is slightly longer. Ardennosteus 
Leliévre, 1982b also has a strong posterolateral lobe 
of the Ce, but this Famennian form differs in its sinuous 
interlocking sutures, broader transverse nuchal 
thickening, and coarse tubercular ornament. 
Development of a posterolateral lobe of the Ce is one 
of three features representing the ‘trilobate’ condition 
of the Ce plates (characters 13, 14, 21 of Carr 1991), a 
widespread condition amongst Middle-Late Devonian 
eubrachythoracids which has proved difficult to define. 
Internally this part of the Ce is more extensive, the 
overlapped portion extending back to the 
endolymphatic thickening, again as in other 
brachythoracids. The visceral surface of the Ce is 
gently concave laterally, with several shallow grooves 
(vg) resembling the vascular grooves described in 
Holonema by Miles (1971: fig.12). This depressed 
region is flanked mesially by the pre-endolymphatic 
thickening (th.pre), which forms a low broad ridge with 
a curved anteromesial orientation. The preserved 
anteromesial edge of the Ce plate is thickened and 
abraded (Fig. 3D). 

Associated with this skull portion was a 
smaller part of the left preobstantic corner of the skull 
roof (Fig. 3B,C), assumed to have belonged to the same 
individual. The specimen includes part of the PNu and 
M plates (Fig. 4C,D), and is crossed by a section of 
the main lateral line (IIc), and the infraorbital (ifc.ot) 
and postmarginal (pmc) sensory canals. Unlike forms 
such as Coccosteus, Holonema and Buchanosteus 
(Miles and Westoll 1968; Miles 1971; Young 1979), 
the M plate carries part of the subobstantic area (soa, 
Fig. 4D). A subobstantic area of similar extent is seen 
in the Gogo brachythoracid Harrytoombsia Miles and 
Dennis (1979: fig. 4), and in all plourdosteids sensu 
Long (1995). The PM plate is missing, but on the 
visceral surface there is a clear contact face for this 
bone (cf.PM, Fig. 4C). The visceral surface also shows 
the inframarginal crista to be strongly developed, 
dorsally as a very prominent irregular knob of bone 
(kb) separated posteriorly by a deep groove from the 
ventrally directed crista (cr.im), which itself carries a 
groove. The free ventral margin of the plate is 
thickened (Icp), representing the ‘lateral consolidated 
part’ of the skull, and a depression between the 
thickening and the inframarginal crista (dep) may 
correspond to similar structures in Coccosteus and 
Buchanosteus Young (1979: 314). 

The external ornament on both specimens 
comprises fine tubercles in some areas, sometimes only 
faintly discernible on a generally smooth surface (Fig. 


Proc. Linn. Soc. N.S.W., 125, 2004 


3B,D). The fine ornament is similar to that on the SO 
plate of Atlantidosteus pacifica, but that form displays 
affinity with the homostiid arthrodires in a range of 
features (Young 2003a), whereas the skull of 
Doseyosteus talenti gen. et sp. nov. lacks various 
specialised characters of Homostius and related forms 
(e.g. elongate Nu and PNu plates, small dorsal orbits, 
etc.). The reduced ornament also distinguishes this new 
form from various ‘coccosteomorph’ arthrodire 
remains known from the early Middle Devonian of 
northern Germany and the Baltic sequence (Otto 1997, 
1999). 

An attempted reconstruction of the skull roof 
of the new taxon based on available information is 
presented in Fig. 5. The skull could have been broader 
across the preobstantic corners than shown, since the 
gap between the two preserved portions is based only 
on a general alignment of sutures and sensory grooves. 
The anterior part of the skull is unknown, and restored 
shape of bones is generally based on various 
coccosteomorph arthrodires (e.g. Denison 1978: fig. 
57). Advanced features depicted (T-shaped R plate, 
Pi plate separating PrO plates, trilobate Ce plates) are 
based on their co-occurrence with preserved skull 
characters in all other known taxa. They need to be 
confirmed with additional material. On the larger 
preserved portion, the breadth and anterior embayment 
of the Nu plate, and the marked posterior lobe of the 
Ce plate separating the Nu and PNu plates, are general 
resemblances to Eastmanosteus and Golshanichthys, 
as noted above. The M and Ce plates retain extensive 
contact to separate the PNu from the PtO, the assumed 
primitive condition for brachythoracids. In contrast, 
the plourdosteid arthrodires, which were widespread 
in the Late Devonian, and apparently replaced the 
largely Middle Devonian coccosteids (Long 1995), 
have a much enlarged PtO reaching back to contact 
both the Ce and PNu plates. In consequence the M 
plate is reduced in size, whereas in Doseyosteus gen. 
nov., although not completely preserved, the M plate 
was clearly a more extensive bone, which apparently 
shows a unique feature in the large mesial process 
embaying the Ce plate in front of the PNu. 

In summary, this new but poorly known 
brachythoracid shows a range of advanced characters 
otherwise only seen in Middle or Late Devonian taxa, 
and it resembles the Frasnian taxa Eastmanosteus and 
Golshanichthys in several features which might 
indicate a close relationship. Eastmanosteus 
yunnanensis Wang, 1991 from the Givetian of China 
would otherwise be the earliest known member of this 
group (family Dinichthyidae). Kiangyousteus Liu, 
1955, also from China (Givetian of Szechuan), may 
be another primitive dinichthyid (Denison 1978). Both 


53 


DEVONIAN ARTHRODIRE SKULL FROM QUEENSLAND 


taxa differ from the new form described here in their 
well-developed coarse tubercular ornament, 
presumably a primitive feature. Doseyosteus talenti 
gen. et sp. nov. displays an unusual shape of the M 
plate which is apparently unique to this new genus 
and species. More material, including the unknown 
trunk armour, which in brachythoracids comprises 17 
separate bones, will clarify the affinities of this new 
taxon. 


ACKNOWLEDGMENTS 


Professor J.S. Jell (University of Queensland) and 
Professor K.S.W. Campbell (ANU) are thanked for making 
the specimen available for study. Mr R.W. Brown 
(Geoscience Australia) assisted in acid preparation. Professor 
J.A. Talent and Dr A. Basden (Macquarie University) advised 
and discussed at length the provenance and age of Broken 
River placoderms, and Dr S. Turner (Queensland Museum) 
provided comparative material. Comparison with European 
and Moroccan arthrodire material was facilitated by a visiting 
professorship at the Muséum national d’Histoire naturelle, 
Paris, in 1999. Professor D. Goujet is thanked for arranging 
this, and for the provision of facilities, and together with Dr. 
H. Leliévre and Dr. P. Janvier discussed at length placoderm 
morphology and relationships. Dr Leliévre arranged for 
arthrodire casts to be sent to Canberra for comparative study. 
B. Harrold is thanked for providing essential computer 
support at ANU, and V. Elder for assistance with specimen 
curation. Dr E. Mark-Kurik and Dr R. Carr discussed 
arthrodire phylogeny, and Dr Carr arranged for a visit to 
Cleveland, Ohio, for study of large arthrodire material. 
Financial support was provided in Canberra by ANU 
Faculties Research Fund Grants FO1083 and FO2059, and 
overseas by the Alexander von Humboldt Foundation, for a 
Humboldt Award in Berlin (2000-2001), and assistance with 
travel to the USA (Flagstaff and Cleveland, 2000). I thank 
Prof. H.-P. Schultze for provision of facilities in the Museum 
fiir Naturkunde, Berlin. Dr P. De Deckker is thanked for 
provision of facilities in the Geology Dept., ANU. This 
research was a contribution to IGCP Projects 328, 406, 410, 
and 491. 


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Proc. Linn. Soc. N.S.W., 125, 2004 


Effects of Slashing and Burning on Thesium australe R. Brown 


(Santalaceae) in Coastal Grasslands of NSW 


JANET S. COHN 


Biodiversity Research and Management Division, NSW National Parks and Wildlife Service, PO Box 1967, 


Hurstville, NSW 2220 (Ganet.cohn@npws.nsw.gov.au) 


Cohn, J. (2004). Effects of slashing and burning on Thesium australe R. Brown (Santalaceae) in coastal 
grasslands of NSW. Proceedings of the Linnean Society of New South Wales 125, 57-65. 


Two studies examined the effects of burning and cutting on aspects of the population dynamics of 
a nationally vulnerable herb, Thesium australe on the central and north coast of NSW. Study sites were 
grasslands dominated by Themeda australis with scattered native shrubs (Banksia integrifolia, Acacia 
sophorae) and the exotic shrub Chrysanthemoides monilifera ssp. rotundata. In the first study (May 1995 to 
December 1996), Thesium australe occurred at high density (1/m?) on exposed, long-unburnt headlands. In 
the second study, (December 1996 to December 1998), Thesium australe was at low density (<1/100m7) on 
more protected and recently burnt hinterland. On the headlands, winter treatments had no significant effect 
on the survival, density and vigour of Thesium australe. In the hinterland, one year after summer treatments, 
seedling recruitment resulted in a higher density of Thesium australe in the cut plots than either the burnt or 
the control. Flowering and fruiting of Thesium australe were not restricted by season. After winter and 
summer treatments, flowering and fruiting occurred within 6 months and 1 year, respectively. Although 
exposed coastal headlands may require no management intervention to increase the occurrence of Thesium 
australe, except where the possibility of shrub invasion exists, a regime of slashing on less exposed hinterlands 
may be needed to reduce competition from Themeda australis. Further research is necessary to determine if 
slashing or burning the more protected hinterland would yield different results if carried out in seasons other 


than summer. 


Manuscript received 1 March 2003, accepted for publication 22 October 2003. 


KEYWORDS: fire, grasslands, headlands, mowing, slashing, Thesium australe. 


INTRODUCTION 


Although Thesium australe is a herb with a 
wide ecological tolerance, extending from tropical to 
alpine climates, it is confined to widely scattered 
locations in open woodlands and grasslands where 
Themeda australis/ T. triandra (Kangaroo Grass) is 
common in the understorey (Scarlett et al. 1994). On 
the north coast of NSW, T. australe occurs on grassy 
headlands used predominantly for passive recreation, 
often adjacent to residential areas (Griffith 1992; Fig. 
1). 

In south-eastern Australia, open woodland and 
grassland communities have largely been modified and 
fragmented by introduced grazers, cultivation and 
changed fire regimes (Stuwe and Parsons 1977; Scarlett 
and Parsons 1990; McDougall and Kirkpatrick 1994; 
Tremont and McIntyre 1994; Prober and Thiele 1995; 
Lunt 1997). As a consequence T. australe is rated as 
nationally vulnerable (Briggs and Leigh 1996) and 
vulnerable in NSW under Schedule 2 of the NSW 
Threatened Species Conservation Act 1995 


Non-coastal, long-unburnt grasslands 
dominated by Themeda australis / triandra, have been 
shown to be species poor (Stuwe and Parsons 1977; 
Kirkpatrick 1986; McDougall 1989), largely as a result 
of the high competitive ability of this tussock grass 
(Groves 1974). With a general recent decline in fire 
frequency on coastal headlands (Griffith 1992), 
dominance by Themeda australis and the recruitment 
of native and exotic shrubs are potential threats to the 
survival of T. australe (Griffith 1992), although Cooper 
(1986) suggested that headlands exposed to salt-laden 
winds may be an exception. He cites the persistence 
of T. australe at Perpendicular Point, 20 years after 
fire, as an example. 

Research on Thesium alpinum in Denmark 
found that it became extinct as a result of shading from 
trees (Lojtnant and Worsoe 1980). Thesium australe 
may be similarly sensitive. In coastal Victoria, an 
increase in native shrub and tree recruitment has been 
linked to a decline in fire frequency (Bennett 1994; 
McMahon et al. 1994; Lunt 1998a b). On the north 
coast of NSW, increased recruitment of native shrubs 


EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 


Figure 1. Grassland habitat of Thesium australe 
at Look at Me Now Headland on the north coast 
of NSW. 


and trees (Acacia, Banksia, Allocasuarina spp.) and 

an invasive exotic shrub, Bitou bush, 

(Chrysanthemoides monolifera ssp. rotundata have 

been observed (Dodkin and Gilmore 1985; Griffith 

1987; Griffith 1992). 

A number of studies have suggested a regime 
of regular burning and/or mowing to maintain species 
richness in grasslands and prevent shrub invasion 
(Groves 1974; Stuwe and Parsons 1977; Kirkpatrick 
1986; McDougall 1989; Lunt 1990a, 1998b). Current 
information on the response of T. australe to fire in 
the field has been based on observations. While Leigh 
and Briggs (1989) suggest that survival and recruitment 
are unaffected by fire, Archer (1984) believed seeds 
were stimulated to germinate. In laboratory trials, 
Scarlett (pers. comm.) found that heat did not stimulate 
seed germination. There have been no studies on the 
effect of mowing or cutting on T. australe (Griffith 
1992). 

On coastal headlands and conservation reserves 
where burning or slashing grasslands may be used for 
conservation or hazard reduction purposes, it is 
important to establish their effect on native species. 
Two separate studies examined the effects of a single 
burning and a single cutting on aspects of the 
population dynamics of T. australe, namely: 

1/ its survival, density, vigour and reproductive status 
where it occurred at relatively high density on 
long-unburnt, exposed headlands (winter 
treatments); 

2/ its density and reproductive status where it was at 
very low density in a more protected and 
recently burnt hinterland (summer treatments). 
These were not intended to be comparative 

studies and indeed the different timing and methods 

of treatment (see Materials and Methods), driven by 


58 


the availability of resources, make this not possible 
anyway. 


MATERIALS AND METHODS 


The studies were located at several sites on the 
north and central coast of NSW (Fig. 2): Perpendicular 
Point (AMGR Easting 485600, Northing 6499200); 
Look at Me Now Headland (E 518000, N 6661300); 
and Old Bar Park (E 461300, N 6462800). 
Perpendicular Point and Look at Me Now Headland 
are within respectively, Kattang Nature Reserve (NR) 
and Moonee Beach NR. Both are managed by the New 
South Wales National Parks and Wildlife Service 
(NSW NPWS). Old Bar Park is managed by The 
Greater Taree City Council. 

All three sites are used for recreation, 
predominantly by walkers. Although no motor 
vehicular access is allowed in the NRs, there was 
evidence of their past usage at Look at Me Now 
Headland, where at the time of this study wheel ruts 
were still very obvious. Vehicles were used on 
Perpendicular Point as recently as 1986 (Cooper 1986). 
There is some use of motor vehicles in Old Bar Park, 
but this is mostly on the pre-existing tracks and the 
airstrip (author’s personal observations). 

Perpendicular Point and Look at Me Now 
Headland are characterised by black headland soils, 
which are loamy soils high in organic matter (Parbery 
1947). Yellow podzolic soils predominate at Old Bar 
Park (Long 1996). Aspects and slopes of the study sites 
varied. At Perpendicular Point the site was located on 
a north-western aspect with a slope of 9°, whilst at 
Look at Me Now Headland the site was on a more 
exposed southerly aspect with a slope of 6°. The site at 
Old Bar Park was flat. 

The study sites were in grassland communities 
dominated by Themeda australis. Scattered shrubs at 
Perpendicular Point included native (e.g. Acacia 
sophorae, Banksia integrifolia) and exotic (e.g. 
Chrysanthemoides monilifera ssp. rotundata) taxa. 
Another nationally endangered herb, Zieria prostrata 
(Briggs and Leigh 1996) also occurred on a number 
of the headlands with T. australe (Griffith 1992; NPWS 
1998). 

Thesium australe was found at Perpendicular 
Point in 1957 (Cooper 1986) and at Look at Me Now 
Headland and Old Bar Park after 1992 (Griffith 1992). 
Although at relatively high density at Perpendicular 
Point and Look at Me Now Headland (approximately 
1/m*), at Old Bar Park it occurred mostly as very 
scattered plants (approximately <1/100 m7’). Thus, the 
focus at this latter site was more on recruitment 


Proc. Linn. Soc. N.S.W., 125, 2004 


J.S.COHN 


Moonee Beach 
Nature Reserve 


Coffs Harbour 


kilometres 


Port Macquarie 


Kattang Nature 
Reserve 
Map 
location 


¢ 


Figure 2. Locality of study sites within Moonee Beach NR (Look at Me Now Headland), Kattang NR 
(Perpendicular Point) and Old Bar Park on the north and central coast of NSW. Stippled areas represent 
estate managed by NSW National Parks and Wildlife Service. 


responses to treatments. 

There was little information on the fire history 
at the three sites. Griffith (1992) believed that 
Perpendicular Point may not have burnt for a 
considerable period of time. In 1985, Cooper (1986) 
believed that Perpendicular Point had not burnt for at 
least 20 years. There was no record of the last fire at 
Look at Me Now Headland. Old Bar Park was last 
burnt in 1991 by a low intensity fire (T. Cross pers. 
comm.). Approximately 1 year prior to this study Old 
Bar Park was slashed (S. Griffith pers. comm.), 
presumably for hazard reduction purposes. 


Headlands (high density plants) 

Treatments were applied in winter (July 1995) 
at Perpendicular Point and Look at Me Now Headland 
(Table 1). There were 15 replicate plots of each 
treatment (burnt, cut, control). Each treatment was 
allocated randomly to a 0.5 m x 0.5 m plot laid out in 
rows, over a total area of 75 m? at Perpendicular Point 
and 112 m? at Look at Me Now Headland. Plots were 


Proc. Linn. Soc. N.S.W., 125, 2004 


burnt using a gas burner. Because of the heavy dew, 
each burnt plot was subjected to heat for 5 minutes, 
until all of the grasses and herbs had been burnt and 
the bare ground had been heated and scorched. This 
simulated a high intensity burn (R. Bradstock pers. 
comm.). In the cutting treatment all grasses and herbs, 
including T. australe were cut to within 0.5 cm of the 
ground with shears. 

At both sites, in all plots, individual T. australe 
plants were tagged and numbered and the fates of the 
original and emerged plants were surveyed over 1.5 
years (Table 1). Data on plant vigour (number of stems/ 
plant; Perpendicular Point only) and the incidence of 
flowering or fruiting were also collected. 

Analyses of the proportion of T. australe plants 
surviving 6 and 16 months after treatment, were made 
using Generalised Linear Modelling (GLIM), with a 
binomial error structure (Crawley 1993). The effects 
of the factors, treatment (burnt, cut, control) and site 
(Perpendicular Point, Look at Me Now Headland) and 
their interactions were examined using the chi-squared 


59 


EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 


Table 1. The dates of treatment applications and monitoring at the study sites. 


Study Site Treatment (date) 


burn, cut (26/7/95) 
burn, cut (27/7/95) 


Perpendicular Point 
Look at Me Now Headland 
Old Bar Reserve 


statistic. 

The density of T. australe plants (0.25 m*) was 
examined using fully factorial analyses of variance 
(ANOVA) and Tukey tests for pairwise comparisons. 
The effects of treatments (burnt, cut, control) and sites 
(Perpendicular Point, Look at Me Now Headland) were 
examined at pre- and post-treatment dates (0, 6 and 
16 months). To satisfy Cochran’s test of homogeneity 
of variances, data were square root transformed and if 
necessary a more conservative level of significance 
(p<0.01) was applied (Underwood 1981). 

Analyses of the vigour of T. australe plants 
(number of stems/plant) at Perpendicular Point were 
made using one-way ANOVAs. The effects of 
treatments (burnt, cut, control) were examined at pre- 
and post-treatment dates (0, 6 and 16 months). Data 
from all plots and cohorts within each treatment were 
pooled. 


Hinterland (low density plants) 

At Old Bar Park, where T. australe occurred at 
very low density, large plots were subjected to burning 
or slashing. Each treatment (burnt, slashed, control) 
was allocated to a 10 m x 10 m plot within an overall 
area of 40 m x 50 m. There were 2 replicates of each 
treatment. Whilst for practical purposes the two burnt 
plots were placed together, replicates of the cut 
treatment and control were randomly allocated to the 
remaining plots. Burning took place in hot conditions 
during summer (December 1996). Two plots were 
slashed the next day to within 5 cm of the ground. The 
resulting cuttings were removed from the plots. 
Individual T. australe plants were tagged, numbered 
and followed for 2 years (Table 1). 

Although not measured quantitatively at 
Perpendicular Point and Look at Me Now Headland, 
observations indicated that the measurement of bare 
ground may be useful in discussing trends in the data. 
The cover of grasses/herbs and bare ground were 
measured at each census (<5 replicates) in classes (1=1- 
10%, 2=11-20%.,.....,10=91-100%) within randomly 
allocated quadrats (1 m2), located within each treatment 


60 


burn, slash (16/12/96) 


Monitoring Dates 

(pre and post treatment) 
12/5/95, 14/2/96, 4/12/96 
26/7/95, 11/2/96,18/12/96 
3/12/96, 2/12/97, 16/12/98 


plot. Rock cover was negligible. 

Analysis of the density of T. australe plants in 
each plot (number/100 m?) was made using fully 
factorial ANOVAs and Tukey tests for post hoc 
comparisons. The effects of the treatments (burnt, cut, 
control) were examined on each day of sampling. 

Analyses of the cover classes of bare ground 
were made using a two-way fully factorial ANOVA. 
The effects of treatment (burnt, cut, control) and 
sampling date (pre-treatment, 1 and 2 years post- 
treatment) were examined. 


RESULTS 


Headlands (high density plants) 

Site, but not treatment, had a significant effect 
on the proportion of plants surviving 6 and 16 months 
after the start of the study (Fig. 3). At both times 
survival was higher at Perpendicular Point than at Look 
at Me Now Headland (respectively X7=10, df=1, 
p<0.005; X°=8.5 df=1, p<0.025). By the end of the 
study between 80% and 100% of the original plants 
had suffered mortality. 

Six months after the application of treatments 
there was no significant difference in the density of T. 
australe (0.25 m) with respect to treatment and site 
(p>0.05; Fig. 4). Sixteen months after treatment, 
however, there was a significant effect of site (F=4.72, 
df = 1,84, p<0.05). Look at Me Now Headland had a 
higher density of plants than Perpendicular Point. 

At Perpendicular Point there was no significant 
difference in the vigour of T. australe (number of 
stems/plant) with respect to treatment either prior to 
treatment, or 6 months and 16 months after treatment 
(p>0.05; Fig. 5). There appeared to be a general 
increase in plant vigour over this period. 

Within 6 months of applying treatments at 
Perpendicular Point and Look at Me Now Headland, 
flowering and fruiting of original plants and new 
recruits of T. australe were recorded in summer (11 
February 1996). 


Proc. Linn. Soc. N.S.W., 125, 2004 


J.S.COHN 


Hinterland (low density 
plants) 
The density of T. 


1.0 australe plants at Old Bar 
0.8 Park was _ significantly 
0.6 Treatments affected by treatment 2 years 
0.4 applied after application. The slashed 
ee areas had a higher density 


than either the burnt or the 


Proportion of plants alive 


oO ra oa o & a a a ox a control which were not 

> Qa > : = | eer ip significantly different from 
Se Oo) AO gerade Tes ° 8 y 

Swan wae NS ey Ayo one another (F=15.5, df=2,3, 

p<0.05; Fig. 6). Pre-treatment 

Date (month and year) and | year after treatment, 


there was no significant 


Figure 3. Proportional survival of 7. australe (mean, se) at Perpendicular difference in the density of T. 
Point (#) and Look at Me Now Headland () following treatments (pooled; 9 75!” ale between treatments 


burnt, cut, control) applied in July 1995. (p>0.05). 
There was a 


significant interactive effect 

of treatment and time of 
a/_ Perpendicular Point sampling on the cover of bare 
ground at Old Bar Park 
(F=3.12, df=4, 27, p<0.05; 
Fig. 7). Whilst there was no 
significant difference 
between the plots prior to the 
imposition of treatments, 1 
and 2 years (no significant 


Treatments applied 


So FF ND W 


N 
VOnmeOu ae ta SO IOP NO KONO NS 
& Nn NHN DH DR DN 2 a ON Oh o difference) after burning, the 
N Ze i Q, > =| S jor oan 
. S 3 =) 5) bare ground was significantl 
Oo oS) ) S) a R= fe) 8 8 y 
a a ea SE eZ higher than in the slashed or 
= the control at any time (except 
a. b/ Look at Me Now Headland burn at 2 years = pre-burn and 
Seeing cut at 2 years). 
A Within 1 year of 
2 treatment application, 
1 flowering and fruiting of new 
recruits of 7. australe were 
0 recorded in summer (2 Dec 
1998). 
PRR Re Fe FR S 
eee 6 FP EBS 
a A 2 A DISCUSSION 
Date (month and year) Headlands (high density 
plants) 
Figure 4. The density (mean, se) of 7. australe plants (0.25 m7) before Burning or cutting T. 


and after treatments at Perpendicular Point and Look at Me Now = 2#trale plants in winter, did 


Headland. Treatments (burnt @, cut M, control A) were applied in July —-°t_ Significantly affect their 
1995. survival. Similarly, Leigh and 


Briggs (1989) found the 
survival of a population of T. australe near Canberra, 
was unaffected by a trial burn in autumn. Indeed, the 


Proc. Linn. Soc. N.S.W., 125, 2004 61 


EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 


—_ 


ONO O 


Treatments applied 


! 


No. stems/plant 


May-96 
Jul-96 
Sep-96 
Nov-96 


WD NH NY NH OO 
Di Ai A CN genes 
Ste EG StS ec 
S DZ) ee 


Date (month and year) 


Figure 5. The size of T. australe plants before and after treatments at 
Perpendicular Point. Size was measured as the number of stems per 
plant (mean, se). Treatments (burnt @, cut @, control A) were applied 
in July 1995. 


— 
Nn 


Treatments 
applied 


— 
(=) 


No. plants/1 00m? 
Nn 


=) 


1996 1997 1998 


Date (December in year) 


Figure 6. The density (100 m7) of 7. australe plants (mean, s.e.) in 
the hinterland at Old Bar Park before and after treatments were 
applied (burnt @, slashed @, control A). Treatments were applied 
in December 1996. 


Zz 
S| 

5 Wu Treatments 
bb —~ 8 

o a 

He 6 

Oo Oo 

ems 

5 2 

> 

S 0 


1996 1997. 1998 


Date (December in year) 


Figure 7. Cover classes (mean, se) of bare ground before and after 
treatments (burnt @, slashed ™, control A), in the hinterland at Old 
Bar Park. Treatments were applied in December 1996. Cover 
classes:1=0-10%.....10=11-100%. 


existence of buds in the 
immediate vicinity of the soil 
surface (McIntyre et al. 1995) 
allows the species to resprout 
after disturbance. In subalpine 
and tableland climates, it is the 
habit of T. australe to die back to 
the rootstock during winter and 
resprout in spring (Cooper 1986; 
Archer 1987; Gross et al. 1995; 
Cohn 1999). This is not the case 
in coastal areas, where the species 
persists all year round (Cohn 
1999). 

Whilst a study on the 
southern tablelands of NSW 
(Leigh and Briggs 1989), 
describes T. australe as an annual 
or a biennial, this study suggests 
that the species may live longer 
on the coast. After 6 and 16 
months, respectively, 
approximately 30% and 17% of 
plants were still alive. Since it is 
likely that these plants originated 
at least 9 months previously in 
spring, their ages were more than 
likely 15 months and 25 months, 
respectively. Certainly, Prober 
and Thiele (1998) believe it 
possible that T. australe lives 
longer in less severe climates. 

Although there was no 
significant effect of treatment on 
the density of T. australe, there 
was a higher density at the more 
exposed Look at Me Now 
Headland than at Perpendicular 
Point 2 years after treatments. 
This agrees with Cooper’s (1986) 
hypothesis that competition from 
Kangaroo Grass (T. australis) on 
exposed headlands is reduced by 
salt laden winds. It is also possible 
that the experimental burn, which 
was hotter than would be 
experienced naturally, even in 
extreme conditions (R. Bradstock 
pers. comm.), could have led to 
some mortality of T. australe 
seeds near the soil surface, thus 
reducing the effectiveness of this 
treatment. The small size of the 
plots may also have reduced 


62 Proc. Linn. Soc. N.S.W., 125, 2004 


J.S.COHN 


treatment effectiveness. Finally, more time may have 
been required for the T. australe populations to respond 
to a reduction in competition brought about by the 
experimental treatments. 

Thesium australe is able to grow and reproduce 
very quickly following disturbance in winter. In 
December, 6 months after burning or cutting, there 
was no significant difference in the vigour of plants in 
the treated plots and the control. At the same time 
resprouting plants and new recruits were flowering and 
beginning to fruit. Indeed, flowering and fruiting of T. 
australe at both Perpendicular Point and Look at Me 
Now Headland occurred throughout the year (Cohn 
1999). By contrast, flowering and fruiting has been 
found to be seasonal at inland locations, occurring from 
spring to autumn (Stanley and Ross 1983; Briggs and 
Leigh 1985; Gross et al. 1995; Cohn 1999). 


Hinterland (low density plants) 

At the more protected Old Bar Park, where T. 
australe was mostly absent from the plots prior to 
treatment, summer slashing rather than burning led to 
significant seedling recruitment of T. australe, 2 years 
after treatment (Fig. 6). Although it is generally 
recognised that burning provides the bare ground for 
seedling establishment that slashing does not (Lunt 
1990a), other factors seemed to be at play in this study. 

The comparable cover of bare ground in all 
treatments at the time of the high numbers of T. australe 
in the slashed plots (Fig. 7), indicates that a reduction 
in grass height may have been responsible. In Victoria, 
Lunt (1990b) believed that selective grazing of 
tussocks to a height of 5 cm by rabbits and kangaroos 
may have contributed to the maintenance of species 
richness by reducing competition from perennial 
grasses. Thus, it is probable that a reduction in the 
height of the dominant species Themeda australis 
rather than an increase in bare ground, led to significant 
recruitment of T. australe seedlings in the slashed 
treatment. 

Given that the post-fire conditions reduced 
competition from Themeda australis, it is curious that 
there was not a significant effect on T. australe 
numbers. In the same summer following burning, 
seedlings of 7. australe were observed in these plots 
(S. Long pers. comm.). Their low numbers throughout 
the study, however, may have resulted from the more 
exposed conditions, reflected by higher cover of bare 
ground, experienced during the first and second 
summers (Fig. 7). In addition, T. australe’s 
hemiparasitic dependence on other herbs and grasses, 
(Scarlett et al. 1994), may have made it difficult for T. 
australe to survive, given that its hosts were also 
recovering from the effects of the fire. Indeed summer 


Proc. Linn. Soc. N.S.W., 125, 2004 


“dying back’ of T. australe in times of water stress has 
previously been recorded by Leigh and Briggs (1989). 
Whilst older T. australe plants may have the resources 
to recover, seedlings, such as those observed in this 
experiment soon after the burning, may not have had 
that capability. 


Management implications 

The results from this study, coupled with the 
long-term persistence of T. australe on exposed 
headlands in the absence of active management 
(Cooper 1986; Griffith 1992), indicate that there is 
apparently no need for a change in this regime, except 
where shrub recruitment (native or exotic), may be 
competing with the survival of T. australe. 

By contrast, in the more protected hinterlands, 
where T. australe also occurs, active management may 
be required to reduce competition from Kangaroo 
Grass (T. australis). Although, results from this study 
indicate that early summer slashing of a grassland (5 
cm height) resulted in recruitment of T. australe plants, 
further research is required to determine if this is the 
most appropriate time of the year and method. Of 
particular concern is that disturbance of the summer 
growing Kangaroo Grass (T. australis) at this crucial 
time could result in the introduction of weed species 
(Griffith 1992). There is also a need to determine if 
burning outside summer, especially in autumn or early 
spring, would yield different results. This is important 
in the light of other work, which indicates that fire 
rather than mowing in grasslands is preferable to 
maintain species richness (Kirkpatrick 1986; Lunt 
1991; James 1994). 

If shrub encroachment becomes a threat to the 
survival of T. australe, studies have recommended 
various fire intervals of between 2 and 10 years to 
reduce dominance of native shrubs (Groves 1974; Lunt 
1998b) or a regime of frequent fire and mechanical 
disturbance to reduce exotic shrub frequency (e.g. 
Chrysanthemoides monolifera; Kirkpatrick 1986). 
Although this study did not examine an appropriate 
disturbance interval for 7. australe, its quick growth 
and reproductive development and its continued 
presence at Hat Head (S. Griffith pers. comm.), which 
has burnt every 2 to 4 years for the past 15 years (NSW 
NPWS Records), indicates it can apparently cope with 
a relatively frequent disturbance regime. Studies in 
Victoria (Scarlett and Parsons 1982, 1993) suggest that 
the absence of T. australe and other late-flowering 
species along railway lines has resulted from annual, 
late-season burning. Further research is required in 
coastal areas to determine an appropriate disturbance 
interval for the long-term conservation of T. australe. 


63 


EFFECTS OF SLASHING AND BURNING ON THESIUM AUSTRALE 


ACKNOWLEDGEMENTS 


Thanks to Steve Griffith, Shirley Cohn and Steve 
Clemesha for their assistance in the field. National Parks 
and Wildlife Service staff from Port Macquarie District office 
provided me with encouragement, information and 
equipment. Thanks to Old Bar Bushfire Brigade, who carried 
out the burn at Old Bar Park. Financial assistance was 
provided by Environment Australia and New South Wales 
National Parks and Wildlife Service. Thanks to Andrew 
Denham and Mark Tozer who kindly provided comments 
on the manuscript. 


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65 


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Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the 
Botanical Significance of Insects Host-specific to Austral 
Bracken Fern (Pteridium esculentum) 


LAURENCE Mounp! AND Masami Masumoro? 


‘CSIRO Entomology, GPO Box 1700, Canberra 2601 (laurence.mound @csiro.au); 
*MAFF, Yokohama Plant Protection Station, Shin’ yamashita, 1-16-10, Yokohama, 238-0801, Japan. 


Mound, L. and Masumoto, M. (2004). Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the 
botanical significance of insects host-specific to Austral bracken fern (Pteridium esculentum). 
Proceedings of the Linnean Society of New South Wales, 125, 67-71. 


Austral bracken fern, Pteridium esculentum, differs from its European counterpart in supporting one 
species of both thrips and aphid. The previously undescribed species of thrips, Trichromothrips veversae 
sp.n. (Thripidae), is widespread and locally abundant in southern Australia breeding on the youngest fronds 
of bracken but not on other ferns. It is unique among nearly 30 species of this Old World tropical genus in 


lacking long setae on the pronotum. 


Manuscript received 18 June 2003, accepted for publication 17 September 2003. 


KEYWORDS: aphids, bracken, Pteridium, thrips, Trichromothrips, 


INTRODUCTION 


Common bracken fern is often considered to 
be a single, cosmopolitan species Pteridium aquilinum 
(Dennstaedtiaceae). In retaining this view, the major 
reference work on botanical nomenclature (Mabberley, 
1997) recognised two subspecies, the nominate one 
from the Northern Hemisphere and Africa, and P. 
aquilinum caudatum from the Southern Hemisphere. 
In Australia, in contrast, Brownsey (1989) recognised 
three species of Pteridium: P. aquilinum introduced 
to a small area of South Australia in the Adelaide Hills; 
P. revolutum native to north-eastern Queensland but 
extending widely across New Guinea and South East 
Asia; and P. esculentum native to southern and eastern 
Australia but extending to South East Asia and the 
Pacific. More recently, Thomson (2000) has concluded 
from an extensive study of both structural and 
molecular characters that several of the Pteridium 
varieties distinguished worldwide, including 
esculentum, “might best be treated as species”. 

These differences in opinion concerning the 
botanical status of bracken fern are not without 
entomological significance. No species either of aphid 
(Homoptera) or of thrips (Thysanoptera) is known to 
live on bracken in Europe, where this plant is 
widespread and abundant and often an invasive weed. 
In contrast, the aphid species Shinjia orientalis 
(Mordwilko) (= S. pteridifoliae Shinji) has been 
reported widely on Preridium from northern India and 
Japan to eastern Australia. Moreover, populations of 


bracken in eastern North America support another 
aphid species, Mastopoda pteridis Oestlund, and in 
western North America five aphid species in the genus 
Macrosiphum have been reported from Pteridium (V. 
F. Eastop, 2003 pers. comm.). If Pteridium were truly 
monotypic, comprising one worldwide panmictic 
species, then different populations might be expected 
to support similar, if not identical insect species. The 
description here of a new species of Thripidae that is 
widespread on bracken in Australia would thus appear 
to provide further support for the recognition of distinct 
species within this ubiquitous plant genus. Presumably 
these insects are reflecting diversity within the genus 
Pteridium that botanists have been reluctant to 
acknowledge. 

The existence of this thrips species had been 
suspected for many years. In 1967, the wife of the 
eminent Australian insect ecologist H.G. Andrewartha, 
Hattie Vevers-Steele after whom the new species 
described below is named, drew the attention of one 
of us (LAM) to some specimens of a thrips species 
taken from bracken near Adelaide during her studies 
on Australian Thysanoptera (see Mound, 1996). The 
specimens were in poor condition, and efforts at that 
time to locate the species in the field were not 
successful. However, during the past 10 years this 
thrips has been found to be widespread across southern 
Australia, but breeding only in the curled apices of the 
youngest fronds of bracken. This species was listed 
by Shuter and Westoby (1992) from a population of 
bracken near Sydney as “Anaphothripinae gen. et sp. 


THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN 


indet”’, but is here recognised as a new species of the 
widespread Old World genus Trichromothrips. 
However, within that genus it exhibits one remarkably 
deviant autapomorphy — the absence of any long setae 
on the pronotum. This thrips has been found only on 
Pteridium esculentum, as defined by Brownsey (1998), 
even when this has been found growing in association 
with other ferns that are superficially similar, such as 
the closely related Hypolepis  muelleri 
(Dennstaedtiaceae), or young specimens of the more 
distantly related tree fern Dicksonia antarctica 
(Dicksoniaceae). No thrips have been found on any 
species of Hypolepis, although Scirtothrips frondis 
Hoddle and Mound breeds abundantly on the youngest 
fronds of Dicksonia and has also been taken on a 
species of Cyathea (Hoddle and Mound, 2003). 


Trichromothrips Priesner 


Trichromothrips Priesner, 1930: 9. Type species T. 
bellus Priesner. 

Bhatti (2000) has fully defined and reviewed 
this genus, synonymising the genus Dorcadothrips 
Priesner and providing a key to identify the 27 included 
species. Of these, 24 are from the Old World, between 
Africa and Queensland but mostly from South East 
Asia. The other three species, two from Hawaii and 
one widespread, may also have come originally from 
the Oriental region. The collection data for most of 
the species are probably not reliable indicators of the 
plants on which these thrips breed, but two species (T. 
billeni Strassen and T. bilongilineatus Girault) are 
associated with ferns (Mound, 2002b), and in the 
region of Japan around Tokyo and Yokahama, T. alis 
Bhatti or a closely related species is found on a species 
of Polystichum (Dryopteridaceae). Finally, three 
related genera of Thripidae are also associated with 
ferns, Laplothrips Bhatti, Octothrips Moulton and 
Pteridothrips Priesner (Mound, 2002b). 

Members of these four genera are unusual in 
bearing a pair of setae on the dorsal apical margin of 
the first antennal segment. This character state is also 
shared by species in the following genera of Thripidae, 
although none involves fern-living species: Alathrips 
Bhatti, Bregmatothrips Hood, Ceratothripoides 
Bagnall, Craspedothrips Strassen, Diarthrothrips 
Williams, Furcithrips Bhatti, Megalurothrips Bagnall, 
Mycterothrips Trybom, Odontothrips Amyot and 
Serville, Odontothripiella Bagnall, Pezothrips Karny, 
Sorghothrips Priesner, Watanabeothrips Okajima, 
Yoshinothrips Kudo. Moreover, although the two 
species comprising the Oriental genus Bathrips Bhatti 
lack this pair of setae on the dorsal apical margin of 


68 


the first antennal segment, they share many other 
character states with Trichromothrips species, and 
these two genera are possibly closely related. 


Trichromothrips veversae sp.n. 

Holotype 2 macroptera, Australian Capital 
Territory, Woods Reserve, from young fronds of 
Pteridium esculentum, 6.xii.2002 (LAM 4244), in 
ANIC, CSIRO Entomology, Canberra. 

Paratypes: 2 males, 17 females, same host, 
date and locality as holotype (Masumoto, Mound and 
Wells); 3 females at same locality but 16.1.1999 (LAM 
3664). 

Specimens excluded from the type series 
were collected widely in southern Australia, 
including Tasmania, Western Australia, New South 
Wales, and the Australian Capital Territory (see 
Distribution below). 


Female macroptera 
Colour: body yellow with orange pterothorax, 


ocelli bright red, antennae brown, abdomen with 
transverse light brown markings, wings shaded; colour 
of cleared and mounted specimens yellow, tergites 
shaded anteromedially and along antecostal line, IX 
and X shaded, mesonotum and metanotum weakly 
shaded; head and antennal segment I pale, segments 
If to VIL almost uniformly dark brown with extreme 
base of segments III to V slightly paler, I paler than 
segment III; all legs greyish brown; fore wing and scale 
greyish brown, but base of fore wing paler. 
Structure: Head slightly wider than long, not 
prolonged in front of eyes, with a few transverse striae 
posteriorly on vertex (Fig. 1); ocellar setae I absent, 
setae III no longer than length of an ocellus and arising 
between anterior margins of posterior ocelli; three pairs 
of postocular setae, pairs I and II close together behind 
ocelli; ventral surface of head with 5 pairs of setae 
between compound eyes anterior to anterior tentorial 
pits; mouth-cone rounded, maxillary palpi 3- 
segmented; compound eyes without pigmented facets. 
Antenna 8-segmented (Fig. 3); forked sense-cones on 
Ill and IV exceptionally stout; segment I with 2 dorsal 
apical setae; II with weak microtrichia laterally only, 
Ill to VI with a few large microtrichia on dorsal and 
ventral surfaces; III with 2 dorsal and 2 ventral setae. 
Pronotum medially with few or no lines of 
sculpture and 4 to 10 discal setae; posterior margin 
with five pairs of setae, none of which is longer than 
the discal setae. Mesonotum with weak transverse lines 
of sculpture, without campaniform sensilla near 
anterior margin, median pair of setae far ahead of 
posterior margin. Metanotum (Fig. 2) medially without 
sculpture and one pair of small setae far from anterior 


Proc. Linn. Soc. N.S.W., 125, 2004 


L. MOUND AND M. MASUMOTO 


Figure 1. Trichromothrips veversae, head and 
pronotum. 


margin, without campaniform sensilla. Prosternal ferna 
not divided; mesothoracic sternopleural suture not 
developed; meso- and metasternum each with well- 
developed spinula. All tarsi 2-segmented. Forewing 
veinal setae short, less than half width of wing in 
length; first vein with about 8 setae near base and 2 
(rarely 3) setae near apex; second vein with about 10 
setae; posterior fringe cilia wavy; forewing scale with 
4 marginal setae. 

Abdominal tergites without posteromarginal 
craspeda or lateral ctenidia; tergites II to VIII without 
sculpture medially, lateral to seta S2 with about 7 
anastomosing transverse lines bearing tuberculate 
microtrichia; tergite VIII without posteromarginal 
comb; tergite IX with paired campaniform sensilla 
posteromedially; tergite X undivided; pleurotergites 


without discal seta, sculpture similar to lateral areas 
of tergites. Sternites without discal setae; sternite II 
with two pairs of posteromarginal setae, sternites III 
to VII with three pairs, on VII all three pairs arise in 
front of sternal posterior margin. 

Measurements (holotype female in um with 
small paratype female in parentheses): Body length 
1400 (1100). Head, length 90 (85); width 125 (105). 
Pronotum, length 105 (95); width 160 (130); 
posteromarginal setae 15 (12). Forewing, length 750 
(650). Antennal segments 25, 32, 50, 57, 40, 43, 10, 
17 (25, 30, 40, 47, 35, 37, 7, 15). 


Figure 2. Trichromothrips veversae, mesonotum 
and metanotum. 


Male aptera 
Colour paler than female. Structure similar 


to female except: forked sense-cones on antennal 
segments III and IV small and slender; one of three 
available males lacks ocellar setae II; mesonotum 
transverse with 4 or 5 setae near lateral margins; 
pleurotergal sutures weakly developed; tergite IX 


Figure 3. Trichromothrips veversae, antenna. 


Proc. Linn. Soc. N.S.W., 125, 2004 


69 


THRIPS AND APHIDS ON AUSTRAL BRACKEN FERN 


posterior margin with horn-like paired drepanae 
extending beyond segment X; sternites III to VIII each 
with about 50 small, irregularly arranged, glandular 
areas, marginal setae arising at margin on all sternites. 

Measurements (paratype male in um). Body 
length 1000. Head, length 83; width 100. Pronotum, 
length 85; width 130; posteromarginal setae 15. Tergite 
IX drepanae length 60. Antennal segments 25, 30, 37, 
AO; 3253775 15. 


Larva Il. 

Colour pale yellow with red eyes, 
progressively developing extensive pale red 
hypodermal pigment in meso- and metathorax and 
anterior abdominal segments, body usually turning 
deep yellow progressively; major dorsal setae parallel- 
sided with bluntly square apices, 3 pairs on head, 6 
pairs on pronotum, 3 pairs on abdominal tergites II — 
VIII, 2 pairs on IX, antennal II with 2 pairs of similar 
but smaller setae; setae on tergite X and abdominal 
sternites with apices acute; sternite IX posterior margin 
with row of about 30 small tooth-like tubercles. 


Systematic relationships 
Currently, this new species cannot be placed 


in any of the 10 species-groups distinguished by Bhatti 
(2000) within Trichromothrips, although it shares with 
the other 27 species the many character states listed 
by that author in his diagnosis of the genus. In contrast 
to those species, it lacks any long pronotal setae, the 
metasternal spinula is well developed not weak, and 
females have unusually stout antennal sense cones. 

In Australia, only one other species of 
Trichromothrips has been collected in good numbers: 
T. bilongilineatus (Girault) from ferns near Gosford 
(Mound, 2002a). Of the other two members of the 
genus listed from Australia, the record of T. xanthius 
(Williams) is based on one female taken in quarantine 
in North America but labelled as coming from 
Australia (Mound, 1996), and T. obscuriceps (Girault) 
is known from a single sample apparently taken on 
Crinum lilies near Brisbane. The genus is probably 
well established in northern Australia, but only a few 
specimens are available, representing two further 
unidentified species, swept from grasses near Darwin. 
All of these species have long pronotal posteroangular 
setae. 

The lack of long pronotal setae gives T. 
veversae the superficial appearance of an Anaphothrips 
species. This is another example of the ineffective 
supra-generic classification within the subfamily 
Thripinae, in which traditional subtribal names such 
as Aptinothripina do not refer to definable groups 
(Mound, 2002c), despite their continued use by various 


70 


authors (eg. Vasiliu-Oromulu et al. 2001). There are 
several unrelated Thripinae genera in which species 
usually have two pairs of long pronotal setae, but in 
which one or more species have these setae no longer 
than the discal setae and are thus “Anaphothripine” in 
appearance, eg. Dichromothrips  Priesner, 
Pseudanaphothrips Karny and Thrips Linnaeus. 

The presence or absence of long setae on the 
pronotum was recognised as a poor indicator of 
phylogenetic relationships by Mound and Palmer 
(1981), who proposed a series of informal genus- 
groups within the Thripinae. These authors included 
Scolothrips Hinds, a genus of predatory thrips, in their 
Dorcadothrips genus-group (Mound and Palmer, 
1981). Scolothrips species resemble some 
Trichromothrips species in general appearance, for 
example the pale slender body and bulging compound 
eyes, but they have very long ocellar setae and the 
pronotum bears six pairs of elongate setae. Moreover, 
the dorsal apical margin of the first antennal segment 
does not bear a pair of setae, and the mesosternal 
sternopleural sutures are weakly developed. The 
character state on the first antennal segment discussed 
above suggests that the genus-groups recognised by 
Mound and Palmer (1981) require reappraisal. 


Distribution and host records 

T. veversae has been found to be locally 
abundant in many parts of southern Australia, 
including Western Australia near Albany, Tasmania 
near Hobart, and various sites in South Australia 
(Adelaide Hills; Cox’s Scrub south of Adelaide; and 
Kangaroo Island). It is abundant in the mountains of 
the ACT, and is widely distributed in the eastern forests 
of New South Wales from near Eden to the Blue 
Mountains. It possibly occurs even further north, but 
a sample taken from Pteridium at Beerwah, north of 
Brisbane, yielded only Scirtothrips dobroskyi Moulton 
(Hoddle and Mound, 2003). In a survey of the insects 
associated with bracken in New Guinea, Kirk (1977) 
does not mention thrips, but since thrips on ferns are 
associated only with very young fronds, or even with 
croziers that are not yet fully expanded, these minute 
insects are often difficult to detect. Similarly, the list 
given by Balick et al. (1978) of insects taken from 
ferns worldwide is based on a survey of published 
records, derived mainly from general collecting, and 
some of the thrips species listed are fungus-feeders, 
not fern-feeders. Mound (2002b) emphasised that 
several published records of thrips on ferns are based 
on single samples or even single specimens, and thus 
cannot be relied on to indicate a host relationship. 

In Japan, the common species of bracken fern 
is considered also to represent Preridiium esculentum 


Proc. Linn. Soc. N.S.W., 125, 2004 


L. MOUND AND M. MASUMOTO 


and, as indicated above, the aphid species Shinjia 
orientalis has been recorded from this plant in Japan 
as well as Australia. However, searches for thrips on 
substantial populations of bracken in Japan, 
particularly near Narita City, have failed to discover 
Trichromothrips veversae. 

At Crafers in the Adelaide Hills, South 
Australia, a substantial population of adults and larvae 
of Thrips imaginis Bagnall was found on bracken 
fronds in an open field during December 2002, together 
with a few larvae of Trichromothrips veversae. 
However, this seems to be a rare host association for 
the highly polyphagous Australian Plague Thrips. 

At several sites, near Adelaide and on 


Kangaroo Island, larvae of T. veversae were found | 


bearing up to 12 larval Eucharitidae (Hymenoptera). 
This is presumably a phoretic association, but no 
observations were made on associated ants, the 
probable host of these small wasps. 


ACKNOWLEDGEMENTS 


The authors are grateful to the Chief, CSIRO 
Entomology, for providing research facilities at Canberra, 
to Victor Eastop of the Natural History Museum, London, 
for information on aphids associated with ferns, and to Alice 
Wells for field assistance and comments on the manuscript. 
John Thomson of the Royal Botanic Gardens Sydney kindly 
drew our attention to several references. Mr. Kenji Morita, 
Head of Yokohama Plant Protection Station, and Mr. Tetsuo 
Imamura, Chief of Identification Section, Yokohama Plant 
Protection Station, kindly facilitated a study visit to Canberra 
by M. Masumoto, and Mr. Shigeo Aochi in Narita City, and 
Mr. Yuji Yoshida in Chiba City provided help in searching 
for thrips on Preridium in Japan. 


REFERENCES 


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89. 

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Brownsey, P.J. (1989). The taxonomy of Bracken 
(Pteridium: Denstaedtiaceae) in Australia. 
Australian Systematic Botany 2, 113-128. 

Brownsey, P.J. (1998). Denstaedtiaceae, pp. 214-228 in 
Flora of Australia volume 48. Ferns, 
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Melbourne. 

Hoddle, M.S. and Mound, L.A. 2003. The genus 
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Thripidae). Zootaxa 268, 1-40. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Kirk, A.A. (1977). The insect fauna of the weed Prteridium 
aquilinum (L.) Kuhn (Polypodiaceae) in Papua 
New Guinea: A potential source of biological 
control agents. Journal of the Australian 
Entomological Society 16, 403-409 

Mabberley, D.J. (1997). The Plant Book. Second edition. 
Cambridge University Press, Cambridge. 

Mound, L.A. (1996). Thysanoptera, pp 249-336, 397-414 
(Index), in Wells, A., Zoological Catalogue of 
Australia. Volume 26. Psocoptera, Phthiraptera, 
Thysanoptera. Melbourne. CSIRO Australia. 

Mound, L.A. (2002a). Thrips and their host plants: new 
Australian records (Thysanoptera: Terebrantia). 
Australian Entomologist 29, 49-60. 

Mound, L.A. (2002b). Octothrips lygodii sp.n. 
(Thysanoptera, Thripidae) damaging weedy 
Lygodium ferns in south-eastern Asia, with 
notes on other Thripidae reported from ferns. 
Australian Journal of Entomology 41, 216-220. 

Mound, L.A. (2002c). The Thrips and Frankliniella 
genus-groups: the phylogenetic significance of 
ctenidia, pp. 379-386 in Mound L.A. and 
Marullo, R. [eds] Thrips and Tospoviruses. 
Proceedings of the 7 International Symposium 
on Thysanoptera. Australian National Insect 
Collection, Canberra. 

Mound, L.A. and Palmer, J.M. (1981). Phylogenetic 
relationships between some genera of Thripidae 
(Thysanoptera). Entomologica Scandinavica 15, 
153-17. 

Priesner, H. (1930). Contribution towards a knowledge of 
the Thysanoptera of Egypt, II. Bulletin de la 
Société Royal Entomologique d’Egypte 14, 6- 
13). 

Thomson, J.A. (2000). Morphological and genomic 
diversity in the genus Pteridium 
(Dennstaedtiaceae). Annals of Botany 85, 77-99. 

Vasiliu-Oromulu, L. zur Strassen, R. and Larsson, H. 
(2001). The systematic revision of Thysanoptera 
species from the Swedish fauna and their 
geographical distribution. Entomologia romana 
6, 93-101. 


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Cyst Shell Morphology of the Fairy Shrimps (Crustacea: 
Anostraca) of Australia 


BrIAN V. Tiwms!, WILLIAM D. SHEPARD? AND RICHARD. E. Hiti3 


' School of Environmental and Life Sciences, University of Newcastle, Callaghan NSW 2308. 
* Department of Biology, California State University, Sacramento California 95819, USA. 
33900 Central Avenue, Fair Oaks California 95628, USA. 


Timms, B.V., Shepard, W.D. and Hill, R.E. (2004). Cyst shell morphology of the fairy shrimps (Crustacea: 
Anostraca) of Australia. Proceedings of the Linnean Society of New South Wales 125, 73-95. 


Cyst shell morphology is described for 31 species and 4 genera of endemic Australian fairy shrimp. All 


four genera have distinctive cyst shell structure. 


Manuscript received 3 May 2003, accepted for publication 23 July 2003. 


KEYWORDS: Branchinella, branchiopodid, Parartemia, Streptocephalus. 


INTRODUCTION 


Australia has a relatively simple fauna of 
anostracans, with about 50 species in five genera. The 
list is currently growing because of recent discoveries, 
but presently stands at 29 described and one 
undescribed species of Branchinella 
(Thamnocephalidae), 8 described and 7 undescribed 
species of Parartemia (Parartemidae), two species of 
Artemia (Artemidae), and two described and perhaps 
many undescribed species of Streptocephalus 
(Streptocephalidae) and a species of an undescribed 
branchipodid genus (Timms, in press). All species are 
endemic, except for the two species of Artemia, one 
of which is certainly introduced and the other possibly 
introduced (see McMasters et al., in press; Timms, in 
press). 

Little is known on the biology of Australian 
species (Timms, in press) and even less on detailed 
morphology and of adults and of cysts. Some general 
observations on cyst morphology of some species have 
been made by Herbert and Timms (2000), Timms 
(2002) and Timms and Geddes (2003). 

Cyst shell morphology is much better known 
in fairy shrimp that occur in Europe, Africa and North 
America, and even the species occurring on some 
oceanic islands have been described. Political units 
from which cysts of fairy shrimp have been described 
include: France (Thiéry and Gasc 1991); Italy (Mura 
1986, 1991b); California (Hill and Shepard 1997, 
Shepard and Hill 2001); Africa (Brendonck and 
Coomans 1994a, b; Brendonck and Riddoch 1997); 
and the Galapagos Islands of Ecuador (Brendonck et 
al. 1990). Some cyst studies have centered on certain 


families, genera, subgenera or species groups: 
streptocephalids and thamnocephalids (Mura 1992, 
DeWalsche et al. 1991); streptocephalids (Hamer et 
al. 1994a, b); Branchinecta (Mura 1991a), 
Chirocephalus (Mura 2001, Mura et al. 2002), 
Linderiella (Thiéry and Chanpeau 1988); 
Streptocephalus (Brendonck et al. 1992; Brendonck 
and Coomans 1994a, b; Streptocephalus 
(Parastreptocephalus) (Brendonck et al. 1992); the 
Streptocephalus sealii group (Shepard 1999). Cyst 
descriptions are also now becoming a common part of 
descriptions of new species. Identification keys based 
solely on cyst morphology have been produced for 
some branchiopod groups (Brendonck and Coomans 
1994a, b; Shepard and Hill 2001; Thiéry and Gasc 
1991). Cyst shell morphology is useful for identifying 
all genera so far examined, and many, but not all, 
species. Besides a perceived lack of differences 
between some related species, intraspecific variation 
associated with various predation levels (Mura et al. 
2002, Dumont et al. 2002) and perhaps incipient 
speciation (Mura et al. 2002) may lead to confusion in 
some species. 

Cyst morphology includes both external and 
internal structure (Hill and Shepard 1997). The 
outermost part of the shell is represented by the cortex 
surface. This surface often has spines, spinules, ridges 
of several different forms, buttons and ropey folds. 
The cortex is generally relatively thin. Below the cortex 
is the thick alveolar layer, which may be divided into 
various distinct sublayers. The alveolar layer is usually 
highly vacuolated with the vacuoles separated by 
curved or straight, solid struts of varying length. The 
innermost layer is the tertiary base which is relatively 
thin, solid and sometimes lamellate throughout. Often 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


the interior surface has an impressed polygonal pattern. 
An additional cyst shell character is inpocketing of 
the entire shell. Inpocketing is only typical of cyst shells 
which have a fairly uniform alveolar layer. 

The goal of this paper is to describe the 
external and internal structure of the cyst shell in as 
many as possible of the endemic Australian fairy 
shrimp fauna. This is a prerequisite for understanding 
differences between genera and species, as a base for 
understanding any intraspecific differences and for the 
construction of identification keys. Comparative 
information on Artemia cysts is provided by Shepard 
and Hill (2001) and is not repeated here. This is of 
overseas material, but is unlikely to be different from 
Australian cysts, as all Artemia cysts are the same (G. 
Mura, pers. com.; W. Shepard, pers. com.), except for 
A. monica, which does not occur here. 


METHODS 


Specimens of fairy shrimps were collected 
in the field and preserved in 70-80% ethyl alcohol and 
identified by BVT using Geddes 1981 and Timms in 
press. Females with the most mature looking cysts 
(those with a regular pattern all over the surface and 
which retain their sphericity) present in the brood 
pouch(es) were selected for dissection of the cysts. 
Cysts were removed and air-dried on filter paper. Then 
the cysts were mounted on SEM stubs using double- 
stick tape and gold-coated. A Zeiss DSM 940 SEM 
was used for measuring and photographing the cysts. 
Cyst morphology is described as in Hill and Shepard 
(1997), Shepard (1999) and Shepard and Hill (2001). 
Cyst stubbs have been stored in the private collection 
of REH. 


RESULTS 


Cysts were obtained for 31 of the 50 or so 
species of fairy shrimp known to exist in Australia. 
Those species include: Branchinella affinis Linder 
1941, B. arborea Geddes 1981, B. australiensis 
(Richters 1876), B. basipina Geddes 1981, B. 
buchananensis Geddes 1981, B. budjiti Timms 2001, 
B. campbelli Timms 2001, B. complexidigitata Timms 
2002, B. dubia (Schwartz 1917), B. frondosa Henry 
1924, B. hattahensis Geddes 1981, B. kadjikadji 
Timms 2002, B. lamellata Timms and Geddes 2003, 
B. longirostris Wolf 1911, B. lyrifera Linder 1941, B. 
nana Timms 2002, B. nichollsi Linder 1941, B. 
occidentalis Dakin 1914, B. pinnata Geddes 1981, B. 
proboscida Henry 1924, B. simplex Linder 1941, B. 


74 


wellardi Milner 1929, Parartemia contracta Linder 
1941, P. cylindrifera Linder 1941, P. informis Linder 
1941, P. minuta Geddes 1973, P. zietziana Sayce 1903, 
Streptocephalus queenslandicus Herbert and Timms 
2000, Streptocephalus n. sp. a, Streptocephalus n. sp. 
b and an undescribed branchiopodid anostracan (all 
three from the Paroo, nw NSW-sw Qld - Timms and 
Sanders 2002 and Timms unpublished data. Two 
different cyst types were found in Branchinella 
occidentalis. Full descriptions could not be made for 
Branchinella affinis, B. nana and Parartemia informis 
due to the presence of only immature cysts. Locality 
and collection details for each species are listed in 
Appendix 1. The described Australian fairy shrimp 
species for which we did not have cysts include the 
following: Branchinella apophystata Linder 1941, B. 
compacta Linder 1941, B. denticulata Linder 1941, 
B. halsei Timms 2002, B. insularis Timms and Geddes 
2003, B. latzi Geddes 1981, B. tyleri Timms and 
Geddes 2003, Parartemia extracta Linder 1941, P. 
longicaudata Linder 1941, P. serventyi Linder 1941 
and Streptocephalus archeri Sars 1896. 

The cyst characteristics of Australian 
anostracans are as follows. 


Family Parartemidae 
Parartemia contracta Linder 

Cyst shape spherical (Fig. 1). Cyst size 250- 
275 um (mean = 263.3; n= 10). External surface with 
some inpocketing, adjacent inpockets separated by 
indistinct ridges; individual cysts with 2-6 inpockets; 
surface smooth. Alveolar layer (Fig. 2) uniform; short 
struts; vesicles small rounded and subequal. Tertiary 
layer thin. 


Parartemia cylindrifera Linder 

Cyst shape spherical (Fig. 3). Cyst size 204- 
225 um (mean = 214.6; n = 10). External surface with 
some inpocketing; adjacent inpockets separated by 
ridges; individual cysts with several inpockets each; 
surface smooth. Alveolar layer uniform, mostly solid 
(Fig. 4). Tertiary layer thin. 


Parartemia informis Linder 
Immature cyst shape spherical. Cyst size 162- 
176 um (mean = 167.5; n = 10). 


Parartemia minuta Geddes 

Cyst shape spherical (Fig. 5). Cyst size 180- 
232 um (mean = 208.0; n = 30). External surface with 
inpocketing; adjacent inpockets separated by ridges; 
individual cysts with several inpockets each; surface 
smooth. Alveolar layer uniform and solid (Fig. 6). 
Tertiary layer thin. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Parartemia zietziana Sayce 

Cyst shape spherical to hemispherical (Figs. 
7-8). Cyst size 180-208 um (mean = 192.7; n = 11). 
External surface with inpocketing; inpocketing usually 
at one end; adjacent inpockets separated by ridges; 
ridges usually with sloping walls and rounded tops; 
surface smooth. Alveolar layer uniform and solid (Fig. 
9). Tertiary layer thin. 


Family Branchiopodidae 
undescribed branchiopod genus and species 

Cyst shape very irregularly spherical (Fig. 
10), with projecting flanges that interlock with those 
of other cysts binding the cyst mass together (Fig. 11). 
Cyst diameters 208-250 um (mean = 229.7; n = 20). 
External surface with ridges defining irregular 
polygons; intersecting ridges often extending as 
flanges; entire surface covered with shallow, rounded 
to polygonal depressions; polygon size variable and 
larger on flanges and ridges (Fig. 12). Cortex relatively 
thick (» 2 um); one layer of short columns, columns 
parallel to each other and perpendicular to the surface. 
Alveolar layer (Fig. 13) thin; with small and uniform 
vesicles. Tertiary base thin. 


Family Streptocephalidae 
Streptocephalus queenslandicus Herbert and Timms 
Cyst shape tetrahedral (Fig. 14-15); cyst size 
169-208 um (mean = 195; n = 20). External surface 
with ridges dividing surface into 4 triangular faces; 
ridges each with 2 longitudinal grooves in middle half 
producing 3 short ridge crests; intersecting ridges flare 
outward; triangular faces gently convex with 1 or 2 
small bumps midface; surface granular. Alveolar layer 
(Fig. 16) with 2 sublayers; inner sublayer with short 
straight struts and medium-sized vesicles and hollows; 
outer sublayer uniform and solid; vesicular layer 
extends outward into ridge intersections. Tertiary layer 
thin. 


Streptocephalus n. sp. a (locality: Pine Tree Pool, 
Budgerie Paddock, Bloodwood Station, 130 km NW 
of Bourke, NSW) 

Cyst shape tetrahedral (Fig. 17); cyst size 254- 
300 um (mean = 283.0; n = 20). External surface with 
ridges dividing surface into 4 triangular faces; ridges 
with nearly vertical sides, top rounded (Fig. 18), middle 
half of ridge along each face with a series of large 
punctae on each side, intersecting ridges flare outward; 
triangular faces flat to convex; surface bumpy. Cortex 
thick (2-3 um). Alveolar layer (Fig. 19) with 2 
sublayers; inner sublayer with thin short struts defining 
variably-sized, interconnecting vesicles; outer sublayer 
uniform and solid. Tertiary layer thin. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Streptocephalus n. sp. b (locality: Box Hole, on road 
3 km south of homestead, Currawinya National Park, 
SW QLD) 

Cyst shape tetrahedral (Fig. 20); cyst size 240- 
268 um (mean = 249.3; n = 20). External surface with 
ridges dividing surface into 4 triangular faces; ridges 
each with 2 longitudinal grooves in middle producing 
3 short ridge crests (Fig. 21); intersecting ridges 
producing rounded corners; triangular faces depressed, 
flattened, with small bumps midface; surface mildly 
bumpy. Alveolar layer (Fig. 22) vesicular; inner 
vesicles smaller; vesicles under ridges larger. Tertiary 
layer thin. 


Family Thamnocephalidae 
Branchinella affinis Linder 

Cyst shape spherical (Fig. 23). Cyst diameter 
95-134 um (mean = 113.2; n = 20). External surface 
with steep-walled, arcuate ridges defining pinched 
polygons. 


Branchinella arborea Geddes 

Cyst shape spherical (Fig. 24). Cyst size 183- 
201 um (mean = 191.6; n = 10). External surface with 
ridges defining flat-bottomed polygons; ridges straight- 
sided, surface covered with small punctae (Fig. 25). 
Alveolar layer (Fig. 26) with 2 sublayers; inner 
sublayer with small subequal vesicles; outer sublayer 
with long struts and large hollows. Tertiary base thin. 


Branchinella australiensis (Richters) 

Cyst shape spherical (Fig. 27). Cyst size 197- 
222 um (mean = 213.5; n= 10). External surface with 
ridges defining polygons; ridges straight-walled, top 
pinched into a bead running along ridge, bead with 
pores (Fig. 28); polygons with flat bottoms. Alveolar 
layer (Fig. 29) with variably-sized vesicles; inner 
vesicles usually smaller than outer ones. Tertiary base 
thin. 


Branchinella basipina Geddes 

Cyst shape spherical (Fig. 30). Cyst size 225- 
275 wm (mean = 253.5; n = 10). External surface with 
ridges defining pinched polygons; ridges with sloping 
sides, tops evenly rounded; polygons with concave 
bottoms; surface rugose. Alveolar layer (Fig. 31) with 
small, subequal vesicles. Tertiary base thin. 


Branchinella buchananensis Geddes 

Cyst shape spherical (Fig. 32). Cyst size 204- 
232 um (mean = 217.9; n= 10). External surface with 
ridges defining pinched polygons; ridges with sloping 
sides; polygons with concave bottoms; surface rugose. 
Alveolar layer (Fig. 33) with 2 sublayers; inner 


15 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


sublayer with small subequal vesicles; outer sublayer 
with long branching struts and large unequal vesicles. 
Tertiary base thin. 


Branchinella budjiti Timms 

Cyst shape spherical (Fig. 34). Cyst size 141- 
155 um (mean = 144.7; n = 23). External surface with 
sinuous ridges defining tightly pinched polygons; 
ridges steep-sided with rounded tops; polygons with 
sinuous valley-like bottoms; surface slightly bumpy. 
Alveolar layer (Fig. 35) with 2 sublayers; inner layer 
with small vesicles; outer sublayer with long parallel 
struts and large hollows. Tertiary layer thin. 


Branchinella campbelli Timms 

Cyst shape spherical (Fig. 36). Cyst size 162- 
194 um (mean = 172.3; n = 20). External surface with 
broad ridges defining pinched polygons; ridges with 
sides sloped to vertical, ridge tops very broad; surface 
scaley. Alveolar layer (Fig. 37) thick, with small to 
moderate sized vesicles. Tertiary layer thin. 


Branchinella complexidigitata Timms 

Cyst shape spherical (Fig. 38). Cyst size 211- 
307 um (mean = 251.0; n = 40). External surface with 
ridges defining irregular polygons; ridges narrow, 
steep-sided, midline of ridges with sharp-pointed 
projections (Figs. 39-40); polygons with bottom flat 
to concave; surface smooth to scaley and porous. 
Alveolar layer (Fig. 41) thin; small to medium sized 
circular vesicles. Tertiary base thin. 


Branchinella dubia (Schwartz) 

Cyst shape spherical (Fig. 42). Cyst size 187- 
215 um (mean = 197.1; n= 32). External surface with 
ridges defining polygons; ridges straight, steep-walled 
and narrow; polygons with bottoms flat to slightly 
concave; surface covered with short wrinkles, pores 
abundant (Fig. 43). Alveolar layer (Fig. 44) with 2 
sublayers; inner sublayer with small rounded subequal 
vesicles; outer sublayer with long sometimes branching 
struts and large hollows. Tertiary base thin. 


Branchinella frondosa Henry 

Cyst shape spherical (Fig. 45). Cyst size 185- 
211 um (mean = 196.1; n= 10). External surface with 
ridges defining polygons; ridges with sloping sides, 
tops rounded; polygons with bottom flat to slightly 
concave; surface smooth. Alveolar layer (Fig. 46) with 
3 sublayers; inner sublayer with small vesicles; middle 
sublayer with long sometimes branching struts and 
large hollows; outer sublayer with medium-sized 
vesicles. Tertiary base thin. 


76 


Branchinella hattahensis Geddes 

Cyst shape spherical (Fig. 47). Cyst size 254- 
289 um (mean = 268.9; n = 20). External surface with 
ridges defining polygons; ridges with steep sloping 
sides, ridge tops with a sinuous bead and sharp-pointed 
and sometimes recurved projections (projections 
sometimes with film between) (Figs. 48-49); polygons 
with bottoms flat to slightly concave; surface strongly 
rugose or bumpy. Alveolar layer (Fig. 50) with 2 
sublayers; inner sublayer with small- to medium-sized 
rounded vesicles; outer sublayer with narrow curving 
struts and large hollows. Tertiary base thin. 


Branchinella kadjikadji Timms 

Cyst shape spherical (Fig. 51). Cyst size 254 
um (n = 2). External surface with ridges defining 
polygons; ridges steep-sided and sinuous, tops with 
rounded bead; polygons irregularly shaped, bottoms 
flat; surface smooth (Fig. 52). Alveolar layer (Figs. 
53-54) with at least 2 sublayers; outer sublayer with 
medium-sized rounded vesicles; next sublayer inside 
with short struts and large hollows. Tertiary base not 
visible in available cross-sections. 


Branchinella lamellata Timms & Geddes 

Cyst shape spherical (Fig. 55). Cyst size 124- 
180 um (mean = 147.6; n = 31). External surface with 
ridges defining polygons; ridges with sloping sides and 
rounded tops; polygons irregular and highly pinched, 
bottoms concave; surface bumpy. Alveolar layer (Fig. 
56) with narrow struts and very large hollows. Tertiary 
base thin. 


Branchinella longirostris Wolf 

Cyst shape spherical (Fig. 57). Cyst size 264- 
300 -um (mean = 276.9; n= 29). External surface with 
ridges defining polygons; ridges very narrow, not 
always connecting to other ridges, ridge junctions 
extended into spines with tips that are bifid, trifid or 
recurved (Fig. 58); polygons with concave bottoms; 
surface bumpy to scaley. Alveolar layer (Fig. 59) 
mostly hollow with very narrow struts. Tertiary base 
thin. 


Branchinella lyrifera Linder 

Cyst shape spherical; inpocketing common 
(Figs. 60). Cyst size 158-183 um (x = 171.5; n= 20). 
External surface with numerous short flat-topped 
columns, columns often connected by ridges; ridges 
of variable widths (Figs. 61-62). Alveolar layer solid 
(Fig. 63). Tertiary base thin. 


Branchinella nana Timms 


Immature cyst shape spherical . Cyst size 144- 
158 um (mean = 152.3; n= 19). 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Branchinella nichollsi Linder 

Cyst shape spherical (Fig. 64). Cyst size 187- 
247 um (mean = 202.3; n = 30). External surface with 
ridges defining polygons; ridges narrow, steep-sided, 
with zig-zag patterns, tops with narrow bead (Fig. 65); 
polygons tightly pinched, bottoms flat to concave; 
surface slightly bumpy. Alveolar layer (Fig. 66) mostly 
hollow, with very thin struts. Tertiary base thin. 


Branchinella occidentalis Dakin (two cyst morphs) 
Type I cyst: Cyst shape spherical; inpocketing 

common (Fig. 67). Cyst size 257-328 um (mean = 
283.5; n = 20). External surface with narrow sinuous 
ridges defining small, irregular, pinched polygons; 
ridges steep-sided and short. Alveolar layer (Fig. 68) 
not well developed. Tertiary base not seen. 

Type II cyst: Cyst shape spherical (Fig. 69). 
Cyst size 550-571 um (mean = 565.3; n= 20). External 
surface with ridges defining rounded depressions; 
ridges narrow except where intersecting, sides vertical, 
tops flat; rounded depressions with bottoms slightly 
concave and extremely porous (Fig. 70); surface 
smooth. Alveolar layer (Fig. 71) with 3 sublayers; inner 
sublayer with small equal rounded vesicles; middle 
sublayer with few struts, mostly open hollows, pores 
connecting to outside; outer layer with stringy 
vermiform struts, connecting to inner side of external 
surface. Tertiary layer thin. 


Branchinella pinnata Geddes 

Cyst shape spherical (Fig. 72). Cyst size 173- 
190 um (mean = 181.1; n = 10). External surface with 
ridges defining pinched polygons; ridges with sides 
nearly vertical, tops rounded; polygons irregular in 
shape, bottoms flat to concave; surface very bumpy. 
Alveolar layer (Fig. 73) with 3 sublayers; inner 
sublayer with small rounded subequal vesicles; middle 
layer with long branching struts and large hollows; 
outer sublayer with medium-sized irregularly shaped 
vessicles. Tertiary layer thin. 


Branchinella probiscida Henry 

Cyst shape spherical (Fig. 74). Cyst size 158- 
187 um (mean = 174.9; n = 19). External surface with 
ridges defining polygons; ridges vermiform with sides 
nearly vertical, tops smooth and rounded; polygons 
irregular in shape, some pinched, bottoms slightly 
concave and with pores; surface generally smooth (Fig. 
75). Alveolar layer (Fig. 76) without distinct sublayers; 
struts usually short; vesicles irregular in shape and 
variable in size. Tertiary layer thin. 


Branchinella simplex Linder 
Cyst shape spherical (Fig. 77). Cyst size 144- 


Proc. Linn. Soc. N.S.W., 125, 2004 


201 um (mean = 176.4; n = 32). External surface with 
ridges defining polygons; ridges with sides slightly 
sloping to vertical; polygons pinched, bottoms 
concave; surface smooth with irregularly spaced 
punctae. Alveolar layer (Fig. 78) uniform; short 
branching struts and medium-sized hollows. Tertiary 
layer thin. 


Branchinella wellardi Milner 

Cyst shape spherical (Fig. 79). Cyst size 158- 
176 um (mean = 168.4; n = 30). External surface with 
ridges defining polygons; ridges narrow and steep- 
sided, tops sharp except broader at intersections (Fig. 
80); polygons irregular in shape, some pinched, 
bottoms concave to flat; surface punctate to porous. 
Alveolar layer (Fig. 81) uniform, thick; short struts 
with small interconnected vesicles. Tertiary layer thin. 


DISCUSSION 


Australia is currently experiencing a surge of 
interest in its fairy shrimps, as indicated by the large 
percentage of newly described species and species 
awaiting description (Timms, in press). Species 
descriptions omit information on cyst shells, but 
suitable descriptions for many species are provided 
here. Although about 40% of known species are yet to 
be studied (and for most of these, data are unlikely to 
be available in the near future), enough descriptions 
of cyst shell morphology are available for comparison 
between Australian and overseas fauna and among the 
Australian species. 

Cysts of the species examined could be easily 
separated at the genus level, but only with difficulty 
or not at all at the species level. The most distinctive 
are those of Streptocephalus with their overall 
tetrahedral shape and distinct facet-edge ridges and 
convex to concave faces with bumps. This feature 
places them within the sudanicus species group of 
Maeda-Martinez et al. (1995) and subgenus 
Parastreptocephalus of Brendonck et al. (1992); 
Streptocephalus species with mature spherical cysts 
have yet to be found in Australia. 

Cysts of the new branchiopodid genus are 
also distinctive because of flanges from ridges defining 
irregular polygons. Almost all of the species of 
Branchinella have cysts with a polygonal surface 
largely reminiscent of those of overseas members of 
the genus (Belk and Sisson 1992, Mura 1992, 
Brendonck and Riddoch 1997; Sanoamuang et al. 
2003). Cysts of B. lyrifera are discordant for the genus 
because they are covered with flat-topped columns 
with little evidence of polygons, though the columns 


Wl 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


are connected by uncoordinated ridges. 

Cysts of Parartemia, an Australian endemic 
genus, are described for the first time. They are smooth, 
rather like those of Artemia (Hill and Shepard 1997), 
but with some inpocketing. Given possible undetected 
variation within species, particularly in those subjected 
to predation (Mura et al. 2002), it is unwise to compare 
species within genera. 

None of the Australian cysts examined here 
have extreme anti-predation devices such as 
honeycombing or long spines (Dumont et al. 2003). 
Some species have minor protruding structures which 
may assist against predation. The most developed are 
the bifid/trifid/recurved spines at ridge junctions in B. 
longirostris. Interestingly this species lives in gnammas 
(rock pools) in which many potential predators live 
including flatworms (Pinder et al. 2000). Other species 
of Branchinella with small spines include B. 
complexidigitata in which they are sharp pointed and 
B. hattahensis where the spines are recurved. The 
flanges on the cysts of the undescribed branchipodid 
genus could be antipredation devices, but another 
explanation is that they assist in holding the egg mass 
together (as observed when trying to separate the 
cysts). The value of such a strategy is questionable, 
given the decreased possibly of dispersal of a clumped 
egg mass, but perhaps it helps to deter egg predators. 

The cyst shell structure (especially the cross- 
sectional structure) in this study ranged from relatively 
simple (e. g., in Parartemia) to complex (e. g., in 
Branchinella budjiti and an undescribed branchipodid 
genus). During the examination of Branchinella 
occidentalis the first set of cysts appeared to be not 
completely mature so a second set was examined. This 
second set was of a very different size and form. Thus, 
it appears that either the species has two different cyst 
morphs or there may be a cryptic species in what is 
now known as B. occidentalis. At least other species 
are known to have two cyst types: Eubranchipus 
serratus (Hill and Shepard 1997) and Streptocephalus 
sealii (R. Hill, pers.com.). During the course of this 
study the only cyst structure found that has not been 
seen in any other fairy shrimp before was the external 
flanges and the parallel columns in the cortex of the 
undescribed genus of branchipodid fairy shrimp. 
Therefore, with minor exception, the maximum range 
of structural diversity in cysts may have been 
described. Whether or not this is true will be decided 
by examination of the remaining undescribed cysts of 
other world genera. 

The structure of the cyst shell, especially in 
the alveolar layer, begs a functional explanation. It is 
likely that the structure represents a compromise 
between crush-resistance and rehydration ability. But 
other functions that have been suggested include: 


78 


protection from abrasion and ultraviolet light (Belk 
1969); a float for cysts (Morris and Afzelius 1967); a 
barrier against physio-chemical stresses and an aid to 
gaseous exchange (Tommasini and Sabelli 1989); an 
anti-predation device (Dumont et al. 2003) and, 
dispersal aids or inhibitors (Brendonck et al. 1992). 


ACKNOWLEDGEMENTS 


We thank Bruce Scott for his masterful supervision of 
the scanning electron microscope. 


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Geddes, M.C. (1973). Studies on Australian Anostracans 
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Hamer, M., Brendonck, L. Coomans, A. and Appleton, C. 
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Hamer, M., Brendonck, L. Coomans, A. and Appleton, C. 
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Henry, M. (1924). A monograph on the freshwater 
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Herbert, B. and Timms, B.V. (2000). A new species of 
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Hill, R.E. and Shepard, W.D. (1997). Observations on the 
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Linder, F. (1941). Contributions to the morphology and the 
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Maeda-Martinez, A.M., Belk, D., Obreg6n-Barboza, H and 
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Morris, J. E. and Afzelius, B. A. (1967). The structure of 
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Mura, G. (1986). SEM morphological survey on the egg 
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Mura, G. (1991a). SEM morphology of resting eggs in the 
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Mura, G., Zarattini, P. and Petkowski, S. (2002). 
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(2000). Granite outcrop pools in south-western 
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Sanoamuang, L., Saenghan, N., and Murugan, G. (2003). 
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descriptions of some new genera and species. 
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Schwartz, K.Y. (1917). Results of Dr. E Mjobergs Swedish 
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Shepard, W.D. and Hill, R.E. (2001). Anostracan cysts found 
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Linderiellidae), a fairy shrimp from south-eastern 
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Thiéry, A. and Gasc, C. (1991). Resting eggs of Anostraca, 
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259. 

Timms, B.V. (2001). Two new species of fairy shrimp 
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Timms, B.V. (2002). The fairy shrimp genus Branchinella 
(Crustacea: Anostraca: Thamnocephalidae) in 
Western Australia, including a description of four 
new species. Hydrobiologia 486, 71-89. 

Timms, B.V. (In press). A key to the fairy shrimps 
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Timms, B.V. and Geddes, M.C. (2003). The fairy shrimp 
genus Branchinella Sayce, 1903 (Crustacea: 
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Vol 3, pp 251-276. 


80 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


APPENDIX 1 
Locality data for species examined. 


Branchinella affinis: Far North Bulla claypan, 
Rockwell Station, 180 km SW of 
Cunnamulla, Queensland. 28° 52’ S, 144° 
56’E, 27-11-2000. 

Branchinella arborea: Marsilea Pool, Bloodwood 
Station, 130km NW of Bourke, NSW. 29° 
33’S, 144° 52’E, 1-vi-1999. 

Branchinella australiensis: Lower Crescent Pool, 
Bloodwood Station, 130 km NW of 
Bourke, NSW, 29° 33’S, 144° 52’E, 1-vi- 
1999. 

Branchinella basispina: Homestead Dam, 
Balladonia Station, 220 km E of Norseman, 
WA. 32° 28’S, 123° 52’E, 3-iii-1975. 

Branchinella buchananensis: Gidgee Lake, 
Bloodwood Station, 130 km NE of Bourke, 
NSW, 29° 33’S, 144° 50’E, 28-ix-1998. 

Branchinella budjiti: a claypan on Muella Station, 
128 km NE of Bourke, NSW, 29° 31’S, 
144° 56’E, 6-xii-1999. 

Branchinella campbelli: Muella Lake, Tredega 
Station, 140 km NW of Bourke, NSW, 29° 
31’S, 144° 53’E, 27-vi-1998. 

Branchinella complexidigitata: Lake Arro near 
Eneabba, 300 km N of Perth, WA, 29° 
44’S, 115° 10’E, 2-1x-1999. 

Branchinella dubia: pool 89 km from Derby on the 
Gibb R Road, Kimberley, WA, 17° 26’S, 
124° 36’E, 31-1-1985. 

Branchinella frondosa: Steves Pool, Muella Station, 
128 km NW of Bourke, NSW, 29° 33’S, 
144° 55’E, 24-xi-1998. 

Branchinella hattahensis: Mid Kaponyee Lake, 
Currawinya National Park, SW 
Queensland, 28° 50’S, 144° 19’E, 5-vii- 
19978 

Branchinella kadjikadji: claypan on Kadyji Kadji 
Station, 35 km ENE of Morewa, WA, 29° 
08’S, 116° 24E, 14-viii-1998. 

Branchinella lamellata: a claypan near Warburton 
Crossing, Clifton Hills Station, NE South 
Australia, 27° 02’S, 138° 53’E, 5-xii-2000. 

Branchinella longirostris: Warrdagga Rock, via 
Paynes Find, WA, 29° 24’S, 117° 30‘E, 26- 
viii-2001. 

Branchinella lyrifera: Turkey claypan, Bloodwood 
Station, 130 km NE of Bourke, NSW, 29° 
34’S, 144° 50’E, 1-iv-1999. 

Branchinella nana: Lake Arrow, near Kalgoorlie, 
WA, 30° 32’S, 121° 24’E, 14-v-1995. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Branchinella nichollsi: Lake Hannan near 
Kalgoorlie, WA, 30° 40’S, 121° 28’E, 17- 
iii- 1937. 

Branchinella occidentalis: Freshwater Lake, 
Bloodwood Station, 130 km NW of 
Bourke, 29° 29’S, 144° 50’E, 2-vii-1997 
(Type I); Plover claypan, Bloodwood 
Station, 130 km NW of Bourke, 29° 29’S, 
144° 48’E, 29-vi-1998 (Type II). 

Branchinella pinnata: a blackbox swamp, Tredega 
Station, 140 km NW of Bourke, NSW, 29° 
29°S, 144° 52’E, 30-vi-1999. 

Branchinella probiscida: Darko claypan, 
Currawinya National Park, SW 
Queensland, 28° 52’S, 144° 18’E, 4-x1- 
1999. 

Branchinella simplex: Lake Arrow, near Kalgoorlie, 
WA, 30° 32’S, 121° 24’E, 14-v-1995. 

Branchinella wellardi: Marsilea Pool, Bloodwood 
Station, 130km NW of Bourke, NSW. 29° 
33’S, 144° 52’E, 1-vi-1999 

Parartemia contracta: small lake adjacent to Lake 
O’Grady, WA, 30° 25’S, 117° 25’E, ?-viii- 
1978. 

Parartemia cylindrifera: Lake Grace, WA, 33° 06’S, 
18° 24’E, 24-viii-1978. 

Parartemia informis: Lake Ballard, WA, 29° 37’S, 
121° OVE, 18-viii-1978. 

Parartemia minuta: Lower Bell Lake, Bloodwood 
Station, 130 km NW of Bourke, NSW, 29° 
00’S, 144° 48’E, 28-ix-1998. 

Parartemia zietziana: small lake on “Flowerfield’ 
via Beeac, Victoria, 38° 10’S, 143° 09’E, 
15-viii- 1970. 

Streptocephalus queenslandicus: fish rearing ponds, 
Walkamin Research Station, via Atherton, 
Queensland, 17° 08’S, 145° 26’E, ii-1997. 

Streptocephalus n.sp. a: Pine Tree Pool, Bloodwood 
Station, 130 km NE of Bourke, NSW, 29 
29’S, 144 49’E, 2-xii-1999. 

Streptocephalus n.sp.b: Box Hole, 4 km S of 
Homestead, Currawinya National Park, 
Qld., 28 51’S, 144 29’E, 1-iv-1999. 

Undescribed new branchiopod genus: Marsilea Pool, 
Bloodwood Station, 130km NW of Bourke, 
NSW. 29° 33’S, 144° 52’E, 24-viii-1998. 


81 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


Figure 1. Parartemia contracta cyst. Bar = 50 um. 

Figure 2. Parartemia contracta cyst shell cross section. Bar = 10 um. 
Figure 3. Parartemia cylindrifera cyst. Bar = 50 um. 

Figure 4. Parartemia cylindrifera cyst shell cross section. Bar = 10um. 
Figure 5. Parartemia minuta cyst. Bar = 50 um. 

Figure 6. Parartemia minuta cyst shell cross section. Bar = 5 um. 


82 Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 7. Parartemia zietziana cyst. Bar = 50 um. 

Figure 8. Parartemia zietziana cyst. Bar = 5O um. 

Figure 9. Parartemia zietziana cyst shell cross section. Bar = 5 um. 

Figure 10. Undescribed branchipodid species cyst. Bar = SO um. 

Figure 11. Undescribed branchipodid species cysts clumped together. Bar = 50 um. 
Figure 12. Undescribed branchipodid species cyst surface. Bar = 10 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 83 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


84 


Undescribed branchipodid species cyst shell cross section. Bar = 5 um. 
Streptocephalus queenslandicus cyst. Bar = 50 um. 

Streptocephalus queenslandicus cyst. Bar = 20 um. 

Streptocephalus queenslandicus cyst shell cross section. Bar = 5 um. 
Streptocephalus n. sp. a cyst. Bar = 50 um. 

Streptocephalus n. sp. a cyst ridges. Bar = 10 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 19. Streptocephalus n. sp. a cyst shell cross section. Bar = 5 wm. 
Figure 20. Streptocephalus n. sp. b cyst. Bar = 50 um. 

Figure 21. Streptocephalus n. sp. b cyst ridges. Bar = 10 um. 

Figure 22. Streptocephalus n. sp. b cyst shell cross section. Bar = 10 um. 
Figure 23. Branchinella affinis cyst (not quite mature). Bar = 20 um. 
Figure 24. Branchinella arborea cyst. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 85 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


86 


Figure 25. Branchinella arborea cyst ridge. Bar = 5 um. 

Figure 26. Branchinella arborea cyst shell cross section. Bar = 5 um. 
Figure 27. Branchinella australiensis cyst. Bar = 20 um. 

Figure 28. Branchinella australiensis cyst ridge structure. Bar = 5 um. 
Figure 29. Branchinella australiensis cyst shell cross section. Bar = 10 um. 
Figure 30. Branchinella basipina cyst. Bar = 50 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


BUCHAN 


Figure 31. Branchinella basipina cyst shell cross section. Bar = 10 um. 
Figure 32. Branchinella buchananensis cyst. Bar = 50 um. 

Figure 33. Branchinella buchananensis cyst shell cross section. Bar = 10 um. 
Figure 34. Branchinella budjiti cyst. Bar = 20 um. 

Figure 35. Branchinella budjiti cyst shell cross section. Bar = 5 wm. 

Figure 36. Branchinella campbelli cyst. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 87 


88 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


BCONMPL 


© 


”s 

is 

4 
‘F) 


a ~ 
5 
f 

Ba i 


Figure 37. Branchinella campbelli cyst shell cross section. Bar = 5 um. 

Figure 38. Branchinella complexidigitata cyst. Bar = 50 um. 

Figure 39. Branchinella complexidigitata cyst surface structure. Bar = 10 um. 
Figure 40. Branchinella complexidigitata cyst surface structure. Bar = 10 um. 
Figure 41. Branchinella complexidigitata cyst shell cross section. Bar = 10 um. 
Figure 42. Branchinella dubia cyst. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 43. Branchinella dubia cyst surface. Bar = 10 um. 

Figure 44. Branchinella dubia cyst shell cross section. Bar = 10 um. 
Figure 45. Branchinella frondosa cyst. Bar = 20 um. 

Figure 46. Branchinella frondosa cyst shell cross section. Bar = 10 um. 
Figure 47. Branchinella hattahensis cyst. Bar = 50 um. 

Figure 48. Branchinella hattahensis cyst surface structure. Bar = 10 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 89 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


Figure 49. Branchinella hattahensis cyst shell cross section with ridge spines. Bar = 10 um. 

Figure 50. Branchinella hattahensis cyst shell cross section. Bar = 10 um. 

Figure 51. Branchinella kadjikadji cyst. Bar = 50 um. 

Figure 52. Branchinella kadjikadji cyst surface. Bar = 10 um. 

Figure 53. Branchinella kadjikadji cyst shell cross section showing outer sublayer of alveolar layer pulled 
away from inner sublayer. Bar = 5 um. 

Figure 54. Branchinella kadjikadji cyst shell fracture showing inner sublayer of alveolar layer. Bar = 2 um. 


90 Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 55. Branchinella lamellata cyst. Bar = 20 um. 

Figure 56. Branchinella lamellata cyst shell cross section. Bar = 5 um. 
Figure 57. Branchinella longirostris cyst. Bar = 50 um. 

Figure 58. Branchinella longirostris cyst surface structure. Bar = 10 um. 
Figure 59. Branchinella longirostris cyst shell cross section. Bar = 10 um. 
Figure 60. Branchinella lyrifera cyst. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 91 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


92 


Figure 61. Branchinella lyrifera cyst ridges, polar view. Bar = 10 um. 
Figure 62. Branchinella lyrifera cyst ridges, side view. Bar = 10 um. 
Figure 63. Branchinella lyrifera cyst shell cross section. Bar = 5 um. 
Figure 64. Branchinella nichollsi cyst. Bar = 50 um. 

Figure 65. Branchinella nichollsi cyst ridge. Bar = 10 um. 

Figure 66. Branchinella nichollsi cyst shell cross section. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 67. Branchinella occidentalis cyst (type 1). Bar = 50 um. 

Figure 68. Branchinella occidentalis cyst (type I) shell cross section. Bar = 10 um. 
Figure 69. Branchinella occidentalis cyst (type Il). Bar = 100 um. 

Figure 70. Branchinella occidentalis cyst (type II) surface. Bar = 10 um. 

Figure 71. Branchinella occidentalis cyst (type I) shell cross section. Bar = 20 um. 
Figure 72. Branchinella pinnata cyst. Bar = 20 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 93 


94 


FAIRY SHRIMP CYST SHELL MORPHOLOGY 


PINNATA 


Figure 73. Branchinella pinnata cyst shell cross section. Bar = 10 um. 
Figure 74. Branchinella probiscida cyst. Bar = 20 um. 

Figure 75. Branchinella probiscida cyst surface. Bar = 10 um. 

Figure 76. Branchinella probiscida cyst shell cross section. Bar = 5 um. 
Figure 77. Branchinella simplex cyst. Bar = 20 um. 

Figure 78. Branchinella simplex cyst shell cross section. Bar = 5 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 


B.V. TIMMS, W.D. SHEPARD AND R.E. HILL 


Figure 79. Branchinella wellardi cyst. Bar = 20 um. 
Figure 80. Branchinella wellardi cyst ridges. Bar = 10 pm. 
Figure 81. Branchinella wellardi cyst shell cross section. Bar = 10 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 95 


Figure 7a j Branchinella pinhead yt re 
Figure 75. Brinchinella pribascida 


The Yule Island Fauna and the Origin of Tropical Northern 
Australian Echinoid (Echinodermata) Faunas 


I.D. LINDLEY 


Department of Geology, Australian National University, Canberra, A.C.T. 0200. 
(lindley @ geology.anu.edu.au) 


Lindley, I.D. (2004). The Yule Island Fauna and the Origin of Tropical Northern Australian Echinoid 
(Echinodermata) Faunas. Proceedings of the Linnean Society of New South Wales 125, 97-109. 


Systematic description of the rich Lower Pliocene echinoid fauna from Yule Island, Papua New 
Guinea, and recently available palaeogeographic data from onshore Papua, the Gulf of Papua and Torres 
Strait have provided new insights into the origins of the extant tropical northern Australian echinoid fauna. 
Previous studies of echinoderm origins, hindered by a lack of fossil evidence, concluded that tropical northern 
Australian echinoderms were derived predominantly by Recent migrations from East Indian and West Pacific 
stocks. However, 47 per cent of species from the Yule Island fauna are extant in northern Australian waters, 
indicating that present faunistic patterns were to a large extent, in-place by at least the Lower Pliocene. 
Palaeogeographic evidence supports the earlier observations of H. Barraclough Fell, that migration of echinoid 
stock into (and out of) eastern New Guinea and tropical northern Australia probably occurred during the 
Lower to Middle Miocene, when widespread tropical to sub-tropical reef development occurred across a 
5600 km belt from SE Asia through New Guinea and into the SW Pacific as far as Fiji. This favourable 
pathway for exchange between echinoid stocks disappeared during the Upper Miocene, when the onset of 
tectonic instability throughout the region, and the establishment of a discontinuous volcanic arc, resulted in 
influx of terrigenous sediments and may have caused the death of the reef complex. This pattern of 


sedimentation has persisted to the present. 


Manuscript received 11 April 2003, accepted for publication 20 August 2003. 


KETWORDS: Biogeography, East Indies, Echinoidea, Great Barrier Reef, Palaeogeography, Papua New 


Guinea, Pliocene, Queensland. 


INTRODUCTION 


Echinoderms are one of the most intensively 
studied shallow-water faunas of tropical northern 
Australia (Endean 1982). Their specific composition 
and general distribution in the region is well known 
because of the work of A.H. Clark (1908, 1911a, 1915, 
1915-1967), H.L. Clark (1907, 1909, 1915, 1921, 
1925, 1926, 1928, 1932, 1938, 1946), Endean (1953, 
1956, 1957, 1961, 1965), A.M. Clark (1970), A.M. 
Clark and Rowe (1971), and Gibbs et al. (1976). 
However, studies of the origins of these echinoderm 
faunas have been hindered by the lack of outcropping 
marine Tertiary sequences with macro-invertebrate 
faunas (Endean 1957). McNamara and Kendrick 
(1994) described the Middle Miocene echinoid fauna 
from the Poivre Formation on Barrow Island, off the 
Western Australian coast, about 1 500 km WSW of 
the northern Great Barrier Reef (GBR). The Poivre 
Formation represents the northern-most exposure of 
Miocene marine deposits in Australia, and contains 


the only tropical Miocene macro-invertebrate fauna 
in Australia (McNamara and Kendrick 1994). The 
diverse Mio-Pliocene echinoid fauna described by 
Jeannet and R. Martin (1937) from Java, about 1 000 
km WNW of the northern GBR, provides closer 
(geological) time links with the region than does the 
Barrow Island fauna. However, caution is necessary 
in reviewing Jeannet and R. Martin’s (1937) fauna, 
with the validity of some species identifications in need 
of re-evaluation and name changes required, in 
accordance with present taxonomic nomenclature. The 
rich Pliocene Yule Island, Papua New Guinea, echinoid 
fauna comprising 19 species, is located less than 300 
km from the northern end of the GBR, and represents 
the nearest Tertiary echinoid fauna to the GBR region 
(Lindley 2003a-c; Fig. 1). This paper examines the 
importance of the Yule Island fauna with respect to 
the origins of extant echinoid faunas of tropical 
northern Australia. 

The taxonomic classification used herein 
follows that of Fell (1966), Fell and Pawson (1966), 
Durham (1966) and Fischer (1966). 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


ARAFURA 
SEA 


AUSTRALIA 


Pacific south 
equatorial current 


Capricorn- 
‘\ Bunker Groups 


Figure 1. Locality map showing proximity of Yule Island to tropical northern Australia. 


FAUNAL PROVINCES AND MAINLAND AND 
REEF ECHINODERM FAUNAS 


Tropical northern Australian seas embrace 
two faunistic provinces, namely, the Tropical 
Australian Province and the Solanderian Province 
(Endean 1957). 

The Solanderian Province (Whitley 1932) is 
restricted to the fauna of the GBR, and includes 
echinoderms which Endean (1957) designated as ‘reef? 
species (Fig. 2). These echinoderms are considered to 
have strong affinities with West Pacific stocks, with 
their pelagic larval stages transported onto the GBR 
by the Pacific south equatorial current Endean (1957). 
Endean (1957) considered that the lack of reef 
structures to the west of Torres Strait has proved a 
major barrier to the migration of reef stocks into the 
Solanderian Province. 

The Tropical Australian Province of Endean 
(1957) incorporates the Banksian Province of Whitley 
(1932) and the Dampierian Province of Hedley (1904, 
1926) and extends from Geraldton, on the Western 


98 


Australian coast, to Wide Bay (26°S) on the southern 
Queensland coast (Fig. 2). As originally proposed, the 
Banksian Province included the marine faunas of 
coastal districts of Queensland, distinct from those of 
the GBR, and the Dampierian Province along the 
Western Australian coast. However, Endean (1957) 
concluded that Torres Strait does not present a major 
biogeographical boundary separating the Dampierian 
and Banksian Provinces, and that they should be 
considered one single faunistic unit. The Tropical 
Australian Province contains an echinoderm fauna 
designated by Endean (1957) as ‘mainland’ species, 
typically found in habitats dominated by terrigenous 
sediments. Mainland echinoderms have strong 
affinities with East Indies stocks, with the principal 
exchange route being via Torres Strait and the Arafura 
Sea (Endean 1957). He noted that for Queensland 
waters, there is little intermingling between reef and 
mainland stocks. 

The East Indies Province lies to the north of 
Australia (Fig. 2). H.L. Clark (1946) defined the 
geographic limits of this faunal province, to include 
the sea and its islands between 90° and 155°E, witha . 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


oo & 
aes 


East Indies Province 


AUSTRALIA 


TROPIC OF CAPRICORN 


\\ Geraldton 


Figure 2. Australia and New Guinea showing the distribution of 


biogeographical zones mentioned in the text. 


southern limit coinciding with the northern limit of 
what is now know as the Tropical Australian Province. 
At its eastern extent, it includes the island of New 
Guinea with the eastern boundary following longitude 
155° between the Bismarck Archipelago and the 
Solomon Islands to 5°N, where it turns west to 130°E. 


PREVIOUS BIOGEOGRAPHICAL STUDIES 


Austin Hobart Clark and Hubert Lyman Clark 
Much of our knowledge of shallow-water 
Australian echinoderms is based on the intensive work 
of A.H. Clark and H.L. Clark. A.H. Clark (1911b) 
concluded that ‘The crinoids of Australia have come 
from the north, from the great East Indian 
Archipelago’. H.L. Clark (1946) extended A.H. Clark’s 
observations to other echinoderm groups, and 
concluded that the ‘evidence is overwhelming that 
Australian echinoderms have come southward from 
the East Indian area, either around the eastern end of 
New Guinea or across the Timor Sea’. H.L. Clark 
(1921, 1946) thought that the extant Queensland fauna 
postdated the “depression of land areas east of New 
Guinea which led to the connection of the Coral Sea 
with the western Pacific and the East Indian region’. 
The Dampierian fauna had migrated from farther west 
and was well established when formation of the Torres 
Strait made mingling of the two faunas possible. 


Proc. Linn. Soc. N.S.W., 125, 2004 


H. Barraclough Fell 

Fell (1953) also 
concluded that there had 
been a mainly southward 
migration along the Indo- 
Malayan archipelago, but 
noted that northward 
movements of Australasian 
echinoderms into the Indo- 
Pacific could be detected 
from the Miocene onward. 
He believed that the 
archipelago, or its Tertiary 
equivalents, provided the 
shallow-water migration 
route, both into and out of 
Australia. Fell (1953) noted 
that profound changes in 
fossil Australian echinoid 
faunas occurred during the 
Upper Miocene, an event he 
attributed to cooling climate. 


BISMARCK 
SEA 


Robert Endean 

Endean (1957) saw several obstacles to H.L. 
Clark’s (1946) echinoderm migration patterns around 
New Guinea. Firstly, he believed there was little 
likelihood of any significant exchange of mainland 
species via the northern shores of New Guinea. Deep 
waters are present immediately offshore of the north 
coast of the island, and the large Sepik and 
Mamberamo Rivers, by contributing large volumes of 
freshwater and silt, acted as natural barriers to the 
spread of littoral echinoderms. He argued that in 
eastern New Guinea there was a lack of suitable 
habitats for mainland echinoderms, with the Papua- 
Louisiade Barrier Reef, extending along the 
southeastern coast of New Guinea and to the east of 
the island, representing an additional barrier to the 
migration of mainland species. Endean (1957) 
considered that the Fly River, which empties large 
amounts of silt and freshwater into the Gulf of Papua, 
acted as an ecological barrier to the spread of reef 
species to the west of Torres Strait. 

Endean (1957) concluded that extant tropical 
northern Australian echinoderm faunas were derived 
predominantly from Recent migrations of East Indian 
and West Pacific stocks. With the Queensland 
mainland species having strong affinities with those 
of the East Indies, the principal exchange route of these 
forms was via the Torres Strait and the Arafura Sea, 
the deepwater of the Coral Sea serving as a barrier to 
the spread of mainland species to the West Pacific. 


99 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


Endean (1957) believed that reef species have strong 
ties with those of the West Pacific. Given that very 
few of the reef species which occur outside of 
Australian waters are present in waters west of Torres 
Strait, he concluded that there was insignificant 
exchange of reef species in Queensland waters with 
those of the East Indies by way of the Torres Strait. 
Rather, the main route of exchange of Queensland and 
West Pacific echinoderms must have been by way of 
either the south coast of eastern New Guinea or the 
Coral Sea. However, since eastern New Guinea and 
the GBR are separated by deep water from West Pacific 
localities, Endean (1957) believed the most likely route 
of exchange of Queensland’s reef species and West 
Pacific stocks must have been via pelagic larval stages 
swept into the Coral Sea by the unidirectional Pacific 
south equatorial current. 


ANALYSIS OF THE YULE ISLAND ECHINOID 
FAUNA 


The echinoid faunas from the Lower Pliocene 
of Yule Island, Middle Miocene of Barrow Island and 
the Mio-Pliocene of Java each contain mixed 
assemblages of clypeasteroid, regular and spatangoid 
echinoids, representative of very different 
palaeoecologies. Because none of these faunas is 
dominated by an echinoid group unique to a particular 
palaeoecology, valid comparisons can be made 
between these fossil faunas. Using a similar argument, 
these comparisons are extended to the extant faunas 
of tropical northern Australia. 

Of the 19 echinoid species described from 
Yule Island, a group of seven species is not recorded 
from either the Barrow Island and Javanese faunas or 
the extant echinoid fauna of tropical northern Australia 
(Table 1 - located after the reference list). These species 
include Schizechinus cf. tuberculatus (Pomel), 
Phyllacanthus sp., Laganum depressum sinaiticum 
Fraas 1867, and L. depressum delicatum Mazzetti 
1894, Palaeostoma kairukuensis Lindley 2003c, 
Maretia cordata Mortensen 1948, and Schizaster 
(Schizaster) alphonsei Lindley 2003c. Only two of 
these echinoids are endemic, viz. S.(S.) alphonsei and 
P. kairukuensis. Laganum depressum sinaiticum, and 
L. depressum delicatum are notable in that they are 
otherwise only recorded extant in the western Indian 
Ocean (Persian Gulf). Similar observations of western 
Indian Ocean affinities have been made amongst fossil 
reef corals from the nearby Upper Pliocene-Lower 
Pleistocene Era Beds, northwest of Yule Island, notably 
the faviid coral Parasimplastrea simplicitexta 
Umbgrove 1942 (Veron and Kelley 1988). 


100 


From an analysis of the remaining 12 species, 
nine occur in tropical waters of northern Australia 
either in mainland or reef stocks. These species include 
Cyrtechinus verruculatus (LUutken), Prionocidaris 
verticillata (Lamarck 1816), Parasalenia poehli 
Pfeffer 1887, Clypeaster humilis (Leske), C. latissimus 
(Lamarck), C. reticulatus (Linnaeus 1758), Laganum 
depressum Lesson in L. Agassiz 1841, L. decagonale 
(de Blainville 1827) and Maretia planulata (Lamarck). 
Nine species are known as fossil in the Mio-Pliocene 
of Java, viz. Prionocidaris verticillata, Phyllacanthus 
imperialis var. javana K. Martin 1885, Temnotrema 
macleayana (Tenison-Woods 1878), L. depressum, L. 
decagonale, C. reticulatus, C. humilis, M. planulata 
and Eupatagus (Eupatagus) pulchellus (Herklots). The 
presence of these species as fossil in the Mio-Pliocene 
of Java, and the observed low levels of species 
endemism, suggests that the two populations were 
well-connected prior to and during the Pliocene. By 
contrast, the Yule Island fauna, as with the Javanese 
fauna, does not have any species in common with the 
Middle Miocene fauna of Barrow Island (Table 1). 

Significantly, 47 per cent of echinoid taxa 
(nine species) from the Yule Island fauna, now inhabit 
northern Australian waters. Clearly, this observation 
conflicts with Endean’s (1957) proposal that tropical 
Australian echinoderms were derived from Recent 
migrations from East Indian and West Pacific stocks. 
Of these nine species, four can be included as mainland 
stock, viz. C. verruculatus, C. reticulatus, C. humilis 
and C. latissimus; three are of reef stock, viz. L. 
decagonale, P. verticillata and P. poehli; and two 
species occur in both reef and mainland waters viz. L. 
depressum and M. planulata. Endean (1957) noted a 
similar, but limited intermingling of echinoderm stock, 
including the echinoids M. planulata and Salmacis 
sphaeroides, in waters surrounding the Low Isles, a 
coral-dominated locality relatively close to the 
mainland (about 11 km distant), north of Cairns. He 
regarded these species as having ‘strayed into the coral 
environment’. Such a near-shore intermingling is 
envisaged during the Pliocene of Yule Island (Fig. 3). 


MIO-PLIOCENE PALAEOGEOGRAPHY OF THE 
GULF OF PAPUA AND ADJACENT AREAS 


Advances in the knowledge of the geological 
evolution of onshore Papua and the western Coral Sea 
have emerged with the synthesis of extensive oil well 
and outcrop data gathered during hydrocarbon 
exploration of the past 25 years (Home et al. 1990; 
Carman 1993; Struckmeyer et al. 1993 amongst 
others). These advances have permitted a detailed - 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


depocentre 


Probably locally 
emergent 


% 


FLUVIAL-ALLUVIAL 


Se 
> 
4FYLNAIIDOdId 2 


GNVINDIGLS 


understanding of the development of Mio-Pliocene 
sedimentary facies and the region’s tectonic history. 
Tectonic stability during the Lower to Middle 
Miocene was associated with the development of an 
extensive tropical to sub-tropical platform carbonate 
reef complex, not only in the Papuan Basin, but in a 
region extending from SE Asia to Fiji, a distance of 
some 5 600 km, and including northern Australia 
(Coleman and Packham 1976: Home et al. 1990; 
McNamara and Kendrick 1994). During the late 
Miocene, the onset of tectonic compression along the 


Proc. Linn. Soc. N.S.W., 125, 2004 


Volcanolithic sediments 
Data points 


C3 Biostromal/algal reefs 


SEDIMENTS 


© 
=} 
2. 
© 
oO. 
— 
fo) 
= 
=) 
1) 


DELTAIC/PRO-DELTAIC 


Great Barrier Reef 


Reefal precursor of 
in Torres Strait, reef development (including the Kairuku Formation at Yule Island) along the northern coastline of the 


Figure 3. Pliocene (c. N20) palaeogeography of onshore Papua-Gulf of Papua-Torres Strait region showing emergence 
Gulf of Papua and reefal precursors to the Great Barrier Reef. Modified after Carman (1993). 


edge of the Pacific Plate across much of the region 
resulted in major uplift and the establishment of a 
discontinuous volcanic arc. In New Guinea, as 
elsewhere throughout the region, the influx of 
terrigenous sediments, to the north and south of an 
uplifting central cordillera, resulted in the death of the 
Miocene reef complex (Home et al. 1990; Carman 
1993; Struckmeyer et al. 1993). The western end of 
the Gulf of Papua became emergent during this time 
and sedimentation was predominantly on a fluvial/ 
alluvial plain adjacent to a volcanic province (Carman 


101 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


1993; Fig. 3). South of a fluctuating Pliocene shoreline, 
poorly sorted deltaic/pro-deltaic sediments were 
deposited across a greater part of the rapidly subsiding 
Gulf of Papua (Home et al. 1990; Carman 1993). More 
than 4 000 m of sediment were deposited in the Vailala- 
Purari depocentre, near the mouth of the Purari River 
(Carman 1993), indicative of very rapid subsidence. 
Subsidence and the large quantities of clastic sediment 
were deterrents to coral growth across a large part of 
the region. However, biohermal reef limestone 
(foraminiferal zone c. N20: Lower Pliocene/Upper 
Pliocene boundary), surrounded by interpreted 
argillaceous micritic limestone, penetrated in the 
Anchor Cay 1 Well, only 250 km WSW of Yule Island, 
is associated with reefal precursors of the present day 
GBR which existed farther south (Carman 1993). 
These patterns of sedimentation have continued to the 
present day (Home et al. 1990). 


ORIGIN OF TROPICAL AUSTRALIAN 
ECHINOID FAUNAS 


Preamble 

Migration of adult littoral echinoderms may 
occur via connected shallow-water, typically near- 
shore habitats. Deep water, a lack of interconnected 
reefal structures, and large influxes of freshwater and 
silt poured out from large coastal rivers, all act as 
ecological barriers to their spread (Endean 1957). 
However, for echinoderms possessing prolonged larval 
stages, transport of pelagic young by currents, may 
distribute species across deep water (Endean 1957; 
Nichols 1969). 


Lower to Middle Miocene origin of the Yule Island 
fauna 

The Yule Island echinoid fauna comprises 
representatives of reef and mainland species and 
species common to both reef and mainland faunas. 
Comparisons show that, with the relatively low level 
of species endemism evident in the Yule Island 
echinoid fauna, prior to the Lower Pliocene the fauna 
was well-connected with at least the population in Java, 
having not developed in isolation. Palaeogeographic 
evidence indicates that an excellent interconnected 
exchange route for echinoderms existed during the 
Lower to Middle Miocene when an extensive tropical 
to sub-tropical reef complex existed across a 5 600 
km belt extending from SE Asia through New Guinea 
to Fiji. Both reef and mainland species migrated along 
this reef complex. 

Endean’s (1957) West Pacific influence on 
extant reef echinoid populations of tropical northern 
Australia may be more apparent than real. For all 


102 


echinoids, reef and mainland, only one, Rhynobrissus 
hemiasteroides Agassiz, is endemic to the West Pacific. 
Eleven species occur in the Indian Ocean and East 
Indies, and 21 are widely distributed throughout the 
Indo-Pacific. The origins of the reef stock at Yule 
Island, with its apparent ‘West Pacific affinity’, is 
readily accounted for by the interconnected pathway 
afforded by the 5 600 km long Lower to Middle 
Miocene reef complex that extended well into the SW 
Pacific. 

Fell (1953) noted that a great extinction of 
the Australasian echinoid stocks occurred during the 
late Miocene, an event he and Fleming (1949) 
attributed to a cooling climate. The cooling event 
coincided with the previously noted retreat of 
widespread platform carbonate deposition from SE 
Asia and the SW Pacific. However, the demise of the 
Miocene carbonate platform in the western Coral Sea 
was accompanied by a resurgence of tectonic instability 
and associated volcanism. This tectonic instability is 
evident in the post-Miocene geological record that 
succeeds carbonate deposition throughout the region 
(Coleman and Packham 1976) and it is possible that 
an increase in volcanic activity, and in-turn 
atmospheric volcanic aerosols, was responsible for the 
late Miocene cooling climate. 

The late Miocene extinction saw the 
disappearance of, for example, the warm-water 
echinoid genera Schizaster L. Agassiz 1836 and 
Phyllacanthus Brandt 1835 from New Zealand (Fell 
1953). Fell (1953) considered that the impact of the 
extinction event in low latitudes was reduced. A 
comparison of the Yule Island fossil fauna with low 
latitude Miocene faunas from India, Java and Fiji 
supports this view. Eight species from the Yule Island 
fauna, P. imperialis var. javana, P. verticillata, T. 
macleayana, L. decagonale, L. depressum, C. 
reticulatus, C. humilis and E.(E.) pulchellus are known 
from the Miocene faunas of these regions (Jeannet and 
R. Martin 1937; Mortensen 1948; Lindley 2001, 
2003a-c). 


Origin of Tertiary and extant northern Australian 
echinoid faunas 

As previously noted, mainland and reef 
echinoid populations are believed to have established 
themselves in the region to the north of Australia and 
adjacent areas during the Lower to Middle Miocene 
when platform carbonate sedimentation prevailed 
throughout much of SE Asia and the SW Pacific. Fell 
(1953) proposed a broadly similar outline for 
Australasian echinoid origins. 

The Middle Miocene echinoid fauna from the 
Poivre Formation on Barrow Island is a far from 
complete representation of the original fauna 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


(McNamara and Kendrick 1994). The fauna (at a 
generic level) has much in common with the Miocene 
faunas of Java and India (McNamara and Kendrick 
1994), but its relative geographic isolation during the 
Miocene, 15-20°S of Yule Island and Java, may 
explain the faunal mismatch at species level. Barrow 
Island during the early Miocene was located about 
40°S, Yule Island 25°S and eastern Indonesia 20°S 
(Veevers et al. 1991: Fig. 12). McNamara and 
Kendrick (1994) noted that, at a generic level, the 
echinoid fauna has strong affinities with modern 
communities of the region. 

Endean (1957) considered that the extant NW 
Australian echinoderm fauna, dominated by mainland 
species, has strong affinities with East Indies stocks, 
but exhibits a high degree of endemism. The deep water 
of the Timor and Arafura Seas appears to have served 
as a barrier to any significant post-Miocene exchange 
between the East Indies and NW Australian mainland 
faunas, and exchange with Queensland mainland 
faunas has only been possible since the opening of 
Torres Strait. The strait was emergent during at least 
the late Miocene-early Pliocene (Fig. 3). Ekman (1953) 
noted that the average age of echinoid species was at 
most 4-6 million years, and the origin of many endemic 
species in the NW Australian mainland fauna may have 
resulted from the long period of geographic isolation 
following the demise of the Lower to Middle Miocene 
(10-20 Ma) reef. 

The Lower Pliocene coral growth at Yule 
Island (foraminiferal zones N18/N19-N20: 4-6 Ma) 
was part of a chain of interconnected reefs extending 
NW along the northern margin of the Gulf of Papua, 
remnants of the formerly extensive Miocene reef 
complex (Fig. 3). Reef species were confined to these 
coral structures, which were in-turn flanked by 
terrigenous sediments, a habitat dominated by 
mainland species. In the southern Gulf of Papua, reef 
precursors to the GBR did not appear until 3-4 Ma 
(Lower Pliocene/Upper Pliocene boundary: 
foraminiferal zone c. N20: Carman 1993; Haig 1996). 
These precursor reefs were situated only 250 km WSW 
of Yule Island and extended farther south (Carman 
1993), indicating that, although large quantities of 
terrigenous sediment at this time were deterrents to 
coral growth, reefs were able to establish themselves 
in parts of the Gulf of Papua. Of the nine echinoid 
species (47 per cent) of the Yule Island fauna that are 
recorded extant in northern Australian waters, five are 
recorded on the GBR, suggestive of ties between both 
faunas. With major barriers to echinoderm migration 
existing to the west (Torres Strait was emergent) and 
along northern New Guinea (deep waters and 
terrigenous sedimentation), it is likely that species 


Proc. Linn. Soc. N.S.W., 125, 2004 


exchange occurred from existing Pliocene Gulf of 
Papua echinoid stocks to proto-GBR structures and 
farther south. 


ACKNOWLEDGMENTS 


The author kindly acknowledges Prof. K.S.W. 
Campbell for his constructive review of and improvements 
to the manuscript. The comments of anonymous referees 
and Dr M.L. Augee also improved the manuscript. 


REFERENCES 


Carman, G.J. (1993). Palaeogeography of the Coral Sea, 
Darai and Foreland megasequences in the 
eastern Papuan Basin. In ‘Petroleum Exploration 
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151. 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


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Linnean Society of New South Wales 124, 153- 
162. 

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Australia. Australian Naturalist 8, 166-167. 


Proc. Linn. Soc. N.S.W., 125, 2004 


105 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


TABLE 1: Distribution of selected shallow-water tropical echinoid groups in the East Indies and Australia. 

Compiled from Endean (1957); A.M. Clark and Rowe (1971); Gibbs et al. (1976); Jeannet and R. Martin 

(1937); Philip (1963); Mortensen (1943); H.L. Clark (1946); Lindley (2001, 2003a,b,c); McNamara and 
Kendrick (1994); and McNamara and Philip (1980). 


Species Java, Mio- Yule Is, Barrow Is, Australian Australian 
Pliocene Lower Middle ‘mainland’ ‘reef’ 
Pliocene Miocene species species 


CIDARIDAE 
Phyllacanthus dubius 


Phyllacanthus dubius var. 
sundaica 

Phyllacanthus imperialis 
Phyllacanthus imperialis var. 
javana 


Phyllacanthus sp. _ 
Phyllacanthus cf. clarkeii 
Prionocidaris bispinosa 
Prionocidaris verticillata 


Prionocidaris baculosa 
Prionocidaris baculosa var. 
annulifera 

Stylocidaris_reini 

Cidaris mertoni 

Cidaris aculeata 


Cidaris_sp. 

Chondrocidaris sundaica 
Eucidaris_ sp. 

Goniocidaris cf. murrayensis 


DIADEMATIDAE 
Diadema setosum 
Diadema savignyi 
Echinothrix calamaris 
Echinothrix diadema 
Astropyga radiata 


PARASALENIIDAE 
Parasalenia 


Parasalenia gratiosa 


Parasalenia_ sp. 


TOXOPNEUSTIDAE 
Schizechinus cf. tuberculatus 
Cyrtechinus verruculatus 
Nudechinus darnleyensis 
Nudechinus multicolor 

| Tripneustes gratilla 
Tripneustes pregratilla 
Gymnechinus epistichus 
Toxopneustes pileolus 


106 Proc. Linn. Soc. N.S.W., 125, 2004 


Species 


Java, Mio- 


Pliocene 


ECHINOMETRIDAE | 


I.D. LINDLEY 


Yule Is, 
Lower 
Pliocene 


Barrow Is, 
Middle 
Miocene 


Australian 
‘mainland’ 
species 


Echinometra mathaei 
Heterocentrotus mammillatus 


Australian 
‘reef’ 
species 


Echinostrephus aciculatus 


= 


Echinostrephus molaris 


mK 1K [XK LX 


ECHINONEIDAE 


Echinoneus cyclostomus 
Echinoneus abnormalis 


TEMNOPLEURIDAE 


Temnopleurus alexandri 


Temnopleurus toreumaticus 
Salmacis belli 


Salmacis sphaeroides 


KM 1K [XK LX 
x< 


Salmacis sphaeroides belli 
Salmacis rarispina 


Salmacis bicolor 
Temnotrema macleayana 


mK [MS [XS [>< 


Temnotrema_bothryoides 


Temnotrema siamense 


Temnotrema phoenissa 


Opechinus cf. collignoni 


Opechinus cf. cheribonensis 


Opechinus madurae 


Opechinus percultus 


Opechinus percultus var. 
oligoporus 


Martinechinus_molengraaffi 


mM (dK LK LK [X< 


Microcyphus_ sp. 


mM [>< |< [>< 


Desmechinus rembangensis: 
Desmechinus erbi 
Mespilia globulus 


Temnopleurid sp. 


ARACHNOIDIDAE 
Arachnoides placenta 


CLYPEASTERIDAE 


Clypeaster reticulatus 
Clypeaster humilis 1 


Clypeaster latissimus 


Clypeaster telurus 


Clypeaster blumenthali 
Clypeaster brevipetalus 


Clypeaster butleri 


Clypeaster cf. malumbang- 
ensis 


K |< [xX |< 


Clypeaster sp. A 


Proc. Linn. Soc. N.S.W., 125, 2004 


107 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


Species Java, Mio- Yule Is, Barrow Is, Australian Australian 
Pliocene Lower Middle ‘mainland’ ‘reef' 
Pliocene Miocene species species 


CLYPEASTERIDAE (Cont) 
Clypeaster sp.B 


LAGANIDAE 

Laganum decagonale 
Laganum depressum 
Laganum depressum var. 
sinaiticum 

Laganum depressum var. 
delicatum 

Laganum herklotzi 
Peronella lesueuri 
Peronella orbicularis 
Sismondia javana 


ASTRICLYPEIDAE 
Echinodiscus angulosus 
Echinodiscus tenuissimus 
Echinodiscus_ sp. 


FIBULARIIDAE 

Fibularia crispa 

Fibularia cf. scabra 
Fibularia rhedeni 

Fibularia ovulum 

Fibularia volva 

Fibularid sp. 

Echinocyamus cf. cribellum 
Echinocyamus sp. 


ECHINOLAMPADIDAE 
Echinolampas ovatus 
Echinolampas elevatus 
Echinolampas depressus 
Echinolampas tumulus 


PLIOLAMPADIDAE 


Pliolampas minutus 
Pliolampas javanus 


Pliolampas elevatus 


HEMIASTERIDAE 
Hemiaster_cf. eupetalum 


Opissaster_sp. ea 


SPATANGIDAE 
Maretia planulata 
Maretia cordata 


Maretia bandaensis 
Maretia mojsvari 


108 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Java, Mio- Yule Is, Barrow Is, Australian Australian 
Pliocene Lower Middle ‘mainland’ ‘reef’ 
Pliocene Miocene species species 


PALAEOSTOMATIDAE 
Palaeostoma kairukuensis 


BRISSIDAE 

Metalia spatagus 
Metalia sternalis 
Brissus latecarinatus 
Eupatagus (Eupatagus) 
pulchellus 

Eupatagus affinis 
Eupatagus sp. 


LOVENIIDAE 
Breynia_aff. carinata 
Breynia australasiae 
‘Breynia paucituberculata 
Breynia sp.a 

Breynia sp. b 


Lovenia elongata 


SCHIZASTERIDAE 
Schizaster (Schizaster) 
lacunosus 

Schizaster (Schizaster) 
alphonsei 

Schizaster (Schizaster) 
compactus 

Schizaster (Schizaster) aff. 
compactus 

Schizaster (Schizaster) sp. A 
Schizaster subrhomboidalis 
Schizaster progoensis 
Schizaster cf. pratti 
Schizaster excavatus 
Schizaster jeanneti 
Schizaster_ sp. 1 
Schizaster sp. 2 
Schizaster_sp. 3. 
Schizaster sp. _ 
Proraster jukesii 

Moira lethe 

Hemifaorina tuber 


mK LK [KL LK LK LK LK LX [xX 


Proc. Linn. Soc. N.S.W., 125, 2004 109 


ECHINOIDS OF NEW GUINEA AND TROPICAL AUSTRALIA 


ERRATA 


The following figures are replacements for: 


Figure 2 (page 128) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule 
Island, Papua New Guinea: Clypeasteroida. Proceedings of the Linnean Society of New South Wales 124, 
125-136. 


Figure | (page 155) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule 
Island, Papua New Guinea: Spatangoida. Proceedings of the Linnean Society of New South Wales 124, 153- 
162. 


110 Proc. Linn. Soc. N.S.W., 125, 2004 


EDFEINDEBY. 


Replacement Figure 2 (page 128) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower 


Pliocene), Yule Island, Papua New Guinea: Clypeasteroida. Proceedings of the Linnean Society of New 
South Wales 124, 125-136. 


Proc. Linn. Soc. N.S.W., 125, 2004 111 


ECHINOIDS OF NEW GUIN 


wold) nomimey (eho ints ene «eoney ALL alba { 
wa \o vision? nosed sii {a onan. aaloeame nei ii oH 


om 


I.D. LINDLEY 


Replacement Figure 1 (page 155) of Lindley, I.D. (2003). Echinoids of the Kairuku Formation (Lower 


Pliocene), Yule Island, Papua New Guinea: Spatangoida. Proceedings of the Linnean Society of New South 
Wales 124, 153-162. 


Proc. Linn. Soc. N.S.W., 125, 2004 113 


PURE 


PE 


Some living and fossil echinoderms from the Bismarck 
Archipelago, Papua New Guinea, and two new echinoid species 


1D. LINDLEY 


Department of Geology, Australian National University, Canberra, A.C.T. 0200. 
(lindley @ geology.anu.edu.au) 


Lindley, I.D. (2004). Some living and fossil echinoderms from the Bismarck Archipelago, Papua New 
Guinea, and two new echinoid species. Proceedings of the Linnean Society of New South Wales 125, 


115-139. 


Starfish and sea-urchin records of the Bismarck Archipelago, Papua New Guinea, are scattered throughout 
the literature of the past 160 years. This paper lists the region’s valid starfish and sea-urchin species records 
contained in the literature. In addition, records of 17 species of starfish and sea-urchins from material in the 
Department of Geology, Australian National University and the East New Britain Historical and Cultural 
Centre collections are included, with descriptions of two new sea-urchin species, the schizasterid Schizaster 
(Paraster) ovatus sp. nov. and the echinometrid Heliocidaris robertsi sp. nov. Some Tertiary echinoids 
from the region are described for the first time, namely Stereocidaris cf. squamosa Mortensen 1928 (Lower- 
Middle Miocene: Manus Island), Stereocidaris sp. (Pliocene: east New Britain), Phyllacanthus sp. (Pliocene: 
east New Britain) and Echinoneus sp. (Pleistocene-Holocene: Tanga Group, New Ireland). 


Manuscript received 22 August 2003, accepted for publication 22 October 2003. 


KEYWORDS: Asteroidea, Bismarck Archipelago, East Indies, Echinoidea, Extant, Fossil, Papua New 


Guinea, West Pacific. 


INTRODUCTION 


The Bismarck Archipelago, northern Papua 
New Guinea (PNG), encompasses the islands of New 
Britain, Bougainville, New Ireland and adjacent groups 
(Tabar, Lihir, Tanga and Feni), St. Matthias Group, 
the Admiralty Group, including Manus Island, and the 
surrounding waters of the Bismarck Sea (Fig. 1). It 
lies along the easternmost boundary of the East Indian 
Faunal Province. To the east and southeast lies the West 
Pacific Ocean or Melanesia faunal province (Endean 
1957 and A.M. Clark and Rowe 1971, respectively). 

Knowledge of the extant starfishes and sea- 
urchins (Echinodermata: Asteroidea and Echinoidea, 
respectively) from the Bismarck Archipelago 
comprises records scattered throughout a diverse 
literature of the past 160 years. The earliest described 
asteroid is Echinaster eridanella Miller and Troschel 
1842 (= Echinaster luzonicus Gray 1840) with a type 
locality in New Ireland. Sladen (1889) and A. Agassiz 
(1879 1881) described the asteroids and echinoids, 
respectively, collected during the 1873-76 voyage of 
H.M.S. Challenger. This expedition passed through the 
Admiralty Group and retrieved two new deep-water 
echinoids in the Bismarck Sea (the arbaciid 


Pygmaeocidaris prionigera (A. Agassiz 1879) and the 
temnopleurid Prionoechinus sagittiger A. Agassiz 
1879), at a site between the Admiralty Group and New 
Guinea. Loriol (1891) described additional asteroids 
from the archipelago, including Nardoa finschi de 
Loriol 1891 (= Nardoa tuberculata Gray 1840) and 
Nardoa mollis de Loriol 1891 (= Nardoa 
novaecaledoniae Perrier 1875), both with type 
localities in New Britain. Bell (1899) described the 
non-holothurian echinoderms collected by Arthur 
Willey during his 1895-97 visit to New Britain and 
the Loyalty Islands (Willey 1902). H.L. Clark (1925) 
redescribed several of Willey’s Bismarck Archipelago 
echinoids and erected two new species (the arbaciid 
Coelopleurus elegans (Bell 1899) and the diadematid 
Micropyga nigra H.L. Clark 1925) with type localities 
in New Britain. H.L. Clark (1946) and A.H. Clark 
(1954) recorded additional asteroids and echinoids 
from the archipelago. Struder (1876, 1880) and H.L. 
Clark (1908) provided descriptions of the extant 
echinoderm fauna of west New Guinea, a region 
contiguous with the Bismarck Archipelago. 

During the past 20 years echinoderm research 
in the region has concentrated on the biology of 
asteroids and comatulid crinoids at Hansa Bay and 
Madang, on the southern shores of the Bismarck Sea 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Saonek 
Sorong 


Admiralty Gp 
(Manus Is) 
= 


ARAFURA 


SEA 
Torres ,, 


Strait f 
" 


Darwin 


AUSTRALIA 


New Ireland 
@ Tanga Gp 
Rabaul/Blanche Bay/Cape Gazelle 


ii} | 
nae Is 
New Britain | 


Solomon 
Islands 


CORAL SEA 


x Swain Reefs 
.? 
\ Capricorn- 
*, Bunker Groups 


Figure 1. Locality map showing the Bismarck Archipelago, Papua New Guinea, and other localities 


discussed in text. 


(Britayev et al. 1999; Bouillon and Jangoux 1984; 
Eeckhaut et al. 1996; Messing 1994). 

This paper describes some extant asteroids and 
echinoids from the Bismarck Archipelago. It also 
provides for the first time, a tabulation of previously 
reported asteroid and echinoid occurrences (Tables 1 
and 2 respectively; tables located after the reference 
list) in the region. Several Tertiary echinoids from the 
archipelago are also described. The author is not aware 
of any previous description of the region’s fossil 
echinoderm fauna. 

The specimens described in this paper were 
collected between 1981-2003, and do not necessarily 
represent the results of thorough, methodical site 
collections. The Cape Gazelle, east New Britain, 


116 


locality encompasses any one of a number of nearby 
localities, including Tovarur Plantation, Reiven Beach 
and southeastern Tokua Airport. Full systematic 
descriptions are provided for all fossil species while 
in most cases, only brief remarks concerning the 
significance of occurrences are provided for extant 
species. Specimens prefixed ANU are housed in the 
Department of Geology, The Australian National 
University; specimens prefixed B are housed in the 
East New Britain Historical and Cultural Centre, 
Kokopo, East New Britain Province, PNG. 
Terminology and classification used herein follows that 
of the Treatise on Invertebrate Paleontology and A.M. 
Clark and Rowe (1971). 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


SYSTEMATIC DESCRIPTIONS 


Class STELLEROIDEA Lamarck 1816 
Subclass ASTEROIDEA de Blainville 1830 
Order VALVATIDA Perrier 1884 

Suborder GRANULOSINA Perrier 1894 
Family OPHIDIASTERIDAE Verrill 1867 
Genus LINCKIA Nardo 1834 


Synonymy 
Cribella Agassiz 1835 (non Forbes 1841). 
Acalia Gray 1840. 
Catantes, Undina Gistl 1847. 


Type species 
Linkia typus Nardo 1834 (= Asterias laevigatus Linnaeus 1758) by original designation. 


Linckia multifora (Lamarck 1816) 


Synonymy 
Asterias multifora Lamarck 1816, p. 565. 
Linckia leachi Gray 1840, p. 285: Mauritius. 
Linckia costae Russo 1894, p. 163: Daret Is., Red Sea. 


Materials and locality 
Two specimens, ANU 60651-2, collected at Ralum, Blanche Bay, East New Britain Province, PNG, 


Remarks 
Linckia multifora (Lamarck 1816) is widely distributed throughout the Indo-Pacific, from the Red Sea to 
the Hawaiian Islands (A.M. Clark and Rowe 1971). This record is the first from New Guinea. 


Family OREASTERIDAE 
Genus PROTOREASTER Doderlein 1916 


Type species 
Asterias nodosa Linnaeus 1758, p. 420, by subsequent designation. 


Protoreaster nodosus (Linnaeus 1758) 


Synonymy 
Oreaster nodosus, Bell 1884, p. 70; H.L. Clark 1908, p. 280; Fisher 1911, p. 346; H.L. Clark 1921, p. 
31. 
Pentaceros nodosus, Bell 1899, p. 136. 
Protoreaster nodosus, Doderlein 1916, p. 420; H.L. Clark 1946, p. 106; A.M. Clark and Rowe 1971, 
p. 34, 54; Rowe and Gates 1995, p. 106. 


Material and locality 
Single beach worn specimen, ANU 60650, collected at Ralum, Blanche Bay, East New Britain 
Province, PNG. 


Remarks 


Protoreaster nodosus (Linnaeus 1758) is a common East Indian starfish with a range extending to the 
West Pacific (Caroline Islands) (H.L. Clark 1946; A.M. Clark and Rowe 1971). Bell (1899) previously described 


Proc. Linn. Soc. N.S.W., 125, 2004 117 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


the species from the collections of Arthur Willey in Blanche Bay. H.L. Clark (1908, p. 280) described variations 
in specimens from several west New Guinea localities (Humboldt Bay, Sorong, Ansus, Jappen Island). Although 
the present specimen (R/r = 20/9 mm) has lost most of its granules and abactinal plates, it is identified as a 
juvenile P. nodosus (L.M. Marsh, pers. comm.). It is similar to a specimen of P. nodosus in the Western Australian 
Museum (WAM 599-76: R/r = 27/11 mm) collected by L.M. Marsh from Pulau Langkai, off south Sulawesi, 
Indonesia. Two juvenile specimens (R = 11-12 mm) from the Andaman Islands, figured and described by 
Koehler (1910: plate XVI, fig. 1) as Anthenea sp., are also very similar to ANU 60650. These specimens may 


also be P. nodosus (L.M. Marsh, pers. comm.). 


Class ECHINOIDEA Leske 1778 

Subclass PERISCHOECHINOIDEA M’Coy 1849 
Order CIDAROIDA Claus 1880 

Family CIDARIDAE Gray 1825 

Subfamily STEREOCIDARINAE Lambert 1900 
Genus STEREOCIDARIS Pomel 1883 


Synonymy 
Typocidaris Pomel 1883 
Phalacrocidaris Lambert 1902 
Anomocidaris Agassiz and Clark 1907 


Type species 


Cidaris cretosa Mantell 1835; subsequent designation Lambert and Thiéry 1909 (Feb., p. 31; non 


Mar., where C. merceyi was designated, p. 152). 


Stereocidaris cf. squamosa Mortensen 1928 


Figs 2, 3a 


Synonymy 
Stereocidaris indica Bell 1909, p. 21; H.L. Clark 1925, p. 26. 


Stereocidaris squamosa Mortensen 1928b, p. 70; Mortensen 1928a, p. 245. 


Figure 2. Stereocidaris cf. squamosa Mortensen 1928. Lower-Middle 
Miocene, Manus Island, Manus Province. 2a-b, plating diagrams at 
ambitus for ambulacrum, interambulacrum. Abbreviations: btr basal 
terrace; it inner tubercle; m mamelon; mt marginal tubercle; r ridge; 
ser scrobicular ring; w wall. 


118 


Material 

An incomplete specimen 
ANU 60638, with an 
interambulacral plate and portion 
of ambulacral series. 


Locality and horizon 

Village of Drankei, west 
bank of Wari River, central 
southern Manus Island, Manus 
Province, PNG. Grid reference 
060612 Lorengau 1: 100 000 Sheet 
8393 (Edition 1). The collection 
horizon is an outlier of the (lower) 
Mundrau Limestone. A sample of 
limestone from a nearby outlier of 
the Mundrau Limestone at 
Metawarei village, 0.5 km 
northwest of Drankei village, 
contained a foraminiferal 
assemblage of mid or upper Tfl 
age, and suggests a late Lower 


Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Figure 3. Tertiary echinoids from the Bismarck Archipelago. Stereocidaris cf. sgquamosa Mortensen 1928. 
Lower-Middle Miocene, Manus Island, Manus Province. 3a, ANU 60638, incomplete interambulacral 
plate with large tubercle and part of adjacent ambulacral plating (refer to Figs 2a, b for plating diagram). 
Bar scale = 2.5 mm. Stereocidaris sp. Pliocene, Sikut River area, East New Britain Province. 3b, ANU 
60639, primary spine. Bar scale = 2.5 mm. Phyllacanthus sp. Pliocene, Mevelo River area, East New 
Britain Province. 3c, ANU 60637, proximal portion of primary spine. Bar scale = 5 mm. Echinoneus sp. 
Pleistocene-Holocene, Boang Island, Tanga Group, New Ireland Province. 3d, ANU 60640, aboral view 
of worn specimen. Bar scale = 5 mm. 


Miocene or earliest Middle Miocene age for the unit (Francis 1985). 


Description 

Test size and shape unknown. 

Ambulacra sinuate, rather broad, ca. 39% of width of interambulacra. Interporiferous zone about twice 
width of a pore-zone. Interporiferous zone with distinctly vertical series of marginal tubercles and inner tubercles; 
marginal tubercles slightly larger than those of inner series. Pores are rather small, circular, nonconjugate and 
separated by a broad wall; ridge low and narrow. The arrangement of pores and tubercles in ANU 60638 is 
strikingly similar to that described for Stereocidaris squamosa Mortensen 1928 by Mortensen (1928a: 245 and 
Plate LXX, fig. 7). 

Interambulacral plate higher than broad (height:width = 8.0:5.5) with aureole of moderate size but not 
very deep, well separated. The very high plate of ANU 60638 exhibits a broad miliary covered space between 


Proc. Linn. Soc. N.S.W., 125, 2004 119 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


successive aureoles, one unequal to the other, indicating it to be from an upper interambulacral, or aboral, 
position on test (Fig. 2). Mamelon apparently large, the part of plate is damaged in ANU 60638. Edge of 
aureole is not raised, and the scrobicular tubercles are not prominent. Outside the scrobicular ring, the 
interambulacral plate has a sparse to moderate covering of tubercles of similar size to scrobicular tubercles. On 
the adradial edge of plate there are a few secondary tubercles outside the scrobicular ring. 

Apical and periproctal systems unknown. 

Details of primary and secondary spines unknown. 


Remarks 

Stereocidaris Pomel 1883 is a well characterised genus with both fossil and extant species (Mortensen 
1928a; Chapman and Cudmore 1934). The genus is distinctive for its usually very high interambulacral plates, 
ambulacra that are generally conspicuously sinuate and nonconjugate pores (Mortensen 1928a; Fell 1966). The 
oldest occurrence of the genus is from the Cretaceous of Europe. In the Tertiary it is known from the Eocene of 
Europe and Australia, Oligocene of New Zealand, Miocene of Australia and Indonesia, and Pliocene of Australia 
and New Zealand (Mortensen 1928a; Chapman and Cudmore 1934; Fell 1966). Mortensen (1928a) noted the 
lack of fossil Stereocidaris from the Indo-Pacific, with K. Martin’s (1918) record of the occurrence of a spine 
of Dorocidaris papillata (= Stereocidaris) from the Miocene of Java the only known fossil. This may well 
represent a collection bias. Extant species of Stereocidaris, numbering 15, with nine subspecies, are distributed 
throughout the Indo-Pacific, including southeast Africa (Mortensen 1928a). Notably the genus has not been 
recorded from Australasian seas (H.L. Clark 1925; Fell 1966). 

The Manus Island specimen, represented by a small fragment of plating from an adapical position, is 
tentatively assigned to Stereocidaris squamosa Mortensen 1928. As already noted, the striking resemblance of 
the ambulacral and interambulacral plating of this single specimen to that described by Mortensen (1928a) for 
S. squamosa cannot be ignored. Stereocidaris squamosa is an extant species recorded from 270 m depth on the 
Saya de Malha Bank (10° 30’S), about 800 km southeast of the Seychelles in the Indian Ocean (Mortensen 
1928a). The species has a small-moderate sized test that ranges in diameter from 30-47 mm, with height from 
18.5-29 mm (Mortensen 1928a). Longest spines range in length from ca. 50-59 mm. The late Lower Miocene/ 
earliest Middle Miocene Manus Island occurrence represents the first fossil occurrence of test remains of 
Stereocidaris from the Indo-Pacific. 


Stereocidaris sp. 
Fig. 3b 


Material 
An isolated fragmentary spine, ANU 60639. 


Locality and horizon 

Collected ‘5 km east of the intake structure of the Warangoi hydro-scheme’ (Lindley unpubl. field 
notes), in the headwaters of Matuli Creek, a tributary of the Warangoi River, Sikut area, northeastern Gazelle 
Peninsula, East New Britain Province, PNG. Grid reference 081961 Merai 1: 100 000 Sheet 9388 (Edition 1). 
The collection horizon is from the Sinewit Formation, of Mio-Pliocene age (Lindley 1988). However, fossil 
evidence and a K-Ar radiometric age from the Sikut and adjacent areas, indicates the formation in this area is 
restricted to the Pliocene (Read 1965; B. McGowran in Lindley 1988; Lindley 1988; Corbett et al. 1991). 


Description 

No test fragments which belong to this species have been identified. 

Primary spine cylindrical, distinctly fusiform, tapering, point not widened. Spine length 15 mm, with 
maximum diameter of 2 mm occurring about 1/2 distance from proximal end. The shaft with about 16 series of 
low rounded warts; only towards the point do they assume the shape of low rounded ridges. The collar is only 
0.75-1 mm long, slightly increasing in thickness towards inconspicuous milled ring. Neck is equal in length to 
collar. 


Remarks 
The primary spines of cidaroids possess a distinctive structure with a compact outer or cortex layer 


120 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


covering all except the collar and enveloping a central core consisting of an irregular calcareous meshwork 
(Mortensen 1928a). The cortex layer is found only in a few other echinoids, mainly the salenids, and spinule 
and wart ornament along the shaft is formed by this alone (Mortensen 1928a). Mortensen (1928a: 50) considered 
that primary spine shape and structure is of considerable use in cidaroid classification, both at specific and 
generic levels. 

The primary spine ANU 606339 is identified as that of a cidaroid by its spine shape and its possession of 
an outer cortex layering. The nature of the inner central core meshwork is clearly visible on the spine collar. The 
nature of wart development, the number of longitudinal series, and their distal transition to low rounded ridges, 
bears a strong resemblance to that seen in the primary spines of some extant species of Stereocidaris, including 
Stereocidaris grandis (Déderlein) and Stereocidaris hawaiiensis Mortensen 1928b, found only in Japanese 
seas and Hawaiian seas, respectively (cf. Mortensen 1928a: Plate XIX, fig. 5 and XXI, fig. 5, respectively). 


Subfamily RHABDOCIDARINAE Lambert 1900, emended Fell 1966 
Genus PHYLLACANTHUS Brandt 1835 


Synonymy 
Leiocidaris Desor 1885, p. 48. 


Type species 
Cidarites (Phyllacanthus) dubia Brandt 1835, p. 67, by original designation. 


Phyllacanthus sp. 
Fig. 3c 


Material 
One isolated fragmentary spine, ANU 60637. 


Locality and horizon 

Collected in stream float from an unnamed large western tributary of Mevelo River, Lakit Range, 
southwestern Gazelle Peninsula, East New Britain Province, PNG. Grid reference 660623 Pondo 1:100 000 
Sheet 9288 (Edition 1). Lakit Limestone, Pliocene (Lindley 1988). 


Description 

No test fragments which belong to this species have been identified. 

Proximal portion of primary spine moderately thick, cylindrical, fusiform, with a maximum diameter of 
8.0 mm. Details of distal shaft unknown. Details of spine base, milled ring and collar unknown. Spine swells 
rapidly above the collar. Surface of shaft is finely and uniformly granulated (not visible to the naked eye), the 
granules forming numerous (> 50) longitudinal series along length of spine. 


Remarks 

Lindley (2003b) described the spines of Phyllacanthus imperialis var. javana K. Martin 1885 and 
Phyllacanthus sp. from the Lower Pliocene Kairuku Formation, Yule Island. Unfortunately, the characters 
diagnostic of these species, including spine collar length and the number of ridges on the distal part of the spine, 
are not visible on ANU 60637. 


Subclass EUECHINOIDEA Bronn 1860 
Superorder ECHINACEA Claus 1876 

Order TEMNOPLEUROIDA Mortensen 1942 
Family TOXOPNEUSTIDAE Troschel 1872 
Genus TOXOPNEUSTES A. Agassiz 1841 


Synonymy 
Boletia Desor 1846, p. 362. 


Proc. Linn. Soc. N.S.W., 125, 2004 121 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Type species 
Echinus pileolus Lamarck 1816, p. 45, by original designation. 


Toxopneustes pileolus (Lamarck 1816) 


Synonymy 
Echinus pileolus Lamarck 1816, p. 45. 
Toxopneustes pileolus, A. Agassiz 1841, p. 7; H.L. Clark 1925, p. 123; Mortensen 1943a, p. 472; 
A.M. Clark and Rowe 1971, p. 156; Rowe and Gates 1995, p. 258. 
Mortensen (1943a: 472) lists additional synonymies. 


Material and locality 
Single naked test, B20022, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 


Remarks 
Toxopneustes pileolus (Lamarck 1816) is widely distributed throughout the Indo-West Pacific 
(Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). 


Genus TRIPNEUSTES L. Agassiz 1841 


Type species 
Echinus granularis Lamarck 1816, p. 44, by original designation. 


Tripneustes gratilla (Linnaeus 1758) 


Synonymy 
Echinus gratilla Linnaeus 1758, p. 664. 
Tripneustes gratilla, H.L. Clark 1925, p. 124; Mortensen 1943a, p. 500; A.M. Clark and Rowe 1971, 
p. 156; Rowe and Gates 1995, p. 259. 
Mortensen (1943a: 500) lists additional synonymies. 


Material and locality 
Single naked test, B20023, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 


Remarks 

Tripneustes gratilla (Linnaeus 1758) is widely distributed throughout the Indo-West Pacific (Mortensen 
1943a; A.M. Clark and Rowe 1971). Previous records from the Pacific include the Marshall Islands, Norfolk 
Island, Hawaiian Islands, Kermadec Islands, Solomon Islands, Fiji and Hood Lagoon, south coast of Papua 
(H.L. Clark 1925; Mortensen 1943a; A.M. Clark and Rowe 1971; Miskelly 2002). 


Order ECHINOIDA Claus 1876 
Family ECHINOMETRIDAE Gray 1825 
Genus ECHINOMETRA Gray 1825 


Synonymy 
Ellipsechinus Liitken 1864, p. 165. 
Plagiechinus Pomel 1883, p. 78. 
Mortensenia Déderlein 1906, p. 233. 


Type species 
Echinus lucunter Linnaeus 1758, p. 665, by original designation. 


122 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Echinometra mathaei (de Blainville 1825) 


Synonymy 
Echinus lucunter Lamarck 1816, p. 50 (non E. lucunter Linnaeus). 
Echinometra mathaei, H.L. Clark 1925, p. 143; H.L. Clark 1932, p. 216; Mortensen 1943b, p. 381; 
H.L. Clark 1946, p. 332; A.M. Clark and Rowe 1971, p. 157; Rowe and Gates 1995, p. 211. 
Mortensen (1943b: 381) lists additional synonymies. 


Material and localities 

Fourteen naked tests from Gargaris village, northern coast of Malendok Island, Tanga Group, New 
Ireland Province, PNG; one partly naked test from beach at Ralum, Blanche Bay, East New Britain Province, 
PNG; one naked test from Penlolo village, south coast of New Britain, West New Britain Province, PNG; one 
naked test, B 20016, from Cape Gazelle, New Britain, East New Britain Province, PNG. 


Remarks 

Echinometra mathaei (de Blainville 1825) is a long ranging species, recorded from late Lower Miocene- 
early Middle Miocene rocks in the western and eastern Mediterranean Sea (Negretti et al. 1990). Extant E. 
mathaei is one of the most widely distributed echinoids, occurring throughout tropical-subtropical waters of the 
Indo-West Pacific (Mortensen 1943b; A.M. Clark and Rowe 1971). H.L. Clark (1908) recorded the species 
from Sorong, west New Guinea and Miskelly (2002) recorded it from the Solomon Islands. This record indicates 
a wide distribution throughout the Bismarck Archipelago (Tanga Group, New Ireland; Blanche Bay, New 
Britain; and south coast New Britain). 


Genus HETEROCENTROTUS Brandt 1835 


Synonymy 
Acroladia L. Agassiz and Desor 1846, p. 373. 


Type species 
Echinus mamillatus Linnaeus 1758, p. 664, by subsequent designation of Pomel 1883, p. 77. 


Heterocentrotus mammillatus (Linnaeus 1758) 


Synonymy 
Echinus mamillatus Linnaeus 1758, p. 664. 
Heterocentrotus mammillatus, H.L. Clark 1925, p. 147; Mortensen 1943b, p. 409; H.L. Clark 1946, p. 
333; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 213. 
Mortensen (1943b: 409) lists additional synonymies. 


Material and locality 

A single naked test, B 20017, and unlabelled isolated spines (housed in the East New Britain Historical 
and Cultural Centre, Kokopo) from Cape Gazelle, New Britain, East New Britain Province, PNG; an isolated 
primary spine, ANU 60648, from Nosnos village, Boang Island, Tanga Group, New Ireland Provine, PNG. 


Remarks 

Heterocentrotus mammillatus (Linnaeus 1758) is widely distributed throughout the Indo-Pacific, from 
the Gulf of Suez and Madagascar to the Hawaiian Islands and Fiji (Mortensen 1943b). It is recorded from the 
Solomon Islands by Miskelly (2002). The largest test of H. mammillatus noted by Mortensen (1943b) has a 
long diameter of 82 mm, with most individuals having diameters of 72 mm or less. The long diameter of the 
Cape Gazelle test is 72 mm. The Tanga spine has a length of 74 mm and, given that the primary spines of H. 
mammillatus usually do not exceed the long diameter of the test (Mortensen 1943b), appears to have come from 
a relatively large individual. 


Proc. Linn. Soc. N.S.W., 125, 2004 123 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Genus HELIOCIDARIS L. Agassiz and Desor 1846. 


Synonymy 
Toxocidaris A. Agassiz 1863, p. 22. 


Type species 
Echinus tuberculatus Lamarck 1816, p. 50, by original designation. 


Diagnosis 

Low hemispherical echinoids, widest at circular ambitus. Ambulacral plates with 7 or more pore-pairs to 
each plate; arcs may be irregularly double; expanded poriferous tracts of the flattened adoral surface are petaloid. 
Oculars I and IV usually insert. Gill-slits are shallow (Philip 1965; Fell and Pawson 1966). 


Remarks 

Heliocidaris L. Agassiz and Desor 1846 is distributed along the southern coasts of Australia, northern 
New Zealand, Kermadec Islands and Lord Howe Island (Mortensen 1943a). Two species are included in the 
genus by Mortensen (1943a), viz: Heliocidaris tuberculata (Lamarck 1816) and Heliocidaris erythrogramma 
(Valenciennes 1846) and, given their similar morphologies, he has questioned whether they are really 
conspecific. Anthocidaris Liitken 1864 is a closely allied genus (only known species Anthocidaris 
crassispina [A. Agassiz 1863]) from the coasts of southern Japan and China, distinguished from Heliocidaris 
by the spicules of the tubefeet (Mortensen 1943a). On the status of Anthocidaris, Mortensen (1943a: 328) 
questioned whether the genus should be merged into Heliocidaris. Philip (1965) described the only known 
fossil representative of the genus, Heliocidaris ludbrookae Philip 1965 from the Lower-early Middle 
Miocene (Longfordian-Batesfordian) of southeastern Australia. 


Heliocidaris robertsi sp. nov. 
Figs 4, 5a-e 


Diagnosis 

Test low hemispherical, somewhat inflated above. Ambulacral plates with 12 pore-pairs per plate; ambital 
and aboral pore-arcs doubled. Ambulacral and interambulacral plates relatively large; each bearing a primary 
tubercle and numerous secondary tubercles; aureoles of primaries not in contact. Primary tubercles of ambital 
and aboral ambulacral plates with an aborally positioned secondary tubercle. 


Etymology 
Named for Mr Michael Roberts, amateur conchologist of Kokopo, East New Britain Province, PNG. 


Material and locality 

Single naked test, ANU 
60654, from the vicinity of 
Cape Gazelle, New Britain, 
East New Britain Province, 
PNG. 


Description 

Test low hemispherical, 
somewhat inflated above, 
widest at circular ambitus. The 
oral side is flattened, scarcely 
sunken towards the peristome. 


Only specimen of 38 mm 
Figure 4. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain —gjameter. 


Province. 4a-b, plating diagrams at ambitus for interambulacrum, The pore zones are 
ambulacrum. conspicuously petaloid on the 


124 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Figure 5. Heliocidaris robertsi sp. nov. Cape Gazelle area, East New Britain Province. 5a-e, ANU 60654, 
aboral, oral, lateral views. Bar scale = 10 mm; ambulacral plating at ambitus (refer to Fig. 4b for plating 
diagram). Bar scale = 5 mm; apical disc. Bar scale = 2.5 mm. 


Proc. Linn. Soc. N.S.W., 125, 2004 125 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


oral surface, about 1.5-2 times the width of interporiferous zone. The pore-series in this area are almost horizontal 
and are separated by secondary tubercles forming a single prominent vertical series; scattered miliary tubercles 
are also present. In the ambital region there are 12 pore-pairs arranged in double arcs (Fig. 4). Above the 
ambitus the pore-zones become much narrower. Primary tubercles in the ambital zone are large, almost as large 
as the interambulacral primaries; aureoles of adjacent primaries in each vertical series widely separated. Sutures 
between adjacent plates are seen very distinctly on the outer adoral side of the boss. Each ambulacral plate at 
and above the ambitus has a prominent secondary tubercle positioned aborally to the primary tubercle; 4-5 
other secondary tubercles are also present. Miliaries tend to be arranged along the perradial sutures of ambital 
and superoambital ambulacrals; elsewhere on each plate only a sparse covering of miliaries is present. 

The interambulacral primaries are large, forming prominent series aborally; their aureoles are distinctly 
separated, leaving a broad space at the upper edge of each plate, occupied by several small tubercles and 
miliaries. Usually sutures between adjacent plates are close to, but do not cross, aureole of successive tubercle. 
In the median space there is on the oral surface and in the ambital region a conspicuous double series of 
secondary tubercles about half the size of the ambulacral primaries. Below the ambitus all the tubercles decrease 
rapidly in size, with the secondaries disappearing, and only the primaries continuing to the peristome. 

The apical system is small, only about 18 percent of the test diameter. There is typically one large 
tubercle on each genital plate, except the very large madreporite, and a scattering of small tubercles over the 
remainder (Fig. 5e). Ocular I and IV are broadly insert. The peristome is very small, about 29 percent of test 
diameter. Gill-slits shallow. 

Details of spines and pedicellarie unknown. 


Remarks 

Heliocidaris robertsi sp. nov. is readily distinguished from H. tuberculata and H. erythrogramma and 
the closely allied A. crassispina by its possession of double pore-arcs on the adoral surface. The double pore- 
arcs of H. robertsi are very similar to those of Heterocentrotus trigonarius (Lamarck 1816), figured by Mortensen 
(1943a: fig. 132c) and Fell and Pawson (1966: fig. 324, 7c). However, any resemblance between the new 
species and H. trigonarius is easily discounted because of the latter’s possession of a distinctly elongated test 
and a significantly larger peristome (51 percent of test diameter). 

The biogeographical position of H. robertsi is noteworthy in that it is the tropical representative of two 
closely allied temperate water genera, Heliocidaris, a very common form restricted to southern Australia and 
New Zealand, and Anthocidaris, an equally common form restricted to Japan and China. 

Pore-arc doubling is almost as strongly developed in other echinometrids including Colobocentrotus 
Brandt 1835 and Zenocentrotus A.H. Clark 1931, and incipient development may also been seen in Echinometra 
Gray 1825 (Mortensen 1943a: 281). All three genera possess an elliptical or oblong ambitus. The functional 
significance of doubling of pore-arcs in compound plates relates to (a) increasing the area over which tube-feet 
are spread, and thereby increasing respiratory and feeding efficency (Mortensen 1943a; Woods 1958; Durham 
1966; A.M. Clark 1968) and (b) strengthening of the test (Durham 1966). The doubling of pore-arcs on the 
aboral surface of H. robertsi greatly increases the number of tube-feet in this area, not only aiding in improved 
respiration, but allowing it to catch food particles falling onto its upper surface. With such adaptations to its 
upper surface, the echinoid may have been a reef rock borer, inhabiting a hole perhaps several centimetres deep. 


Superorder GNATHOSTOMATA Zittel 1879 
Order HOLECTYPOIDA Duncan 1889 

Suborder ECHINONEINA HLL. Clark 1925 
Family ECHINONEIDAE Agassiz and Desor 1847 
Genus ECHINONEUS Leske 1778 


Synonymy 
Echinanaus Gray 1825, p. 7 (nom. van.). 
Pseudohaimea Pomel 1885, p. 118. 
Koehleraster Lambert and Thiéry 1921, p. 331. 


Type species 
Echinoneus cyclostomus Leske 1778, by subsequent designation of H.L. Clark 1917, p. 101. 


126 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Remarks 

Echinoneus Leske 1778 is an Oligocene-Recent form, with some ten fossil species described from the 
Oligocene and Miocene of Europe (Mortensen 1948a; Wagner and Durham 1966). Two Recent species are 
known, viz. Echinoneus cyclostomus Leske 1778 and Echinoneus abnormalis de Loriol 1883, distinguished by 
the presence or absence of imperforate primary tubercles and well developed glassy tubercles. Recent forms are 
distributed throughout the West Indies, Indo-Pacific and Australia. Mortensen (1948a) considered that many of 
the fossil species are very difficult to distinguish and may in fact be Recent E. cyclostomus. 


Echinoneus sp. 
Fig. 3d 


Material 
One poorly preserved test, ANU 60640. 


Locality and horizon 

Nosnos village, Boang Island, Tanga Group, New Ireland Province, PNG. Grid reference 296246 Tanga 
1:100 000 Sheet 9591 (Edition 1). Unnamed poorly compacted bioclastic limestone, Pleistocene-Holocene 
(Wallace et al. 1983). 


Description 

Test ovoid, moderate size, measuring 23 x 17 x 11.5 mm; oral surface weakly concave. Ambulacra 
narrow, not petaloid. Other details of ambulacra unknown. Details of interambulacra unknown. Details of 
tubercles unknown. Apical and periproctal systems unknown. 


Remarks 
The lack of well preserved tubercles on this specimen makes it difficult to assign a species. 


Kchinoneus cyclostomus Leske 1778 


Synonymy 
Echinoneus cyclostomus Leske 1778, p. 173; H.L. Clark 1925, p. 177; H.L. Clark 1946, p. 353; 
Mortensen 1948a, p. 75; A.M. Clark and Rowe 1971, p. 158; Rowe and Gates 1995, p. 215. 
Mortensen (1948a: 75) lists additional synonymies. 


Material and locality 

Twelve naked tests, including ANU 60641, from Gargaris village, northern coast of Malendok Island, 
Tanga Group, New Ireland Province, PNG; one naked test, B 20021, from Cape Gazelle, New Britain, East 
New Britain Province, PNG. 


Remarks 

Echinoneus cyclostomus Leske 1778 is the only known case of a (tropical) cosmopolitan echinoid, having 
been recorded from the West Indies, Ascension (but not the African west coast) and the Indo-Pacific-East 
Africa (Zanzibar, Natal), Madagascar to the Pacific islands (Funafuti, Palmyra, Hawaiian Islands), and from 
Japan to Queensland (Great Barrier Reef) and Lord Howe Island (Mortensen 1948a). Miskelly’s (2002) record 
of E. cyclostomus from the Solomon Islands represents the nearest previous record to that from the Tanga 
Group and Cape Gazelle. 


Echinoneus abnormalis de Loriol 1883 
Synonymy 
Echinoneus abnormalis de Loriol 1883, p. 41; H.L. Clark 1917, p. 102; H.L. Clark 1925, p. 176; 


Mortensen 1948a, p. 80; A.M. Clark and Rowe 1971, p. 158. 
Koehleraster abnormalis Lambert and Thiéry 1921, p. 331. 


Proc. Linn. Soc. N.S.W., 125, 2004 127 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Material and locality 
One naked test, ANU 60641, from Gargaris village, northern coast of Malendok Island, Tanga Group, 
New Ireland Province, PNG. 


Remarks 

This species is represented by a single naked test measuring 30 x 22.5 x 15 mm. Echinoneus abnormalis 
de Loriol 1883 is distinguished from E. cyclostomus by possessing perforated, non-glassy spine tubercles. The 
apical system of the Tanga specimen is distinctly anterior to that of co-occurring specimens of the much more 
common E. cyclostomus. Echinoneus abnormalis has a restricted distribution, known from Mauritius (type 
locality), Kei Islands, Palmyra Island, Banda, Ellice Islands and the Hawaiian Islands (Mortensen 1948a; A.M. 
Clark and Rowe 1971). The recent record of E. abnormalis from the vicinity of Raine Island on the northern 
Great Barrier Reef (Gibbs et al. 1976) represents the first from Australasian waters. The record from the Tanga 
Group is the second from the East Indies. The species is observed to be sympatric with the much more common 
E. cyclostomus in many localities, a fact Gibbs et al. (1976) suggested may have resulted in it having gone 
unrecognised in samples. Mortensen (1948a: 81) considered that the two species probably didn’t live together 
at the same localities. Of the 15 specimens of Echinoneus collected from the Malendok Island locality, only one 
was an E. abnormalis, suggesting that in this case, the species’ apparent rarity may be related to different niches 
within the same locality. 


Order CLYPEASTEROIDA A. Agassiz 1872 
Suborder CLYPEASTERINA A. Agassiz 1872 
Family CLYPEASTERIDAE L. Agassiz 1835 
Genus CLYPEASTER Lamarck 1801 


Type species 
Clypeaster rosaceus (Linnaeus 1758), by subsequent designation of Desmoulins 1835. 


Clypeaster reticulatus (Linnaeus 1758) 


Synonymy 
Lindley (2003a) lists previous synonymies. 


Material 
Single naked test, B20020, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 


Remarks 

Clarification of Lindley’s (2003a) statement on the distribution of Clypeaster reticulatus (Linnaeus 1758) 
is needed. The species is a very common Indo-West Pacific echinoid, distributed in the western Indian Ocean 
and the Red Sea, throughout the East Indies and east into the Pacific Ocean to the Hawaiian Islands (A.M. Clark 
and Rowe 1971). Previous south Pacific records of the species have been made by A. Agassiz (1863), Mortensen 
(1948b) and A.H. Clark (1954) from the Gilbert Islands, New Caledonia and Marshall Islands, respectively. 
Mortensen’s (1948b) New Caledonian record has not been confirmed by De Ridder (1986: 29). McNamara and 
Kendrick (1994) have also recorded the species from Barrow Island, northwestern Australia. The species is 
known from fossil in Java (Lower Miocene), Yule Island, PNG (Lower Pliocene), East Africa (Pliocene- 
Pleistocene) and the New Hebrides (Pleistocene) (Mortensen 1948b; Lindley 2003a). 


Family ARACHNOIDAE Duncan 1889 
Subfamily ARACHNOIDINAE Duncan 1889 
Genus ARACHNOIDES Leske 1778 


Synonymy 
Echinarchinus Leske 1778, p. 217. 


128 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Type species 
Echinus placenta Linnaeus 1758, p. 666, ICZN 1954. 


Arachnoides placenta (Linnaeus 1758) 


Synonymy 
Echinus placenta Linnaeus 1758, p. 666. 
Arachnoides placenta (Linnaeus 1758): L. Agassiz 1841, p. 94; Bell 1899, p. 136; H.L. Clark 1925, p. 
154; H.L. Clark 1946, p. 340; A.M. Clark and Rowe 1971, p. 161; Rowe and Gates 1995, p. 
176. 
Mortensen (1948b) lists additional synonymies. 


Material and locality 
Single naked test, B20018, from the vicinity of Cape Gazelle, New Britain, East New Britain 
Province, PNG. 


Remarks 

Arachnoides placenta (Linnaeus 1758) is a common littoral species throughout the East Indies and the 
south Pacific (Mortensen 1948b; A.M. Clark and Rowe 1971). The first record of the species from the Bismarck 
Archipelago is that of Bell (1899) from an unspecified locality in New Britain. 


Suborder LAGANINA Mortensen 1948 
Family LAGANIDAE A. Agassiz 1873 
Genus LAGANUM Link 1807 


Synonymy 
Lagana Gray 1825, p. 427. 


Type species 
Laganum petalodes (= Echinodiscus laganum Leske 1778, p. 204), by original designation. 


Laganum laganum (Leske 1778) 


Synonymy 
Laganum Bonani Klein 1734, p. 25. 
Echinodiscus laganum Leske 1778, p. 204. 
Laganum laganum, Mortensen 1948b, p. 312. 
Laganum depressum, Lindley 2001, p. 130. 
Mortensen (1948b: 312) list previous synonymies. 


Material and locality 
Single test, ANU 60649, from Penlolo village, south coast of New Britain, West New Britain 
Province, PNG. 


Remarks 

Laganum laganum (Leske 1778) is distinct with its pentagonal test with thick, swollen edges, and an 
oblong-elongate periproct situated midway between the mouth and test edge. The species is common in the East 
Indies, and is also recorded from Port Jackson and Tasmania (Mortensen 1948b). Mortensen (1948b) also 
recorded it from the Bismarck Archipelago (Table 2). H.L. Clark (1908) recorded the species from Saonek, 
Waigiou Island, in west New Guinea (Fig. 1) 


Suborder SCUTELLINA Haeckel 1896 
Family ASTRICLYPEIDAE Stefanini 1911 


Proc. Linn. Soc. N.S.W., 125, 2004 129 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Genus ECHINODISCUS Leske 1778 


Type species 
Echinodiscus bisperforatus Leske 1778, p. 196. 


Echinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) 


Synonymy 
Lobophora tenuissima L. Agassiz and Desor 1847, p. 136. 
Echinodiscus tenuissimus, Gray 1855, p. 20; H.L. Clark 1914, p. 71; H.L. Clark 1925, p. 171; 
Mortensen 1948b, p. 411; A.M. Clark and Rowe 1971, p. 144 162; Rowe and Gates 1995, p. 
185. 
Mortensen (1948b: 411) lists additional synonymies. 


Material and locality 
Two tests, B 20024 (naked) and B 20025 (with spines), from the vicinity of Cape Gazelle, New 
Britain, East New Britain Province, PNG. 


Remarks 

Echinodiscus tenuissimus (L. Agassiz in Agassiz and Desor 1847) is a widely distributed Indo-West 
Pacific form, occurring throughout the East Indies, northern Australia, southern Japan and the south Pacific 
(Mortensen 1948b; A.M. Clark and Rowe 1971). In the south Pacific, the species is recorded from Tanna, 
Vanuatu, (H.L. Clark 1925) and from New Caledonia (A.M. Clark and Rowe 1971). However, De Ridder 
(1986) only noted the occurrence of Echinodiscus bisperforatus Leske 1778 from New Caledonia. H.L. Clark 
(1925) observed that New Caledonian specimens of EF. tenuissimus in the British Museum (Natural History) 
have a form more like E. bisperforatus. The Cape Gazelle specimens have very short lunules, about one quarter 
the length of the radius taken through them, and there is no difference in the tuberculation and spines of the 
ambulacral and interambulacral areas of the oral surface, both diagnostic characters of E. tenuissimus (Mortensen 
1948b; A.M. Clark and Rowe 1971). 


Superorder ATELOSTOMATA Zittel 1879 
Order SPATANGOIDA Claus 1876 
Suborder HEMIASTERINA Fischer 1966 
Family SCHIZASTERIDAE Lambert 1906 
Genus SCHIZASTER L. Agassiz 1836 


Type species 
Schizaster studeri L. Agassiz 1836, p. 185, by subsequent designation ICZN 1948. 


Remarks 

McNamara and Philip (1980a, b) questioned the familial classification of the spatangoids used by 
Mortensen (1951) and Fischer (1966) and, in particular, the Family Schizasteridae. Within the Schizasteridae 
McNamara and Philip recognized genera sharing the gross morphological test features of Schizaster, viz. a 
posteriorly located apical system, with the apex of the test posterior to this; a long, typically sunken, poriferous 
frontal ambulacrum; and sunken petals, of which the posterior pair are markedly shorter than the anterior ones. 
Within this group, McNamara and Philip (1980a, b) included the genus Schizaster L. Agassiz 1836 (with its 
subgenera Dipneutes Arnaud 1891; Paraster Pomel 1869 and Ova Gray 1825 [= Diploraster Mortensen 1951)); 
Brisaster Gray 1855; Kina Henderson 1975; Moira L. Agassiz 1872 (= Moiropsis L. Agassiz 1881); and Proraster 
Lambert 1895 (= Hypselaster Clark 1917). The author accepts their emended diagnosis for Schizaster. 


Subgenus PARASTER Pomel 1869 


Type species 
Schizaster gibberulus L. Agassiz 1847, by original designation of Pomel 1869, p. 14. 


130 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Diagnosis 

Species of Schizaster with a small to moderate sized test, with a shallow frontal sinus. Apical system 
slightly posterior of centre. Frontal ambulacrum shallow with pore pairs inclined at about 45° and arranged in 
single rows. Anterior petals almost straight, diverging at an angle up to 110° (McNamara and Philip 1980a). 


Remarks 

There is difficulty in placing the Cape Gazelle species firmly within McNamara and Philip’s (1980a) 
subgenus Paraster Pomel 1869. This is particularly in relation to details of the anterior petals, their flexed 
nature and 80° angle of divergence, both characters diagnostic of subgenus Schizaster L. Agassiz 1836. The 
frontal ambulacrum does not possess the steeper sided walls typical of species referred to Schizaster (Schizaster) 
(McNamara and Philip 1980a). Furthermore, McNamara and Philip (1980a) noted that species referred to 
Schizaster (Schizaster) possess a more elongate, narrower test than those assigned to Paraster. The Cape Gazelle 
species is assigned to Schizaster (Paraster) by its possession of a small test, shallow frontal sinus, apical system 
slightly posterior of centre and shallow frontal ambulacrum with pore pairs inclined at about 45°. The species 
is probably morphologically transitional between the Paraster and Schizaster morphotypes. 


Schizaster (Paraster) ovatus sp. nov. 
Figs 6a-d 


Diagnosis 

A small species of Schizaster (Paraster) with a moderately depressed, ovoid test; apical system is 55 
percent of test length from anterior, with four genital pores. Anterior ambulacrum relatively narrow and shallow; 
pore pairs inclined at about 45° and arranged in single rows; outer pores elongate, with similarly sized inner 
pores comma-shaped. Frontal sinus shallow. 


Etymology 

Ovatus L. egg- 
shaped, in reference to 
the form of the test, 
distinctive amongst the 
Schizasteridae. 


Material and locality 
Holotype ANU 
60653, a complete naked 
test from the vicinity of 
Cape Gazelle, New 
Britain, East New Britain 
Province, PNG. 


Description 

Test of small size, 
elongate ovoid, with 
length x width x height 
measuring 34 x 28 x 18 
mm; test length:width = 
1.21, width:height = 1.55. 
Test moderately 
depressed, with apical 
system located 55 percent 
of test length from 
anterior; test highest 
posterior to apical system, 


along keel of ambulacrum 
Figure 6. Schizaster (Paraster) ovatus sp. nov. Cape Gazelle, East New Britain. vy. Oral surface is gently 


6a-d, ANU 60653, aboral, oral, lateral, posterior views. Bar scale = 2.5 mm. 


Proc. Linn. Soc. N.S.W., 125, 2004 131 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


convex. Apical system ethmolytic, depressed, with four genital pores, posterior pair being larger than anterior 
pair. Frontal ambulacrum long, shallow and narrow (12 percent of test length); pore pairs inclined at about 45° 
and arranged in a single row. Outer pores elongate, with similarly sized inner pores comma-shaped. Frontal 
sinus broad and shallow. Interambulacra II and II form sharp, high keels. Anterior petals diverging at angle of 
80°; flexed distally and shallow, bearing pore pairs which are elliptical, widely spaced and conjugate; 26 pairs 
are present. Posterior petals are moderately long (occupying 21 percent of test length), bearing 18 pore pairs. 

Peripetalous fasciole is distinct, passing transversely between posterior petals and thickening at petal 
ends; the fasciole describes a concave arc between the extremities of the posterior and anterior petals, with an 
outwards flexure, corresponding with a constriction, forward of the apical system. Fasciole reaches maximum 
thickness at the extremities of the anterior petals. Peripetalous fasciole passes forward from anterior petals at 
about 60° before curving strongly to close with frontal ambulacrum; constrictions occur on interambulacral 
keel and adjacent to the abrupt curvature. Lateroanal fasciole is narrower than peripetalous fasciole and of more 
constant width. Lateroanal fasciole extends abaxially posteriorly from peripetalous fasciole at constriction between 
posterior and anterior petals; at ambitus it runs far below periproct, close to adoral surface. 

Peristome oval and slightly sunken; situated anteriorly, anterior tip of labrum 15 percent of test length 
from anterior. Anteriorly labrum is strongly curved; bounded by thick rim that degenerates laterally. Labrum as 
long as broad; posterior extension triangular, about as long as broad. Labrum carries several small tubercles 
anteriorly. Plastron is pear-shaped and broad, maximum width being 3/4 length. Plastron tubercles are arranged 
in curving rows. 

Periproct at mid-level on sub-truncate end of test. Periproct longitudinally elliptical, with a prominent 
narrow slit extending a short distance axially and aborally towards interambulacrum V, nearly reaching apical 
surface (Fig. 6d). 


Remarks 

Schizaster (Paraster) ovatus sp. nov. can be distinguished from other Schizaster-like heart urchins by its 
small, distinctively narrower and less inflated test, and long, shallow and narrow frontal ambulacrum. The test 
L:W and L:H ratios of 1.21 and 1.88 are larger than for most other echinoids of this group. The presence of four 
genital pores would suggest that the holotype is a mature specimen. McNamara and Philip (1980b) noted that in 
Schizaster (Ova) myorensis McNamara and Philip (1980b) the onset of maturity, occurring at a test length of 
about 25 mm, followed the sequential opening of the first, second, third and fouth genital pores. 

Morphological adaptations in Schizaster-like heart urchins are related to a need to produce a more efficient 
current flow over the aboral surface in sediment of low permeability (McNamara and Philip 1980a). The posterior 
migration of the apex meant more water would flow over over the frontal sinus to the peristome; the deepening 
of the frontal ambulacrum and the frontal sinus assisted in channelling water to the peristome; and the deep and 
long frontal ambulacrum further enabled more-funnel-building tube feet to be accommodated, presumably in 
response to finer-grained sediment (McNamara and Philip 1980a). The weakly vaulted test of S.(P.) ovatus 
with its shallow, open frontal ambulacrum and shallow frontal sinus suggests the species was a shallow-burrower 
in coarse (permeable) shell gravel. 


Suborder MICRASTERINA Fischer 1966 
Family BRISSIDAE Gray 1855 
Genus BRISSUS Gray 1825 


Synonymy 
Bryssus Martens 1869, p. 128 (nom. van.). 
Brissus (Allobrissus) Mortensen 1950, p. 162. 


Type species 
Spatangus brissus unicolour Leske 1778, p. 248 by subsequent designation of ICZN, Op. 290 1948. 


Brissus (Brissus) latecarinatus (Leske 1778) 
Synonymy 


Brissus carinatus Gray 1825, p. 431; A. Agassiz 1872-74, p. 96, 596. 
Brissus latecarinatus (Leske 1778): H.L. Clark 1921, p. 153; H.L. Clark 1925, p. 219; H.L. Clark 


132 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


1946, p. 375; Mortensen 1951, p. 514; A.M. Clark and Rowe 1971, p. 165; Gibbs et al. 1976, 
p. 135. 

Brissus (Brissus) latecarinatus: Rowe and Gates 1995, p. 187. 

Spatangus Brissus latecarinatus Leske 1778, p. 249. 

Mortensen (1951: 514) lists additional synonymies. 


Material and locality 
Three naked tests, ANU 60643-5, from Nosnos village, Boang Island, Tanga Group, New Ireland 
Province, PNG; one naked test, B 20014, from Cape Gazelle, New Britain, East New Britain Province, PNG. 


Remarks 

Brissus (Brissus) latecarinatus (Leske 1778) is a widely distributed species throughout the Indo-Pacific 
(Mortensen 1951; A.M. Clark and Rowe 1971). It is present on Australian coasts, from Queensland to Port 
Jackson, and is also known from Lord Howe Island (H.L. Clark 1946). Miskelly’s (2002) record of the species 
from the Solomon Islands is nearest to the present record in the Tanga Group. The largest specimen, ANU 
60644 from the Tanga Group, measures 70 x 60 x 39 mm, considerably smaller than the largest known specimen, 
from Hawaii, measuring 130 x 108 x 74 mm (H.L. Clark 1946). The shape of the periproct of the Tanga Group 
and Cape Gazelle specimens, somewhat pointed above and below, differs from the rounded periproct evident in 
specimens figured by Mortensen (1951: Plate XXXII, fig. 7) and Miskelly (2002). In this respect, the Bismarck 
Sea specimens closely resemble Brissus (Allobrissus) agassizii Doderlein 1885 (Mortensen 1951: Plate XXXII, 
fig. 7). Gibbs et al. (1976) noted the similarity of a Pelican Island, Great Barrier Reef, specimen of B. (B.) 
latecarinatus with B. (A.) agassizii. The posterior end of this particular specimen, like that of B. (A.) agassizii, 
is vertically truncated, with the posterior interambulacrum being only slightly carinate aborally (and not prolonged 
backwards to overhang the periproct and conceal it from dorsal view). 


Genus METALIA Gray 1855 


Synonymy 
Xanthobrissus Agassiz 1863, p. 28. 
Prometalia Pomel 1883, p. 34. 
Eobrissus Bell 1904, p. 236. 
Metaliopsis Fourtau 1913, p. 68. 


Type species 
Spatangus sternalis Lamarck 1816, p. 326, by original designation. 


Metalia spatagus (Linnaeus 1758) 


Synonymy 
Echinus spatagus Linnaeus 1758, p. 665. 
Metalia spatagus (Linnaeus 1758): H.L. Clark 1925, p. 216; H.L. Clark 1932, p. 219; H.L. Clark 
1946, p. 372; Mortensen 1951, p. 540; A.M. Clark and Rowe 1971, p. 166; Gibbs et al. 1976, 
p. 136; Rowe and Gates 1995, p. 190. 
Mortensen (1951: 540) lists additional synonymies. 


Material and locality 
Two naked tests, ANU 60646-7, from Nosnos village, Boang Island, Tanga Group, New Ireland 
Province, PNG. 


Remarks 

Metalia spatagus (Linnaeus 1758) is widely distributed through the Indo-Pacific (Mortensen 1951; A.M. 
Clark and Rowe 1971). H.L. Clark (1932) provided the first record of this species from Australasian waters 
(Low Isles, Great Barrier Reef), recording the largest known specimen, measuring 110 x 93 x 52 mm. By 
comparison, the largest Tanga specimen measures 54 x 40 x 29 mm. Miskelly (2002) records the species from 
the Solomon Islands. 


Proc. Linn. Soc. N.S.W., 125, 2004 133 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


ACKNOWLEDGMENTS 


The author is grateful to Alistair Norrie and 
Richard Joycey, East New Britain Historical and Cultural 
Centre, for the loan of specimens from the Kokopo 
Museum, Kokopo, PNG. Prof. Ken Campbell kindly 
provided his thoughts on the erection of new species 
Schizaster (Paraster) ovatus sp. noy. and Heliocidaris 
robertsi sp. nov. and Loisette Marsh, Western Australian 
Museum, kindly provided an opinion on the identification 
of the juvenile Protoreaster nodosus from New Britain. 
Dr. Richard Barwick and Dr. Maité LeGleuher, both of 
the Department of Geology, Australian National 
University, kindly photographed all specimens, and 
provided translation from French of sections from Koehler 
(1910) and De Ridder (1986), respectively. The comments 
of Geoff Francis and an anonymous reviewer improved 
the manuscript. 


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Miskelly, A. (2002). ‘Sea urchins of Australia and the 
Indo-Pacific’. (Capricornica Publications: 
Sydney). 179 pp. 

Mortensen, T. (1928a). A Monograph of the Echinoidea I, 
Cidaroidea. (C.A. Reitzel, Copenhagen). 551 pp. 

Mortensen, T. (1928b). Papers from Dr. Th. Mortensen’s 
Pacific expedition 1914-16. New Cidaridae. 
Videnskabelige Meddelelser fra Dansk 
Naturhistorisk Forening 1 K¢kenhayn 85, 65-74. 

Mortensen, T. (1943a). A Monograph of the Echinoidea 
I1I.2 Camarodonta I: Orthopsidae, 
Glyphocyphidae, Temnopleuridae and 
Toxopneustidae. (C.A. Reitzel, Copenhagen). 553 
pp. 

Mortensen, T. (1943b). A Monograph of the Echinoidea 
IlI.3 Camarodonta II: Echinidae, 
Strongylocentrotidae, Parasalenidae, 
Echinometridae. (C.A. Reitzel, Copenhagen). 446 
PP: 

Mortensen, T. (1948a). A Monograph of the Echinoidea 
IV.1 Holectypoida, Cassiduloida. (C.A Reitzel, 
Copenhagen). 371 pp. 

Mortensen, T. (1948b). A Monograph of the Echinoidea 
IV.2 Clypeasteroida: Clypeasteridae, Arachnoidae, 
Fibulariidae, Laganidae and Scutellidae. (C.A. 
Reitzel, Copenhagen). 471 pp. 

Mortensen, T. (1951). A Monograph of the Echinoidea 
V.2 Spatangoida II: Amphisternata II, 
Spatangidae, Loveniidae, Pericosmidae, 
Schizasteridae, Brissidae. (C.A. Reitzel, 
Copenhagen). 593 pp. 

Miller, J. and Troschel, F.H. (1842). “System der 
Asteriden’. (Papier, Druck and Verlag: 
Braunschweig). 134 pp. 

Negretti, B., Philippe, M., Soudet, H.J., Thomassin, B.A. 
and Oggiano, G. (1990). Echinometra miocenica 
Loriol, echinide Miocéne, synonyme 
d’Echinometra mathaei (Blainville), actuel: 
Biogéographie et paléoécologie. Geobios 23, 445- 
456. 

Philip, G.M. (1965). The Tertiary echinoids of south- 
eastern Australia Ill Stirodonta, Aulodonta, and 
Camarodonta (1). Proceedings of the Royal 
Society of Victoria 78(2): 181-196. 

Read, J.R.L. (1965). Preliminary geological investigation 
of the lower Warangoi hydro-electric scheme. 
New Britain August-November 1964. Bureau of 
Mineral Resources, Geology and Geophysics, 
Record 1982/15. 

Rowe, F.W.E. and Gates, J. (1995). Echinodermata. In 
‘Zoological Catalogue of Australia’, Vol. 33. 510 
pp. (CSIRO Australia: Melbourne). 

Sladen, W.P. (1889). Report on the scientific results of the 
voyage of H.M.S. Challenger during the years 
1873-76: Zoology - vol. 30, Report on the 


135 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Asteroidea. 893pp. (H.M. Stationery Office: 
London). 

Spencer, W.K. and Wright, C.W. (1966). 

Asterozoans. In “Treatise on Invertebrate 
Paleontology, Part U, Echinodermata 3’ (Ed. 
R.C. Moore) pp. U4-107. (Geological Society 
of America and University of Kansas Press: 
Lawrence). 

Sluiter, C.P. (1895). Die Asteriden Sammlung des 
Museums zu Amsterdam. Die Echiniden 
Sammlung des Museums zu Amsterdam. 
Bijdragen tot de Dierkunde 17, 49-64, 65-74. 

Struder, T. (1876). Uber Echinodermen aus dem 
antarktischen Meere und zwei neue Seeigel von 
den Papua-Inseln gesammelt auf der Reise S.M.S. 
‘Gazelle’ um die Erde. Monatsberichte der 
Koniglich Preussischen Akademie der 
Wissenschaften zu Berlin 1876, 452-465. 

Struder, T. (1880). Ubersicht iiber die wahrend der Reise 
S.M.S. Corvette ‘Gazelle’ um die Erde 1874-76 
gesammelten Echinoiden. Monatsberichte der 
Koniglich Preussischen Akademie der 
Wissenschaften zu Berlin 1880, 861-885. 

Wagner, C.D. and Durham, J.W. (1966). Holectypoids. In 
“Treatise on Invertebrate Paleontology, Part U, 
Echinodermata 3’ (Ed. R.C. Moore) pp. U440- 
450. (Geological Society of America and 
University of Kansas Press: Lawrence). 

Wallace, D.A., Johnson, R.W., Chappell, B.W., Arculus, 
R.J., Perfit, M.R. and Crick, I.H. (1983). 
Cainozoic volcanism of the Tabar, Lihir, Tanga 
and Feni Islands, Papua New Guinea: Geology, 
whole-rock analyses, and rock-forming mineral 
compositions. Bureau of Mineral Resources, 
Geology and Geophysics, Report 243. 

Willey, A. (1902). Zoological results based on material 
from New Britain, New Guinea, Loyalty Islands 
and elsewhere, collected during the years 1895 
1896 and 1897. Pt. 2. (Cambridge University 
Press: Cambridge). 253pp. 

Woods, H. (1958). ‘Palaeontology Invertebrate’. 
(Cambridge University Press; Cambridge). 477pp. 


136 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Table 1. Reported starfishes from the Bismarck Archipelago, Papua New Guinea. 


ASTERIINAE 
Tarsaster stoichodes Sladen 1889: Fisher 1919, p. 491: north of the Admiralty Group (150 fathoms). 


ASTERINIDAE 
Asterina cephus (Miller and Troschel 1842): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. 
Patiriella exigua (Lamarck 1816): A.H. Clark 1954, p. 258: Admiralty Group; Seleo Island, Aitape district. 


ASTEROPSEIDAE 
Asteropsis carinifera (Lamarck 1816): A.H. Clark 1954, p. 258: Seleo Island, Aitape district. 


ASTROPECTINIIDAE 
Astropecten monacanthus Sladen 1883: Bell 1899, p. 136: New Britain. 
Astropecten polyacanthus Miiller and Troschel 1842: Fisher 1919, p. 64: Admiralty Group. 


ECHINASTERIDAE 
Echinaster luzonicus (Gray 1840): Rowe and Gates 1995, p. 59. (= Echinaster eridanella Miiller and 
Troschel 1842, p. 24; Bell 1899, p. 138): New Ireland; New Britain. 


LUIDIIDAE 
*Luida aspera Sladen 1889: Fisher 1919, p. 171: north of Admiralty Group (150 fathoms). 


OPHIDIASTERIDAE 

Linckia laevigata (Linnaeus 1758): Bouillon and Jangoux 1984, p. 249: Laing Island reef, Hansa Bay. 

Nardoa novaecaledoniae (Perrier 1875): Rowe and Gates 1995, p. 88. (= Nardoa mollis de Loriol, 1891, 
H.L. Clark 1946, p. 115; A.H. Clark 1954, p. 255): New Britain; Seleo Island, Aitape district. 

Nardoa tuberculata Gray 1840: Rowe and Gates 1995, p. 88. (= Nardoa finschi de Loriol 1891; Nardoa 
pauciforis von Martens 1866, H.L. Clark 1946, p. 115): New Britain. 

Ophidiaster granifer Liitken 1871: A.H. Clark 1954, p. 256: Seleo Island, Aitape district. 


OREASTERIDAE 

+Anthenea sidneyensis Déderlein 1915: Rowe and Gates 1995, p. 98: Manus Island (Admiralty Group). 

Culcita novaeguineae Miiller and Troschel 1842: A.H. Clark 1954, p. 254: Seleo Island, Aitape district. 

Pentaster obtusatus (Bory de St. Vincent 1827). [= Pentaceropsis obtusata (Bory de St. Vincent 1827) Bell 
1899, p. 136]: Blanche Bay, New Britain. 

Protoreaster lincki (de Blainville 1830): Oreaster lincki (= Pentaceros lincki, Bell 1899, p. 136): Blanche 
Bay, New Britain. 

Protoreaster nodosus (Linnaeus 1758): H.L. Clark 1946, p. 106; A.H. Clark 1954, p. 254. (= Pentaceros 
nodosus, Bell 1899: p. 136; Oreaster nodosus H.L. Clark 1908): Blanche Bay, New Britain; Seleo 
Island, Aitape district. 


PTERASTERIDAE 
Hymenaster pullatus Sladen 1889: Fisher 1919, p. 467: southwest of the Admiralty Group (1,070 fathoms). 


NOTES 
+ the writer follows Spencer and Wright (1966) and Rowe and Gates (1995) in placing Anthenea in Family 
Oreasteridae. H.L. Clark (1946) and A.M. Clark and Rowe (1971) placed the taxon in Family Goniasteridae. 


* Denotes type locality in Bismarck Archipelago. 


Proc. Linn. Soc. N.S.W., 125, 2004 137 


FOSSIL AND LIVING ECHINODERMS FROM PAPUA NEW GUINEA 


Table 2. Reported shallow and deep-water sea-urchins from the Bismarck Archipelago, Papua New 
Guinea. 


ARACHNOIDIDAE 
Arachnoides placenta (Linnaeus 1758): Bell 1899, p. 136; H.L. Clark 1925, p. 154: New Britain. 


ARBACIIDAE 

*Pygmaeocidaris prionigera (A. Agassiz 1879): A. Agassiz 1881, pl. XXXIV, figs 14 and 15; H.L. Clark 
1925, p. 73 (= Podocidaris prionigera A. Agassiz 1879, p. 199): between New Guinea and Admiralty 
Group (1,070 fathoms). 

*Coelopleurus elegans (Bell 1899): H.L. Clark 1925, p. 73. (= Salmacis elegans Bell 1899, p. 135): New 
Britain. 


CIDARIDAE 

Eucidaris metularia (Lamarck 1816): H.L. Clark 1925, p. 20. (= Cidaris metularia de Blainville, 1830, Bell 
1899, p. 134): New Britain. 

Prionocidaris baculosa var. annulifera (Lamarck): Mortensen 1928a, p. 437, 446. (= Schleinitzia crenularis 
Struder 1876, p. 463; 1880, p. 865): west New Guinea. 

Stylocidaris reini (Déderlein): H.L. Clark 1925, p. 24; Mortensen 1928a, p. 342, 347, 474 (= Phyllacanthus 
annulifera Bell 1899, p. 134): New Britain; Milne Bay. 


DIADEMATIDAE 
Echinothrix calamaris (Pallas 1774): A.H. Clark 1954, p. 250: Bougainville Island. 


*Micropyga nigra H.L. Clark 1925: A. Agassiz 1879, p. 200; H.L. Clark 1925, p. 47. (= Astropyga elastica 
Struder, Bell 1899, p. 135): New Britain. 

Micropyga tuberculata A. Agassiz 1879, p. 200: A. Agassiz 1881, pl. VI; H.L. Clark 1925, p. 48: Blanche 
Bay, New Britain. 


ECHINOMETRIDAE 
Echinometra mathaei (de Blainville 1825): A.H. Clark 1954, p. 251: Bougainville Island; Seleo Island, 
Aitape district; Normanby Island. (= Echinometra lucunter Bell 1899, p. 136). 


ECHINOTHURIIDAE 
Araeosoma gracile (A. Agassiz 1881): A. Agassiz 1881, p. 89; H.L. Clark 1925, p. 61: Admiralty Group 
(150 fathoms). 


LAGANIDAE 

Laganum decagonale (de Blainville 1827): A. Agassiz 1881; H.L. Clark 1925, p. 156; Mortensen 1948b, p. 
332, 336; Lindley 2003a, p. 133: near Admiralty Group (150 fathoms). 

Laganum depressum vat. tonganense (Quoy and Gainard): Mortensen 1948b, p. 324: Admiralty Group. 

Laganum laganum (Leske): Mortensen (1948b), p. 312: Bismarck Archipelago. 


SPATANGIDAE 
Maretia ovata (Leske 1778): A. Agassiz 1881; H.L. Clark 1925, p. 226: Admiralty Group. 


TEMNOPLEURIDAE 

Prionechinus agassizii Wood-Mason and Alcock 1891: H.L. Clark 1925, p. 78. (= Echinus elegans, A. 
Agassiz 1881): near Admiralty Group. 

*Prionechinus sagittiger A. Agassiz 1879, p. 202: A. Agassiz 1881, pl. IVa, figs 11-14; H.L. Clark 1925, p. 
79: between New Guinea and Admiralty Group (1,070 fathoms). 

Temnopleurus sp., Bell 1899, p. 135: New Britain. 

Temnopleurus reevesii (Gray 1855): A. Agassiz 1881; H.L. Clark 1925, p. 81: near Admiralty Group (150 
fathoms). 


138 Proc. Linn. Soc. N.S.W., 125, 2004 


I.D. LINDLEY 


Temnotrema scillae (Mazetti 1894): Mortensen 1904, p. 86; H.L. Clark 1925, p. 91 (= Pleurechinus 
reticulatus in H.L. Clark 1925, p. 91): New Britain. 


TOXOPNEUSTIDAE 
Tripneustes gratilla (Linnaeus 1758): A.H. Clark 1954, p. 250: Bougainville Island; Seleo Island, Aitape 


district. 


INVALID RECORDS 

Astriclypeus manni Verrill, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 416, 418. 
Colobocentrotus mertensi Brandt 1835, Sluiter 1895, p. 69, New Ireland; Mortensen 1943b, p. 433. 
Mellita longifissa Michelin 1858, Sluiter 1895, p. 73, New Ireland; Mortensen 1948b, p. 427, 428. 
Taxonomic reason: Erroneous labelling (Mortensen 1948b, p. 418; Mortensen 1943b, p. 433; Mortensen 


1948b, p. 428, respectively). 


NOTES 
* Denotes type localities in Bismarck Archipelago 


Proc. Linn. Soc. N.S.W., 125, 2004 


139 


; : - - a j 
-AND LIVING ECS gaging (PUR NEW GUINEA 


somes omer AS de raged MR i cabeakle. ( eae 
aM 412 .q 2891 del LH or 


7 (HDAI 


Suit habizh oni s2 ohaatak reg a snes _ peas 


rrnti lem A Tien sere TA ern Lite x, Agareis Ty et. ATV fier: Me 
(925-5 73a Le :d8dP}osteorOM danke we AT Ly De Bhy 2 
(oD) pee GiameannoM -bakled wort 00 .q 208! ronal? 2EBT oar rset 

HBESe Sten SREP) yesencitid  Chantl tt dolt AY sy ORB sotikeye 3 
tt Sesiinid Efhig ERR noananoM, -Bi bk .q debe] asensniM) gnitledal: cungnorn :anatan 
| ‘ene 
IDARIDA! _ migy 
Cueidarismenionid (larmerch 146): BL Clark £925, p. 26 ‘i Cmbiels owetiulea risa he Eittgervito, 
Sow 1A): Wew Brie. cailitnA fume a nilgol om 2 
renjdarly Darcie var oniiiers Camicks: Mortensen 19784. p 437. 446, t= ; 
sander 1? 6. 0.403; (REQ, 9. BAS): weer Now fines >. Vn = ‘Oder 2 ; 
tori rit (Dieter tein) Hi. Ciark 1035, fi 24: Muwtennest IDR TR, 3, A te 

innalilera BEIT 1899, 6 1545 | New Britain ithe Ray 

MADEMA TIDAL, « ) fp AS Waeeig 
Adviwvthrix <itlaradtess (Pallas 7 Tas 3 AOC wath | 1954, p. 250: avi Ilene. ih 
“Micropwga nigra Vb. Chars BS: Ac Agana 187% p 2007 Chur 1925p) ADA =A i 


vy Arn gs 

Sirieder, Bel 1 8969..45\) New Britain, A Lee Ve @ se 
Micropven tuberewlaa A Agassis WTI p. 200.8. Aguinia INU pL ns 725, Ae Bi 
Bay, New Arita rk ea 7 rare ees 


ECHINOMETRIDAE- _ = | 
Exhinometramatiae tide Risinvi le T825) AS. Clark 1234.9, 231: oo << eo. 4s 
Ajtape cisthict: Nonnanhy Island, (= Echinamrtg fhecoener Seely: bis e 136% S 
: i ine - 
ECHINGTHUBIEDAE as ae a) ee a 
pacesnmna sracte TAS Sgensie PhAty & Neeeoe’ ee p. 35 1. Chek 825.006 


Pane Fath} ' : Te. alin 


4 


LAG ANIA a : ti | | ms 

Lagann decdgenile (de Bittosie LRET LM Agnasih: 185A iri 

$32; 436; Lindley 20030, py (25, aeigr Aepiehiytipokay (50.6 ine 
Satya Mepresser vee. des aypeati ne (CMe ald Fated): MonecenatShs “pk 
Lagann layanun (Leske) Moreowen — r itis Bimask Aig 


SPATANGIDAE ; <2 oa ey 
Waretid ovdh (Lteke 178): A, isu 188: i HL Won 9, biog, 


TEMNOPLEURIDABR. 2” <u Yeaitig 
Prinnerhings aqisvigit Woresfastn ane sot: Cat 
Agassia IRB} near Adinjcalty Gram. 
Prone: ties wightiger A. Agaeste ree 2 
79: teeweocn New Guinea asl 
Termmaplearas Sp, Bell 189%, p, 13%- 
Termvaplenras: peetestl (Gray less A 
fattusunm). 


Conodont Faunas from the Mid to Late Ordovician Boundary 
Interval of the Wahringa Limestone Member (Fairbridge 
Volcanics), Central New South Wales 


Y.Y. ZHEN"’, I.G. PERcIvVAL” AND B.D. WeEBBy* 


‘Division of Earth and Environmental Sciences, The Australian Museum, 6 College Street, Sydney, N.S.W. 
2010 (yongyi@austmus.gov.au); "Geological Survey of New South Wales, Department of Mineral 
Resources, P.O. Box 76, Lidcombe, N.S.W. 2141 (ian.percival@minerals.nsw.gov.au); *Centre for 

Ecostratigraphy and Palaeobiology, Department of Earth and Planetary Sciences, Macquarie University, 


N.S.W. 2109, Australia (bwebby @laurel.ocs.mq.edu.au); Honorary Research Associate, Centre for 


Ecostratigraphy and Palaeobiology, Macquarie University, N.S.W. 


Zhen, YY. Percival, I.G. and Webby, B.D. (2004). Conodont faunas from the Mid to Late Ordovician 
boundary interval of the Wahringa Limestone Member (Fairbridge Volcanics), central New South 
Wales. Proceedings of the Linnean Society of New South Wales 125, 141-164. 


Twenty-nine conodont species are documented from the Wahringa Limestone Member and other isolated 
limestone pods of the Fairbridge Volcanics, in the Bakers Swamp area between Wellington and Orange, 
central New South Wales. Three conodont assemblages are recognised within the Wahringa Limestone 
Member. The oldest is characterised by the occurrence of Pygodus protoanserinus and Pygodus serra, 
indicative of a late Darriwilian age (Da3 to early Da4). The overlying assemblage B, bearing Belodina 
monitorensis, probably ranges across the Mid to Late Ordovician boundary. Assemblage C with abundant 
Belodina compressa in the upper part of the Wahringa Limestone Member is of late Gisbornian (Gi2) age. 
The conodont faunas are significant in being the first described from the Lachlan Orogen in New South 
Wales spanning the Mid to Late Ordovician interval, although resolution of the actual boundary level is 


limited in the section measured. 


Manuscript received 17 September 2003, accepted for publication 17 December 2003. 


KEYWORDS: Conodonts, Fairbridge Volcanics, Late Ordovician (Gisbornian), Mid Ordovician 


(Darriwilian), Wahringa Limestone Member. 


INTRODUCTION 


The Molong Volcanic Belt (MVB) is a 
meridionally-aligned tectonic feature of Ordovician 
age within the east Lachlan Orogen in central New 
South Wales (Glen et al. 1998). Ordovician strata in 
the northern sector of the MVB near Bakers Swamp 
between Orange and Wellington (Fig. 1) are 
represented by the Early Ordovician Mitchell 
Formation and Hensleigh Siltstone — the latter yielding 
conodonts of Bendigonian age (upper Prioniodus 
elegans Zone) from allochthonous limestones (Zhen 
et al. in press) — and the Fairbridge Volcanics of Mid 
to early Late Ordovician age. Two autochthonous 
limestone units occur in the Fairbridge Volcanics. The 
lower one, known as the Wahringa Limestone 
Member, is the subject of this paper. Representative 
conodonts were illustrated earlier in a preliminary 


report defining this unit, and their age connotations 
discussed (Percival et al. 1999). Subsequent detailed 
sampling has yielded many more elements and species, 
enabling broad confirmation of the original age 
determination and providing increased precision for 
the upper age limit of the limestone. The faunas are 
systematically described here for the first time. They 
are significant in spanning the Mid to Late Ordovician 
boundary, an interval which is otherwise poorly 
represented in shallow water settings of eastern parts 
of the Lachlan Orogen. 


STRATIGRAPHY 


Much of the Fairbridge Volcanics consists of 
andesitic lava flows, with subsidiary boulder 
conglomerates (Morgan, Scott & Percival, in Meakin 
& Morgan 1999). Allochthonous limestones are 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


CATOMBAL 
GROUP 
Vas 
C1474, C1483 
C1472 
C1695-C1700 
1 eae C1471, C1693, C1694 
eitk : C1429 
oso nig WAHRINGA cy45g 
ob Eales Pd = LIMESTONE a bans 
x x x wo 
= MEMBER ©1456 [Et 
vag aa ATI. S| crass, c1sa7, c1ase 
4 RA = : , 
616982 Visit o C1463 
C1695 y = 
C1699, 4 A458" “A A“ uw < 
38 Re ON Qa 
17 Abe DV = 
“Ei6er CSO care” c1463 E 
~ ©1450° C1488 = 
SIGE. 5 
“g'Wohringa" 
HENSLEIGH 
SILTSTONE 
MITCHELL 
FORMATION 
REFERENCE 
Ty N 
aedoael a [sal Quaternary alluvium _#2 Dip, and strike 
« RY \ 
| | ’ ; : 
; yo AUSTRALIA ae [: < :] Unditferentiated Devonian —{-> Synciine, with plunge 
| a 7s} \ 
a ———- of as Undifferentiated Silurian + ___ Anticline 
\ os | nsw i 
Ch pe Vip if . a 
- A | Neae/ cr C1458e Fossil local 
Loy 85 Fairbridge Volcanics is 
~ ges 
A 2 > allochthonous —_a__ Thrust fault 
FS 56 J jimestones 
£990 Wahringa Limestone REE 
NEW SOUTH WALES : Member 5 
» tO Waren 3 Type section 
tudy Area—a i * A 
ae Orange * ys aney c Hensleigh Siltstone 
xcs zo 
See ee 5 8 | limestones ) 1 2km 
Noe co} ee ed 
~ = 
A 2 L Mitchell Formation 


Figure 1. Locality maps. A. location of Wahringa area, between Wellington and Orange, central New 
South Wales; B. simplified geological map of the Wahringa area; C. generalised stratigraphic column for 
this area, showing spot sampled horizons within the Wahringa Limestone Member and the allochthonous 
limestone blocks within the Fairbridge Volcanics. For further details of the sampled section, see Figure 2. 


142 Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


uncommon in the lower Fairbridge Volcanics below 
the Wahringa Limestone Member and conodont yields 
are disappointingly low. Numerous allochthonous 
limestone pods emplaced within the Fairbridge 
Volcanics, stratigraphically overlying the Wahringa 
Limestone Member, were also processed for 
conodonts. With the exception of six samples that 
contained Belodina compressa (indicating an early 
Late Ordovician or Gisbornian age), these were either 
barren or yielded only sparse, non-diagnostic elements. 


Wahringa Limestone Member of Fairbridge 
Volcanics (Fig. 2) 

The name derives from the “Wahringa” 
property, located approximately 28 km south of 
Wellington on the Mitchell Highway. Here the 
Wahringa Limestone Member is exposed along strike 
for approximately 400 m and attains a thickness of 88 
m in its type section (situated just north of a bend in 
Bakers Swamp Creek). Invertebrate macrofossils 
described or illustrated from the Wahringa Limestone 
Member comprise brachiopods, gastropods, nautiloids, 
crinoids, demosponges, stromatoporoids, and a species 
of tabulate coral (Percival et al. 2001). The unit is 
subdivisible into three parts: lower beds rich in 
oncolites, ooids and volcaniclastic detritus, a middle 


C1687 4S 
C1683 3 
C1678 ‘= assemblage C 
C1677 ==, g 
C1676/44a ot e = 
C1645), ———— | Mg 9 
C1674 sae 
C1673 = é 9 
0 3 
a seo 
3 $ E 5 
B| eo8 
C1672 Soles 2 
2 2 3 g 
C1668 & = 20 
C1667 | ?ae 
3 
C1664 8 assemblage B 
® 
ane ce 
1 ek) a Se 
ane Ei Eo oe ¢ 
2 eee & © 
os 
a a § 
Vee € 
& 
& 
| 
aS) 
Cy 
rea) 


10m 


assemblage A 


e Pygodus protoanserinus 
e@ Erraticodon balticus? 


e@ Pygodus serra 
© Belodina sp. B 


0 


C1652 
(C1450) 


Figure 2. Stratigraphic section measured through 
the Wahringa Limestone Member showing sampled 
horizons and ranges of selected, age-significant 
conodont taxa. 


Proc. Linn. Soc. N.S.W., 125, 2004 


part of muddy, thinly bedded limestones rich in 
brachiopods, and an upper section that is more massive. 
The variation in lithologies reflects an increase in water 
depth from shallow subtidal at the base, to below 
normal wave base in the middle and upper beds. 
However, this depth increase does not have a 
controlling influence on the three distinct conodont 
assemblages recognised, which represent age- 
significant rather than biofacies-distinct assemblages. 

Lithologies in the lower part of the member, 
which consist mainly of red ooidal grainstones, 
calcarenites and oncolitic grainstone-packstones, are 
particularly characteristic of very shallow deposition. 
Fauna present in these beds (not observed at other 


-levels of the measured section) include the large 


gastropod Maclurites cf. M. florentinensis, the 
siphonotretoidean brachiopod Multispinula, and a 
Calathium-like receptaculitid. Demosponges, 
including Archaeoscyphia? sp. B, Malongullospongia? 
sp. and Hindia cf. H. sphaeroidalis, are more widely 
distributed but are especially common at this level. 
Fossil grains are subangular to rounded, frequently 
algal-coated and include dasycladacean and 
solenoporid algae. Large oncolites with well-preserved 
cyanobacterial Girvanella filaments and ooids are also 
abundant. 

The middle part of the Wahringa Limestone 
Member is characterised by fine to coarse grained 
skeletal grainstone (interbedded with silty layers) that 
is dominated by remains of echinoderms, brachiopods, 
dasycladacean algae, molluscs, trilobites, and 
ostracods. Brachiopods, including Sowerbyites?, 
Leptellina and rare Sowerbyella are concentrated in 
thin-bedded packstones in the middle part of the unit. 
Stromatoporoids (Labechia, Labechiella), mostly 
preserved in growth position, occur slightly above this 
level and range into the uppermost limestone beds of 
the unit. 

The most common lithologies in the upper 
part of the Wahringa Limestone Member are fine to 
coarse grained skeletal, oncolitic grainstone and lesser 
packstone to wackestone lacking internal lamination. 
Grains include ostracods, dasycladacean algae, rare 
ooids, and oncolites with associated Girvanella. 


CONODONT BIOSTRATIGRAPHY 


Twenty-nine conodont species based on 897 
individual specimens were recovered from 44 samples 
(Fig. 3), collected from the Wahringa Limestone 
Member and various limestone pods within the 
enclosing Fairbridge Volcanics. The faunas range in 
age from late Darriwilian (Da3, lower Eoplacognathus 
suecicus Zone) to late Gisbornian (Gi2, Belodina 


143 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Isolated limestone pods in Fairbridge 
Voics above Wahringa Lst type section 


Wahringa Lst Mbr 
NE of type section: 


Qq1aqlra 
alala 
| 3/8 


Wahringa Limestone Member type section 


e}e/e/ele/elelelelelelelelelele 

R18) 8) 8) 219) 8) 3) 8) 3) 3] 3) 9) 3) 3/8 

EE a a | 
MEAN SUSE lee ae S| Sane 
SEE Ree a 
eee 
EERE RASS 
Ea Wea Ts TR TEE I 


' Ist pods below 
State vege 
Samples fol Mell Mell me 

> > > 
$/ 8/8] 8 


-_ 


_ w 


Figure 3. Distribution chart of conodont species from 44 samples through the Wahringa Limestone Member, and allochthonous limestone blocks in the 
Fairbridge Volcanics, from above and below the Wahringa Limestone Member. Column with asterisk depicts occurrence of species in sole sample 
(C1429: middle or upper beds) from southwestern extremity of outcrop of the Wahringa Limestone Member. 


L68 


o 
Qq1aql1rala ie) Q;1aqrTa;a alalalal & 
SIS/S1 S/S] ele = sl] ele Bleleis 
S| S)=/ 5) 2/8/51) 8/3] 8/8 8) 3/$ia 


Proc. Linn. Soc. N.S.W., 125, 2004 


144 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


compressa Zone). 

The oldest fauna is represented by a small 
assemblage, including Appalachignathus delicatulus, 
Protopanderodus nogamii, ?Periodon aculeatus, 
Ansella sp., Erraticodon sp., and Stiptognathus sp. A 
from a single sample C1463, which was obtained from 
several small limestone clasts within the Fairbridge 
Volcanics at a stratigraphic level some 120 m below 
the Wahringa Limestone Member (Percival et al. 
1999). This fauna is comparable with that recently 
described from allochthonous limestones of Da3 age 
in the Oakdale Formation of the Bell River valley 
(Zhen and Percival in press), situated approximately 
23 km southeast of the “Wahringa” area. Of the six 
species recognised in sample C1463, four also occur 
in the Bell River valley fauna. Stiptognathus sp. A is 
rare, but the other three species (Appalachignathus 
delicatulus, Protopanderodus nogamii, and 
Erraticodon sp.) dominate in all five samples from the 
Oakdale Formation (Zhen and Percival in press). On 
this basis, the age of sample C1463 can now be revised 
downwards to the E. suecicus Zone from the 
previously-interpreted level near the Mid/Late 
Ordovician boundary (Zhen et al. 2001). 

In its type section, the Wahringa Limestone 
Member consists of three laterally continuous outcrops 
separated by two intervals of poor or negligible 
exposure (Fig. 2). Initially, one spot sample was 
collected from each of these three major outcrops, 
representing the lower, middle, and upper beds of the 
unit. Subsequent more intensive collecting during 
section measuring produced 17 samples that yielded 
conodonts. Although most of these samples have low 
yields and diversity, three conodont assemblages 
(herein referred to as A, B, and C from oldest to 
youngest) can be distinguished. 

Assemblage A was recovered from sample 
C1652 at the base of the Wahringa Limestone Member 
and spot sample C1450 within the lower part of this 
unit (essentially an equivalent stratigraphic level to 
C1652) in the type section. The fauna consists of 15 
species including Acodus sp., Ansella nevadensis, 
Ansella biserrata, Belodina sp. B, Dapsilodus 
variabilis, Drepanoistodus sp., Erraticodon balticus?, 
Oistodus? sp. cf. venustus, Panderodus gracilis, 
Periodon aculeatus, Phragmodus flexuosus, 
Protopanderodus nogamii, Protopanderodus 
varicostatus, Pygodus serra, and Pygodus 
protoanserinus. Most of these species are widely 
distributed and relatively long ranging, but the two 
species of Pygodus are important biostratigraphically. 
Pygodus protoanserinus has a range from the upper 
E. robustus Subzone to the E. lindstroemi Subzone 
(of the upper Pygodus serra Zone). One specimen (see 
Fig. 9K, L) referrable to the middle form of the Pa 


Proc. Linn. Soc. N.S.W., 125, 2004 


element of Pygodus serra (Zhang 1998a) was also 
recovered in the sample C1652. Co-occurrence of P. 
serra and P. protoanserinus places the base of the 
Wahringa Limestone Member precisely within the 
upper E. robustus Subzone (upper Da3 to lowest Da4) 
of the P. serra Zone. 

Closest correlations are with successions in 
China. In the top Guniutan Formation (upper P. serra 
Zone) of Hunan Province, Zhang (1998b) recorded 
the co-occurrence of Pygodus protoanserinus with 
Erraticodon balticus?, Protopanderodus varicostatus, 
and Periodon aculeatus. Pygodus serra, Periodon 
aculeatus, Protopanderodus  varicostatus, 
Protopanderodus nogamii, and Panderodus gracilis 
also occur in the lower part (P. serra Zone) of the 


-Pingliang Formation of the Ordos Basin (An and Zheng 


1990). 

Only five samples from the middle section 
of the Wahringa Limestone Member have yielded 
conodonts. These faunas are of very low diversity and 
productivity, and are referred to herein as Assemblage 
B. They include Ansella nevadensis, Ansella sp., 
Belodina monitorensis, Panderodus gracilis, and 
Periodon aculeatus. Of these, only B. monitorensis is 
significant for age determination, occurring widely 
within the late Darriwilian to Gisbornian interval 
(Sweet in Ziegler 1981). Assemblage B is likely very 
close to the Mid/Late Ordovician boundary, probably 
within the Cahabagnathus sweeti Zone, although the 
precise recognition of the boundary within the middle 
Wahringa Limestone Member is not determinable on 
current evidence. 

The upper part of the type section of the 
Wahringa Limestone Member is relatively more 
productive, with 12 samples yielding an assemblage 
(designated as Assemblage C) of 15 species including 
Acodus sp., Ansella nevadensis, Belodina compressa, 
Belodina monitorensis, Besselodus sp., Dapsilodus 
variabilis, Dapsilodus viruensis, Drepanoistodus sp., 
Oistodus? sp. cf. venustus, Panderodus gracilis, 
Periodon aculeatus, Protopanderodus cooperi, 
Protopanderodus varicostatus, Protopanderodus 
liripipus and Stiptognathus sp. B. As a zonal index 
species of late Gisbornian equivalents in the North 
American Midcontinent zonal scheme, the occurrence 
of Belodina compressa in the upper part of the 
Wahringa Limestone Member indicates that top of this 
unit may be as young as Gi2. This species also occurs 
in six samples from limestone pods within the 
Fairbridge Volcanics above the Wahringa Limestone 
Member. The presence of two elements confidently 
identified as this species in limestone pods in the 
Fairbridge Volcanics slightly below the base of the 
Wahringa Limestone Member (samples C1487 and 
C1488) cannot be explained at present, as this 


145 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Figure 4. A-G, Acodus sp.; A, B, P element, MMMC2639, C1450, A, inner lateral view, B, outer lateral 
view; C, D, P element, MMMC2640, C1652, C, outer lateral view, D, anterior view; E-G, Sa element, 
MMMC2641, C1680, lateral views. H-P, Ansella nevadensis (Ethington and Schumacher 1969); H-J, Pa 
element, MMMC2642, C1450, H, inner lateral view, I, outer lateral view, J, showing surface striation; K, 
L, Pb element, MMMC2643, C1450, K, outer lateral view, L, inner lateral view; M, N, Sa element, 
MMMC2644, C1683, lateral views; O, P, Sc element, MMMC2645, C1450, O, outer lateral view, P, inner 
lateral view. Q, Ansella biserrata Lehnert and Bergstrom in Lehnert et al. 1999; Pa element, MMMC2646, 
C1652, outer lateral view. R, S, Ansella sp.; R, Pb element, MMMC2647, C1486, inner lateral view; S, Pb 
element, MMMC2648, C1672, outer lateral view. T, Appalachignathus delicatulus Bergstrom et al. 1974; 
Pb element, MMMC2649, C1463, inner lateral view. Unless otherwise indicated scale bars are 100 um. 


146 Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


contradicts the species succession in all known global 
occurrences. 


LOCALITIES AND SAMPLES 


Details of the localities and measured section 
are shown in Figures | and 2, and summarised in the 
Appendix. Distribution of conodont species is 
presented in Figure 3. All illustrations of conodont 
elements are presented as SEM photomicrographs 
(Figs 4-9). Figured specimens bearing the prefix 
MMMC (“Mining Museum microfossil catalogue’’) 
are deposited in the collections of the Geological 
Survey of New South Wales, Sydney. Individual 
samples are referred to by the prefix “C”’. Although 
all species recorded are documented by illustration, 
only those where adequate material was recovered, or 
which are of biostratigraphic significance, are 
described. Unless otherwise mentioned, all specimens 
are from the Wahringa Limestone Member. 


SYSTEMATIC PALAEONTOLOGY 


Class CONODONTATA Pander 1856 
Genus ANSELLA Fahraeus and Hunter 1985a 


Type species 
Belodella jemtlandica Lofgren 1978. 


Ansella nevadensis (Ethington and Schumacher 
1969) 
Fig. 4H-P 

Synonymy 

Roundya sp. Sweet and Bergstrom 1962, p. 1244, 
1245, text-fig. 5. 

New Genus A Ethington and Schumacher 1969, p. 
478, 479, pl. 68, fig. 12, text-fig. 4J. 

Oepikodus copenhagenensis Ethington and 
Schumacher 1969, p. 465, pl.68, figs 5, 9, 
text-fig. 4L. 

Oistodus nevadensis Ethington and Schumacher 
1969, p. 467, 468, pl. 68, figs 1-4, text-fig. 
5C; Tipnis et al. 1978, pl. 6, fig. 7. 

Belodella nevadensis (Ethington and Schumacher); 
Bergstrom 1978, pl. 79, figs 9, 10; Bauer 
1987, text-fig. 5D. 

Ansella nevadensis (Ethington and Schumacher); 
Fahraeus and Hunter 1985a, p. 1175, 1176, 
pl. 1, figs 7, 10, pl. 2, figs 11a, b, 13a, b, 
14, text-fig. 2A-C; Bergstrom 1990, pl. 1, 
figs 11-14; McCracken 1991, p. 47-49, pl. 
3, figs 3, 4, 8, 9, 13, 14, 19-31 (cum syn.); 


Proc. Linn. Soc. N.S.W., 125, 2004 


?Bauer 1990, pl. 1, fig. 1; ?7Bauer 1994, fig. 
3.4, 3.5. 


Material 
Ten specimens (1 Pa, 4 Pb, 4 Sa, 1 Sc). 


Description 

The P elements are characterised by a prominent 
median costa on each side, and display a sharply inner 
laterally curved anterior margin. The Pa element has a 
row of denticles along the posterior edge (Fig. 4H-J). 
The denticle next to the cusp is the largest, and the 
others become gradually smaller towards the base. A 
sharp costa on each side extends from the tip of the 


cusp and disappears a short distance away from the 


basal margin. The Pb element has a sharp posterior 
margin without any denticles, a weaker and broader 
costa on the inner lateral side, and a sharp, strong costa 
on the outer lateral side (Fig. 4K, L). The Sa element 
is symmetrical with a row of closely spaced small 
denticles along the posterior margin, and bears a sharp 
antero-lateral costa on each side (Fig. 4M, N). The 
asymmetrical Sc element has an inner laterally curved 
anterior margin, and a row of small closely spaced 
denticles along the posterior margin (Fig. 40, P). 
Specimens are ornamented with fine striation. 


Discussion 

Originally proposed as the form species Oepikodus 
copenhagenensis Ethington and Schumacher 1969 and 
New Genus A Ethington and Schumacher 1969 (found 
in association with Oistodus nevadensis in the 
Copenhagen Formation of Nevada), these elements 
were considered as part of the species apparatus of A. 
nevadensis by Fahraeus and Hunter (1985a), and are 
herein assigned to the Pa and Sb positions. Fahraeus 
and Hunter (1985a) also illustrated a symmetrical Sa 
element (Fahraeus and Hunter 1985a, pl. 2, fig. 14) 
from the Cobbs Arm Limestone of Newfoundland. 
Specimens from the Wahringa Limestone Member 
permit differentiation of denticulate Pa and 
adenticulate Pb elements. The latter has not been 
recognised previously, but its assignment to the Pb 
position is consistent with the apparatus composition 
of comparable species such as A. jemtlandica and A. 
crassa Bauer 1994, from central New South Wales 
(Zhen and Percival in press). 


Ansella biserrata Lehnert and Bergstrom in Lehnert 
et al. 1999 
Fig. 4Q 
Synonymy 
Ansella biserrata Lehnert and Bergstr6m in Lehnert 
et al. 1999, p. 210, 212, pl. 1, figs 4, 7, pl. 
3, figs 1-3, 5 (cum syn.). 


147 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Material 
One specimen (Pa). 


Discussion 

Lehnert and Bergstrom in Lehnert et al. (1999) 
recognised a quadrimembrate apparatus for A. 
biserrata including biserrate, planoconvex, 
oistodiform, and triangular elements. We refer the 
biserrate and planoconvex elements to the Pa and Pb 
positions respectively, whereas the triangular element 
is regarded as taking either the Sa (symmetrical) or Sb 
(asymmetrical) position. The sole specimen from the 
lower Wahringa Limestone Member (C1652), with 
smooth lateral faces and fine denticles along its anterior 
and posterior margins, strongly resembles the holotype 
(a biserrate element) of A. biserrata from the basal 
Lindero Formation (Pygodus serra and P. anserinus 
zones) of west central Argentina (Lehnert et al. 1999). 


Ansella sp. 
Fig. 4R, S 


Synonymy 

Serraculodus? sp. Zhen and Webby 1995, p. 286, 
only pl. 5, figs 1, 3, 4. 

Ansella sp. Zhen et al. 2003a, p. 38, fig. 4A, 4B. 


Material 
Three specimens (Pb). 


Discussion 

These specimens are similar to the Pb element of A. 
nevadensis, but they lack the prominent lateral median 
costae of that species. They are identical with some 
specimens from the Fossil Hill Limestone of Eastonian 
age at Cliefden Caves previously assigned to 
Serraculodus? sp. (Zhen and Webby 1995). They can 
be distinguished from the Pb elements of both A. 
jemtlandica (Lofgren 1978) and A. crassa Bauer 1994 
in lacking the posteriorly more expanded base 
displayed in the latter two species (Zhen and Percival 
in press). 


Genus BELODINA Ethington 1959 


Type species 
Belodus compressus Branson and Mehl 1933. 


Belodina compressa (Branson and Mehl 1933) 
Fig. 5A-I 
Synonymy 
Belodus compressus Branson and Mehl 1933, p. 
114, pl. 9, figs 15, 16. 
Belodus grandis Stauffer 1935, p. 603-604, pl. 72, 
figs 46, 47, 49, 53, 54, 57. 


148 


Belodus wykoffensis Stauffer 1935, p. 604, pl. 72, 
figs 51,52, 55, 58;.59: 

Oistodus fornicalus Stauffer 1935, p. 610, pl. 75, 
figs 3-6. 

Belodina dispansa (Glenister); Schopf 1966, p. 43, 
plod, fie: 7- 

Belodina compressa (Branson and Mehl); Bergstrom 
and Sweet 1966, p. 321-315, pl. 31, figs 
12-19; Webers 1966, p. 24, pl. 1, figs 2, 6, 
7, 13, 15; Sweet in Ziegler 1981, p. 65-69, 
Belodina - plate 2, figs 1-4; An et al. 1983, 
only pl. 25, figs 13, 14; Moskalenko 1983, 
fig. 3Q-S; Leslie 1997, p. 921-926, figs 
2.1-2.20, 3.1-3.4 (cum syn.). 

Belodina confluens Sweet; Percival et al. 1999, p. 
13, fig. 8.21. 


Material 

255 specimens, including eobelodiniform, 
compressiform, grandiform and dispansiform 
elements, mainly from the upper part of the Wahringa 
Limestone Member; some specimens from 
allochthonous limestones in the Fairbridge Volcanics 
above the Wahringa Limestone Member, and two 
elements from limestone pods (C1487, C1488) in the 
Fairbridge Volcanics which apparently underlie the 
Wahringa Limestone Member. 


Discussion 

Of the known species of Belodina, three including B. 
compressa, B. confluens Sweet 1979, and B. 
monitorensis Ethington and Schumacher 1969, are 
morphologically very similar to each other, reflecting 
their close phylogenetic relationship. Well- 
documented successions in the U.S.A. (Sweet 1979) 
show that the oldest species, B. monitorensis, preceded 
B. compressa which was succeeded by B. confluens. 
Sweet (in Ziegler 1981, p. 65) revised all three species 
as consisting of trimembrate apparatuses, and 
emphasised that the type species of the genus, B. 
compressa, was characterised by having a distinct 
flattening (in lateral view) of the anterior margin near 
the antero-basal corner. This feature is more prominent 
in the compressiform element, as shown by the types 
(Branson and Mehl 1933) and also the specimens 
figured by Webers (1966); also see Sweet in Ziegler 
(1981). Both B. confluens and B. compressa are 
commonly found in association with a more slender, 
rastrate element bearing smaller denticles. Bergstrom 
(1990) suggested that these dispansiform elements 
might represent juveniles of the rastrate elements. 
Many other workers included these dispansiform 
elements in a separate species (dispansa) assigned 
either to Pseudobelodina (Sweet in Ziegler 1981, 
Nowlan and McCracken in Nowlan et al. 1988, 


Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


Figure 5. A-I, Belodina compressa (Branson and Mehl 1933); A, inner lateral view, B, outer lateral view, 
grandiform element, MMMC2650, C1472; C, dispansiform element, MMMC2651, C1429, inner lateral 
view; D, compressiform element, MMMC2652, C1472, inner lateral view; E, compressiform element, 
MMM(C2653, C1458, outer lateral view; F, compressiform element, MMMC2654, C1683, outer lateral 
view; G, dispansiform element, MMMC2655, C1429, outer lateral view; H, inner lateral view, I, outer 
lateral view, eobelodiniform element, MMMC2656, C1683. J-N, Belodina monitorensis Ethington and 
Schumacher 1969; J, outer lateral view, K, inner lateral view, grandiform element, MMMC2657, C1687; 
L, inner lateral view, M, outer lateral view, compressiform element, MMMC2658, C1456; N, 
eobelodiniform element, MMMC2659, C1456, outer lateral view. O, P, Belodina sp. B; O, eobelodiniform 
element, MMMC2660, C1450, inner lateral view; P, eobelodiniform element, MMMC2661, C1652, outer 
lateral view. Q, Belodina sp. A; grandiform element, MMMC2662, C1429, outer lateral view. R-T, 
Besselodus sp.; R, S, Sa element, MMMC2663, C1676, lateral views; T, M element, MMMC2664, C1675, 
anterior view. Scale bars are 100 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 149 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


McCracken and Nowlan 1989, Trotter and Webby 
1995, Leslie 1997, McCracken 2000) or to Belodina 
(Schopf 1966, Barnes 1977, Nowlan and Barnes 1981, 
Sansom et al. 1995). In comparing the apparatus 
architecture of Panderodus, Sansom et al. (1995) 
suggested that these slender dispansiform elements, 
which were previously included in Belodina dispansa 
(Glenister 1957) and Belodina arca Sweet 1979, might 
belong to the species apparatus of co-occurring B. 
confluens. Based on twelve well preserved, fused 
clusters of B. compressa recovered from the Plattin 
Limestone of Missouri and Iowa, Leslie (1997) 
demonstrated that B. compressa consisted of a 
quadrimembrate apparatus including the M 
(eobelodiniform), Sl (compressiform), S2 
(grandiform) and S2 (dispansiform) elements. He 
further suggested that the dispansiform element — 
although superficially similar in morphology to 
Pseudobelodina dispansa — was apparently not 
conspecific. Leslie also rejected the possibility that 
such elements represented the juveniles of 
compressiform and grandiform elements in 
consideration of the range of sizes and growth series 
preserved in the dispansiform elements. 

The material of B. compressa and B. 
confluens from central New South Wales shows 
recognisable differences between the two species. 
Compressiform elements of B. compressa display in 
lateral view a straight segment of the anterior margin 
near the antero-basal corner. In comparison, the 
anterior margin of the compressiform element of B. 
confluens is regularly curved near the antero-basal 
corner. Hence specimens of B. compressa previously 
reported from the Fork Lagoons Beds of central 
Queensland (Palmieri 1978), and from the Trelawney 
Beds of the New England Fold Belt (Philip 1966) were 
subsequently reassigned to B. confluens (Zhen and 
Webby 1995). Specimens from the Wahringa 
Limestone Member and various limestone pods within 
the Fairbridge Volcanics are the first confirmed records 
of B. compressa from eastern Australia. 


Belodina monitorensis Ethington and Schumacher 
1969 
Fig. 5J-N 

Synonymy 

Belodina monitorensis monitorensis Ethington and 
Schumacher 1969, p. 456, pl. 67, figs 3, 5, 
8, 9, text-fig. SD. 

Belodina monitorensis marginata Ethington and 
Schumacher 1969, p. 456, pl. 67, figs 1, 2, 
4, 6, text-fig. 5E. 

Eobelodina occidentalis Ethington and Schumacher 
1969, p. 456, pl. 67, figs 16, 20, text-fig. 
5H. 


150 


Belodina monitorensis Ethington and Schumacher 
1969, p. 455, 456; Sweet in Ziegler 1981, 
p. 79-81, Belodina - plate 1, figs 10, 11; 
Belodina - plate 2, figs 5-7; Bauer 1987, p. 
12, pl. 1, figs 10, 13, 14; Bauer 1990, pl. 1, 
fig. 9; Bauer 1994, fig. 3.16, 3.17, 3.20, 
Se 


Material 
17 specimens including eobelodiniform, 
compressiform and grandiform elements. 


Discussion 

Belodina monitorensis was originally defined as having 
prominent antero-lateral costae on either side of the 
grandiform element and generally four or five denticles 
on both grandiform and compressiform elements. A 
similar antero-lateral costa is also commonly found in 
the grandiform elements of B. compressa (Fig. 5B; 
also see Leslie 1997, fig. 2.3), and in the grandiform 
elements of B. confluens (McCracken 1987, pl. 1, fig. 
1; Zhen and Webby 1995, pl. 1, figs 17, 20; Zhen et 
al. 1999, fig. 5.8). Therefore, this character appears to 
be unreliable in characterising B. monitorensis. 
Although B. confluens and B. compressa typically have 
a greater number of denticles (five to nine), it seems 
rather arbitrary to split B. monitorensis (typically four 
or five denticles) from B. confluens based solely on 
the former having fewer denticles. 

Though the species status of B. monitorensis 
is uncertain in our view, stratigraphically it occurs 
much earlier than typical B. confluens. In the type 
section of the Wahringa Limestone Member, B. 
monitorensis occurs lower than B. compressa, but it is 
also found in association with the latter species in 
several samples in the upper part of the type section. 
The Wahringa Limestone Member specimens are 
comparable with the type material of B. monitorensis 
in having only three or four denticles, and in having 
an antero-lateral costa on the furrowed side of the 
grandiform element (Fig. 5J), also shown by the 
holotype (Ethington and Schumacher 1969, pl. 67, fig. 
5). Therefore, the species is tentatively retained here 
pending further detailed studies on B. monitorensis and 
other related species. 


Belodina sp. A 
Fig. 5Q 
Material 
One specimen from sample C1429 (upper beds of the 
Wahringa Limestone Member at the southwestern 
extremity of its outcrop). 


Discussion 
This compressiform element has a squat cusp and two 


Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


short and stout denticles along the posterior margin. 
The specimen is not as strongly compressed laterally 
as other known species of Belodina. 


Belodina sp. B 
Fig. 50, P 
Material 
Three eobelodiniform specimens. 


Discussion 

From its association with Pygodus protoanserinus in 
two samples (C1450 and C1652) from the lower part 
of the Wahringa Limestone Member this species has a 
late Darriwilian age (upper Da3 to lowest Da4, upper 
P. serra Zone), making it one of the earliest 
representatives of the genus Belodina. With a less 
extended heel it shows some morphological 
resemblance to the eobelodiniform element of 
Belodina beiyanhuaensis Qiu in Lin, Qiu and Xu 1984, 
but no rastrate elements have been recovered to 
confirm such an assignment. 


Genus ERRATICODON Dzik 1978 


Type species 
Erraticodon balticus Dzik 1978. 


Erraticodon balticus? Dzik 1978 
Fig. 6P, Q 

Synonymy 

Erraticodon balticus Dzik 1978, p. 66, pl. 15, figs 1- 
3, 5, 6, text-fig. 6; ?Stouge 1984, p. 84, pl. 
17, figs 9-19; Watson 1988, p. 113, pl. 5, 
figs 2-10, pl. 8, figs 1, 2, 5, 6, 8-13 (cum 
syn.); Dzik 1991, p. 299, fig. 12A; Ding et 
al. in Wang 1993, p. 176, pl. 37, only figs 
19-28, non fig. 18; ?Pohler 1994, pl. 3, figs 
3-5; Lehnert 1995, p. 87, pl. 10, figs 13, 16, 
pl. 12, figs 3-5; ?Zhang 1998b, p. 71, pl. 9, 
figs 6-13; ?Albanesi in Albanesi et al. 
1998, p. 176, pl. 4, figs 16-18; ?Johnston 
and Barnes 2000, p. 19, pl. 4, figs 18, 20, 
23, 24, 29; Zhao et al. 2000, p. 203, pl. 36, 
figs 1-16; ?Pyle and Barnes 2002, p. 111, 
pl.20, figs 8, 9. 


Material 
One specimen (M). 


Discussion 

Dzik (1978) originally defined the species as consisting 
of a seximembrate apparatus, but later (Dzik 1991) 
determined a septimembrate apparatus with digyrate 
Pa and Pb elements as typical of the species (Zhen et 
al. 2003b). Erraticodon balticus is characterised by 


Proc. Linn. Soc. N.S.W., 125, 2004 


having an accentuated denticle on the posterior process 
of the Sa, Sb and Sc elements (Dzik 1991, fig. 12A). 
The specimen from the Wahringa Limestone Member 
is broadly comparable with the M element of the 
illustrated type material (Dzik 1978, pl. 15, fig. 5), 
except that the latter has a reclined cusp; as our 
specimen has an erect cusp, it is only questionably 
referred to this species. 

Specimens ascribed to Erraticodon balticus 
from the Guniutan Formation of South China (Zhang 
1998b), the San Juan Formation of the Precordillera 
in Argentina (Albanesi in Albanesi et al. 1998), the 
Ospika Formation of British Columbia (Pyle and 
Barnes 2001), and the Cow Head Group of western 
Newfoundland (Johnston and Barnes 2000), all 
apparently lack the distinctive larger denticle on the 
posterior process of the S elements, and therefore 
should only be doubtfully assigned to the species. 


Erraticodon sp. 
Fig. 60 
Material 
One specimen (Sa) from sample C1463, a limestone 
pod in the Fairbridge Volcanics stratigraphically below 
the Wahringa Limestone Member. 


Discussion 

This alate element is identical with the Sa element of a 
new species of Erraticodon under description from 
allochthonous limestones within the Oakdale 
Formation of central New South Wales (Zhen and 
Percival in press). It has a prominent cusp with a 
flange-like costa on each side, which extends basally 
to define the upper margin of the lateral processes. 
The lateral processes bear four widely spaced, peg- 
like denticles. Comparison with other species of 
Erraticodon are discussed elsewhere (Zhen and 
Percival in press). 


Genus PERIODON Hadding 1913 


Type species 
Periodon aculeatus Hadding 1913. 


Periodon aculeatus Hadding 1913 
Figs 6R, S, 7A-K 

Synonymy 

Periodon aculeatus Hadding 1913, p. 33, pl. 1, fig. 
14; Lindstr6m 1955b, p. 110, pl. 22, figs 
10, 11, 14-16, 35; Lofgren 1978, p. 74, pl. 
10, fig. 1; pl. 11, figs 12-26, Fig. 29 (cum 
syn.); Sweet in Ziegler 1981, p. 237, 
Periodon - plate 1, figs 1-6; Nowlan 1981, 
pl. 4, figs 1-9; An 1987, p. 167, pl. 24, figs 
7-17; Bergstré6m 1990, pl. 1, figs 15, 16, pl. 


151 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Figure 6. A-D, Dapsilodus variabilis (Webers 1966); A, B, symmetrical distacodontiform element, 
MMMC2665, C1675, lateral views; C, symmetrical distacodontiform element, MMMC2666, C1450, lateral 
view; D, acodontiform element, MMMC2667, C1652, outer lateral view. E-J, Dapsilodus viruensis 
(Fahraeus 1966). E, F, Sa element, MMMC2668, C1675, lateral views; G, outer lateral view, H, inner 
lateral view, Sb element, MMMC2669, C1675; I, outer lateral view, J, inner lateral view, Sc element, 
MMMC2670, C1675. K-M, Drepanoistodus sp.; K, outer lateral view, L, inner lateral view, Sc element, 
MMMC2671, C1652; M, P element, MMMC2672, C1450, inner lateral view. N, Oistodus? sp. cf. venustus 
Stauffer 1935; anterior view, MMMC2673, C1450. O, Erraticodon sp.; Sa element, MMMC2674, C1463, 
postero-lateral view. P, Q, Erraticodon balticus? Dzik 1978; M element, MMMC2675, C1450, P, posterior 
view, Q, anterior view. R, S, Periodon aculeatus Hadding 1913; Pb element, MMMC2676, C1450, R, 
inner lateral view, S, outer lateral view. Scale bars are 100 Lm. 


is) Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


2, fig. 15; An and Zheng 1990, pl. 12, figs 
12-17; McCracken 1991, p. 50, pl. 1, figs 
13, 20, 22, 25-28, pl. 2, figs 24-27, 31, 34, 
35 (cum syn.); Pohler 1994, pl. 4, figs 30- 
32; Dzik 1994, p. 111, pl. 24, figs 10-13, 
text-fig. 31b; Lehnert 1995, p. 110, pl. 10; 
fig. 2, pl. 11, figs 10, 11, pl. 13, figs 9, 11, 
12, pl. 16, figs 8, 9, 11-13; Armstrong 
1997, p. 774, pl. 2, figs 13-21; text-fig. 3; 
Albanesi in Albanesi et al. 1998, p. 170, pl. 
15, figs 16-17, pl. 16, figs 19, 20 (cum 
syn.); Zhang 1998b, p. 80, 81, pl. 14, figs 
1-8 (cum syn.); Johnston and Barnes 2000, 
pys2-so, pl loetiss§ 2135177185 20-31, 
pl. 14, figs 1-7, text-figs 4, 5 (cum syn.); 
Rasmussen 2001, p. 110, pl. 13, figs 8-11 
(cum syn.); Pyle and Barnes 2002, p. 107, 
pl. 21, figs 7-9. 


Material 
97 specimens. 


Description 

Both Pa and Pb elements are angulate with a prominent 
cusp which is laterally compressed with a median costa 
on each side. The Pb element differs from the Pa 
element in having a twisted posterior process and a 
strongly inner laterally curved and downwardly 
extended anterior process (Fig. 6R, S). The M element 
is makellate with an adenticulate outer lateral process, 
and with 4-6 closely spaced denticles along the inner 
lateral margin. The alate Sa element has a long 
posterior process bearing closely spaced denticles. The 
sixth denticle away from the cusp is much larger and 
more robust (Fig. 7E, F). The lateral process on each 
side is blade-like, bearing small closely spaced 
rudimentary denticles along the edge. The basal cavity 
is shallow with a recessive basal margin zone. The Sb 
element is tertiopedate and asymmetrical, and bears a 
long denticulate posterior process with closely spaced, 
strongly reclined denticles, a long inner lateral process 
with more than seven small confluent denticles, and a 
short outer lateral process with only two small denticles 
(Fig. 7H, G). The Sc element is bipennate with a long, 
denticulate posterior process, bearing closely spaced, 
strongly reclined denticles, and with an inner laterally 
curved anterior process bearing small confluent 
denticles (Fig. 7I-K). 


Discussion 

Following the Treatise definition of the genus (Clark 
et al. 1981, p. W128), Sweet (1988) proposed a 
seximembrate apparatus for P. aculeatus, consisting 
of angulate Pa and Pb, makellate M, and ramiform 


Proc. Linn. Soc. N.S.W., 125, 2004 


alate Sa, tertiopedate Sb and bipennate Sc, elements. 
Albanesi (in Albanesi et al. 1998, text-fig 31, pl. 9, 
fig. 10) suggested a septimembrate apparatus for the 
species by recognizing a lozognathiform Sd element, 
which bears a long denticulate, twisted posterior 
process, a short, denticulate outer lateral process and 
an inner laterally curved, sharp, blade-like anterior 
costa. Rasmussen (2001, pl.13, fig.11) also recognised 
a modified tertiopedate Sd element, and described it 
as characterised by a multidenticulate, twisted, 
posterior process and weakly denticulated anterior 
process, and process-like extension of the outer-lateral 
costa or carina, but only a poorly preserved specimen 
was illustrated. In the Wahringa collections no Sd 


_ elements have been recognised. 


Lofgren (1978, p. 75) suggested that the 
number of small denticles between the cusp and the 
biggest denticle increases from a mean of 4.7 to 5.6 in 
successively younger samples. Specimens from the 
Wahringa Limestone Member may therefore represent 
more advanced forms of the species within its 
Stratigraphic range, as shown by the M elements, which 
are strongly geniculate with a sinuous basal margin 
and bear 4-6 denticles (mean 5.5) along the inner lateral 
margin. 


Genus PHRAGMODUS Branson and Mehl 1933 


Type species 
Phragmodus primus Branson and Mehl 1933. 


Phragmodus flexuosus Moskalenko 1973 
Fig. 7L, M 

Synonymy 

Phragmodus sp. Moskalenko 1972, p. 48-50, text- 
fig. 1, table 2. 

Phragmodus flexuosus Moskalenko 1973, p. 73, 74, 
pl. 11, figs 4-6; Sweet in Ziegler 1981, p. 
255-258, Phragmodus - plate 2, figs 1-6 
(cum syn.); Bauer 1987, p. 24, 25, pl. 3, 
figs 10, 14, 15, 17, 18, 20, 24, text-fig. 8; 
Ethington and Clark 1982, p. 79-82, pl. 9, 
figs 2-7 (cum syn.); Bauer 1994, p. 367, 
SOS, 11859-2955 9-20 9228,,9:50-9,99- 
Percival et al. 1999, fig. 8.15. 


Material 
One specimen (Sa). 


Discussion 

This specimen is alate with a suberect cusp, a long 
denticulated posterior process, and with a flange-like 
costa on each lateral side. The straight posterior process 
supports more than seven widely spaced denticles, 


153 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Figure 7. A-K, Periodon aculeatus Hadding 1913; A, outer lateral view, B, inner lateral view, Pa element, 
MMM(C2677, C1450; C, posterior view, D, anterior view, M element, MMMC2678, C1450; E, F, Sa 
element, MMMC2679, C1450, lateral views; G, outer lateral view, H, inner lateral view, Sb element, 
MMMC2680, C1450; I, outer lateral view, J, inner lateral view, Sc element, MMMC2681, C1450; K, Sc 
element, MMMC2682, C1652, outer lateral view. L, M, Phragmodus flexuosus Moskalenko 1973; Sa 
element, MMMC2683, C1450, lateral views. N-U, Panderodus gracilis (Branson and Mehl 1933); N, 
posterior view, O, inner lateral view, P, outer lateral view, graciliform element, MMMC2684, C1450; Q, 
posterior view, R, basal view, S, lateral view, aequaliform element, MMMC2685, C1458; T, falciform 
element, MMMC2686, C1697, outer lateral view; U, falciform element, MMMC2687, C1458, inner lateral 
view. Scale bars are 100 um. 


which are reclined, similar in size, with V- or U-shaped or less equal-sized denticles, and a few small, 
spaces between. It is comparable to the type material rudimentary denticles on the lower edge of the lateral 
from Siberia, except that the Wahringa specimen _ processes. Although Moskalenko (1972) initially 
exhibits a rather straight posterior process with more recognised nine morphotypes for the species, she-later 


154 Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


(Moskalenko 1973) formally described the species as 
a form species. Based on a detailed revision, Ethington 
and Clark (1982) redefined P. flexuosus as consisting 
of a seximembrate apparatus. However, the illustrated 
S elements from Utah display more pronounced 
undulations and twisting of the posterior process and 
size variation of the denticles on the posterior process 
(Ethington and Clark 1982, pl. 9, figs 3, 6) than the 
type material from Siberia (Moskalenko 1973). 


Genus PROTOPANDERODUS Lindstrém 1971 


Type species 
Acontiodus rectus Lindstrom 1955a. 


Protopanderodus cooperi (Sweet and Bergstro6m 
1962) 
Fig. 8A-E 
Synonymy 
Scandodus rectus Lindstr6m 1955a, p. 593, only pl. 
4, figs 22, 23. 
~Acontiodus cooperi Sweet and Bergstr6m 1962, p. 
1221, pl. 168, figs 2, 3, text-fig. 1G. 
Scandodus sp. Sweet and Bergstr6m 1962, p. 1246, 
pl. 168, figs 13, 16. 
Protopanderodus cooperi (Sweet and Bergstrém); 
Zhang 1998b, p. 81, 82, pl. 14, figs 13-17 
(cum syn.). 


Material 
Seven specimens (6 Sa, 1 Sb). 


Discussion 

This species is rare in the Wahringa collections. Two 
morphotypes are recognised as representing the Sa and 
Sb elements, all with sharp anterior and posterior 
margins, and a suberect cusp and one costa on each 
lateral side. The Sa element is symmetrical with a sharp 
median costa (Fig. 8A, B). The Sb element resembles 
the Sa, but is slightly asymmetrical with a more 
strongly developed costa on the inner side (Fig. 8C- 
E). No scandodiform P elements and no laterally 
compressed Sc elements, as characterising P. cooperi 
of previous authors, were recovered. Based on the 
original definition of the species given by Sweet and 
Bergstrom (1962) and more recent revision (Zhang 
1998b), elements of P. cooperi exhibit sharp anterior 
and posterior margins, a well developed anticusp-like 
extension at the antero-basal corner, deep antero-lateral 
recesses in the basal margin, and no more than one 
costa on each lateral face. Protopanderodus cooperi 
can be differentiated from P. rectus (Lindstr6m) in 
having an anticusp-like extension at the anterobasal 
cormer,and from P. varicostatus in displaying no more 


Proc. Linn. Soc. N.S.W., 125, 2004 


than one costa on each lateral side. 

Zhang (1998b) provided a comprehensive 
synonymy list, and illustrated what she recognised as 
P, M, Sa and other undifferentiated S elements; 
however, Zhang provided neither diagnosis nor 
descriptions of the constituent elements of the species 
apparatus of P. cooperi. This species was originally 
proposed as a form species from the Ferry Formation 
of Alabama. The holotype (Sweet and Bergstrom 1962, 
pl. 168, figs 2, 3) is slightly asymmetrical, defined here 
as taking the Sb position. Zhang (1998b) included the 
form species Scandodus sp. Sweet and Bergstr6m 1962 
in the P position of P. cooperi. Based on their 
illustrations and brief discussion, the P element is 


_ inferred to be a scandodiform element with broad costa 


on the inner lateral face and with a smooth outer lateral 
face. Zhang (1998b) also included the holotype of 
Scandodus rectus Lindstr6m 1955a as occupying the 
M position in P. cooperi. This scandodiform element 
is similar to the P element previously defined, except 
that only a broad carina is developed on the inner lateral 
face. For consistency, these two scandodiform 
elements are tentatively taken to represent the Pa and 
Pb positions in Protopanderodus. The symmetrical Sa 
of P. cooperi was illustrated from the Guniutan 
Formation of South China (Zhang 1998b, pl. 14, fig. 
13), and was also recovered from the Wahringa 
samples (Fig. 8A, B). 


Protopanderodus robustus (Hadding 1913) 


Fig. 8J-M 
Synonymy 
Drepanodus robustus Hadding 1913, p. 31, pl. 1, 
fig.5. 


Protopanderodus robustus (Hadding); Lofgren 
1978, p. 94, 95, pl. 3, figs 32-35, text-fig. 
31G-J (cum syn.); An 1987, p. 173, pl. 11, 
figs 7-10 (cum syn.); McCracken 1989, p. 
20-22, pl. 1, figs 1-10, text-fig. 3E (cum 
syn.); Albanesi in Albanesi et al. 1998, p. 
129, 130, pl. 11, figs 17-20, text-fig. 14A 
(cum syn.). 


Material 
Two specimens (Sa, Sc). 


Discussion 

One specimen in the Wahringa collection (Fig. 8L, 
M) which has sharp anterior and posterior margins and 
is laterally compressed with a postero-lateral costa on 
each side, is regarded as representing the Sa element 
of Protopanderodus robustus. The other specimen with 
a single costa on the outer lateral face is referred to the 


155 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Figure 8. A-E, Protopanderodus cooperi (Sweet and Bergstrom 1962); A, B, Sa element, MMMC2688, 
C1683; lateral views; C, upper view, D, outer lateral view, E, inner lateral view, Sb element, MMMC2689, 
C1682. F-I, Protopanderodus nogamii (Lee 1975); F-H, Sa element, MMMC2690, C1450, lateral views; I, 
Sa element, MMMC2691, C1463, lateral view. J-M, Protopanderodus robustus (Hadding 1913); J, outer 
lateral view, K, inner lateral view, Sc element, MMMC2692, C1680; L, M, Sa element, MMMC2693, 
C1458, lateral views. N-X, Protopanderodus varicostatus (Sweet and Bergstrém 1962); N, Pa element, 
MMMC2694, C1652, inner lateral view; O, outer lateral view, P, inner lateral view, Pb element, 
MMMC2695, C1675; Q, R, Sd element, MMMC2696, C1675, lateral views; S, T, Sd element, MMMC2697, 
C1682, lateral views; U, outer lateral view, V, inner lateral view, Sb element, MMMC2698, C1675; W, 
inner lateral view, X, outer lateral view, Sc? element, MMMC2699, C1675. Y, Z, Protopanderodus liripipus 
Kennedy et al. 1979; Sa element, MMMC2700, C1458, lateral views. Scale bars are 100 pm. 


156 Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


Sc element of the same species (Fig. 8J, K). Although 
Lindstr6m (1971), and Johnston and Barnes (2000) 
maintained the original generic assignment of the form 
species Drepanodus robustus Hadding 1913, most 
other authors followed the revision of Lofgren (1978) 
who regarded it as a species of Protopanderodus. 
Lofgren (1978) recognised three morphotypes: 
symmetrical acontiodiform (an Sa element) with a 
postero-lateral costa on each side (Fig. 8L, M), 
asymmetrical acontiodiform (an Sc element) with a 
strong costa on the outer lateral face and a non costate 
inner lateral face (Fig. 8J, K), and scandodiform 
(interpreted here as a P element). Based on this 
multielement species definition, P. robustus is 
morphologically very close to P. cooperi. Specimens 
referable to D. robustus were also recorded from the 
Pratt Ferry Formation of Alabama, where the type 
material of P. cooperi was described (Sweet and 
Bergstr6m 1962). Zhang (1998b) suggested that the 
holotype of D. robustus might be an element of an 
uncertain species of Protopanderodus, but she included 
all the material described from Sweden by Lofgren 
(1978) as P. robustus (Hadding) in her synonymy of 
P. cooperi. This implies that P. cooperi may be a junior 
synonym of P. robustus. Given that the base of the 
holotype of Drepanodus robustus is apparently broken 
(see also Lindstrom 1955b), it remains difficult to 
separate these two species. 


Protopanderodus varicostatus (Sweet and 
Bergstrom 1962) 
Fig. 8N-X 

Synonymy 

Scolopodus varicostatus Sweet and Bergstr6m 1962, 
p. 1247, pl. 168, figs 4-9, text-fig. 1A, C, 
K. 

Scandodus unistriatus Sweet and Bergstr6m 1962, 
p. 1245, pl. 168, fig. 12, text-fig. 1E. 

Protopanderodus varicostatus (Sweet and 
Bergstr6m); Dzik 1976, only text-fig. 16f, 
g; Fahraeus and Hunter 1985b, p. 183, text- 
fig. 2; Bauer 1987, p. 27, pl. 3, figs 19, 21- 
23; An 1987, p. 173, pl. 11, figs 2, 3; Dzik 
1994, p. 74, pl. 14, figs 1-5, text-fig. 11b; 
Zhang 1998b, p. 83, 84, pl. 15, figs 14-19 
(cum syn.). 


Material 
Seven specimens from sample C1675, and one 
specimen from C1652. 


Discussion 

Sweet and Bergstrom (1962) originally recognised 
three form-groups for the species, namely symmetrical, 
slightly asymmetrical, and markedly asymmetrical. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Fahraeus and Hunter (1985b) proposed a 
quinquimembrate apparatus for this species, with ’ 
elements referred to as groups A to E. Group A is a 
symmetrical multi-costate element with two costae on 
each lateral face. Group B is an asymmetrical tri- 
costate element (= the markedly asymmetrical form 
group of Sweet and Bergstr6m 1962) with two costae 
on the inner lateral face and a postero-lateral costa on 
the outer lateral face. Group C is an asymmetrical 
multi-costate element (= slightly asymmetrical form 
group of Sweet and Bergstrom 1962) with a twisted 
cusp and two costae on each side. Group D is a tri- 
costate element, similar to group B but less laterally 
compressed with costa on the outer lateral face situated 
more towards the middle. Group E is a scandodiform 
element represented by the form species Scandodus 
unistriatus Sweet and Bergstrom 1962 (here assigned 
to the Pb position). 

Zhang (1998b) illustrated one of the multi- 
costate specimens as the P element, and two 
morphologically different scandodiform specimens as 
the M element. One of the latter specimens (Zhang 
1998b, pl. 15, fig. 19) is comparable with the form 
species S. unistriatus Sweet and Bergstrom 1962, and 
is regarded here as representing the Pb element of P. 
varicostatus (Fig. 80, P). The other specimen 
illustrated as the M element (Zhang 1998b, pl. 15, fig. 
14), which was recovered from the same sample with 
other illustrated specimens of P. varicostatus, has a 
multi-costate inner lateral face with three costae 
bordering two grooves and a few, shorter and weaker 
secondary costae near the base. It is designated here 
as occupying the Pa position of the species. 

Similar specimens (arcuatiform) referrable to 
the Pa element of P. varicostatus were also reported 
from allochthonous limestone clasts within the Shinnel 
Formation of Scotland (Armstrong 1997, pl. 3, figs 3, 
4). Morphologically it resembles the Pa element of 
Protopanderodus cf. calceatus Bagnoli and Stouge 
1996, recovered from the allochthonous limestones in 
the Oakdale Formation of central New South Wales 
(Zhen and Percival in press, pl. 17, figs A, C). 
However, this latter element has one larger, open 
groove on the inner lateral face, while the Pa element 
of P. varicostatus from South China (Zhang 1998b, 
pl. 15, fig. 14) and from the Wahringa area (Fig. 8N) 
has two equally developed, narrower grooves with a 
sharp costa between them. Protopanderodus liripipus 
Kennedy et al. 1979 is also multi-costate, but with a 
more posteriorly extended base (Fig. 8Y, Z). 

Three morphotypes of multicostate (S) 
elements with two costae on each side are recognised 
from sample C1675 and possibly should be assigned 
to the Sd, Sb and Sc? positions, as no tri-costate 
elements have been recovered. The Sd element is 


7 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


symmetrical with a reclined cusp and a short base. The 
Sb element is weakly asymmetrical with suberect cusp 
and a longer base. The Sc? element is asymmetrical 
and laterally strongly compressed with a suberect cusp 
and a short base. 


Genus PYGODUS Lamont and Lindstr6m 1957 


Type species 
Pygodus anserinus Lamont and Lindstrom 1957. 


Pygodus protoanserinus Zhang 1998a 
Fig. 9B-J 

Synonymy 

Pygodus anserinus Lamont and Lindstrom 1957, p. 
68, only fig. 1d. 

Pygodus serrus (Hadding); Bergstrom 1971, p. 149, 
pl. 2, figs 22, 23; An 1981, pl. 4, figs 1-3; 
An 1987, pl. 24, fig. 25, pl. 26, figs 3, 6, 
13, pl. 29, figs 2, 3; Nicoll 1980, fig. 3H-L; 
Ding et al. in Wang, 1993, p. 198, pl. 30, 
figs 10, 13, 15-18, 20-22, 24, pl. 35, 24, 26. 

Pygodus protoanserinus Zhang 1998a, p. 96, Fig. 
2D, pl. 3, figs 9-18 (cum syn.); Zhang 
1998b, p. 86, 87, pl. 16, figs 6-8 (cum 
syn. ). 

Pygodus serra (Hadding); Percival et al. 1999, fig. 
8.18; Pickett and Percival 2001, fig. 4C. 


Material 
Four Pa (stelliscaphate), five Pb (pastinate), and one 
Sb (tertiopedate) elements. 


Discussion 

Five species were assigned to Pygodus in the recent 
study of the genus by Zhang (1998a). They have short 
stratigraphic ranges and hence are very useful 
biostratigraphic index fossils. Sweet and Bergstrom 
(1962) and Bergstrém (1971) initially suggested that 
the apparatus of Pygodus anserinus, the type species 
of the genus, included elements represented by the 
form species Pygodus anserinus Lamont and 
Lindstr6m 1957, and Haddingodus serrus (Hadding). 
Bergstr6m (1971) also raised the possibility that the 
Pygodus apparatus might include elements represented 
by the form species Tetraprioniodus lindstroemi Sweet 
and Bergstrom 1962 and Roundya pyramidalis Sweet 
and Bergstrom 1962. This quadrimembrate Pygodus 
apparatus composition has been widely accepted 
(Lofgren 1978, Clark et al. 1981, Sweet 1988). 
Subsequently, Armstrong (1997) has implied a 
septimembrate apparatus for Pygodus, but with only 
confirmed elements occupying the Pa, Pb, Pc, M and 
Sc positions. By including two pygodiform elements 
in the apparatus, Armstrong (1997) suggested that the 


158 


P. anserinus apparatus consisted of the stelliscaphate 
Pa, pastiniscaphate Pb, bipennate Pc (= pastinate Pb 
of other authors’ usage - see Zhang 1998a, b), 
tertiopedate M (termed an S element by other authors 
- see Zhang 1998a, b, and herein), and the ramiform 
Sc element. More recently Zhang (1998a, 1998b) 
proposed a quinquimembrate apparatus for Pygodus, 
including stelliscaphate Pa, pastinate Pb, and three 
ramiform S elements (alate, tertiopedate and 
quadriramate). 

Distinctions between P. serra and P. 
protoanserinus were discussed in detail by Zhang 
(1998a). Pygodus protoanserinus ranges from the E. 
robustus Subzone to the E. lindstroemi Subzone of the 
upper serra Zone in Baltoscandia, Scotland, North 
America, China, and Australia. The stelliscaphate Pa 
element from the lower part of the Wahringa Limestone 
Member is identical with the type material of P. 
protoanserinus, being characterised by having the 
middle denticle row situated more towards the outer 
denticle row on the upper surface. Specimens 
illustrated by Nicoll (1980) as P. serrus from the 
Pittman Formation at Black Mountain, Canberra, ACT, 
are here reassigned to P. protoanserinus on this same 
basis. A single specimen from the lower part of the 
Wahringa Limestone Member has the middle row of 
the denticles positioned centrally, and is therefore 
referred to P. serra (Fig. 9K, L), being more 
comparable with the middle form of the Pa element of 
that species as defined by Zhang (1998a). 


Genus STIPTOGNATHUS Ethington, Lehnert, and 
Repetski 2000 


Type species 
Reutterodus borealis Repetski 1982. 


Stiptognathus sp. A 


Fig. 90, P 
Synonymy 
Stiptognathus sp. Zhen and Percival in press, fig. 
21L-O. 
Material 


Two specimens from sample C1463 from an 
allochthonous limestone within the Fairbridge 
Volcanics, stratigraphically below the Wahringa 
Limestone Member. 


Discussion 

The symmetrical Sa and geniculate M elements 
recovered from sample C1463 are identical with those 
from the allochthonous limestones of the Oakdale 
Formation (Zhen and Percival in press). Denticles on 

these specimens are small, closely spaced and blunt. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Y.Y. ZHEN, I.G. PERCIVAL AND B.D. WEBBY 


Figure 9. A, Pseudooneotodus mitratus (Moskalenko 1973); upper view, MMMC2701, C1474. B-J, Pygodus 
protoanserinus Zhang 1998a; B, Pa element, upper view, MMMC2702, C1450; C, Pa element, upper 
view, MMMC2703, C1652; D, upper view, and E, enlargement showing surface structure, Pa element, 
MMMC2704, C1652; F, inner lateral view, G, outer lateral view, H, anterior view, Pb element, 
MMMC2705, C1652; I, outer lateral view, J, showing surface structure, Sb element, MMMC2706, C1652. 
K, L, Pygodus serra (Hadding 1913); K, upper view, L, basal view, Pa element, MMMC2707, C1652. M, 
N, Stiptognathus sp. B; M, antero-lateral view, N, posterior view, Sa element, MMMC2708, C1675. O, P, 
Stiptognathus sp. A; O, M element, MMMC2709, C1463, posterior view; P, Sa element MMMC2710, 
C1463, antero-lateral view. Unless otherwise indicated scale bars are 100 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 159 


ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


Stiptognathus sp. B 
Fig. 9M, N 
Material 
Two specimens (Sa) from sample C1675. 


Discussion 

The cusp of this symmetrical element is triangular in 
cross section with a gently curved, wide anterior face, 
posterior costa, and an antero-lateral costa on each side. 
Three sharp costae extend basally into three blade-like 
processes, which bear small, upward-pointed denticles 
along the edges. They are easily distinguishable from 
the blunt denticles of Stiptognathus sp. A. 


ACKNOWLEDGMENTS 


This study was supported by a Science Fellowship 
provided by the Sydney Grammar School to Zhen. Initial 
field collecting was undertaken with the support of the 
Australian Research Council during 1996 to 1999 (grant 
A39600788 to B.D. Webby). Gary Dargan from the 
Geological Survey of N.S.W. assisted in acid leaching, 
separation and other laboratory work. A grant to Y.Y. Zhen 
from the Betty Mayne Scientific Research Fund of the 
Linnean Society of New South Wales defrayed costs of some 
of the SEM work. The scanning electron microscope 
illustrations were prepared in the Electron Microscope Unit 
of the Division of Life and Environmental Sciences, 
Macquarie University and in the Electron Microscope Unit 
of the Australian Museum. I.G. Percival publishes with the 
permission of the Director General, New South Wales 
Department of Mineral Resources. 


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ORDOVICIAN CONODONT FAUNAS FROM CENTRAL N.S.W. 


APPENDIX 
LOCALITY DATA 


All grid references are AMG66 co-ordinates and relate to the Cumnock 8632-S 1:50,000 topographic sheet 
(first ed., 1978). 


Allochthonous limestones stratigraphically below Wahringa Limestone Member 
C1463: GR 679150 mE 6371900 mN 
C1486: GR 678750 mE 6371800 mN 
C1487: GR 678720 mE 6371800 mN 
C1488: GR 678700 mE 6371800 mN 


Wahringa Limestone Member, type section 
C1450, C1652 (basal beds): GR 678650 mE 6372000 mN 
C1456, C1664, C1667-C1668, C1672 (middle beds): GR 678700 mE 6372050 mN 
C1458, C1464, C1673-1683, C1687 (upper beds): GR 678700 mE 6372100 mN 


Wahringa Limestone Member, northeast extremity of outcrop 
C1707, C1709-1713: centred on GR 679330 mE 6372620 mN 


Wahringa Limestone Member, southwest extremity of outcrop (middle or upper beds) 
C1429: GR 677570 mE 6370800 mN 


Allochthonous limestones stratigraphically above Wahringa Limestone Member 
C1693, C1694: GR 678000 mE 6371310 mN 
C1695, C1696: GR 678100 mE 6372000 mN 
C1697, C1698: GR 678050 mE 6372000 mN 
C1699: GR 678025 mE 6372000 mN 
C1700: GR 678010 mE 6372000 mN 
C1471: GR 678500 mE 6372200 mN 
C1472: GR 679250 mE 6373550 mN 
C1474: GR 678100 mE 6372600 mN 


C1483: GR 678500 mE 6372800 mN 


164 Proc. Linn. Soc. N.S.W., 125, 2004 


Wenlock (Early Silurian) Brachiopods from the Orange District 
of New South Wales 


A.J. Wricut! and D.L. Strusz? 


‘School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522, 
tony_wright @uow.edu.au; *Department of Earth and Marine Sciences, Australian National University, 
Canberra ACT 0200; dstrusz@ems.anu.edu.au 


Wright, A.J. and Strusz, D.L. (2003). Wenlock (Early Silurian) brachiopods from the Orange District of 
New South Wales. Proceedings of the Linnean Society of New South Wales 125, 165-172. 


Two late Wenlock (Early Silurian) brachiopod species from the Ulah Formation near Orange, New 
South Wales, are closely associated with graptolite faunas. Visbyella cumnockensis occurs in the testis 
Biozone on Wallace Creek in the Four Mile Creek area, and Strophochonetes melbournensis is recorded 
from the Judensis Biozone on Spring Creek. Poorly preserved but similar Visbyella? and Strophochonetes? 
From the Pridoli Wallace Shale at Cheesemans Creek are also illustrated. These occurrences provide significant 
new stratigraphic and distributional data for the species. 


Manuscript received 19 March 2003, accepted for publication 22 October 2003. 


KEYWORDS: Brachiopods, New South Wales, Pgidoli, Silurian, Strophochonetes melbournensis, Ulah 
Formation, Visbyella cumnockensis, Wallace Shale, Wenlock. 


INTRODUCTION 


The Silurian strata of the area west and 
southwest of Orange, NSW, in the valleys of Spring 
Creek and Four Mile Creek (Fig. 1), have yielded a 
diversity of fossils, but very few shelly fossils have 
ever been described, apart from corals described by 
authors including Etheridge and McLean (full 
references to these works can be found in Pickett 1982). 
The most abundant and important fossils in the region 
are graptolites, which have been known for more than 
50 years and were reported by Packham and Stevens 
(1955) and Jenkins (1978, 1986). 

Jenkins recorded (but did not describe) 
brachiopod faunas from limestones in the Four Mile 
Creek area, but few brachiopods have been reported 
from clastic strata common in the area. Rickards and 
Wright (1997) described two brachiopod species from 
late Wenlock strata (Judensis Biozone) in Cobblers 
Creek (Fig. 1), and in the section at ‘Mirrabooka Park’ 
brachiopods were noted in Wenlock strata during field 
work by L. Muir, R.B. Rickards, G.H. Packham and 
A.J. Wright. A diverse and abundant shelly fauna 
occurring with the late Wenlock graptolite 


Testograptus testis on the Cadia gold mine access road, 
several kilometres to the east of Four Mile Creek, was 
illustrated by Rickards et al. (2001). 

The two species described here are recorded 
for the first time from the region near Orange. One, 
Visbyella cumnockensis Walmsley et al., 1968, was 
originally described from near Cumnock, 55 km 
northwest of Orange, where it occurs with T. testis 
(Walmsley et al. 1968:315). Visbyella has been 
reported also, but not illustrated, by Pickett (1982) and 
Pogson and Watkins (1998). The other species, 
Strophochonetes melbournensis (Chapman 1903), was 
previously known only from Wenlock and Ludlow 
strata in the Melbourne Trough, Victoria. Pickett’s 
report was based on the record of Visbyella cf. 
cumnockensis by Sherwin (1971). Sherwin’s locality 
is younger, and contains a sparse and poorly preserved 
fauna including also a chonetoide similar to 
Strophochonetes? savagei Strusz, 2000 from 
Cumnock. These taxa are illustrated but not described. 
Documented brachiopod occurrences in the Orange 
region are still insufficient, however, to permit any 
notion of a regional brachiopod zonation. 


EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 


to Wellington 


Borenore™ 


:gaMirrabooke Park’, 


Spring Creek - Mt. Canobolas 
tens Creek area A 


148° 45° 


Figure 1. Map of the area southwest to west of Orange, central 


Wenlock to Pridoli; the age of the strata 
at this locality is late Wenlock. 


W940. 

The somewhat more abundant 
specimens of Strophochonetes 
melbournensis were collected from 
dark siltstones of the Ulah Formation 
on the southern side of Spring Creek 
at ‘Mirrabooka Park’, directly opposite 
One Tree Hill. There are also 
occasional poorly preserved 
brachiopods, including pentamerides, 
in beds at about the same level on One 
Tree Hill itself. The shells at W940 
occur with a graptolite fauna that 
includes Monograptus ludensis (R.B. 
Rickards, pers. comm.). Only 
disarticulated valves are known at this 
locality; small phosphatic brachiopods 
are quite common, and there are rare 
specimens of other brachiopods 
including strophomenides and 
atrypides. Most specimens of 
Strophochonetes melbournensis at this 
locality retain shelly material and the 
spines on the pedicle valve hinge line 
are often preserved. The environment 
was most probably a low-energy one. 


~ 
o 
3 
<= 
= 
Gy 
o 
o 
= 


New South Wales, showing the geographic context of the two 


localities, LM3 on Wallace Creek east of Cargo and W940 near 
‘Mirrabooka Park’ east-southeast of Cudal. Inset: the location 


of Orange within Australia. 


LOCALITIES 


LMG. 

Visbyella cumnockensis was collected from 
Wallace Creek in the Four Mile Creek area, in grey- 
brown siltstones assigned by Jenkins (1978) to the 
Wenlock-Ludlow Ulah Formation. These beds have 
also yielded the graptolites Cyrtograptus and a new 
species of Monograptus (L. Muir, pers. comm.), and 
overlie beds containing T. testis. The brachiopod 
specimens are moulds of a single pedicle and a single 
brachial valve on the same bedding surface, which 
could represent the disarticulated valves of a single 
shell. No other macrofossils have been found at this 
locality. In contrast, the type material of V. 
cumnockensis is entirely of specimens in the “butterfly” 
position, with the shell opened so that the conjoined 
valves lie on the bedding surface. The age assigned to 
the Ulah Formation by Chapman et al. (2003) is late 


166 


MO/1/27. 

A few poorly preserved orthide 
and chonetoide specimens have been 
collected from this outcrop of fine thin- 
bedded siltstone low in the Wallace 
Shale, about 600 m east of ‘Mirrabooka’ homestead. 
The fauna also includes occasional trilobites. The lo- 
cality lies within the Monograptus transgrediens 
Biozone. 


SYSTEMATIC PALAEONTOLOGY 


Suprageneric taxonomy follows that in Kaesler 
(2000); references to authorship of suprageneric taxa 
are therefore not repeated here. Specific diagnoses have 
been rephrased to accord with currently accepted 
terminology (Kaesler 1997). Details of localities are 
given in the descriptive section below. 


Abbreviations. 
Ls - shell length 
Ld - dorsal valve length. 
Ws - shell width 
Wh - hinge line width 


Proc. Linn. Soc. N.S.W., 125, 2004 


A.J. WRIGHT AND D.J. STRUSZ 


Figure 2. a-g, Visbyella cumnockensis Walmsley et 
al., 1968. a-c, ventral valve counterparts; a, latex 
cast of exterior, AM F124331. b-c, internal mould 
and latex cast, AM F124332. d-g, dorsal valve 
counterparts; d, latex cast of exterior, AM F124333. 
e-g, internal mould and latex cast (in ventral and 
postero-ventral views), AM F124334. h-k, cf. 
Visbyella cumnockensis, Pridoli, Wallace Shale. h, 
latex cast of ventral valve, MM F37431. i, latex cast 
of shell in ‘butterfly’ position, MM F21132. j, 
external mould of dorsal valve plus internal mould 
of ventral valve, MM F21125. k, latex cast of 
incomplete interior of shell in ‘butterfly’ position, 
MM F37428. Scale bar 2 mm. 


AM - Australian Museum 

MM -— Mining Museum Collection, 
Geological Survey of NSW 

CPC - Commonwealth Palaeontological 
Collection 

NMV - Museum of Victoria 

SU - Sydney University (Geology 
Department) 


Suborder DALMANELLIDINA Moore 1952 
Superfamily DALMANELLOIDEA Schuchert 1913 
Family DALMANELLIDAE Schuchert 1913 
Subfamily RESSERELLINAE Walmsley and 
Boucot 1971 

Genus VISBYELLA Walmsley, Boucot, Harper and 
Savage 1968 


Type species 
Orthis visbyensis Lindstr6m 1861, by original 
designation; late Llandovery, Gotland. 


Diagnosis 

Subcircular, small valves with apical deltidium 
and hypercline dorsal interarea; ventral interior with 
recessive dental plates and cordate muscle scar; dorsal 
interior with trilobed, dorsally-facing cardinal process 
and median septum (Harper p. 797 in Kaesler 2000). 


Visbyella cumnockensis Walmsley, 
Boucot, Harper and Savage 1968 
Fig. 2 (a-g) 


Synonymy 


1968 Visbyella cumnockensis sp. nov.; 
Walmsley et al., pp. 313-315, pl. 61 figs 6-12. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Type material 

Holotype AM F67781; paratypes AM F67782- 
67788 (formerly SU P19511, 19512-19518; all 
renumbered when collections were transferred from 
the University of Sydney to the Australian Museum). 


New material 
External and internal moulds of a ventral valve 
(AM F124331, F124332) and a dorsal valve (AM 


167 


EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 


F 124333, F124334) from one bedding plane at locality 
LM3 (Grid reference 782 988, Cudal 8631 II and III 
50 000 topographic sheet, Wallace Creek, Four Mile 
Creek area south of Orange, N.S.W.); Ulah Formation, 
Testograptus testis Biozone; late Wenlock (Early 
Silurian). 


Diagnosis 

Relatively small, weakly sulcate, coarsely 
multicostellate Visbyella with semicircular outline. 
Dorsal median ridge broad and low posteriorly, 
becoming narrower and higher to form an anterior 
median septum (after Walmsley et al. 1968) 


Description 

Shell small, almost plano-convex. Ventral valve 
broadly naviculate, with low suberect beak; dorsal 
valve weakly convex with shallow but distinct sulcus. 
Outline suboval, moderately transverse, with straight 
hinge, obtuse slightly rounded cardinal angles; greatest 
width at about 0.4Ls. Ventral interarea strongly 
apsacline, almost flat, apical angle about 120°; 
delthyrium open, apical angle about 70°, rimmed by 
narrow crescentic deltidium. Dorsal interarea low, 
concave, catacline, apical angle about 150°; 
notothyrium filled by cardinal process, apical angle 
about 80°. Ribs rather angular, stronger medially than 
laterally, increasing by bifurcation on the ventral valve, 
intercalation on the dorsal valve; about 30 counted at 
ventral valve margin. 

Ventral interior with prominent subtriangular 
muscle field, impressed posteriorly but slightly raised 
anteriorly, length 1/3Ls and width 1/4Ws. Diductor 
scars elongate oval, divergent, depressed a little below 
slightly shorter flat adductor field. Raised anterior 
margin to adductor field distinctly denticulate, extends 
forward to about 3/4Ls as low ridge. Vascula media 
flank this ridge as broad, shallow furrows extending 
from the diductor scars. Muscle field flanked by stout 
dental plates, divergent forward at about 100° and 
slightly divergent ventrally, not extending beyond 
muscle field. Teeth strong, wide, triangular, with 
distinct crural fossettes on antero-median faces. Valve 
floor faintly radially furrowed, marginally strongly 
crenulated. 

Dorsal interior with prominent oval muscle 
field extending to 2/3Ld, width 1/3Ws, defined by 
strong ridges arising just in front of brachiophores and 
increasingly raised anteriorly, which converge to abut 
on median septum. Diductor scars impressed, elongate 
oval, subequal, posterior scars subparallel, anterior 
scars convergent forward; scars separated by tapering 
ridge from which rises the stout median septum. 
Septum highest a little in front of muscle field, and 
extends to valve margin. Cardinal process large, 


168 


directed posterodorsally, continuous with well 
developed notothyrial platform; no_ shaft. 
Brachiophores stout, blade-like, divergent ventrally, 
supported by low, thick plates. Sockets oval, diverging 
from valve axis at about 75°, deeply excavated into 
thick triangular socket pads. Valve floor radially 
grooved, marginally strongly crenulated. 


Dimensions 
AM F124332 AM F124334 

valve ventral dorsal 

Ls, Ld est 2.85 2.59 

Ws 3.90* 3.73 

Wh 3.60* BB2 

Ls/Ws est. 0.73 0.69 

Wh/Ws est. 0.92 0.89 


* values obtained by doubling exposed half-width, 
assuming a symmetrical shell. 


Remarks 

The Wallace Creek occurrence of this species 
is almost exactly the same age as the original 
occurrence at Cumnock, and our admittedly limited 
new material corresponds closely in all specific 
characters to the type material. The specimens are 
slightly larger than shells of the type series (the 
maximum length and width of any specimens of the 
type series are 2.1 mm and 3.1 mm respectively), but 
the ratio Ls:Ws is close to the 2:3 cited for the type 
material; while the marginal crenulations in the ventral 
valve are less extensive. The internal moulds of the 
disarticulated valves are somewhat better than the 
types, and features of the hinge line can be seen more 
clearly. 

The species was also tentatively recorded by 
Sherwin (1971, p. 223) from the Pridoli Wallace Shale 
at locality MO/I/27 in the Cheesemans Creek area north 
of Quarry Creek; his report was the basis for 
subsequent reports by Pickett (1982, pp. 154-155) and 
Pogson and Watkins (1998, p. 131). This occurrence 
is in significantly younger strata than the two other 
occurrences noted herein. Sherwin’s report was based 
on several specimens from one locality; we were 
recently guided to this locality by Dr Sherwin, and 
collected a further seven specimens of the ‘orthid’ 
species, which is very rare at the locality (also collected 
were a few poor specimens of a chonetide, identified 
as Strophochonetes? cf. savagei Strusz, 2000, and 
illustrated in Fig. 4 for comparison with 
Strophochonetes melbournensis). 

Unfortunately the only internal mould of a 
dorsal valve of the Wallace Shale orthide (Fig. 2k) is 
incomplete, and appears to lack a median septum, 
although its presence anteriorly cannot be completely 
ruled out. It was initially thought that the absence of a 


Proc. Linn. Soc. N.S.W., 125, 2004 


A.J. WRIGHT AND D.J. STRUSZ 


septum would rule out the presence of Visbyella. 
However, one specimen (AM F125552) of Visbyella 
cumnockensis on one of the type slabs is close in size 
to the Wallace Shale material and, unlike all the other 
type specimens, lacks a median septum, so this is not 
an infallible character of this species. Other 
morphological features of the Wallace Shale material 
are not well preserved; there appear to be more than 
30 costellae, and the internals of both valves, in so far 
as they are preserved, are similar to those of the 
Wallace Creek material (compare Figs 2h-i with Fig. 
2a, and Fig. 2j with Fig. 2b). 

Hence no conclusive argument can be presented 
to refute the presence of Visbyella at this locality, 
unlikely as it might seem. This opinion is slightly 
supported by the presence of a similar orthide 
(probably Resserella), but definitely lacking a median 
septum, in the late Ludlow Cardinal View Shale (Bauer 
1994) at Bungonia, NSW. Unfortunately, our 
experience gives us no reason to expect more definitive 
material at this very unproductive Wallace Shale 
locality. 


Suborder CHONETIDINA Muir-Wood 1955 

Superfamily CHONETOIDEA Bronn 1862 

Family STROPHOCHONETIDAE Muir-Wood 1962 

Subfamily STROPHOCHONETINAE Muir-Wood 
1962 

Genus STROPHOCHONETES Muir-Wood 1962 


Type species 
Chonetes cingulatus Lindstrom 1861, by 
original designation; Wenlock, Gotland. 


Diagnosis 

Shell small, plano- to moderately concavo- 
convex; well developed median enlarged costa; long, 
symmetrically arranged high-angled spines varying 
from intraverse cyrtomorph proximally to orthomorph 
vertical distally; cardinal process strongly bilobed 
internally, anteriorly bounded by cardinal process pit; 
no median septum; anderidia long, narrow, anteriorly 
divergent at about 60° and isolated on valve floor; inner 
socket ridges short, thin, as two rounded ridges almost 
parallel to hinge (after Racheboeuf p. 369 in Kaesler 
2000). 


Strophochonetes melbournensis (Chapman 1903) 
Fig. 3 
Synonymy 


1903 Chonetes melbournensis sp. nov.; 
Chapman, pp. 74-76, pl. XI, fig. 2 only. 


Proc. Linn. Soc. N.S.W., 125, 2004 


1945 Chonetes (Chonetes) melbournensis 
Chapman; Gill, pp. 132-133. 

1953 Chonetes infantilis n. sp.; Opik; p. 15, 
pl. UI, figs 19-22. 

2000 Strophochonetes melbournensis 
(Chapman, 1903); Strusz, pp. 249-251, figs 
2-3. 


Type material 

Lectotype NMV P1419, paralectotypes NMV 
P615-6, 619, 623, 625-7, 630-633, 637-43 designated 
by Strusz (2000); Melbourne Formation, Melbourne 
and South Yarra, Victoria; Ludlow (Late Silurian). 
Type material of Chonetes infantilis Opik, 1953: 


_ holotype CPC 661, paratypes CPC 662-663, I/laenus 


Band, Wapentake Formation, Heathcote, Victoria; late 
Wenlock (Early Silurian). 


New material 

AM F124306 - 124330, locality W940 (grid 
reference 743 123, Cudal 8631-II and III 50 000 
topographic sheet; south bank of Spring Creek, 
‘Mirrabooka Park’, southwest of Orange, central 
N.S.W.); Ulah Formation, with Monograptus ludensis; 
Late Wenlock (Early Silurian). 


Diagnosis 

Small, weakly concavo-convex, subquadrate 
Strophochonetes with up to 5 pairs of gently intraverse- 
cyrtomorph hinge spines, and finely multicostellate 
ornament with median rib on ventral valve usually 
strongly enlarged. Valve floors heavily papillose, 
ventral muscle field distinct, anderidia short and 
diverging at 60-80 (after Strusz 2000). 


Description 

Shell small, plano-convex, ventral valve of very 
low convexity. Outline subquadrate, lateral margins 
gently sigmoid, with shallow re-entrants in front of 
small triangular ears; hinge width usually slightly less 
than greatest width (mean Wh/Ws 0.93). Ventral 
protegulum posteromedially furrowed, variably raised 
above remaining shell surface; distinct protegular lobe, 
weaker lateral lobes on dorsal valve. Maximum 
observed width 9.8 mm, length 6.5 mm, most 
specimens being much smaller; mean Ls/Ws 0.75, ratio 
decreasing with increasing shell size. Interareas mostly 
obscure; ventral interarea apparently low, apsacline, 
flat, delthyrium wide, beak very low; pseudodeltidium 
not seen; dorsal interarea linear, attitude unclear. 
Myophore small, projecting posteroventrally, bifid, 
each lobe less strongly bifid, flanked by small but 
distinct cardinal crests. Chilidium obscure, might be 
present as very narrow ridge wrapped around base of 
myophore. Hinge spines fine, relatively long, upright 


169 


EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 


Figure 3. Strophochonetes melbournensis (Chapman, 1903). a-g, ventral valves; some hinge spines are 
visible in b-e, only spine bases in f-g. a, juvenile valve AM F 124320. b, juvenile with particularly prominent 
protegulum, AM F124317. c, juvenile AM F124322. d, AM F124324. e-f, internal mould and latex cast, 
AM F124312. g, AM F124326. h-j, dorsal valves; h-i, incomplete external mould and latex cast showing 
well developed protegular and lateral nodes, AM F 124318. j, latex cast of incomplete interior, AM F124307. 


Scale bar 3 mm. 


or nearly so (initial angle with hinge line about 60- 
80°), straight (particularly in small specimens) to gently 
cyrtomorph intraverse, symmetrically placed; up to 4 
each side of beak (AM F124324). Ornament of fine, 
rounded radial ribs, 29-34 counted in 5 mm at 5 mm 
radius, separated by narrower furrows; increase is by 
bifurcation only. Median rib on ventral valve 
prominent, arises within protegulum; remaining ribs 
arise at or in front of margins of concentrically 
wrinkled protegular regions. 

Ventral interior with low, narrow median 
septum, reaching forward to about 0.2Ls; septum 
posteriorly raised and slightly widened. Teeth small, 
widely divergent. Muscle field generally obscure other 
than for weak or absent endospines; in one specimen 
(AM F124312) the field is weakly impressed, with 


170 


small, elongate subtriangular, slightly divergent 
adductor scars further impressed posteriorly. 
Remainder of valve floor densely covered by fine 
endospines radially arranged beneath ribs, weakest 
towards cardinal margin and ears. 

Dorsal interior still not well known. Cardinal 
process small, internally bifid, fused to short but strong 
inner socket ridges which are curved parallel to hinge 
margin. Short, shallow furrow in front of cardinal 
process, but no median ridge developed. Anderidia 
visible in only one specimen (AM F124307); they are 
short (0.2Ld), fine, low, diverging at about 60°. Muscle 
field obscure. Distal two-thirds of valve floor with 
numerous small radially arrayed endospines, as in 
ventral valve. . 


Proc. Linn. Soc. N.S.W., 125, 2004 


A.J. WRIGHT AND D.J. STRUSZ 


Dimensions 
valve Ls, Ld Ws 
AM F124307 dorsal 4.9 = 
F124312 ventral est. 4.8 5.6* 
F124318 dorsal 5.5 — 
F124322 ventral 3.6 47 
F124324 ventral 5.3 WP 
F124326 ventral 5.6 8.4 


Wh _ = Ls/Ws Wh/Ws 
Sain = - 

5.4* est. 0.86 0.96 
44 0.77 0.94 
V2 Qg4! 1.00 
7.2 0.67 0.86 


* values obtained by doubling exposed half-width, assuming a symmetrical shell. 


Discussion 

Although preservation is not particularly good, 
the Wenlock specimens from Spring Creek conform 
in all important aspects (very low ventral convexity, 
rib increase only by bifurcation, and less prominent 
protegular and lateral lobes on the dorsal valve) with 
S. melbournensis rather than S. kemezysi Strusz, 2000. 
Some of the minor differences could be related to the 
small size of most of the specimens (several are clearly 
juvenile, none approaches the maximum size recorded 
for the Victorian material). Some could be of age 
significance, but without better and more abundant 
material from older levels in Victoria this remains 
unclear. Thus no ventral valves show the anterior 
sulcus seen in some Victorian Late Silurian shells, and 
no more than 4 spines have been seen to either side of 
the ventral beak. The NSW specimens tend also to be 
more elongate (Ls/Ws very variable, mean 0.76; for 
Victorian specimens the mean is 0.61). Internally, the 
ventral muscle field is less obvious, and there are no 
coarser endospines near the hinge. In this last respect, 
and in a greater tendency for spines on small specimens 
to be straight, the Late Wenlock Spring Creek 
specimens are more like the few poor specimens from 
the Early Wenlock of Heathcote than the Ludlow 
material from Melbourne. Dorsal interiors, while still 
few and inadequate, do add some information, 


Figure 4. Strophochonetes? cf. savagei Strusz, 2000. 
Latex cast of ventral valve, MM F21133. Scale bar 
3 mm. 


Proc. Linn. Soc. N.S.W., 125, 2004 


particularly the form of the cardinal process and its 
flanking cardinal crests. The presence internally of a 
weak posteromedian dorsal furrow instead of a low 
ridge places these specimens closer to typical 
Strophochonetes than are the type specimens. 

Three similar chonetoide specimens (MM 
F21133, 37435, 37436) are available from the Wallace 
Shale locality - the best of them is figured (Fig. 4). All 
are small and weakly convex. In the absence of internal 
data, particularly of the dorsal valve, generic identity 
must remain uncertain. The long more or less upright 
hinge spines, low ventral valve convexity, fine ribbing 
and accentuated median rib all indicate 
Strophochonetes, however, and of the Australian taxa 
described by Strusz (2000) the closest is undoubtedly 
S? savagei from the Early Lochkovian of Manildra, 
northwest of Orange. S. melbournensis and S. kemezysi 
Strusz, 2000, while superficially similar, are both larger 
and more coarsely ribbed; the latter has very prominent 
protegulae. In only one respect these specimens appear 
unlike typical Strophochonetes, and that is in the 
alternating pattern of hinge spine insertion described 
for instance by Strusz (2000, p. 259) for the strongly 
convex and fairly coarsely ribbed Australian species 
of Johnsonetes Racheboeuf, 1987 (all of which lack 
spine 1’). However it is not clear that spine 1' is 
undeveloped in the Wallace Shale specimens. 
Moreover, the Manildra species show considerable 
variation in spine form, and some asymmetry cannot 
be ruled out. 


ACKNOWLEDGMENTS 


We gratefully acknowledge access graciously made 
available by Ian Street to ‘Mirrabooka Park’ and Ken 
Williams to ‘Ashburnia’, and thank Dr L. Sherwin for 
drawing our attention to the report of Visbyella cf. 
cumnockensis from the Wallace Shale and subsequently 
guiding us to the locality. Prof. Barrie Rickards and Dr Lucy 
Muir kindly allowed us to cite identifications of the 
graptolites. Robert Jones readily made the type material of 
Visbyella cumnockensis available for study. Strusz wishes 
to thank Dr Patrick DeDeckker for providing facilities at the 


171 


EARLY SILURIAN BRACHIOPODS FROM ORANGE NSW 


Australian National University; Wright’s research has been 
supported by the University of Wollongong and the Linnean 
Society of NSW. 


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Proc. Linn. Soc. N.S.W., 125, 2004 


Early Silurian Graptolites from Cadia, New South Wales 


R.B. Rickarps! AND A.J. WricurT? 


‘Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, England 
and *School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522. 


Rickards, R.B. and Wright, A.J. (2004). Early Silurian graptolites from Cadia, New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 173-175. 


A low-diversity graptolite fauna is reported from the Ulah Formation at Cadia, central western New South 
Wales. The assemblage includes Testograptus testis, Monoclimacis flumendosae, fragments of Monograptus 
flemingii, possible Cyrtograptus and unidentifiable retiolitid meshworks, and is correlated with the Jungreni- 


testis Biozone, of late Wenlock (Early Silurian) age. 


Manuscript received 3 May 2003, accepted for publication 23 July 2003. 


KEY WORDS: Cadia, graptolites, Lower Silurian, Wenlock. 


INTRODUCTION 


Three Silurian faunas were documented by 
Rickards et al. (2000) from the vicinity of Cadia open 
cut, south of Orange, New South Wales. One of these 
faunas, of late Wenlock-early Ludlow aspect, consisted 
of shelly fossils and graptolites collected by Dr Ian 
Percival from a slumped mudstone at a locality on the 
access road to the Cadia open cut. This fauna was 
discussed and illustrated by Rickards et al. (2000), who 
figured but could not determine the poor graptolite 
material to genus or species because of the poor 
preservation of the fragmentary material. The locality 
(grid reference 687240E, 6295047 N, Canowindra 
8360-N 1:50 000 topographic sheet) is on the eastern 
face of the access road to the Cadia open cut, about 1 
km from the entrance gates; a map of the region 
showing the location of this and other fossil localities 
was provided by Rickards et al. (2000, Fig. 1). The 
fossiliferous strata are considered to correlate with the 
Ulah Formation, at Four Mile Creek west of Cadia (see 
Rickards et al. 2000, Fig. 1), in which the Testograptus 
testis fauna occurs. 


NOTES ON THE GRAPTOLITE FAUNA 


Since the publication of Rickards et al. 
(2000), we have made a further but small graptolite 
collection from the Cadia mine shelly fossil locality 
which permits fuller identification of the low-diversity 
fauna and determination of its age. The Cadia graptolite 
fauna consists of Testograptus testis (Barrande), 


Monoclimacis flumendosae (Gortani), fragments of 
Monograptus flemingii (Salter), fragmentary stipes 
possibly belonging to Cyrtograptus, and fragmentary 
retiolitid meshworks which cannot be assigned, even 
approximately, to a genus. 

In discussing this as ‘the Cadia graptolite 
fauna’ we are mindful of the presence of other 
graptolites in Silurian strata in the vicinity of the Cadia 
mine. Full documentation of any such graptolite faunas 
as that documented here is important as graptolite 
localities in the vicinity of Cadia mine (such as the 
Pridoli ‘borrow pit’ locality, W910 of Rickards et al. 
2000) are very much less common than at Four Mile 
Creek, and are under threat. A brief review of 
graptolites previously reported from Cadia by 
Offenberg (1963) was given by Rickards et al. (2000). 

We have not provided here any systematic 
descriptions of the fauna, but limited comments on 
the morphological detail are included in the 
explanatory text for Figure 1. The Cadia specimens 
have undergone soft sediment deformation, with a 
considerable amount of twisting and breakage, in 
contrast to the Rodds Creek black shale specimens 
(Rickards et al. 2000) which were undeformed other 
than by diagenetic flattening. 


AGE OF THE CADIA GRAPTOLITE FAUNA 


The dominant species is Testograptus testis 
(Barrande), which normally indicates the late Wenlock 
(Early Silurian) /undgreni-testis Biozone. Testograptus 
testis has been recorded, very rarely, from the ludensis 


SILURIAN GRAPTOLITES FROM NEW SOUTH WALES 


Figure 1. (A) Monoclimacis flumendosae (Gortani), AM F114926, distal thecae, undeformed, low relief. 
(B-E) Testograptus testis (Barrande). (B) proximal end, AM F114928, showing some soft sediment 
deformation distally; (C) AM F114925, a proximal end with spines visible on th1; (D) AM F114930, 
spines on several thecae; (E) AM F114929, distal thecae with a growing end visible. (F) Monograptus 
flemingii (Salter), AM F114927, subscalariform view of mesial thecae. 


All figures x10, scale bar Imm; heavy bar indicates deformation stretching direction, possibly not tectonic. 
All specimens from locality W 937, grid reference 687240E, 6295047 N, Canowindra 8360-N 1:50 000 
topographic sheet. Unfigured specimens are AMF 114931-940. 


Biozone (Rickards et al. 1995) but, as the Cacia 
specimens are abundant and occur with Monoclimacis 
flumendosae (Gortani), a pre-/udensis Biozone is 
indicated for this fauna. 

The Cadia fauna is probably slightly younger 
than the Rodds Creek fauna (Rickards et al. 2000). 
Although this latter assemblage included some 
lundgreni-testis Biozone indicators, the presence of 
Cyrtograptus ex gr. rigidus Tullberg indicated a 
probable middle rather than late Wenlock for the Rodds 
Creek fauna. The Cadia fauna is thus significantly older 
than the Pridoli fauna from the ‘borrow pit’ locality 
(W910) 2 km to the southeast (Rickards et al. 2000). 


174 


Correlation with the Four Mile Creek 
sequence is probably with festis-bearing beds of the 
Ulah Formation in Wallace Creek; in Spring and 
Quarry Creeks, the festis-bearing beds of the same 
formation are largely green and black mudstones 
(Packham, Rickards and Wright, unpublished data). 


SHELLY FAUNAS 
The disarticulated and fragmental shelly 


fauna in this slump unit is unusually abundant and 
diverse for the region, in contrast with clastic units of 


Proc. Linn. Soc. N.S.W., 125, 2004 


R.B. RICKARDS AND A.J. WRIGHT 


this age in the Four Mile Creek area and the Spring- 
Quarry Creek areas which are singularly poor in shelly 
fossils. The faunas at Cadia have undergone soft- 
sediment deformation and are clearly transported. 
Described shelly faunas (other than corals) from the 
Four Mile Creek area and the Spring Creek areas are 
limited to two species of Judensis Biozone brachiopods 
described by Rickards and Wright (1997) from 
Cobblers Creek (see Fig. 1 of Rickards et al. 2000) 
and by Wright and Strusz (2004) from Spring Creek 
and Wallace Creek (see Fig. 1 of Rickards et al. 2000: 
ludensis Biozone and lundgreni-testis Biozone 
respectively). Other brachiopod faunas from the region 
were listed by Jenkins (1978, 1986), but the only rich 
faunas cited by him are from Llandovery (Early 
Silurian) limestones. 


CONCLUSIONS 


Graptolites identified from the Cadia Mine 
access road locality are Testograptus testis, 
Monoclimacis flumendosae, fragments of 
Monograptus flemingii, ?Cyrtograptus and retiolitids. 
The fauna is late Wenlock (Early Silurian) and is 
probably best correlated with a level high in the 
lundgreni-testis Biozone. It appears to be slightly 
younger than the probably middle Wenlock Rodds 
Creek black shale fauna (Rickards et al. 2000), and is 
assumed to correlate with the testis fauna of the Ulah 
Formation in the Four Mile Creek area to the west of 
Cadia. 


ACKNOWLEDGEMENTS 


We are grateful to Ian Tedder (Newcrest Mining 
Limited) for allowing access to, and guiding us to, this 
locality in November 2001. The universities of Cambridge 
and Wollongong have provided financial support for 
participation by RBR and AJW in this study, and funds were 
provided to AJW by the Linnean Society of New South 
Wales through the Betty Mayne Fund. 


Proc. Linn. Soc. N.S.W., 125, 2004 


REFERENCES 


Jenkins, C.J. (1978). Llandovery and Wenlock stratigraphy 
of the Panuara area, central New South Wales. 
Proceedings of the Linnean Society of New South 
Wales 102, 109-130. 

Jenkins, C.J. (1986). The Silurian of mainland Australia: a 
field guide. 82 p. UGS Silurian Subcommission, 
Sydney. 

Offenberg, A.C. (1963). Geology of the Panuara-Cadia- 
Errowanbong area, south of Orange, New South 
Wales. BSc (Hons) thesis, University of Sydney 
(unpublished), 112 p. 

Rickards, R.B., Packham, G.H., Wright, A.J. and 
Williamson, P.L. (1995). Wenlock and Ludlow 
graptolite faunas and biostratigraphy of the Quarry 
Creek district, New South Wales. Association of 
Australasian Palaeontologists, Memoir 17, 1-68. 

Rickards, R.B., Percival, I.G., Simpson, A:J. and Wright, 
A.J. (2000). Silurian biostratigraphy of the Cadia 
area, near Orange, New South Wales. Proceedings 
of the Linnean Society of New South Wales 123, 
173-191. 

Rickards, R.B. and Wright, A.J. (1997). Graptolite 
zonation in the late Wenlock (Early Silurian), with 
a new graptolite-brachiopod fauna from New 
South Wales. Records of the Australian Museum 
49, 229-248. 

Wright, A.J. and Strusz, D.L. (2004). Wenlock (Early 
Silurian) brachiopods from the Orange district of 
New South Wales. Proceedings of the Linnean 
Society of New South Wales 125, 165-172. 


175, 


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Silicified Early Devonian Trilobites from Brogans Creek, New 
South Wales 


Grecory D. EDGECOMBE! AND ANTHONY J. WRIGHT? 


"Australian Museum, 6 College Street, Sydney, NSW 2010 (greged@austmus.gov.au); 
*School of Earth and Environmental Sciences, University of Wollongong, Wollongong, NSW 2522 
(tony_wright@uow.edu.au) 


Edgecombe, G.D. and Wright, A.J. (2004). Silicified Early Devonian trilobites from Brogans Creek, New 
South Wales. Proceedings of the Linnean Society of New South Wales 124, 177-188. 


Trilobites in an Emsian silicified fauna from the Carwell Creek Formation at Brogans Creek SE of Mudgee, 
NSW, include Acanthopyge (Jasperia) bifida, Dentaloscutellum hudsoni and Proetus nemus, all originally 
described from the Taemas area of NSW, together with Sthenarocalymene. Proetus nemus was known from 
limited material at Taemas, but is the most abundant species at Brogans Creek. Fuller description substantiates 
membership in Proetus (=Devonoproetus), rather than Ryckholtia, Longiproetus or Rhenocynproetus. Early 
ontogenetic stages of the trilobites are lacking at Brogans Creek, in contrast to Taemas. Conodonts co- 
occurring with the shelly fauna at Brogans Creek and at Taemas include Polygnathus nothoperbonus, which 
indicates the Polygnathus perbonus Conodont Zone (medial Emsian). 


Manuscript received 16 October 2003, accepted for publication 8 January 2004. 


Keywords: Trilobita, Devonian, New South Wales, Acanthopyge (Jasperia), Dentaloscutellum, Proetus, 


Sthenarocalymene. 


INTRODUCTION 


The presence of Devonian limestone at 
Brogans Creek (Fig. 1), located SE of Mudgee in the 
central tablelands of NSW, was first noted by Carne 
and Jones (1919) and later by Lishmund et al. (1986). 
Fossils from the limestone were discussed in detail by 
Colquhoun (1998) and Colquhoun and Meakin in 
Colquhoun et al. (in Meakin and Morgan 1999). 
Colquhoun (1995) illustrated the conodonts 
Pandorinellina e. exigua and Polygnathus 
nothoperbonus from Brogans Creek, the latter species 
considered (after Mawson 1987) to be characteristic 
of the medial Emsian (Polygnathus perbonus zone). 

Here we provide the first descriptions of any 
of the well-preserved and abundant fossils from the 
quarry at Brogans Creek. A silicified trilobite fauna is 
of low diversity, but it provides new data on some taxa 
described from the Taemas area by Chatterton (1971), 
in particular the proetid Proetus nemus. 


Stratigraphic assignment and age 

The Devonian strata at Brogans Creek were 
considered part of the Carwell Creek Formation by 
Colquhoun et al. (1999). The limestones that have 
yielded the trilobites and other fossils documented here 
have also yielded (Colquhoun 1995) the medial Emsian 


conodont Polygnathus nothoperbonus, so this 
limestone is significantly younger than most limestones 
occurring in the area between the Mudgee and Brogans 
Creek, with the principal exception of those reported 
by Pickett (1972) and Colquhoun (1998) from the 
Mount Knowles Limestone Member of the Carwell 
Creek Formation and by Pickett (1978) from the Mount 
Frome Limestone, both located to the E of Mudgee. 
Little is known about the sequence of the 
Devonian strata in the vicinity of the Brogans Creek 
quarry, and recent land reclaimation operations have 
concealed formerly productive parts of the abandoned 
quarries. Colquhoun (1998) stated that the sequence 
grades upwards from the fossiliferous limestone 
through crinoidal sandstone into massive shale and 
volcarenite. The sequence of beds that yielded silicified 
fossils is about 10 m in thickness. Beds immediately 
overlying these strata have yielded the tetracorals 
Xystriphyllum mitchelli and Embolophyllum, both also 
described from the Receptaculites Limestone Member 
at Taemas and Wee Jasper by Pedder et al. (1970). 
The similarity of the macrofauna to that from 
the Receptaculites and Warroo limestone members of 
the Taemas Formation in the Burrinjuck Dam area of 
NSW (see Pedder et al. 1970) necessitates some 
consideration of the ages of these units. Conodont data 
summarised by Talent et al. (2000) for the Taemas 


EARLY DEVONIAN TRILOBITES FROM N.S.W. 


Mudgee. 


Taemas , 


> 
a eae all 
\ 


148°50' 


Rylstone 


Figure 1. Location of trilobite collection at Brogans Creek. Map of NSW indicates Taemas, where the 
same species have been described (Chatterton 1971). Shading in inset map (after Colquhoun 1995) shows 
distribution of Lower Devonian platform sediments. 


178 Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE AND A.J. WRIGHT 


area of NSW indicate that the Receptaculites and 
Warroo limestones at Taemas, which overlie the Cavan 
Formation with its Polygnathus pireneae to P. 
dehiscens fauna, are probably early Emsian. Lindley 
(2002) recorded Polygnathus nothoperbonus from the 
Warroo Limestone Member, further confirming the 
assignment of this limestone to the medial Emsian 
Polygnathus perbonus Conodont Zone. However, 
Basden et al. (2002) concluded that the Warroo 
Limestone should be correlated with the Polygnathus 
inversus to P. serotinus Conodont Zones. On balance 
the co-occurrence of P. nothoperbonus in both areas 
of NSW seems to indicate unequivocally a medial 
Emsian age for the macrofaunas. This supports the 
conclusions of Garratt and Wright (1988:Fig. 3), who 
correlated their Malurostrophia-Taemostrophia- 
Howittia fauna (essentially the shelly fauna discussed 
here) with the Polygnathus gronbergi (=P. perbonus) 
Conodont Zone. 


Faunal characters and affinities 

The fossiliferous limestones have yielded 
very rich and well-preserved invertebrate faunas, 
dominated by brachiopods, tabulate corals and 
tetracorals, trilobites, gastropods, ostracodes, 
cephalopods, tentaculitids, crinoid debris and sponges; 
bivalves are subordinate at this locality. Most of the 
trilobites and brachiopods at Brogans Creek are 
conspecific with those described from Emsian 
limestones in the Lake Burrinjuck sequence at Taemas 
and ‘Bloomfield’ by Chatterton (1971, 1973). With 
respect to the trilobites, the faunal composition of the 
Brogans Creek assemblage is best matched in the lower 
half of the Receptaculites Limestone at Locality I of 
Chatterton (1971). The three species identified here, 
Proetus nemus, Dentaloscutellum hudsoni and 
Acanthopyge bifida, are represented in the lower 
Receptaculites Limestone at Locality I and at that 
locality as well as Brogans Creek they occur with 
Sthenarocalymene. Silicified residues from Brogans 
Creek yield the following for minimal number of 
individuals per species, based on the most abundant 
Skeletal element: Proetus nemus (N=54), 
Dentaloscutellum hudsoni (N=16), Acanthopyge bifida 
(N=7), and Sthenarocalymene sp. (N=2). About 120 
kilograms of limestone have been etched to produce 
our fauna. 

In terms of diversity, the silicified assemblage 
consists additionally of more than 15 brachiopod 
species (Malurostrophia flabellicauda reverta 
Chatterton; Salopina kemezysi Chatterton and other 
dalmanellids; Schuchertella murphyi Chatterton; 
Coelospira dayi Chatterton; Howellella sp.; 
Ambothyris runnegari Chatterton; Howittia sp.; 


Proc. Linn. Soc. N.S.W., 125, 2004 


?Buchanathyris sp.; reticulariid indet.; Cydimia parva 
Chatterton; Parachonetes flemingi Chatterton; P. sp. 
cf. P. konincki Chatterton; rhynchonellids). Some 30 
gastropod species are under study by Dr A.G. Cook. 
Tetracoral species are dominated numerically by an 
abundant solitary Plasmophyllum, as well as other 
solitary corals (?acanthophyllids) and rare fragments 
of ?Calceola. The sponge Amphipora is locally 
abundant, and presumably represents lagoonal phases 
of deposition or influx of lagoonal debris; several 
biofacies are evident. Colquhoun (1998) indicated that 
the Brogans Creek limestone was deposited in a well- 
oxygenated, normal salinity environment. The trilobite 
material is represented by disarticulated sclerites, but 
many brachiopods shells are articulated. Scolecodonts 


are at least as common as conodonts in residues; this 


is also a feature of limestones in the Capertee Valley 
(S of Brogans Creek) where the strata are highly 
deformed and preservation is poor. Despite the 
disarticulated nature of parts of the Brogans Creek 
shelly fauna, their excellent preservation indicates that 
postmortem transportation was minimal. 


SYSTEMATIC PALAEONTOLOGY 


Figured material is in the Palaeontology collection, 
Australian Museum, Sydney (prefix AMF). 


Order PROETIDA Fortey and Owens, 1975 
Family PROETIDAE Salter, 1864 
Subfamily PROETINAE Salter, 1864 
Genus PROETUS Steininger, 1831 


Type species 
Calymmene concinna Dalman, 1827; by original 
designation. 


Proetus nemus Chatterton, 1971 
Fig. 2a-p, Fig. 3a-t 


Proetus nemus Chatterton, 1971:65-67, Pl. 16, Figs 
18-32. 
Ryckholtia? nemus (Chatterton). Liitke, 1990:21. 


Material 
39 cranidia, 103 librigenae, 3 hypostomes, 62 
thoracic segments, 50 pygidia. 


Diagnosis 

Proetus with relatively elongate, tapering glabella, its 
posterior two thirds with dense, mostly moderate sized 
tubercles, its anterior third granulate. Facial suture 
divergent between y and B. Genal ridge strong along 


179 


EARLY DEVONIAN TRILOBITES FROM N.S.W. 


Figure 2. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Emsian), Brogans Creek, 
NSW. Scale bars 1 mm. a-c, AMF 124700, cranidium, dorsal, anterior and lateral views; d-f, AMF 124701, 
cranidium, dorsal, anterior and lateral views; g, AMF 124702, cranidium, dorsal view; h, AMF 124703, 
cranidium, dorsal view; i, AMF 124704, cranidium, lateral view; j, AMF 124705, cranidium, anterior 
view; k, AMF 124706, cranidium, dorsal view; l-m, AMF 124707, cranidium, dorsal and lateral views; n, 
AMF 124708, librigena, dorsal view; 0, AMF 124709, librigena, dorsal view; p, AMF 125485, cranidium,. 
dorsal view. 


180 Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE AND A.J. WRIGHT 


Figure 3. Proetus nemus Chatterton, 1971. Carwell Creek Formation (medial Emsian), Brogans Creek, 
NSW. Scale bars 1 mm. a, AMF 124710, librigena, internal view; b, AMF 124711, librigena, dorsal view; 
c, AMF 124712, librigena, dorsal view; d-e, AMF 124713, hypostome, ventral and lateral views; f, AMF 
124714, thoracic segment, dorsal view; g, AMF 124715, thoracic segment, dorsal view; h-j, AMF 124716, 
pygidium, posterior, lateral and dorsal views; k, AMF 124717, thoracic segment, dorsal view; 1, AMF 
124718, thoracic segment, anterior view; m, AMF 124719, thoracic segment, anterior view; n, AMF 124720, 
pygidium, dorsal view; o-p, AMF 124721, pygidium, lateral and dorsal views; q-r, AMF 124722, pygidium, 
posterior and dorsal views; s, AMF 124723, pygidium, ventral view; t, AMF 124724, pygidium, dorsal 
view. 


Proc. Linn. Soc. N.S.W., 125, 2004 181 


EARLY DEVONIAN TRILOBITES FROM N.S.W. 


all but posteriormost part of librigenal field, distinct 
but less prominent on preocular fixigena; small caecal 
pits abundant on librigenal field; genal spine relatively 
long. Pygidium with seven axial rings and lunate 
terminal piece (7+1); anterior three or four pleural 
furrows well impressed, fifth and sixth faint. 


Description 
Cranidial length about equal to maximum width at @; 
width at 6 slightly more than 80% width at @; width at 
B 85-95% width at 5. Axial furrow narrow, moderately, 
evenly deep. Glabella widest basally, length (excluding 
LO) 1.1-1.2 times basal width, with moderate taper 
anteriorly, slightly constricted at S2, gently convex 
(sag., tr.); frontal lobe rounded; terminating at but not 
overhanging anterior border furrow. S1 originating 
opposite midlength of palpebral lobe, shallow, directed 
posteromedially, distally birfucate, with posterior 
branch terminating well in front of SO; S2 parallel with 
S1, more weakly incised, originating just behind 
anterior edge of palpebral lobe; S3 obscure. Posterior 
two thirds of glabella with mostly moderate sized 
tubercles, some small tubercles, densely packed so as 
to nearly touch; anterior third of glabella granulate, 
non-tuberculate. SO transverse medially, narrow (sag., 
exsag.), deep, flexed forwards abaxially against lateral 
occipital lobes. LO distinctly wider than basal part of 
glabella, length about 20% its width; lateral occipital 
lobes large, drop-shaped, isolated from remainder of 
LO by deep furrows; LO, including lateral lobes, 
covered with tubercles as on posterior part of glabella, 
including moderately large median tubercle behind 
midlength. Preglabellar region 13-15% of cranidial 
length; in large specimens, composed of an inclined, 
medially flat posterior half and moderately convex 
(sag.) anterior half bearing 5-6 terrace lines in dorsal 
view; in small specimens, posterior half forms a wide 
(sag., tr.) depressed field with a broad (tr.), gently 
inflated transverse median swelling. Genal ridge well 
developed on preocular fixigena, anteromedially 
directed, terminating at juncture of preglabellar and 
anterior border furrows, stronger in small specimens. 
Postocular fixigena 25-35% width (tr.) and about 60% 
length (exsag.) of LO. Palpebral lobe arcuate, 35-45% 
length of glabella; palpebral furrow faint or indistinct. 
Anterior sections of facial suture diverging from each 
other at 45-62° between y and B, running subparallel 
against anterior border furrow, then strongly 
converging between B and a. Posterior sections of 
facial suture running subparallel or gently diverging 
between € and €, close to axial furrow, then sharply 
turned outwards to 0. 

Librigenal field moderately wide, gently 
convex (tr.); genal ridge strong along all but 


182 


posteriormost part of field, closer to eye socle than to 
lateral border furrow; most of field with abundant, 
small caecal pits, least distinct at posterolateral corner 
of field. Eye socle narrow, separated from visual 
surface and librigenal field by shallow furrows. 
Posterior border furrow narrow, deep; lateral border 
furrow wider, the two merging at genal angle, 
extending along a variable extent of the genal spine, 
usually along about half its length. Lateral border 70- 
80% as wide as narrowest part of librigenal field in 
dorsal view, strongly convex (tr.); terrace lines well 
defined along entire length and width of lateral border 
and along genal spine. Genal spine relatively long, its 
inner margin straight or faintly concave. Panderian 
notch large, semicircular. Connective suture with 
straight, diagonal course along most of its length, its 
extent relative to cranidium indicating that rostral plate 
is trapezoidal or triangular, fairly wide anteriorly (cf. 
P. concinnus: Owens 1973:Text-fig. 1B). 

Hypostomal width across shoulders about 
65% sagittal length. Anterior margin weakly convex 
medially, flexed backward abaxially. Anterior lobe of 
middle body strongly inflated (tr.), anteromedial part 
raised but not forming discrete rhynchos; middle body 
gently convex (sag.) along most of length, fairly steeply 
turned up anteromedially; anterior lobe bearing many 
sinuous terrace lines. Middle furrow moderately deep, 
directed posterolaterally across abaxial third of middle 
body then abruptly effacing. Border furrow narrow, 
distinctly impressed around entire middle body, 
shallowest against anterior wing. Anterior border 
uniformly narrow (sag., exsag.); lateral border gently 
converging between anterior wing and shoulder; 
shoulder rounded; posterolateral margin straight 
between shoulder and pair of blunt spines at lateral 
edge of posterior border; posterior border narrow (sag., 
exsag.), about 10% length of hypostome, with gently 
convex posteromedian margin. 

Number of thoracic segments unknown. 
Axial furrow narrow, shallow. Axis strongly convex 
(tr.), 32-41% width of thorax. Articulating half ring 
varying from equal in width (sag.) to 1.6 times as wide 
as preannulus along length of thorax, 70-90% length 
of ring; preannular furrow transverse to gently concave 
medially, sharply impressed but much shallower than 
articulating furrow; ring covered with small, dense 
tubercles or coarse granules. Pleural furrow narrow, 
about as deep as articulating furrow, gently flexed 
forward at fulcrum, abruptly shallowing then effacing 
on inner part of articulating facet; anterior and posterior 
pleural bands equal in width (exsag.) proximal to 
fulcrum; pleurae moderately declined abaxial to 
fulcrum, at midwidth (tr.) of rib. Pleural tips with 
curved anterolateral margin, blunt rounded posterior 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE AND A.J. WRIGHT 


projection. Panderian notch deep, U-shaped. 
Pygidium subsemicircular, length (excluding 
articulating half ring) 55-60% width. Axial width about 
35% pygidial width anteriorly; axial furrows narrow, 
uniformly impressed along most of length. Seven axial 
rings and short, lunate terminal piece (7+1); first one 
or two ring furrows lengthened medially as short 
preannulus; more posterior ring furrows shallower but 
with moderately deep incision across axis, posterior 
few gently convex backwards; axis raised strongly 
above pleurae, gently convex in sagittal profile, 
moderately arched (tr.); rings with dense small 
tubercles or coarse granules. Postaxial region about 
20% length of pygidium. Pleural furrows narrow 
(exsag.), anteriorly convex, anterior three or four well 
impressed, fifth and variably sixth faintly discernible; 
first pleural furrow terminates near pygidial lateral 
margin, others terminate at shallow posterior border 
furrow; interpleural furrows narrower and shallower 
than pleural furrows; pleural ribs with sculpture of 
dense, medium sized granules. Border widening back 
to its intersection with third pleural furrow, then 
maintaining even width, occupying most of postaxial 
region, weakly convex. Doublure extending in nearly 
as far as border furrow, bearing several terrace lines. 


Discussion 

The sample from Brogans Creek resembles that from 
Taemas in that the largest cranidia (Fig. 2a-c, h, i, p; 
Chatterton 1971:Pl. 16, fig. 28) have the anterior end 
of the glabella abutting the inclined posterior part of 
the anterior border, whereas small specimens have a 
broad depression between the frontal lobe and the 
convex, terraced part of the anterior border (Fig. 2d-f, 
j, k; Chatterton 1971:Pl. 16, fig. 25). The latter 
morphology, associated with a more pronounced 
fixigenal ridge (Fig. 2d, k versus 2a, h, p) is confined 
to small specimens. This difference in the preglabellar 
region is bridged by intermediate sized specimens, and 
is ascribed to ontogenetic variation. The transverse 
median swelling in the depression of small specimens 
(Fig. 2e, j) retains a faint expression in large cranidia. 
No bimodality can be detected in the strength of the 
librigenal ridge (Figs. 2n, 0, 3b, c), which is 
consistently pronounced. 

In assigning this species to Proetus, 
Chatterton (1971) acknowledged its distance from the 
type species, the Wenlock P. concinnus (Dalman). 
However, several other Australian Emsian and Eifelian 
Proetinae are validly assigned to that genus. These 
include Proetus talenti Chatterton, 1971 (type of 
Devonoproetus Liitke, 1990), P. sparsinodosus Feist 
and Talent, 2000, and P. latimargo Feist and Talent, 
2000, the latter two originally assigned to 


Proc. Linn. Soc. N.S.W., 125, 2004 


Devonoproetus at the subgeneric level. Devonoproetus 
is ajunior synonym of Proetus s.s. (Adrain 1997; Zhou 
et al. 2000). 

Proetus nemus was reassigned, with question, 
to the otherwise Ludlow-Lochkovian Ryckholtia 
Snajdr, 1980 (type Proetus ryckholti Barrande, 1846) 
by Liitke (1990). The new material described herein 
conflicts with this reassignment. Membership in 
Ryckholtia is precluded by the pronounced tuberculate 
sculpture on the glabella and axial rings of P. nemus, 
the strongly defined lateral occipital lobes, and sagittal 
elimination of the preglabellar field. 

This species displays characters that suggest 
alternative assignments. The elongate, tapering 
glabella of Proetus nemus and its pattern of sculpture 
(strong tuberculation posteriorly, becoming subdued 
anteriorly), together with the profile of the preglabellar 
region, including the wide (sag., exsag.) anterior 
cranidial border furrow, and the divergence of the 
facial suture between y and B resemble Longiproetus 
tenuimargo (Richter, 1909) (type of Longiproetus 
Cavet and Pillet, 1958). Longiproetus has been 
regarded as a synonym of Gerastos Goldfuss, 1843 
(Owens 1973), a valid subgenus of Gerastos (Snajdr 
1980), restricted to its type species on the basis of a 
distinctive shape of the rostral plate (Ltitke 1990), or 
slightly expanded to include a small group of 
Rhenohercynian mid Eifelian to early Givetian species 
(Basse 1996, 2002). Liitke (1990) reassigned the 
Bohemian species that had been referred to 
Longiproetus (e.g., Snajdr 1980) to Coniproetus 
Alberti, 1966, and other genera, whilst the inadequately 
known Emsian species referred to Longiproetus by 
Pillet (1972) defy classification. Despite the similarities 
in the glabella and preglabellar region, several 
characters conflict with an alliance between P. nemus 
and Longiproetus. Notably, the strong genal ridge of 
P. nemus is lacking in L. tenuimargo and other certain 
congeners (sensu Basse 2002), the prominent lateral 
occipital lobes contrast with the inconspicuous lobes 
in Longiproetus s.s., LO is wider than the basal part of 
the glabella, the cephalon is much less vaulted, the 
palpebral lobe is situated more posteriorly, and the 
pygidium is relatively paucisegmented (7+1 rings 
versus 8+1). The course of well preserved connective 
sutures on librigenae suggests that the rostral plate of 
P. nemus is more regularly trapezoidal or triangular 
than is that of L. tenuimargo (Liitke 1990:Text-fig. 8). 

Affinities to species that have been assigned 
to Devonoproetus by recent workers better account for 
the large occipital lobes, width of LO relative to the 
glabella, and 7+1 pygidial segmentation. Among these, 
Proetus latimargo Feist and Talent, 2000 (Eifelian, 
Queensland) and P. zhusilengensis Zhou et al., 2000 


183 


EARLY DEVONIAN TRILOBITES FROM N.S.W. 


(Emsian, Inner Mongolia) resemble P. nemus in having 
a tongue-shaped glabella (narrowest in P. nemus) with 
dense, pronounced tuberculation, and P. latimargo 
shares the divergence of the facial suture between 
and B. 

Among those species that have been referred 
to Devonoproetus, the strong genal ridge of Proetus 
nemus is developed in a group recognised by Basse 
(2002) as a separate genus, Rhenocynproetus, from 
which the Australian “Devonoproetus” species were 
explicitly excluded. The presence of a genal ridge in 
other genera of Proetinae [e.g. Gerastos: Snajdr 
1980:Pl. 3, Fig. 13, Pl. 4, Fig. 17; Coniproetus 
(Bohemiproetus): Snajdr 1980:Pl. 6, Figs 5, 6, 14; 
Lieberman 1994:Fig. 9.3) demonstrates that this feature 
is not an infallible indicator of relationships. Characters 
cited by Basse (2002) as excluding Australian species 
of Proetus from Rhenocynproetus also distinguish P. 
nemus; these include the large size of the lateral 
occipital lobes and weaker outer edge of the eye socle. 
Proetus nemus possesses (plesiomorphic) features 
considered by Basse (2002) to more generally 
distingish Proetus from Rhenocynproetus, such as a 
less inflated glabella, the lateral occipital lobes wider 
than the base of the glabella, terrace lines developed 
on the dorsal as well as lateral extent of the cranidial 
border, and the well developed librigenal spine. The 
presence of a pair of posterior border spines on the 
hypostome (Fig. 3d) is shared with Proetus (e.g. 
Whittington and Campbell 1967:PI1. 1, Fig. 17; Schrank 
1972:Pl. 4, Fig. 7), including P. talenti, but is likely 
symplesiomorphic (Adrain 1997). 


Order CORYNEXOCHIDA Kobayashi, 1935 
Suborder SCUTELLUINA Hupé, 1953 
Family STYGINIDAE Vogdes, 1890 
Genus DENTALOSCUTELLUM Chatterton, 1971 


Type species 
Dentaloscutellum hudsoni Chatterton, 1971; by 
original designation. 


Dentaloscutellum hudsoni Chatterton, 1971 
Fig. 4a-i 


Dentaloscutellum hudsoni Chatterton, 1971:12-22, 
Pl. 1, Figs 1-24, Pl. 2, Figs 1-24, Pl. 3, Figs 1-12, Pl. 
24, Fig. 15, Text-figs 4-5. 


Material 


4 cranidia, 29 librigenae, 1 hypostome, | thoracic 
segment, 5 fragmentary pygidial margins. 


184 


Discussion 

This species was fully described based on specimens 
from the Receptaculites Limestone near Taemas 
(Chatterton 1971). The Brogans Creek material is 
considered to be conspecific, the only possible 
difference being slightly more numerous cranidial 
tubercles (Fig. 4b, c) than in the type material. 


Order LICHIDA Moore, 1959 
Family LICHIDAE Hawle and Corda, 1847 
Subfamily TROCHURINAE Phleger, 1936 
Genus ACANTHOPYGE Hawle and Corda, 1847 


Type species 
Acanthopyge leuchtenbergii Hawle and Corda, 
1847; by subsequent designation of Reed (1902). 


Subgenus JASPERIA Thomas and Holloway, 1988 


Type species 
Acanthopyge (Mephiarges) bifida Edgell, 1955; by 
original designation. 


Acanthopyge (Jasperia) bifida Edgell, 1955 
Fig. 4j-t 


Acanthopyge (Mephiarges) bifida Edgell, 1955:138; 
Chatterton, 1971:30-41, Pl. 6, Figs 1-24, Pl. 7, Figs 
1-27, Pl. 8, Figs 1-17, Text-figs 8-10. 


Material 
7 cranidia, | rostral plate, 7 librigenae, 3 
hypostomes, | thoracic segment, 2 pygidia. 


Discussion 

The Brogans Creek specimens are indistinguishable 
from those described from Wee Jasper (Edgell 1955) 
and Taemas (Chatterton 1971). The species was fully 
described by Chatterton (1971), rendering description 
of the Brogans Creek material unnecessary. A few 
specimens are illustrated (Fig. 4j-t) in support of the 
conspecificity of the collections. 


Order PHACOPIDA Salter, 1864 
Suborder CALYMENINA Swinnerton, 1915 
Family CALYMENIDAE Milne Edwards, 1840 
Genus STHENAROCALYMENE Siveter, 1977 


Type species 


Sthenarocalymene lirella Siveter, 1977; by original 
designation. 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE AND A.J. WRIGHT 


Figure 4. a-i, Dentaloscutellum hudsoni Chatterton, 1971. Scale bars Imm. a, AMF 124725, librigena, 
dorsal view; b-d, AMF 124726, cranidium, dorsal, anterior and lateral views; e, AMF 124727, librigena, 
dorsal view; f, AMF 124728, librigena, ventral view; g, AMF 124729, fixigena, dorsal view; h, AMF 
124730, incomplete pygidium, ventral view; i, AMF 124731, incomplete pygidium, ventral view. j-t, 
Acanthopyge (Jasperia) bifida Edgell, 1955. Scale bars 1 mm. j, AMF 124732, rostral plate, ventral view; 
k, AMF 124733, cranidium, dorsal view; l-m, AMF 124734, cranidium, dorsal and anterior views; n, 
AMF 1247335, librigena, dorsal view; o-q, AMF 124736, pygidium, lateral, dorsal and ventral views; r, 


AME 124737, librigena, ventral view; s-t, AMF 124738, hypostome, ventral and dorsal views. 


Proc. Linn. Soc. N.S.W., 125, 2004 185 


EARLY DEVONIAN TRILOBITES FROM N.S.W. 


Sthenarocalymene sp. 


Material 
Two cranidial fragments, one fragmentary librigena. 


Discussion 

A few calymenid cephalic fragments indicate the 
presence of a species lacking a buttress between the 
fixigena and L2. On this basis the material is assigned 
to Sthenarocalymene, the non-buttressed calymenid 
in many Australian Lower Devonian faunas [see 
Sandford (2000) for discussion of this genus, its 
synonym Apocalymene Chatterton and Campbell, 
1980, and Gravicalymene Shirley, 1936]. The Brogans 
Creek material may be identical with S. quadrilobata 
(Chatterton, 1971), which co-occurs with the other taxa 
described herein in the lower Receptaculites Limestone 
at Locality [ of Chatterton (1971), but specific identity 
requires better specimens. 


ACKNOWLEDGEMENTS 


Alex Cook (Queensland Museum) assisted AJW 
in the field, and processed and picked much of the material 
studied here. Yongyi Zhen (Australian Museum) 
photographed the specimens and assembled the plates, and 
processed several blocks of limestone. Robert Owens 
(National Museum of Wales) provided helpful suggestions 
on an earlier version of the mansucript. 


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188 Proc. Linn. Soc. N.S.W., 125, 2004 


A New Species of the Henicopid Centipede Dichelobius 
(Chilopoda: Lithobiomorpha) from Southeastern Australia and 
Lord Howe Island 


GreGcory D. EDGECOMBE 


Australian Museum, 6 College Street, Sydney, NSW 2010 
(greged @ austmus.gov.au) 


Edgecombe, G.D. (2004) A new species of the henicopid centipede Dichelobius (Chilopoda: Lithobiomorpha) 
from southeastern Australia and Lord Howe Island. Proceedings of the Linnean Society of New South 


Wales 125, 189-203. 


The genus Dichelobius Attems, 1911, based on D. flavens Attems, 1911, from the southwest of Western 
Australia, has its only other previously assigned species in New Caledonia and Chile. The Tasmanian type 
species of the monotypic Tasmanobius Chamberlin, 1920, is regarded as a member of Dichelobius. 
Dichelobius giribeti n. sp. represents the genus in eastern mainland Australia (southeastern New South 
Wales, the Australian Capital Territory, and northeastern Victoria) and on Lord Howe Island. Dichelobius 
bicuspis Ribaut, 1923, is widely distributed in New Caledonia. 


Manuscript received 1 March 2003, accepted for publication 8 January 2004. 


KEYWORDS: Anopsobiinae, Chilopoda, Dichelobius, Henicopidae, Lithobiomorpha. 


INTRODUCTION 


The subfamily Anopsobiinae is a group of 
minute centipedes (Chilopoda) in the predominantly 
southern temperate family Henicopidae. Anopsobiinae 
is distributed chiefly in the Southern Hemisphere, with 
species described from Patagonian Argentina and Chile 
(Silvestri 1899, 1909a-b; Verhoeff 1939; Chamberlin 
1962), the Falkland Islands (Eason 1993), New 
Zealand (Silvestri 1909a; Archey 1917, 1937), New 
Caledonia (Ribaut 1923), Tasmania (Chamberlin 
1920), New South Wales (Edgecombe 2003), 
southwest Western Australia (Attems 1911), and the 
Cape region of South Africa (Attems 1928). Four 
Gondwanan genera have been named: Anopsobius 
Silvestri, 1899, Catanopsobius Silvestri, 1909b, 
Dichelobius Attems, 1911, and Tasmanobius 
Chamberlin, 1920. Four additional anopsobiine genera, 
all monotypic, occur in the Northern Hemisphere, 
namely Anopsobiella Attems, 1938, Ghilaroviella 
Zalesskaja, 1975, Shikokuobius Shinohara, 1982, and 
Rhodobius Silvestri, 1933. In total, 17 species and 
subspecies of Anopsobiinae have been described. 

Silvestri (1909a) cited the occurrence of an 
anopsobiine from Sydney, but formal descriptions of 
Anopsobiinae in eastern Australia are limited to 
Tasmanobius relictus Chamberlin, 1920, based upon 
a single specimen from Tasmania, and Anopsobius 
wrighti Edgecombe, 2003, from northern New South 


Wales. Mesibov (1986) indicated the presence of two 
species of Anopsobiinae in Tasmania. The present 
study continues a systematic treatment of 
Anopsobiinae of Australia by documenting a new 
species of Dichelobius from New South Wales, the 
Australian Capital Territory, Victoria, and Lord Howe 
Island (Fig. 1). 

For electron microscopy, specimens were 
photographed on a Leo 435VP using a Robinson 
backscatter detector. Digital images were assembled 
into plates with Photoshop. Morphological 
terminology is as summarised by Edgecombe 
(2001:203), with terminology for the mandible as in 
Edgecombe et al. (2002:40, Fig. 4). 

The following abbreviations are used for 
repositories of specimens examined: 

AM - Australian Museum, Sydney 

ANIC — Australian National Insect Collection, 
Canberra 

MCZ — Museum of Comparative Zoology, Harvard 
University, Cambridge, MA 

MNHN — Museum National d’ Histoire Naturelle, 
Paris 

NMW — Naturhistorisches Museum Wien 

QM — Queensland Museum, Brisbane 

WAM — Western Australian Museum, Perth. 

Other abbreviations: Berl., ANIC Berlesate; CBCR, 
Australian Museum Centre for Biodiversity and 
Conservation Research; Ck, Creek; Mt, Mountain; NP, 
National Park; rf, rainforest; SF, State Forest. 


A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 


Brisbane 


Lord Howe 
Island © 


Canberra 
|e 


Melbourne 


Sydney 


ht 


Figure 1. a, southeastern Australia and Lord Howe 
Island. Inset shows location of map in b, indicating 
records of Dichelobius giribeti n. sp. (open dots) in 
New South Wales, the Australian Capital Territory, 
and Victoria. 


Collectors: GBM — G.B. Monteith; JFL — J.F. 
Lawrence; RJB — R.J. Brooks; RWT — R.W. Taylor. 


Order LITHOBIOMORPHA Pocock, 1902 
Family HENICOPIDAE Pocock, 1901 
Subfamily ANOPSOBIINAE Verhoeff, 1907 
Genus DICHELOBIUS Attems, 1911 


Tasmanobius Chamberlin, 1920 n. syn. 
Type species 


Dichelobius flavens Attems, 1911; by 
monotypy. 


190 


Assigned species 

Dichelobius relictus (Chamberlin, 1920) n. 
comb.; D. bicuspis Ribaut, 1923; D. schwabei 
Verhoeff, 1939; D. giribeti n. sp. 


Diagnosis 
Anopsobiinae with spiracle on segments 3, 
10 and 12, variably present on segment 14. 


Discussion 

The Gondwanan genera Dichelobius, 
Tasmanobius and Anopsobius share several 
apomorphic characters relative to Northern 
Hemisphere Anopsobiinae. These include coxal pores 
confined to legs 14 and 15, a ventrodistal spur on the 
prefemur of legs 14 and 15, an elongate longitudinal 
median furrow on the head shield, the basal article of 
the female gonopod extended as a short process bearing 
the spurs, and indistinct scutes on the proximodorsal 
part of the pretarsal claws (Edgecombe and Giribet 
2003). Considering previous concepts of Dichelobius 
(Attems 1928; Verhoeff 1939; Shinohara 1982), 
reduced spiracles are the only morphological character 
that unites its members to the exclusion of Anopsobius 
as delimited by Chamberlin (1962) and Edgecombe 
(2003). The Dichelobius distribution of spiracles is 
shared by the eastern Australian species D. giribeti. 
The cladistic reliability of a diminished number of 
segments with spiracles can be questioned because 
other genera of Anopsobiinae have been diagnosed 
based on having spiracles confined to segments 3, 10 
and 12 (Tasmanobius), 3, 12 and 14 (Rhodobius) or 3 
and 10 only (Catanopsobius). However, molecular 
sequence data provide independent support for a close 
relationship between D. flavens and D. giribeti, with 
the implication that their shared spiracle distribution 
can be considered a synapomorphy (Fig. 2a). 
Parsimony analysis of five molecular loci as well as 
combination of the molecular data and morphology 
unite D. flavens and D. giribeti to the exclusion of 
Anopsobius species under many explored gap costs 
and transversion:transition ratios (Edgecombe and 
Giribet 2003) (Fig. 2c). An alternative relationship 
between D. giribeti and Anopsobius (Fig. 2b) is 
discussed below. 

Verhoeff (1925) cited the presence of a 
median suture in the maxillipede pleural band as an 
additional character by which Dichelobius is 
distinguished from Anopsobius. The presence of a 
median suture (see Fig. 6}) is a plesiomorphic character, 
shared with Henicopinae, and is thus not useful for 
defining Dichelobius as a clade. 

Tasmanobius relictus Chamberlin, 1920, is 
considered to be a member of Dichelobius as grouped 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


Shikokuobius 
Anopsobius TAS 
Anopsobius wrighti 


japonicus 
Anopsobius 
neozelanicus 


Dichelobius 
giribeti 


Dichelobius 
flavens 


Shikokuobius 


japonicus 
Anopsobius wrighti 


Dichelobius 
Anopsobius TAS 
Anopsobius 
neozelanicus 


flavens 
Dichelobius 


giribeti 


1 


oo 
1 2 


Gap cost 


4 


inf. 


c Tv : Ts cost 


Figure 2. a, b, alternative cladograms for 
Anopsobiinae based on combined morphological 
and molecular data (Edgecombe and Giribet 2003). 
Character 1, absence of spiracles on segment 8; 
character 2, short posteroventral spine on pretarsal 
claw; c, summary of 12 analyses for combined 
morphological and molecular data with different 
gap costs (gap:substitution = 1:1, 2:1, 4:1) and 
transversion:transition costs (1:1, 2:1, 4:1, infinity). 
Black squares, parameters that resolve cladogram 
a (Dichelobius monophyletic); white squares, 
parameters that resolve cladogram b (Dichelobius 
paraphyletic); grey square, cladograms a and b of 
equal length. 


Proc. Linn. Soc. N.S.W., 125, 2004 


herein (with Tasmanobius consequently being a junior 
subjective synonym of Dichelobius). Tasmanobius 
relictus was described as having spiracles on segments 
3, 10, and 12, as in Dichelobius. Mesibov (1986) 
suggested that a widespread Tasmanian anopsobiine 
species (Anopsobiine sp. 2 of Mesibov 1986) may be 
Tasmanobius relictus, and that species closely 
resembles Dichelobius giribeti. The holotype and sole 
type specimen of T. relictus (MCZ 14533) is in poor 
condition, and lacks locality data more specific than 
Tasmania, making the identification of any other 
specimen as this species problematical. The description 
by Chamberlin did not note a spiracle on segment 14 
which is present in the Tasmanian Dichelobius, though 
this is not obvious in contracted specimens, as noted 
by Mesibov (1986). A spiracle being absent on segment 
eight in T. relictus and the colour being “nearly 
chestnut’ (Chamberlin 1920) make it probable that this 
species is identical with the Tasmanian Dichelobius 
(=Anopsobiinae sp. 2 of Mesibov 1986) rather than 
the northwestern Tasmanian Anopsobius 
(=Anopsobiinae sp. 1 of Mesibov 1986), which has a 
spiracle on segment 8 and is more orange-yellow than 
orange-brown. Accordingly, the name Dichelobius 
relictus (Chamberlin, 1920) is applied to Anopsobiinae 
sp. 2 of Mesibov (1986). 

Attems’ (1928:74) key to anopsobiine genera 
followed Chamberlin’s (1920) in distinguishing 
Dichelobius and Tasmanobius based on the former 
having a 1-jointed tarsus 13 and the latter a 2-jointed 
tarsus 13. This distinction is inconsistent with the 
referral of D. bicuspis, which has a 2-jointed tarsus 13 
(even fide Attems 1928:77). The supposed difference 
between these species seems to be nothing more than 
a terminological difference in what constitutes a 
‘Joint’, since D. flavens, D. bicuspis and D. relictus 
are, upon direct comparison, identical with respect to 
the segmentation of leg 13. All have a distinct 
articulation on the tarsus of leg 13, though it is less 
flexed than is the articulation on leg 14. 

Other ambiguities concerning Attems’ 
description and illustrations of Dichelobius flavens 
have plagued previous interpretations of the genus, and 
exaggerated differences between D. flavens and other 
species. Interpretation of D. flavens is based on 
examination of syntypes from Lion Mill (WAM), 
Freemantle and Eradu (NMW), and large new 
collections from the southwest of Western Australia 
(AM, ANIC, WAM). Dichelobius bicuspis and D. 
schwabei were distinguished from D. flavens by the 
first two species having two coxal pores on legs 14 
and 15 in the female, versus a single pore on each of 
the coxae in D. flavens. This cannot be upheld, since 
large females of D. flavens characteristically have two 


191 


A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 


coxal pores on both legs 14 and 15. Attems’ (1911:157, 
Fig. 10) described and figured a single spur on the 
female gonopod in D. flavens, which Ribaut (1923) 
and Verhoeff (1939) cited as a distinction from the 
pair of spurs in D. bicuspis and D. schwabei, 
respectively. Large specimens of Dichelobius flavens 
resemble congeners (and indeed all other 
Anopsobiinae) in having a pair of spurs. The specimen 
drawn by Attems, with a single spur and single coxal 


pore, is typical of immature stadia of all Dichelobius 
species (see Archey 1937:pl. 23, fig. 6, for a 
comparable stage in Anopsobius neozelanicus). Ribaut 
(1923:27) distinguished D. bicuspis by its plumose 
setae along the length of the inner margin of the distal 
article of the telopodite of the first maxilla versus only 
three plumose setae confined to the distal end of this 
article in D. flavens (Attems 1911:Fig. 3). Either 
Attems’ drawing is erroneous or else the illustrated 


Figure 3. Pretarsal claws in Anopsobiinae. a, Dichelobius relictus (Chamberlin, 1920). Leg 14, posterior 
side. b, c, Dichelobius flavens Attems, 1911. Leg 14, posterior and anterior sides. d, Anopsobius neozelanicus 
Silvestri, 1909. Leg 14, posterior side. e, f, Shikokuobius japonicus (Murakami, 1967). Leg 13, posterior 
and anterior sides. Scales 10 [um except b, 5 um. 


192 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


specimen is anomalous, because D. flavens has 
plumose setae all along the inner margin of this article, 
the same as D. bicuspis (Ribaut 1923:Figs. 30, 31) and 
other congeners. 

Certain characters of the pretarsus (claws) 
conflict with the monophyly of Dichelobius as grouped 
herein. Dichelobius flavens (Fig. 3b, c) and D. bicuspis 
differ from D. giribeti (Fig. 8b) and D. relictus (Fig. 
3a) in having a long, needle-like spine (=’sensory spur” 
of Eason 1964:Fig.486) originating ventrally on the 
posterior side of the main claw. In the latter two species, 
the posteroventral spine is short, and a short spine is 
shared by species of Anopsobius, such as A. 
neozelanicus Silvestri, 1909a (Fig. 3d) and A. wrighti 
(Edgecombe 2003:Figs.30, 31). The short spine 
appears to be apomorphic within the Gondwanan group 
of Anopsobiinae (i.e., a clade composed of Anopsobius 
+ Dichelobius) because the Japanese anopsobiine 
Shikokuobius japonicus resembles Dichelobius flavens 
and D. bicuspis in possessing a greatly elongated 
posteroventral spine (Fig. 3e, f). The cladogram 
implied by this character, in which D. giribeti is more 
closely related to Anopsobius than to D. flavens (Fig. 
2b), is retrieved under several parameter sets for 
combined morphological and molecular data (Fig. 2c). 
This cladogram would favour the assignment of D. 
giribeti to another genus. Should this topology find 
further support from additional data, Tasmanobius 
Chamberlin, 1920, could be rediagnosed to receive D. 
giribeti. A rediagnosed concept of that genus might 
emphasise the shared 14-15 antennal articles, short 
pretarsal posteroventral spine, absence of a distal 
spinose projection on the tibia of leg 12, and lack of 
spiracles on segments 5 and 8. 


Key to Dichelobius species 


la. Dental margin of maxillipede coxosternite 
lacking median notch ....... schwabei Verhoeff, 
1939 [Chile] 

1b. Dental margin of maxillipede coxosternite with 
median notch...... 2 


2a. 14-15 (usually 15) antennal articles; pretarsus 
with short posteroventral spine, not more than one- 
eighth length of main claw (Fig. 8b)...... 3 

2b. 17 antennal articles; pretarsus with needle-like 
posteroventral spine nearly as long as main claw 
(Fig. 3c).....4 


3a. Spiracle absent on segment 14...... giribeti n. sp. 
[southeastern Australia, Lord Howe Island] 
3b. Spiracle present on segment 14...... relictus 


Chamberlin, 1920 [Tasmania] 


Proc. Linn. Soc. N.S.W., 125, 2004 


4a. Tibia of leg 12 with short, blunt distal 
projection...... flavens Attems, 1911 [Western 
Australia] 

4b. Tibia of leg 12 with spinose distal 
projection...... bicuspis Ribaut, 1923 [New 
Caledonia} 


Dichelobius giribeti n. sp. 


Dichelobius sp. Edgecombe, 2004:Fig. 38A. 
Dichelobius sp. ACT. Edgecombe and Giribet, 
2003:Figs. 1-3. 


Etymology 

For Gonzalo Giribet, my collaborator in 
henicopid phylogeny, who sequenced DNA from this 
species. 


Diagnosis 

Dichelobius usually with 15 antennal articles; 
head pale orange, tergites orange-yellow; four to six 
(most commonly five) teeth on each dental margin of 
maxillipede; spiracle lacking on segment 14; two coxal 
pores on legs 14 and 15 in females, one or two pores 
on both legs in males; short posteroventral spine on 
pretarsus. 


Type material : 
Holotype: AM KS 82628, female (Fig. 4b), 

Badja SF, NSW, Peters Rd, 36°08’52"S 149°32’09"E, 

J. Tarnawski and S. Lassau, 13.ii1.1999; length of body 

5.1 mm. Paratypes, all from type locality, same 

collection: AM KS 82629, male (Fig. 4c), KS 82630, 

male (Fig. 5b-e), KS 82631, female (Figs. 6a-g, 7a, b, 

d, h, j-l, 8k), KS 82632, female (Fig. 831, j,n), KS 82633, 

male (Fig. 81), KS 82634, 10 females, 1 male. 


Other material 

NSW: AM KS 82635, Kanangra-Boyd NP, 
Empress Fire Trail turnoff, 33°59’S 150°08’E, M. 
Gray, G. Hunt and J. McDougall, 27.111.1976, 
Eucalyptus pauciflora, AM KS 82636, female (Figs 
4a, 5a), KS 82637, female (Fig. 8b, e), KS 82638, male 
(Fig. 61, j), Monga SF, NSW, Link Rd, 35°34’04"S 
149°54’ 14"E, R. Harris and H. Smith, 16.11.1999; AM 
KS 82639, Buckenbowra SF, Macquarie Rd, 70 m S 
from junction with Milo Rd, 35°38’ 15"S 149°53’27"E, 
1020 m, L. Wilkie and R. Harris, 16.11.1999; AM KS 
82640, Tallaganda SF, South Forest Way, 35°42’50"S 
149°32’?20"E, J. Tarnawski and S. Lassau, 15.11.1999; 
AM KS 82641, Dampier SF, Coomerang Rd, 
36°04’01"S 149°54’57"E, R. Harris and H. Smith, 
11.11.1999; AM KS 82642, Badja SF, Wiola Ck Fire 
Trail, 36°05.56’S 149°35.09’E, J. Tarnawski and S. 
Lassau, 13.11.1999; AM KS 82643, Badja SF, Burkes 


193 


A NEW SPECIES OF HENICOPID CENTIPEDE D/CHELOBIUS 


Figure 4. a-c, Dichelobius giribeti n. sp. a, AM KS 82636, female, Monga SF, NSW. b, holotype AM KS 
82628, female, Badja SF, NSW, terminal segments and gonopods; c, AM KS 82629, male, Badja SF, 
NSW, terminal segments and gonopods. All scales 100 um. 


194 Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


Rd, 36°10733"S 149°31’58"E, J. Tarnawski and S. 
Lassau, 13.11.1999; AM KS 82644, Badja SF, Burkes 
Rd, approx. 1.3 km E from junction with Peters Rd, 
36°10.55’S 149°31.97°E, 992 m, J. Tarnawski and S. 
Lassau, 13.11.1999; AM KS 82645, Bodalla SF, 300 
m along Reservoir Link Rd from junction with Big 
Rock Rd, 36°07.25’S 150°2.82’E, 121 m, L. Wilkie 
and R. Harris, 09.111.1999; AM KS 82646, Bodalla SF, 
Orange Ridge Rd, 36°16’55"S 149°53’31"E, R. Harris 
and H. Smith, 12.11.1999; AM KS 82647, Wadbilliga 
NP, 9.6 km N on Bumberry Ck Fire Trail, 36°14.33’S 
149°33.60’°E, 1059 m, L. Wilkie and R. Harris, 
13.11.1999. 

ANIC (ex. Berl. 855), Kanangra-Boyd NP, 
W Morong Creek, 33°58’S 150°04’E, 1200 m, L. Hill, 
03.x.1982; ANIC (ex. Berl. 829), Kanangra-Boyd NP, 
Kanangra Brook and Rocky Spur, 34°00’S 150°06’E, 
L. Hill, 20.11.1982, closed forest; ANIC (ex. Berl. 852) 
Twin Falls, 14 km SE Moss Vale, 34°39’S 150°28’E, 
600 m, L. Hill, 11.vii.1982; ANIC (ex. Berl. 663), 
Pigeon House Range via Nerriga, 35°02’S 150°08’E, 
J.C. Cardale, 22.x1.1979; ANIC (ex. Berls 2, 18, 34, 
78A, 206A, 222, 246, 468, 657, 851), Clyde Mt, 
35°33’S 149°57°E, 500-c. 800 m, various collections 
1966-1982, dry sclerophyll, wet sclerophyll, rf; ANIC 
(ex. Berl. 877), 2 km N Monga, 35°34’S 149°56’E, 
M.S. Harvey, 18.1x.1983, wet sclerophyll; ANIC (ex. 
Berl. 594), Monga, 35°35’S 149°55’E, JFL and T. 
Weir, 10.111.1978, wet sclerophyll; ANIC (ex. Berl. 
739), Tallaganda SF, 7 km ENE Captains Flat, 35°34’S 
149°31’E, W. Allen, 29.vili.1981; ANIC (ex. Berl. 
1069), Kioloa SF, 35°35’S 150°18’E, JFL and N. 
Lawrence, 4-5.111.1986; ANIC (ex. Berl. 927), Milo 
Forest Preserve, 1.6 km S Monga, 35°36’S 149°55’E, 
L. Hill, 25.x11.1983; ANIC (ex. Berl. 218), 8.8 km ESE 
Captains Flat, 35°38’S 149°31’E, 940 m, RWT, 
10.1.1970, dry sclerophyll; ANIC (ex. Berl. 891), 
Rosedale, 35°49’S 150°14’E, R.J. Moran, 20.xi.1983, 
eucalypt litter; ANIC (ex. Berl. 933), Kosciusko NP, 
1 km ENE Mt Sunrise, 36°22’S 148°29’E, L. Hill, 
411.1984; ANIC (ex. Berl. 935), Kosciusko NP, 4 km 
NNE Mt Perisher, 36°22’S 148°29’E, L. Hill, 411.1984; 
ANIC (ex. Berl. 10), Brown Mt, 36°36’S 149°23’E, c. 
3000 ft., RWT, 5.1.1967, wet sclerophyll; ANIC (ex. 
Berl. 20), Brown Mt, c. 2800 ft., RWT and R.J. Bartell, 
30.11.1967, rf; ANIC (ex. Berl. 24), Brown Mt, 2500- 
3000 ft., RWT and R.J. Bartell, 11.iv.1967; ANIC (ex. 
Berl. 41), Brown Mt, Rutherford Creek, 2700 ft., RWT 
and RJB, 9.x11.1967, rf; ANIC (ex. Berl. 42), Brown 
Mt, c. 3000 ft., RWT and RJB, 9.xii.1967, rf. 

ACT: ANIC (ex. Berl. 283), Black Mt, 
eastern slope, 35°16’S 149°06’E 750 m, J. Simmons, 
26.v.1970, dry sclerophyll; ANIC (ex. Berl. 228), 
Uriarra to Piccadilly Circus, 35°19’S 148°51’E, 700 
m, RWT, 27.1.1970, dry sclerophyll; ANIC (ex. Berl. 


Proc. Linn. Soc. N.S.W., 125, 2004 


225), Uriarra to Piccadilly Circus, 35°20’S 148°50’E, 
500 m, RWT, 16.1.1970, wet sclerophyll; ANIC (ex. 
Berl. 231), Uriarra to Piccadilly Circus, 35°20’S 
148°50’E, 1000 m, RWT, 16.1.1970, wet sclerophyll; 
ANIC (ex. Berl. 999), Wombat Creek, 6 km NE 
Piccadilly Circus, 35°19’S 148°51’E, 750 m, JFL, T. 
Weir and M.-L. Johnson, 30.vi.1984, open forest; 
ANIC (ex. Berl. 1001), Piccadilly Circus, 35°22’S 
148°48’E, 1240 m, JFL, T. Weir and M.-L. Johnson, 
30.vi.1984, subalpine eucalypt litter; ANIC (ex. Berl. 
1000), Blundells Creek, 3 km E Piccadilly Circus, 
35°22’S 148°50’E, 850 m, JFL, T. Weir and M.-L. 
Johnson, 30.vi.1984, open forest; ANIC (ex. Berl. 821), 
Brindabella Range, Franklin Rd, N end Moonlight 
Hollow, 2 km SW Bulls Head, 35°24’S 148°48’E, M.S. 
Harvey and R.J. Moran, 3.iv.1983; ANIC (ex. Berl. 
926), Ginini Flat, 2 km NE Mt Ginini, 35°31’S 
148°46’E, 1580 m, L. Hill, 20.viti.1983; ANIC (ex. 
Berl. 659), Mt Ginini, 35°32’S 148°46’E, 1660 m, JFL 
and T. Weir, 16.x.1979; ANIC (ex. Berl. 1068), 1 km 
S Mt Ginini, 35°33’S 148°46’E, JFL, 11.xi.1986; ANIC 
(ex. Berl. 704, 705), 1 km N Mt Gingera, 35°33’S 
148°47°E, A.A. Calder, 18.11.1981; ANIC (ex. Berl. 
26), Mt Gingera, 35°34’S 148°47’E, c. 5500 ft., E.B. 
Britton, 13.1v.1967, wet sclerophyll; ANIC (ex. Berl. 
50), Mt Gingera, summit, E.B. Britton and Misco, 
19.vii.1967; ANIC (ex. Berl. 661), Mt Gingera, E.C. 
Zimmerman, 20.x1.1979; ANIC (ex. Berl. 830, 831), 
Mt Gingera, 1620-1700 m, L. Hill, 6.11.1982; ANIC 
(ex. Berl. 1084), Snowy Flat Creek, 0.5 km NE Mt 
Gingera, 35°35’S 148°47°E, A.A. Calder, 28.vi.1988. 

VIC: ANIC (ex. Berl. 1045), Cobb Hill, 14 
km SE Bonang, Goonmirk Ra, 37°18’S 148°50’E, JFL 
and N. Lawrence, 24.xi.1985. 

LORD HOWE ISLAND: AM KS 35592, 
NE area of Mt Gower summit, moss forest near 
campsite, 31°35.2’?S 159°04.7°E, 855 m, M.R. Gray, 
12-15.11.1971; AM KS 35589, creek crossing above 
Boat Harbour, 31°33.5’’"S 159°05.5’E, 60 m, M.R. 
Gray, 8.11.1971; AM KS 82998, female (Figs. 6h, 8a, 
d, f, g), KS 82999, male (Figs. 6k, 1, 0, 7g, m, 8c, h, 
m), KS 83000, male (Figs. 6m, n, 7c, e, f, 1), west end 
of Mt Gower summit on south edge, 31°35.32’S 
159°04.2’E, I. Hutton, 15.v.2001; AM KS 84206- 
84233, additional localities/samples on Mt Gower, AM 
KS 84234-84237, four localities on Mt Lidgbird, I. 
Hutton and CBCR, 2000-2002; AM KS 84238, North 
Hummock, trail to Intermediate Hill, 31°32’54"S 
159°04’58"E, CBCR, 3.x1i.2000, mixed rf; AM KS 
84239, western slope of Malabar Ridge, 31°30’57"S 
159°03’31"E, CBCR, 24.xi.2000, broad megaphyllous 
closed sclerophyll forest; AM KS 84240, Transit Hill, 
31°32’01"S 159°04’40"E, I. Hutton, 14.1v.2002; AM 
KS 84241, Little Island, below Far Flats, 31°34’08"S 
159°04’32"E, I. Hutton, 10.viii.2001, under Ficus 


195 


A NEW SPECIES OF HENICOPID CENTIPEDE D/CHELOBIUS 


columnaris. 


Description 

Length (anterior margin of head shield to 
telson) up to 6.6 mm; length of head shield up to 0.7 
mm; leg 15 33-40% length of body. Colour: head shield 
and maxillipede pale orange; antenna and most tergites 
orange-yellow, T14 and tergite of intermediate 
segment deeper orange; legs 1-13 pale yellow to pale 
orange, legs 14 and 15 may be deeper orange. 

Head shield (Fig. 5a) smooth, of equal length 
and width, slightly wider than Tl, median notch 
contributing to biconvex anterior margin; longitudinal 
median furrow incised to transverse suture, about one- 
third length of head shield; posterior two-thirds of 


region distal to antennocellar suture desclerotised; setae 
on head shield arranged with bilateral symmetry, four 
larger pairs anterior to antennocellar suture, ten pairs 
behind suture, including four evenly spaced 
submarginal pairs; head shield lacking posterior and 
lateral borders. 

Antenna 27-32% length of body, 2.5-3.3 
times length of head shield, composed of 14 or 
(usually) 15 articles; basal two articles enlarged, most 
articles in distal half moniliform, sclerotised part 
generally of subequal length and width; ultimate article 
about twice length of penultimate. Basal article bearing 
about a dozen sensilla microtrichoidea proximally on 
dorsal side (Fig. 6a). Trichoid sensilla arranged in three 
whorls per article; one or occasionally two curved, 


Figure 5. a-e, Dichelobius giribeti n. sp.a, AM KS 82636, female, Monga SF, NSW, head shield, maxillipede 
segment and T1; b-e, AM KS 82630, male, legs 12-15, Badja SF, NSW. All scales 100 um. 


196 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


Figure 6. Dichelobius giribeti n. sp. Scanning electron micrographs. a-g, Badja SF, NSW; h, k-o, Mt 
Gower, Lord Howe Island; i, j, Monga SF, NSW. a-g, AM KS 82631, female. a, cluster of sensilla 
microtrichoidea on proximal part of antenna, dorsal side, scale 10 um; b, clypeus, scale 50 um; c, posterior 
part of clypeus and labrum, scale 50 tm; d, labral margin, scale 10 pm; e, antennal articles 10-13, dorsal 
side, scale 30 tum; f, basiconic sensillum at anterior edge of antennal article 12, dorsal side, scale 5 um; g, 
tip of terminal antennal article, scale 10 um. h, AM KS 82998, female, dental margin of maxillipede, 
scales 100 pm, 30 um. i, j, AM KS 82638, male, dental margin and ventral view of maxillipede, scales 50 
pum, 100 pm. k, I, o, AM KS 82999, male. k, porodont, scale 10 um. |, dental margin of maxillipede, scale 
50 um. 0, anterior angle of telopodite of first maxilla, scale 10 tm. m, n, AM KS 83000, male, telopodite of 
maxillipede and detail of tarsungulum, showing sensilla coeloconica, scales 50 um, 5 um. 


Proc. Linn. Soc. N.S.W., 125, 2004 197 


A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 


digitiform sensilla near anterior edge on dorsomedial 
side of a few, variable antennal articles (Fig. 6e); four 
or five articles with a single, short, fusiform sensillum 
at anterior edge on dorsal side (Fig. 6f), most consistent 
on articles 11, 12 and 14; digitiform and fusiform 
sensilla sometimes cooccur on a single article (article 
7 or 9); ultimate article with cluster of 8 or 9 trichoid 
sensilla at apex, one or two curved, digitiform sensilla 
behind apical cluster (Fig. 6g). 

Clypeus with apical cluster of three setae on 
ventral side near lateral margin, single seta medially 
(Fig. 6b); transverse band of four setae in front of 
labrum, outer pair slightly to distinctly smaller than 
inner (Fig. 6c); transverse seta projecting from 
sidepiece; labral margin moderately concave where 
cluster of 7-13 bristles projects; bristles with numerous 
short, spine-like projections along lateral margins and 
on ventral surface along their lengths (Fig. 6d). 
Tomosvary organ large, longitudinally ovate, outer 
edge at lateral margin of cephalic pleurite (Fig. 8k). 

Maxillipede (Figs 6h-n): coxosternal width 
across dental margin 39-44% maximum width; lateral 
margin flexed inward at base of dental projections and 
less convergent than against posterior part; each dental 
margin convex, usually with 5+5, 4+5 or 5+4 teeth, 
sometimes 4+4, 6+5, 5+6 or 6+6; inner tooth smaller 
than others, its apex well posterior to base of outer 
tooth; median notch varying from broadly V-shaped 
(Fig. 6h) to deeply parabolic (Fig. 61); porodont of 
similar length and thickness to largest coxosternal 
setae, its socket at posterolateral edge of outermost 
tooth (Fig. 6k); setae relatively sparsely, fairly evenly 
scattered on coxosternite; tarsal and pretarsal parts of 
tarsungulum of about equal length (Fig. 6m). Dorsal 
and ventral sides of tarsungulum with several sensilla 
coeloconica (Fig. 6n). Bands of pleural collar separated 
by longitudinal median suture (Fig. 6j). 

Mandible: Six curved aciculae (Fig. 7j), all 
with many (up to 18) short, blunt denticles along both 
margins (Fig. 71) on distal half to two-thirds. Four 
paired teeth, dorsal three with accessory denticle field 
delimited by deep groove; dorsalmost tooth and basal 
part of second and third teeth composed of densely 
tuberculate rhomboid and polygonal scales (Fig. 71), 
becoming denticulate near furry pad (Fig. 7m). Fringe 
of branching bristles terminates against dorsalmost 
acicula (Fig. 7f); ventralmost bristles in fringe with 
flattened bases lacking spines, distal two-thirds with 
short spines along both margins and on outer face; 
bristles multifurcating at their distal tips, with three or 
four spines that are longer and thicker than those more 
proximally (Figs 7f, k); more dorsal bristles gradually 
become more uniformly spinose to their broader bases, 
with more numerous distal spines (Fig. 7k), grading 


198 


into wide scales that form a nearly continuous double- 
fringe of hair-like spines, each scale composed of a 
narrow outer fringe and a wider inner fringe, each with 
12-15 spines per scale (Fig. 71); fringe terminates at 
edge of dorsalmost tooth, against a large, smooth scale 
that separates dentate lamina from furry pad (Fig. 7m). 
Furry pad composed of a few scales with distal spines 
and cluster of six or seven mostly simple, elongate 
spines. 

First maxilla: sternite indistinctly delimited 
from coxa (Fig. 7a), short, wide. Coxal projections 
tapering, with rounded apex bearing four or five simple 
setae; one small seta along inner margin near base of 
coxal projection. Telopodite strongly delimited from 
coxal projection; basal article of telopodite with single 
marginal seta anterolaterally or lacking setae; distal 
article with one or two setae near outer margin, anterior 
angle terminating as a long, stout spine; entire inner 
margin fringed with row of six or seven plumose setae 
(Fig. 7b), paired in posterior part of row, with slender 
branchings along more than half of their length (Fig. 
7c); five shorter simple setae inserting near bases of 
plumose setae on ventral side; anterior plumose setae 
fringed on dorsal side by a few elongate spines. 

Second maxilla: anterior margin of coxa 
gently concave; band of four or five small setae across 
anterior part of coxa. Inner edge of tarsus with a row 
of five or six brush-like setae with abundant, slender 
branchings nearly to their bases (Fig. 7d, h). Claw 
composed of up to five digits with concave, scoop- 
like inner surfaces (Fig. 7g); large, curved medial digit 
with furrows or sutures running along its length (Fig. 
7e); outer digits shorter, separated from medial digit 
by a slender, spine-like digit. 

Tergites smooth, all with rounded posterior 
angles, lacking projections; T1 about 85% width of 
widest tergite (TT 10 or 12). Posterior margins of TT1, 
3, 5 and 7 transverse (Fig. 4a); TT8, 10 and 12 gently 
concave; TT9, 11, 13 and 14 transverse to weakly 
concave; tergite of intermediate segment transverse or 
gently concave, posterior angle rounded. Two or three 
moderately long setae on lateral margins of long 
tergites, usually with short setae between these; 
posterior margins of tergites fringed with four to twelve 
setae, generally more abundant on more posterior 
segments (maximal number typically on T13); setae 
on inner part of long tergites include transverse band 
of up to six setae across anterior third, two or three 
pairs in two bands behind this. 

Legs 12-15 (Fig. 5b-e) with length ratios 1: 
1.1 : 1.3-1.4 : 1.7. Leg 15 basitarsus 85-115% length 
of distitarsus (Fig. 5e); basitarsus 70-75% length of 
tibia; tibia 2.9-3.4 times longer than maximal width, 
basitarsus 3.4-4 times, distitarsus 5.2-5.7 times. 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


Figure 7. Dichelobius giribeti n. sp. Scanning electron micrographs. Scales 10 um except where indicated. 
a, b, d, h, j-l, AM KS 82631, female, Badja SF, NSW; c, e-g, i, m, Mt Gower, Lord Howe Island. a, ventral 
view of first maxillae, scale 50 pm; b, distal article of telopodite of first maxilla; d, h, tarsus and claw of 
second maxilla, scales 10 um, 30 um; j, aciculae; k, 1, ventral and dorsal parts of fringe of branching 
bristles on mandible. c, e, f, i, AM KS 83000, male. c, plumose setae on inner margin of telopodite of first 
maxilla; e, claw of second maxilla; f, aciculae and fringe of branching bristles on mandible; i, aciculae. g, 
m, AM KS 82999, male. g, claw of second maxilla, scale 10 um; m, dorsalmost tooth of mandible and 


furry pad. 


Basitarsus 90% length of distitarsus on leg 14 (Fig. 
5d). Coxal projections on leg 15 tapering (in ventral 
view) at about 25-30 degrees; terminal spine with 
distinct (Fig. 8e) or indistinct (Fig. 81) basal joint, its 


Proc. Linn. Soc. N.S.W., 125, 2004 


surface with fine longitudinal grooves and ridges like 
those on pretarsal claws. Trochanter of leg 15 with 
small ventrodistal spur (Figs 5e, 8h). Prefemur of legs 
14 and 15 with large ventrodistal spur; leg 15 spur 


ey) 


A NEW SPECIES OF HENICOPID CENTIPEDE DICHELOBIUS 


with basal width about 25% maximum width of 
prefemur (Fig. 4b). Sharp distal spinose projections 
on tibiae of legs 1-11, absent on legs 12-15. Two 
tarsomeres of leg 13 defined by distinct constriction 
in width and weak articulation without flexure; 
articulation between tarsomeres stronger on leg 14. 
Setae fairly evenly distributed on all podomeres along 
leg, tarsal setae only slightly more slender than those 
on prefemur-tibia; proximo-distal gradient in setal 
thickness enhanced on legs 14 and, especially, 15, with 
distinctly thickened prefemoral setae, including on 
dorsal side of leg. Anterior and posterior accessory 
claws present on all legs, 25-40% length of main claw 
(Fig. 8a, b); accessory claws with closely-spaced linear 
ridges on their surface except for pitted proximoventral 
part separated by a shallow suture (Fig. 8c). Main claw 
curved, subdivided by sutures; deepest sutures define 
an elongate scute on both lateral sides of claw, proximal 
end of this scute at about distal end of shorter accessory 
claw; large pore or pair of pores at proximal end of 
scute on both sides of leg (Fig. 8c); strong suture 
extends from lateral pore across ventral surface of main 
claw (Fig. 8d), defining proximal end of an elongate, 
triangular ventral scute (Fig. 8g). Proximal part of main 
claw densely pitted; on ventral side of claw, ornament 
changes abruptly at suture delimiting lateral scute, 
becoming linear grooves and ridges as on accessory 
claws (Fig. 8d), with these lineations well developed 
on lateral scute and along length of claw on dorsal 
side; change from pitted to linear ornament gradual 
on dorsal side of claw, with pits irregular proximally, 
becoming aligned as rows of pits, then linear grooves. 
Pair of distally-directed spines proximoventrally, at 
distal end of a curved suture (Fig. 8d); larger spine not 
more than not more than one-eighth length of main 
claw, with tiny subsidiary spine at its base (Fig. 8b). 

Coxal pores: on legs 14 and 15; 2,2/2,2 in 
females (Fig. 4b), 1,1/1,1 in small males, either 1,1/ 
1,1 or 2,2/2,2 (Fig. 4c) in large males, occasionally 
one and two pores on opposing sides of either leg or 
1,2/1,2; pores round, separated by less than their 
diameter when paired; inner pore often smaller than 
outer pore in male, inner pore sometimes larger than 
outer pore in female. 

Female (Fig. 4b): Sternite of segment 15 
gently convex posteromedially, fringed by a 
submarginal setal band that extends along entire 
posterolateral and posterior margin; several setae 
scattered on inner part of sternite. Posterior margin of 
first genital sternite moderately embayed between 
gonopod articulations, sternite bearing 6-11 setae. 
Gonopod with pair of spurs at terminus of a short (Fig. 
8n) to moderately long (Fig. 8e, f) projection; bases of 
spurs nearly touching each other; inner spur 


200 


substantially shorter and narrower than outer spur, both 
bullet-shaped, pointed (Fig. 8n); four or five setae on 
basal article of gonopod, three large setae on second 
article, one large seta on third (Fig. 8j); second and 
third articles variably with one and two smaller setae, 
respectively, on ventromedial face (Fig. 8n); claw 
simple. 

Male (Fig. 4c): Posterior margin of sternite 
15 evenly convex; 10-13 setae fringing margin of 
sternite, 10-12 additional setae scattered over its ventral 
surface; first genital sternite entire medially, bearing 
6-12 setae aligned in two imprecisely-defined 
transverse rows; gonopod bearing two or three setae 
on first article, two on second article, none or one on 
third article, which grades into long, flagelliform 
terminal process, up to 80% length of rest of gonopod 
(Fig. 81); terminal process bearing numerous slender 
spines proximally (Fig. 8m). 

Larvae: five larval stadia (ANIC Berl. 18 and 
231) identified as LO-LIV by comparison to limb 
development in other Lithobiomorpha (Table 1). LI 
with 11 antennal articles; LII-LIV all with 14 articles. 
LII and LIII with 2+2 teeth on dental margin of 
maxillipede; LIV with 3+3 teeth. 


Discussion 

Specimens from Lord Howe Island resemble 
those from the Australian mainland in all meristic 
characters and in fine detail. Intrapopulation variation 
is observed with respect to the number of teeth on the 
maxillipede coxosternal margin, the depth of the 
median notch in the maxillipede coxosternite 
(relatively shallow in Fig. 6h, relatively deep in Fig. 
61), the concavity of the posterior margins of the short 
tergites, and the length of the spur-bearing process on 
the female gonopod. Samples vary in the frequency 
with which large males have either one or two coxal 
pores on legs 14 and 15 (usually two in Lord Howe 
specimens versus one in the large sample from Clyde 
Mountain, NSW, but also two in large specimens from 
the type locality and in the Brindabella Range, e-.g., 
Piccadilly Circus, Mt Gingera and Mt Ginini). 

Distinction from other congeners is indicated 
in key above. Dichelobius relictus and D. giribeti are 
consistently distinguished by the presence of a spiracle 
on segment 14 in the former, and D. relictus is 
generally a deeper brown colour. The two species share 
minute details of mandibular and maxillary structure, 
indeed to the extent that description of the mouthparts 
for D. giribeti serves for D. relictus as well. 

The early larval stadia of Dichelobius giribeti 
differ in detail from those of Lithobiidae and 
Henicopinae (see Table 1) with respect to limb 
development. Segmentation of LO is matched by 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


Figure 8. Dichelobius giribeti n. sp. Scanning electron micrographs. a, c, d, f-h, m, Mt Gower, Lord Howe 
Island; b, e, Monga SF, NSW; i-l, n, Badja SF, NSW. a, d, f, g, AM KS 82998, female. a, pretarsus of leg 
14, scale 10 pm. d, g, ventral views of pretarsus of leg 14, scales 10 um; f, gonopods, scale 30 um. b, e, AM 
KS 82637, female. b, pretarsus of leg 14, posterior view, scale 10 [1m; e, ventrolateral view of first genital 
sternite and gonopods, scale 100 tum. c, h, m, AM KS 82999, male. c, pretarsus of leg 15, detail of anterior 
accessory claw, scale 5 um; h, prefemur of leg 15, anterior side, scale 100 um; m, terminal process on 
gonopod, scale 10 um. i, j, n, AM KS 82632, female. i, leg 15 coxal process, scale 30 Lum; j, n, lateral and 
ventral views of gonopod, scales 50 tm, 10 pm. k, AM KS 82631, female, cephalic pleurite with Tomoésvary 
organ, scale 50 um. 1, AM KS 82633, male, gonopod, scale 30 [um. 


Proc. Linn. Soc. N.S.W., 125, 2004 201 


A NEW SPECIES OF HENICOPID CENTIPEDE D/ICHELOBIUS 


Table 1. Comparison of limb development in larval stadia of Lithobiomorpha. Modified from Andersson 
(1979:Table I), adding data for Dichelobius giribeti. 


Lamyctes 
emarginatus 
Lithobius 8 spp. 


oe. a io ee 


Lamyctes coeculus, but larval stadium LI has a unique 
combination of half-developed legs and limb-buds in 
D. giribeti. Segmentation of stadia LI-IV is as in other 
lithobiomorphs. Four larval stadia identified by Eason 
(1993) for Anopsobius macfaydeni have seven, eight, 
ten and twelve pairs of legs, the last three obviously 
being LII-LIV. The taxonomic significance of the 
distinction between six- and seven-legged first larval 
stages in Dichelobius giribeti and Anopsobius 
macfaydeni is unclear without additional data for 
Anopsobiinae. 


Dichelobius bicuspis Ribaut, 1923 


Dichelobius bicuspis Ribaut, 1923:24, Figs. 27-34. 
Dichelobius bicuspis: Wiirmli, 1974:526. 


Material 

NEW CALEDONIA: PROV. NORD: AM 
KS 83001, 1 female, 1 male, Mt Panié, nr summit, 
20°34’S 164°46’E, 1500 m, C. Burwell, 9.xi.2001, rf; 
MNHN, | female, 1 larval stadium LIV, Mt Panié, 
20°34’53"S 164°45’38"E, 1350 m, J. Chazeau, A. & 
S. Tillier, 18.xi1.1986, wet Agathis forest; QM S60653, 
1 female, Pic d’Amoa, N slopes, 20°58’S 165°17’E, 
500 m, GBM, 10.xi.2001, rf; QM S60654, 1 male, Me 
Maoya, summit plateau, 21°12’S 165°20’E, 1400 m, 
GBM, 12.xi.2002, rf. PROV. SUD: MNHN, 3 females, 
Mt Do, 21°45’37"S 165°59°33"E, 840 m, A. & S. 
Tillier & Monniot, 2.iv.1987, wet Araucaria forest; 
QM S60655, 1 male, Mt Humboldt refuge, 21°53’S 
166°24’E, 1300 m, GBM, 7-8.xi.2002, rf; AM KS 
83002, 1 male, R Bleue, Pourina Track, 22°04’S 
166°38’E, 900 m, GBM, 18.xi.2001, rf; AM KS 83003, 
1 male, Mt Ouin, 22°01’S 166°28’E, 1100 m, GBM, 
9.xi.2002, rf; AM KS 83004, 1 female, 1 male, QM 


202 


Lamyctes 
coeculus 


Dichelobius 
giribeti 


S60656, 1 male, Mt Mou base, 22°05’S 166°22’E, 200 
m, GBM, 30.x.2001, 15.xi.2001, rf; MNHN, 3 females, 
1 juvenile, Riviére Bleue, 22°06’13"S 166°39°16"E, 
160 m, A. & S. Tillier, 1.viii.1986-30.iv.1987; QM 
S60657, 1 male, Mt Koghis, 22°11’S 166°01’E, 750 
m, GBM, 29.xi.2000, rf; AM KS 83005, 1 female, 
Yahoué, 22°12’S 166°30’E, 100 m, GBM, 4.x1.2001, 
rf. 


Remarks 

Dichelobius bicuspis was based on a few 
specimens from Mt Humboldt (the type locality) and 
Mt Canala, New Caledonia, with Wiirmli (1974) 
adding a record at Nékliai. New collections are listed 
above to indicate that the species has a more 
widespread distribution. 


ACKNOWLEDGEMENTS 


I thank Suzanne Bullock (Scientific Interface) and Sue 
Lindsay and Yongyi Zhen (Australian Museum) for 
assistance with illustrations and electron microscopy, and 
the referees for useful suggestions. Geoff Monteith 
(Queensland Museum) and Jean-Paul Mauriés (Museum 
National d’ Histoire Naturelle, Paris) kindly provided material 
from New Caledonia. Collection study was hosted and loans 
were arranged by Matthew Colloff (Australian National 
Insect Collection), Gonzalo Giribet and Laura Leibensperger 
(Harvard University), and Mark Harvey and Julianne 
Waldock (Western Australian Museum). Verena Stagl 
(Naturhistorisches Museum Wien) is thanked for arranging 
loan of C. Attems’ types. 


Proc. Linn. Soc. N.S.W., 125, 2004 


G.D. EDGECOMBE 


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203 


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A Survey of Ectoparasite Species on Small Mammals During 
Autumn and Winter at Anglesea, Victoria 


Haylee J Weaver!” and John G Aberton’. 


'School of Ecology and Environment, Deakin University, Geelong VIC 3217; *Present address: School of 
Biological and Environmental Sciences, Central Queensland University, Rockhampton QLD 4702 
(h.weaver @cqu.edu.au) 


Weaver, H.J. and Aberton, J.G. (2004). A survey of ectoparasite species on small mammals during autumn 
and winter at Anglesea, Victoria. Proceedings of the Linnean Society of New South Wales 125, 205-210. 


A survey of the ectoparasites of small native mammals was carried out between April and August 2002, 
in heathlands surrounding Anglesea, Victoria. Antechinus minimus, A. agilis, Rattus lutreolus, R. fuscipes, 
Sminthopsis leucopus and Isoodon obesulus were the dominant host mammal species examined. A total of 
921 ectoparasites were collected and identified as five flea species, seven mite species and two species of 
tick. Isoodon obesulus was found to have the highest ectoparasite species richness, with eleven of the 
fourteen species present; while S. leucopus displayed the lowest ectoparasite species richness with only 
three species found on the hosts examined. The flea Pygiopsylla hoplia was the only ectoparasite species in 
this study to have a distribution across all host mammal species. A new distribution record was made for a 


Haemaphysalis tick species. 


Manuscript received 16 October 2003, accepted for publication 8 January 2004. 


KEYWORDS: Anglesea, ectoparasites, host specificity, marsupials, rodents, species richness. 


INTRODUCTION 


The main groups of ectoparasitic arthropods 
encountered on Australian mammals include fleas 
(order Siphonaptera), mites (order Acariformes), ticks 
(order Parasitiformes) and lice (order Phthiraptera). 
These ectoparasites, as a group, have evolved 
specialised piercing and sucking mouthparts, designed 
for the extraction of blood from a host, with the degree 
of host specificity displayed by ectoparasites varying 
amongst species (Kemp et al. 1982; Dunnet and 
Mardon 1991). 

Many species of ectoparasites are of 
considerable medical and veterinary importance. Fleas 
are capable of transmitting various rickettsial, filarial 
and protozoan diseases (Dunnet and Mardon 1991), 
and ticks can transmit pathogenic filariae, bacteria, 
protozoa, rickettsiae and viruses to wild and domestic 
animals and humans (Obenchain and Galun 1982; 
Aeschlimann 1991). Previous research on ectoparasites 
in the Anglesea region has been limited to flea surveys 
as a precursor to the introduction of myxomatosis 
(Dunnet and Mardon 1991) and calicivirus (F. 
Bartholomaeus pers. comm.), and basic natural history 
of ticks (Roberts 1970). Ectoparasites also negatively 
impact on the health of both domestic and wild animals 
through large infestations, which are of importance in 


management considerations of rare or endangered 
small mammal species present at Anglesea as increases 
in host densities may increase ectoparasite loads. 

The objective of this study was to survey 
ectoparasite species on small native mammals near 
Anglesea, Victoria because an awareness of the 
ectoparasites is important for the potential transmission 
of disease to humans, domestic animals and livestock. 
It is also of interest to the general ecology of small 
mammals in the region. 


METHODS 


Ectoparasites were removed from small 
mammals trapped at two sites at Anglesea, Victoria 
(Fig. 1). The sites chosen for study were the Eumeralla 
Scout camp (38°24’0”S, 144°12’36”E) and Bald Hills 
Road (38°23’24”S, 144°8’24”E) at the Alcoa Lease. 
Both sites were selected using knowledge that they 
contained many host species, and these species were 
all relatively abundant. The Eumeralla Scout camp 
consisted of a coastal tea tree, Leptospermum 
continentale shrub layer, with plants varying from 20 
centimetres to over two metres in height and 
Eucalyptus obliqua at a height of over two metres 


ECTOPARASITES ON SMALL MAMMALS 


Victoria 


Bald Hills Rd 


Eumeralla 


Alcoa Lease 
boundary —> 


Anglesea 


Great Ocean Rd 


Airey's Inlet 


Figure 1. Location map of study area. 


forming the canopy. The site was a flat open heathland 
with woodland dispersed through it, and a swamp 
consisting mainly of Gahnia radula and also L. 
continentale. The Bald Hills Rd site on the Alcoa Lease 
was situated on a slope of approximately 30°in a 
southwesterly direction. The heathland was dominated 
by L. continentale, Epacris impressa, Conospermum 
mitchelli, L. myrsinoides, Platylobium obtusangulum 
and G. radula were the main species present in the 
understorey. Stands of Eu. willisi and Banksia 
marginata were present at the study site. 

Trapping of small mammals was carried out 
during Autumn and Winter 2002, due to the study 
being an honours project requiring completion during 
an academic year. Trapping sessions of three nights 
each were carried out at Eumeralla in June (3-6.6.02), 
July (8-11.7.02) and August (6-9.8.02), with a total of 
131 mammals captured over the three sessions and at 
the Bald Hills Rd site in April/May (29.4-2.5.02), July 
(22-25.7.02) and August (19-22.8.02), with 161 


206 


captures recorded. Any previously 
trapped mammals captured again in 
following sessions were re-examined 
for ectoparasites and were counted 
accordingly. Fifty aluminium Elliott 
traps (32 x 9 x 10 cm) were placed in 
transects across the Eumeralla site. 
The site at Bald Hills Rd consisted of 
100 traps set in a grid pattern (100 m 
x 100 m) at ten metre intervals. Traps 
were baited using a rolled oats, peanut 
butter and honey mix and were cleared 
within three hours of sunrise. 

Upon capture, mammals 
were transferred from the trap into a 
lightweight mesh bag, identified, ear 
notched for identification purposes, 
weighed, sexed and inspected for 
ectoparasites. As ticks were physically 
attached to the host, they were 
removed using fine forceps to grip the 
tick as close to the host’s skin as 
possible and flipping it over to remove 
the tick while leaving the mouthparts 
intact. Fleas and mites were removed 
by ruffling the host’s pelage with 
fingers in order to dislodge the 
ectoparasites, or the host was combed 
using Licemeister combs or animal 
flea combs. Numbers of each 
ectoparasite taxa were recorded from 
each mammal and all ectoparasites 
collected were placed in labelled 
containers of 70% ethanol. 

Identification of fleas, mites 
and ticks were carried out using descriptions provided 
by Dunnet and Mardon (1974), Domrow (1987, 1991) 
and Roberts (1970) respectively. 

A linear regression on host mammal body 
weight and ectoparasite species richness was carried 
out using log transformed data. 


RESULTS AND DISCUSSION 


A total of 292 individual mammals were 
trapped over 1350 trap nights from the two sites. The 
host mammals trapped included Antechinus minimus 
(74), A. agilis (69), Sminthopsis leucopus (4) 
(Dasyuriomorphia: Dasyuridae), Isoodon obesulus 
(10) (Peramelemorphia: Peramelidae), Rattus fuscipes 
(50) and R. lutreolus (85) (Rodentia: Muridae). 

Examination of 296 host mammals yielded 
364 fleas and 557 acari (mites and ticks) in total. From 
this, five flea species were identified, along with seven 


Proc. Linn. Soc. N.S.W., 125, 2004 


H.J. WEAVER AND J.G. ABERTON 


mite species and two tick species. Of these, two species 
of mites were unable to be identified to species level; 
these were referred to by their family names as 
unidentified Laelapidae and _ unidentified 
Trombiculidae. Table 1 shows the number of 
examinations of each host mammal species and the 
species of ectoparasites removed from the host species. 

The most common host examined for 
ectoparasites was Rattus lutreolus, with 85 
examinations and the host examined least was 
Sminthopsis leucopus with only four examinations. 
Sminthopsis leucopus is an uncommon mammal in the 
Anglesea area. Lunney (1995) states although it has a 
wide distribution throughout southern Australia, it 
prefers sparse ground to forage, whereas the sites in 
this study had very dense ground cover. 

Figure 2 shows J. obesulus as having the 
greatest ectoparasite species richness and S. leucopus 
the smallest. A significant linear association was found 
between host weight and ectoparasite species richness 
(MS=0.098, F=7.966, df=1, P=0.048) with 64.32% of 
the variation in ectoparasite species richness accounted 
for by mean body weight of the hosts. This is consistent 
with previous studies showing that host body size 
determines ectoparasite species richness (Kuris et al. 
1980, cited in Stanko et al. 2002). Another factor that 
can influence ectoparasite species richness is the social 
behaviour of the host. Stanko et al. (2002) found that 
higher host densities generally equated to lower species 
richness on individuals, possibly because of anti- 
parasitic behaviours such as grooming. As bandicoots 
have a reputation of “pugnacious behaviour between 
conspecifics’ (Lobert 1990) and indicate a low social 
tolerance (Thomas 1990), it could be that the 
bandicoots examined in this study had a higher species 
richness of ectoparasites and a higher abundance of 
each species in part due to a combination of larger 
body size and lack of social grooming. 

The most common ectoparasite collected was 
the flea Pygiopsylla hoplia, which was recorded on 
every host mammal species. According to Dunnet and 
Mardon (1974), P. hoplia is the most commonly 
collected Australian species of flea. It has a distribution 
across Australia, excluding the Northern Territory, and 
has been recorded on many species of peramelids, 
dasyurids and rodents (Dunnet and Mardon 1974). In 
contrast, Stephanocircus dasyuri was mostly recorded 
on J. obesulus, and occasionally on A. minimus. The 
similar foraging nature of both these mammal species 
may be the reason why this species of flea was not 
recorded on any other hosts. Macropsylla hercules was 
only recorded on Rattus spp. and I. obesulus, perhaps 
due to the size of the host animals, as this flea is very 
large. Macropsylla hercules is commonly collected 


Proc. Linn. Soc. N.S.W., 125, 2004 


from various native Rattus species from southern 
Australia (Dunnet and Mardon 1974). The other 
species of flea collected, Acanthopsylla rothschildi 
rothschildi and Bibikovana rainbowi appeared to 
display little host specificity, as they were recorded 
from the majority of the host species. 

Host specificity for acarine ectoparasites 
collected varied. The highly host specific Androlaelaps 
marsupialis was only found between the groove of the 
tibia and fibula on the hind legs of J. obesulus where 
grooming is difficult (pers. obs.). Similarly, 
Mesolaelaps anomalus was recorded only on J. 
obesulus. In contrast, the trombiculid mites and the 
tick Ixodes tasmani showed a broad host range, being 
found on all host species except for S. leucopus and A. 
minimus respectively. The trombiculids were found 
most frequently inside the ears of hosts during this 
study, but can be found on any exposed skin including 
legs, feet and tails (pers. obs.). Trombiculid mites are 
parasitic during their larval stage and later live in the 
soil as free living adults (Domrow 1962). One small 
infestation was recorded in the pouch of a female /. 
obesulus, and it has been suggested that larval 
trombiculids occurring in the pouches of A. minimus 
can directly infest any pouch young present (B. Wilson, 
Deakin University, pers. comm.). Jxodes tasmani is a 
common species of tick with a distribution widespread 
across southern Australia with a wide range of hosts 
(Roberts 1970). 

The species of Haemaphysalis collected from 
I. obesulus was identified as H. humerosa, but 
differences in the spiracular plate between the Anglesea 
specimens and specimens from known populations in 
Queensland have been observed. An alternative 
identification is H. ratti. Further research is being 
carried to provide a definite identification of the 
specimens (I. Beveridge, University of Melbourne, 
pers. comm., D. Kemp, CSIRO, pers. comm). 

Other ectoparasitic arthropods were collected 
from host mammals studied. Lice (Phthiraptera, species 
unknown) were collected from R. lutreolus on three 
occasions; but were not observed on any other host 
mammals examined. The rove beetle species 
Myotyphlus jansoni (Coleoptera: Staphylinidae) was 
collected from Rattus lutreolus on five occasions. 
However, M. jansoni is not an obligate ectoparasite. 
Myotyphlus jansoni has only been recorded on a very 
small number of individual native Rattus species 
previously (Hamilton-Smith and Adams 1966). The 
beetles are usually collected near the anus or tail (as 
they were in this study) and have also been recorded 
in bat guano in a cave near Warrnambool, Victoria; 
thus it may be assumed that the beetles feed on the 
excreta of the rats, which is not strictly an ectoparasitic 


207 


ECTOPARASITES ON SMALL MAMMALS 


Host species No. of mammals Body weight (g) Siphonaptera Number Acari Number 
examined (Mean + SD) 
Antechinus minimus 74 50 + 13 Pygiopsylla hoplia 36 Andreacarus tauffliebi 5 
(swamp antechinus) Acanthopsylla rothschildi 5 Mesolaelaps sminthopsis 1 
Bibikovana rainbowi 1 Androlaelaps telemachus 5 
Stephanocircus dasyuri 3 Trombiculidae 17 
Antechinus agilis 67 Sire 9 Pygiopsylla hoplia 11 Mesolaelaps sminthopsis 4 
(agile antechinus) Acanthopsylla rothschildi 39 Androlaelaps telemachus 3 
Trombiculidae 8 
Ixodes tasmani 6 
Rattus fuscipes 50 106 + 23 Pygiopsylla hoplia 4 Androlaelaps telemachus 1 
(bush rat) Bibikovana rainbowi 5 Trombiculidae 23 
Macropsylla hercules 7 Ixodes tasmani 7 
Rattus lutreolus 85 95+21 Pygiopsylla hoplia 3 Andreacarus tauffliebi 50 
(swamp rat) Bibikovana rainbowi 7 Mesolaelaps sminthopsis 2 
Macropsylla hercules 6 Trombiculidae 12 
Ixodes tasmani 2 
unidentified Laelapidae 10 
Sminthopsis leucopus 4 Dit 3 Pygiopsylla hoplia 1 Ixodes tasmani 1 
(white footed dunnart) Acanthopsylla rothschildi 1 
Isoodon obesulus 10 400 + 173 Pygiopsylla hoplia 90 Androlaelaps marsupialis 10 
(southern brown bandicoot) Acanthopsylla rothschildi 2 Mesolaelaps anomalus 55 
Bibikovana rainbowi 2 Mesolaelaps sminthopsis 317 
Macropsylla hercules 1 Trombiculidae 5) 
Stephanocircus dasyuri 140 Ixodes tasmani 2 
Haemaphysalis sp. 11 


Table 1. Species of ectoparasites collected from host mammals. 


Proc. Linn. Soc. N.S.W., 125, 2004 


208 


H.J. WEAVER AND J.G. ABERTON 


Total parasite spp. 


y = 0.017x + 5.722 
R’ = 0.643 


0 50. ~=—-:100 


150 200 250 


300 


350 400 


Mean body weight (g) 


Figure 2. Relationship between ectoparasite species richness and body weight of host mammals. SI = 
Sminthopsis leucopus, Aa = Antechinus agilis, Am = Antechinus minimus, R\ = Rattus lutreolus, Rf = Rattus 


fuscipes, lo = Isoodon obesulus. 


relationship (Hamilton-Smith and Adams 1966; 
Lawrence and Britton 1991). 

The ectoparasite species collected during this 
study were all considered to be common throughout 
the region (Roberts 1970; Dunnet and Mardon 1974) 
and all are theoretically able to transmit pathogens to 
animals or humans. Generally, fleas are known to be 
intermediate hosts for the cosmopolitan rodent 
tapeworm, Hymenolepis diminuta and the canine 
tapeworm Dipylidium canium, along with being able 
to transmit various filarial, rickettsial and protozoan 
pathogens, however native Australian fleas have not 
been found to contribute epizootics in the field (Dunnet 
and Mardon 1991). Ixodes tasmani has been recorded 
as an intermediate host of various rickettsiae, including 
Rickettsia australis, the organism which causes 
Queensland tick typhus (Campbell and Domrow 1974, 
cited in Cavanagh 1999). Haemaphysalis ticks are 
vectors of Coxiella burnetii (Q fever), in bandicoots 
and macropods and domestic livestock (Kettle 1995). 
Therefore it is recommended that care be taken when 
in areas where ticks are present, especially at the 


Proc. Linn. Soc. N.S.W., 125, 2004 


Eumeralla Scout Camp where groups of scouts may 
come into contact with ticks while carrying out 
activities in the area. 

In conclusion, it was found that there was a 
significant relationship between ectoparasite species 
richness and body weight of host mammal species. 
There was no difference in the species of ectoparasites 
collected from both study sites, except for M. jansoni, 
which was only found on R. lutreolus at the Bald Hills 
Rd site. As there have been no other studies carried 
out of this type in the region, it is recommended that a 
study over a longer time frame be carried out in order 
to accurately assess seasonal variations of ectoparasite 
numbers. 


ACKNOWLEDGEMENTS 


The authors are grateful to F. Bartholomaeus 
(South Australian Research and Development Institute) and 
M. Shaw (University of Queensland) for additional flea and 
mite identification respectively, and Dr lan Beveridge 
(University of Melbourne) and Dr David Kemp (CSIRO) 


209 


ECTOPARASITES ON SMALL MAMMALS 


for assistance with the identification of Haempahysalis sp. 
We appreciate Philip Barton’s comments on the manuscript. 
The study formed part of the principal author’s BEnvSe 
Honours thesis and was carried out in accordance with 
Deakin University Animal Ethics committee guidelines and 
conducted under Victorian Department of Natural Resources 
and Environment wildlife permit no. 10001759. 


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(Oxford University Press: Oxford). 

Campbell, R.W. and Domrow, R. (1974) Rickettsioses in 
Australia: Isolation of Rickettsia tsutsugamushi 
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Cavanagh, F.A. (1999) The common marsupial tick, 
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degree of infestation of the common brushtail 
possum, Trichosurus vulpecula. BSc Honours 
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Domrow, R. (1962). The mammals of Innisfail. 2: Their 
mite parasites. Australian Journal of Zoology 2, 
268-307. 

Domrow, R. (1987). Acari Mesostigmata parasitic on 
Australian vertebrates: An annotated checklist, 
keys and bibliography. Invertebrate Taxonomy 1, 
817-948. 

Domrow, R. (1991). Acari Prostigmata parasitic on 
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keys and bibliography. Invertebrate Taxonomy 4, 
1283-1376. 

Dunnet, G.M. and Mardon, D.K. (1974). A monograph of 
Australian fleas (Siphonaptera). Australian 
Journal of Zoology Supplementary Series 30, 1- 
PIB 

Dunnet, G.M. and Mardon, D.K. (1991). Siphonaptera. In 
‘The insects of Australia: a textbook for students 
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716. (Melbourne University Press: Melbourne). 

Hamilton-Smith, E. and Adams, D.J.H. (1966). The 
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jansoni (Mathews) (Coleoptera: Staphylinidae) 
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Secretions and the Host Response. In ‘Physiology 
of Ticks’. (Eds F.D. Obenchain and R. Galun) 
pp.1 19-168. (Pergamon Press: Oxford). 

Kettle, D.S. (1995) ‘Medical and Veterinary Entomology 
(2" edn.)’. (CAB International: United Kingdom). 

Kuris, A.M., Blaustein, A.R. and Alio, J.J. (1980). Hosts 
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Lawrence, J.F. and Britton, E.B. (1991). Coleoptera. In 
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683. (Melbourne University Press: Melbourne). 

Lobert, B. (1990). Home range and activity period of the 
Southern brown Bandicoot (Jsoodon obesulus) in 
a Victorian heathland. In ‘Bandicoots and Bilbies.’ 
(Eds J.H. Seebeck, P.R. Brown, R.L. Wallis and 
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Lunney, D. (1995). White-footed Dunnart. In “The 
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Stanko, M., Miklisova, D., Goiiy de Bellocq, J. and 
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Thomas, L.N. (1990). Stress and population regulation in 
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Proc. Linn. Soc. N.S.W., 125, 2004 


Occurrence and Conservation of the Dugong (Sirenia: 
Dugongidae) in New South Wales 


StmMon ALLEN!, HELENE MArRSH2? AND AMANDA HopDGSsoNn?3 


1Graduate School of the Environment, Macquarie University, NSW 2109; 2School of Tropical Environment 
Studies and Geography, James Cook University, Townsville, Qld 4811; 3CRC Reef Research Centre, PO Box 


772, Townsville, Qld 4810 


Allen, S., Marsh, H. and Hodgson, A. (2004). Occurrence and conservation of the dugong (Sirenia: 
Dugongidae) in New South Wales. Proceedings of the Linnean Society of New South Wales 125, 211- 
216. 


Recent sightings of dugongs well beyond the southern limit of their accepted range (~27°S) on the Australian 
east coast prompted a review of past records of dugongs and their current conservation status in New South 
Wales. While archaeological analyses have identified bones of Dugong dugon in Aboriginal middens at 
Botany Bay (~34°S) and colonial records indicate stranded animals as far south as Tathra (~36.5°S), there 
were no verified sightings of live individuals in NSW waters for some years; however, five separate sightings 
of individuals and pairs were documented in the austral summer of 2002/03 in estuaries on the NSW central 
coast (~32-33.5°S). It is suggested that conditions such as warm sea temperatures and low rainfall (promoting 
seagrass growth) may be facilitating explorative ranging south by dugongs. 

The IUCN lists dugongs as ‘vulnerable’ at a global scale and they are also classified ‘vulnerable’ under 
the Threatened Species Conservation Act NSW 1995, yet they are not routinely considered in risk assessments 
for inshore development in this State. Threatening processes such as shark meshing persist. The importance 
of considering dugongs in future impact assessments for inshore marine and estuarine developments is 


emphasized. 


Manuscript received 17 October 2003, accepted for publication 8 January 2004. 


KEYWORDS: conservation, distribution, dugong, Dugong dugon, risk assessment, sightings, status, 


vulnerable. 


INTRODUCTION 


The dugong (Dugong dugon), along with all 
other extant Sirenians, is regarded as a shallow water, 
tropical and sub-tropical species (Martin and Reeves 
2002; Rice 1998). Dugongs are thought to be strictly 
marine, inhabiting the coasts of some 37 countries and 
territories (Marsh et al. 2002). Despite their widespread 
distribution, dugong numbers have declined in most 
of their known range and they are believed to be 
represented by fragmented, relic populations in most 
countries. Likely causes for this decline and continuing 
threats include: large-scale destruction of seagrass as 
a result of sedimentation, dredging, mining, trawling, 
and pollution; incidental take as by-catch in 
commercial and recreational gill and mesh nets as well 
as shark nets set for bather protection; direct takes from 
indigenous hunting, and vessel strikes and disturbance 
(Marsh et al. 1999, 2002; Hodgson 2003). 

Australian waters are the dugong’s 
stronghold, where their distribution is described as 
extending from Shark Bay in Western Australia (25°S) 
around northern Australia to Moreton Bay in southern 
Queensland (27°S) (Marsh et al. 2002). Dugongs are 


a ‘listed marine species’ under the Australian 
Environment Protection and Biodiversity Conservation 
Act 1999 (EPBC Act). The EPBC Act reflects 
Australia’s commitments under various international 
conventions including the Bonn Convention on the 
Conservation of Migratory Species of Wild Animals, 
which lists the dugong on Appendix 2. Dugongs are 
also considered ‘vulnerable’ under the Threatened 
Species Conservation Act NSW 1995 and under the 
Nature Conservation Act Qld 1992. 

Evidence of a decline in dugong numbers 
along the urban coast of Queensland (Marsh et al. 
2001) led to the establishment of a series of dugong 
protection areas in some key dugong habitats in 
Queensland (Marsh et al. 1999; Marsh 2000). No 
similar protection has been afforded dugongs in NSW, 
presumably on the assumption that only vagrants of 
the species range into NSW waters. Dugongs have been 
considered in some impact assessments for aquaculture 
developments in NSW (e.g. Anon. 2001a), but not 
others (e.g. Anon. 2001b). These assessments occurred 
in the same location, suggesting consideration of 
dugongs and potential impacts thereon is inconsistent 
in NSW. 


DUGONGS IN NEW SOUTH WALES 


In this paper, we highlight past and present 
evidence that the dugong’s range on the east coast of 
Australia extends into NSW waters, including 
estuaries, when environmental conditions are suitable. 
Given their conservation status under both international 
conventions and national acts, we suggest that 
occasional visitation warrants adherence to the legal 
obligation of considering dugongs and their preferred 
habitats in future impact assessments. 


EARLY RECORDS TO RECENT SIGHTINGS 


Dugong bones have been found associated 
with edge-ground hatchet heads in Aboriginal middens 
near Sydney, indicating that at least small numbers of 
dugongs have utilized NSW waters for many centuries 
(Etheridge et al. 1896). In 1799 Flinders described the 
catching of dugongs by Aborigines in Moreton Bay, 
southeast Queensland (Mackaness 1979). Aborigines 
in NSW also caught dugongs in more recent times, 


guava 


146° 


Ae 


joones: al 


with bones having been found in middens as far south 
as Botany Bay in the late 18" Century (Troughton 
1928). 

There are currently two sources of dugong 
sightings in NSW: the Atlas of NSW Wildlife and 
records of by-catch from shark meshing supervised 
by NSW Fisheries. The Atlas of NSW Wildlife yields 
83 reports of live, stranded and dead animals for the 
period 1788 to 2003 (Anon. 2003b; Fig. 1). 

A significant portion of these reports (63) 
occurred in late 1992 and throughout 1993. This influx 
of animals occurred after the loss of 1,000 km? of 
seagrass from Hervey Bay in southeast Queensland 
following floods (Preen and Marsh 1995). Two 
dugongs were caught in NSW shark meshing during 
this time (Swansea in November 1992 and January 
1993). Three earlier captures were also made in shark 
nets (Bronte in July 1951, Bondi in July 1951, 
Queenscliff in April 1971) (Krogh and Reid 1996). 

Only two records of dead and stranded 
individuals have been reported to the NSW National 


148° 150° 152° 


‘isang 


Figure 1. Past records of dugongs on the NSW coast from 1788 to 2003 (open circles; Anon. 2003b) and 
dugong sightings in central NSW estuaries during summer 2002/03 (filled circles). 


212 


Proc. Linn. Soc. N.S.W., 125, 2004 


S. ALLEN, H. MARSH AND A. HODGSON 


32°11.0° 


152°30.2° 


Late Oct. 
2002 


Wallis 
Port 
Stephens 

| Lake 24" Jan. 
Port 1* Feb. 
Brisbane 3 Feb. 


32°42.8° 


~ 152°06.7” 
33°20.5° 
150°29.8” 


32°41.8° 


152°03.27 


33°30.1° 


152°20.3° | beach 


Peete Cee | ae 


Kayak tour operator reports dugong/s over 
seagrass beds within Wallis Lake 
Dolphin watch operators report two adult 


dugongs near Manton Bank 


Recreational 
travelling seaward out Swansea Channel 
Dolphin watch operator report dugong/s in 
upper estuary west of Soldiers Point 


Resident reports dugong/s off Orange Grove 


S. Smith, 


pets. comm. 


D. Aldritch, 


pets. comm. 


fishers report cow-calf pair | B. Roche, 


pers. comm. 


D. Aldritch, 


pets. comm. 


Anon. 2003b 


Table 1. Dugong sightings in central NSW estuaries in the austral summer of 2002/2003. 


Parks and Wildlife Service (NPWS) in the last decade, 
with no live sightings occurring until late 2002/03. 
Between late October 2002 and early February 2003, 
five separate sightings of individuals and pairs within 
(or swimming out of) central coastal estuaries were 
reported to NPWS and/or the authors (Table 1; Fig. 
1). These occurred along ac. 200km stretch of coastline 
and we do not know if these sightings include repeat 
sightings of the same individual(s). 


SEAGRASS DISTRIBUTION AND WATER 
TEMPERATURES 


All the estuaries in which dugongs were 
sighted are known to support seagrass meadows (Table 
2). Dugongs have been recorded eating the seagrasses 
listed in Table 2, with the exception of Ruppia spp. 
(Anderson 1986, Marsh et al. 1982, Lanyon et al. 
1989). Species of the genus Halophila are preferred. 
The distribution of dugongs has been reported as being 
constrained to water temperatures >~18°C (Anderson 
1986, 1994; Marsh et al. 1994; Preen et al. 1997). 
However, the water temperatures at the sites in Table 
2 were above this thermal threshold in summer 2002/ 
03. 


DISCUSSION 


The low abundance of dugongs in NSW 
waters may be the result of a number of factors 


Proc. Linn. Soc. N.S.W., 125, 2004 


including limited availability of seagrass in the region, 
relatively low water temperatures during winter months 
and in open coastal waters between estuary and bay 
habitats, and/or human pressures. The entire NSW 
coast supports only 155 km? of seagrass (West et al. 
1989), the major portion of which would be Posidonia 
australis and species of the Zosteraceae family, which 
are not favoured by dugongs. In relative terms, the 
amount of seagrass in NSW is much less than the total 
area of seagrass in Moreton Bay alone (250 km?: Abal 
et al. 1998) and would contain correspondingly small 
cover of Halophila spp. Troughton (1928) interpreted 
historical records as suggesting that dugongs may have 
occurred in greater numbers in NSW prior to European 
settlement. It has also been suggested (MacMillan 
1955) that dugong populations on the tropical east coast 
were again beginning to expand into the northern rivers 
region of NSW. Any expansion of the dugong’s range 
into NSW waters further south than this region may 
have been inhibited by the loss of seagrass beds in 
areas such as Port Macquarie and Botany Bay to 
anthropogenic influences (Pointer and Peterkin 1996). 

The dugong observations in 2002/03 (Table 
1) were in areas of NSW which have some of the largest 
seagrass beds, at least two of which include Halophila 
species — part of the preferred diet of dugongs (Marsh 
et al. 1982; Table 2). The increasing evidence that 
individual dugongs embark on movements over many 
hundreds of kilometres within tropical waters (N. Gales 
pers. comm; Marsh and Lawler 2001, 2002; Marsh 


ANS) 


DUGONGS IN NEW SOUTH WALES 


Estuary (latitude) Seagrass species and approximate area coverage 


Zosteraceae, Posidonia australis, Ruppia and Halophila 


Wallis Lake 
(~32.2S) spp. ~30.785km” 
Port Stephens 
(~32.79S) Halophila spp. ~7.453km? 
Lake Macquarie 
(~33.1S) spp. ~13.391km? 
Brisbane Water 


(~33.4) Halophila spp. ~5.490km? 


Zosteraceae, Posidonia australis and 


Zosteraceae, Posidonia australis, Ruppia and Halophila 


Zosteraceae, Posidonia australis and 


Water temp. (°C) 

October mean: 18.9 
October 2002: 21.0 
January mean: 24.1 


February mean: 24.6 


January mean: 21.6 
\ 


February mean: 22.1 


Table 2. Extent of seagrass meadows and water temperatures at sighting ocations. Sources for seagrass 
coverage and water temperature data: West et al. (1985) and Anon. (2003a) respectively. Water 
temperatures are means from 1987-2002, unless otherwise stated. 


and Rathbun 1990; Marsh et al. 2002) suggests it is 
possible that dugongs explore and utilize these southern 
seagrass beds. Warm water temperatures during the 
summer months of 2002/03 may have encouraged this 
behaviour. 

Although only five dugongs have been 
reported drowned in shark nets in NSW over the last 
c. 50 years (Krogh and Reid 1996), such deaths are 
not inconsequential since few dugongs are commonly 
found south of Moreton Bay. Two of these mortalities 
coincided with a seagrass dieback event (Preen and 
Marsh 1995) and further impact on Queensland 
seagrass beds or increase in water temperature in NSW 
may see an increase in shark net capture of dugongs 
off NSW beaches. Such events will highlight negative 
effects on populations of non-target species, and the 
efficacy of shark control programs for bather protection 
in NSW and Queensland will again be called into 
question (Anon. 2002). 

The dugong is classified as ‘vulnerable’ at a 
global scale on the IUCN Red List of Threatened 
Species. As the only extant species in the family 
Dugongidae, the extinction of the dugong will result 
in biodiversity loss at the family and generic levels as 
well as at the species level. In the light of 
inconsistencies evident in risk assessments for inshore 
development in NSW, we re-iterate that dugongs 
should be considered occasional visitors to NSW 
coastal waters. Their limited numbers warrant the 
dugongs’ consideration in future impact assessments 
for estuarine and inshore marine developments. The 
estuarine nature of recent sightings suggests that 


214 


explorative ranging by dugongs is not necessarily 
limited to strictly marine environments, rather to areas 
where seagrass beds occur. This also adds weight to 
the importance of assessing potential impacts on 
seagrass habitats. 


ACKNOWLEDGMENTS 


We gratefully acknowledge all those that provided 
prompt and unambiguous reports of recent sightings. We 
would also like to thank Mick Murphy of Hunter Coast Area 
NSW National Parks and Wildlife Service for providing 
access to the relevant wildlife database and Jeanine Almany 
for information on seagrass distribution and water 
temperatures in NSW. SA was supported by an ARC SPIRT 
grant, HM and AH by funding from the CRC Reef Research 
Centre. This manuscript was greatly improved by comments 
from Robert Williams, John Merrick and two anonymous 
reviewers. 


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216 Proc. Linn. Soc. N.S.W., 125, 2004 


Captures, Capture Mortality, Age and Sex Ratios of Platypuses, 
Ornithorhynchus anatinus, During Studies Over 30 Years in the 
Upper Shoalhaven River in New South Wales 


T.R. GRANT 


School of Biological, Earth and Environmental Sciences, University of NSW, NSW 2052 
Email t.grant@unsw.edu.au 


Grant, T.R. (2004). Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus 
anatinus, during studies over 30 years in the upper Shoalhaven River in New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 217-226. 


Data collected during a number of studies over a period of 30 years in the upper Shoalhaven River, New 
South Wales, are presented. A total of 700 individual platypuses (Ornithorhynchus anatinus) were captured 
during the studies, consisting of 137 juvenile females, 94 juvenile males, 292 adult females and 177 adult 
males. The overall sex ratios of both adults (1.65:1) and juveniles (1.46:1) were significantly biased towards 
females. Females were found to live up to 21 years. Very few recaptures of juvenile males made estimates 
of longevity equivocal, but three individuals were at least 7 years old when last captured. Capture and 
handling mortality during the various studies was low (0.86%). Sixty-two percent of platypuses marked in 
the study area were never recaptured, fewer adult males were recaptured than females (36% and 51% 
respectively) and recaptures of juveniles were much lower than for adults (32% females and 14% males). 
Recapture data suggest considerable mobility by adults and dispersal by juvenile platypuses along the upper 


Shoalhaven River and its tributaries. 


Manuscript received 2 September 2003, accepted for publication 8 January 2004. 


Keywords: Age, Capture, Marking, Mortality, Ornithorhynchus anatinus, Platypus , Sex Ratio 


INTRODUCTION 


Over the past 30 years a number of research 
projects has been carried out in a study area in the 
upper Shoalhaven River in New South Wales. 
Individual projects have included investigation of 
temperature physiology (Grant and Dawson 1978a,b; 
Grant 1983; Hulbert and Grant 1983a,b), diet (Faragher 
et al. 1979), movements and home ranges (Grant 1983, 
1992), haematology and pathology (Munday et al. 
1998; Whittington and Grant 1983, 1984, 1995; 
Whittington et al. 2002), lactation and milk 
composition (Grant et al. 1983, Gibson et al. 1988; 
Grant and Griffiths, 1992) and population genetics 
(Gemmell 1994; Akiyama 1999). During these studies, 
long-term data have been collected on recaptures, 
capture mortality, longevity and sex ratios of 
platypuses (Ornithorhynchus anatinus). 

During the later studies from 1987, the 
investigation and use of Passive Integrated 
Transponder (PIT) tags or “microchips” was begun, 
probably the first time that this marking method was 
used on a wild mammalian species in Australia (Grant 
and Whittington 1991). The long-term success of this 


method in the mark and recapture studies of the 
platypus is reported below. 

Collins (1973) tabulated the ages of eight 
platypuses kept in captivity in a variety of locations, 
including the Bronx and Budapest Zoos. These ranged 
from four to 17 years, although anecdotal information 
from zoos and sanctuaries indicates that the species 
may survive in captivity for up to 21 years (Whittington 
1991). Concerning the longevity of platypuses in the 
wild, the naturalist Harry Burrell (1927) wrote that “the 
length of life of the platypus is not known. It is my 
intention to ring-mark some fully furred young as 
opportunity offers, and it may be that we shall gain 
some information on this point at a later date, if these 
marked individuals are captured”. Burrell did not later 
report the ages of platypuses he may have “ring- 
marked”. However, Grant and Griffiths (1992) 
reported the ages of platypuses marked in the upper 
Shoalhaven River as being between as much as 4 years 
for males and 8 for females. Since that report, a further 
12 years of research has resulted in the data presented 
in this paper on the ages and sex ratios in this 
population of platypuses. Mortality of capture and 
handling of platypuses in previous studies has not 


CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 


previously been discussed in the literature and this 
aspect of the studies is presented in the current paper. 


METHODS 


Between June 1973 and January 2004, 
platypuses were captured in 16 pools in the upper 
Shoalhaven River in New South Wales using the 
unweighted “gill” net methods outlined in Grant and 
Carrick (1974). Until 1987, individuals were marked 
using stainless steel leg bands (Grant and Carrick 
(1974) but these were phased out after trials on the 
use of Passive Integrated Transponder (PIT) tags 
proved to be successful (Life Chip tags; Destron 
Fearing Corporation Scanner; Grant and Whittington 
1991). 

Sex was determined using the presence or 
absence of the adult spur or the morphology of juvenile 
spurs. Absolute ages were determined from individuals 
initially captured as juveniles and minimum ages for 
adults were estimated using the time of loss of the 
female spur, the morphological changes in the males 
spur (Temple-Smith 1973; Grant 1995), and 
subsequent recaptures. Females possessing a spur were 
categorised as being in their first year of life (0 years 
of age) and males could be assigned to their first or 
second year of life (0 or 1 year of age). As the females 
in this area lose the spur between October and 
December in their first year after emergence from the 
nesting burrows, any female lacking a spur at first 
capture was considered to be = 1 year of age (i.e. in 
their second year of life). It should be noted that two 
female juvenile platypuses bred in captivity at Taronga 
Zoo in the 2002/2003 breeding season apparently lost 
their spurs within only 4 months after emergence from 
the nesting burrow (Adam Battaglia, Taronga Zoo, 
pers. comm.). Males with adult spur morphology were 
considered to be at least in their third year of life, or = 
2 years old. Subsequent recaptures permitted minimum 
ages to be assigned to individuals, beginning with a 
minimum age at first capture of one year for adult 


Juvenile 
Males 94 
Females 137 
Total Agi. 
Sex Ratio (F:M) 1.46:1 
Chi? 8.00 
p ** < 0.005 


females and two years for adult males (Temple-Smith 
1973; Grant and Griffiths 1992; Grant 1995). 

Recaptures were recorded for animals in all 
of the 16 pools of the 12.5 km section of the upper 
Shoalhaven River and 3.9 km of an adjacent creek. 
However, by 1987 a number of these pools had filled 
with sand and were no longer netted. By 1993, the 
previously largest and deepest pool (1 km long x 2-5 
m deep) was completely filled with sand and was no 
longer sampled, although many of the platypuses 
originally captured in this pool were captured in the 
pools downstream. From 1988, when PIT tagging had 
become the predominant method of marking, sampling 
was mainly restricted to three pools in the Shoalhaven 
River itself and one in the adjacent creek. In most years 
after that time these pools were sampled late in the 
year (mainly December) when lactating animals were 
most likely to be caught and at the end of summer 
(mainly February or March) when juveniles had newly 
emerged (Grant and Griffiths 1992). 


RESULTS 


Sex ratios 

During the studies from June 1973 to January 
2004, 700 individual platypuses were captured. Table 
1 shows the numbers in each of four age/sex classes. 
All sex ratios were significantly biased towards 
females. The ratio of females to males was 1.58 
females:1 male (Chi? = 34.87; p < 0.001) for all 
animals, 1.46:1 for juveniles (Chi* = 8.00; p < 0.01) 
and 1.65:1 for adults (Chi? = 27.38; p < 0.001). 


Age 

Only 45 individuals (41 females and 5 males), 
first marked as juveniles, were subsequently 
recaptured. Figure 1 shows the distribution of ages of 
these individuals at their latest recapture. Two juvenile 
females were recaptured regularly over periods of 13 
and 16 years but were not captured again in 5-6 
subsequent years. These animals were assumed to have 


Adult Total 
177 2/4 
292 429 
469 700 

1.65:1 1.58:1 

27.38 34.87 

** < ().001 ** < 0.001 


Table 1. Numbers and sex ratios of platypuses captured in the upper Shoalhaven River study area. 


** significant at < 0.01 level. 


218 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


- Year Juvenile Juvenile Year Adult Adult 
(Actual) Female Male (Minimum) — Female Male 
0 96 89 0 - - 
1 2A 4 1 181 28 
2 3 1 2 33 iis 
3 3 - 3 18 11 
4 3 - 4 18 9 
5 2 - 5 10 6 
>5 9 - >5 32 6 
37 94 Total 292 177 


Total 1 


Table 2. Numbers of platypuses allocated to actual and estimated minimum age categories in the upper 
Shoalhaven River study area. Actual = ages of animals initially captured as newly-emerged juveniles; 
Minimum = minimum ages; calculated from years between initial and last capture of individuals first 


captured as adults. 


died. However, one was again recaptured and lactating 
at the end of the study, when her age was 21 years. As 
indicated in the Methods section, females without spurs 
are at least in their second year of life (>/=1 year old) 
and it is possible to attribute males to either their first 
second or third year of life (0, 1 or 2 years old) based 
on spur morphology changes. Ages of >/= 2 years 
could be attributed to 111 female adults caught and 
subsequently recaptured. Similarly 32 adult males were 
attributed to the >/= 3 year age category. The 
distribution of these minimum ages are shown in Table 
2 and Fig. 1. 

While most platypuses caught in the study 
could only be attributed to the >/= 1-2 year age 
category, 9 females first captured as juveniles survived 


between 5-21 years and 32 of those initially captured 
as adults survived 5-15 years. One juvenile male was 
subsequently recaptured at 2 years of age but 32 males, 
first captured as adults, survived to minimum ages of 
3-7 years. 


Recaptures 

The numbers of juvenile, adult male and adult 
female platypuses recaptured at least once in the latter 
12 years (when most animals were marked with PIT 
tags) were not significantly different from those of the 
first 18 years of the study (when the majority were 
marked with leg-bands)(Table 3). Table 4 presents 
combined recapture data for both leg-banded and PIT 
tagged animals for the whole study period. 


Adult 


Juvenile Juvenile Adult 

Female Male Female Male 
Leg-banded 
Total recaptures 33 105 53 
€ 1 recapture) c 
Total captures oF 69 214 134 
% recapture 34.0% 13.0% 49.1% 39.5% 
PIT tagged 
Total recaptures 24 47 ils) 
(= | recapture) 
Total captures 54 SD 86 41 
% recapture 42.9% 6.3% 54.7% 36.6 
Chi? 1.60 1.04 0.77 0.12 
p NS <0.20 NS <0.31 NS <0.38 NS <0.73 


Table 3. Comparison between total number of recaptures (>/= 1) for leg-banded and PIT tagged platypuses 
in the upper Shoalhaven River study area. Animals marked with both bands and PIT tags between 1987 
and 1991 are included in both sets of data. NS = not significant. 


Proc. Linn. Soc. N.S.W., 125, 2004 


ANY) 


CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 


Juvenile Females 


200 
180 
160 
140 
120 
100 


Frequency 


12068 Fe ws HORS FF Sey Oey 10611 21S a 6 7) 1G 9 2O eat 


Age (years) 


Adult Females 


Frequency 


ba Di Sic bie SO Sok ONO ntl 21S. eel Sees 
Age (years) 


Figure 1. Actual and minimum age frequencies of platypuses in the study. Actual ages are shown for 
those animals initially caught as juveniles (Juvenile Females and Juvenile Males). Estimated minimum 
ages shown are for animals caught first as adults (Adult Females and Adult Males). 


CONTINUED ON FACING PAGE. 


Considerable numbers of both male and female adults 
and juveniles were captured only once. 

Table 4 shows that total recaptures (>/= 1 
times) and recaptures in the categories of 1, 2-5 and 
>5 times were lower for adult males than for females 
and that total recaptures of juvenile males was less 


220 


than half (14%) that of juvenile females (32%). The 
majority of recaptured males were caught within the 
first months after initially emerging from the nesting 
burrows (0 years of age), while recaptures of juvenile 
females were spread across 0-21 years after emergence 
(Figure 1 and Table 4). 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


Juvenile males 


160 5 

140 
> 120 4 
S 100 - 
= 
3 80 - 
Se 60 - 

40 5 

20 - 

(0) ee T T T T T T T T T | T | 
eee oA enon 7. 8 9 0 Tl 12-13" 1415 16 
Age (year's) 
Adult Males 

a) 
oO 
| 
cB) 
=) 
i=_ 
x 
cs 


ves Aaa LO 7 8 OO L 2 1314. 1S 16 
Age (years) 


Capture mortality 

Of the 700 platypuses captured during the 
various research projects six died as a result of capture 
and handling (0.86%). Two drowned as a result of 
netting, two died suddenly within a few hours of 
capture (sudden death; animals appeared healthy and 
no obvious cause of death was identified from post- 
mortem examination by veterinarians), one succumbed 
to anaesthesia and one became caught in submerged 
vegetation by its transmitter attachment and drowned 
during telemetry work. Details are in Table 5. 


Proc. Linn. Soc. N.S.W., 125, 2004 


DISCUSSION 


Sex Ratios 

As reported by Grant and Griffiths (1992) for 
the first 18 years of the study, the sex ratios for both 
adults and juveniles were significantly biased towards 
females. Table 6 compares the sex ratios for captured 
adult platypuses in three other areas (Grant 
unpublished). Although based on much smaller sample 


221 


CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 


Juvenile Juvenile Adult Adult 

Female Male Female Male 
1 recapture 17 10 66 34 
2-5 recaptures 19 3 58 20 
>5 recaptures 7 0 18 6 
Total recaptures 43 15 142 60 
(> 1 recaptures) » 
% recapture 32% 14% 51% 36% 
Total captures’ 135 94 278 165 


Table 4. Total recaptures of male and female juvenile and adults platypuses in the upper Shoalhaven 
River study area. ‘Mortalities and some animals which would have been unlikely to have been 
recaptured after the netting of some pools was discontinued are not included in this total captures 


figure. 


sizes, none of these were significantly different from 
parity. Grant and Griffiths (1992) also reported no 
significant difference between males and females in 
total numbers of platypuses (juvenile and adults not 
specified) captured in various rivers of New South 
Wales and the Australian Capital Territory (Table 6). 
Like the situation in the upper Shoalhaven River, 
during the earlier years of a study (1986-90) in the 
Duckmaloi River on the central tablelands of New 
South Wales, a bias towards females in both adult and 
juvenile platypuses was found. However, in the later 
years (1991-2000) more adult males than females were 
recorded (David Goldney, University of Sydney, 
Orange, pers. comm.). 

The recapture data discussed below seems to 
indicate that female platypuses in the upper Shoalhaven 
River survived for significantly longer periods than 
males. Over time, this longer survival of females would 
presumably have led to a sex ratio weighted towards 
females in the population. However, this explanation 
does not account for the disparity between numbers 
of male and female juveniles in this population. 

Most juvenile male platypuses disappeared 


from the upper Shoalhaven River population in their 
first year (86%; Table 4). However, 13% of juvenile 
females were recaptured in the area up to age one year, 
19% were recaptured later than two years after 
emerging from the nesting burrows and two even 
remained in the area up to age 13 and 21 years 
respectively. Twelve juvenile females (9%) bred in the 
area, eight of these over a number of breeding seasons. 
Differential dispersal may contribute to the difference 
in the adult sex ratio, but again this does not explain 
the significant bias to females in the numbers of 
juveniles captured at the time they were becoming 
independent (late January-late March), unless most 
male juveniles dispersed immediately after 
independence, with females dispersing later. 
Unfortunately the data from this study do not permit 
this hypothesis to be rigorously tested, as most 
sampling only occurred early and late each year. 

The possibility also exists that the uneven sex 
ratios are determined by differential fertilisation of 
eggs, development of embryos or pre-emergence 
survival of young but no explanation arises from the 
data collected in this study concerning the significant 


Cause of Death Juvenile Juvenile Male Adult Female Adult - Total 
Female Male 
Drowned in net 1 0 1 0 2 
Sudden death 0 0 0 2 2; 
Anaesthesia (ether) 1 0 0 0 A 
Snagged transmitter 0 0 1 0 1 
Total Dy 0 2 2 6 


Table 5. Capture and handling mortalities in platypuses during work on various projects in the upper 


Shoalhaven River study area 


py L)2 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


Location Adult Female Adult Sex Ratio Chi” 
Male (F:M) Probability 
Various streams NSW/ACT’ 101 117 1.15:1 IL-7) 
p< 0.28 
Various streams NSW/ACT" 47 47 1:1 - 
Barnard River, NSW 24 22; 1.10:1 0.09 
p< 0.77 
Thredbo River, NSW 14 10 1.4:1 0.67 
p< 0.41 
Wingecarribee River, NSW 29 30 1:1.03 0.02 
ps 0.90 
Shoalhaven River, NSW 285 173. 1.85:1 27.38** 


p< 0.001 


Table 6. Comparison of sex ratios of adult platypuses in various studies in New South Wales and the 
Australian Capital Territory (ACT). Collected by ‘Temple-Smith and * Griffiths (from Grant and Griffiths 


1992); ** significant at < 0.01 level. 


bias towards females in the sex ratio of juvenile 
animals. In the Barnard River study referred to in Table 
6, where the adult sex ratio was not different from 
parity, the sex ratio for juveniles was heavily male- 
biased (14 males; 3 females) during the single breeding 
season studied. A similar result was also obtained once 
during a single breeding season (12 males to 3 females; 
1978/79) in the current study, indicating differences 
between individual years. However, in the 24 years 
during the study in which juveniles were captured, the 
numbers of females exceeded males in 87.5% of those 
years. Considerable annual differences in recorded 
annual sex ratios were also found in the Duckmaloi 
River study (David Goldney, University of Sydney, 
Orange pers. comm.). 


Age 

The minimum estimated age for male 
platypuses in this study (7 years) was considerably less 
than for females (up to 21 years), with nine adult 
females surviving for a minimum of 10 years. These 
data suggest that females live longer than males in the 
wild. However, determination of actual age, or the 
estimated minimum age, depended on recapture data 
and, as discussed, recapture of males was much lower 
(36% recaptured >/= once) than for females (51% 
recaptured >/= once). After being regularly captured 
previously, 13 and 21 year old females (first marked 
as juveniles) had not been recaptured for the last 6 and 
5 years respectively of the study. Except for one adult 
female, which had been captured in December 2002 


Proc. Linn. Soc. N.S.W., 125, 2004 


and was not caught in March 2003, all the other adult 
females >/= 10 years of age had also not been 
recaptured in the latter years of the study. These data 
appeared to indicate a life span of 10-16 years may 
represent an expected upper range of longevity for 
female platypuses in the wild. However, the final 
recapture of one female at the age of 21 years showed 
a maximum female longevity in the wild comparable 
to that in captivity. While the data for males in the 
wild appeared to show shorter life spans (up to 7 years), 
this could equally represent non-recapture of older 
males. Reports do not suggest differing longevity 
between the sexes in captivity (Collins 1973; 
Whittington 1991). 


Recaptures 

PIT tags were initially used because of the 
occurrence of notched and broken male spurs as a result 
of bands abrading the spur base. However, it was also 
suspected that the lower capture rates in males, 
particularly juveniles may have been attributable to 
band losses. Bands were normally fitted more loosely 
to males to permit the much greater radial growth of 
the hind legs in this sex. No significant differences 
between captures for banded and PIT tagged males 
(Table 3) indicated that band loss could not fully 
explain the lack of recaptures, although four female 
animals, initially marked with bands and PIT tags, were 
found to have lost their bands during the latter part of 
the study, indicating some band loss. Only one PIT 


223 


CAPTURE, MORTALITY AND SEX OF PLATYPUSES IN THE SHOALHAVEN RIVER 


Home Maximum distance Source Location 
range(km) (km) : 
0.2-2.0 5.6 (24 hr max.=4.0) Grant, 1983 Shoalhaven River 
0.4-2.3 23 Grant, 1983, Grant et al. 1992 Thredbo River 
0.3-2.3 2.3 Serena, 1994 Badger Creek 
2.9-7.0 15.0 Gardner and Serena, 1995 Watts River and Badger Creek 
0.4-2.6 2.6 Gust and Handasyde, 1995 Goulburn River 
Adult: 24 hr max. = 10.4 Serena et al. 1998 Yarra River, Mullum Mullum Creek, 
2.9-7.3 (male) Diamond Creek 
Juvenile: 24 hr max. = 4.0 
1.4-1.7 (female) 
40 in 18 months Australian Platypus Yarra catchment 
Guvenile) Conservancy 1999 Andersons to Steels Ck 
48 in 7 months Australian Platypus Wimmera River 


(young male) 


Conservancy 2001 


Table 7. Home ranges and maximum distances moved by platypuses in various studies, including the 


Shoalhaven River (bold). 


tag failure or loss was confirmed in 220 tags applied 
to animals during the studies. The lack of spur damage, 
some evidence of band loss and no significant 
differences being found between recaptures of leg- 
banded and PIT tagged animals confirmed PIT tagging 
as the preferred method of marking platypuses (Grant 
and Whittington 1991). 

Large numbers of both adult male (64%) and 
female (49%) platypuses were not recaptured in the 
study area after being marked either with leg bands, 
PIT tags or both. This observations suggests one, or a 
combination of the following: 


Loss of marks. Double marking indicated that some 
band loss did occur during the study but there was 
little indication that PIT tags were lost or failed. 


Mortality. Little is known about the causes and 
incidence of mortality in platypuses. 

Foxes (or dogs) will take platypuses on land, 
from shallow riffle areas and by digging into burrows 
(Serena 1994; Grant 1993; Anon. 2002). Large eagles 
may also be possible natural predators of platypuses 
(Rakick et al. 2001). The remains of a platypus near a 
burrow excavated by a fox or dog, an isolated skull in 
a pool and part of a skull in a pile of other mammalian 
bones (mainly cattle and sheep) were the only observed 
evidence of mortality found during the studies in the 
upper Shoalhaven River. 

While 50% of 131 individuals tested positive 
to leptospirosis antibodies (Leptospira interogans 
serovar hardjo)(Munday et al. 1998), no clinical 
symptoms of the disease were observed and nothing 
is known of any disease organism, resulting in 
significant mortality in this population. The Mucor 
fungus, which has caused mortality in Tasmanian 


224 


populations, has not so far been detected in mainland 
populations of platypuses, including those in the upper 
Shoalhaven River (Whittington et al. 2002). 


Mobility. Diurnal and longer-term mobility over 
distances of up to 5.6 kilometres have been previously 
reported in individuals in the upper Shoalhaven River 
(Grant 1983, 1992, 1995; unpublished) and in other 
studies. These data are summarised in Table 7. 

After the marking of 700 individual 
platypuses (including significant numbers of new 
juveniles) during the 30 years, it was expected that the 
majority of the population would eventually be marked 
and that unmarked dispersing juveniles or adults might 
still enter the area but would be in fairly small numbers. 
In fact, considerable numbers of new adult animals 
were captured throughout the study. Some individuals 
were captured as many as 20 times over periods of up 
to 21 years and yet the times between recaptures of 
these individuals was often quite variable. For example, 
despite the pools being regularly netted during the 
study, two adult females were only subsequently 
recaptured nine and 10 years respectively after their 
initial capture in those pools. Even for females 
identified as breeding in particular pools during 
different breeding seasons, periods of time between 
some recaptures of these animals ranged from 1 to 10 
years. 

These latter observations suggest that a great 
deal of mobility probably characterises the platypus 
populations in the upper Shoalhaven River, although 
the effects of mortality and/or dispersal cannot be ruled 
out as reasons for the influx of new animals and the 
lack of recapture of a significant proportion of the 
platypuses in the upper Shoalhaven River study area. 
All of these possibilities need further study. 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


While there was capture and handling 
mortality during the studies in the upper Shoalhaven 
River population, this was quite low (< 1%) due to the 
utilisation of methods developed through considerable 
experience by the author and other researchers over 
the past three decades. 


ACKNOWLEDGMENTS 


The many friends and colleagues who assisted with 
the field work over the years are too numerous to thank 
individually but all are gratefully acknowledged. The last 
10 years of the study would not have been possible without 
the help and support of Colin, Kate, Sue and Tom Heath 
and Paul Anink, Marie-Louise Lissone and Gina Grant. The 
late Athol MacDonald and the Izzard and Laurie families 
are acknowledged for their permission for access to the river 
and creek, and for their friendship and assistance in various 
aspects of the field work. Unfortunately Bill and Ron Izzard 
both died in 2003 and this paper is dedicated to their memory. 
Adam Battaglia and David Goldney are acknowledged for 
their personal communications. Richard Whittington and 
Joanne Connolly carried out the post-mortem examinations 
of the two animals which died suddenly after capture. Peter 
Temple-Smith, Michael Augee and an anonymous referee 
provided valuable comments on the manuscript. Some of 
the work reported was done while in receipt of funding from 
the Environment Australia (then Australian National Parks 
and Wildlife Service) and the Australian Research Council 
(then Australian Research Grants Committee). This work 
was carried out under NSW National Parks and Wildlife 
Service Scientific Investigations Licence A184, New South 
Wales Fisheries Scientific Research Permit F84/1245 and 
University of New South Wales Animal Care and Ethics 
Approvals 94/91, 97/46 and 00/45. 


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Collins, L-R. (1973). ‘Monotremes and marsupials. A 
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Proc. Linn. Soc. N.S.W., 125, 2004 


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Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. 
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Gemmell, N.J. (1994). Population and evolutionary 
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Grant, T.R. (1983). Body temperature of free-ranging 
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Grant, T.R. (1992). Captures, movements and dispersal of 
platypuses, Ornithorhynchus anatinus, in the 
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2nd edition. Sydney: University of NSW Press: 
Sydney 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist 18: 
133-135 

Grant, T.R. and Dawson, T.J. (1978a). Temperature 
regulation in the platypus, Ornithorhynchus 
anatinus, maintenance of body temperature in 
air and water. Physiological Zoology 51:1-6. 

Grant, T.R. and Dawson, T.J. (1978b). Temperature 
regulation in the platypus, Ornithorhynchus 
anatinus, production and loss of metabolic heat 
in air and water. Physiological Zoology 51: 315- 
332. 

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a free-ranging population of platypuses, 
Ornithorhynchus anatinus, in the Shoalhaven 
River, New South Wales. In. Platypus and 
Echidnas. (Ed M.L.Augee). pp. 80-89. (Royal 
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Aspects of lactation in the platypus, 
Ornithorhynchus anatinus (Monotremata), in 
waters of eastern New South Wales. Australian 
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freeze-branding and implanted transponder tags 
as a permanent marking method for platypuses, 
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Ornithorhynchidae). Australian Mammalogy 14: 
147-150. 

Gust, N., Handasyde, K. (1995). Seasonal variation in the 
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River; Victoria. Australian Journal of Zoology 
43, 193-208. 

Hulbert, A.J. and Grant, T.R. (1983a). A seasonal study of 
body condition and water turnover in a free- 
ranging population of platypuses, 
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of Zoology 31: 109-116. 

Hulbert, A.J. and Grant, T.R. (1983b). Thyroid hormone 
levels in the egg-laying monotreme, the 
platypus, Ornithorhynchus anatinus. General 
and Comparative Endocrinology 51: 401-405. 

Munday, B.L., Whittington, R.J. and Stewart, N.J. (1998). 
Disease conditions and subclinical infections of 
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Observations of a platypus foraging in the sea 
and hunting by a wedge-tailed eagle. Tasmanian 
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226 


Serena, M., Thomas, J.L., Williams, G.A., Officer, R.C.E. 
(1998). Use of stream and river habitats by the 
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fringe habitat. Australian Journal of Zoology 
46, 267-282 

Temple-Smith, P.D. (1973). Seasonal breeding biology of 
the platypus, Ornithorhynchus anatinus Shaw 
1799, with special reference to the male. PhD 
Thesis. Australian National University: 
Canberra. 

Whittington, R.J. (1991). the survival of platypuses in 
captivity. Australian Veterinary Journal 68, 32- 
39: 

Whittington, R.J. and Grant, T.R. (1983). Haematology 
and blood chemistry of free-living platypuses, 
Ornithorhynchus anatinus 
(Shaw)(Monotremata: Ornithorhynchidae). 
Australian Journal of Zoology 31: 475-482. 

Whittington, R.J. and Grant, T.R. (1984). Haematology 
and blood chemistry of the conscious platypus, 
Ornithorhynchus anatinus 
(Shaw)(Monotremata: Ornithorhynchidae). 
Australian Journal of Zoology 32: 631-635. 

Whittington, R.J. and Grant, T.R. (1995). Haematological 
changes in the platypus (Ornithorhynchus 
anatinus) following capture. J. Wildlife Diseases 
31: 386-390. 

Whittington, R.J., Connolly, J.H., Obendorf, D.L., 
Emmins, J., Grant, T.R. and Handasyde, K.A. 
(2002). Serological responses against the 
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Proc. Linn. Soc. N.S.W., 125, 2004 


Breeding in a Free-ranging Population of Platypuses, 
Ornithorhynchus anatinus, in the Upper Shoalhaven River, New 
South Wales - a 27 Year Study 


T.R. Grant!, M. GrirFitHs* AND P.D. TEMPLE-SMITH? 


' School of Biological, Earth and Environmental Sciences, University of NSW, Sydney 2052, 
t.grant@unsw.edu.au; 780 Dominion Circuit, Deakin, ACT, 2600 
3Department of Conservation and Research, Zoological Parks Board of Victoria and 
The University of Melbourne 


Grant, T.R., Griffiths, M. and Temple-Smith, P.D. (2004). Breeding in a free-ranging population of 
platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven River, New South Wales - a 27 year 
study. Proceedings of the Linnean Society of New South Wales 125, 227-234. 


A total of 150 captures of lactating platypuses (97 individuals) were made over a period of 27 years in the 
study area. The proportion of lactating females from December samples ranged from 18 to 80% (mean 
43.4+17.7%; n = 21 breeding seasons). The percentage of juveniles in samples taken at the seasons when 
young were leaving the nesting burrows varied from 0-63% (mean 34.4+17.9%; n = 22 breeding seasons). 
Only 8.8% percent of captured juvenile females went on to breed in the area; one bred in its second breeding 
season after emergence but two others did not breed until at least their 4" breeding season. Some females 
bred during at least 2-3 consecutive breeding seasons but others failed to breed in consecutive years. The 
percentages of females lactating in the months of September to April indicated a spread in the breeding 
season. Lactation in the wild was apparently shorter than reported in captivity, lasting more than 3 but less 
than 4 months. The majority of variation in breeding activity and recruitment could not be explained in 
terms of drought or observed riverine and riparian changes during the study. 


Manuscript received 2 September 2003, accepted for publication 8 January 2004. 


KEYWORDS: Breeding, Drought, Lactation, New South Wales, Ornithorhynchus anatinus, Platypus, 


Recruitment, Sedimentation 


INTRODUCTION 


Platypuses (Ornithorhynchus anatinus) mate 
in late winter or early spring. Eggs are laid and the 
developing young are nourished on milk in the nesting 
burrows for several months, after which juveniles leave 
these burrows, become independent and most disperse 
from natal sites. There is a north-south cline in the 
timing of the breeding season, which begins earliest 
in north Queensland and latest in Tasmania (Temple- 
Smith and Grant 2001). The current study was carried 
out near the centre of this cline, on the southern 
tablelands of New South Wales in the upper 
Shoalhaven River. It began with the investigation of 
the nature of lactation and the composition of the milk 
of the platypus (Griffiths et al. 1973; Grant et al. 1983; 
Messer et al. 1983; Parodi and Griffiths 1983; Griffiths 
et al. 1984; Griffiths et al. 1985; Gibson et al. 1988; 
Teahan et al. 1991; Grant and Griffiths 1992; Joseph 
and Griffiths 1992). However, in the mid-1980s there 
was considerable change to the habitat of the platypus 


within the study area, with sand slugs encroaching into 
many of the pools and considerable bank erosion 
occurring as a result of poor past and present riparian 
and catchment land management practices. On 
completion of the initial studies early in the 1990s, the 
investigation continued by sampling in December, 
when females captured would be most likely to be 
lactating, and in February or March when juveniles 
had left the nesting burrows but had not yet dispersed 
(see also Grant, 2004 this volume). 

While the study has permitted general aspects 
of lactation to be further considered since the work of 
Grant and Griffiths (1992), it has also investigated the 
effects of stream degradation and drought on the 
platypus population in the upper Shoalhaven River. 
With regard to this latter aspect of the study, the 
hypothesis being tested was that successful breeding, 
as indicated by females breeding and young being 
recruited to the population each year, would be 
adversely affected by stream degradation and/or by 
droughts. 


BREEDING IN FREE-RANGING PLATYPUSES 


MATERIALS AND METHODS 


Study Site 

The study area consisted of a series of 16 
pools in agricultural land, separated by riffle areas 
along 12.5 kilometres of the upper Shoalhaven River 
and 3.9 kilometres of an adjacent tributary stream, near 
Braidwood on the southern tablelands of New South 
Wales. A narrow discontinuous strip of riparian 
vegetation, consisting of both introduced and 
indigenous species of trees and shrubs, interrupted by 
numerous gaps, which were normally eroded as a result 
of access by sheep and/or cattle from the surrounding 
pasture land to the river. During the period of the study 
(late 1977 to early 2004), some of these pools suffered 
in-filling by sand slugs. For some pools, the effects on 
the habitat from sand in-filling was so severe that they 
were deleted from the sampling program. During the 
study period, three significant droughts occurred. 


Sampling Periods 

Two to four pools representative of the area 
(core area) were sampled during December, then again 
in February and/or March of 21 and 22 breeding 
seasons respectively over the 27 years of the study. 
Other pools within and outside these core area pools 
were sampled intermittently at various times during 
research in associated projects (Grant, 2004 this 
volume). 


Capture, Marking and Possessing 

Animals were captured using the unweighted 
“gill” net methods outlined in Grant and Carrick 
(1974). Until 1987, individuals were marked using 
stainless steel leg bands (Grant and Carrick 1974) but 
these were phased out after trials on the use of Passive 
Integrated Transponder (PIT) tags proved to be 
successful (Life Chip tags; Destron Fearing 
Corporation Scanner; Grant and Whittington 1991; 
Grant 2004; this volume). After removal from the nets, 
animals were weighed, measured and age and sex were 
determined (Temple-Smith 1973; Grant 2003 this 
volume). Females were injected intramuscularly with 
0.1-0.2 mL of synthetic oxytocin (1-2 International 
Units; Syntocinon, Novartis) to induce milk “let down” 
(Griffiths et al. 1972, 1973, 1984; Grant and Griffiths 


1992). In females that were lactating, milk could be 
expressed from the mammary gland, using gentle 
pressure along the flanks towards the areolae, 5 minutes 
after injection. 


Data Collection 

The percentage of lactating females captured 
in each December sample and the percentage of 
juveniles caught in relation to the total numbers of 
animals captured at each sampling in February and/or 
March were calculated. These provided indices of 
breeding and recruitment success for each breeding 
season. The timing and duration of lactation were 
determined from these data and from the capture and 
recapture of females in other pools of the study area. 


RESULTS 


Timing and duration of lactation 

During the 27 years of the study, captures of 
150 lactating platypuses were made. A total of 97 
individuals were lactating at least once during the study 
(Table 1). Only a single individual was found lactating 
in late September, with the highest proportions of 
lactating animals being captured in December and 
January. Sequential recaptures of three individuals 
within the same breeding season showed lactation in 
the field lasted at least 70-98 days (2.3-3.3 months) 
(Table 2). Other sequential data showed that 97% (30 
from 31) of females found lactating in December or 
January, had ceased lactation when recaptured in 
March. Of five individuals lactating in December, three 
were no longer lactating when recaptured in February 
(Table 2). 


Breeding ages of juvenile platypuses 

Of 137 female platypuses captured as 
juveniles, only 12 were later recaptured as breeding 
females in the study area (8.8%, Table 3). One of these 
individuals was lactating in its second breeding season 
after emergence from the nesting burrow, but three 
others did not breed until at least their 3" or 4" breeding 
season. The individual (FJ222) which bred in the 
second breeding season (1983/84) failed to breed the 
following year (1984/85). This animal was not captured 


Sept. Oct. Nov. Dec. Jan Feb. Mar. Apr 
Total 26 5 25 256 60 59 179 5 
Lact. 1 1 7 106 24 10 1 0 
% 3.8 20.0 28.0 41.3 40.0 17.6 0.6 0 


Table 1. Numbers and percentages of individual female platypuses lactating in all samples in the upper 


Shoalhaven River study area 


228 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 


Animal December January 
FAO15 
FAO019 
FA046 
FA126 
FA133 
FA158 
FA158 
FA161 
FA161 
FA185 
FA185 
FA185 
FA209 
FA212 
FA214 
FA276 
FA335 
FA368 
FA368 v 
FA370 
FA391 
“-FA462 
FA514 
FA535 
FA530 v 
FA547 
FJ157 

FJ222 

FJ248 - 
FJ248 v 
FJ272 
FJ272 
FJ272 
FJ436 
FJ469 


SONS NON NN NN NS NGAI NONI 


SNe AN 


iN I 


Noe! 


February 


IND ON Nee 
a 


Lactation 
duration 


March April 


mM 


a >72 days 


xX >98 days 


mM! 


- >70 days 


PS PS PS PS PS Pd OPS PS PS PS PS PS PS Pd Pd OP PS 


PS PS Pd PS PS PS PS PS 


Table 2. Recaptures of lactating platypuses within given breeding seasons V Lactating; X Not 


lactating; - not recaptured. 


in the breeding season of the next year (1985/86) but 
was lactating again in the subsequent breeding season. 


Breeding success and recruitment 

Twenty-eight females were captured in 
successive breeding seasons. While some females were 
captured lactating in up to three consecutive breeding 
seasons, many failed to breed in consecutive seasons, 
with 39% not lactating in a season immediately 
following one in which they did breed. (Table 4). 

The mean percentage of lactating (breeding) 
animals in December of 21 breeding seasons in the 
core section of the study area was 43.4+17.7% but the 
numbers and proportions fluctuated considerably 
between breeding seasons from 80% down to 18% of 
the numbers of females captured (Figure 1). 


Proc. Linn. Soc. N.S.W., 125, 2004 


The data showed a general relationship 
between the numbers of juvenile platypuses captured 
in February and/or March in the core area of the study 
site and the total numbers of lactating animals captured 
in each of the breeding seasons sampled (Figure 1). In 
some years recruitment of juveniles was low after 
reasonable numbers of lactating females had been 
captured in December and in other years higher than 
expected recruitment levels were observed in February/ 
March after relatively low numbers of lactating females 
had been captured in the previous December. However, 
in general higher percentages of juveniles were 
captured at the end of breeding seasons when the 
percentage of lactating females sampled was also high 
(Figure 1). 


229 


BREEDING IN FREE-RANGING PLATYPUSES 


Animal 
FJ157 


FJ217 
FJ222+ 


FJ230 
FJ235# 
FJ248 
FJ272# 
FJ273 
FJ409 
FJ436 
FJ469# 


FJ496 


79/ 80/ 81/ 82/ 83/ 84/ 85/ 86/ 87/ 88/ 89/ 90/ 91/ 92/ 93/ 94/ 95/ 96/ 97/ 98/ 99/ O00/ O1/ 02/ 03/ 


80 


* 


81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 «~=97:~ «©=298—(99) 600) h6lOl) 602) (603) hl 


i yi Vi e > AX : a , a = 2 : = = 5 : z E z 2 : E 
Xi 

#E Z af v = e = = x E, 4 . : 4 = a S = = : 2 F 3 
a '¢ v ¥- ax” 2% z = 5 5 = = 3 a - z : a = 3 2 2 2 
x” 
#E OX oe v - = = 2 = 5 2 & ; 3 é e x x 2 F; & B A 
*k XK xX e A = v - is 2 : 3 : = ‘ 5 a E 3 2 é . 2 
kt oes ce) XK Se . Ke’ OXE AES KE Sx YS ow 2 a oe 2 P a 5 vj 

x” x" 
ee OX “4 ame Ms v b Yi: XY . 2 xe a = : & E F 3 a 

Nm te OX eee 
x 
EK z z xX 5 xe Z z v “ = Z 5 2 2 Z 7 5 E “ 5 
EE 4 z 5 3 = = v = 2 2 E B q 4 
ek OX g Vie XS xX é = 2 Z < “ 
xt 

** x’ xX xX vi x # xe :: s e 

x™ 
EE a x a a e 5 ¥ v z 

Ker 


Table 3. Juvenile platypuses originally captured in the study which later bred within the study area. ** 
emerged; X not lactating; V lactating ‘ month of capture (eg November); - not captured/sampled; + First 
breeding in second breeding season after emergence from nesting burrow; # First breeding at 3-4 breeding 
seasons after emergence. 


Proc. Linn. Soc. N.S.W., 125, 2004 


230 


T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 


Animal 77/ 78/ 79/ 80/ 81/ 82/ 83/ 84/ 85/ 86/ 87/ 88/ 89/ 90/ 91/ 92/ 93/ 94/ 95/ 96/ 97/ 98/ 99/ O00/ O1/ 02/ 03/ 
7879 80 92. 93 94 95 96 97 +98 +99 #00 OFF 02 «203 ~=04 

FA019 - - 

FA124 

FA126 

FA133 

FA147 

FA158 

FA161 

FA185 

FA188 

FJ222 

FJ235 

FJ248 

FJ272 

FA280 

FA282 

FA370 

FA333 

FA335 

FA370 

FA391 

FA423 

FJ469 wes ory, OK 

FA513 

FAS14 

FA529 

FA534 2 

FA535 Xx 

FA559 


i) 
pay 
le} 
N 
oO 
(*) 
i>’) 
db 
oO 
a) 
Oo 
N 
ic.) 
| 
oo 
-') 
i) 
=) 
‘oe 
(—) 
K-) 
= 


RN 
<M NS 


LAK MK 
2 
2 
2 
S 


mK <K 
i KKK 
eK << 
bs Sa 
Ko 


A 
Stat 
NN Ne 
NN 

\ AK 


wo 
SSi4 
KK NAM 


AK 
\ 


iN a 


Table 4. Female platypuses captured in November to January in consecutive breeding seasons in the 
whole study area during the study. ** newly emerged juveniles; V lactating; X not lactating; ? caught 
during February, March or April (could have bred but have finished lactating; see Table 2). 


231 


Proc. Linn. Soc. N.S.W., 125, 2004 


BREEDING IN FREE-RANGING PLATYPUSES 


100 4 


90 - 


% Lactating Females or Juveniles in Samples 
to (eS) = un fo) I oo 
oO (>) (>) oO Oo Oo So 
I J 1 1 4 1 


1979/80 SS Ee ae ee Se SS 

1981/82 Ee ESS ae ae SS SSS See 

1983/84 ER a eS 
1984/85 eee eR ae ee 

1986/87. S| 

1987/88 See ee ee ree 

(Cy 0) | =| 


1999/00 Se 
1990/91 eee 
CM) | Sa ae A 


fon = ise) \o On oS ty SO? co TON Oe est SIN oe 
= 20 ge 2 Ds 3. Bi. Wea Se oO oS Fe 
225 8 2 A 2-2 8 2S ese Soe 
< 
Breeding Seasons 


Figure 1. Percentages of lactating females captured in relation to total adult females caught in each 
December sample (n = 21) and the percentages of juveniles in each February and/or March sample (n 
= 22) in the core study area. Open bars = lactating females (5 seasons not sampled); solid bars = 
juveniles (4 seasons not sampled; 1988/89 and 1991/92 seasons no juveniles were caught). 


DISCUSSION 


The low overall percentages of lactating 
animals in samples caught in September (3.8%), 
October (20%), November (28%) and in February 
(17.6%), March (0.6%), April (0%), contrasted with 
higher percentages in December (41.3%) and January 
(40%). This indicated a spread in the breeding season. 
However, it appears that the majority of animals were 
breeding around the same time, with a few individuals 
breeding earlier (eg. one already lactating by the end 
of September) and a few later (eg. one still lactating in 
March; Table 1). 

The sequential recaptures of lactating females 
within the same breeding seasons provided evidence 
that lactation in the wild can last at least 98 days (3.3 
months) but is unlikely to exceed four months. This 
suggestion is supported by the distribution of lactating 
females in the various months (Table 1), combined 
with the observation that all but one of 31 females 
lactating when captured in December or January (97%) 
had ceased lactation on their subsequent recapture in 
March (Table 2). 


i) 
oS) 
ie) 


Nestlings in the wild may be weaned more 
rapidly than those bred in captivity. Lactation in captive 
animals has been reported to continue as long as 145 
days (4.8 months; Holland and Jackson 2002; 
Healesville Sanctuary and Taronga Zoo, unpublished) 
and requires lactating females to consume up to 100% 
of their body weight in food during peak lactation 
(Holland and Jackson 2002). It may be that the young 
are weaned more quickly in the wild depending on the 
local availability of macroinvertebrate food items 
(Faragher et al. 1979) for the breeding females. 
Certainly in some years of this study, lactating females 
were in poorer body condition, based on observations 
of tail fat reserves (Temple-Smith 1973; Grant and 
Carrick 1978) and general body condition. 
Interestingly, both the lactating females and the 
captured juvenile animals in the 2002/03 breeding 
season, at the end of a very severe drought, were judged 
to still be in good body condition. However, four 
lactating females captured at the beginning of January 
2004 appeared to be in poorer condition, despite the 
river flows being an improvement on those of the: 
previous breeding season, when surface flows in the 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M. GRIFFITHS AND P.D. TEMPLE-SMITH 


study area stopped for several weeks (pers. comm. 
from local residents). 

Recapture rates of juvenile platypuses during 
the various studies in the area was quite low. Thirty- 
two percent of female platypuses first captured as 
juveniles were recaptured compared to only 14% for 
males (Grant 2004; this volume). In spite of this, only 
12 female juveniles were recruited into the breeding 
population. 

Although the breeding season was 
predictable, the breeding of individuals in any one 
season was much less predictable, with varying 
numbers of non-breeding females in any sample, 
individuals not breeding until later in life and breeding 
animals failing to breed in consecutive seasons (Table 


4). Similar observations have been made with > 


platypuses in captive conditions, where no individuals 
have so far bred in successive breeding seasons 
(Holland and Jackson 2002; Healesville Sanctuary and 
Taronga Zoo, unpublished). Temple-Smith and Grant 
(2001) have speculated whether resource availability, 
social organisation or genetic factors are involved in 
this uncertain breeding in the species but little is known 
of any of these aspects of platypus biology. 

A decline was expected in the number of 
platypuses breeding and/or the number of juveniles 
recruited to the population after the mid 1980s, when 
sand slugs began to reduce the pools available for 
foraging and the provision of refuge areas during 
drought. Surprisingly, no such overall trend occurred 
in either numbers breeding or in recruitment data and 
there is no ready explanation for the considerable 
variation in the numbers breeding between the seasons 
covered by the study. Such variations must be 
attributable to more subtle changes occurring in the 
environment and/or to unexplained sampling effects. 

Both breeding success and recruitment fell 
sharply in the 1982/83 breeding season, corresponding 
to the end of a long and severe drought, which lasted 
from October 1978-February 1983. The effects of the 
drought provided an explanation for the observation 
that three females lactating in the 1981/82 season did 
not breed in the 1982/83 season. However, there were 
no similar trends recorded in the 1993-95 or 2001-03 
droughts, although during the latter, lactation and 
recruitment percentages were slightly below the mean 
values for each (Fig. 1). As discussed above, all 
lactating females and juveniles captured in the 2002/ 
03 sampling were considered to be in good body 
condition in spite of the severe drought conditions 
which existed at the time. 

During the 1988/89 and 1991/92 breeding 
seasons no juvenile platypuses were captured. There 
is no obvious reason for the observed lack of 
recruitment in 1988/89, but two local over-bank flood 


Proc. Linn. Soc. N.S.W., 125, 2004 


events in late December/early January (pers. comm. 
from local residents) of 1991/92 may have drowned 
many nestlings confined to burrows during that season. 
This would explain the failure to capture juveniles in 
a year when the percentage of lactating females in the 
previous December had been slightly higher the mean 
value (Fig. 1). 

Prior to enactment of legislation protecting 
platypuses in all states of Australia (1892 in Victoria 
to 1912 in South Australia; Grant and Denny 1991), 
thousands were hunted for their fur. Their numbers 
are reputed to have declined dramatically, although 
rebounding since protection has been enforced (Grant 
and Denny 1991; Grant and Temple-Smith 1978). The 
species is currently listed as protected, but is either 
regarded as ‘common’ or not threatened, in all states 
(except for South Australia, where it is probably now 
extinct, except for an introduced population on 
Kangaroo Island). In spite of this, there is concern at 
the fragmentation of populations in some river systems 
and in small local populations as a result of habitat 
degradation, illegal and recreational fishing and 
encroaching effects of urban and regional development 
(Grant and Temple-Smith 2003). While this study 
demonstrates that the species has continued to survive 
and reproduce in the upper Shoalhaven River in spite 
of considerable riparian and riverine degradation, the 
effects of drought and the combination of both of these 
perturbations, further investigation leading to a 
complete understanding of the factors determining the 
uncertain breeding in the species is critical to its 
conservation. Many questions regarding the population 
biology and reproduction of Ornithorhynchus anatinus 
still remain unanswered but the long-term studies 
reported here and in Grant (2004, this volume) have 
gone some way to providing a greater understanding 
of some aspects of the species’ field biology, which 
could not have been achieved by a study of shorter 
duration. 

ACKNOWLEDGMENTS 


Merv Griffiths, a friend, colleague and our co- 
author died on 06 May 2003. The many other friends and 
colleagues who were instrumental in the success of field 
work, in often severely inclement conditions, over the years 
are too numerous to name individually but the Heath family, 
Paul Anink, Marie-Loiuse Lissone, David Read and Gina 
Grant deserve special mention. All are gratefully 
acknowledged. Some of the work reported was done while 
in receipt of funding from the Environment Australia (then 
Australian National Parks and Wildlife Service) and the 
Australian Research Council (then Australian Research 
Grants Committee). The late Athol MacDonald and the 
Izzard and Laurie families are acknowledged for their 
permission to access the river and creek on the properties 
managed or belonging to them, and for their friendship and 


UES) 


BREEDING IN FREE-RANGING PLATYPUSES 


assistance in various aspects of the field work. The work 
was carried out under NSW National Parks and Wildlife 
Service Scientific Investigations Licence A184, New South 
Wales Fisheries Scientific Research Permit F84/1245 and 
University of New South Wales Animal Care and Ethics 
Approvals 94/91, 97/46 and 00/45. 


REFERENCES 


Faragher, R.A., Grant, T.R. and Carrick, F.N. (1979). 
Food of the platypus, Ornithorhynchus 
anatinus, with notes on the food of the brown 
trout, Salmo trutta, in the Shoalhaven River, 
New South Wales. Australian Journal of 
Ecology 4: 171-179. 

Gibson, R.A., Neumann, M., Grant, T.R. and Griffiths, M. 
(1988). Fatty acids of the milk and food of the 
platypus (Ornithorhynchus anatinus). Lipids 23, 
377-379. 

Grant, T.R. (2004). Captures, capture mortality, age and 
sex ratios of platypuses, Ornithorhynchus 
anatinus, during studies over 30 years in the 
upper Shoalhaven River in New South Wales. 
Proceedings of the Linnean Society of New 
South Wales 125, 217-226. 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist 18: 
133-135. 

Grant, T.R. and Carrick, F.N. (1978). Some aspects of the 
ecology of the platypus, Ornithorhynchus 
anatinus in the upper Shoalhaven River, New 
South Wales. Australian Zoologist 20: 181-199. 

Grant, T.R. and Denny, M.J.S. (1991). Historical and 
Current Distribution of the Platypus in 
Australia, with Guidelines for the Management 
and Conservation of the Species. Unpublished 
Report to Australian National Parks and 
Wildlife Service by Mt. King Ecological 
Surveys. 

Grant, T.R., Griffiths, M. and Leckie, R.M.C. 1983. 
Aspects of lactation in the platypus, 
Ornithorhynchus anatinus (Monotremata), in 
waters of eastern New South Wales. Australian 
Journal of Zoology 31, 881-889. 

Grant, T.R. and Griffiths, M. (1992). Aspects of lactation 
and determination of sex ratios and longevity in 
a free-ranging population of platypuses, 
Ornithorhynchus anatinus, in the Shoalhaven 
River, New South Wales. In “Platypus and 
Echidnas’. (Ed M.L.Augee). pp. 80-89. (Royal 
Zoological Society of NSW: Sydney). 

Grant, T.R. and Temple-Smith, P.D. (1998). Field biology 
of the platypus (Ornithorhynchus anatinus) - 
historical and current perspectives. Transactions 
Royal Society London Series B 353, 1081-1091. 

Grant, T.R. and Temple-Smith, P.D. (2003). Conservation 
of the platypus, Ornithorhynchus anatinus: 
Threats and challenges. Aquatic Health and 
Management 6, 1-15. 


234 


Grant, T.R. and Whittington, R.J. (1991). The use of 
freeze-branding and implanted transponder tags 
as a permanent marking method for platypuses, 
Ornithorhynchus anatinus (Monotremata: 
Ornithorhynchidae). Australian Mammalogy 14: 
147-150. 

Griffiths, M., Elliott, M.A., Leckie, R.M.C. and Schoefl, 
G.I. (1973). Observations on the comparative 
anatomy and ultrastructure of mammary glands 
and on the fatty acids of the triglycerides in 
platypus and echidna milk fats. Journal of 
Zoology [London] 169, 255-275. 

Griffiths, M., Green, B., Leckie, R.M.C., Messer, M. and 
Newgrain, K.W. (1984). Constituents of 
platypus and echidna milk with particular 
reference to the fatty acid complement of the 
triglycerides. Australian Journal of Biological 
Science 37, 323-329. 

Griffiths, M., McIntosh, D.L. and Leckie, R.M.C. (1972). 
The mammary glands of the red kangaroo with 
observations on the fatty acid components of the 
milk triglycerides. Journal of Zoology [London] 
166, 265-275. 

Griffiths, M., McKenzie, H.A., Shaw, D.C. and Teahan, 
C.G. (1985). Monotreme milk proteins: echidna 
and platypus “whey” proteins. Proceedings of 
the Australian Biochemical Society. 17, 25. 

Holland, N. and Jackson, S.M. (2002). Reproductive 
behaviour and food consumption associated 
with captive breeding of platypus 
(Ornithorhynchus anatinus). Journal of Zoology 
[London] 256, 279-288. 

Joseph, R and Griffiths, M. (1992). Whey proteins in early 
and late milks of monotremes (Monotremata; 
Tachyglossidae, Ormithorhynchidae) and of the 
tammar wallaby (Macropus 
eugenii)(Marsupialia; Macropodidae). 
Australian Mammalogy 15, 125-127. 

Messer, M., Gadiel, P.A., Ralston, G.B. and Griffiths, M. 
(1983). Carbohydrates of the milk of the 
platypus (Ornithorhynchus anatinus). 
Australian Journal of Biological Science. 36, 
129-138. 

Parodi, P.W. and Griffiths, M. (1983). A comparison of 
the positional distribution of fatty acids in milk 
triglycerides of the extant monotremes platypus 
(Ornithorhynchus anatinus) and echidna 
(Tachyglossus aculeatus). Lipids 18, 845-847. 

Teahan, C.G., McKenzie, H.A. and Griffiths, M. (1991). 
Some monotreme milk “whey” and blood 
proteins. Comparative Biochemistry and 
Physiology. 91B, 99-118. 

Temple-Smith, P.D. (1973). Seasonal breeding biology of 
the platypus, Ornithorhynchus anatinus (Shaw 
1799) with special reference to the male. PhD 
thesis, Australian National University, Canberra. 

Temple-Smith, P.D. and Grant, T.R. (2001). Uncertain 
breeding: A short history of reproduction in 
monotremes. Reproduction, Fertility and 
Development 13, 487-497. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Depth and Substrate Selection by Platypuses, Ornithorhynchus 


anatinus, in the Lower Hastings River, New South Wales 


Tom GRANT 


School of Biological, Earth and Environmental Sciences, University of New South Wales, NSW 2052 


t.grant@unsw.edu.au 


Grant, T. (2004). Depth and substrate selection by platypuses, Ornithorhynchus anatinus, in the lower 
Hastings River, New South Wales. Proceedings of the Linnean Society of New South Wales 125, 235- 
241. 


Platypuses were observed foraging most frequently in water >1 metre in depth during normal (91.3%) and 
drought (82.1%) conditions. Mean water depth in the study pools was 1.08+0.66 and 0.86+0.61 metres 
during normal and drought conditions respectively. The distribution of depths in the study area was 
significantly different from the distribution of depths where platypuses were observed during normal (Chi? 
= 90.2; p < 0.01) and drought conditions (Chi? = 37.35; p < 0.01). Platypuses were apparently not simply 
utilising depths in relation to their occurrence but preferring to forage in water deeper than 1.5 metres and 
avoided depths < 1 metre. Overall distribution in numbers of platypuses observed foraging over different 
benthic substrate types was not significantly different (Chi? = 12.9; p > 0.05) from the distribution of these 
substrate categories in the study area. However, when the substrates were considered separately, significant 
preference was shown for cobbled substrate (Chi? = 18.4; p < 0.01) and avoidance of gravel (Chi? = 9.7; p 
< 0.01). These observations have implications for catchment, stream and riparian management, where activities 
leading to sedimentation and reduced flushing flows may reduce depths and/or alter the distribution of 


preferred foraging substrates. 


Manuscript received 2 September 2003, accepted for publication 7 January 2004. 


KEYWORDS: depth, foraging, Hastings River, Ornithorynchus anatinus, platypus, substrate. 


INTRODUCTION 


During foraging in the wild, platypuses dive 
to feed almost exclusively on small benthic invertebrate 
animals (Faragher et al. 1979; Grant 1982), which are 
normally unevenly and often sparsely distributed in a 
variety of substrates and depth zones (Boulton and 
Brock 1999; Elhott 1977; Young 2001). The platypus 
is small, has a high metabolic demand to regulate its 
body temperature in water and has an estimated 
maximum aerobic capacity for diving of only 40-60 
seconds (Bethge 2002; Bethge et al. 2001; Evans et 
al. 1994; Grant and Dawson, 1978). Consequently its 
foraging is restricted to relatively shallow depths and 
the species is seldom reported occurring in deep lakes 
or impoundments (Bryant 1993; Grant 1991; McLeod 
1993; Ellem et al. 1998; Ellem and McLeod 1998). 
The current study reports on observations of depths of 
diving and foraging over different substrates by 
platypuses in a coastal river in New South Wales during 
drought and normal flow conditions. Grant and Bishop 
(1998) discussed the importance of the measurement 
of physical habitat variables associated with platypus 
occurrence as a means of assessing possible impacts 


of stream use and management activities. As part of 
the monitoring and detection of possible environmental 
effects of the Hastings District Water Supply 
Augmentation Scheme, the utilisation of depth and 
substrate categories by foraging platypuses was 
investigated. 


METHODS 


Study area 

The study was undertaken in two separate 1.5 
kilometre sections of the lower Hastings River near 
Wauchope in New South Wales. Immediately above a 
large riffle separating the riverine section from the 
upper estuary tidal influence, the study area consisted 
of a series of pools, riffles and runs, with the banks 
predominantly consisting of earth consolidated by the 
roots of riparian vegetation, but with a number of 
gravel/cobble bars and sections of bedrock also present 
. Predominantly surrounded by agricultural land, 
especially pastures for dairy cattle, much of the stream 
bank supported a narrow strip of vegetation consisting 
of river oaks (Casuarina sp), rainforest species (e.g. 
Waterhousea floribunda, Ficus coronata) and 


DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 


introduced weed species (e.g. willows, Salix sp; 
Lantana camara; privet, Ligustrum spp, wild tobacco, 
Solanum mauritianum). A range of macrophyte species 
also occurred in the stream (especially Myriophyllum 
verrucosum), although these were reduced to low 
incidences after several flood events. The aquatic grass, 
Potamophila parviflora, was also common along 
several sections of bank and occurred in island clumps 
within several sections of the stream. 


Sampling 

The 3 kilometres of river surveyed consisted 
of four riffle areas and five pools. Each section was 
surveyed in both directions during the two hours prior 
to darkness and immediately after first light in winter 
(late May to early July) and spring (September to 
October) over six years from 1998-2003 (88 
longitudinal transects x 2 river sections = 176 
longitudinal transects; i.e. the whole 3 kilometres was 
surveyed 88 times). During 1998-2000 the same 
number of longitudinal transects (16) was surveyed in 
both winter and spring but from 2001 to July 2003 
fewer were surveyed in winter (8) and more in spring 
(24), as lower numbers of platypuses were normally 
observed during the winter period. Depth and the 
predominant substrate type were recorded at the point 
where each platypus was first seen foraging. It should 
be noted that visibility, due to turbidity and/or poor 
light conditions, often meant that a determination of 
substrate could not be made at all of these points. 
During the 2001 and 2002 sampling periods, visibility 
was so low (probably due to the high abundance of 
phytoplankton) that the substrate could be observed 
only in few instances where platypuses were foraging. 


Physical habitat analysis 

The stream was paced out into 60 x 50 metre 
sections (3 km) and marked at each point with brightly- 
coloured flagging tape. At each of these points depth 
measurements were made at both edges (approximately 
2 metres from bank) and in the middle of the river 
(using a weighted line or the kayak paddle graduated 
in 25 cm units). These depth measurements (n = 174) 
provided a measure of the distribution of depth 
categories in the study area (Figs 1 and 2). The 
occurrence of benthic substrates (mud, sand, gravel, 
cobble and bedrock) was scored on a scale of 1-5 (using 
the following estimated percentage cover of each 
substrate type; 0 = 0%; 1 = 0-5%; 2 = 5-25%; 3 = 25- 
50%; 4 = 50-75%; 5 = >75%) along three transects 
parallel to the stream bank between corresponding 
depth measurement points at the beginning and end of 
each 50 metre section. Substrates along these transects 
were not homogeneous, often with some of each type 
within a single transect. However, the predominant 


236 


substrate types (score of 4 or 5; i.e. >50% estimated 
coverage) for each transect (n = 174) were used as a 
measure of the distribution of the occurrence of 
substrates (Fig. 3). Data on depths and substrate 
distribution were collected once during July 2000 but 
depth measurements were repeated in October 2002 
when the river was under severe drought conditions 
and was barely flowing. 


Data analysis 

The null hypothesis being tested was that the 
occurrence of platypuses across depth and substrate 
categories was the same as the occurrence of these 
categories in the study area. The overall distribution 
of numbers of platypuses observed foraging within the 
various depth and substrate categories and the recorded 
numbers of occurrence of these physical attributes in 
the stream (n = 174 samples), were compared using 
Chi’ analysis (Statistica, StatSoft Inc.) with expected 
values calculated using 2 x 5 contingency tables (Bailey 
1969). Comparisons between numbers of platypuses 
observed foraging at specific depths or on specific 
substrates compared to those not foraging at these 
specific depths or substrates (i.e. all other depths or 
substrate categories) were made using Chi? for 2 x 2 
contingency tables. Comparisons between drought and 
non-drought measurements of depth were made using 
Student’s t-tests for unpaired samples (Bailey 1969; 
Statistica; StatSoft Inc). Indices of selection/avoidance 
(Response Index) of depth or substrate categories by 
platypuses were calculated as: 
Response Index = 

% Occurrence of platypuses in a 
depth or substrate category 
% Occurrence of that depth or 
substrate category in the stream 

An index of greater than unity suggested a selection 
response and less than one an avoidance response to a 
depth or substrate category. All means given are + 
Standard Deviation. 


RESULTS 


Depth selection 

The mean depths of the stream during non- 
drought and drought were significantly different, being 
1.08+0.66 and 0.86+0.61 metres respectively (t = 3.48; 
p < 0.001). The maximum water depth recorded by 
these transect-based measurements was 2.75 metres 
but platypuses were recorded foraging at depths of up 
to 3.2 metres and the maximum depth recorded 
opportunistically (not along transects or at foraging 
sites) was just under 4 metres. 

Figure | shows the numbers of observations 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


40 5 


pall 


0.0-0.5 0.6-1.0 1.1-1.5 1,5-2.0 
Depth Category (metres) 


% Occurrence of platypuses or Depths 
recorded/Depth category 
WwW 


Figure 1. Percent occurrence of platypuses observed 
foraging in various depth categories (n = 127 
observations; white bars) and the percentage of 
occurrence of these depth categories (n = 174 
observations; black bars) in the Hastings River 
study area during non-drought conditions. 


Halbho 


0.0-0.5 0.6-1.0 leila) 1.5-2.0 
Depth Category (metres) 


% Occurrence of platypuses or Depth: 
recorded/Depth category 
Ww 
So 
4 


Figure 2. Percent occurrence of platypuses observed 
foraging in various depth categories (n = 28 
observations; white bars) and the percentage of 
occurrence of these depth categories (n = 174 
observations; black bars) in the Hastings River 
study area during drought conditions. 


of platypuses foraging in the various depth categories 
during a range of non-drought conditions. These data 
show that 91.3% of the platypuses (total n = 127) were 
observed foraging in depths greater than 1 metre, 
despite this depth category occurring in only 39.1% 
of the study area. The distribution of numbers of 
platypuses foraging within the depth categories and 
the recorded numbers of occurrence of these categories 
were significantly different (Chi* = 90.2; p < 0.01). 
Platypuses showed significant preferences for 
water deeper than 1.5 metre (Response Indices 3.1- 


Proc. Linn. Soc. N.S.W., 125, 2004 


60 


50 


40 


30 


20 4 


% Occurrence of platypuses or substrates/categoi 


Mud Sand Gravel Cobbles Bedrock 


Substrate Category 


Figure 3. Percent occurrence of platypuses observed 
foraging in various substrate categories (n = 56 
observations; white bars) and the percentage of 
occurrence of these substrate categories (n = 174 
observations; black bars) in the Hastings River 
study area during non-drought and drought 
conditions. 


3.2) and avoidance of depths of less than 1 metre 
(Response Indices 0.2-0.7). Foraging within the 1.1- 
1.5 metre category showed no significant preference 
or avoidance by platypuses (Table 1a). 

During severe drought conditions (July and 
October 2002) there was a significant difference 
between the distribution of foraging platypuses (n = 
28) and the distribution of recorded depth categories 
(Chi? = 37.35; p = < 0.01; Fig. 2). Response Indices 
showed a similar, but apparently more marked pattern 
towards preference for depths > 1 metre (Response 
Indices 2.6-6.2) and avoidance of shallower depths 
(Response Indices from 0-0.7). Considering the small 
sample sizes of platypuses foraging in specific depth 
categories (n = 0-10) no Chi’ analyses were attempted 
on these data collected during the drought. 


Substrate selection 

The numbers of platypuses observed foraging 
on particular benthic substrate types (n = 56) are shown 
in Figure 3. The overall distribution of numbers of 
platypuses foraging over the various substrates was 
not significantly different from the distribution of these 
substrates (Chi? = 12.9; p > 0.05). However, only 
26.8% of platypuses were found foraging over gravel 
substrates, while this substrate type was the most 
abundant in the study area (50.9%). While cobbles 
made up only 12.7% of the available substrate, 28.6% 
of the platypuses observed were foraging over this 
substrate. Response Indices of 0.5 and 2.3 respectively 


237 


DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 


Table 1. Chi’ and probability values (2x2 contingency tables; Statistica; StatSoft Inc.) for comparisons 


of: 


a. platypuses foraging at specific depths and those not foraging at each of these depths 


Depth Category —0.0-0.5 m 0.6-1.0 m 
Chi * 27.1 38.8 
<10,01* < 0.01* 


1.1-1.5 m 1.6-2.0 m >2m 
43 18.8 4.3 
> 0.05 < 0.01* < 0.01* 


b. platypuses foraging on specific substrates and those not foraging on each of these substrates 


Substrate Category Mud Sand Gravel Cobbles Rock 
Chi 2 0.01 0.96 9.70 18.70 0.26 
> 0.05 > 0.05 < 0.01* < 0.01* > 0.05 


* indicates statistical significance. 


for gravel and cobbles, suggested avoidance of the 
former and preference for the latter. Individual 
comparisons between platypuses foraging over specific 
substrates compared to all other substrates showed 
significant differences from expected for gravel and 
cobbles but not for the other substrate types (Table 
1b). 


DISCUSSION 


The study suggested that platypuses observed 
in the early morning and late afternoon/evening were 
selecting the deeper sections of their habitat, with 
91.3% of the observed individuals foraging in water > 
| metre in depth and 33.1% foraging in water of greater 
than 2 metres, which constituted only 39.1% and 10.3% 
of the area respectively during normal flow conditions. 
Even during the severe drought conditions, 82.1% of 
platypuses were still observed foraging in water deeper 
than | metre, despite the fact that the proportion of 
recorded depths > 1 metre had decreased by 11.5%. 
During the drought observations, 48.3% of the area 
had a depth of 0.0-0.5 metres but no platypuses were 
observed forging in this depth category. Thus, 
platypuses appeared to show preference for foraging 
in deeper areas and an avoidance of shallower depths 
within the area during the study in both drought and 
non-drought conditions. 

These observations were similar to those 
reported for a study in a small alpine lake in Tasmania 
(Lake Lea). While reporting a maximum dive depth 
of 8.77 metres, Bethge (2002) and Bethge et al. (2003) 
found that the majority of dives recorded for platypuses 
fitted with data loggers were to depths of less than 
three metres (98% of dives), with a mean diving depth 
of 1.28 metres. These workers also found a large 


238 


proportion of the foraging dives were to depths of less 
than one metre (48%), although, during winter (when 
the lake level was higher than in summer), most dives 
were to depths greater than 1 metre. Platypuses were 
monitored foraging in the same area, rather than 
moving to the shallower parts of the lake during the 
winter, suggesting that foraging was determined by 
factors other than depth preference, possibly including 
substrate type and/or availability of benthic food 
species. However, the workers in this study did not 
report on these possibilities. 

Rohweder (1992), Bryant (1993) and 
McLeod (1993) also reported platypuses foraging in 
water less than 5 metres in depth. While Ellem et al. 
(1998) found increasing depth of pools (up to 2 metres) 
to be positively related to the observed presence of 
platypuses in 36 pools on the Macquarie River system 
in the Bathurst area of the central tablelands of New 
South Wales, Ellem and McLeod (1998) found radio- 
tacked platypuses using shallower parts of some 
sections of a weir pool in the Duckmaloi River near 
Oberon in New South Wales. 

Bethge (2002) also reported platypuses 
foraging in deeper areas and spending less time on the 
surface of the water in Lake Lea during daylight hours 
than at night. He speculated that these behavioural 
changes may have been related to avoidance of 
predators. Little is known regarding predation by 
indigenous predators which could take foraging 
platypuses from the water. Predation by the introduced 
red fox (Vulpes vulpes) on platypuses moving through 
or foraging in shallow water, such as riffle areas, has 
been reported (Serena 1994; Grant 1993; Anon. 2002) 
and the species has been included as a possible food 
item of wedge-tailed eagles (Aquila audax)( Rakick 
et al. 2001; Marchant and Higgins 1993). 

Several white-breasted sea eagles (Haliaeetus 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT 


leucogaster) and several ospreys (Pandion haliaetus) 
were observed at the Hastings River study site. Both 
of these species were seen taking fish from the surface 
of the water. Although a grey goshawk (Accipiter 
novaehollandiae) has been reported attacking a 
juvenile platypus on land (Richards 1986), it seems 
unlikely that either this species or the osprey would 
be large enough to take even a juvenile platypus (which 
are around 65-70% adult weight when they first leave 
the nesting burrows; Grant and Temple-Smith 1998) 
from the water. It is possible however, that the sea 
eagle may represent a potential predator of the 
platypus. During the study a sea eagle was observed 
retrieving a large dead Australian bass (Macquaria 
novemaculeata) of 485 mm in length, and estimated 
to weigh 2.5 kg (Harris 1987), from the bottom of a 
pool. The eagle was unable to fly with the fish and 
dragged it to a nearby gravel bar, where part of the 
flesh was eaten before darkness fell. Soon after first 
light the following morning the eagle was seen carrying 
off the remaining carcass of the bass. Interestingly, 
the platypus does not seem to have been recorded as a 
food item of this species of large eagle (Marchant and 
Higgins 1993; Olsen 1999). 


Substrate selection 

Higher invertebrate productivity is often 
associated with areas where logs, roots and vegetation 
provide a range of habitats for an array of types of 
benthic invertebrate species and coarse substrates 
(gravel, cobbles, rocks) provide fixed habitat, rather 
than a shifting substrate, such as sand and fine sediment 
(Young 2001; Boulton and Brock 1999; Smith and 
Pollard 1998). Data in the study were restricted by a 
small sample size due to the inability to observe the 
type of substrate over which platypuses were seen 
foraging in times of high turbidity and/or poor light 
conditions. However, there was some indication that 
platypuses were avoiding sections of the study area 
which consisted mainly of mud or sand (Fig 3) but 
this was not statistically significant (Table 1b). There 
appeared to be marked avoidance of gravel (the most 
abundant substrate; 50.9% of the area) and preference 
shown for areas where cobbles were the predominant 
substrate (12.7% of the area). Both of these trends were 
statistically significant (Table 1b). 

The complexity of benthic habitat has been 
previously identified as being positively related to the 
occurrence of platypuses (Rohweder 1992) and Serena 
et al. (2001) found a positive relationship between 
numbers of radio-tracked platypuses and the 
occurrence of coarser substrates, including gravel, 
pebbles, cobbles, large rocks and coarse particulate 
organic matter. These observations may be related to 
the distribution of benthic food organisms but this was 


Proc. Linn. Soc. N.S.W., 125, 2004 


not investigated in the present study. No explanation 
of the apparent avoidance of gravel substrate in the 
present study is suggested as the species has been 
observed by the author foraging on gravel substrates 
in other areas. 


Implications for stream management 

The development of adaptive management 
strategies for streams, particularly with regard to water 
extraction and the operation of impoundments, should 
consider flows which maintain pool depth and benthic 
habitat diversity by preventing the accumulation of 
sand and fine sediments. The removal of riparian 
vegetation, erosion as a result of unrestricted stock 
access to stream banks and poor catchment 
management practices have also resulted in the infilling 
of pools by sand ‘slugs’ in many streams in eastern 
Australia (Brooks and Brierley 1996; Boulton and 
Brock 1999; Brierley et al. 1999; Grant et al. 2003). 

Grant and Bishop (1998) encouraged the use 
of physical habitat analysis, considering broad habitat 
variables normally associated with platypus 
occurrence, in any attempts to monitor and/or predict 
effects of human activities impinging on streams and 
their catchments. More recently a habitat simulation 
model was used by Davies and Cook (2001) to generate 
weighted useable habitat area estimates for the platypus 
at various proposed discharge regimes in a regulated 
river in Tasmania. This model used more specific 
habitat requirements of the species in terms of depth, 
velocity and substrate, calculating habitat preference 
curves based on available information from the 
literature and from experts in the field. These authors 
observed that: “Platypus[es] are known to feed in very 
shallow water and up to ca 1-3 m” and “foraging is 
optimal at depths of <2 m” and “platypus[es] actively 
feed in silt, sands, finer gravel substrates, and are 
known to forage on coarse gravel to smaller cobble 
substrate. Feeding activity is not deemed to be efficient 
or to frequently occur on coarse cobble, boulder or 
bedrock substrates” 

While the important modelling work of 
Davies and Cook (2001) drew upon the information 
available to the authors at the time, the data from the 
current study and that from Serena et al. (2001) do not 
totally support the information used to generate their 
habitat preference curves for the species in mainland 
sites. It is vitally important that studies seeking to 
predict possible impacts of human activities on the 
platypus (or any other species) must consider the 
widest range and the most currently available 
information on which to base assumptions. 

Too often, assessment of possible 
environmental impact is based on ‘conventional 
wisdom’ which may be enshrined in publications 


239 


DEPTH AND SUBSTRATE SELECTION BY PLATYPUSES 


which are either not current or poorly researched. It is 
not acceptable, for example for one Environmental 
Impact Statement to become the main reference for 
statements or predictions made in another such 
document, without reference to the wider and most 
current scientific literature. The following example 
from the assessment of dam development on the 
Burnett River in Queensland sharply illustrates these 
concerns. 

Arthington (2000) suggested that 
“platypus[es] feed by scooping prey items and mud 
into cheek pouches in the mouth and grinding the 
mixture to a sludge before digesting it”. Based on this 
suggestion, the resultant Environmental Impact 
Statement concluded that “the deposition of sediments 
in the shallower areas of the dam would provide extra 
foraging area for Platypus[es]” as “an increase of 
available muddy substrate would provide more 
foraging area. Hard substrates offer less feeding 
opportunities because prey cannot be as easily scooped 
and ground up if they are on hard substrates or if the 
scooped material contains large pebble material” 
(Anon. 2003). Neither the literature nor the current 
study support the original suggestion by Arthington 
(2000) which consequently has led to a very equivocal 
prediction regarding the possible impact of dams on 
platypus foraging. It is crucial that such equivocal 
predictions do not become established in the literature 
consulted by those carrying out environmental impact 
assessment studies. 


ACKNOWLEDGEMENTS 


This work was carried out during monitoring 
studies associated with the Hastings District Water Supply 
Augmentation Scheme and was funded by Hastings Council 
and the New South Wales Department of Commerce Offices 
of Government Procurement and Government Business 
(formerly Department of Works and Services). Keith Bishop 
provided valuable advice during this study and comments 
on drafts of this paper. Michael Augee and another 
anonymous referee are thanked for their comments and 
recommendations on the paper. 


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241 


DEPTH AND SUBS ors ASE 


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Distribution of the Platypus in the Bellinger Catchment from 
Community Knowledge and Field Survey and its Relationship 


to River Disturbance 


DaAntEL LuNNEY'!, ToM GRANT? AND ALISON MatTHEws! 


‘Department of Environment and Conservation (NSW), PO Box 1967, Hurstville, New South Wales 2220, 
Australia; "School of Biological, Earth and Environmental Sciences, University of New South Wales, New 


South Wales 2052, Australia. 


Lunney, D., Grant, T. and Matthews, A. (2004). Distribution of the platypus in the Bellinger catchment 
from community knowledge and field survey and its relationship to river disturbance, Proceedings of 
the Linnean Society of New South Wales 125, 243-258. 


Platypus distribution in the Bellinger catchment was investigated using a combination of field and 
community surveys. The field survey in 1996 consisted of netting and observations from the river bank and 
a canoe. The community-based wildlife survey consisted of a questionnaire and colour maps on which 
respondents were asked to mark the locations of sightings. Platypuses were observed or caught in 36 locations 
from all three rivers of the catchment. Two of the three platypuses captured were lactating females. The 
community recorded 123 locations of platypuses. The fact that the wildlife survey yielded similar results to 
the field surveys in identifying the location of individuals highlights the value of community records for 
platypus surveys. There were major floods in 2001, after which we contacted respondents who had reported 
seeing platypuses three years before. Of the 21 respondents who had been near the river since the flood, 7 
had seen platypuses, principally in the tributaries of the Bellinger River. The habitat quality of the rivers 
was evaluated for platypuses and records were related to disturbance and rehabilitation. The species has 
survived in this system, but its future can only be assured by strategies which prevent further degradation of 


its habitat and institute proactive rehabilitation of the damaged sections of these streams. 


Manuscript received 18 August 2003, accepted for publication 8 January 2004. 


KEYWORDS: Catchment management, community wildlife survey, distribution, platypus, river 


management, wildlife. 


INTRODUCTION 


Grant (1992) and Grant et al. (2000) reported 
the distribution of the platypus Ornithorhynchus 
anatinus in New South Wales as having changed very 
little since the occupation of Australia by Europeans. 
Platypuses are considered common in the river systems 
of the coastal, tableland and western slopes (Grant 
1991, 1992) and are frequently reported from streams 
flowing through agricultural land in these areas. In 
three separate surveys in New South Wales, 52-76% 
of recorded platypus sightings were from agricultural 
land (Grant 1991; Lunney et al. 1998; Rohweder and 
Baverstock 1999). The current study investigated the 
distribution of the platypus in a typical north coast river 
system, the Bellinger catchment, where the highland 
headwater streams arise in forested areas but grade 
into predominantly agricultural land (especially cattle 
grazing) towards the coast. 

The distribution of platypuses in the 
Bellinger-Kalang river system was investigated using 


both field and community surveys. Community 
surveys have been successful in identifying locations 
of platypuses (Lunney et al. 1998; Turnbull 1998; 
Rohweder and Baverstock 1999; Otley 2001). This 
survey was part of a wider community-based survey 
of the distribution of a number of key native wildlife 
species in the Bellinger and Kalang valleys adjacent 
to Bellinger River National Park. This study sought to 
assess the co-existence of typical rural community 
activities and wildlife species, including the platypus. 
The field observations and capture of platypuses were 
compared with the questionnaire reports for this and 
other native species in the catchment with the aim of 
testing the hypothesis that information gained from 
survey data provided by the community would be a 
reliable indicator of the presence of wildlife species in 
the area. 

Since our field and questionnaire study, 
Cohen et al. (1998) assessed the Bellinger-Kalang 
catchment using the River Styles framework (Brierley 
et al. 2002) and assigned conservation and 


PLATYPUS IN THE BELLINGER CATCHMENT 


rehabilitation priorities to various stream reaches. 
Using analysis of aerial photography and field 
observations, Cohen et al. (1998) assigned various 
sections of the Bellinger-Kalang catchment to a 
number of River Styles which have particular 
geomorphic attributes. The authors of the study stress 
that these categories are a “record of the character and 
behaviour of sections of river” and “are not a direct 
measure of river condition”. A separate set of 
procedures has been developed to appraise geomorphic 
river condition, building on attributes of river character 
and behaviour that are pertinent to any given River 
Style (Fryirs 2003). The attributes used in discerning 
the River Styles are shown in Table | and include 
channel planform and stability, morphology and 
geometry (depth and width), as well as descriptions of 
geomorphic units (e.g. pools, riffles, point bars), bed 
character (e.g. sand, gravel, cobbles) and vegetation 
character (including riparian vegetation and woody 
debris in the stream). Some of these attributes have 
been identified with the occurrence and foraging 
activity of platypuses and include: 


Channel geometry: pool depth has been 
positively related to the occurrence of the 


species (Ellem et al. 1998; Grant 2004), with 
platypuses often being observed foraging in 
water of greater than one metre depth and less 
than 5 metres. It has been suggested that 
foraging in shallow water can expose 
individuals to predation, especially from the 
introduced fox Vulpes vulpes (Grant and Denny 
1993; Serena 1994; Anon. 2002). 
Geomorphic units: pool/riffle sequences have 
also been found to be associated with the 
presence (Rohweder 1992; Bryant 1993) and 
foraging (Serena et al. 2001) of platypuses, this 
probably being related to the benthic 
productivity of such geomorphic units (Hynes 
1970; Logan and Brooker 1983; Boulton and 
Brock 1999). 

Bed character: the complexity of the bed 
substrate, including large particle sizes (rocks, 
cobbles, pebbles and gravel), has been 
positively related to both occurrence 
(Rohweder 1992) and foraging activities 
(Serena et al. 2001; Grant 2004) of platypuses, 
again probably resulting from greater benthic 
productivity (Hynes 1970; Marchant et al. 
1984; Boulton and Brock 1999). 

Vegetation character: medium-to-large trees, 
especially indigenous species, are associated 
with the use of river reaches by foraging 
platypuses (Serena et al. 2001) and overhanging 


244 


vegetation has also been identified as a variable 
found in areas where platypuses are found 
(Rohweder 1992; Bryant 1993; Serena et al. 
1998). This association between riparian 
vegetation and platypus occurrence is related 
to a number of important functions of such 
vegetation, including stabilisation of the bank, 
provision of cover from predators, supply of 
organic material to the food chains of the stream 
and shade moderating temperature variations, 
especially in summer (Riding and Carter 1992; 
Boulton and Brock 1999). 


The abundance of woody debris, included in 
the “vegetation character” attribute of Cohen et al. 
(1998), is also known to be positively associated with 
platypus occurrence (Rohweder 1992) and foraging 
(Serena et al. 2001), again probably being related to 
the complexity of habitats available for 
macroinvertebrates (Benke et al. 1985; Anon. 1998; 
Anon. 2000a). 

Cohen et al. (1998) also sorted sites in the 
Bellinger-Kalang catchment, grouping river reaches 
into five categories based on procedures outlined in 
Brierley and Fryirs (2000). These are summarised in 
Table 2 and are generally ranked from the least 
(conservation) to the most disturbed sites (degraded), 
although the “strategic” priority #2 sites were identified 
as being more disturbed than the priority #3 sites and 
were given a higher priority as they may impact on 
other sites downstream. This paper analyses these 
Rivers Styles and conservation/rehabilitation 
categories in relation to the data on occurrence of the 
platypus. We propose priorities for conservation and 
rehabilitation of the river system for the future survival 
of platypuses in the Bellinger and Kalang river system 
and in rural areas in general. 

Major floods in the Bellinger catchment in 
early 2001 provided an opportunity to assess the impact 
of floods on a known population of platypuses. A 
follow-up community survey was undertaken to 
determine the survival of platypuses post-flood. 


METHODS 


Study area 

The Bellinger is a fertile river valley on the 
north coast of New South Wales just south of Coffs 
Harbour, and includes the main townships of Bellingen 
and Urunga (Figure 1). The valley extends 
approximately 5O km inland from the coast at Urunga, 
and is approximately 20 km wide from Tucker’s Nob 
range in the north to the Bellbucca ridge in the south. 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


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245 


Proc. Linn. Soc. N.S.W., 125, 2004 


PLATYPUS IN THE BELLINGER CATCHMENT 


Priority 
1. Conservation sites 
2. Strategic sites 
3. High recovery potential 
4. Moderate recovery potential 
5. Degraded 


Nature of Sites 


least disturbed; river structure and vegetation relatively intact 
may be sensitive to disturbance or may affect sites downstream 
may show signs of natural recovery 

moderately degraded with reasonable potential for recovery 
highly degraded reaches with little natural recovery potential 


Table 2. Priority ranking of sites for river rehabilitation in the Bellinger-Kalang catchment (Cohen et al. 


1998). 


Forested lands rise steeply from the valley, forming 
the extremely rugged fringe of the New England 
Plateau. The Bellinger Valley comprises the catchment 
areas of the Bellinger and Kalang Rivers, referred to 
in earlier maps as the North and South Arms of the 
Bellinger River. The third major river of the valley, 
the Never Never River, is a tributary of the Bellinger 
River, which it joins near Gordonville, about 10 
kilometres upstream of Bellingen township. Tidal 
influence extends to Bellingen on the Bellinger River 
and as far as Spicketts Creek on the Kalang River 
(Cohen et al. 1998). The valley and floodplain was 


rapidly cleared by the cedar-getters and during 
settlement in the mid to late 1800s (Anon. 1978; 
Lunney and Moon 1997). Now, the land is used 
primarily for dairying and beef cattle grazing, with 
small areas being planted for crops. The population of 
the valley was 12,253 in 1996, representing a 
population growth of 21 per cent over the previous 10 
years (Anon. 2001a). 

The upper reaches of the streams of the 
Bellinger and Kalang valleys flow through steep 
forested areas in their headwaters, but degradation due 
to human activities, particularly clearing for 


a =a, 


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Figure 1. Location map of the Bellinger catchment, showing main features of the study area and the. 


sections surveyed in the field work. 


246 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


Location Kalang River 
Jamisons Creek 

Date 14.12.96 

Sex Female 

Age adult 

Length (cm) 39.0 

Bill length (cm) 4.9 

Bill width (cm) 4.2 

Weight (g) 730 

Spur 0 

Milk (oxytocin) No 


-Kalang River Bellinger River 
Jamisons Creek Justins Bridge 
14.12.96 IS AALS 
Female Female 
adult adult 
43.5 43.0 
49 Sel 
4.4 43 
905 900 
0 0 
Yes Yes 


Table 3. Details of platypuses captured during field survey in December 1996. 


agriculture, increases from the middle reaches to lower 
reaches upstream of the tidal limits at Bellingen and 
Spicketts Creek (Figure 1). Access of cattle to river 
banks has resulted in bank damage, especially in the 
lower Kalang River and the Bellinger River 
downstream of Thora. Parts of these sections of the 
rivers and their tributaries have good bank habitat for 
platypuses, but other sections are of lower quality due 
to the occurrence of natural gravel bars, to the effects 
of past gravel extraction, and to earth banks being 
cleared and/or damaged by cattle. The upper reaches 
of both these rivers and the Never Never River provide 
good platypus bank habitat, although some cattle 
damage to banks in parts of the upper sections of the 
Bellinger River and lower Never Never River was 
present at the time of the survey. 

Riparian vegetation is generally continuous 
on both banks of the upper reaches of all the streams 
in the system but becomes less continuous in the lower 
reaches of most streams. The Bellinger River between 
the Never Never River junction and Bellingen was 
considered to be the most degraded section of the 
system. River oak Casuarina cunninghamiana was the 
main native riparian species found, while exotic species 
— willows Salix sp., camphor laurel Cinnamomum 
camphora, privet Ligustrum sp. and lantana Lantana 
camara — were widely distributed in the riparian zones 
of most streams at the time of the study. 


Field sampling 

Field sampling was carried out over 10 days 
during December 1996. The sections of the system 
surveyed by canoe, bank observation and netting are 
shown in Figure 1. The dates of platypus captured are 
given in Table 3. 

Canoe and bank observations were made 
either in the two hours prior to darkness and/or the 


Proc. Linn. Soc. N.S.W., 125, 2004 


two hours after dawn. Most of the Bellinger River from 
the mid-catchment gorge to Bellingen was surveyed 
by canoe either in the late afternoon or early morning 
(Figure 1). The Kalang River was unsuitable for canoe 
transects along much of its length due to its smaller 
size and the presence of obstacles in the channel. The 
section of river downstream from Duffys Bridge was 
suitable for use of the canoe and was surveyed a 
number of times. 

Live trapping of platypuses was carried out 
at four sites, one on the Never Never River, one on the 
upper Bellinger River and two on the Kalang River 
(Figure 1) using the methods of Grant and Carrick 
(1974). All captured females were injected with 0.2 
ml of synthetic oxytocin (Syntocinon) to indicate the 
presence of lactation (Grant and Griffiths 1992). 

The Kalang River was less intensively 
sampled than the Bellinger section of the catchment 
due to its unsuitability for canoe transects and difficulty 
of access for observation at sites which appeared to 
represent good platypus habitat. However, netting was 
carried out at one downstream and one upstream site 
on the Kalang River to assess the accuracy of reports 
obtained from local residents during the survey and to 
compare with community reports from this part of the 
river system. 


Habitat assessment 

At a number of accessible sites (mainly at 
road crossings) on the Bellinger (15), Never Never 
(8) and Kalang (15) rivers the following data or rank 
scores were collected to provide an assessment of 
habitat characteristics known to be associated with the 
occurrence of platypuses and their use of an area 
(Rohweder 1992; Bryant 1993; Ellem et al. 1998; Grant 
and Bishop 1998; Serena et al. 1998, 2001). This 
scoring procedure was based on both published and 


247 


PLATYPUS IN THE BELLINGER CATCHMENT 


unpublished field observations of platypus habitat: 


Habitat Category - this was a broad scoring of habitat 
suitability (1 best to 5 worst). Note: in the following 
categories, shade/shelter is usually provided by 
overhanging vegetation, but shade did not have to be 
present at time of observation as long as vegetation 
would provide shade/shelter at some times of the day. 
This is important not only to the platypus itself but to 
benthic invertebrate prey species: 
Category 1. EXCELLENT HABITAT - pools 
and/or riffle areas with >75% earth banks 
consolidated by roots of vegetation and 
providing significant shade/shelter, on both 
sides of river. 
Category 2. GOOD HABITAT - pools and/or 
riffle areas with 50-75% earth banks 
consolidated by roots of vegetation and 
providing significant shade/shelter, on at least 
one side of river or evenly distributed on both 
sides. 
Category 3. MODERATE HABITAT - pools 
and/or riffles with 25-50% earth bank 
consolidated by vegetation and providing a little 
shade/shelter. 
Category 4. POOR HABITAT - pools and/or 
riffles with 5-25% earth banks consolidated by 
roots of vegetation and providing little or no 
shade/shelter. 
Category 5. MARGINAL - pools and/or riffles 
with < 5% earth banks consolidated by roots 
of vegetation and providing no shade/shelter 


Riparian characteristics - these were features of banks 
that had been associated with platypus occurrence in 
other studies and were expressed as a percentage of 
sites at which they were present: 

- bank damage attributable to stock access; 

- bank damage attributable to floods; 

- presence of riparian vegetation; 

- presence of C. cunninghamiana (the most 

predominant native riparian tree species); 

- presence of introduced plant species in the 

riparian zone (especially willows, lantana, 

privet and camphor laurel). 


Community-based survey 

A community-based wildlife survey, in which 
the platypus was one of the target species, was posted 
to residents of the Bellinger-Kalang valley in 
December 1997. A total of 3000 survey forms was 
distributed by post to every household. There was a 
free-post return. The survey consisted of a 
questionnaire and colour maps on A3 size paper. The 


248 


first colour map was a user-friendly map of the area 
where respondents to the survey could mark on it the 
locations of fauna, including the platypus, they had 
seen in the area. A grid was included on this map so 
that grid references could be determined with ease. 
These locations were then transferred to the 
geographical information system, ArcView, for 
analysis. The survey form, including the maps of the 
catchment, appear in Figure 2A&B. 


Relationship to River Styles 

To investigate the possibility that analysis of 
River Styles may be a useful method of predicting 
platypus occurrence or relative abundance in sections 
of a river system, platypus records from the field and 
community surveys were allocated by one of the 
authors (TRG, who has Provisional River Styler 
accreditation) to the various River Styles identified in 
the Bellinger System by Cohen et al. (1998). Stream 
reaches representing various River Styles from Cohen 
et al. (1998: Figures 1A and 9) were transposed onto 
the relevant 1:25000 topographical maps and the 
distances calculated using a manual map measure 
(Uchida Curvimeter). 

As only two platypus records were obtained 
from the mountain headwater streams and upland 
stream River Styles, these stream categories and 
observations were not included in the analysis. 
Observations of platypuses in streams which were not 
classified by Cohen et al. (1998) or were in the tidal 
sections of the rivers (one observation only) were also 
not considered. 


Relationship to river disturbance 

As was carried out for the River Styles 
categories, conservation/rehabilitation sections from 
Cohen et al. (1998: Figures 1A and 9) were transposed 
onto the relevant 1:25000 topographical maps and the 
distances calculated using a manual map measure 
(Uchida Curvimeter). Platypus records from the field 
and community surveys were allocated to the various 
conservation/rehabilitation categories. 


Post-flood survey 

In September 2001 we contacted those 
community members who had reported platypuses in 
the wildlife survey conducted three years previously. 
A letter was individually addressed to each respondent 
and contained a covering note, a questionnaire to gather 
information on post-flood platypus sightings and a map 
showing the results of the community and field 
locations of platypuses on which each respondent could 
mark recent sightings. 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


No Postage stamp required 
if posted in Australia 


Bellinger Valley 
Wildlife Survey 


Please fold and return to: 


Reply Paid 100 


Bellinger Valley Wildlife Survey Agen XN 
c\- Dan Lunney > TO THE oY 
Biodiversity Survey and Research Division HOUSEHOLDER 
NSW National Parks and Wildlife Service 4 Oo 
PO Box 1967 OOAGE Ca 


HURSTVILLE NSW 2220 


Dear Shire Resident or Visitor, 


We are seeking your co-operation in conducting a wildlife survey of the Bellinger 
Valley. Its purpose is to locate wildlife populations as well as the habitats that 
are important for them. The long-term aim is to improve wildlife management of 
the valley by knowing which animals inhabit the area, where they occur, and the 
possible threats to their survival. This survey has the endorsement of Bellingen 
Shire Council and is supported by grants from the Heritage Assistance Program 
and the Foundation for National Parks and Wildlife. 


We would like you to fill out this survey even if you have only one wildlife 
sighting to record or you can only complete a part of the form. Also, if you have 
any historical information, this would help us understand the changes that have 
occurred to local wildlife populations over time in the Bellinger Valley. 


Please post your completed survey form (no stamp required) by 16 February 


1998. 


Thank you for taking the time to assist us in compiling this community-based 
survey. If you would like a souvenir copy of this form, please tick the box on 


page 4. 

Dan Lunney Alison Matthews Dionne Coburn 

(02) 9585 6489 (02) 9585 6559 (02) 9585 6558 
NSW 
NATIONAL 

New South Wales National Parks and Wildlife Service PARKS AND 
December 1997 WILDLIFE 

SERVICE 


Figure 2A. The Bellinger Wildlife Survey form. 


Proc. Linn. Soc. N.S.W., 125, 2004 249 


PLATYPUS IN THE BELLINGER CATCHMENT 


DORRIGO. 


Aas and Wikile Service and its erg 


; are 


Please show fusing a aces %) cn the map above those places where you bexe sam ay cf the wildlife listed belay. 
Peres wite the iitials gom met to axes to ddrrify th es. TE pasiile, plese deo pt te yer of the 
sicthing nee to tte imtials fc. =81 1963) Plesse also nerk the locatio cf where you Live or bolicey (atic) . 


Green Tree Fing W adyetailed Sule rc ailed fle BP 


Blus-btorqued Lizard 

Kastem Water Draom 

Carpet or Diarond Pythn CP 
loakeet a 


any axt done oF omission made on 


16 LANDSAT data supplied by Australian G 


This map Is not guaranteed to be {ree of error of omission, Therefore, the NSW Navional Parks and Wildlife Service and tes employees aitclal 


the mformation In the map and any consequences of such acts or omissions 


© NSW National Parks and Wildlife Service. t'rinted by GIS Division December 


© plzcss with a ood wmiay of wildlife © plaves vhere you see wildlife reqilarly 
© places wee jo te visiting frierk to se wildlife =e plas with Its of fog als 
© places Were you sae uneued wildlife 

Please draw the Lins using a dark pen oe pencil thet will not srunbe. 

Tt vould als help us if you give the reasons for your dices in the scace below: 


Blue-tongued Enzard 


Baxeacions © NPWS Dey P-Sherran No. 11970305 


250 Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


RESULTS 


Field survey 

Platypuses were observed or caught in all 
three rivers at a total of 36 locations (Figure 3). Two 
platypuses were captured at one of the two sites (the 
upstream site) on the Kalang River but none was 
observed in the limited sampling of this river by foot 
or by canoe. None was caught at the Never Never River 
netting site, but one was captured at the site on the 
upper Bellinger River. All the individuals captured 
were female, two of which were lactating, indicating 
the occurrence of breeding populations in both the 
Bellinger and Kalang rivers. All animals captured were 
within expected dimensions and body condition (Table 
3). Platypuses were found to be common and 
continuously distributed along the Bellinger and Never 
Never Rivers, being captured or observed at 35 sites 
(Figure 3). 

The canoe transect survey method was most 
successful, yielding 2.2 animals per hour of 
observation, compared with 0.17 for both netting and 


observations by foot from river banks (Table 4). 


Community reporting of the occurrence of 
platypuses 

A total of 522 replies (17.4% return) was 
received to the Bellinger valley wildlife survey. 
Platypuses were recorded at 123 sites by the 
community-based survey. Only two platypus records 
were obtained from the headwater streams of the 
catchment and the field survey did not sample these 
streams. These data showed a much lower number of 
sightings (13) in the Kalang River and its tributaries 
than in the Bellinger River (110) and its tributary 
streams. 

The field and community-based data showed 
that the platypus is commonly found throughout the 
Bellinger River catchment, including the Never Never 
and Rosewood Rivers, and its distribution is probably 
continuous above the tidal limit at Bellingen to the 
headwater streams, which were not surveyed in this 
study. There was one report of a platypus downstream 
of the tidal limit on the Bellinger River. As well as 
being reported less often along the Kalang River 


a a, 


“porrigo. - 


f/f River 
(J Bellingen LGA 
[3] National Parks 
State Forests 
4 Platypus field records 
* Platypus community records 
All community records 


Figure 3. The location of field and community-based records of platypuses in the Bellinger catchment. 


Proc. Linn. Soc. N.S.W., 125, 2004 


251 


PLATYPUS IN THE BELLINGER CATCHMENT 


Method Kalang River Bellinger River Never Never River Total 

Hrs No. CPU arts No. CPU Hrs No. CPU *ePU 
* 

Canoe EM By all. 0 95 29 3:05) 21>: 6 DID) 2.20 

observations | 

Bank 4.0 0 0 0.25 1 40 15 0 0 0.17 

observations 

Netting 85 2 0.23 45 1 O2Z20rS 0 0 0.17 

Total 1G 25ee2 OZ "14-25" Sil DIST S252 0.65 0.98 


*CPU: Catch/Observation per Unit Effort 


Canoe = individuals seen/hour observation in each observation period 

Observation = individuals seen/hour in each observation period 

Netting = individuals captured per net hour (1x50m net in water for 1 hour) 

Total = division of total individuals seen/caught by total hours of observation or net hours 


Table 4. Success of various field survey methods used for recording platypuses in December 1996. 


system, platypuses seemed to be more discontinuous 
in their distribution in this part of the river system 
(Figure 3). 


Reliability of the community-based data set 

The distribution of community-based reports 
of platypuses in the Kalang and Bellinger components 
of the river system showed considerable overlap with 
the field records (Figure 3). There were few 
observations by the community outside the areas in 
which the field work identified the occurrence of the 


% per Category 


E 
Yy 
/ 
] 
] 
j 


ZA 


species. One exception to this was the section of the 
Kalang River between Moodys Bridge and Sunny 
Corner, where no sightings or captures were made 
during the field work, but where platypuses were 
reported by the community. 


Habitat assessment 

There was little difference among the 15 sites 
sampled on the Bellinger and Kalang rivers in terms 
of habitat suitability (Figure 4), although on the Kalang 
River, 13% of the sites sampled were classified as 


@ Kalang 
0 Bellinger 
Never Never 


Bank Categories 


—— 


and Never Never Rivers. 


252 


Figure 4. Percentage of sites at which bank suitability categories were recorded on the Kalang, Bellinger © 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


120 
100 
£80 
2 i Kalang 
5 604 DO Bellinger 
ey Never Never 
rw «405 
20 
0 
Cattle Flood Riparian Casuarinas Introduced 
Damage Damage __- Vegetation. Plants 
Bank Characteristics 


Figure 5. Percentage of sites exhibiting various bank characteristics on the Kalang, Bellinger and Never 


Never Rivers. 


category 1, whereas none of the sites on either the 
Bellinger River or its major tributary, the Never Never 
River, fell into this category. 

All sites sampled had some riparian 
vegetation present but fewer sites on the Bellinger 
River had introduced species of riparian plants and 
more had C. cunninghamiana trees on the riverbank. 
More sites on the Kalang and Never Never Rivers than 
on the Bellinger River exhibited cattle and flood 
damage (Figure 5). 


Relationship to River Styles 

Table 5 details the lengths of each River Style, 
the total numbers of platypuses recorded in the field 
and community surveys and the numbers of platypus 
records per kilometre of each River Style. 

The River Styles of the Bellinger and Kalang 
sections of the catchment differ, with almost all (97%) 
of the Kalang River catchment being classified as 
confined bedrock with discontinuous alluvial 
floodplains, while the Bellinger River catchment 
contained a variety of Rivers Styles, ranging from 57% 
confined bedrock with discontinuous alluvial flood 
plains, through 24% alluvial with a meandering gravel 
bed to 10% and 9% of alluvial stream with a wandering 
gravel or discontinuous bed (Table 5). 


In the Bellinger River, there were 


Proc. Linn. Soc. N.S.W., 125, 2004 


significantly more platypus records in the alluvial 
meandering gravel bed sections of river and fewer in 
the discontinuous alluvial stream than expected if 
platypuses were distributed uniformly across River 
Styles (y7=26.64, 3d.f., P<0.01). In the Kalang River, 
platypus records were distributed evenly across River 
Styles. Expressed on the basis of platypus records per 
kilometre of river represented by each River Style, the 
Bellinger River had 0.95 records/km in the confined 
bedrock with discontinuous flood plains River Style, 
while the Kalang River (where this River Style made 
up 97% of the river downstream of the mountainous 
headwater reaches) had only 0.23 records/km (less than 
25% of the value for the Bellinger). In the only other 
River Style represented in the Kalang River, alluvial 
river with meandering gravel bed (3% of the river), 
there were no platypus records and yet this was the 
River Style on the Bellinger River which had most 
records (2.1/km). 


Relationship to river disturbance 

The lengths of each conservation and 
rehabilitation priority reaches proposed by Cohen et 
al. (1998), along with the total numbers of platypus 
records from the field and community-based data, as 
well as the number of records per kilometre for each 
priority category in the Bellinger and Kalang 


253 


PLATYPUS IN THE BELLINGER CATCHMENT 


Bellinger River 


Distance 
(km) 


River Style Platypus 


Confined bedrock 56 
with discontinuous 
floodplain 

Alluvial meandering 
gravel bed river 
Alluvial wandering 
gravel bed 
Discontinuous 
alluvial stream 
Total 


Kalang River 
Distance Platypus 
(km) 


Sh) 


Platypus 
per km 


Platypus 
per km 
0.23 


14 


Dell 0 0 
0 0 0 
0 0 0 


Table 5. River Style distances, numbers of platypus records and numbers of platypuses reported per 
kilometre of river in the Bellinger and Kalang rivers and their tributaries in December 1996. 


catchments, were compared (Table 6). The number of 
platypus records per kilometre of river in the Bellinger 
River increased from the “strategic” category (0.74/ 
km) through the “high recovery potential” (0.97/km) 
and “moderate recovery potential” (1.80/km) 
categories to be highest in the most “degraded” (2.25/ 
km) section of the river. There were significantly more 
platypus records than expected in the degraded and 
moderate recovery potential categories (y’=17.99, 
3d.f., p<0.01). In addition, the number of records of 
platypuses in the Bellinger River was much higher 
(1.22/km) than in the Kalang River (0.23/km), in spite 
of the fact that the latter system appears to be less 
disturbed than the Bellinger River. 


River Style Distance Platypus 
Conservation 
Strategic 

High recovery 
potential 
Moderate recovery 
potential 
Degraded 

Total 


Bellinger River 


Post-flood survey 

A total of 43 replies was received from 
respondents who had reported platypuses in the 1997 
survey. Twenty-one respondents had been near the 
river since the 2001 floods and of these, 7 had seen 
platypuses. Sightings of platypuses post-flood were 
in Hydes Creek (6 sightings), Boggy Creek (1 
sighting), the Never Never River (1 sighting), the 
Kalang River between Duffys and Moodys bridges (1 
sighting) and the upper Bellinger River between 
Diehappy and Bishops Creeks (2 sightings). 


Kalang River 
Platypus 


Distance 
(km) 


Platypus 
per km 


Platypus 
per km 


20.5 5 0.24 
139 4 0.29 
2ABS. 5 0.22 
Set 0 0 
14 0.23 


Table 6. Conservation/rehabilitation priority distances, numbers of platypus records and numbers of 
platypuses reported per kilometre of river in the Bellinger and Kalang rivers and their tributaries in 


December 1996. 


254 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


DISCUSSION 


Data from the field and community surveys 

The results strongly support the hypothesis 
that the community-based data from this study are 
reliable. The field observations and captures of 
platypuses closely corresponded to community 
records. As a result of this correspondence, the field 
and community data were combined in the analysis of 
platypus occurrence in relation to River Styles and 
conservation and rehabilitation priorities. Further, a 
post-flood survey was able to be conducted because 
of the reliability of community records. 

The lack of sightings from the headwater 
streams of the whole catchment suggest a lack of 
observation, rather than the species not occurring in 
them. This is supported by the low reporting of other 
wildlife species in the community wildlife survey in 
these areas (Figure 3). It is assumed that platypuses 
would almost certainly occupy these sections of the 
catchment streams, although this was not determined 
by the field survey. 

The field and community-based data rank the 
Bellinger River part of the system as being more 
suitable for occupation by platypuses than the Never 
Never and Kalang rivers. The limited habitat data 
collected during the survey point to the Kalang River 
having less suitable platypus habitat than either the 
Never Never or Bellinger Rivers. The discontinuous 
distribution of the platypus in the Kalang River, 
especially between Moodys Bridge and Rosewood 
Creek, almost certainly identified poorer habitat 
conditions. As community reports of other wildlife 
species were made along the Kalang River, the lack of 
platypus records from these sections means that the 
species is not found or is uncommon in these sections 
of the river. Further, any temporary loss of individuals 
from an area, such as from a flood, should not affect 
an accurate determination of distribution if the 
community had observed them in these sections of the 
river at other times. This is one of the values of the 
community survey, namely it was not restricted to one 
point in time and it also considered historical records. 
Qualitative field observations of the Kalang River 
between Moodys Bridge and Rosewood Creek 
confirmed that this section had poorer habitat quality 
than other sections of the river, with considerable 
disturbance of the river banks due to depletion of the 
riparian vegetation and cattle access, as well as 
accumulation of sand in the river bed. 

The field survey did not allow an adequate 
explanation to be made of the differences between the 
Bellinger and Kalang sections of the river system in 
terms of the observed platypus distribution. While parts 


Proc. Linn. Soc. N.S.W., 125, 2004 


of the Kalang were more degraded than sections of 
the Bellinger and Never Never rivers, platypus reports 
and field sightings were common in the Bellinger River 
between the Never Never River junction and the town 
of Bellingen, a river section which was also highly 
altered by bank clearing, stock damage to banks and 
past gravel extraction. 

One report was obtained of a platypus in the 
tidal section of the Bellinger River, close to the entrance 
of Connells Creek. Three reports of platypuses in the 
tidal section of the river were also made to the authors 
during the field study; one at Fernmount in the 1940s, 
another at the mouth of Hydes Creek in 1996 and one 
near the Old Butter Factory in Bellingen (which was 
said to be “recent’’). Platypuses have occasionally been 
found in the sea (Fleay 1980; Connolly and Obendorf 
1998) and in estuarine habitats, but such occurrences 
are irregularly reported and are considered unusual 
(Stone 1983; Grant 1991, 1999; Rohweder 1992; Hird 
1993; Menkhorst 1995; Connolly and Obendorf 1998; 
Rakick et al. 2001). It seems unlikely that the species 
regularly occupies the brackish or saline waters of 
estuarine environments. Nothing is known of its 
abilities to osmoregulate under marine or brackish 
conditions or any need by the species to have access 
to fresh water to groom salt from the fur, as occurs in 
several species of otters (Kruuk 1995). Platypuses are 
known to consume a range of benthic invertebrates as 
food but insect larvae are the most common prey items 
(Faragher et al. 1979; Grant 1982). In a number of 
rivers along the coast of New South Wales, tidal 
influences and/or saline intrusion into the lower reaches 
results in the diversity of benthic macroinvertebrates 
beginning to change at the tidal limit from being 
numerically dominated by insect fauna to being 
dominated by Crustacea, including amphipods and 
isopods, with oligochaetes worms and gastropod 
molluscs also having greater representation 
(Anon.1993; Simon Williams, then of Australian Water 
Technologies, pers. comm.). This could affect platypus 
distribution in the lower reaches of rivers of coastal 
New South Wales. It is also known that increased 
conductivity impairs the ability of the platypus to locate 
moving prey items, particularly small invertebrates, 
using the electrosensory mechanisms in its bill 
(Pettigrew et al. 1998). Competition with benthic- 
feeding fish species, which do not enter the freshwater 
sections of rivers, and possible predation by larger fish 
species, may also be involved in the occurrence of 
platypuses being unusual in tidal areas. 


Relationship to River Styles 


The differences in distribution and numbers 
of platypus records between the two rivers were not 


255 


PLATYPUS IN THE BELLINGER CATCHMENT 


found to be related to the differences in River Styles 
between the two parts of the system. On the basis of 
our findings in the Bellinger catchment we consider 
that analyses using the River Styles framework 
(Brierley et al. 2002) will not successfully predict the 
occurrence of platypuses. However, methods 
integrating geomorphic and biological considerations 
could lead to a framework which may be capable of 
predicting the suitability of streams for occupation by 
the platypus. Such integration could also provide a 
better basis for river management and rehabilitation 
than arises from the consideration of either geomorphic 
or biological considerations in isolation. This approach 
has been called the “landscape ecology approach” by 
Tockner et al. (2002). 


Relationship to river disturbance 
This study has shown platypuses to be present 


in degraded habitat of the Bellinger catchment. 
However, it would be a mistake to be complacent about 
these observations and regard disturbances of rivers 
to be benign with respect to the platypus. 

Despite the common occurrence of platypuses 
in agricultural areas, there are strong indications that 
platypus distribution has been fragmented and/or their 
numbers reduced in the streams of the Eden area 
(Lunney et al. 1998) and in the Bega (Brooks and 
Brierley 1997), Thredbo (Goldney 1998) and 
Richmond (Rohweder and Baverstock 1999) rivers of 
New South Wales and in the Wimmera River system 
in Victoria (Anon. 1999, 2000b, 2001b). In each of 
these instances the changes have been mainly attributed 
to the effects of agricultural practices. Lack of reports 
of platypuses from the Kalang River in the disturbed 
section between Moodys Bridge and Rosewood Creek 
also point to a fragmentation of platypus distribution 
within this part of the Bellinger catchment. 

Lunney et al. (1998) attributed fragmentation 
of platypus populations in the Eden region (Bega 
Valley Shire) of New South Wales to the effects of 
farming, particularly cattle grazing and the clearing 
of the riparian vegetation since 1830. Brooks and 
Brierley (1997) and Brierley et al. (1999) have detailed 
the effects of early agricultural practices in the Bega 
River valley of New South Wales, confirming that 
these practices were almost certainly responsible for 
the irreversible changes to that river system. However, 
Turnbull (1998) recorded the occurrence of platypuses 
in most of the rivers around Bombala, in the tableland 
headwater streams of the Bega and Snowy Rivers in 


256 


New South Wales, in spite of the area having been 
utilised for both cattle and sheep grazing for the past 
160 years. 

Of the 11 platypus sightings made by 
respondents to the post-flood questionnaire in the 
Bellinger catchment, 8 were in tributary streams. This 
suggests that the tributaries act as refuge areas during 
extreme floods. The tributaries could also be important 
for this population if the main streams of the Bellinger 
River system experience further degradation. This 
latter suggestion is based on our data from the Bega 
River (Lunney et al. 1998) where historically 
platypuses were found in the lower reaches, but it is 
now so degraded, shallow, sandy and exposed, that it 
no longer supports viable platypus populations. Instead 
platypuses occur only in the more protected and less 
developed tributary streams of the Bega River system. 


Conservation and rehabilitation 

Considering that the distribution of this 
unique Australian species overlaps extensively with 
activities of rural communities, its conservation 
depends on the adaptive management of these 
activities. The species has survived the current 
environmental disturbances so far, but its future 
conservation can only be assured by strategies aimed 
at preventing any further degradation of its habitat in 
these areas and by proactive rehabilitation of damaged 
sections of streams and a recognition of the possible 
importance of the tributary streams in retaining refuge 
populations of platypuses. 


ACKNOWLEDGEMENTS 


The authors are indebted to many people, 
particularly those who took the time to respond to the survey. 
We wish to acknowledge the contribution of P. Sherratt for 
the design of the community survey form; K. Weinman for 
production of the map and extra digitising of road and river 
systems that were not previously available on the National 
Parks and Wildlife Service geographic information system; 
and I. Dunn and G. Brierley for critical comments on the 
manuscript. Funding was provided by the Foundation for 
National Parks and Wildlife and a National Estate Grant. 
Platypuses were captured under licences issued by the NSW 
National Parks and Wildlife Service (A184) and NSW 
Fisheries (Scientific Research Permit F84/1245) and under 
Ethics Approval from the University of NSW Animal Care 
and Ethics Committee (Animal Research Authority ACE 94/ 
Wily); 


Proc. Linn. Soc. N.S.W., 125, 2004 


D. LUNNEY, T. GRANT AND A. MATTHEWS 


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(Hons) Thesis, University of New England 
Northern Rivers, Lismore). 

Rohweder, D.A. and Baverstock, P.R. (1999). Distribution 
of platypus, Ornithorhynchus anatinus, in the 
Richmond River Catchment, northern New 
South Wales. Australian Zoologist 31, 30-37. 

Serena, M. (1994). Use of time and space by platypus 
(Ornithorhynchus anatinus: Monotremata) 
along a Victorian stream. Journal of Zoology 
(London) 232, 117-131. 

Serena, M., Thomas, J.L., Williams, G.A. and Officer, 
R.C.E. (1998). Use of stream and river habitats 
by the platypus, Ornithorhynchus anatinus, in 
an urban fringe habitat. Australian Journal of 
Zoology 46, 267-282. 

Serena, M., Worley, M., Swinnerton, M. and Williams, 
G.A. (2001). Effect of food availability and 
habitat on the distribution of platypus 
(Ornithorhynchus anatinus) foraging activity. 
Australian Journal of Zoology 49, 263-277. 

Stone, G.C. (1983). ‘Distribution of the platypus, 
Ornithorhynchus anatinus, in Queensland’. 
(Queensland National Parks and Wildlife 
Service, Brisbane). 

Tockner, K. Ward, J.V., Edwards, P.J. and Kollman, J. 
(2002). Riverine landscapes: an introduction. 
Freshwater Biology 47, 497-501. 

Turnbull, R.W. (1998). Distribution of the platypus 
(Ornithorhynchus anatinus) in the Bombala 
River catchment, South-Eastern New South 
Wales. Australian Mammalogy 20, 251-256. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Reducing the By-catch of Platypuses (Ornithorhynchus 
anatinus) in Commercial and Recreational Fishing Gear in New 
South Wales 


T.R. Grant!, M.B. Lowry’, BRucE PEASE”, T.R. WALFORD? AND K. GRAHAM? 


'School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington NSW 
2052. email: t.grant@unsw.edu.au * NSW Fisheries P.O. Box 21 Cronulla, NSW 2230. 


Grant, T.R., Lowry, M.B., Pease, B., Walford, T.R. and Graham, K. (2004). Reducing the by-catch of 
platypuses (Ornithorhynchus anatinus) in commercial and recreational fishing gear in New South Wales. 
Proceedings of the Linnean Society of New South Wales 125, 259-272. 


The problem of platypus by-catch mortality in the eel, yabby and carp trap fisheries in New South 
Wales is reviewed, and the results of several experiments to determine the effectiveness of gear modifications 
to reduce platypus by-catch are presented. Entrance screens with 50-60 mm openings prevented the entry of 
platypuses into eel or yabby traps. Larger screens were not effective as a deterrent to platypuses entering 
traps. By-catch of platypuses in the eel fishery can be minimised by restricting traps to estuarme areas, 
where platypuses seldom occur, and by providing air spaces in the cod ends of traps used in impoundments 
and farm dams. Prohibiting the use of yabby traps in areas where platypuses are known to occur provides 
the most practical protection against by-catch of platypuses in this fishery. Platypuses were unable to exit 
from prototype carp traps, designed to permit escape of air-breathing species, but the provision of 
appropriately-sized openings at the base of the entrance funnels in these drum traps permitted platypuses to 


escape. 


Manuscript received 4 September 2003, accepted for publication 24 November 2003. 


KEYWORDS: by-catch, carp, eel, fishing, Ornithorhynchus anatinus, platypus, yabby. 


INTRODUCTION 


By-catch mortality of air-breathing 
vertebrates, including several species of freshwater 
turtles and diving birds, water rats (Hydromys 
chrysogaster) and platypuses (Ornithorhynchus 
anatinus), has been recognised for some time as a 
significant problem in various inland fisheries in 
Australia (Jackson 1979; Beumer et al. 1981; Grant 
1991, 1993; Grant and Denny 1991; Leadbitter 2001). 
Such by-catch mortality of platypuses is of particular 
concern in small streams, where multiple drownings 
of breeding individuals have the potential to impact 
severely on small local populations. For example, an 
abandoned fyke net in a tributary of the Gellibrand 
River in Victoria was found to contain the skeletons 
of 17 platypuses (Serena 2003). 

There has often been conflict between the 
desires of fishers to maximise catches of their target 
species, and the implementation of effective methods 
to reduce non-target by-catch. This has resulted in a 
diverse range of regulations enacted by fishery 
authorities and voluntary gear modifications by fishers 
aimed at reducing the mortality of non-target species 
(e.g. Leadbitter 2001). Unfortunately, little research 


or monitoring has been done to assess the effectiveness 
of voluntary and regulated gear modifications. 

An historical assessment of inland fishing in 
New South Wales showed that commercial fishing 
probably resulted in significant platypus mortality 
when small-mesh nets were used (Grant 1991, 1993; 
Grant and Denny 1991). No commercial or recreational 
fishery using nets or traps to capture native fish species 
or salmonids in freshwater sections of coastal rivers is 
now permitted in New South Wales (NSW), but there 
is a commercial eel fishery based on the use of baited 
traps in estuaries, farm dams and a few large 
impoundments. West of the Great Dividing Range, the 
commercial fishery for native fin-fish species was 
phased out in 2001. Fishers previously involved in that 
industry have been encouraged to fish for yabbies, 
mainly (Cherax destructor), using “Opera house” traps 
(Rankin 2000). The introduced carp (Cyprinus carpio) 
is also targeted by commercial fishers using a variety 
of gear, including traps, mesh and haul nets and 
electrofishing. 

There are a number of options to prevent or 
minimise mortality of air-breathing wildlife species 
in traps. The most direct way is to ban fishing in areas 
where these potentially vulnerable species occur. 


REDUCING BY-CATCH OF PLATYPUS 


UNREGULATED 
FISHERY 


PREVENT 
BY-CATCH 


REGULATE 
TO CLOSE 
WATERS 


TRAP 
MODIFICATION 
NECESSARY 


BY-CATCH 


REDUCTION 
DEVICE 
MINIMISES 
BY-CATCH 
MORTALITY 


MODIFY TRAP 
TO PERMIT 
NON-TARGET 
SPECIES TO 
ESCAPE 


MINIMISE 
BY-CATCH 


REGULATE 
FISHING 
METHODS 


STANDARD 
TRAP 
MINIMISES 


BY-CATCH 
MORTALITY 


BY-CATCH 
REDUCTION 
DEVICE NOT 

APPROPRIATE 
OR IS 
INEFFECTIVE 


MODIFY TRAP 
TO PROVIDE 
NON-TARGET 


SPECIES WITH 
AIRSPACE 


Figure 1. Schematic diagram of possible options available to achieve by-catch reduction of air-breathing 


species in fisheries operations. 


However, maintaining a commercial fishery, while still 
addressing the issue of by-catch mortality, is to adopt 
capture methods which minimise by-catch. Mortality 
of air-breathing non-target species can be reduced or 
prevented by trap modifications, such as fitting devices 


260 


to keep non-target species out (By-catch Reduction 
Device - BRD), providing a route to let them escape 
or permitting access to an airspace once they have 
entered a trap. Figure 1 summarises these possible 
options, which need to be explored in relation to the 
following issues: 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


Fishery requirements. The practicalities and economics 
of the fishery, in terms of trap design and cost, catch 
per unit effort, size of target species, and even the 
necessity to hide traps from possible interference and/ 
or vandalism must be considered. For instance, a device 
which reduces by-catch but unduly restricts the entry 
of the target species into a trap may be economically 
unviable. 

Behaviour of target species. It is necessary to know 
the reactions of the target species to trap modifications 
provided for non-target species. For example, the target 
species may escape via holes provided for the non- 
target species, or its behaviour could prevent the non- 
target species from utilising air spaces or escape routes 
provided. 

Behaviour of non-target species. In fishing areas where 
a number of potential by-catch species occur, escape 
holes, BRDs or air spaces in traps may not be suitable 
for all potential non-target species. For example one 
species may use an escape hole in a trap which will 
not be used by another species. 

This paper reviews past efforts to reduce the 
mortality of platypuses in the eel, yabby and carp 
fisheries and reports on a number of recent studies 
carried out to assess the effectiveness of trap 
modifications designed to reduce by-catch mortality 
of this species in these fisheries. The three fisheries 
are reviewed in separate sections of the paper and the 
experiments pertinent to each are discussed within 
these sections. 


THE EEL FISHERY IN NEW SOUTH WALES 


Freshwater eels were initially captured in 
upper estuarine waters of NSW as a by-catch of other 
fisheries. A fledgling industry targeting eels, based on 
the use of traps, was established in the early 1980s. At 
that time prices for eels were low but in the late 1980s 
and early 1990s a high-value export market to Asia 
was established. This increased interest in the fishery 
and the adoption of potentially more productive fishing 
methods. Requests were made by fishers to extend their 
operations into freshwaters using fyke nets (Figure 2a), 
which were known to be involved in the mortality of 
air-breathing wildlife species in the eel fisheries both 
in Tasmania and in Victoria (Jackson 1979; Beumer 
et al. 1981; Grant 1991). The potential fishers drew 
attention to a brief experiment in Lake Crescent and 
Dee Lagoon in Tasmania, where two fyke nets 
screened with 100 mm square mesh grids, and two 
unscreened control nets, were deployed in those lakes 
for six days. During that time, two platypuses were 
captured in the unscreened nets but none were captured 


Proc. Linn. Soc. N.S.W., 125, 2004 


in the ones with the screens in place (Grant 1991). 
While it appeared from this very limited experiment 
that a 100 mm mesh screen may have been effective 
in reducing platypus by-catch in Tasmania, an 
experiment done in the upper Shoalhaven River did 
not support this contention (Grant, unpublished data). 
Six platypuses (two female and four male) were placed 
separately between the river bank and the wing of a 
fyke net with a 100 mm mesh entrance screen in place. 
Two of these animals moved off after bumping the 
mesh and did not enter the fyke net but the other four 
either passed straight through into the net, or did so 
after first investigating the screen. 

At the time it was also known that elevating 
the cod end of fyke nets above the surface was effective 
in permitting platypuses to breathe and survive capture 
(Jackson 1979; Beumer et al. 1981; Grant 1991; Figure 
2b). Unfortunately professional fishers were 
unprepared to do this, as they feared their catch could 
be stolen and/or their equipment vandalised if it was 
visible above the surface. 

As a result of the brief experiment with the 
Shoalhaven River platypuses described above, and 
advice from experts in the other states regarding the 
poor compliance of fishers to fit BRDs and/or to raise 
the cod-ends of their nets above the water level, the 
request by fishers to use fyke nets for eels, and to extend 
the fishery to freshwater streams was denied by NSW 
Fisheries. Instead, the fishery was restricted to estuarine 
waters, a limited number of impoundments and private 
farm dams, using baited traps without wings to direct 
animals into the traps (NSW Fisheries Eel Policy 
Document, May 1992). 

The standard eel traps used in the fishery are 
shown in Figure 2c. They consist of a metal rod frame 
50 cm wide by 40 cm high by 90 cm long covered 
with 30 mm mesh polyethylene netting. The single 
entrance funnel (or ‘valve’) is located in one end of 
the trap. The opening in the funnel consists of a hole 
in the netting stretched firmly into a 100 mm wide 
slot, and pulled approximately 20 cm into the trap. 
The traps used in estuaries have a 1.5 m long cod end 
(bag with a draw-string) on the opposite end of the 
trap from the entrance funnel. Those used in freshwater 
impoundments and farm dams are similar to the estuary 
trap, but have a 5 m long cod end. A 150-200 mm 
diameter float is fastened inside the cod end near the 
draw-string and from one to three 50 cm diameter 
aluminium hoops are fastened to the inside of the cod 
end to keep the passage to the surface open. These 
traps are normally baited with frozen pilchards or 
mullet to attract eels. 

In the late 1990s anecdotal reports to the 
National Parks and Wildlife Service, NSW Fisheries 
and one of the authors (TRG) indicated that platypuses 


261 


REDUCING BY-CATCH OF PLATYPUS 


cod end 


entrance funnel 


_ Y 
: Ny 
=] il XX) 
= NYYXY) 
/ NN 
~ 


TR | 


Figure 2. (a) Fyke net used in eel fisheries in Tasmania and Victoria. (b) Commercial eel trap used in the 
impoundment or farm dam eel fishery, showing the elevated cod-end creating an air space. (c) Typical eel 
trap used in the tidal estuary fishery in New South Wales. Note the entrance funnel or ‘valve’ which 
permits animals to enter the traps in one direction (wide outside to narrow inside). Animals are unable to 
locate the narrow inside entrance to escape. In Experiment 1 grids were placed at the narrow end of the 
funnel and in Experiment 2 at the wide end. 


262 Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


were being drowned in eel traps, not only in the upper 
reaches of some estuaries (where tidal influence 
changed with river discharges) but also in farm dams 
and impoundments (where air spaces were not 
consistently being maintained in the cod ends of traps). 
As aresult, the following experiments were undertaken 
to determine if it was possible to reduce this mortality 
of platypuses by trap modification. 


EXPERIMENT 1 - Investigation of grid sizes for a 
platypus exclusion device 


The objective of this experiment was to 
determine the optimum grid size for excluding most 
platypuses from eel traps. The experiment was 
conducted in two pools on the Wingecarribee River in 
New South Wales from 17-19 February 2000. 


Methods 

The entrance funnels in eight standard eel 
traps were fitted with grids of different sizes. Each 
grid was a square divided into four equal openings; 
the openings in these grids ranged from 55 to 90 mm, 
in 5 mm increments. The plastic material used to make 
the grids was reinforced with lengths of 3 mm wire. 
The traps were fastened end to end (in order of 
decreasing grid size) and placed on a flat sandy area 
in the pools where platypuses were to be captured 
(Figure 3a). Water depth varied between traps but all 
had an airspace to allow the platypuses to breathe 
during the experiment. 

Trials were done in different pools on two 
days. Platypuses were captured using unweighted gill 
nets (Grant and Carrick 1974) during the evening or 
morning. Once the required numbers of platypuses 
were captured, each individual was measured and 
weighed, then tested individually in the experiment. 
Platypuses were placed through an access door into 
the first trap leading into an entrance funnel with the 
90 mm grid in place (Fig. 3a). Red-filtered lights were 
used to observe the animals at night, as observations 
in captivity indicated that platypuses are less 
responsive to disturbance under red light illumination 
(Grant, personal observation). The time that animals 
remained in each trap before passing through each grid 
was recorded, along with the number of attempts that 
each animal made to pass through the entrance funnel 
into the next trap in the series. Animals were removed 
from the experiment and released immediately if they 
remained in any trap for more than 15 minutes. 


Results 


A total of ten platypuses were used in the 
trials, comprising two adult males (1190 and 1760 g), 


Proc. Linn. Soc. N.S.W., 125, 2004 


six adult females (890-1060 g) and two juvenile 
females (700 and 760 g). Data are summarised in Table 
Ile 

Trial 1: Animals tested at night were reluctant 
to pass through the 85 mm grid and none passed 
through the 75 mm grid, while a single female captured 
in the morning, and tested in daylight readily, passed 
through all grid sizes, although exhibiting some delay 
at the 80 and 70 mm grids. However, it was noted that 
the traps with 85-70 mm grids, which were apparently 
difficult for the animals to negotiate, were located in 
slightly shallower water than the rest of the traps. The 
water level in these traps was located at or just above 
the top of the grid, whereas the water level in the other 
traps was well above the top of the entrance grids. It 
was thought that this difference in water depth may 
have influenced platypus behaviour. Subsequently, all 
traps were placed in deeper water (well over the top of 
the grid) during the second trial. 

Trial 2: The largest male (1760 g) could not 
pass through the 65 mm grid, but the smallest female 
(700 g) passed through each grid in less than | minute. 
The two slightly larger females did not initially pass 
through the 55 mm grid. However, it was found that, 
due to some unevenness on the bottom of the pool, the 
trap with this grid was in slightly shallower water than 
the preceding traps in the series. After moving this 
last trap to a position in slightly deeper water, animals 
passed through the 55 mm grid almost immediately. 

The data from Trial 1 indicated that there was 
a greater reluctance for platypuses to negotiate the grids 
when the traps were less submerged. However, Trial 
2 confirmed that female platypuses of up to 1 kilogram 
in weight could pass through a 55 mm grid. Animals 
smaller than 1 kg passed through easily, while the 1 
kg female had a tighter squeeze. Only one male 
platypus was captured for use in Trial 2. This was the 
largest animal tested (1760 g) and was stopped by the 
65 mm grid. A grid between 55 and 65 mm would 
apparently be required to exclude most adult male 
platypuses. 


EXPERIMENT 2 - Investigation of possible 
avoidance of entrance grids by free-swimming 
platypuses 


In Experiment 1, each platypus was closely 
confined inside the traps so there was an imperative to 
find an escape route. However, two of the four animals 
in Trial 2 hesitated, and made more than one attempt 
to pass through the 70 mm grid, indicating possible 
deterrent effect of this grid size. Experiment 2 was 
designed to test whether grids across the outer end of 
the entrance funnel (Figure 3b) deterred foraging 


263 


REDUCING BY-CATCH OF PLATYPUS 


entrance funnels 


AX) 
water level | NU 


netting 
enclosure 


ANN 
: 
a) 


Figure 3. Set up used in Experiments 1 and 2 to test the effectiveness of by-catch reduction devices (BRDs) 
on entry of platypuses into eel traps. (Top) Experiment 1. Traps were attached together in a line with 
grids of different sizes at the narrow end of each entrance funnel or ‘valve’. (Bottom) Experiment 2. 
Mesh enclosure in a river pool with trap entrance attached. Note the position of the replaceable rectangular 
grid across the outer (wide) entrance of the funnel. 


platypuses from entering traps. The experiment was 
done in a pool on the upper Shoalhaven River in the 
southern tablelands of New South Wales from 17-19 
March 2000. 


Methods 

A circular enclosure, 1.5 m high x 3 m 
diameter, made from 10 mm mesh monofilament gill 
net material, was constructed in a pool between the 
two netting sites where platypuses were captured for 
the experiment. The enclosure was designed so that 
the only possible escape for a platypus was through 
the grid of the entrance funnel of a trap inserted in the 
enclosure wall. Square grids, made from 4 mm steel 
rods, with 50, 60, 70 and 80 mm openings were used 
in this experiment. Each trial was done by attaching a 


264 


replaceable grid to the entrance funnel of the trap, then 
placing a platypus into the enclosure (Figure 3b). At 
night, red-filtered lights were used to observe the 
animals. The time each animal remained in the 
enclosure before passing through the grid was 
recorded, along with the number of attempts that each 
made to pass through the grid. If an animal did not 
pass through a particular grid in the test series, this 
was replaced by the next larger grid in the series and 
the observations repeated. After the first animal was 
obviously unable to exit the 50 mm grid, the trials on 
all others were begun with either the 60 or 70 mm 
grid. 


Results 
Eight relatively small platypuses (ranging in 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


Table 1. Details of platypuses exiting through the various grid sizes within the funnels of 
eel traps in the two trials of Experiment 1. + = animal exited specific grid size; 
X = platypus did not exit through specific grid size. 


Sex/ Weight 90mm 85mm 80mm 
age g) Grid Grid Grid 
Trial 1 

Male 

Adult 1190 + Xx xX 
Female 

Adult 1060 + + Xx 
Female 

Adult 1030 + xX x 
Female 

Adult 1020 + + + 
Female 

Adult 920 + + + 
Female 

Adult 890 + + + 
Exited 6/6 4/6 3/6 
Trial 2 

Male Adult 1760 + + + 
Female 

Adult 1000 + + + 
Female 

Juvenile 760 + + + 
Female 

Juvenile 700 + + + 
Exited 4/4 4/4 4/4 


size from 500 to 940 g) were tested in the enclosure at 
night. Results of the grid-deterrent trials are shown in 
Table 2. 

The first platypus was initially placed in the 
enclosure with the 50 mm grid. After six attempts to 
go through the grid it was apparent that the animal 
would not fit through the spaces. After several tentative 
attempts at the 60 mm grid it appeared to stop trying 
to escape through the subsequent grids and remained 
in the enclosure even after the largest grid was 
completely removed. The test with the second platypus 
was started with the 60 mm grid in place, but this 
platypus was less active than the first animal and made 
only one tentative attempt to pass through this grid. It 
then readily passed through the 70 mm grid after only 
one attempt. Trials with the next three platypuses were 
all started with the 60 mm grid. All three of these 
animals swam past the grid at least once before 
escaping through it. The last three animals were 
initially trialed with the 70 mm grid, and all passed 


Proc. Linn. Soc. N.S.W., 125, 2004 


75mm 70mm 65mm 60mm 55mm 


Grid Grid Grid Grid Grid 
xX x xX Xx xX 
x xX xX xX xX 
Xx X xX xX xX 
X X xX x X 
+ + + + + 
X x xX x x 
1/6 1/6 1/6 1/6 1/6 
+ + x x x 
+ + + + + 
+ + + + x 
+ + + + + 
4/4 4/4 3/4 3/4 2/4 


through it at the first attempt. Overall, two animals 
out of five appeared to be deterred by a 60 mm grid 
(40%) and only a single animal was deterred by a 70 
mm grid (Table 2). 


EXPERIMENT 3 - Platypus behaviour in the 
elevated cod ends of traps modified for use in farm 
dams and impoundments 


The objective of this experiment was to record 
the behaviour of platypuses in modified eel traps used 
in impoundments and farm dams (Figure 2c) and to 
investigate their ability to negotiate the long cod end 
extension to the air space. The experiment was done 
in a pool on the upper Shoalhaven River from 17-19 
March 2000. 


Method 


Two impoundment eel traps, with 5 m cod 
ends (Figure 2c) were placed in a pool of 0.5 m depth. 


265 


REDUCING BY-CATCH OF PLATYPUS 


Table 2. Details of platypuses deterred from entering the 
various grid sizes across the entrances of eel traps in 
Experiment 2. Animals are arranged in the order in which 
they were used in the experiment. + = animal passed through 
specific grid size; X = platypus did not pass through specific 


grid size i.e. deterred; - no data; 


Sex/Age Weight 50 mm 60 mm 70 mm 80 mm 
(g) Grid Grid Grid _ Grid 

Female 

Adult 800 xX x x x 

Female 

Juvenile 500 - x + - 

Male 

Juvenile 800 - + - - 

Male 

Juvenile 740 - + - - 

Male 

Juvenile 640 - + - - 

Female 

Adult 940 =- - + - 

Female 

Adult 900 —s- - + - 

Female 

Juvenile 690s; - + - 


One trap had three evenly spaced hoops in the cod end and the 
other had only one hoop near the airspace. The cod end of each 
trap was stretched and tied off above the surface of the water to 
a star-picket. Three platypuses (one male and two females) were 
placed consecutively in the trap with three hoops, and one 
female platypus was placed in the trap with one hoop. Each 
platypus was observed for 15 to 20 minutes before being 
released. 


Results 

In each case the platypus spent several minutes searching the 
inside of the trap before travelling up the cod end to the airspace. 
Each took several breaths then travelled to the trap where it 
again searched around or ‘wedged’ itself under the entrance 
funnel. Within five to eight minutes each would again travel 
up to the airspace for several breaths before returning to the 
trap. Platypuses travelled back and forth from the trap to the 
airspace 2-3 times during the 15-20 minutes they were confined 
in the trap. 


DISCUSSION - Eel Trap Experiments 


The results of Experiments 1 and 2 indicated that a 
grid of 50-55 mm would be necessary to exclude platypuses 
from entry into eel traps. Such a by-catch reduction device 
(BRD) would almost certainly affect the catch rates and sizes 
of eels (Koed and Dieperink 1999). This would be unacceptable 


266 


to commercial fishers, particularly those 
fishing for adults of the long-finned species 
(Anguilla reinhardtii). Free-ranging 
platypuses may be deterred from entering 
traps fitted with external grids of 70 mm or 
less across the entrance funnels but such 
screening would be unlikely to significantly 
reduce platypus by-catch in eel traps. 

Raising the cod end to provide an 
air space would facilitate the survival of 
platypuses captured in eel traps fitted with 
elongated cod ends. Platypuses captured in 
these traps were reluctant to stay at the 
surface and preferred to remain submerged 
in the trap between taking breaths. This 
behaviour, which minimises the time spent 
at the surface, may be a mechanism to avoid 
natural predation. Because platypuses must 
breathe at least every 2-10 minutes (Bethge 
2002), captured individuals would need to 
travel back and forth to the airspace many 
times during any extended period of 
confinement after capture. This would be 
stressful and energetically demanding. It is 
essential that captured animals be released 
as soon as possible after capture. Studies 
using fyke nets (with elevated cod ends) to 
capture fish have shown that platypuses can 
survive for periods of up to 24 hours (Grant 
and NSW Fisheries, unpublished data). 
However, hypothermia has been reported in 
platypuses restrained in fyke nets after a few 
hours in cold conditions (Serena, personal 
communication). The current regulations in 
New South Wales demand that eel traps be 
inspected at least every 24 hours. 

The platypus forages aerobically 
for short periods by holding its breath, 
following a comparatively large inspiration 
of air after each dive (Evans et al. 1994; 
Bethge 2002). The behaviour observed in 
this study of ‘wedging’ themselves under an 
object, and reducing energetic demands by 
remaining stationary, has been reported in 
captivity to last up to 11 minutes (Evans et 
al. 1994; Bethge et al. 2001; Bethge 2002). 
The function of this behaviour and its 
occurrence in the wild has not been 
determined. However, from the perspective 
of by-catch mortality this behaviour would 
not prevent platypuses from being drowned 
in completely submerged traps during 
normal fishing operations, which demand a 
period of trap submergence of hours rather’ 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


than minutes. 

Observation of traps with airspaces 
maintained only by the use of a float has shown that 
the cod end can easily become twisted or bunched. 
This situation would undoubtedly prevent a captured 
air-breathing species from reaching the airspace. This 
can be avoided by stretching the cod end tightly to a 
fixed point, either on the bank or a star picket driven 
into the bottom of the water body. It should be noted 
however, where traps are set with elevated cod ends 
attached to a fixed point, allowance needs to be made 
for anticipated rises in water level as a result of rainfall 
and/or tidal influences. Attachment of the cod ends of 
eel traps to a fixed point is mandatory under regulations 
for the use of eel traps in impoundments and farm dams 
in NSW. 


THE COMMERCIAL AND RECREATIONAL 
YABBY FISHERY IN NEW SOUTH WALES 


The results of the experiments done to 
evaluate the effectiveness of devices to prevent or deter 
platypuses from entering eel traps are also directly 
applicable to both the commercial and recreational 
“‘yabby’ [freshwater crayfish] fisheries. Based on the 
lack of adverse reports and on the assumption that the 
traps used to capture yabbies were small and did not 
have mesh wings to direct foraging platypuses into 
them, Grant (1993) suggested that “yabby fishing poses 
little threat to platypuses”. This conclusion is now 
thought to be incorrect, as anecdotal reports from a 
number of states suggest that yabby traps were 
affecting some local platypus populations. These traps 
have also been implicated in the mortality of other non- 
target species, especially freshwater turtles. The 
drowning of as many as five platypuses in a single 
yabby trap has been reported, although the species’ 
attraction to these traps is not fully understood. 
Platypuses are known to locate their prey by sensing 
the electrical fields generated by muscular activity of 
the prey species, especially large food items such as 
yabbies (Pettigrew et al. 1998). A trap containing live 
yabbies may therefore attract platypuses during their 
normal foraging activities. Once there is a dead 
platypus in a trap, more yabbies may feed on the 
decomposing carcass, which could in turn attract other 
platypuses into the trap. 

Rankin (2000) suggested that a fixed ring 60- 
70 mm in diameter may prevent platypuses from 
entering traps and also facilitate their escape. Some 
commercially available yabby traps are fitted with 90 
mm entrance rings, which are effective in excluding 
larger turtles but which are still reported to have 


Proc. Linn. Soc. N.S.W., 125, 2004 


drowned platypuses. The experiments described above 
for eel traps indicate that a 90 mm diameter ring is too 
large to exclude platypuses. Similarly, neither the 
experiments reported here nor anecdotal observations 
support Rankin’s (2000) suggestion that platypuses 
could escape by returning through a fixed entrance 
ring. 

Allanson and Thurstan (1999) evaluated the 
effect of entrance rings of different diameters in yabby 
traps using relatively small captive-bred yabbies 
(Cherax destructor). These trials showed that the 
smallest ring tested (63 mm) still permitted yabbies of 
the same size to enter the experimental traps as were 
entering the control traps with no rings fitted. However, 
the experimental traps caught substantially fewer 
yabbies. When the results of Allanson and Thurston’s 
(1999) experiments were discussed with commercial 
fishers, it was concluded that the use of such a small 
entrance ring was not a viable option for the 
commercial yabby fishery. 

Current regulations in New South Wales 
exclude the use of traps in commercial and recreational 
yabby fishing from known platypus waters and 90 mm 
rings are required in all yabby traps to exclude most 
turtles. Closed waters are located east of the Newell 
Highway, from the Victorian border (Murray River) 
to the Queensland border (Macintyre River), along with 
local closures around Deniliquin on the Edward River, 
Echuca on the Murray River and between Narrandera 
and Darlington Point on the Murrumbidgee River, 
where platypuses are also know to occur. 


THE CARP FISHERY IN NEW SOUTH WALES 


Carp (Cyprinus carpio) were probably first 
introduced into Australia around 1850 but did not 
spread until the introduction of the “Boolarra’ strain 
in the 1960s. Ecological effects of high densities of 
carp are poorly understood, but increased bank 
damage, disturbance of aquatic macrophytes and 
turbidity are all possible consequences. The overall 
disruption of riverine food webs by the large biomass 
of carp is thought to be detrimental to freshwater 
ecosystems (Schiller and Harris 2001). Carp are 
harvested in New South Wales using a variety of gear, 
including traps, haul and mesh nets, and electrofishing 
equipment. There is considerable overlap between the 
distribution of carp and platypuses (Boulton and Brock 
1999), making the use of submerged traps a concern 
in this fishery. 

A drum trap was constructed by NSW 
Fisheries (Fig. 4), which was designed to permit the 
escape of air-breathing vertebrate species, including 


267 


REDUCING BY-CATCH OF PLATYPUS 


wy 
his WY 
i 
; 


hed 


platform 


platypus escape 
holes ~ 


entrance funnel 


escape hole 


platform 


entrance funnel 


Figure 4. (Top) Modified drum trap showing escape hole in the roof 
above the mesh platform. Note the entrance funnel (or ‘valve’) on the 
left end of the drum. The entrance was sealed in the experiments and 
the triangular escape holes were made at the base of this funnel. 
(Bottom) Inside the trap showing the position of the steel mesh platform 
below the escape opening. 


vertebrate species could pass 
through the 8 cm gap between it 
and the roof of the trap and exit 
through the escape hole, while 
larger carp would not be able to 
escape. Carp are also inclined to 
congregate near the bottom of a 
trap. The design assumed that air- 
breathing species would tend to 
swim towards the surface and 
search along the roof of the trap 
for a means of escape (surface/ 
search behaviour). The objective 
of the following experiment was 
to test the effectiveness of the 
escape device for platypuses. 


EXPERIMENT 4 - Assessment 
of escape of platypuses from a 
prototype carp trap 


Platypuses close their 
eyes, ears and nostrils when under 
water, using the sensory 
mechanisms in their bills to find 
their way around (Pettigrew et al. 
1998). It was expected that 
platypuses in the experiment 
would exhibit surface/search 
behaviour and be able to escape 
from the modified drum trap. The 
experiment was done in several 
pools on the Wingecarribee 
River, New South Wales from 25- 
27 November 2002 to determine 
if this expectation was realised. 


Method 

The trap consisted of a 
90 cm diameter x 170 cm long 
cylinder, covered with black 
plastic mesh (55 mm x 40 mm), 
except at the entrance end, where 
a conical funnel or ‘valve’ made 
from 3 mm diameter braided 
polyethylene trawl netting was 
strung tightly between the 
circular steel frame at one end of 
the trap and an oval ring rigidly 
suspended inside the trap (Figure 
4). 

The trap was fully 


platypuses, water rats, turtles and diving birds, through —_ submerged in the pools from which the platypuses were 
a hole in the trap’s roof. A wire-mesh platform was captured. The trap was oriented with the escape hole’ 
positioned below the escape hole so that small uppermost. A remote lens for a video camera was 


268 Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


Table 3. Results of Experiment 4. Assessment of escape of platypuses from the carp trap in the 


Wingecarribee River. 


Sex/Age Weight (g) Length 
(cm) 

Female 

Adult 1080 48.5 

Male 

Adult 1880 So 

Male 

Juvenile 1790 56.5 

Male 

Adult 1880 57.5 

Male 

Juvenile 1400 53.0 


mounted inside the trap to record the behaviour of the 
animals and these images were stored for later analysis. 

Platypuses were captured using unweighted 
gill nets (Grant and Carrick 1974). Each animal was 
weighed and measured, then temporarily marked with 
a piece of brightly coloured tape attached to the tail, 
making the platypuses more visible to observers and 
to the video camera. Based on observations reported 
above and Bethge (2002), who reported a maximum 
foraging dive duration of 138 seconds, individuals 
were immersed for a maximum of 3 minutes before 
the trap was lifted to permit them to breathe. If they 
exited the trap prior to lifting, the elapsed time was 
recorded. The numbers of times each animal 
approached the platform below the escape hole was 
recorded. All animals were used only once in the 
experiment and remained in the trap for no more than 
3 minutes. 


Results 

Table 3 shows the dimensions of the 
platypuses used, the time in the trap, the number of 
approaches to the platform below the escape hole, and 
whether or not individuals escaped. Only one juvenile 
male platypus managed to find the escape hole (after 
30 seconds in the trap), but showed reluctance to leave 
the steel ring around the hole. It re-entered the body 
of the trap three more times before finally leaving the 
trap completely. This animal repeatedly relocated the 
escape hole after re-entering the trap, taking 30, 50 
and 50 seconds respectively, before finally escaping. 
The other four trial animals failed to find the escape 
hole and were released after 2-3 minutes. 

Contrary to expectation, platypuses 
(including the one which escaped) spent most of the 
time investigating the bottom or ends of the trap, rather 
than exhibiting surface/search behaviour. In fact, they 
seemed to actively avoid the platform area below the 


Proc. Linn. Soc. N.S.W., 125, 2004 


Time in Approaches __ Escape 
trap (sec) to platform 

150 0 No 
165 3 No 
180 0 No 
180 0 No 

30 0 Yes 


escape hole. All animals searched with their bills 
around the corners of the trap between the sides and 
ends. The video showed them frequently investigating 
the acute angled edge between the base of the entrance 
funnel and the sides of the trap. When released, all 
animals were observed to surface and appeared to be 
breathing heavily. 


EXPERIMENT 5. Assessment of escape of 
platypuses from the modified carp traps 


In Experiment 4, platypuses were observed 
continually searching for an escape hole around the 
corners of the trap. It was therefore decided to test the 
effectiveness of escape holes positioned around the 
base of the entrance funnel. Because the gap between 
the funnel and the sides of the trap was quite narrow at 
the base of the funnel, it was considered that most carp 
would be too large to access openings in this position. 
Experiment 5 tested the effectiveness of these 
modifications. The experiment was done in one pool 
on the Wingecarribee River on 27 November 2002 and 
then in four pools on the upper Shoalhaven River from 
21-23 December 2002. 


Methods 

Every third mesh attached to the trap frame 
at the base of the funnel was released and tied back to 
provide 90 x 90 x 90 mm triangular openings (Fig. 4, 
top). In the initial trial in the Wingecarribee River these 
openings were made only in the upper half of the trap, 
but in the later trials in the upper Shoalhaven River, 
openings were made in both the upper and lower halves 
of the trap. 

Fourteen platypuses were individually placed 
in the submerged trap as described in Experiment 4. 
Again observations were made of the number of times 
animals approached the platform below the escape 


269 


REDUCING BY-CATCH OF PLATYPUS 


Table 4. Results of Experiment 5. Assessment of escape of platypuses from the modified carp trap. * not 
observed escaping but were not present in trap when it was lifted after 3 minutes; - escape holes only 


available in upper part of this trap. 


Sex Weight Length Time in 
(g) (cm) trap (sec) 
Female 850 43.0 85 
Female 690 41.0 15 
Female 900 46.0 22 
Female 940 43.5 15 
Female 900 44.0 40 
Female 790 41.0 41 
Female 870 43.5 140 
Female 930 44.0 33 
Female 860 44.0 <180* 
Female 840 43.5 45 
Female 790 43.0 156 
Male 1850 55.2 35 
Male 1740 52.0 <180* 


hole. Escapes through the triangular holes at the base 
of the entrance funnel were partitioned as being from 
the ‘upper’ or ‘lower’ openings in the trap. Some 
underwater video observations were made but the 
turbidity of the pools made viewing difficult. However, 
brightly coloured tape attached to the tails of the 
animals (see Experiment 4) usually permitted their 
movements in the trap to be observed. Again, if the 
platypus was not seen to escape, the trap was lifted 
from the water after a maximum of 3 minutes. 


Results 

Thirteen platypuses escaped from the 
openings around the base of the entrance funnel of the 
trap within 3 minutes (Table 4). As was observed in 
Experiment 4, all animals attempted to find an escape 
route around the bottom or ends of the trap. Another 
individual used in the initial trial located a hole 
inadvertently left at the bottom of the trap (which was 
sealed before subsequent trials). No preference was 
shown for escape location, with six animals exiting 
from the ‘upper’ and 5 from the ‘lower’ openings, 
where both were available (Table 4). One individual 
moved into the space between the platform and the 
escape hole but did not find the hole, submerging again 
and leaving the trap by one of the openings at the base 
of the entrance funnel. Only two individuals 
approached the platform at any time during their 
confinement in the trap. In two instances the platypuses 
could not be seen, but were no longer in the traps when 
they were lifted after 3 minutes. It was presumed that 
they had exited the lower holes, as they were not seen 
leaving the upper ones, which were visible to the 
observers. 


270 


Approaches Escape Escape 
to platform location 
] Yes - 

0 Yes lower 
0 Yes lower 
0 Yes lower 
0 Yes upper 
0 Yes upper 
0 Yes upper 
0 Yes upper 
0 Yes lower 
0 Yes upper 
0 Yes upper 

1 Yes - 

0 Yes lower 


DISCUSSION - Carp trap experiments 


Experiment 4 indicated that the unmodified 
carp trap would probably result in significant mortality 
of platypuses if deployed in areas where their 
distribution overlaps that of carp. However, experiment 
5 indicated that carp traps with appropriate escape holes 
could be used to reduce by-catch of platypuses. 
Platypuses over a size range of 690-1880 grams were 
able to exit quite quickly (15-156 seconds) through 
the 90 mm triangular openings in the modified carp 
trap. 

It should be noted that the platypuses used in 
these experiments were not particularly large. There 
is considerable sexual dimorphism in the species, with 
the average male being around 75% heavier and 20% 
longer than females (Carrick 1995; Grant 1995; 
Connolly and Obendorf 1998). Individuals of up to 
twice the size of those used in current experiments are 
found in some mainland areas (especially west of the 
Great Dividing Ranges; Carrick 1995; Grant 1995) and 
in Tasmania males may reach up to three kg (Connolly 
and Obendorf 1998). Further experiments are required 
to determine the size of escape holes effective for larger 
platypuses. In the interim, the authors recommend 
triangular openings of 100 x 100 mm for east-flowing 
streams in New South Wales and openings of at least 
120 x 120 mm for west-flowing streams in the state. 
Trials would also need to be carried out to assess the 
effectiveness of retaining captured carp in the presence 
larger escape holes. 

The unexpected lack of surface/search 
behaviour in platypuses during Experiments 4 and 5 


Proc. Linn. Soc. N.S.W., 125, 2004 


T.R. GRANT, M.B. LOWRY, B. PEASE, T.R. WALFORD AND K. GRAHAM 


indicates the importance of field trials of fishing 
equipment with regard to specific wildlife species. The 
reason for the unexpected lack of surface/search 
behaviour in water can only be speculated upon. 
Platypuses frequently forage among dense woody 
debris and under submerged overhanging banks (Grant 
1995 and personal observation). It may be that a 
behavioural response of moving down and/or sideways 
away from an obstruction during foraging may be of 
greater survival value than attempting to rise directly 
to the surface when seeking an escape route. No 
‘wedging’ behaviour (Evans et al. 1994; Bethge et al. 
2001; Bethge 2002; Experiment 3) was exhibited by 
animals in the carp traps. Rather, all individuals 
searched constantly for an escape route. 


GENERAL CONCLUSIONS 


The results of the literature reviewed and 
experiments presented in this paper indicate that any 
fishery in freshwaters of New South Wales based on 
the use of traps should not be operated as an 
unregulated fishery (Figure 1) if reducing platypus 
mortality is a priority. By-catch minimisation has been 
possible in the eel fishery by a combination of closures 
of some inland waters and by modifications to provide 
an airspace in traps used in farm dams and 
impoundments. Exclusion devices (e.g. grids across 
the entrance funnels of traps) do not provide a 
commercially viable option for reducing the by-catch 
of platypuses in eel or yabby traps. Banning of yabby 
traps from areas where platypuses occur is currently 
the only available means of avoiding by-catch 
mortality in this fishery. The commercial and 
recreational yabby fisheries in New South Wales are 
currently restricted to waters where platypuses do not 
commonly occur or are very uncommonly reported. 
Trap modifications, which permit the escape of 
platypuses, appear to be the most feasible means of 
by-catch minimisation in the use of traps to capture 


carp. 


ACKNOWLEDGMENTS 


This work was conducted under Animal Research 
Authorities (ACEC 99/13 and ACEC 02/12) from the NSW 
Fisheries Animal Care and Ethics Committee and under NSW 
Fisheries Scientific Research Permit F84/1245 (TRG) and 
NSW National Parks and Wildlife Service Scientific 
Investigation Licence A184 (TRG). Frank Jordan constructed 
the grids used in Experiment 2 and Stuart Scott and the Izzard 
and Laurie families allowed us to work on their properties 
on the Wingecarribee and Shoalhaven Rivers. Dr Melody 


Proc. Linn. Soc. N.S.W., 125, 2004 


Serena, of the Australian Platypus Conservancy is 
acknowledged for her personal communication regarding 
hypothermia in platypuses captured in fyke nets. Funding 
for much of the reported work was provided by NSW 
Fisheries. The authors thank Mike Augee and an anonymous 
referee for their valuable comments and suggestions. 


REFERENCES 


Allanson, M. and Thurstan, S. (1999). “Yabby (Cherax 
destructor) trap by-catch reduction.’ 
Unpublished report NSW Fisheries-Inland 
Management. August/September, 1999. 

Bethge, P. (2002). Energetics and foraging behaviour of 
the platypus Ornithorhynchus anatinus. PhD 
thesis. University of Tasmania, Hobart. 

Bethge, P., Munks, S. and Nicol, S. (2001). Energetics 
and locomotion in the platypus, 
Ornithorhynchus anatinus. Journal of 
Comparative Physiology - B, Biochemical, 
Systematic and Environmental Physiology 171, 
497-506. 

Beumer, J.P., Burbury, M.E. and Harrington, D.J. (1981). 
The capture of fauna other than fishes in eel and 
mesh nets. Australian Wildlife Research 8, 673- 
677 

Boulton, A.J. and Brock, M.A. (1999). “Australian 
Freshwater Ecology. Processes and 
Management’. Gleneagles Publishing, Glen 
Osmond, South Australia. 

Carrick, F.N. (1995). Family Ornithorhynchidae. 
Platypus. In “The mammals of Australia’ (Ed R. 
Strahan) pp. 35-38. (Reed Books: Chatswood). 

Connolly, J.H. and Obendorf, D.L. (1998). Distribution, 
captures and physical dimensions of the 
platypus (Ornithorhynchus anatinus) in 
Tasmania. Australian Mammalogy 20, 231-237. 

Evans, B.K., Jones, D.R., Baldwin, J. and Gabbott, G.R.J. 
(1994). Diving ability of the platypus. 
Australian Journal of Zoology 42, 17-27. 

Grant, T.R. (1991). The biology and management of the 
platypus (Ornithorhynchus anatinus) in New 
South Wales. Species Management Report # 5. 
NSW National Parks and Wildlife Service, 
Hurstville, New South Wales 

Grant, T.R. (1993). The past and present freshwater 
fishery in New South Wales and the distribution 
and status of the platypus, Ornithorhynchus 
anatinus. Australian Zoologist 29, 105-113. 

Grant, T. (1995). “The platypus. Unique mammal’. 
University of New South Wales Press. Sydney. 

Grant, T.R. and Carrick, F.N. (1974). Capture and 
marking of the platypus, Ornithorhynchus 
anatinus, in the wild. Australian Zoologist. 18: 
133-135. 

Grant, T.R. and Denny, M.J.S. (1991). Distribution of the 
platypus in Australia with guidelines for 
management. Report to Australian National 


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Parks and Wildlife Service. Mount King 
Ecological Surveys, Oberon, New South Wales. 

Jackson, P.D. (1979). Survey of fishes in the west branch 
of the Tarwin River above Berrys Creek. 
Victorian Naturalist 97, 11-14 

Koed, A. and Dieperink C. (1999). Otter guards in river 
fyke-net fisheries: effects on catches of eels and 
salmonids. Fisheries Management and Research. 
6: 63-69. 

Leadbitter, D. (2001). Bycatch Solutions. A Handbook for 
Fishers in Non-Trawl Fisheries. Oceanwatch 
and Fisheries Research and Development 
Corporation. Pyrmont, New South Wales, 
Australia. FRDC Project NO. 98201. 

Pettigrew, J.D., Manger, P.R., Fine, S.L.R. (1998). The 
sensory world of the platypus. Transactions of 
the Royal Society of London. Series B. 353, 
1199-1210. 

Rankin, T. (2000). Status of the New South Wales 
commercial yabby fishery. NSW Fisheries 
Fishery Resource Assessment Series No. 9. 
Fisheries Research Institute, Cronulla. May 
2000. 

Schiller, C.B., Harris, J.H. (2001). Native and alien fish. 
In ‘Rivers as Ecological Systems: the Murray- 
Darling Basin’ (Ed W.J. Young) pp. 229-258. 
(Murray-Darling Basin Commission: Canberra). 

Serena, M. (2003). Wanted. Platypus sightings. Australian 
Society for Limnology Newsletter March 41, 36. 


272, Proc. Linn. Soc. N.S.W., 125, 2004 


Platypus Burrow Temperatures at a Subalpine Tasmanian Lake 


Pattie BeTHGe!, SARAH Munks’, HELEN OTLEY’, STEWART NICOL! 


' Division of Anatomy and Physiology, University of Tasmania, GPO Box 252-24, Hobart TAS 7001, 
Australia (Address for correspondence: Brandstwiete 19, 20457 Hamburg, Germany; philip @bethge.org); 
* School of Zoology, University of Tasmania, Private Bag 4,GPO, Hobart TAS 7001, Australia 


Bethge, P., Munks, S., Otley, H. and Nicol, S. (2004). Platypus burrow temperatures at a subalpine 
Tasmanian lake. Proceedings of the Linnean Society of New South Wales 125, 273-276. 


When platypuses are in their burrows, microhabitat is of great importance for energy conservation, especially 
where air temperatures frequently fall below freezing in winter. In this study, we investigated burrow 
temperatures of platypuses (Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Nine individual 
platypuses were equipped with time-depth recorders with integrated temperature sensors measuring ambient 
temperature. Burrow temperatures were recorded in two minute intervals for a total of 61 resting periods 
(duration: 5.45 to 27.20 hours) and were averaged over the period of resting. Mean burrow temperatures 
were 17.5 and 14.2°C (SD=2.76 and 0.89, respectively, n=9) in summer and winter, respectively, and 
ranged between 12.2 and 22.8°C for individual resting periods. In winter, burrow temperatures were held 
fairly constant over the resting period while in summer larger variations were observed. Burrow temperature 
in winter was found to be up to 18°C higher than outside air temperature. 


Manuscript received 26 November 2003, accepted for publication 8 January 2004. 


Key words: Burrow temperature, Energetics, Ornithorhynchus anatinus, Platypus, Tasmania 


INTRODUCTION 


The platypus, Ornithorhynchus anatinus, 
inhabits the lakes, rivers and streams of eastern 
Australia from the Cooktown area in the north to 
Tasmania in the south. Over much of its range, the 
animal is found in alpine and tableland areas where, 
especially in winter, air temperatues fall well below 
freezing and water temperatures approach O0°C (Grant 
1995). Grant (1983a) suggested that under such 
conditions, the microhabitat of platypus burrows is of 
great importance for energy conservation. Even in an 
unoccupied artificial burrow the insulation of layers 
of earth was found to provide significant buffering 
effect against outside ambient temperature changes 
both in winter and in summer (Grant 1976). (Grant 
1983b) suggested a further modifying effect of the 
animal’s presence on the microhabitat temperature, 
elevating it several degrees above that of an unoccupied 
burrow. 

In this study ambient temperatures in 
occupied platypus burrows at a sub-alpine Tasmanian 
lake were investigated. The use of time-depth recorders 
with integrated temperature sensors made it possible 
to determine burrow temperatures during naturally 
occuring resting periods of the equipped animals. 


MATERIALS AND METHODS 


Field experiments were carried out at Lake 
Lea (41°30° S, 146°50° E), a sub-alpine lake in 
northwestern Tasmania. Information on burrow 
temperatures was obtained from nine individual 
platypuses (4 adult males, mass: 2.27 kg + 0.26 (SD), 
5 adult females, mass 1.48 kg + 0.07 (SD)) between 
November 1998 and January 2000. Platypuses were 
captured and processed following the methods outlined 
in Otley et al. (2000) and Bethge et al. (2003). 
Individuals were equipped with combined data logger- 
transmitter packages (max 62 mm x 28 mm x 18 mm, 
weight 50 g, Fig. 1) consisting of a specially designed 
standard transmitter (Faunatech, Eltham, Victoria) and 
a time-depth recorder (LTD 10, Lotek Inc., Canada). 
The packages were attached with glue (5 min-Araldite, 
Selleys Inc., Australia) to the guard fur of the lower 
back of the animals, just above the tail, following the 
method outlined in Serena (1994). Animals were 
released at the site of capture. After approximately two 
weeks the animals were relocated by radiotracking and 
recaptured on emergence and the devices were 
removed. 

The data loggers allowed measurement of 
ambient temperature in the range from 2 to 25°C with 


PLATYPUS BURROW TEMPERATURES 


20 


(tart Po 


10 


Temperature [°C] 


-5 


Tw 


Ta 


15/6 16/6 17/6 18/6 19/6 20/6 21/6 22/6 23/6 24/6 25/6 26/6 27/6 


Figure 1: Winter sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a 
time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature 
is only shown at times when the animal was in the burrow). 


an accuracy of 0.06°C. The devices were calibrated 
by the manufacturer (Equipment for temperature- 
calibration: Neslab RTE-2000 Bath/Circulator and 
Omega HH40 Thermistor/Thermometer). Temperature 
sensors were located at the back end of the devices 
and were facing backwards when the devices were 
fixed on the platypus’s lower backs. Ambient 
temperature was measured in two-minute intervals for 
11 days each. While foraging, the sensors measured 
water temperatures. In resting platypuses, ambient 
temperatures close to the animals’ bodies (approx. 5 
mm from above the fur) were recorded. The resting 
period was defined as the time span between the end 
of the last dive of a foraging trip (detected by the depth 
sensor of the time-depth recorders) and the beginning 
of the first dive of the following foraging trip. Burrow 
temperatures, i.e. ambient temperatures during resting 
periods, were recorded in two minute intervals for a 
total of 61 resting periods and were averaged over the 
period of resting. Resting periods ranged from 5.45 to 
27.20 hours. 

All investigated platypuses occupied burrows 
in consolidated steep or gently sloping earth banks of 
the lake or along associated creeks. Water and air 
temperatures at Lake Lea were recorded in two-hour 


274 


intervals using archival tags (HOBO Thermocouple 
logger and Stowaway Temperature Logger, Onset 
Computer Corp., USA). Water temperature was 
measured in the lake while air temperature was taken 
in a wind shaded forest patch nearby. 


RESULTS 


Mean burrow temperatures were 17.5 and 
14.2°C (SD=2.76 and 0.89, respectively, n=9) in 
summer and winter, respectively, and ranged between 
12.2 and 22.8°C for individual resting periods. In 
winter, burrow temperature was held fairly constant 
over the resting period while in summer larger 
variations were observed. A low but significant 
correlation between air temperature and burrow 
temperature was found with higher air temperatures 
resulting in higher burrow temperatures (p=0.003, 
n=61). Ambient air temperatures ranged between -4°C 
and 31°C and water temperatures between O°C and 
29°C depending on season. Samples of measured 
burrow temperatures and corresponding air and water 
temperatures are shown in Fig. 1 and Fig. 2 for winter. 
and summer, respectively. 


Proc. Linn. Soc. N.S.W., 125, 2004 


P. BETHGE, S. MUNKS, H. OTLEY AND S. NICOL 


Temperature [°C] 


7A 68Nn OA 


104 114 124M 13/4 144 15/1 16 17/1 


18/1 19/1 20/1 


Date 


Figure 2: Summer sample data of water (Tw), air (Ta) and burrow temperatures (Tbur, derived from a 
time-depth recorder with integrated temperature sensor fitted to the back of a platypus; the temperature 
is only shown at times when the animal was in the burrow). 


DISCUSSION 


Grant (1983b) suggested that platypus 
burrows act as microenvironments, buffering the 
animals against the rigours of below-freezing air 
temperatures in winter, and modifying the effects of 
high summer temperatures. Accordingly, we found that 
in winter, burrow temperatures at Lake Lea were up 
to 18°C higher than outside air temperatures (Fig. 1). 
In summer, the burrows at Lake Lea clearly buffered 
high midday temperatures of over 25°C (Fig. 2). These 
findings are in line with results by Grant (1976) and 
Munks (personal communication). In winter, 
unoccupied artificial burrow temperatures in the upper 
Shoalhaven River, NSW, averaged around 14°C (this 
study: 14.2°C) despite the fact that ambient air 
temperatures dropped as low as -5°C. During summer 
the temperature of an unoccupied artificial burrow 
averaged around 18°C (this study: 17.5°C) with air 
and water temperatures being several degrees higher 
(Grant 1976, Grant 1995). Munks (personal 


Proc. Linn. Soc. N.S.W., 125, 2004 


communication), while monitoring the burrow of a 
breeding platypus in lowland Tasmania, recorded a 
mean burrow temperature of 16.5°C (range 12.5 to 
20°C) during late summer/early autumn. 

The consistency of these data from different 
sites suggests that platypus burrow temperatures are 
fairly constant regardless of habitat. Whether this is a 
consequence of the metabolic heat produced by the 
animals or mainly of physical characteristics of their 
burrows, remains unclear. Results of Grant (1976) from 
unoccupied burrows are in line with findings presented 
here from occupied burrows. This suggests that - at 
least in burrows located in consolidated earth banks - 
physical characteristics of the burrow are more 
important for burrow temperature than the absence or 
presence of the animal. This view is supported by the 
significant correlation between air temperature and 
burrow temperature found in this study. 

However, Munks (personal communication) 
reported peak burrow temperatures when the mother 
returned to the nest to suckle her young. Also, Grant 
(1983b) suggested that the animal’s presence further 


275 


PLATYPUS BURROW TEMPERATURES 


elevates the microhabitat temperature of the burrow. 
In captivity, Grant (1976) observed, that the 
temperature in an uninsulated plywood nest box rose 
around | to 2°C above ambient temperature when an 
animal was inside. 

We suggest that these different observation 
are a consequence of different burrow characteristics. 
In this study, all investigated platypuses occupied 
burrows in consolidated earth banks. Under such 
conditions, the insulation properties of the surrounding 
earth and of the nesting material inside the burrow are 
most likely the main factors determining burrow 
temperature. A fairly constant burrow temperature may 
of course be more critical during the breeding period 
(Grant, personal communication), which makes deep 
earth burrows ideal for nesting. 

A different situation, however, might occur 
in burrows, which are closer to the surface or above 
ground. Otley et al. (2000) reported that 25 % of 
burrows at Lake Lea were located within dense 
vegetation, such as sphagnum and button grass. The 
insulation properties of such burrows would be 
expected to be poor compared to underground earth 
burrows. How animals cope with high thermal stress 
in vegetation burrows and if they use this sort of burrow 
site regardless of season or even during nesting requires 
further investigation. 


ACKNOWLEDGMENTS 


This work was supported by the Australian 
Research Council, an Overseas Postgraduate Research 
scholarship by the University of Tasmania and a doctoral 
scholarship by the DAAD (Deutscher Akademischer 
Austauschdienst, Germany ,Hochschulsonderprogramm III 
von Bund und Landern‘). The field work was carried out 
under permit from the Department of Parks, Wildlife and 
Heritage, Tasmania, the Inland Fisheries Commission, 
Tasmania and the University of Tasmania Ethics Committee. 
Thanks to all those who assisted with the field work and to 
Mr H.Burrows for access to private land. 


276 


REFERENCES 


Bethge, P., Munks, S., Otley, H.and Nicol, S. (2003) 
Diving behaviour, dive cycles and aerobic dive 
limit in the platypus Ornithorhynchus anatinus. 
Comparative Biochemistry and Physiology A 
136/4, 799-809. 

Grant, T.R. (1976). Thermoregulation in the Platypus, 
Ornithorhynchus anatinus. PhD thesis, 
University of New South Wales, Australia. 

Grant, T.R. (1983a). Body temperatures of free-ranging 
platypuses, Ornithorhynchus anatinus 
(Monotremata), with observations on their use 
of burrows. Australian Journal of Zoology 31, 
117-122. 

Grant, T.R. (1983b). The behavioural Ecology of 
Monotremes. In ,Advances in the Study of 
Mammalian Behaviour* (Eds J.F. Eisenberg and 
D.G. Klieman). The American Society of 
Mammalogists, Special Publication Vol. 7, pp 
360-394. 

Grant, T.R. (1995). The platypus. A unique mammal. 
University of New South Wales Press, Sydney. 

Otley, H.M., Munks, S.A. and Hindell, M.A. (2000). 
Activity pattern, movements and burrows of 
platypuses (Ornithorhynchus anatinus) in a sub- 
alpine Tasmanian lake. Australian Journal of 
Zoology 48, 701-713. 

Serena, M. (1994). Use of time and space by platypus 
(Ornithorhynchus anatinus: Monotremata) 


along a Victorian stream. Journal of Zoology 
232, 117-130. 


Proc. Linn. Soc. N.S.W., 125, 2004 


Ultrasonography of the Reproductive Tract of the Short-beaked 
Echidna (Tachyglossus aculeatus) 


D. P. Hiccins 


Faculty of Veterinary Science, B01, University of Sydney NSW 2006 
(damienh @ vetp.usyd.edu.au) 


Higgins, D.P. (2004). Ultrasonography of the reproductive tract of the short-beaked echidna 
(Tachyglossus aculeatus). Proceedings of the Linnean Society of New South Wales 125, 277-278. 


We describe a brief investigation of ultrasonography as a tool to monitor reproductive activity and to 
determine the sex of short- beaked echidnas (Tachyglossus aculeatus). We found trans-abdominal ultrasound 
to be of limited use for monitoring ovum development but it appears to be useful for imaging the uterus. We 
also found ultrasonography to be a useful tool to confirm the sex of echidnas by visualizing the testis. 


Manuscript received 18 August 2003, accepted for publication 8 January 2004. 


KEYWORDS: abdomen, echidna, monotreme, reproduction, Tachyglossus, testis, ultrasound, uterus. 


Here we describe a brief investigation of 
ultrasonography as a tool to monitor reproductive 
activity and to determine the sex of short- beaked 
echidnas (Tachyglossus aculeatus). Griffiths (1968) 
described the gross anatomy of the reproductive tract 
of the female echidna. An ovum of 3-4 mm diameter 
is ovulated from one of the two flat, sauropsid-like 
ovaries, which lie ventrocaudal to the kidneys 
(Griffiths 1968). Although only one ovum is ovulated 
in the echidna, Flynn (1930) reported that up to three 
large ova and several much smaller ova may occur on 
the ovary. Hughes and Carrick (1978) concluded from 
Hill and Gatenby (1926), Caldwell (1887) and Flynn 
and Hill (1939) that the ovum has a vitelline membrane, 
a zona pellucida and a proalbumen which may be 
analogous to the liquor folliculi of the graafian follicle, 
but has no follicular antrum. During its passage down 
the fallopian tube, the ovum swells to 5 mm diameter. 
The shell membrane is first laid down in the fallopian 
tube and later thickens in the uterus. The egg absorbs 
fluid in-utero and expands from 6.5mm diameter to 
15 mm x 13 mm. 

Ten short- beaked echidna carcasses were 
placed in dorsal recumbency. A portable ultrasound 
machine with a 7.5 Mhz linear transducer (SSD- 500, 
Aloka, Japan) was used to image the abdomen. Results 
were confirmed by dissection. An additional nine 
echidnas were then anaesthetized and examined in a 
similar fashion. Positioning the transducer on the 
ventral abdominal wall, lateral to the epipubic bones 
avoided the need to shave the hair of the pseudopouch 
and minimised interference by intestinal gas. 


Dissection confirmed that the gonads lie against the 
dorsal body wall, dorsal to the cranial ends of the 
epipubic bones. Ovaries of freshly dead echidnas 
lacked grossly visible developing ova. Of frozen and 
thawed bodies, which generally had poorer tissue 
contrast, ovaries and ova were not visible by 
ultrasonography. A structure in the expected location 
of the ovaries and comprising several 2- 3 mm 
diameter, thin walled, echolucent bodies was 
sometimes visible in living echidnas during the 
breeding season, however, the scarcity of surrounding 
interstitium made repeatable identification of 
individual putative ova very difficult. In addition, the 
small intestine frequently cast gas shadows over the 
gonads, reducing their visibility. It is likely that trans- 
rectal ultrasound would improve visualization of the 
ovaries but may be of limited use in serial observations, 
where the extent of manual or chemical restraint 
required may introduce variations to the reproductive 
cycle (Clarke and Doughton 1983, River and Rivest 
1991). The entire oviducts of reproductively active live 
and dead animals were clearly visible, especially when 
adjacent to a full bladder. Ova were not seen in the 
oviducts of any of our animals. Testes appeared as 15 
to 25mm long, ovoid, homogenous, soft tissue 
structures and, when present, were always visible 
caudal to the kidney and, on the left side, dorso-caudal 
to the mobile, spherical portion of the spleen. Due to 
their similar appearance on ultrasound, both the spleen 
and testis were sighted on the left before the left testis 
was identified. 


ULTRASONOGRAPHY IN ECHIDNA REPRODUCTIVE STUDIES 


in conclusion, we found trans-abdominal 
ultrasound to be of limited use for monitoring ovum 
development but it appears to be useful for imaging 
the uterus. In sexing echidnas, the inability to extrude 
or palpate a phallus does not confirm its absence, and 
other characteristics such as absence of a pseudo-pouch 
or presence of spurs may not be reliable indicators of 
sex, therefore the gender of echidnas in captive 
collections is sometimes mistaken. We found 
ultrasonography to be a useful tool to confirm the sex 
of echidnas in these circumstances. 


ACKNOWLEDGMENTS 


We thank Dr Marianne Offner for her assistance 
in interpretation of ultrasonographs, Medtel for the provision 
of the Aloka SSD-500 ultrasonography machine, and the 
Zoological Parks Board of NSW, Novartis Australia and the 
Winifred Scott Foundation for their financial support. 


REFERENCES 


Caldwell, W. H. (1887) The embryology of Monotremata 
and Marsupialia- Partl. Philosophical 
Transcripts of the Royal Society. 178(B), 463- 
480. 

Clarke, I. J. and Doughton, B. W. (1983) Effect of various 
anaesthetics on the resting plasma 
concentrations of lutienising hormone, follicle 
stimulating hormone and prolactin in 
ovariectomised ewes. Journal of Endocrinology. 
98, 79-89. 

Flynn, T. T. (1930). On the unsegmented ovum of the 
echidna (Tachyglossus) Quarterly Journal of 
Microscopical Science. 74, 119-131. 

Flynn, T. T. and Hill, J. P. (1939) The development of the 
Monotremata PartVI -Growth of the ovarian 
ovum, maturation, fertilisation and early 
cleavage. Philosophical Transcripts of the Royal 
Society (London) XXIV (6), 571-578. 

Griffiths, M. E. (1968) ‘The Echidna.’ (Pergamon Press: 
UK). 

Hill, J. P. and Gatenby, J. B. (1926). The corpus luteum of 
the Monotremata. Philosophical Transcripts of 
the Royal Society (London) I, 715-762. 

Hughes, R. L. and Carrick, F. N. (1978). Reproduction in 
female monotremes. Australian Zoologist 20, 
233-254. 

River, C. and Rivest, S. (1991). Review Article. Effect of 
stress on the activity of the hypothalamic- 
pituitary- gonadal axis: peripheral and central 
mechanisms. Biology of Reproduction 45, 523- 
532. 


278 


Proc. Linn. Soc. N.S.W., 125, 2004 


Excretion Profiles of Some Reproductive Steroids in the Faeces 


of Captive Female Short-beaked Echidna (Tachyglossus 
aculeatus) And Long-beaked Echidna (Zaglossus sp.) 


D.P. Hicains, G. Tosias, G.M. STONE 


Faculty of Veterinary Science, BO1, University of Sydney NSW 2006 (damienh@vetp.usyd.edu.au) 


Higgins, D.P., Tobias, G. and Stone, G.M. (2004). Excretion profiles of some reproductive steroids in the 
faeces of captive female short-beaked echidna (Tachyglossus aculeatus) and long-beaked echidna 
(Zaglossus sp.). Proceedings of the Linnean Society of NSW 125, 279-286. 


We evaluated and applied an existing faecal reproductive steroid extraction and radio-immunoassay (RIA) 
procedure to samples from captive short-beaked (Tachyglossus aculeatus) and long- beaked (Zaglossus sp.) 
echidnas. Steroids were extracted from faeces with diethyl ether, resuspended in 80% methanol and lipids 
removed with petroleum ether. The methanol fraction was purified and assayed for progestins or oestrogens, 
results corrected for procedural losses and converted to ng/ g dry weight of faeces. One T. aculeatus was 
injected with radiolabelled and natural progesterone and faecal extracts were subjected to high- performance 
liquid chromatography (HPLC) to allow partial identification of radiolabelled and RIA- reactive metabolites. 
The major RIA-reactive substance and the major labelled ['*C] compound co-eluted with progesterone. An 
additional RIA-weak compound co-eluted with 208-dihydroxyprogesterone, and three additional RIA-weak, 
radio-labelled compounds eluted but were not identified. Increases in faecal progestin of echidnas occurred 
at 17 + 3 (n=5), 33 + 3 (n =4) and 48 (n = 1) day intervals, supporting a cycle length of approximately 17 
or 33 days. However, further study incorporating more animals, behavioral observations and more frequent 


sampling of faecal oestrogens is required to produce more definitive results. 


Manuscript received, 18 August 2003, accepted for publication 8 January 2004. 


KEYWORDS: Faecal reproductive steroids, HPLC, monotreme, oestrogen, progestin, 
radioimmunoassay, Tachyglossus aculeatus, Zaglossus. 


INTRODUCTION 


The short- beaked echidna (Tachyglossus 
aculeatus) is widespread within Australia and New 
Guinea. The long- beaked echidna (Zaglossus bruijnii) 
is restricted to the highlands of New Guinea where it 
is endangered by human interference (Flannery 1990). 
Despite more than 100 years of captive husbandry, it 
is rare for these animals to breed in captivity (Augee 
et al. 1978; Boisvert and Grisham 1988) and 
knowledge of the timing and hormonal control of the 
reproductive cycles of monotremes is limited. The 
presence of a luteal phase is generally accepted, based 
on histological evidence (Hill and Gatenby 1926; 
Griffiths 1968; Hughes and Carrick 1978; Griffiths 
1984) but its role and duration is unknown. In addition, 
observations of gestation range from greater than 10 
days (Carrick 1977) to 28 days (Broom 1895) after 
mating. Griffiths (1984) speculated that, like some 
reptiles and bats (Racey and Potts 1970), female 
echidnas may store sperm, or that torpor may alter 


gestational length, as in pygmy possums (Cercartetus 
spp), brown antichinus (Antichinus stuartii), eastern 
quolls (Dasyuris viverrinus) (Tyndale- Biscoe 1973) 
and bent-winged bats (Miniopterus spp) (Wimsatt 
1969). 

Longitudinal studies better define 
reproductive cycles and illustrate inter-individual 
variation than cross-sectional studies, which are more 
conveniently applied to wild animals (Lasley 1985). 
However, even captive echidnas are cryptic and curl 
into a tight ball when threatened, making difficult the 
frequent collection of blood, urine or urogenital swabs 
without anaesthesia or forceful restraint, which may 
cause variation of reproductive cycles and behaviour 
(Clarke and Doughton 1983; Rivier and Rivest 1991; 
Cleva et al 1994). Non- invasive faecal reproductive 
steroid assays have been used to describe the 
reproductive cycles of many species. This paper reports 
the initial assessment and application of a faecal 
reproductive steroid assay as a non- invasive technique 
for the first sequential study of female echidna 


FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 


reproductive endocrinology. 


MATERIALS AND METHODS 


Animals and housing 

Study animals were six female T. aculeatus, 
aged between 4 and 7 years and of 3 to 5 kg 
bodyweight, and three female Zaglossus bruijnii 
(probably Z. bartoni of Flannery and Groves 1998), 
aged between 20 and 32 years and of 6.5 to 14 kg 
bodyweight, all from the Taronga Zoo collection. The 
study was conducted in two phases: From May to 
September 1995, two T. aculeatus and all Zaglossus 
sp. were housed in two indoor enclosures with reverse 
cycle seasonal lighting and in the continual presence 
of a male of their species. From June to October 1997, 
four female 7. aculeatus were housed outdoors, in 
adjacent 5.1 x 6.3 m enclosures. These females were 
housed individually to accommodate the solitary nature 
of the animal (Augee et al. 1975; Abensperg-Traun 
1991) and to facilitate identification of the source of 
faeces. A male T. aculeatus had access to all four 
enclosures through magnetically controlled doors until 
they failed, after which he was manually rotated, daily, 
between enclosures. Sixty-centimeter deep woodchip 
substrate, half pipes, tables, tree branches and logs were 
provided as shelter. 


Feeding and sample collection 

Animals were fed daily slurry of minced beef, 
egg, cereal, vitamin and mineral supplements and 
sufficient unprocessed bran to produce firm stools. 
Initially, 1mm x 1mm x 3mm food grade polyethylene 
pellets (Hoechst Industries, Australia) were added to 
the food of the indoor groups to identify the source of 
faeces. It appeared possible that not all pellets were 
being excreted, therefore the feeding of pellets was 
discontinued and for the course of the study each 
animal from the indoor groups was placed in a separate 
room for 24- 48 h once weekly and faeces were 
collected. Blue food dye (8 mg/day, Hexacol Brilliant 
Blue FCF Supra 75328, Pointing Hodgsons Pty Ltd, 
Australia), was added to the food of the outdoor female 
animals to make faeces more visible, and all visible 
faeces were collected daily. Samples were handled 
using latex gloves and stored in plastic zip- lock bags 
at -20°C for up to one year. 


Extraction and purification 

Due to the need to separate echidnas for 
sample collection in the first phase of the study, the 
sampling interval for progestin excretion profiles of 
echidnas housed indoors was 5 to 7 days. Sampling 


280 


interval for progestin and oestrogen excretion profiles 
of echidnas housed individually outdoors in the second 
phase of the study was | to 3 days. The steroid 
extraction technique was based on a procedure used 
by Hindle and Hodges (1990). Each stool was finely 
chopped and mixed, then duplicate 0.5g samples were 
transferred to new glass vials (Econo Glas Vial, 
Packard, USA). Pieces were broken up using a glass 
rod, 5 ml diethyl ether (APS, Ajax Finechem, 
Australia) was added and vials were rotated for 30 min 
then centrifuged at 1500 G for 15 min at 4°C. The 
faecal sediment and aqueous portion were frozen in 
liquid nitrogen. Supernatant was decanted, evaporated 
at 30°C under nitrogen gas and reconstituted in 5 ml 
80%(v/v) methanol (80% MeOH) by rotation for 30 
min. Solutes were partitioned by addition of 5ml of 
petroleum ether (B.P. 40°C to 60°C, APS, Ajax 
Finechem, Australia), rotation for 20 min, and 
centrifugation at 1500 G for 15 min. The 80% MeOH 
fraction was aspirated and then stored at -20°C. 
Following extraction, faecal residue was dried at 100°C 
for 4 hours and weighed to determine dry matter 
content. 

Aliquots of 500 ul faecal extract in 80% 
MeOH were dried at 80°C under vacuum, reconstituted 
in Iml 10% MeOH by agitation at 30°C for 30 min, 
and purified using Sep-Pak C18 Cartridges (Waters 
Scientific, Milford, USA) according to manufacturers 
recommendations. Eluants of 25%, 50%, 75%, 90% 
and 100% MeOH were collected and stored at -20°C. 


Radioimmunoassay (RIA) 

Duplicate 200 ul aliquots of eluates 
(unknowns) were dried and reconstituted in 200ul 10% 
MeOH in 1P buffer (0.031M Na,HPO,, 0.019M 
NaH,PO,.2H,O, 0.154M NaCl and 0.1%w/v gelatin, 
pH 7.4) by agitation at 30°C for 60 min. Radiolabelled 
steroids ([1,2,6,7-7H] progesterone in toluene ([7H]P, 
96 Ci/mmol; Amersham Australia, Sydney, NSW) or 
[2,4,6,7-7H] oestradiol in toluene ((7HJE, 104 Ci/mmol; 
Amersham Australia, Sydney NSW)) were dried and 
reconstituted in 1P buffer to approximately 15000 dpm/ 
100 wl. Our ovine antiserum to progesterone-110- 
hemisuccinate-BSA conjugate (1:55000 final dilution, 
#C-9817 Sirosera™, CSIRO Bioquest, Blacktown, 
Australia), cross-reacted with progesterone (100%), 
11B- hydroxyprogesterone (32.5%), corticosterone 
(18.8%), 20a- hydroxy-4-pregnane-3-one (0.7%), 
17a- hydroxyprogesterone (0.2%), 20B- hydroxy-4- 
pregnane-3-one (0.2%), pregnenolone (0.2%), 
oestradiol (<0.2%), testosterone (<0.2%), cortisol 
(<0.2%) (Curlewis, Axelson and Stone, 1985). Our 
ovine antiserum to 17B-  oestradiol-6- 
carboxymethyloxime-BSA (1:100000, #9757 


Proc. Linn. Soc. N.S.W., 125, 2004 


D.P. HIGGINS, G. TOBIAS AND G.M. STONE 


Sirosera™, CSIRO Bioquest, Blacktown, Australia), 
cross-reacted with oestradiol (100%), oestrone 
(10.8%), oestriol (2.3%), oestradiol- 17a (<0.1%), 
progesterone (<0.1%), testosterone (<0.1%), 
androstenedione (<0.1%), cortisol (<0.1%), 
corticosterone (<0.1%) (Curlewis 1983). Standards 
were generated from two overlapping doubling 
dilutions of progesterone (BDH Chemicals, Australia) 
from 500 - 7.84 pg/100 ul or oestradiol (BDH 
Chemicals, Australia) from 500 - 1.96 pg/100 ul. 

Reactions contained 100 ul radiolabelled 
steroid, 100 wl antiserum and either 200 ul of unknown 
in 10% MeOH in 1P buffer or 100 ul 20% MeOH in 
1P buffer and 100 ul of standard. The resulting 400 ul 
was vortexed for 30 sec and incubated at 4°C for 18 h. 
Triplicate “total” (200 wl 1P buffer, 100 wl 
radiolabelled hormone in 1P buffer, 100 tl 20% MeOH 
in 1P buffer), “non-specific binding” (200 tl 1P buffer, 
100 ul radiolabelled hormone in 1P buffer, 100 ul 20% 
MeOH in 1P buffer) and “B ” (100 ul 1P buffer, 100 
ul antiserum in 1P buffer, 100 wl radiolabelled 
hormone in 1P buffer, 100 ul 20% MeOH in 1P buffer) 
standards were processed simultaneously with 
unknowns and standards. 

Free radiolabelled hormone was removed 
from all except “total” solutions by incubation for 10 
min at 4°C with 500 ul of charcoal/ dextran solution 
(0.25% w/v Norit-A filtered activated charcoal powder, 
Matheson, Coleman and Bell; USA) and 0.025% w/v 
dextran T70 (Pharmacia Fine Chemicals, Sweden) 
suspended in 1P buffer). In place of the charcoal/ 
dextran solution, 500 ul of milli Q water was added to 
“total” solutions. After centrifugation at 1500 G for 
10 min at 4°C, the supernatant was decanted and its 
radioactivity measured as counts per minute (cpm) on 
a Beckmann LS 6500 Liquid Scintillation Spectrometer 
(Beckmann Instuments Inc, CA, USA.), which then 
converted cpm to disintegrations per minute (dpm) 
using an external standard. 


High- performance liquid chromatography (HPLC) 
of excreted metabolites 

One female T. aculeatus was injected intra- 
peritoneally with 5 mCi of [4-'*C] progesterone ({“C]P, 
48.9 mCi/mmol, NEN Dupont, USA) and 2 mg of 
natural progesterone in 30% (v/v) propylene glycol in 
isotonic saline. Eight 0.5 g faecal samples were 
obtained two days after injection. Extracts from these 
samples were pooled into two samples and subjected 
to sep-pak chromatography. Eluates of 2252 dpm and 
2440 dpm were dried under N, gas, reconstituted in 
75% acetonitrile, filtered and subjected to HPLC 
(K65B HPLC system, ETP Kortec, Australia) at a flow 
rate of 0.5 ml per minute, using 61% acetonitrile at a 


Proc. Linn. Soc. N.S.W., 125, 2004 


pressure of 2250 psi at room temperature. Fractions 
were collected every 30 sec for 22 min, then every 
minute for 19 min, then every 10 min for 20 min. 
Absorbance at 240nm was measured, to monitor the 
separation of steroids with a 4-ene-3-ketone structure. 
Elution time of progesterone was identified using [7H]P 
and a progesterone standard and the column was 
calibrated for testosterone, androstenedione, 
progesterone and 20am dihydroprogesterone. 


Assessment of extraction, purification and RIA 
procedures 

Three different solvents were tested for use 
in the extraction process. Faeces containing metabolites 
of injected radiolabelled and natural progesterone were 
agitated in 90% MeOH, 80% MeOH ox diethyl] ether, 
and partitioned with petroleum ether as described 
above. The three solvents and their petroleum ether 
portions were assayed for progestins as above. 

The Sep-pak chromatography elution profile 
for oestrogen calculated by Spanner et al. (1997) was 
assumed for this study. The elution profile for progestin 
was determined by Sep-pak chromatography of 
solutions containing 200 fmol [7H]P, using the series 
of MeOH dilutions described previously or the same 
series of dilutions of ethanol (EtOH). Co-elution of 
metabolites of faecal origin with progesterone was 
determined by adding 25000 dpm [7H]P to duplicate 
faecal extracts from five female 7. aculeatus and 
subjecting these to Sep-pak chromatography. 

Sample steroid recovery was estimated by 
adding 25000 dpm [?H]P or 30000 dpm [?HI]E to 
respective samples and then performing the extraction. 
Duplicate 50 ul aliquots of purified 80% MeOH extract 
were combined with 500 wl Milli-Q water and 5 ml of 
scintillation fluid. Triplicate “total” vials were 
prepared, each containing 100 pl of radiolabelled 
hormone solution, 50 ul 80% MeOH, 400 wl Milli-Q 
water and 5 ml of scintillation fluid. Triplicate “blank” 
vials were prepared, each containing 50 wl 80% MeOH, 
500 wl Milli-Q water and 5 ml of scintillation fluid. 
Radioactivity was measured and percentage recovery 
was calculated by the formula: 


R = [400(d-B)/ (T-B)] x 100 


where R = percentage recovery (%), d = sample dpm, 
B = mean “blank” dpm, T = mean “total” dpm. 

To assess parallelism, faecal extracts from 
three faecal samples were reconstituted and diluted 
twofold and fourfold in 10% MeOH in 1P buffer. 
Standards were similarly diluted and all dilutions were 
assayed for progestins as described above. Spanner et 
al. (1997) estimated parallelism of the oestradiol assay. 


281 


FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 


The inter- assay coefficients of variation for 
progesterone and oestradiol assays were taken as the 
mean of the coefficients of variation of repeated (n=2), 
duplicated extraction and assay of 6 and 4 randomly 
chosen samples, respectively. 

The intra- assay coefficient of variation was 
estimated from the mean of two coefficients of 
variation, each calculated from five concurrent 
replicate extractions and assays of two randomly 
chosen samples. The intra-assay coefficient of variation 
was estimated for two progesterone extraction methods 
to determine the homogeneity of steroid in the stool. 
In the first (unmixed) method, 5 samples were taken 
from an intact stool. In the second (mixed) method, 
the stool was finely chopped and mixed and each of 
the 5 samples consisted of at least 5 randomly chosen 
pieces from the mix. Spanner et al. (1997) estimated 
the intra-assay coefficient of variation of the oestradiol 
assay. 


Data analysis 

Standard curve generation and conversion of 
dpm to pg hormone’ scintillation vial were calculated 
with “Assayzap” (Biosoft, Cambridge). All other 
calculations and graphs were made using “Excel 5.0” 
(Microsoft, USA). Mean steroid recovery was 
calculated from the first 60 samples in each assay. 
Mean recovery was used to correct results of progestin 
assays for procedural losses. As steroid recovery was 
more variable in oestrogen assays, results were 
corrected using a recovery value calculated for each 
individual sample. Faecal steroid peaks were defined 
as those values greater than 1.5 standard deviations 
from the mean of all values from that animal (Graham 
et al 1995). 


RESULTS 


High pressure liquid chromatography 

The eluate with the highest RIA activity and 
moderate radioactivity was collected at 26 min, 
approximating the progesterone standard, which eluted 
at 25.5 min. [H]P eluted at 24 min. A moderately 
radioactive eluate with poor RIA activity that was 
collected at 22 min coincided with a 20B- 
dihydroxyprogesterone standard, which has a low cross 
reactivity with the antiserum. Other ["C]-labelled, 
moderately RIA-reactive compounds that eluted at 30, 
36 and 38 min and one ['4C]-labelled, weakly RIA- 
reactive compound that eluted at 22 min were not 
identified. One RIA-reactive compound that eluted at 
41 min did not co-elute with a ['*C]-labelled metabolite 
and this substance is yet to be identified. 


282 


Assessment of progestin extraction, purification 
and RIA procedures 

As an initial solvent, ether extracted 37.0 + 
4.3% (mean + S.E.) more ['4C] labelled progestin than 
either 80% MeOH or 90% MeOH and was used in all 
subsequent extractions. Less than 10% of extracted 
steroid appeared in the petroleum ether fraction. Mean 
percentage recovery of [*H]P through extraction and 
Sep-pak chromatography was 52% + 7.17 (mean + 
SD). Mean percentage recovery of [HJE was 30% + 
13.7 (mean + SD). 

Almost all [*H]P was recovered during Sep- 
pak chromatography. MeOH was chosen as the 
chromatography solvent as EtOH eluted the [*H]P 
across a greater range of EtOH concentrations. Of 
recovered [7H]P metabolites, 79.1% eluted in the 90% 
MeOH fraction and 93% eluted in the 75% MeOH 
and 90% MeOH fractions combined, with a mean 75% 
MeOH: 90% MeOH ratio (75:90 ratio) of 1:3.5. Of 
the progestin RIA- reactive faecal steroids recovered 
from the column, 85% was measured in the combined 
75% MeOH and 90% MeOH fractions with a mean 
75:90 ratio of 1:2.5. 

Correlation coefficients of parallelism curves 
for the progestin assay ranged from 0.988 to 1.000. 
Dose response curves for standards and extracts did 
not differ significantly (P>0.05) in slope. Sensitivity 
of the assay, as defined by 10% displacement from 
the Bo binding was 10 pg/assay tube. The intra- assay 
coefficients of variation were estimated to be 6.2% 
(mixed) and 25.7% (unmixed), therefore the mixed 
method was employed in all further extractions. The 
inter- assay coefficient of variation was estimated to 
be 14.9% for progesterone assays and 6.8% for 
oestradiol assays. 


Faecal progestins 

Maximum and minimum faecal progestin 
concentrations from each of the six T. aculeatus ranged 
from 480 to 1800 ng/g dry weight faeces (mean 860 
ng/g) and 5 to 100 ng/g dry weight faeces (mean 71 
ng/g), respectively. Intervals between samples that 
contained progestin peaks clustered at 17 + 3 (n = 5), 
33 + 3 (n = 4) and 48 (n = 1) days. Maximum and 
minimum faecal progestin concentrations from each 
of the three Zaglossus sp. ranged from 260 to 500 ng/ 
g dry weight faeces (mean 420 ng/g) and 10 to 70 ng/ 
g dry weight faeces (mean 40 ng/g), respectively. Two 
Zaglossus sp. produced two peaks each and the 
intervals between samples that contained these were 
28 and 70 days. The third produced one peak. 


Faecal Oestrogens 


Maximum and minimum faecal oestrogen. 
concentrations from the four animals ranged from 21 


Proc. Linn. Soc. N.S.W., 125, 2004 


D.P. HIGGINS, G. TOBIAS AND G.M. STONE 


to 45 ng/g dry weight faeces (mean 33 ng/g) and 3 to 
14 ng/g dry weight faeces (mean 7 ng/g), respectively. 
Intervals between adjacent oestrogen peaks were 8, 
19, 24 and 30 days apart. Fluctuations approaching 
1.5 SD above the mean were common and made 
difficult the detection of any possible cyclic activity 
as intervals between these ranged from 4 to 16 days. 


Combined profiles 

Of the 8 oestrogen peaks, 7 were associated 
with progestin increases to concentrations less than 
1.5 SD above the mean. Combined oestrogen and 
progestin profiles from two T. aculeatus are shown 
in Figures 1 and 2. 


DISCUSSION 


Feeding and sample collection 

Plastic pellets were less suitable as a faecal 
marker than the blue food dye. T. aculeatus consumed 
95% of pellets placed in their food while Zaglossus 


sp. consumed less than 50%. Not all ingested pellets 
were recovered, indicating that pellets were being 
retained or faeces were remaining undetected. Food 
containing the blue dye was readily eaten and faeces 
containing the dye were considerably more detectable 
than those without. Passage time of the plastic pellets 
ranged from 12 hours to greater than 48 hours. 


High pressure liquid chromatography 

The strong antiserum cross-reactivity of the 
substance which eluted at 26 min, and its proximity to 
the elution time of the progesterone standard, makes 
progesterone its likely identity, however further 
confirmation of this would be desirable as we are 


unable to explain the elution of H]P 2 min earlier. 


The cross-reactive metabolites that were less 
polar than progesterone were not identified. These 
substances may contribute to the difference in 75:90 
ratio between the Sep-pak elution profiles of [7H]P and 
progestins of faecal origin as the 75%MeOH, or less 
polar, component was less in the HIP profile. As a 
priority, future studies should identify the RIA-reactive 


500 


400 


progestins (ng/g dwt) 
(d>) 
oS 
(eo) 


56 64 65 68 69 70 72 75 76 77 79 81 83 86 89 92 94 96 


day 


oestrogens (ng/g dwt) 


mg Progestin 
—¢—oestrogen 


Figure 1. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 40-day period 
showing alternating progestin and oestrogen peaks greater than 1.5 SD above the mean, suggesting an 
oestrous cycle of 29 days. Also visible are increases less than 1.5 SD above the mean, suggesting concurrent 
progestin and oestrogen rises at 65, 79 and 94 days with interceding raised progestin/ lowered oestrogen 


periods surrounding 70 and 89 days, suggesting two cycles of 15 days. 


= mean faecal progestin/ 


oestrogen concentration; ................. = mean faecal progestin/ oestrogen concentration + 1.5 SD. 


Proc. Linn. Soc. N.S.W., 125, 2004 


283 


FAECAL REPRODUCTIVE STEROIDS IN CAPTIVE ECHIDNAS 


ae Progestins 


—¢— oestrogens 


progestins (ng/g dwt) 


27 33 36 42 45 52 58 61 62 64 66 69 71 72 74 76 78 80 82 84 86 88 90 92 94 
days 


oestrogens (ng/g dwt) 


Figure 2. Combined faecal oestrogen and progestin profiles of one T. aculeatus over a 67-day period 
showing alternating progestin and oestrogen peaks greater than 1.5 SD above their respective mean, 
suggesting an oestrous cycle of approximately 32 days. Also visible are additional fluctuations of oestrogen 
concentration at 52, 78, 82 and 86 days and progestin at 36 days, which hinder clear interpretation of 


oestrous cycles. 
progestin/ oestrogen concentration + 1.5 SD. 


compound that did not correspond to a radiolabelled 
metabolite. A similar examination of oestrogen 
metabolites would assist interpretation of oestradiol 
assays. 


Extraction and recovery 

As homogeneity of steroid in the faeces was 
poor, mixing of the stool before sampling was 
necessary to reduce intra-assay variance. Though 
diethyl ether extracted the most steroid, recovery 
through extraction was low and variable in this study, 
especially in oestradiol assays. We attempted to correct 
for this by correcting for procedural losses using 
individual recovery values for each sample in 
oestrogen assays but progestin assay data were 
corrected using a mean recovery value. Use of 
individual recovery values for progestin samples may 
have improved interpretation of data. 

MeOH concentrations exceeding 5% in the 
RIA incubations considerably reduced steroid- 
antibody binding. At each corresponding 


284 


= mean faecal progestin/ oestrogen concentration; ................00+ = mean faecal 


concentration, EtOH had a greater effect on steroid- 
antibody binding than MeOH. Reconstitution of eluates 
in 10% MeOH to produce a final concentration of 5% 
MeOH in the assay provided adequate steroid solubility 
and minimised interference with steroid-antibody 
binding. 


Progestin and oestradiol profiles 

Mean, maximum and minimum progesterone 
and oestrogen values varied among animals, indicating 
that this technique may be unsuitable for assessing the 
status of an animal from a single measurement. The 
small number of animals available for the study and 
the need for further work to identify antiserum-reactive 
metabolites limits the conclusions that can be drawn 
from the sequential data obtained in this study. 
However, the lack of knowledge in this area makes 
some trends worthy of comment for consideration in 
future work. 

The intervals between subsequent progestin 
peaks in this study suggest a progesterone periodicity 


Proc. Linn. Soc. N.S.W.; 125, 2004 


D.P. HIGGINS, G. TOBIAS AND G.M. STONE 


of 16-17 days. However, as there was no clear pattern 
in oestrogen excretion, we could not determine whether 
this reflects concurrent vitelline progesterone and 
oestrogen peaks at 32- 34 day intervals with an 
interspersed luteal peak at 16-17 days (see figs 1 and 
2), or concurrent vitelline progesterone and oestrogen 
peaks at 16-17 day intervals with an interceding luteal 
phase with progestin increases below our arbitrary 
significance criterion. We expect that daily sampling 
and identification of potentially confounding RIA- 
reactive faecal steroids would be necessary to resolve 
this question. However, observations of fetal 
development add some support to the hypothesis of a 
17-day progesterone cycle. Decreasing blood 
progesterone is a precursor to parturition in many 
species of eutheria (Rowlands and Wier 1984) and 
metatheria (Tyndall-Biscoe and Renfree 1987), and 
to oviposition in many reptilia (Licht 1984). At 17 days 
the tammar wallaby fetus consists of 17- 20 somite 
pairs (Griffiths 1984), similar to the 19-20 somite pairs 
possessed by the echidna at oviposition (Hill and 
Gatenby 1926; Luckett 1976; Hughes and Carrick 
1978). Both young also exhibit similar stages of 
development at parturition or hatching 11 days later 
(Griffiths 1984). The similar rates of development in 
the last third of gestation and incubation suggests that 
the age of the echidna fetus at oviposition is 
approximately 17 days, consistent with a luteal phase 
of 16 to 17 days. 

The many irregular oestrogen fluctuations we 
measured could be inherent in the technique or 
indicative of follicular development and atresia. Hill 
and Gatenby (1926) described channels, from the 
vitellus to a well-developed lymphatic sinus in the 
ovarian medulla and histological features indicative 
of follicular regression in the platypus. 

The authors believe that this study provides 
a starting point for further work and suggest the further 
identification of progesterone and oestrogen 
metabolites and the comparison of faecal steroid 
concentrations with blood hormone concentrations, 
urogenital cytology, ultrasonography of the 
reproductive tract or behaviour in a controlled study 
accounting for the potential confounding effects of 
repeated physical or chemical restraint. 


ACKNOWLEDGEMENTS 


We thank the staff of the faculty of Veterinary 
Science, University of Sydney, in particular Michael Lensen, 
Irene van Ekris, Margaret Byrne and Geoff Dutton; and 
Taronga Zoo, in particular Margaret Hawkins, Kerry Foster, 
and Debbie Pritchard for their assistance. We also thank 
Michele Thums and the journal referees for their comments 


Proc. Linn. Soc. N.S.W., 125, 2004 


on the manuscript. The project would not have been possible 
without the help of the staff of Australian Mammals, Taronga 
Zoo and the Taronga Zoo Friends and without the financial 
assistance of Novartis, Australasia and the Winifred Scott 
Foundation. 


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Proc. Linn. Soc. N.S.W., 125, 2004 


Anatomy of the Central Nervous System of the Australian 
Echidna 


M. Hassiotis!, G. PAxINos? AND K.W.S. ASHWELL!* 


"Department of Anatomy, School of Medical Sciences, The University of New South Wales, 2052, Sydney, 


NSW, Australia. 


Prince of Wales Medical Research Institute, The University of New South Wales, 2052, Sydney, NSW, 


Australia. 
*author to whom correspondence and proofs should be addressed. Postal address as above. 
Fax: 61 2 9385 8016, Phone: 61 2 9385 2482 
Email: k.ashwell @unsw.edu.au 


Hassiotis, M., Paxinos, G. and Ashwell, K.W.S. (2004). Anatomy of the central nervous system of the 
Australian echidna. Proceedings of the Linnean Society of New South wales 125, 287-300. 


Even from their gross appearance, the brain and spinal cord of the Australian echidna show unusual 
features. The spinal cord is one of the shortest ever recorded for any mammal, ending at the mid-thoracic 
level, a feature which may be related to the defensive posture of the echidna. The pattern of termination of 
unmyelinated afferents in the spinal cord as revealed by lectin labelling with the B4 isolectin from Griffonia 
simplicifolia is also quite different from that seen in placental mammals, with termination in patches within 
deeper layers of the dorsal horn. Within the brainstem, specializations of the trigeminal system are apparent 
with great enlargement of all trigeminal nuclei. The mesencephalic trigeminal nucleus also shows an unusual 
aggregation of neurons in a central midline position quite unlike therian mammals. While the dorsal thalamus 
of therian mammals shows compartmentation related to function , the dorsal thalamus of the echidna is 
remarkable for its lack of cytoarchitectural differentiation. Most of the high encephalization in this mammal 
is attributable to the highly gyrified cerebral cortex. This cortex is further distinguished by the positioning 
of the major functional areas (primary motor, somatosensory, visual and auditory areas) towards the caudal 


pole of the brain. 


Manuscript received 21 July 2003, accepted for publication 22 October 2003. 


KEYWORDS: cerebral cortex, echidna, monotreme, spinal cord, thalamus, trigeminal. 


INTRODUCTION 


In this paper we will be reviewing what is 
known about the anatomy of the central nervous system 
of the Australian echidna (Tachyglossus aculeatus), 
with special reference to those features with functional 
relevance. Even at the level of gross inspection, the 
central nervous system of the echidna is remarkable 
for the large size of the brain and the short relative 
length of the spinal cord. 


PERIPHERAL RECEPTORS AND 
ELECTRORECEPTION 


One of the most remarkable features of 
monotreme neurobiology, and one which touches on 
trigeminal nuclei development and cortical 
organization, is the reported presence of 
electroreception in two members of this subclass 


(short-beaked echidna and platypus) (Iggo et al. 1985; 
Scheich et al. 1986; Gregory et al. 1987, 1988, 1989; 
Proske et al. 1998). This sensory modality utilizes the 
trigeminal system in both monotremes studied. 

To date, physiological and anatomical studies 
of peripheral sensory systems in this animal have 
concentrated on peripheral receptors of the trigeminal 
system. The short-beaked echidna is known to use its 
sensitive snout as its major sensory tool. Anatomical 
studies of this snout have revealed a rich distribution 
of unusual receptors on the tip (Andres et al. 1991; 
Manger and Hughes 1992). One of these, the gland 
duct receptor system (Andres et al. 1991) or mucous 
sensory gland (Manger and Hughes 1992), is present 
in both platypus and echidna and is thought to be 
involved in electroreception (Iggo et al. 1985; Scheich 
et al. 1986; Gregory et al. 1987, 1988, 1989). Despite 
this attention to snout receptors, very little attention 
has been given to the structure or function of central 
trigeminal pathways in any monotreme. 


ECHIDNA CENTRAL NERVOUS SYSTEM 


SPINAL CORD ANATOMY 


The spinal cord of the Australian echidna was 
examined by Ashwell and Zhang (1997). Even at the 
gross level, the spinal cord is notable because of its 
relatively short length, terminating at the level of the 
seventh thoracic vertebra (Figure 1a)(cf human spinal 


an adaptation to the pronounced vertebral flexure, 
which this mammal achieves when it adopts its 
defensive posture (Figure 1b). Since the spinal cord 
lies posterior to the vertebral column, extreme flexion 
would place the neural elements (spinal cord and cauda 
equina) under considerable tension, amounting to an 
increase of 15% in length or 6 cm in a large adult. The 


cord which terminates at the intervertebral disc 
between lumbar vertebrae! and 2). This may represent 


cauda equina in this animal is collectively as thick as 
the spinal cord, but consists of multiple nerve bundles 


a Echidna - ambulatory posture 


Cauda equina 
and spinal nerves 


b Echidna - defensive posture 


Cauda equina 
and spinal nerves 


Figure 1. The spinal cord of the Australian echidna is very short, ending opposite the seventh thoracic 
vertebra when the animal is in the ambulatory posture (a). This may be an adaptation which allows 
pronounced flexing of the vertebral column in the defensive posture (b), since the spinal nerves of the 
cauda equina would be more tolerant of the stretching associated with flexing of the vertebral column 
than the much thicker and more vascular spinal cord. Figures c and d show features of the spinal cord 
reported in Ashwell and Zhang (1997). Please see that paper for ethical clearance details and tissue 
preparation methods. Figure 1c shows the large neurons of the median nuclear group (arrowhead) at the 
lower lumbar level of the spinal cord (L4). The small inset indicates the position of the larger image. CC 
— central canal of spinal cord: DC — dorsal column; VC — ventral column; VH — ventral horn. Figure 1d 
shows unmyelinated afferent fibres labelled with a peroxidase conjugated B4 isolectin from Griffonia 
simplicifolia. The small inset indicates the position of the larger image. These afferents enter via Lissauer’s 
zone (LZ) and some descend to deep layers of the dorsal horn (arrowhead) terminating in the nucleus 
proprius (NP), unlike unmyelinated afferents to therian spinal cord, which are confined to the superficial 
layers; e.g. marginal zone — MZ; substantia gelatinosa — SG). 


288 Proc. Linn. Soc. N.S.W., 125, 2004 


M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 


which are free to move independently of each other, 
unlike the spinal cord where individual axons are 
tightly bound together and are surrounded by delicate 
capillaries. Therefore the stretching of neural elements 
associated with extreme vertebral flexure, which is not 
only large of itself but also affects different segmental 
nerve roots to a greater or lesser extent, is perhaps more 
easily accommodated by shifting more of the nerve 
pathway length into the cauda equina. 

At a histological level, the spinal cord was 
found to have similar cytoarchitectural features 
characterising the laminar organization to that seen in 
the spinal cords of eutherian mammals (Ashwell and 
Zhang 1977). Spinal cord nuclei found in eutherians 
were also identified in the monotreme, except for the 
central cervical nucleus. In addition, a distinct group 
of large neurons, named the median nuclear group, 
was identified in the ventral part of Rexed’s lamina X 
and extending into the ventral funiculus at the lower 
lumbar level (Figure 1c). Fibre calibre in the dorsal 
and ventral roots of the echidna was similar to that 
reported in eutheria, suggesting similar proportions of 
afferent fibre classes and a and B motorneurons. 

The distribution of unmyelinated primary 
afferent fibres within the dorsal horn of the echidna 
spinal cord have been examined using lectin labelling 
with Griffonia simplicifolia isolectin B4. When 
conjugated with horseradish peroxidase, GSB4 is 
known to label unmyelinated primary afferents 
terminating in both the dorsal horn and cranial nerve 
sensory nuclei (Streit et al. 1985; Plenderleith et al. 
1989; Ashwell and Zhang 1997). It was seen that the 
pattern of labelling with this lectin within the spinal 
cord differed significantly from that seen in eutheria 
in several respects. Firstly, while labelling was seen 
within layers I and II of the echidna dorsal horn (similar 
to eutheria, Streit et al. 1985; Plenderleith et al. 1989), 
labelled fibre bundles were also seen coursing around 
the lateral margin of the dorsal horn as well as through 
layers I and II to terminate in deeper layers of the 
echidna dorsal horn (Figure 1d). In eutheria, lectin 
labelled primary afferents terminate only in the 
superficial layers of the dorsal horn (Streit et al. 1985; 
Plenderleith et al. 1989). This deeper labelling in the 
echidna was found to consist of discrete patches in the 
central and lateral parts of layers III] and V 
(corresponding to the nucleus proprius). Furthermore, 
in upper cervical segments of the echidna spinal cord, 
labelled axons were identified coursing around the 
margins of the dorsal columns to terminate in the 
internal basilar nucleus (Ashwell and Zhang 1997). 
These two aforementioned features reflect unusual 
primary afferent termination in the echidna, but the 
elucidation of the functional significance of these 
would require electrophysiological studies. Generally 


Proc. Linn. Soc. N.S.W., 125, 2004 


however, spinal cord cytoarchitectural organization 
seems to be highly conserved across mammals. 


CORTICOSPINAL TRACT 


The echidna corticospinal tract (Figure 2a) 
differs from other mammals (Figure 2b, c, d) in both 
its position within the brainstem and in the level at 
which it decussates (Goldby 1939). The tract runs 
through the cerebral peduncle, decussates in the pons, 
and continues in the lateral medulla, dorsal to the spinal 
tract of the trigeminal nerve. At the spinomedullary 
junction it enters the most posterior part of the lateral 
column of the spinal cord and has been traced as far 
caudally as the 24th spinal segment, which corresponds 
to lower lumbar to upper sacral levels. No evidence 
has been found for the presence of a pyramidal tract 
close to the ventral midline of the medulla, nor for a 
decussation in the usual position at the caudal end of 
the medulla, as seen in most eutheria. In no other 
mammal is the pyramidal decussation as high as in the 
echidna, nor does the tract, after decussation, lie in 
such an extreme lateral position as in this monotreme. 
It is of interest to note, however, that a high decussation 
of the pyramidal tract is particularly characteristic of 
a small number of highly specialised mammals, which 
probably developed these corticospinal specialisations 
at a very early period in mammalian evolution (Goldby 
1939). For example, some bats and edentates have a 
decussation just caudal to the pons and there is a 
tendency in some of these mammals for fibres from 
this high decussation to take up a lateral position in 
the medulla, e.g. in an armadillo, Lysiurus unicinictus, 
and the pangolin, Manis tricuspis (Goldby 1939). Since 
both of these eutherians are capable of pronounced 
vertebral flexure, as is the echidna, one is tempted to 
speculate that a high pyramidal tract decussation may 
be advantageous for mammals which use this type of 
defensive posture, although the precise nature of the 
advantage which this may confer is not clear at present. 

In polyprotodontid metatheria e.g. the 
American opossum Didelphis virginiana, the 
corticobulbar and corticospinal tracts have been shown 
to be small and probably extend no further than the 
upper cervical segments of the spinal cord (Turner 
1924, see also review by Heffner and Masterton 1983) 
and yet as noted above the pyramidal tract in the 
echidna is much more extensive. Among eutherians, 
both hedgehogs and tree shrews (Figure 2c) show 
termination of the corticospinal tract at higher 
segmental levels (upper cervical for the hedgehog and 
midthoracic for the tree shrew, for review see Heffner 
and Masterton 1983) than that seen in the echidna. 
These observations have made the extensive and 


289 


ECHIDNA CENTRAL NERVOUS SYSTEM 


unusual corticospinal pathway of the 


a) Echidna b) Marsupial (e.g. Phascolartus surviving monotremes of particular 
or Pseudochirus) interest. Extension of the corticospinal 
tract down the greater length of the spinal 
Re ; 
x cord is usually regarded as a feature of 
s advanced neurological organization, as 
re) seen in primates (Figure 2d) and 


Diencephalon 3 y f 
carnivores, because it allows direct 


control of the cerebral cortex over motor 
units within many levels of the spinal 
cord. 


BRAINSTEM AND 
HYPOTHALAMUS 


The gracile and cuneate nuclei 
are extraordinarily large in the echidna 
(Figure 3a), reflecting the well- 
developed somatosensory pathways for 
the limbs of the echidna. Furthermore, 
estimates of the proportion of white 
matter in the dorsal columns to total 
white matter in the cord gave an average 
result of 25%, suggesting well-developed 
trunk and appendicular somatosensory 
pathways comparable in development to 
carnivores and primates (Ashwell and 
Zhang 1997). In absolute terms the dorsal 
column pathway is as large as that in the 
domestic cat and Macaca fuscata - 
therians of similar body weight. This 
degree of development of the dorsal 
columns ranks the echidna among the 
most neurologically specialized primates 
with well-developed discriminative 
tactile sense. Perhaps the high level of 
somatosensory development can be 
attributed to dense innervation of the 
echidna’s forelimb (Mahns et al. 2003) 


an7x0o0 


c) Edentates and Chiroptera d) Human 


Medulla 


Figure 2 Diagrammatic summary of the course, size and extent of the corticospinal tract (bold) in 
representative mammals. Note that the corticospinal tract in the echidna is large, has a high decussation 
and extends to caudal levels of the spinal cord. Contrast this with the small size of the corticospinal tract 
in marsupials (b) and bats and edentates (c) and restriction of the tract to upper segmental levels of the 
spinal cord in those mammals. In size and extent, the echidna corticospinal tract is more like that seen in 
primates (d) and carnivores: mammals in whom a long and large corticospinal tract is believed to confer 
neurological advantages in the form of direct cortical control of motor units in the spinal cord. The 
corticospinal tract of the echidna also has a relatively high level of decussation (crossing over) compared 
to therian mammals, although some mammals (e.g. edentates and chiroptera - c) with the ability to flex 
their vertebral column also have a high level of decussation. No undecussated ventral corticospinal tract, 
as seen in primates (d) has been reported in the echidna. Data for the echidna is derived from Goldby. 
(1939), while data for other mammals comes from Kappers, Huber and Crosby (1960). 


290 Proc. Linn. Soc. N.S.W., 125, 2004 


M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 


Figure 3. Coronal cryostat section (40 um thickness) through the brainstem of an echidna stained for 
cytochrome oxidase by the Wong Riley technique (Wong-Riley 1979)(a, b) and Nissl substance (c, d). 
Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal acquisition. Strong 
cytochrome oxidase reactivity demonstrates the presence of high densities of mitochondria in axon 
terminals of major sensory pathways for limb and trunk somatosensory pathways (e.g. cuneate nucleus) 
and cranial somatosensory pathway (e.g. nucleus of the trigeminal spinal tract). The mesencephalic nucleus 
of the trigeminal nerve occupies a midline position dorsal to the cerebral aqueduct. The inset in c indicates 
the position of d. 3 — oculomotor nucleus; 4v — fourth ventricle; 12 — hypoglossal nucleus; Aq — cerebral 
aqueduct; Cu — cuneate nucleus; [O — inferior olivary nuclear complex; mcp- middle cerebral peduncle; 
Pn — pontine nuclei; SC — superior colliculus; t5 — trigeminal spinal tract; Vc —caudal part of the nucleus 
of the trigeminal spinal tract ; Ve — vestibular nuclei; Vmes — mesencephalic nucleus of the trigeminal 
nerve; Vo -oralis part of the nucleus of the trigeminal spinal tract. 


or spines, although this has never been studied 
histologically. Alternatively, this specialization may 
have arisen because the echidna spends time in 
subterranean channels, where visual and auditory input 
are of little benefit, and the sense of smell and touch 
are of the most value. At present there are no 
morphological studies of echidna postcranial tactile 


Proc. Linn. Soc. N.S.W., 125, 2004 


receptors available to shed light on this. 

The trigeminal nerve is also greatly enlarged 
in the echidna as are the nuclei of the trigeminal spinal 
tract (Figure 3a, b). This is consistent with the 
impression from behavioural and electroreception 
studies that the echidna’s snout is extremely sensitive 
(see above). The trigeminal system in the echidna 


291 


ECHIDNA CENTRAL NERVOUS SYSTEM 


displays a high degree of specialisation similar in kind 
to that seen in Ornithornychus, but not to such a large 
extent. In other words, it does not appear to be as 
sensitive an electroreceptive tool as the platypus bill 
(Proske et al. 1998). Another note-worthy feature is 
that the motor nucleus of the trigeminal nerve in the 
echidna brainstem is much larger than would be 
expected in an animal whose jaw musculature is so 
poorly developed (Abbie 1934). 

The mesencephalic nucleus of the fifth nerve 
in echidna is very like that seen in reptiles in that it 
adopts an almost exclusively mid-line distribution 
(Abbie 1934, Figure 3c, d). Metatheria exhibit a 
condition intermediate between that of the echidna and 
eutheria with more extensive development of the lateral 
mesencephalic V extensions. The mesencephalic 
nucleus and root of the fifth nerve are generally 
considered as being concerned with proprioceptive 
sensibility of jaw musculature. Since the echidna has 
very poor jaw musculature, such a pronounced 
development of mesencephalic V is inexplicable. 

The echidna auditory and vestibular apparatus 
are also notable. In Ornithorhynchus, the entire 
labyrinth has been described as being typically avian 
(Gray 1908). In the echidna, the inner ear shows 
dissimilarities to therians, in that the echidna cochlea 
is banana shaped and has only half a turn, hence is 
partially coiled, whereas in humans the cochlea has 
two and a half turns, and is fully coiled (Gray 1908). 
The cochlea also shows maximal response to sound of 
about 5kHz, substantially lower than in eutheria 
(Augee and Gooden 1993). It has been proposed that 
when the cochlea evolved from the primitive labyrinth, 
it employed the existing vestibular connections within 
the brain, and that when the cochlear apparatus attained 
a mammalian level of structural specialization, a 
trapezoid body appeared in the brainstem. The 
trapezoid body in the echidna is so rudimentary that it 
reveals its primitive vestibular and primarily trigeminal 
origin, because it consists almost entirely of vestibular 
parts and external arcuate fibres which include a large 
trigeminal element (Winkler 1921; Abbie 1934). In 
therians, the pronounced increase in auditory fibres 
almost completely obscures the original trigeminal and 
vestibular connection. Winkler (1921) has argued that 
the poor cochlear development in the echidna renders 
the vestibular fibres relatively conspicuous. While 
central auditory pathways have never been closely 
examined in the echidna, these observations suggest 
that those pathways are either organized differently or 
not as extensive as in theria. 

The hypothalamus in the echidna has been 
reported to have few striking features (Abbie 1934). 
The mammalian hypothalamus is very old 


phylogenetically (Simerly 1995) and very conservative 
in structure throughout the vertebrate series. One 
peculiarity, which links the echidna hypothalamus with 
that of reptiles, and is in sharp contrast to the majority 
of mammals, is the extremely poor development or 
possible absence of the echidna mammillothalamic 
tract (Abbie 1934). Regidor and Divac (1987) for 
example found no evidence of the mammillothalamic 
tract in the echidna on examination of myelin-stained 
coronal sections. This pathway is a key link in the 
Papez circuit underlying memory and emotions in 
eutheria. Its poor development in the echidna may 
indicate that this mammal has an alternative circuit 
for these functions. 


THE ABSENCE OF A CLAUSTRUM IN THE 
FOREBRAIN 


The absence of a claustrum in the echidna 
was initially noted by Abbie (1940) and by Divac and 
co-workers (Divac et al. 1987a) and was further 
discussed more recently (Butler et al. 2002). Similarly, 
no claustrum has been identified in the platypus brain 
(Butler et al. 2002). This structure has been identified 
in all therian mammals so far examined (Johnson et 
al. 1994) and is believed to have a structural and 
chemical affinity with the neocortex, although its 
precise functional significance is uncertain. It engages 
in reciprocal connections with neocortex and receives 
projections from the non-specific intralaminar nuclei 
of the thalamus (see Butler et al. 2002 for review). 
The question remains open as to whether the claustrum 
was present in ancestral mammals and disappeared in 
the monotremes, or whether its evolution represents 
an exclusively therian brain development. 


THE DISTRIBUTION OF CHEMICALLY 
IDENTIFIED NEURONS 


Manger and co-workers have recently 
examined the distribution of cholinergic, 
catecholaminergic and serotonergic neurons in the 
brains of the platypus and echidna (Manger et al. 
2002a, b, c). Those authors showed that while there 
are many similarities between monotremes and therians 
in the distribution of these neurons, there were also_ 
some evolutionarily and potentially functionally 
significant differences. For example, cholinergic cells 
are present in the monotreme brain, but important cells 
groups identified in theria do not appear to be present 
in the platypus or echidna. These include cholinergic 
cells in the cerebral cortex, nuclei of the vertical and 


Proc. Linn. Soc. N.S.W., 125, 2004 


M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 


Figure 4. Coronal cryostat sections (40 um thickness) through the caudal thalamus (a) and rostral thalamus 
(b) stained for cytochrome oxidase and Nissl substance, respectively. Please see Hassiotis and Ashwell 
(2003) for details of experimental ethics and animal acquisition. Note the large ventral posterior thalamic 
nucleus with lateral (VPL) and medial (VPM) compartments. In theria these two regions serve processing 
of somatosensory (touch) information from the body and head, respectively. Contrast the size of these 
two nuclei with the lateral geniculate nucleus (LG) processing visual information. The reticular thalamic 
nucleus, which is found in all therian mammals external to the ventral tier nuclei (i.e. embedded in the 
external medullary lamina to the left of 4a), appears to be absent from the echidna thalamus. The rostral 
thalamus (b) contains a large nucleus (anteromediodorsal -AMD) with no clear division into subnuclei. 
This may correspond to the mediodorsal nucleus of therians. 3v — third ventricle; eml — external medullary 


lamina; ml — medial lemniscus; ZI — zona incerta. 


horizontal limbs of the diagonal band of Broca, the 
magnocellular preoptic nucleus, the substantia 
innominata, nucleus of the ansa lenticularis, 
hypothalamic nuclei and the parabigeminal nucleus 
(Manger et al. 2002a). They proposed that the absence 
of cholinergic neurons from the hypothalamus might 
be related to the unusual features of monotreme sleep 
(Siegel et al. 1996, 1998). 

The catecholaminergic system of the 
monotreme brain appears to be very similar to that 
found in theria, but there were some minor differences 
in the form of the absence of A4, A3 and C3 groups 
from the locus coeruleus and caudal rhombencephalon. 
It should be noted however, that these are only small 
differences and this great similarity demonstrates the 
high degree of evolutionary conservatism in these 
neurons across amniote species (Manger et al. 2002b). 


Proc. Linn. Soc. N.S.W., 125, 2004 


Serotonergic neurons in monotremes appear 
to fall into three groups: hypothalamic, rostral nuclear 
and caudal nuclear clusters. The rostral and caudal 
nuclear groups are found consistently across all 
mammals while the hypothalamic cluster, although not 
reported in other mammals, is found in most other 
species of vertebrates (Manger et al. 2002c). 


THE THALAMUS AND THALAMOCORTICAL 
PROJECTIONS 


Campbell and Hayhow (1971) identified 
several thalamic nuclei in echidna, which exhibited 
cyto- and myeloarchitectonic features resembling those 
found in other mammals (Figure 4). However, echidna 
thalamic nuclei are not as easily distinguished as those 


293 


ECHIDNA CENTRAL NERVOUS SYSTEM 


in opossums (Bodian 1939, 1942; Oswaldo-Cruz and 
Rocha-Miranda 1968; Benevento and Ebner 1971) or 
other commonly used laboratory mammals (Rose 
1942; Rose and Woolsey 1949). Chemoarchitectural 
characteristics of the thalamus in echidnas and rats have 
been compared in sections stained for myelin, 
acetylcholinesterase (AChE), succinate dehydrogenase 
(SDH) and cytochrome oxidase (CO) by Regidor and 
Divac (1987). Numerous species differences were 
noted, but in general the thalamus is architecturally 
more homogenous in echidnas than in rats, especially 
within the anterior portion (Figure 4b). The large 
structure localized in the anteromediodorsal part of the 
thalamus of the echidna has been found to contain small 
amounts of acetylcholinesterase and oxidative 
enzymes; in this respect resembling the mediodorsal 
nucleus of rats. Regidor and Divac (1987) concluded 
that this brain structure of echidnas corresponds to the 
mediodorsal nucleus in placental species. 

Welker and Lende (1980) defined and 
described the thalamic nuclei that contribute major 
projections to the isocortex in echidna. Their purpose 
was to determine whether the echidna thalamus 
exhibited mammalian thalamocortical relations more 
similar to those found in metatheria, or to those in 
eutherian mammals. Welker and Lende also attempted 
to identify whether an enlarged thalamic nucleus was 
sending afferents to the enlarged frontal cortex. They 
performed a series of partial ablations of the somatic 
sensory, auditory, visual and motor areas, as well as 
in several different portions of the greatly enlarged 
frontal neocortex (see below) and demonstrated that 
the thalamocortical connections in the echidna are 
similar in most respects to those demonstrated in 
eutherian mammals. One unusual feature observed by 
Welker and Lende was a large nuclear mass in the 
dorso-fronto-medial thalamus (presumptive 
anteromediodorsal nucleus discussed above), which 
projects to the enlarged frontal cortex (Divac et al. 
1987a, b). It has been hypothesised that this nuclear 
region is homologous to the eutherian mediodorsal 
nucleus. Their data also revealed that projections to 
separate motor and somatic sensory cortical areas from 
the thalamus were spatially distinct (Welker and Lende 
1980). 


CORTICAL STRUCTURE AND 
ORGANIZATION 


Until the late 1800's it was generally believed 
that all mammals possessed a corpus callosum (Turner 
1890), a major fibre bundle connecting the neocortex 
of the two hemispheres of the brain. Elliott Smith 


294 


(1902, 1903) dispelled this notion in his early studies 
of comparative cortical organization. He showed that 
in monotremes and metatheria, the anterior 
commissure is the major cerebral commissure, being 
the sole connection between all parts of the neo- and 
paleocortex, with only a small archicortical 
commissural connection being present dorsally (the 
hippocampal commissure). 

Several striking aspects of gross cortical 
anatomy have been noted in Tachyglossus aculeatus. 
The most obvious of these is the high degree of 
gyrification (36% of isocortex buried in fissures), 
comparable to that in many eutherian mammals (e.g. 
cat 40%, squirrel monkey 39%). The second is the large 
proportion of the brain volume occupied by the 
cerebral cortex (43%), similar to values in eutheria 
(prosimians — 54%, Pirlot and Nelson 1978). Among 
the brains of eutheria, a highly gyrified cerebral cortex 
is usually considered as an attempt to maximise the 
number of cortical columns available for the processing 
of information. Therefore a highly gyrified cortex is 
considered the hallmark of more neurologically 
advanced mammals such as carnivores, primates and 
cetaceans. This raises the question as to why an animal 
like the echidna, which leads a solitary existence and 
has no known complex social life, has such a highly 
gyrified cortex. One principal difference between the 
brains of the two living Australian monotremes is that 
the platypus’ cortex is quite smooth (lissencephalic), 
whereas the echidna cortex is complexly folded. 

Another most remarkable aspect of echidna 
neurobiology concerns cortical topography. Ziehen 
(1897, 1908), Brodmann (1909) and Schuster (1910), 
all published early observations on the cortex in the 
Monotremata. Brodmann examined the cortex in 
echidna and established that there is a typical six- 
layered distribution. The echidna has been noted to 
have a thinner cortex, perhaps due to its denser packing 
of neurons compared to the platypus (Abbie 1940). In 
both the echidna and the platypus, Ziehen (1897, 1908) 
showed that there was a change in the type of cortex 
between the anterior (olfactory) and posterior 
(sensorimotor) portions of the hemisphere, and 
Schuster (1910) confirmed his observations. 
Nevertheless, these early authors concluded that the 
plan upon which the monotreme brain is constructed 
conforms in every respect to the basic pattern 
prevailing among the vast majority of other mammals 
(Abbie 1940). 

To date, the most detailed anatomical study 
of the echidna cortex was performed in the 1940’s by 
Abbie. Since the 1940’s, no further in-depth anatomical 
studies have been done on the anatomy of the echidna 
cortex as a whole, although specific systems have been 


Proc. Linn. Soc. N.S.W., 125, 2004 


M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 


Welker and Lende (1980) 


Krubitzer et al. (1995) 


Rostral 


Figure 5. Electrophysiology and cytoarchitecture of the echidna cerebral cortex. The earliest study 
illustrated is by Lende (1964)(results shown in a summary diagram redrawn from Welker and Lende 
1980). Greek letters denote major consistent sulci as delineated by Smith (1902). The Welker and Lende 
map shows only the externally visible gyral surface and indicates the position of motor (M), somatosensory 
(S1), visual (V) and auditory (A) cortices. The Ulinski map shows cytoarchitectural fields identified by 
that author (Ulinski 1984). Two coronal sections (i and ii) are illustrated with the positions of rostral (r) 
and caudal (c) fields of the somatosensory cortex (SM1) marked. The small lateral view shows the 
rostrocaudal positions from which these sections were taken. The figure below the two sections shows a 
flattened representation of cortex. Note that Ulinski’s “rv” field lies rostral (and superior) to the deepest 
part of the o sulcus. The lower two illustrations summarize the findings of Krubitzer et al (1995). The 
smaller diagram shows a representation of the entire flattened cortical surface with the sulcal walls opened. 
Solid lines in the Krubitzer map indicate the deepest point of the sulci, while dotted lines indicate the 
sulcal rims at the external cortical surface. The larger illustration shows a drawing of the completely 
flattened cortical surface with the boundaries of functional areas indicated relative to B and o sulci (thick 
grey lines). Note that the rostral somatosensory field in Krubitzer’s map lies caudal to the @ sulcus (cf 
Ulinski map). Ent — entorhinal cortex; Hi — hippocampus; M — manipulation cortex; Pir — piriform cortex; 
PV — parietal ventral somatosensory cortex; R -rostral somatosensory cortex; S1 — primary somatosensory 
cortex. 


Proc. Linn. Soc. N.S.W., 125, 2004 295 


ECHIDNA CENTRAL NERVOUS SYSTEM 


studied. Abbie described the monotreme neocortex as 
comprising two broad components; one related to the 
hippocampus, labelled by him as parahippocampal 
regions and located in the anterior and medial parts; 
and the other related to the piriform cortex, labelled as 
the parapiriform regions, located posteriorly and 
laterally. He also defined sulcal boundaries to these 
regions. When labelling the cortex, Abbie adopted the 
system of Elliot Smith (1902), using Greek letters to 
name the major and deepest sulci (Figure 5). There 
are two pronounced sulci in the monotreme cortex, 
denoted as and B. These divide the frontal cortex 
from the posterior motor and sensory cortices. 

More recent functional studies of the 
isocortex of Tachyglossus aculeatus have indicated that 
the primary motor, somatosensory, auditory and visual 
areas are located in the caudal half of the isocortex 
(Lende 1964)(Figure 5). Aside from the posterior 
location of these areas, the following relationships are 
unlike those described in any other therian mammals: 
the somatic sensory area is confined to the ventral 
portion of the lateral surface; the visual area is located 
dorsal to the somatic sensory area and borders the 
representation for the tail; the auditory area is located 
posterior to the visual and somatic sensory areas and 
borders the latter at the representation of the back. 
These relationships might be described as rotational 
dislocation of the areal relations relative to that found 
in eutherians in that the somatic sensory area has been 
displaced downward and backward, the auditory area 
upward, and the visual area upward and forward 
(Lende 1964). 

The somatosensory representation in the 
echidna is in some respects similar to that of other 
mammals. The area for the tail is found uppermost 
and the areas for hind limb, trunk, forelimb, and head 
are located laterally and ventrally, in that sequence. 
This is the same as the basic mammalian pattern of 
somatosensory area | (S1) as established by Woolsey 
(1952). A relatively large portion of somatosensory 
cortex in the echidna was found by Lende to be devoted 
to the head, and particularly the snout and tongue, as 
might be expected from the ant-eating habits of the 
echidna (Lende 1964). 

Physiological studies have indicated that 
more than 50% of the rostral cortex of the echidna has 
no attributable primary motor or sensory function and 
has been considered as an expanded prefrontal cortex 
(Welker and Lende 1980). If this interpretation is 
correct, then the proportion of isocortex in 
Tachyglossus aculeatus occupied by the prefrontal area 
exceeds that in humans (29%) Divac et al. (1987a, 
b)(see section on thalamus and thalamocortical 
projection in previous pages). 

Ulinski’s study (1984) examined the 


296 


cytoarchitecture and thalamic afferents of the 
somatosensory area (SMI) in the echidna. His findings 
indicated that SMI contains two cytoarchitectonic 
fields. A caudal field with a well-developed layer IV 
present extends across the post ® gyrus and onto the 
floor of sulcus @ The rostral field was reported to 
extend from the floor of sulcus & onto its rostral bank. 
It also was reported to have a well-developed layer IV 
but with a large number of pyramidal neurons in layer 
V. The remainder of the pre & gyrus was reported to 
contain a single cytoarchitectonic field with a thin layer 
IV and layer V heavily populated with larger pyramidal 
cells. This field corresponded to the physiologically 
defined motor area M1. Thalamic afferents to 
somatosensory area were examined by placing pressure 
injections of horseradish peroxidase into the two 
architectonic fields. The results indicated that the 
somatosensory area in Tachyglossus aculeatus contains 
two cytoarchitectonic fields that resemble areas 3a and 
3b in some placental mammals, leading Ulinski to the 
conclusion that the collection of cytoarchitectonic 
fields corresponding to areas 4, 3a, and 3b is a basic 
mammalian character which has been modified in 
metatherian and many eutherian mammals. 

In more recent times, Krubitzer et al. (1995) 
undertook a detailed study of monotreme cortical 
organization as part of a comparative approach to 
determining those features of the isocortex which 
characterise all the major lines of mammalian 
evolution, More specifically, their investigation was 
designed to determine the internal organization and 
number of somatosensory fields in the monotreme 
isocortex. 

The isocortices of both monotremes were 
found to contain four representations of the body 
surface. A large area that contained neurons 
predominantly responsive to cutaneous stimulation of 
the contralateral body surface was identified as the 
primary somatosensory area (S1). This was found 
caudal and ventral to the @ sulcus. Another 
somatosensory field (R) was identified rostral to S1. 
The topographic organization of R was similar to that 
found in S1, but neurons in R were responsive most 
often to light pressure and taps to peripheral body parts. 
Neurons in cortex located rostral to R were responsive 
to manipulation of joints and hard taps to the body. 
This field was termed the manipulation field (M) and 
occupies the position of the motor cortex identified 
by Lende. Note that Krubitzer’s M field occupies an 
area which Ulinski denoted as the rostral 
somatosensory field and Krubutzer’s R field occupies 
at least part of Ulinski’s caudal somatosensory field 
(Ulinski 1984)(Figure 5). Consequently the two studies 
are not easily reconciled. A parieto-ventral’ 
somatosensory field (PV) was also identified by 


Proc. Linn. Soc. N.S.W., 125, 2004 


M. HASSIOTIS, G. PAXINOS AND K.W.S ASHWELL 


Figure 6. Nissl stained cryostat sections in the coronal plane through the cerebral cortex and olfactory 
bulb of the echidna. Please see Hassiotis and Ashwell (2003) for details of experimental ethics and animal 
acquisition. The inset drawings show: i) a lateral view of the echidna cerebral hemisphere indicating the 
planes of section shown in a, b, c and d, e, respectively; and ii) a line drawing of a coronal section showing 
the positions from which a, b, and c were taken. Figure 6a shows a lower power view of motor cortex (M) 
and the rostral field of somatosensory cortex (R). Figures 6b and c show motor cortex and S1 somatosensory 
cortex, respectively with layers indicated by Roman numerals. WM - subcortical white matter. Figures 
6d and e show low and high power views of the olfactory bulb. Rectangle in d indicates the position of e. 
Note the lack of a clear tightly grouped monolayer of mitral cells (Mi) as is seen in therian mammals 
(Switzer and Johnson 1977). E — ependyma of lateral ventricle; Gl — glomerular layer; [Gr — internal 
granular layer; IPL — internal plexiform layer; ON — olfactory nerve fibre layer; Pir — piriform cortex. 


Proc. Linn. Soc. N.S.W., 125, 2004 wg) 


ECHIDNA CENTRAL NERVOUS SYSTEM 


Krubitzer and was thought to be homologous to its 
therian counterparts (Krubitzer et al. 1995). The 
evidence for the existence of four separate 
somatosensory representations in somatosensory 
cortex was taken to indicate that cortical organization 
is more complex in the echidna than had been 
previously thought. Furthermore, although the two 
monotreme families have been quite separate for at 
least 55 million years (Richardson 1987), the similarity 
of cortical field organization in both monotremes 
studied suggested either that the original differentiation 
of sensory fields occurred very early in mammalian 
evolution, or that the potential for division of 
somatosensory cortex into numerous fields was highly 
constrained in evolution, so that both species arrived 
at the same result independently. 

Figure 6 shows the cytoarchitecture of the 
echidna motor and somatosensory cortices. 
Nomenclature for cortical areas is adopted from 
Krubitzer et al (1995). As in eutherian motor cortex 
(Figure 6b), the echidna M cortex is characterised by 
large pyramidal neurons in layer V. The S1 part of 
somatosensory cortex (Figure 6c) is characterised by 
a layer IV rich in densely packed small neurons. 

Several other aspects of cortical function in 
this species are also of note; particularly the apparent 
absence of an SII somatosensory representation and 
the relatively lateral position of the motor 
representation compared to that in eutheria (Krubitzer 
etal. 1995). Until recently, functional studies had failed 
to identify any parietal association cortex, but 
Krubitzer et al. (1995) was able to identify a 
topographically discrete, multimodal area between the 
primary sensory cortical areas, which may represent 
such an area. 


CONCLUDING REMARKS 


There are a number of unusual features of 
the anatomy of the brain and spinal cord of the echidna. 
Some of these are probably indicative of the early 
branching of the monotreme lineage from therian 
mammals (e.g. absence of the claustrum), some have 
potential functional significance for the unusual 
physiology of this mammal (e.g. the absence of 
hypothalamic cholinergic neurons), while some appear 
to be peculiar adaptations to the niche occupied by 
this mammal (e.g. trigeminal nuclei development and 
short spinal cord). 


298 


ACKNOWLEDGEMENTS 


This work was supported in part by a grant from the 
Australian Research Council. We are grateful for the 
assistance of Drs Luan ling Zhang and Hong gin Wang 
in the preparation of the Nissl, cytochrome oxidase 
and lectin stained sections through the echidna brain 
and spinal cord. 


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Proc. Linn. Soc. N.S.W., 125, 2004 


Monotreme Tactile Mechanisms: From Sensory Nerves To 
Cerebral Cortex 


Mark J. Rowe, D.A. MAHNS AND V. SAHAI 


School of Medical Sciences, The University of New South Wales, Sydney 2052, Australia 


Rowe, M.J., Mahns, D.A. and Sahai, V. (2004). Monotreme tactile mechanisms: from sensory nerves to 
cerebral cortex. Proceedings of the Linnean Society of New South Wales 125, 301-317. 


Electrophysiological recordings from single tactile sensory nerve fibres supplying the limb extremities in 
the echidna (Tachyglossus aculeatus) reveal a remarkable resemblance between monotreme peripheral tactile 
mechanisms and those of placental mammals. The similarities apply to a concatenation of attributes, including 
the classification of sensory fibre types and aspects of functional properties and tactile coding capacities. 
The analysis demonstrates that high-acuity tactile signalling from the distal forelimb in the monotreme is 
based upon a triad of major tactile fibre classes as is the case for placental mammals. Furthermore, the 
functional similarity between corresponding classes in monotreme and placental species suggests that 
peripheral tactile sensory mechanisms are highly conserved across evolutionarily-divergent mammalian 
orders. 

Evidence for a unique and striking dependence upon tactile sensory mechanisms in monotremes comes 
from both behavioural observations on the animals and from the exceptional prominence given to the 
representation of tactile inputs in the cerebral cortex of these species. In the platypus, for example, almost 
half of its lissencephalic cortex is allocated to the processing of inputs from the bill. Furthermore, within the 
specialized area of bill representation in the platypus cortex, the receptive fields of individual neurones are 
amongst the smallest ever recorded within the somatosensory areas of cortex (often <lmm in diameter), 
presumably conferring great fidelity and precision on the tasks of tactile localization and discrimination 
involving the bill. However, in both the platypus and the echidna there is a complete and highly ordered 
somatotopic representation of tactile inputs from the contralateral body surface, conforming with the so- 
called primary somatosensory cortex (SI) of other mammals. Controversy applies to the issue of whether 
additional, multiple body representations are present in the monotreme cortex, as neither Lende (1964) nor 
Bohringer and Rowe (1977) found evidence of this, in contrast to Krubitzer et al. (1995a), who have argued 


for four body representations in the cortex of both the echidna and the platypus. 


Manuscript received 24 September 2003, accepted for publication 22 October 2003. 


KEYWORDS: echidna, evolution and sensory nerve function, monotreme, Ornithorynchus anatinus, 
platypus, somatosensory system, Tachyglossus aculeatus, tactile receptors, tactile sensory function. 


EVOLUTIONARY PLACE OF THE 
MONOTREME BRAIN 


The emergence, 100-200 million years ago, 
of monotremes on a separate evolutionary line from 
their placental and marsupial mammalian counterparts 
has contributed, in particular in the 19" century, to the 
hypothesis that living monotremes are closer to 
ancestral mammalian forms than their placental and 
marsupial relatives, and therefore that the monotreme 
nervous system may provide a window on the status 
of the ancestral mammalian brain. These notions were 
implicit in the designation of monotremes as the 
Prototheria, or first mammals, while the marsupial 
mammals were designated the Metatheria and were 


thought to represent the next stage of evolution towards 
the Eutherian, or complete mammals. This hierarchical 
concept of the relations between the three great orders 
of mammals was, in part, influenced by the presence 
in living monotremes of reptilian or avian-like 
reproductive mechanisms. This possession of 
Oviparity, or the capacity to lay eggs, set the 
monotremes aside from mammals of the marsupial and 
eutherian orders. Furthermore, a concatenation of other 
anatomical or functional attributes shared with reptilian 
and avian representatives, but not with other mammals, 
tended to re-inforce these hierarchical concepts. These 
attributes have been documented by Augee and 
Gooden (1993) and include, among others, the 


MONOTREME TACTILE MECHANISMS 


arrangement of the shoulder girdle, the less elaborate 
coiling of the cochlea, and the presence of dwarf 
nephrons in the kidney. Perhaps surprisingly, the 
presence of the single opening from the cloaca for both 
reproductive and excretory activity, which led to their 
monotreme designation, is not entirely diagnostic for 
this mammalian group as some marsupials and other 
vertebrates share this feature (Augee and Gooden 
1993). 

Despite the retention of several plesiomorphic 
attributes in living monotremes there should be no 
expectation that such attributes need be a generalized 
feature of this mammalian order. Thus, in other respects 
of their anatomy or physiology, monotremes need not 
have proved any more conservative or constrained in 
their evolutionary progress than their marsupial or 
placental relatives with whom they have shared 
perhaps 200 million years in which to evolve from the 
ancestral forms of their evolutionary forebears. Indeed, 
we see some hint of this evolutionary metamorphosis 
and divergence within the monotreme ranks 
themselves, even in gross features of central nervous 
system organization. For example, although the 
divergence of the monotreme line into separate 
platypus and echidna strands has occurred ~50-80 
million years ago (Griffiths 1978; Dawson 1983; 
Richardson 1987), or even as recently as ~20 million 
years ago (Belov and Hellman 2003), one encounters 
striking differences between them in the gross 
morphology of the cerebral cortex. The echidna has a 
quite elaborately folded, or gyrencephalic cerebral 
cortex, whereas, in contrast, the platypus possesses a 
smooth, essentially unfolded, lissencephalic cortex. 
The elaborate pattern of the cortical fissures in the 
echidna was analyzed and classified by Elliot Smith 
(1899, 1902) according to a scheme of Greek 
characters. The most consistent fissures form a series 
of mediolaterally oriented sulci designated , B and 6, 
with other less prominent and less consistent sulci 
being present (Elliot Smith 1899, 1902; Hines 1929; 
Burkitt 1934; Lende 1964). The only departure from 
lissencephaly in the platypus cortex is the presence of 
a slight and shallow rhinal sulcus on the ventral surface 
of the cerebrum (Elliot Smith 1902). 

The divergence within monotreme evolution 
that has given rise to the lissencephalic and 
gyrencephalic forms of the cerebral cortex in the 
platypus and echidna is similar to that seen within both 
the placental and marsupial orders. Among the 
placental mammals, lissencephaly is apparent in the 
cortex of the rat and rabbit and even in some primates, 
such as the marmoset monkey (Callithrix jacchus; 
Brodmann 1909; Rowe et al.1996; Zhang et al. 1996, 
2001), whereas in the cat, the human being, and in a 


302 


great many other placental mammals the cortex has 
acquired a prominent gyrencephalic form. However, 
the fact that the echidna and platypus display such 
striking differences in cortical folding serves as a clear 
reminder that the monotremes have had the same 
opportunity for evolutionary change and adaptation 
as have the species within the placental or marsupial 
orders. 


Quantitative indices of brain development in 
monotremes and other mammals 

A variety of quantitative measures of brain 
development also reinforce the conclusion that the 
brain of living monotremes is no more likely to provide 
a guide to the status of the ancestral mammalian brain 
than that of any contemporary placental mammal (for 
review, Rowe 1990). Furthermore, such measures fail 
to identify the living monotremes as being 
systematically less advanced in neurological terms than 
many of their marsupial and placental relatives. The 
quantitative measures invoked for these comparisons 
have been based usually upon indices related to the 
ratio of brain mass to total body mass and include, for 
example, the Encephalization Quotient (EQ) put 
forward by Jerison (1973) and defined for a given 
species as the ratio of actual brain mass (E,) divided 
by the expected brain mass (E.) for a mammalian 
species of a given body mass (P;). The expected brain 
mass was obtained from the equation: 


E, = kP3 


that governs the overall relation between brain mass 
and body mass for the large range of living mammals 
for which Jerison gathered data. An EQ value of 1 
was assigned by Jerison for the average living placental 
mammal with values varying by a factor of 
approximately 25 times, from a low of ~0.25 for basal 
insectivores, such as the hedgehog (Erinaceus 
europaeus) and the Madagascan tenrecs, through to 
values of ~6 - 7 for human beings and cetaceans such 
as the bottlenose dolphin (Tursiops truncatus). Values 
of ~0.5 - 0.75 along the EQ scale were assigned to the 
echidnas by Jerison (1973) who argued that they “are 
well in advance of didelphids like the opossum or 
insectivores like the hedgehog in relative brain size 
and differentation”. He argued that the monotremes 
should be considered, in terms of brain development, 
to be “at almost the same level as living progressive 
mammals” and speculated that they had reached this 
level by parallel evolution. 

It must be emphasized that comparisons of 
brain development based upon measures such as the 
EQ are somewhat arbitrary and that relativities arrived 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAI 


at across species may change if other measures were 
to be used. However, a variety of alternative measures, 
including for example, the relative extent of the 
neocortex, once again fail to identify monotremes as 
being distinctly less developed neurologically than 
eutherian mammals (Pirlot and Nelson 1978; for 
review Rowe 1990). In addition, behavioural tests of 
discriminative learning in the echidna reveal an ability 
in particular, in spatial and positional discrimination 
tasks, that is not inferior to eutherian or metatherian 
species (Buchmann and Rhodes 1978). 


SENSORY AND PERCEPTUAL MECHANISMS 
IN MONOTREMES 


Behavioural observations from as early as the 
19" century have pointed to the pre-eminence of the 
tactile sense in the perceptual life of both the platypus 
and the echidna (for review: Rowe 1990; Rowe et al. 
2003). In the case of the platypus it was apparent that, 
for both navigation and feeding, the animal relies, 
perhaps exclusively, upon sensory information from 
the bill as its eyes, nose and external auditory canals 
remain closed in the course of swimming and foraging 
(Bennett 1877; Burrell 1927; Griffiths 1978; Grant 
1984). While vision may be of more importance in the 
echidna than the platypus (see Gates 1978) it is 
probable that it assumes a lesser importance than the 
trigeminal tactile inputs from the snout and tongue 
which appear to play a similar prominent sensory role 
to that of the bill in the platypus. However, tactile 
information from the limb extremities, in particular, 
from the forelimb, also assumes great importance in 
the digging and burrowing activities of the echidna 
(Griffiths 1978; Augee and Gooden 1993). Because 
of this prominent sensory role for the distal forelimb 
as a tactile exploratory organ in the echidna we have 
recently undertaken an analysis of tactile neural 
mechanisms associated with the forepaw in 
Tachyglossus aculeatus (Mahns et al. 2003; Rowe et 
al. 2003). As tactile neural mechanisms have been most 
intensively investigated for the distal glabrous skin of 
the forelimb in a great variety of species, this analysis 
permitted a comparison with placental mammals 
enabling us to establish the extent to which 
correspondence or divergence had arisen over the 
separate evolutionary paths taken within the different 
mammalian orders. 


Peripheral tactile neural mechanisms associated 
with the distal forelimb in the echidna: 
comparison with placental representatives 

Tactile sensory nerve fibres that arise from 


Proc. Linn. Soc. N.S.W., 125, 2004 


the distal glabrous skin of the limbs in the cat and in a 
variety of Old- and New-World monkeys fall into three 
major functional classes (Lindblom 1965; Lindblom 
and Lund 1966; Janig et al. 1968, Talbot et al. 1968, 
Iggo and Ogawa 1977, Ferrington and Rowe 1980; 
Ferrington et al. 1984; Coleman et al. 2001). These 
include one broad class that responds to static skin 
displacement with a so-called slowly adapting (SA) 
pattern of response that provides the basis for their 
designation as the SA class of fibres which, in both cat 
and primate species, appears to be associated with 
Merkel receptor endings (Janig et al. 1968, Talbot et 
al. 1968; Iggo and Muir 1969; Jaénig 1971; Iggo and 
Ogawa 1977; Ferrington and Rowe 1980; Coleman et 


al. 2001). 


The remaining tactile sensory nerve fibres 
supplying the primate hand or cat footpads are sensitive 
to only the dynamic components of tactile stimuli and 
can be divided into two distinct classes according to 
their sensitivity and responsiveness to cutaneous 
vibration (Janig et al. 1968; Talbot et al. 1968; Johnson 
1974; Iggo and Ogawa 1977; Ferrington and Rowe 
1980; Ferrington et al. 1984; Coleman et al. 2001). 
One class, designated the rapidly adapting (RA) or 
quickly adapting (QA) class is most sensitive to 
vibration at ~20-50Hz, and appears to be associated 
with intradermal, encapsulated receptors known as 
Krause corpuscles in the cat (Jaénig 1971; Iggo and 
Ogawa 1977) and as Meissner corpuscles in primates 
(Talbot et al. 1968; Coleman et al. 2001). The other 
purely dynamically-sensitive class (the PC class) is 
exquisitely sensitive to vibrotactile stimuli at 200- 
400Hz and is presumed to be associated with the 
Pacinian corpuscle (PC) class of receptor (Hunt 1960; 
Hunt and McIntyre 1960; Sato 1961; Janig et al. 1968; 
Talbot et al. 1968; Lynn 1969; Ferrington and Rowe 
1980; Ferrington et al. 1984; Coleman et al. 2001). 

In human subjects where microneurography 
studies have permitted the recording and 
characterization of tactile sensory nerve fibres 
supplying the hand and finger tips, the same broad 
divisions apply, except that the SA group of fibres is 
reported to fall into two classes, designated the SAI 
fibres that appear to correspond with the single broad 
SA class in both the cat and non-human primates, and 
an SAI class that appears to be associated with Ruffini 
receptor endings, principally in the regions of skin 
around nail beds or skin creases near 
metacarpophalangeal joints (Knibest6l and Vallbo 
1970; Johansson and Vallbo 1979). Although the SAT 
class is known to be present in the cat, it appears, in 
that case, to be confined to the hairy regions of skin, 
once again in association with Ruffini receptors 
(Chambers et al. 1972; Gynther et al. 1992). 


303 


MONOTREME TACTILE MECHANISMS 


A 
ee 
oy) +--+ ++ + ++ +H] 4$+- 
ee ee 
—) He} 4+ $+ +H + 
ee ce ee ee 
a ee 
eo AH 


f aga a a a rar | | 


1 Second 


60 
3 40 
a 
£ 
@o 
2 
[o} 
a 
® 
= 20 


0 500 
Step Amplitude (microns) 


1000 1500 


Figure 1. Response traces and stimulus-response relations for representative low-thresholds SA afferent 
fibers supplying the glabrous skin of the echidna forepaw. (A): response traces to step indentations (lasting 
1.5s) at a range of amplitudes (100 to 1,500 um; represented in the two waveforms above and below the 
impulse traces). Stimuli were applied to the RF focus (indicated by the arrow in the inset in B) by means 
of a 2mm diameter stimulus probe. (B): stimulus-response relations for four SA fibers based on the mean 
response (impulses/sec + S.D.) over the first one second from the onset of the 1.5s step indentation, plotted 
as a function of the indentation amplitude (modified from Mahns et al. 2003). 


For perhaps most mammalian species the 
densely innervated glabrous skin of the limb 
extremities represents the skin area of greatest tactile 
acuity (Darian-Smith 1984; Coleman et al. 2001; 
Johnson 2001, 2002) and may be regarded for most 
species as the tactile equivalent of the visual system’s 
fovea. Some exceptions to this, where the tactile 
‘foveal’ role may be served by other structures, could 
include the bill in the platypus (Bohringer and Rowe 
1977; Rowe 1990; Rowe et al. 2003) and the nasal 
appendages of the star-nosed mole (Catania and Kaas 
1997). Nevertheless, it is probable that in the echidna, 
the distal forelimb glabrous skin assumes a similar 
crucial tactile sensory role as the footpads in the cat or 
the hand and fingers in primates. 


Electrophysiological analysis of tactile sensory 
nerve fibres supplying the echidna forepaw 
Microdissection of the ulnar or median nerves 
in the echidna forelimb permitted electrophysiological 
recording from almost 30 individual tactile sensory 
nerve fibres that supplied the glabrous regions of the 
echidna distal forelimb (Mahns et al. 2003). Once an 
individual sensory fibre was isolated, its cutaneous 
receptive field (RF) was mapped by means of gentle 
probing with von Frey hairs of known calibrated force. 
The functional characteristics of the fibre were then 


304 


characterized by applying precise mechanical stimuli 
with a probe (usually 1-2 mm diameter) to the centre 
of the RF. The analysis, based upon the use of a 
feedback-controlled mechanical stimulator, revealed 
that the echidna tactile afferent fibres could be 
classified, like their placental counterparts, into two 
broad groups, one displaying slowly adapting (SA) 
response characteristics to static skin displacement, the 
other displaying a pure dynamic sensitivity. 


Functional characteristics of slowly adapting 
(SA) tactile afferent fibres supplying the echidna 
forepaw 

Tactile fibres in the SA class varied widely in 
their sensitivity to skin displacement. Those with 
lowest thresholds displayed small, circumscribed RFs 
and a sensitive grading of their impulse output as a 
function of skin displacement (Fig. 1A,B) and were 
therefore presumably well able to signal information 
about the location and intensity of skin perturbations 
encountered on the glabrous skin surface. A higher- 
threshold subclass of the SA fibres (Mahns et al. 2003) 
had larger RFs and less-well sustained responses 
(Fig.2A) and may contribute to the signalling of much 
more diffuse pressure encountered by the foot in 
locomotor or digging activity. The lesser sensitivity 
of this subgroup may be explained by the nature of 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAI 


B 


200 1 sepa neers nen 
ooo 
pe UE 
1000 ym See eee 
100 um 


—— 1_second = 


Figure 2. Response traces for a representative of the sensitive SA fibers associated with the echidna 
forepaw (A) and for claw-associated SA fibers. (B). The impulse traces in A and B show responses to the 
ramp indentation applied at a range of amplitudes (200-1,500um) to the glabrous skin (A) or to the base 


of the claw (B) (modified from Mahns et al. 2003). 


the echidna forepaw which is a very much thicker, 
cushioned structure than the footpad in the cat. This 
gross anatomical specialization appears to equip the 
echidna well for its robust digging and burrowing 
activity but may create, as a consequence of the 
viscoelastic properties of the footpad, a mechanical 
housing for many SA sensory endings that imposes 
limitations on the effective mechanical coupling 
between the skin surface and the receptor endings, and 
this, in turn, may account for the higher thresholds 
and more diffuse RF boundaries (Mahns et al. 2003). 

A further subset of SA fibres was associated 
with the base of the powerful claws and displayed a 
gradation in impulse output related to displacement at 
the base of the claw (Fig.2B), attributes that should 
equip them to subserve a kinaesthetic role in signalling 
movements of the powerful claws (Mahns et al. 2003). 

The low-threshold SA fibres appear to 
conform in properties to the SA/ class of tactile afferent, 
already identified in the echidna snout (Iggo et al. 1996) 
and in the skin of placental species where it is generally 
thought to be associated with the Merkel receptor 
endings. Although the less-sensitive and claw- 
associated SA fibres in the echidna (Fig.2A,B) have 
some attributes resembling the placental SAJJ fibre 
class, in particular, a rather regular temporal pattern 
in the impulse activity of the claw-associated SA fibres 
(Mahns et al. 2003), the absence of spontaneous 
activity in these fibres is more consistent with an SAI 
identification. Thus, despite the rather disparate 
behaviour of the different subsets of SA fibres 
associated with the echidna forepaw it appears that, as 
a broad class, they probably conform more closely to 


Proc. Linn. Soc. N.S.W., 125, 2004 


the SAJ class with its implied association with Merkel 
receptor endings. However, any conclusions about the 
receptor associations and precise identity if the echidna 
SA fibres must remain tentative until correlative 
histological data are available for the receptors. 


Functional characteristics of dynamically- 
sensitive tactile afferent fibres supplying the 
echidna forepaw 

Dynamically-sensitive tactile sensory fibres 
associated with the echidna forepaw could be divided 
into two classes principally according to their 
differential sensitivity to vibrotactile stimuli but, in 
addition, could often be distinguished on account of 
RF characteristics and sensitivity to manual stimuli 
(Mahns et al. 2003). One class resembled the rapidly 
adapting (RA) class of tactile afferent identified in 
association with the cat and primate glabrous skin in 
having small, circumscribed RFs and displaying 
maximum vibrotactile sensitivity at frequencies < 50 
Hz (Fig.3). The second class had larger RFs and a 
bandwidth of vibrotactile sensitivity that extended up 
to 300-400 Hz, properties resembling those identified 
for the PC sensory fibres that are associated with 
Pacinian corpuscle (PC) receptors in the glabrous skin 
of placental species, including the cat, marmoset and 
macaque monkeys, and human subjects (reviewed 
above). 


Tactile coding capacities of dynamically-sensitive 
tactile afferent fibres in the echidna 

The RA class of afferents supplying the 
echidna footpad had absolute response thresholds of 


305 


MONOTREME TACTILE MECHANISMS 


A B 

5 um | | : 200 
® 
o 
Ss. 
E 
2) 
= 
re) 
a. 
n 
o 


100 


100 
Vibration Amplitude (microns) 


150 200 


Figure 3. Impulse records and stimulus-response relations for a representative RA afferent fiber supplying 
the echidna forepaw. A): Activation of the fiber as a function of amplitude increases in the 50Hz vibration 
train. The response achieves a 1:1 following pattern over the first ~15 cycles at 20 um, and this response 
pattern is sustained throughout the 1-second duration of the vibrotactile stimulus at 40 um. (B): Stimulus- 
response relations for this RA fiber show 1:1 plateau levels of response at <40 um at frequencies of 30 and 
50 Hz. At 100, 150, and 200 Hz, the 1:1 level was attained only at amplitudes of ~70, 100, and 200 um, 


respectively. The RF for the fiber is indicated in the inset in B (modified from Mahns et al. 2003). 


<10 um at frequencies below ~100 Hz. With increases 
in the intensity of vibrotactile stimuli, their response 
(in impulses/s) progressively increased until they 
attained a regular impulse discharge on successive 
cycles of the vibration stimulus. As the spike 
discharges in this so-called one-to-one pattern of 
response were tightly phaselocked to the vibration 
waveform the interspike intervals approximated the 
vibration cycle period. For example, at 50 Hz the 
interspike intervals approximated 20 ms and at 100 
Hz were ~10ms (Fig. 4). Because of the tight 
phaselocking, the pattern of discharge displayed a 
metronomic regularity that reflected very precisely in 
its temporal pattern the periodicity inherent in the 
vibrotactile stimuli. The impulse sequence thus 
provided a reliable signal, in an impulse pattern code, 
of the frequency, or pitch parameter of the vibrotactile 
stimuli. The tightness of phaselocking could be 
quantified by constructing cycle histograms (CHs; Fig. 
4B) which show the time of occurrence of impulses 
during each vibration cycle. The CHs use a pulse 


306 


associated with the onset of each vibration cycle as a 
stimulus marker and have an analysis time that 
corresponds to the vibration cycle period. Responses 
that are tightly synchronized, or phaselocked, appear 
in the CH as a narrow peak as seen in Fig. 4B. When 
impulses occur independently of the phase of the 
applied vibration waveform the distribution in the 
histogram appears flat (Coleman et al. 2001; Mahns 
et al. 2003). The bandwidth of vibrotactile frequencies 
over which responses in the echidna RA fibres 
remained phaselocked extended from ~5 Hz up to ~200 
Hz, matching the behaviour of their placental 
counterparts in the cat (Ferrington and Rowe 1980; 
Ferrington et al. 1984; Rowe and Ferrington 1986) and 
in the macaque and marmoset monkeys (Talbot et al. 
1968; Coleman et al. 2001). 


Functional capacities of the presumed-PC class of 
tactile sensory fibre in the echidna 

Controlled vibrotactile stimuli permitted the 
distinction to be made between the RA afferent fibre 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAI 


@ 7 soHz 
£ R=0.94 
2 
50 Hz : 
O 0 
1007 400 Hz 
R=0.94 


100 Hz 


Counts (Imp) 


Figure 4. Precision of impulse patterning in the responses of echidna RA afferent fibers to vibrotactile 
stimuli. (A): Impulse traces show the tightly phase-locked pattern of response reflecting the periodicity of 
the 50 and 100 Hz vibrotactile stimuli. Quantitative measures of phase locking based on the vector strength 
_or resultant, R (see Materials and Methods), were derived from the cycle histograms in (B), constructed 
to show the distribution of impulse occurrences throughout successive cycles of the vibration stimulus at 
the two frequencies. The analysis time in each CH corresponds to the cycle period of the vibration (modified 


from Mahns et al. 2003). 


class and a class of dynamically-sensitive tactile 
afferents with a distinctly broader bandwidth of 
sensitivity (Fig.5) reminiscent of placental PC fibres 
(Mahns et al. 2003). These putative PC fibres 
supplying the echidna forepaw had absolute response 
thresholds as low as ~5 um for the broad range of 
frequencies from ~50 to 300 Hz (Fig.5C) which may 
equip the animal to detect small vibratory perturbations 
set up by termites in either the soil or in timber material 
encountered in the animal’s use of the forepaw as an 
exploratory organ. Furthermore, these broad 
bandwidth vibrotactile sensors would be well-suited 
to serve as an early-warning system for the detection 
of ground-borne vibration signalling the movements 
of any predators or other animals in the vicinity 
(McIntyre 1980; Mahns et al. 2003). As the PC fibre 
responses to vibrotactile stimuli remained tightly 
phaselocked at frequencies up to and beyond 400 Hz, 
the metronome-like patterning in their discharge (Fig. 
5A) would ensure that these fibres retained high acuity 
for signalling the temporal details of vibrotactile 
perturbations over a broad bandwidth of frequencies 
(Mahns et al. 2003). Although the bandwidth of 
vibrotactile responsiveness in the echidna PC fibres 
(Fig.5) did not extend to frequencies quite as high as 
those of placental PC fibres (Mahns et al. 2003) the 
explanation probably lies in the lower body 


Proc. Linn. Soc. N.S.W., 125, 2004 


temperature of the echidna (~28-32°C; Grigg et al. 
1992) rather than a fundamental difference in the 
receptors; in particular, as Sato (1961) has 
demonstrated that PC fibres in the cat display a 
displacement to lower frequencies in both bandwidth 
and peak sensitivity as temperature is lowered. 

The use of controlled sinusoidal vibration as 
a dynamic form of tactile stimulation in these studies 
permitted the precise quantification of both the 
frequency and intensive parameters of the stimuli. 
However, in addition, it provides a form of dynamic 
tactile stimulation that mimics in a controlled way the 
vibrational disturbances set up in the skin in association 
with relative movement between the skin and any 
textured surface encountered in the tactile exploratory 
movements of the forelimb. The properties revealed 
for echidna RA and PC fibres indicate that they are 
well suited to underpin the echidna’s capacity to signal 
and code information about textural changes in the 
ground surface or in the coarseness or fineness of 
objects encountered, such as sand, gravel or soil, in 
the tasks of locomotion, digging and burrowing. 

In summary, it appears that for the distal 
forepaw of the echidna, the tasks of tactile exploration 
and perception are based upon a triad of major tactile 
sensory fibre classes, comprising a broad SA class and 
both RA and PC classes with functional capacities 


307 


MONOTREME TACTILE MECHANISMS 


400 H 

400 wg 
a) 
o 
@ 
Q 
£ 
®o 
no 
i= 
[o} 
Qa 
wn 
® 
oo 

0 20 40 60 80 100 
Vibration Amplitude (microns) 
50 
B 
40 en 
: te Gap ed UNG 


—— Absolute 


Vibration Amplitude (microns) 


Vibration Frequency (Hz) 


Figure 5. (A): Temporal patterning in the vibrotactile responses of PC-like fibers in the echidna. Impulse 
traces and peristimulus time histograms (PSTHs) show the metronome-like impulse pattern at the 1:1 
response level at vibrotactile frequencies of 50-400 Hz. Each PSTH was constructed from five consecutive 
responses to 20 cycles of vibration at each of the indicated frequencies. (B and C): Stimulus-response 
relations and vibrotactile frequency bandwidths for the putative PC class of echidna tactile afferent 
fiber. (B): Stimulus-response relations for a single PC-like fiber with RF on the lateral aspect of the 
forepaw glabrous skin, based on plots of the mean response (impulses/second) as a function of vibration 
amplitude at seven frequencies in the range 10-400 Hz. (C): Plots of absolute (solid lines) and 1:1 tuning 
thresholds (dashed lines) derived for five PC-like afferent fibers from stimulus-response data of the type 


shown in (B) (modified from Mahns et al. 2003). 


resembling those of the corresponding classes in 
placental mammals. The issue of whether the broad 
SA class might contain subsets will be resolved only 
with more detailed morpho-functional correlative 
analysis on both the receptor endings and the associated 
sensory nerve fibres. However, the breakdown of the 
echidna tactile sensory fibres into three broad classes 
resembling those in placental mammals suggests that 
peripheral mechanisms for tactile sensation in the distal 
glabrous skin are highly conserved across different 
mammalian orders (Mahns et al. 2003). 


CEREBRAL CORTICAL ORGANIZATION FOR 
TACTILE PROCESSING IN MONOTREMES 


The behavioural evidence for the pre- 
eminence of the tactile sense in the platypus, and 


308 


probably also in the echidna, has been re-inforced by 
electrophysiological studies on the organization of the 
cerebral cortex in these two species. In the platypus in 
particular, the allocation of neocortical space to tactile 
processing is quite spectacular as was demonstrated 
with both evoked potential and single-neuron 
microelectrode recording studies first undertaken in 
our laboratory in the 1970s (Bohringer and Rowe 1977; 
Rowe 1990), and more recently by Krubitzer et al. 
(1995a). With evoked potential mapping, a brief 
electrical stimulus delivered at a point on the skin 
surface activates sensory fibres that generate a 
synchronous input to the areas of cerebral cortex 
involved in processing information from that source 
of sensory input. From the cortical surface overlying 
these areas it is possible to record an evoked potential 
that is usually biphasic, consisting of an initial positive-’ 
going deflection followed by a larger negative-going 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAT 


—— 
-e-— 
~ ~ 
SES 


4 o Bee ee Se 
a ae es 
7 ~ 
oA _— — ee eee ee eee es ee ee ee ee a 
7 
Low a 


“ 
eats im “ -\ + si 
RY 


os 
xo 4 
Aone Re 


Ses 
oF 48 


ers 
Ss 
Ss 


Figure 6. Evoked potentials (positively downwards) recorded from the cortical surface of the platypus 
following bipolar stimulation of the anterolateral margin of the contralateral bill (1V, 100-microseconds 
pulse). Each recording was made from the position indicated by the dot at the left of the trace. Dotted 
lines indicate positions of large blood vessels in frontal region of hemisphere; view of hemisphere and 
whole brain (inset) from dorsolateral aspect. Stippled areas in inset represent focal projection sites and 
include sites at which positive-going responses exceed 100uV for bill (B), 30uV for fore limb (FL) and 
10uV for hind limb (HL). Zones between stippling and continuous lines include sites from which smaller 
responses could be recorded (from Bohringer and Rowe 1977). 


component (Fig.6, and Bohringer and Rowe 1977). 
The initial positive-going component is thought to arise 
from the direct excitatory action of thalamo-cortical 
afferent input on cortical neurons (Mountcastle and 
Poggio 1968), and therefore the cortical region from 
which this component can be recorded is thought to 
represent the projection focus for that source of input. 
The evoked potentials illustrated in Fig.6 were 
recorded in response to stimulation on the anterolateral 
margin of the bill and reveal that a vast area of the 
dorsal surface of the contralateral cerebral hemisphere 
is taken up with the processing of bill inputs. At each 
recording point indicated by the dots on the main 
figure, responses were also recorded to forelimb (FL) 
and hindlimb (HL) stimulation in the platypus allowing 
the isopotential contour maps for these and the bill 


Proc. Linn. Soc. N.S.W., 125, 2004 


(B) inputs to be constructed on the inset figure, with 
the stippled areas indicating the focal projection sites 
for the three sources of input, and the zone between 
the stippling and continuous line a region from which 
smaller evoked potentials could be recorded. 
Single neuron mapping of the somatosensory 
cortex in monotremes 

More detailed single-neuron microelectrode 
recordings from as many as 250 individually- 
discriminated cortical neurons in up to 67 electrode 
penetrations in a given experiment on the platypus 
somatosensory cortex confirmed the highly ordered 
and complete cortical representation of tactile inputs 
from the contralateral body, extending from the mid- 
sagittal region of the hemisphere out to the region of 
the rhinal sulcus (Figs.7 and 8) on the ventrolateral 


309 


MONOTREME TACTILE MECHANISMS 


surface, the one sulcus present as an exception to the 
lissencephalic state of the platypus cerebral cortex. As 
the microelectrode mappings were based on inputs 
generated by light tactile stimulation of the skin surface 
they confirm the remarkable size of the cortical space 
devoted to tactile processing, in particular, from the 
bill. This is important as the electrical stimuli delivered 
to the skin in evoked potential mappings could have 
activated a combination of tactile and the putative 
electroreceptive afferent fibres postulated to be present 
in the bill of the platypus (see below). The continuity 
of tactile representation across the somatosensory 
cortex of the platypus was confirmed in several detailed 
mappings (Bohringer and Rowe 1977) and is apparent 
in Fig.8 which shows the tactile receptive fields 
outlined on the figurines for 60 individual neurons 
isolated in nine electrode penetrations made in a single 
anteroposterior plane in the posterior region of the 
hemisphere. While tactile receptive fields for 
individual neurons are up to ~15 cm? on regions such 
as the tail and trunk, those on the distal glabrous skin 
of the limbs were much smaller, while those on the 
bill, in particular, its anterior and lateral margins, were 
no more than 1mm in diameter. These represent the 


smallest tactile receptive fields ever recorded in the 
cortex and are therefore capable of conferring great 
precision and fidelity upon the tasks of tactile 
localization and discrimination involving the bill. 

Where the electrode penetration was made 
normal to the cortical surface, the neurons encountered 
had very similar receptive field locations (Fig.8) 
indicative of the columnar organization, well described 
for the cerebral cortex of placental mammals (e.g. 
Mountcastle 1957), in which neurons of similar 
functional properties are grouped in columns oriented 
normal to the surface. In penetrations, such as number 
eight in Fig.8, that passed obliquely through successive 
cortical columns in the region of bill representation, 
there was a remarkably orderly and progressive shift 
in the representation of the bill surface that is apparent 
in the enlarged and expanded view of this electrode 
track in Fig.9, emphasizing once again the striking, 
fine-grain spatial resolution available within the area 
of bill representation in the somatosensory cortex of 
the platypus. 

The somatosensory cortex of the echidna, 
mapped by Lende (1964), also has a striking allocation 
of space to the tongue and snout representations but, 


Figure 7. Inset shows entry points (filled circles) for 54 microelectrode penetrations into the platypus 
cortex. The black areas on figurines show the combined receptive field areas for all neurons (up to 15) 
sampled in each of the penetrations. Dotted lines represent positions of large blood vessels in frontal 
region of hemisphere; view of hemisphere from dorsolateral aspect (from Bohringer and Rowe 1977). 


310 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAT 


A 
CoE <a 
Y Cr) nina 
SS ee oO eS 

Cc 


9 
c 


) 
. 


i 
1} 
1 


Ti 
oO 
1 


N 
i 


N 
oO 
1 


DEPTH BELOW CORTICAL SURFACE (mm) 


@ 
a 
1 


Figure 8. (A): Photograph of cortical surface of the platypus from dorsolateral aspect indicating a plane 
in which 9 penetrations were made. (B): Coronal section through hemisphere at the plane indicated in A, 
showing the course of the 9 penetrations. (C): Reconstruction of penetrations 1-9 indicating location of 
each neuron studied (filled circles) and its peripheral receptive field. No fields could be found for the first 
two neurons in penetration 4. Receptive fields for all neurons in each of the penetrations 7-9 were confined 
to the shaded areas on each of the associated figurines (from Bohringer and Rowe 1977). 


in addition, a prominent representation of the distal 
forearm, presumably reflecting its importance in 
digging and burrowing. 


Multiple representation of the body within the 
monotreme cerebral cortex 

For both the platypus and the echidna, the 
early cortical mapping studies in our laboratory 
(Bohringer and Rowe 1977; Rowe 1990) and that of 
Lende (1964), led to the conclusion that there was a 
single body representation in the contralateral cerebral 
hemisphere, conforming to the so-called primary 
somatosensory cortex (SJ) of other mammals. No 
evidence was found for a second representation that 


Proc. Linn. Soc. N.S.W., 125, 2004 


might correspond to either the SII area that is well 
recognized in, for example, the cat and primate species 
(for review, Rowe 1990; Johnson 1990; Rowe et al. 
1996; Zhang et al. 1996, 2001) or indeed, to any other 
areas of somatosensory representation that have been 
reported in some placental species (Kaas 1982, 1987; 
Krubitzer and Kaas 1990; Krubitzer et al. 1995b), and 
now more recently, for both the echidna and platypus 
(Krubitzer et al. 1995a). Krubitzer et al. (1995a) have 
proposed that there are four somatosensory 
representations within the contralateral cerebral cortex 
of both the platypus and the echidna which they 
designate the primary somatosensory cortex (S/), the 
Rostral deep field (R), the Manipulation field (M) and 


SH bhh 


MONOTREME TACTILE MECHANISMS 


SE aes Oe 
reac, Sa aks 


Figure 9. Coronal section of platypus cerebral cortex showing the 
position of a microelectrode penetration (track 8 in Fig.8) made 
obliquely to the cortical surface. The filled circles indicate the locations 
of the 14 neurons studied. The receptive field for each neuron is 
indicated on the right which is an enlargement of the area on the 
ventral surface of the bill (modified from Bohringer and Rowe 1977). 


the Parietal Ventral field (PV). Furthermore, each is 
said to contain “a complete representation of the body 
surface”, a surprising claim to us, considering that the 
four areas are not all defined as being concerned with 
the processing of mechanosensory data from the body 
surface. The area given the S/ designation is concerned 
principally with cutaneous inputs, whereas area R 
immediately rostral to SJ is said to contain neurons 
that respond “most often to stimulation of deep 
receptors” and which required light taps or light 
pressure to the body surface to elicit a response. 
However, as the tactile stimuli employed in their study, 
whether brushing, tapping, or pressure forms of 
cutaneous stimulation, were neither quantified nor 
reproducible, it is difficult to see how reliable 
distinctions were made in terms of neural response 
thresholds between neurons in the different cortical 
representation areas. 


Topographic organization of the putative 
multiple areas of somatosensory representation in 
the monotreme cerebral cortex 

As responsiveness criteria may be insufficient 
to permit an unequivocal division of the monotreme 
somatosensory cortex into four distinct areas 
(Krubitzer et al. 1995a), such a differentation must 
therefore depend upon topographic considerations, in 
particular upon the extent to which the four putative 
areas constitute complete and discrete representational 
maps of the body. In the case of adjacent areas, 


32 


whether, for example, S/ and 
R, or SI and PV, Krubitzer et 
al. (1995a) state that the 
boundaries coincide with a 
reversal in the representation 
of peripheral receptive fields as 
one progresses across a 
sequence of cortical recording 
sites. For example, in both 
platypus and echidna cortex, 
the receptive fields on the 
upper body may shift from 
proximal body locations, 
including the face, shoulder 
and upper limb, to the distal 
forelimb, and then move back 
to more proximal parts of the 
limb and shoulder. Similar 
reversals were seen across 
sequences of recording sites 
involving the hindlimb 
representations, leading to the 
interpretation by Krubitzer et 
al. (1995a) that the separate 
representations of proximal limb and associated trunk 
regions constitute parts of two distinct body 
representations. However, an alternative interpretation 
must be considered, which emerges from the detailed 
studies carried out in the macaque monkey by Werner 
and Whitsel (1968, 1973) and Whitsel et al. (1969, 
1971) for S/ in the postcentral gyrus, and for the more 
laterally-placed second somatosensory area, S/I/. 


| ee 


The representation of the body within the 
cerebral cortex: a reflection of the dermatomal 
trajectory 

The crucial observation emphasized by 
Werner and Whitsel was that the body maps in both S7 
and SII of the cerebral cortex owe their essential 
topographic properties to the serial and overlapping 
projection of the dorsal roots. In the case of the 
macaque postcentral gyrus, the representation of spinal 
roots, from sacral through to cervical, forms a 
succession of antero-posteriorly oriented bands 
progressing from medial to lateral across the cortex 
(e.g., Figs. 6 and 8 in Werner and Whitsel 1973). 

Within the SI area of the macaque monkey, 
and indeed within the macaque SJ/ as well, the 
representations of the postaxial and preaxial arm and 
leg areas are separated by the representation of the 
more distal parts of the limbs, in particular, the digits 
and toes respectively. This effectively gives rise to a 
split representation of the upper parts of the limbs. In 
the case of the postcentral gyrus, one component lies © 


Proc. Linn. Soc. N.S.W., 125, 2004 


M.J. ROWE, D.A. MAHNS AND V. SAHAI 


medial to the distal limb representation, the other lateral 
to it, an arrangement represented schematically to 
illustrate this dermatomal trajectory in Fig.8 of Werner 
and Whitsel (1973). One may observe how the 
dermatomal trajectory generates the split representation 
of the upper regions of either forelimb or hindlimb 
within the cerebral cortex by examining, in a human 
anatomy text (e.g., Williams and Warwick 1980) the 
dermatomal boundaries of successive spinal roots 
associated with either the lower or upper regions of 
the body. In the case of the upper body, the ventral 
and dorsal axial lines of the upper limb mark a border 
between the innervation fields of C5 and T1. Therefore, 
in any representation of the body surface within the 
cerebral cortex, one might expect, with the central map 
being laid down according to the dermatomal trajectory 
(Werner and Whitsel 1968, 1973; Whitsel et al. 1969, 
1971), that the central representation of the upper arm 
will be split along the axial lines with the input from 
the distal limb, carried over the C6, 7 and 8 roots, 
creating a clear separation in the cortical representation 
of lateral and medial surfaces of the upper arm. 

As the same fundamental plan and sequence 
for tactile dermatomes is also found in both the cat 
and monkey (Sherrington 1898; Kuhn 1953; 
Hekmatpanah 1961), one may assume that this 
organizational plan for spinal segmental innervation 
is a general one that would operate across mammalian 
orders, including the monotreme representatives. 


Is there multiple representation of the body 
within the platypus and echidna cerebral cortex? 

If one examines the receptive fields plotted 
for the platypus and echidna somatosensory cortex by 
Krubitzer et al. (1995a) in their Figs. 6 and 16, it might 
be argued that those fields on the proximal parts of the 
limb, on either side of the distal limb representation, 
are not clearly and systematically separated into 
representations of the medial and the lateral surfaces 
of the limb as might be expected in a perfect reflection 
of the dermatomal trajectory. However, this is hardly 
surprising on several grounds that are outlined in a 
recent review (Rowe, in press). 

The fundamental point to be emphasized is 
that the reversals in receptive field representation 
described by Krubitzer et al. are consistent with the 
sequence of representation that might be expected 
within a single body map whose plan is determined 
by the dermatomal trajectory. In view of these 
considerations we would re-emphasize our 1977 
finding (Bohringer and Rowe 1977; Rowe 1990) of a 
single large representation of the contralateral body 
within the platypus cerebral cortex and the similar 
conclusion reached even earlier by Lende (1964) for 


Proc. Linn. Soc. N.S.W., 125, 2004 


the echidna. Furthermore, we wish to emphasize the 
fundamental importance and significance of Werner 
and Whitsel’s studies (1968, 1973; and Whitsel et al. 
1969, 1971) on body representation within the cerebral 
cortex, and the need that arises from their studies, to 
take account of the dermatomal trajectory as the crucial 
determinant of representational topography within 
central neural systems. 


Sensory and perceptual specialization in 
monotremes: trigeminal electroreceptive 
mechanisms 

As the monotremes emerged in mammalian 
evolution on a separate line from therian mammals in 
the early Mesozoic (Dawson 1983; Rowe 1990; Augee 
and Gooden 1993) the possibility that some 
qualitatively different apparatus for neural sensing 
might have emerged was given some credence in the 
1980s with both behavioural and electrophysiological 
studies reporting the presence of electroreception in 
monotremes (Scheich et al. 1986; Gregory et al. 1987, 
1988, 1989a,b). These and subsequent reports 
suggested that electroreception might be associated 
with the bill of the platypus and the snout of the 
echidna, in each case in association with the trigeminal 
nerve rather than the lateral line system that is the 
principal basis of electroreception in certain fish (Cahn 
1967; Bullock 1999). However, neither behavioural 
nor electrophysiological data have provided any 
suggestion of electroreception in association with other 
skin regions or somatosensory nerves of the 
monotremes, such as the median or ulnar nerves of 
the forelimb. 

The first behavioural evidence for 
electroreception in the platypus bill came in a short 
report from Scheich et al. (1986) that the platypus could 
detect weak electric fields with threshold strengths as 
low as 50-200 uV cm'. Furthermore, they reported 
that an electroreceptive processing zone was present 
in the posterolateral region of the contralateral cerebral 
cortex on the caudal side, but next to, the map of bill 
mechanoreceptive input identified by Bohringer and 
Rowe (1977). In our view this is a puzzling finding on 
several grounds (for review, see Rowe 1990). First, in 
an earlier detailed electrophysiological mapping of the 
cerebral cortex we found that the cortical region 
concerned with tactile representation of the bill 
occupied almost the whole of the posterolateral region 
of cortex (Bohringer and Rowe 1977; Rowe 1990). 
Furthermore, we constructed separate cortical maps 
of bill representation based upon mechanical 
stimulation which was specific for tactile inputs, and 
electrical stimulation which would have activated both 
tactile and the putative electroreceptive afferents. 


SS) 


MONOTREME TACTILE MECHANISMS 


However, comparison of the two maps (Figs.6 and 7; 
and Bohringer and Rowe 1977) reveals no evidence 
of a separate area of bill representation in the map based 
upon the electrical stimulation, from that obtained with 
pure tactile stimulation. In contrast to the Scheich et 
al. (1986) report that electroreceptive-induced cortical 
evoked potentials were found next to the map of bill 
mechanoreceptor input, those by Iggo et al. (1992) and 
Krubitzer et al. (1995a) indicated that the 
electrosensory area of cortical representation lay 
entirely within the border of the tactile representation 
area defined for the bill in our earlier study (Bohringer 
and Rowe 1977; Rowe 1990). However, Krubitzer et 
al. (1995a) have described a “clear functional 
parcellation” within this SI region whereby regions 
responsive to just tactile input were interdigitated with 
regions responsive to both tactile and electroreceptive 
inputs (e.g. Fig.10 in Krubitzer et al. 1995a). 
Furthermore, the regions of pure tactile sensitivity 
coincided with myelin and cytochrome oxidase-dense 
regions of SI while the regions of putative bimodal 
sensitivity coincided with myelin-light regions. What 
the significance of such differences in myelin density 
might be in this circumstance remains unclear. 
However, there may not be universal agreement with 
this assertion anyway, judging by the allocation of light 
and dark areas in their Fig.10B; for example, the 
myelin-dark region drawn to contain four recording 
sites of pure mechanosensitivity in part B of their figure 
appears rather paler in the adjacent tangentially- 
oriented photomicrograph than some areas represented 
as myelin-light in Fig.10B. It is likely to be difficult to 
make these distinctions reliably, in the tangentially- 
cut cortical sections, firstly without objective analysis 
of image density, and secondly, in the absence of 
systematic control for laminar depth across the extent 
of the cortical section under study. 

Electrosensory field-strength thresholds 
illustrated for cortical neurons in Fig.10 by Krubitzer 
et al. (1995a) ranged up to 900 uV cm; however, it 
was not clear what field strength might have activated 
neurons in the purely tactile-sensitive regions. 


Behavioural and afferent fibre thresholds for 
electroreception 

A further concern in relation to claims for 
electroreception in monotremes arises over the 
thresholds reported for the phenomenon. At the 
behavioural level, Scheich et al. (1986) reported field 
strength values as low ~50-200 uV cm’! for the 
platypus, and, more recently, values of 20uVcm'! have 
been reported (Manger and Pettigrew 1996; Pettigrew 
1999). However, individual trigeminal afferents, 
believed to be of the electrosensitive class, had 


314 


threshold field strengths of ~4m¥V cm’! (Gregory et al. 
1988, 1989b). As these values were based on a 
substantial fibre sample, and as these values are vastly 
higher than reported behavioural thresholds for the 
platypus, one might infer that any putative 
electroreceptive sense must depend upon some other 
source of afferent input. Proposals that spatial 
summation, based upon convergence of a number of 
trigeminal electroreceptive afferents onto central 
neurons, may confer the observed behavioural 
thresholds upon the animal are difficult to accept when 
none of the sampled afferent fibres has thresholds low 
enough to account for this. 

To the extent that there are contentious issues 
associated with the claims for electroception within 
the monotreme order of mammals, it is important that 
further rigorously controlled investigations be pursued 
to resolve such issues, in particular, more detailed 
behavioural studies based upon objective analysis 
rather than anecdotal accounts of the movement 
patterns of the platypus and echidna in relation to 
presumed electrosensory stimuli. 


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The Role of Push Rods in Platypus and Echidna — Some 
Speculations 


U. PROSKE AND J.E. GREGORY 


Department of Physiology, Monash University, Clayton, VIC 3800 
(uwe.proske @ med.monash.edu.au) 


Proske, U. and Gregory, J.E. (2004). The role of push rods in platypus and echidna — some speculations. 
Proceedings of the Linean Society of New South Wales 125, 319-326. 


This is a review of the structure and innervation of the mechanosensory organ, the push-rod, in skin of the 
platypus bill and echidna snout. Four receptor types can be identified in association with push rods in 
platypus and echidna: (i) central vesicle chain receptors, (11) peripheral vesicle chain receptors, (iii) Merkel 
endings and (iv) paciniform corpuscles. Function of the vesicle chain receptors remains unknown. Merkel 
endings are known to be slowly adapting with irregular discharge (SAI) while paciniform corpuscles are 
rapidly adapting vibration-sensitive (RA). Recordings made from echidna nose skin have identified both 
SAIs and RAs. In addition, responses typical of SAII endings (regular discharge) and rapidly adapting, but 
vibration insensitive, responses were observed. It was concluded that the push rod in monotremes is not 
associated with mechanoreceptors unique to the group. Skin of the platypus bill and the echidna nose 
contains erectile tissue. It is conjectured that blood engorgement inflates the skin to facilitate contact between 
push rods and the external environment. In addition platypus push-rods have a ring of contractile tissue 
around their tips which, on contracting, restricts mobility of the rod, perhaps when the platypus leaves the 


water. Possible cooperative roles between electroreceptors and mechanoreceptors are discussed. 


Manuscript received 3 September 2003, accepted for publication 22 October 2003. 


KEYWORDS: Cutaneous receptors; electroreceptors; mechanoreceptors; Merkel receptors; push rod; 


sensation; slowly adapting; vibration. 


INTRODUCTION 


In a recent publication we presented some 
speculations about the role of electroreceptors in the 
detection of prey items by platypus and echidna 
(Proske and Gregory 2003). Here we want to extend 
these speculations to the mechanoreceptors. This, 
therefore, is a review account of our own work and 
that of others and speculations based on those 
observations. 


THE PLATYPUS PUSH ROD 


The bill of the platypus has two prominent 
sense organ structures, the electroreceptors associated 
with sensory-innervated mucous glands, and 
mechanoreceptors associated with the push rods. They 
are distributed differently, the electroreceptors being 
lined up in rostro-caudally directed rows, the push rods 
distributed more or less uniformly across the bill, with 
particularly high concentrations on the edge of the bill 
(Fig. 1). In the platypus, although it has not been stated 


explicitly, it appears that push rods are absent from 
skin, including glabrous skin, of other parts of the body. 
In other words, push-rods seem to be specifically 
associated with the bill, as are the electroreceptors. 

There have been estimated to be 46,500 push 
rods in the platypus (Manger and Pettigrew 1996), 
distributed in the skin covering the outside of the upper 
and lower bill and lining the mouth (Fig. 1a). They are 
especially numerous around the margins of the upper 
bill. 

The platypus push rods (Fig. 2a) are about 
70 um in diameter and 400 um long, spanning nearly 
the full thickness of the epidermis. They consist of a 
column of flattened spinous cells attached rather 
loosely to the surrounding epidermis, which allows 
them relatively free movement in all directions. The 
rounded tip of the push rod protrudes slightly above 
the surrounding skin surface. Each push rod is 
innervated by 25 to 40 myelinated nerve fibres. These 
terminate in 4 types of sensory ending. 

The first type is the central vesicle chain 
receptor, supplied by 5 to 8 medium sized myelinated 
axons. The receptor consists of axon terminals running 


ROLE OF PUSH RODS IN MONOTREMES 


Figure 1. Dorsal and ventral view of the platypus bill (A) and lateral 
view of the echidna snout (B). Dots indicate the distribution pattern 
of push rods, which are more numerous near the edge of the bill 
and tip of the snout. (A, rearranged from Andres and von During, 


1984; B, from Proske, 1997.) 


vertically up the push rod near the centre, with a regular 
array of vesicle-like protrusions along their length. 

The second type, the peripheral vesicle chain 
receptor, is similar, but the myelinated axons (as many 
as 18) supplying it are thinner and their terminals are 
located towards the periphery of the push rod. 

The function of the vesicle chain receptors is 
still unknown. Their structure is somewhat reminiscent 
of Meissner corpuscles found in the glabrous skin of 
many mammals. Meissner corpuscles consist of a 
coiled arrangement of endings from a number of 
myelinated axons that terminate between layers of 
flattened Schwann cells, and they are known to be 
rapidly adapting mechanoreceptors. If, as is likely, 
vesicle chain receptors are mechanosensitive, their 
disposition in the centre and around the periphery of 


320 


the push rod appears favourable for 
detecting compression or bending of 
the rod, and perhaps in this way the 
direction in which a stimulus is acting 
can be signalled. The receptors 
extend to within just a few cells of 
the skin surface, giving rise to the 
suggestion that they would also be 
well placed to function as 
thermoreceptors (Catania 1995). 
The third type of receptor in the 
push rod is the Merkel cell. Up to 12 
Merkel cells are located at the base 
of each push rod, and each 
myelinated afferent nerve fibre 
supplying them branches to supply 6 
to 8 Merkel cells. The functional 
properties of Merkel cell receptor 
complexes have been well 
documented in other mammalian and 
avian species, where they are one of 
the common receptor types found. 
They are slowly adapting 
mechanoreceptors, giving rise to 
what in other mammals are termed 
Type I responses characterised by a 
high degree of variability in the train 
of impulses discharged when 
stimulated. Figure 3 shows an 
example, recorded from the echidna. 
The fourth type of ending is a 
group of 3 to 6 paciniform 
corpuscles, lying in the dermis 
immediately underneath the column 
of epidermal cells comprising the 
push rod. These are one of the most 
intensively studied types of 
cutaneous receptor and like the 
Merkel cell, they are found ubiquitously amongst birds 
and mammals. They are a rapidly adapting 
mechanoreceptor giving highly phasic responses to 
stimulation and able to signal faithfully vibratory 
stimuli at frequencies up to about 1,000 Hz. Figure 3 
shows an example, again recorded from the echidna. 
The paciniform corpuscles at the bottom of 
the platypus push rods have been reported to be 
arranged with their long axes oriented either strictly 
parallel or perpendicular to the skin surface, and at 
right angles to each other, thus building up a three 
dimensional system for transducing any direction of 
movement of the push rod (Andres & von During 
1984). This further implies that the push rods are free 
to move and not constrained to displacement in any 
particular direction. 


Proc. Linn. Soc. N.S.W., 125, 2004 


U. PROSKE AND J.E. GREGORY 


Figure 2. Schematic representations of push rods in the platypus bill (A) and echidna snout (B), showing 
the 4 types of sensory terminals: lamellated, paciniform corpuscles (p, Imr), Merkel cells (m, Mc), central 
vesicle chain receptors (cvc) and peripheral vesicle chain receptors (pvc), the latter 2 having a single label 
(ver) in B. The platypus push rod is attached rather loosely to the surrounding epidermis, allowing relatively 
independent movement in all directions, while the echidna push rod appears more firmly anchored in the 
epidermis. (A, modified from Andres and von During, 1984; B, modified from Andres et al, 1991.) 


THE ECHIDNA PUSH ROD 


In the echidna push rods are scattered across 
the surface of the snout, becoming fewer at its base, in 
the region where the hairs begin (Fig. 1b). They are 
especially dense near the tip of the snout, which is 
also the region where the electroreceptors are found. 
As for the platypus, there is no evidence of push rods 
in skin of other parts of the body. 

The structure of the echidna push rod follows 
the same plan as that in the platypus with two 
differences. Its nerve supply is less dense and the 
column of cells comprising the rod is less mobile than 
in the platypus — there is a less clearly differentiated 


Proc. Linn. Soc. N.S.W., 125, 2004 


boundary between the column of spinous cells with 
tonofibrils and the surrounding epidermis. 

The echidna push rods (Fig. 2b) are smaller 
than those in the platypus and are innervated by only 
about half as many myelinated nerve fibres. However, 
they contain larger numbers of Merkel cells and 
paciniform corpuscles (Andres et al. 1991). 


RESPONSE PROPERTIES OF THE RECEPTORS 


What emerges is that the push rod is a 
specialised mechanoreceptor complex, having separate 
receptor systems for signalling steady stimuli and 
phasic or vibratory stimuli, and if the speculations 


321 


ROLE OF PUSH RODS IN MONOTREMES 


Vibration receptor 


53 Hz 


433 Hz 


tet) 


WWWWWWE 


Slowly-adapting Type I 


Figure 3. Responses of single receptors in the echidna snout, shown as action potentials recorded in 
dissected nerve filaments. The vibration sensitive mechanoreceptor is shown responding in a 1:1 entrained 
fashion to stimulation at just above threshold amplitude and at the frequencies indicated. The stimulus is 
represented in the traces below the action potentials. This type of response is characteristic of paciniform 
corpuscles. The slowly adapting Type I response in the lower part of the figure is typical of Merkel cell 
receptors. It is characterised by an absence of resting or unstimulated discharge, a sensitivity to stimulus 
velocity and an adapting, irregular discharge during constant skin indentation. The stimulus is shown in 
the trace below the train of action potentials, and consisted of a linearly increasing skin indentation to an 
amplitude that was then held constant. (From Iggo et al, 1996.) 


about vesicle chain receptors are correct, a third system 
for signalling the direction in which a mechanical 
stimulus is acting. 

In an electrophysiological study of tactile 
receptors in the echidna, Iggo et al. (1996) recorded 
from 44 mechanoreceptor afferents with receptive 
fields in the skin of the snout. The receptors were not 
identified morphologically but some attempt was made 
to associate a response with the site of a push rod. 


While it could not be ascertained with certainty that 
particular responses arose from within a push rod, two 
types of responses observed were typical of endings 
found at the base of push rods and identified as such 
in other mammals. In our study we classified responses 
into 4 types, based on similarities to the 4 main types 
of response seen in other mammals and for which the 
morphological identity of the receptors has been 
established. 


Proc. Linn. Soc. N.S.W., 125, 2004 


U. PROSKE AND J.E. GREGORY 


The first type of receptor was characterised 
by a rapidly adapting response to mechanical 
stimulation of the bill skin and a high sensitivity to 
vibration at frequencies up to about 800 Hz (Fig. 3). 
This type of response is characteristic of Pacinian and 
paciniform corpuscles in other mammals and can 
confidently be ascribed to the paciniform corpuscles 
found in the snout skin and especially prominent at 
the base of pushrods. 

The second type of response was an irregular, 
slowly adapting discharge (Fig. 3). These SAI 
responses have been known for some time to be 
generated in other mammals by Merkel cell receptors, 
the second ending type found prominently at the base 
of push rods. The examples encountered in the echidna 


typically had very small receptive fields, with a © 


diameter well below 100 um and a threshold for a 
response to skin displacement of 4 um. Such small, 
low threshold fields are consistent with their belonging 
to Merkel receptors at the base of push rods that are 
somewhat shielded from stimuli not directly applied 
to the push rod tip. 

The most numerous type of response to 
mechanical stimulation was a slowly adapting, regular 
discharge, termed a Type II response in other 
mammals. These have been identified with Ruffini 
endings in the dermis, signalling preferentially stretch 
of the skin. A similar morphological type is present in 
echidna snout skin, but not in direct association with 
push rods. 

The fourth type of response was a rapidly 
adapting discharge to skin stimulation. These receptors 
were distinguished from the other rapidly adapting 
group by not showing a sensitivity to vibration at high 
frequencies, and they responded well only to 
frequencies below 300 Hz. 

The study shed no light on the function of 
vesicle chain receptors. Perhaps they are the rapidly 
adapting receptors unresponsive to high frequencies 
of vibration, but the number found seemed too few, at 
only about 15% of the sample studied. It may be that 
one or both types of vesicle chain receptor are 
responsible for at least some of the Type II slowly 
adapting responses, which accounted for nearly half 
the sample studied. What can be said is that no 
completely new mechanoreceptive responses, 
unknown in other species, were seen in the echidna, 
either in this study or in an earlier one by the same 
group (Iggo et al. 1985). 

There is some evidence that the sensitivity of 
the push rod complex to external stimuli is not fixed, 
but can be varied, and in two ways. First, in both 
echidna and platypus, there is a venous cavernous 
system in the bill or snout, beneath the skin. It is 
postulated that engorgement of the venous sinus may 


Proc. Linn. Soc. N.S.W., 125, 2004 


change the mechanical properties of the skin, affecting 
transmission of mechanical stimuli to the receptors at 
the base of the push rod and effectively changing its 
sensitivity. It may also lead to protrusion of rod tips 
beyond the skin surface, to allow them to present more 
effectively to environmental stimuli. 

Secondly, Manger et al. (1998) have 
described a ring of contractile material around the tip 
of the platypus push rod. Perhaps this is used to restrict 
the mobility of the push rod and in this way also change 
its effective sensitivity. Why might it be desirable to 
change the sensitivity of the push rod by altering the 
mechanical coupling between the stimulus and the 
receptors? Perhaps the answer for the platypus lies in 
the different environments, air and water, the animal 
inhabits. An appropriate sensitivity for one medium 
may not be optimal for the other and the platypus may 
have evolved a means of adjusting between the two. 
A similar argument could apply to the echidna, which 
uses the tactile receptors in the snout both in the ground 
and above it. However there are no reports of the 
presence of contractile cells in association with echidna 
push rods, which, anyway, seem to be much less 
mechanically independent of the surrounding skin. 


COMPARISONS WITH OTHER ANIMALS 


The push rod theme reaches its most 
exuberant expression in the star-nosed mole. 
Protruding from the snout of this animal is an 
extraordinary array of 22 radiating fleshy pink fingers 
entirely covered with thousands of small domes. These 
domes are the tips of Eimer’s organs, which have a 
remarkable similarity to the push rods in monotremes, 
except for containing fewer sensory endings and being 
innervated by fewer sensory nerve fibres than push 
rods (Fig. 4). There is a single Merkel receptor at the 
base and a single lamellated receptor immediately 
underneath in the dermis. A single vesiculated nerve 
terminal runs up the centre of the Eimer’s organ and 5 
- 10 at the periphery (Catania 1995). 

Eimer’s organs are believed to be tactile sense 
organs, responsive to the onset and offset of depression 
of the papilla and to sustained compression (Catania 
& Kaas 1995). 

The star-nosed mole is a burrowing 
insectivore living almost entirely underground in 
swamps and bogs. As in the monotremes, the function 
of the vesiculated receptor terminals has not been 
conclusively established. It is assumed that the star is 
a sensitive tactile organ used by the mole to explore 
its environment for prey. Some other members of the 
family Talpidae, moles, shrew moles and desmans, 
have Eimer’s organs (Catania 2000), but only the star- 


323 


ROLE OF PUSH RODS IN MONOTREMES 


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Figure 4. Schematic of an Eimer’s organ in the star-nosed mole, showing the single Merkel cell and 
lamellated corpuscle, as well as the central and peripheral neural processes similar to the central and 
peripheral vesicle chain receptors of the monotreme push rod. (Modified from Catania, 1995.) 


nosed mole has developed such an elaborate structure 
to deploy them. 


FUNCTIONAL CONSIDERATIONS 


If it is accepted that push rods subserve a 
mechanosensory function and are designed to raise 
fidelity of transmission of static and dynamic stimuli, 
it poses the question of why such an arrangement is 
needed. Speculation about the mechanoreceptors in 
monotremes has focussed on two aspects. 


324 


In the platypus, there is a close association 
between the electrosensory and tactile senses, and cells 
in the cerebral cortex have been found that receive 
inputs from both modalities (Iggo et al. 1992; Manger 
et al. 1996). Pettigrew et al. (1998) have postulated a 
system for determining the distance and perhaps 
direction of live objects underwater, based on a 
cooperation between the two senses. An animal like a 
shrimp would generate both an electrical pulse and a 
mechanical pressure wave when it flicked its tail. The 
electrical pulse would arrive at the platypus first and 


Proc. Linn. Soc. N.S.W., 125, 2004 


U. PROSKE AND J.E. GREGORY 


be detected by the electroreceptors. After a delay 
depending on the shrimp’s distance, the mechanical 
pressure wave would arrive and the receptors in the 
pushrods would be stimulated. Suitably tuned cells in 
the cortex would detect the delay between the two 
inputs, and thus distance could be computed. 
Directional information could also be derived if the 
push rods have a directional sensitivity as speculated, 
and this would be combined with the hypothesised 
directional information derived from _ the 
electroreceptors (Manger et al. 1996). 

A second possible role for push rods concerns 
our speculation about close-range electrolocation 
(Proske and Gregory, 2003). As the platypus fossicks 
about on the bottom of a stream or pond, pushing its 
bill into the detritus looking for live prey, its 
electroreceptors will allow it to distinguish animate 
from inanimate objects. It is conceivable that the 
alerting signal from the electroreceptors receives 
confirmation of the presence of a prey item from the 
detailed mechanosensory signals provided by push 
rods. Similarly for the echidna, there may be a 
functional cooperativity between inputs from 
electroreceptors and mechanoreceptors as the animal 
pushes its nose into the soil in its search for prey. 

Speculating more broadly about push rods as 
vehicles for mechanoreceptor stimulation, it is of 
interest that they are located exclusively in bill skin of 
the platypus and in nose skin of the echidna. Being at 
the front end of the animal, perhaps they subserve some 
kind of teletactile function, that is, detection of 
disturbances in the water or in the soil created by prey 
at some distance ahead of the animal. 

Vertebrates have evolved a number of ways 
of stimulating, at a point, a population of receptors 
with both static and dynamic properties. An example 
that comes to mind is the vibrissae of mammals. Here 
it is known that the various afferents supplying each 
vibrissa project to the same area of cerebral cortex to 
form identifiable ‘barrels’ (Miller et al. 2001). It 
suggests that central processing of tactile information 
coming from these structures requires the presence of 
both static and dynamic components of the stimulus 
and the processing is done by neurones lying in close 
proximity to one another. 

For push rods, the presence of a mobile 
column of epidermal cells that is able to excite 
paciniform and Merkel cell receptor types at the same 
time, provides the opportunity for similar central 
processing of static and dynamic stimuli. It remains to 
determine why it is necessary to process the 
information in this way. Presumably the nature of the 
sensory experience evoked by stimulation of a push 
rod is dependent on the simultaneous presentation of 
static and dynamic features of the stimulus. 


Proc. Linn. Soc. N.S.W., 125, 2004 


REFERENCES 


Andres, K. and Von During, M. (1984). The platypus bill. 
A structural and functional model of a pattern- 
like arrangement of different cutaneous sensory 
receptors. In Sensory Receptor Mechanisms (ed. 
W. Hamann and A. Iggo). World Scientific, 
Singapore. 

Andres, K.H., Von During, M., Iggo, A. and Proske, U. 
(1991). The anatomy and fine structure of the 
echidna Tachyglossus aculeatus snout with 
respect to its different trigeminal sensory 
receptors including the electroreceptors. 
Anatomy and Embryology 184, 371-393. 

Catania, K.C. (1995). Structure and innervation of the 
sensory organs on the snout of the star-nosed 
mole. Journal of Comparative Neurology 351, 
536-548. 

Catania, K.C. (2000). Epidermal sensory organs of moles, 
shrew moles, and desmans: a study of the family 
talpidae with comments on the function and 
evolution of Eimer’s organ. Brain, Behavior & 
Evolution 56, 146-174. 

Catania, K.C. and Kaas, J.H. (1995). Organization of the 
somatosensory cortex of the star-nosed mole. 
Journal of Comparative Neurology 351, 549- 
567. 

Iggo, A., Gregory, J.E. and Proske, U. (1992). The central 
projection of electrosensory information in the 
platypus. Journal of Physiology 447, 449-465. 

Iggo, A., Gregory, J.E. and Proske, U. (1996). Studies of 
mechanoreceptors in skin of the snout of the 
echidna Tachyglossus aculeatus. 
Somatosensensory and Motor Research 13, 129- 
138. 

Iggo, A., McIntyre, A.K. and Proske, U. (1985). 
Responses of mechanoreceptors and 
thermoreceptors in skin of the snout of the 
echidna Tachyglossus aculeatus. Proceedings of 
the Royal Society of London - Series B: 
Biological Sciences 223, 261-277. 

Manger, P.R., Calford, M.B. and Pettigrew, J.D. (1996). 
Properties of electrosensory neurones in the 
cortex of the platypus (Omithorhynchus 
anatinus): implications for processing of 
electrosensory stimuli. Proceedings of the Royal 
Society of London - Series B: Biological 
Sciences 263, 611-617. 

Manger, P.R., Keast, J.R., Pettigrew, J.D. and Troutt, L. 
(1998). Distribution and putative function of 
autonomic nerve fibres in the bill skin of the 
platypus (Ornithorhynchus anatinus). 
Philosophical Transactions of the Royal Society 
of London - Series B: Biological Sciences 353, 
1159-1170. 

Manger, P.R. and Pettigrew, J.D. (1996). Ultrastructure, 
number, distribution and innervation of 
electroreceptors and mechanoreceptors in the 
bill skin of the platypus, Ornithorhynchus 
anatinus. Brain, Behavior & Evolution 48, 27- 
54. 


325 


ROLE OF PUSH RODS IN MONOTREMES 


Miller, B., Blake, N.M.J. and Woolsey, T.A. (2001). 
Barrel cortex: structural organisation, 
development and early plasticity. In Plasticity of 
Adult Barrel Cortex (Ed. M. Kossut), pp. 3-45. 
(F.P. Graham, Johnson City TN, USA). 

Pettigrew, J.D., Manger, P.R. and Fine, S.L. (1998). The 
sensory world of the platypus. Philosophical 
Transactions of the Royal Society of London - 
Series B: Biological Sciences 353, 1199-1210. 

Proske, U. (1997). Echidna on the nose. Nature Australia 
25, 58-63. 

Proske, U. and Gregory, J.E. (2003). Electrolocation in 
the platypus — some speculations. Comparative 
Biochemistry and Physiology 136, 821-825. 


326 Proc. Linn. Soc. N.S.W., 125, 2004 


Obituary 


MERVYN EDWARD GRIFFITHS 
1914-2003 


Mervyn Griffiths (who always preferred to be 
called Merv”) was born in Sydney on 8" July 1914, 
was educated at North Sydney Boys’ High School and 
obtained his Bachelor Degree in Zoology with first 
Class Honours in 1937, followed by his Master of 
Science in 1938 at Sydney University 

Merv first began publishing in the scientific 
literature in 1936 with a paper on The colour changes 
in batoid fishes in the Society’s Proceedings and 
contributed six further papers to this journal between 
1936 and 1942. 
After completing his Master studies, Merv was 
awarded the Travelling Research Scholarship for the 
1851 Exhibition, which took him to McGill University 
in Montreal, Harvard University and the National 
Institute for Medical Research in London. From this 
work he produced 10 papers, including research on 
diabetes mellitus and the secretory functions of the 
pituitary. In 1941 Merv returned to the University of 
Sydney where he continued his work on diabetes in 
the Department of Medicine, where he was a Linnean 


Macleay Fellow in Physiology in 1941. The fellowship - 


was renewed in 1942, but only a few months into his 
second fellowship year he joined the Royal Australian 
Air Force. 

Merv was in the Empire Air Training Scheme 
during 1943 in Edmonton, Canada and became a Pilot 
Officer in 1944. He held the post of Commanding 
Officer of the 3rd Malaria Control Unit in Darwin from 
1944-45 and left the RAAF in February 1946 at the 
age of 31, continuing his involvement in diabetes 
research as a Zoologist in the Institute of Anatomy in 
Canberra. He became the Senior Biochemist at the 
Institute in 1949, publishing his work on the 
biochemistry of diabetes through to 1957, when he 
returned to zoology, joining the C.S.I.R.O. Wildlife 
Survey Section [which later became the Division of 
Wildlife Research] as a Senior Research Officer in June 
WT 

In 1959 he was awarded his Doctor of Science 
Degree by Sydney University for his thesis entitled 
The Relationship of the Pituitary Gland to 
Experimental Diabetes and the Action of Insulin. 

Although his initial published works at the 
C.S.LR.O. were concerned with rabbits, Merv became 


interested in the biology of marsupials, particularly 
macropods. In 1965 he published his first paper on 
the biology of the group for which his research is best 
known, and which became his consuming passion - 
the Monotremes. His monograph on the Echidnas was 
published in 1968. During his time at the Division of 
Wildlife Research, Merv was the scientific director of 
two films, The Echidna and the Comparative Biology 
of Lactation, both of which won awards. 

Merv retired in October 1975 from the 
C.S.LR.O. Division of Wildlife Research as Senior 
Principal Research Officer, but not from the field of 
wildlife research. His interests broadened, although the 
monotremes remained his prime focus. His classic 
work The Biology of the Monotremes in 1978 pulled 
together all of the disparate research carried out on 
the group to that time. 

Merv was variously honoured by scientific 
societies, including being awarded the Peter Aitken 
Medal by the South Australian Museum in 1988 and 
becoming a Fellow of the Royal Zoological Society 
of N.S.W. in 1991. In his “retirement” Merv researched 
and published widely in a number of fields, where his 
own work and collaboration with colleagues and 
friends resulted in a further 33 publications to add to 
his pre-retirement total of forty-three. 

Merv Griffiths died on 6" May 2003. The above 
summary of his academic life cannot adequately 
describe his contribution to biological science in 
Australia. Throughout his academic career Merv 
remained a great “generalist” in a world of increasing 
numbers of scientific “specialists”. His encouragement, 
generous advice, support and sometimes cajoling, are 
deeply appreciated by many of these specialists. 


Tom Grant 
Sydney 
May 2003 


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BOOK REVIEW 


Admiral Doenitz’s Legacy 


Paul Adam 
School of Biological, Earth and Environmental Sciences, University of New South Wales, NSW 2052 


Murray, D.R. 2003 Seeds of concern. The genetic manipulation of plants. UNSW Press.156pp. ISBN 0 
86840 460 8. $34.95. 


Over the last decade there has been increasing 
public concern about the development and use of 


genetically modified organisms. This has been | 


manifest in media coverage involving ‘shock-horror’ 
headlines such as ‘Frankenfoods’, litigation, product 
boycotts, increased sales of ‘organic’ products, illegal 
destruction of genetically modified crops and, recently, 
the rejection by Zambia of food aid which may have 
contained genetically modified grains. Opposition has 
been global, but at its most intense in Europe. 

What is the basis for the concern and is it 
justified? 

Some of the concerns are clearly unnecessary, 
reflecting an unfortunate lack of basic scientific 
knowledge amongst the media and general public. 
Alarmist stories about how eating genetically modified 
organisms involves consuming DNA, as if non- 
modified organisms lack DNA, do not add to the 
credibility of journalists or their editors. For products 
which are extracted, highly refined and purified, 
whether their origin is from modified or unmodified 
organisms is irrelevant in terms of the end qualities 
and properties. 

Other issues have more substance, but at least 
in Europe, the GM debate is only a symptom of a much 
broader concern about the nature of modern 
agriculture. Public confidence has been disturbed by 
the outbreaks of both BSE and Foot and Mouth disease, 
which are seen as components of a broader malaise. 
Neither disease of course has anything to do with 
genetic modification, although particularly in the case 
of BSE, significant portions of the media and the public 
believe it does. 

To understand the origins of the perceived 
malaise it is necessary to go back to the First World 
War. For the first time submarines proved to be an 
important weapon, and convoys of food supplies were 
disrupted. Between the wars submarine technologies 
were substantially developed. Agriculture was little 
changed, indeed for much of the period European 
agriculture was in decline, as cheap imports from north 
America and the southern hemisphere satisfied the 


market. During the Second World War U-boats 
maintained a blockade which almost brought Britain 
to defeat. 

The post war response was — ‘never again’, 
and the still prevailing policy of self sufficiency was 
developed. Governments plan to win the last war, so 
the fact that the weapons of mass destruction which 
brought the Second World War to an end changed the 
nature of any future global war, did not influence policy 
development. 

From some perspectives the self sufficiency 
policy could be judged a success. Who in 1945 would 
have anticipated the vast European Union surpluses, 
or that in 2003 British livestock-feed grains would be 
exported to Australia? Nevertheless it is the cost of 
this success which is now being questioned. 

The drive for increased production and 
efficiency was powered by subsidies, both direct and, 
indirect, and the developing agribusiness companies. 
Synthetic pesticides and herbicides (such as DDT and 
MCPA) were first used on a large scale towards the 
end of the war, and in the immediate post war years 
usage burgeoned, being proclaimed as an example of 
the new scientific approach to farming. There was little 
external scrutiny of government programs and the 
administrative bureaucracies were captive to their 
clients — farmers and agribusiness. 

The first expressions of concern surfaced with 
the publication of Rachel Carson’s Silent Spring 
(1962). Both the publication and its author were subject 
to sustained attack by both government and 
agribusiness, but the basic thesis was increasingly 
supported by independent evidence. The publication 
of Silent Spring was one of the key events leading to 
the modern environmental movement, resulted in the 
banning in the west, if not globally, of some pesticides 
and the institution of greater scrutiny of new chemicals. 
Nevertheless these were only minor hiccups on the 
way to industrialization of agriculture. 

Other changes included, in northern Europe, 
the decline of mixed farming in favour of 
specialization, the loss of woodlands, hedgerows and 


BOOK REVIEW - SEEDS OF CONCERN 


wetlands, the loss of genetic diversity amongst crops 
as many local varieties were replaced by a few new 
cultivars, and increased use of nitrogen fertilizers 
resulting in greener, but floristically simpler, basically 
Lolium monocultures, pastures. New crops came to 
prominence, most notably oilseed rape (known in more 
sensitive nations such as Australia as canola) 
converting the landscapes of Constable to ones more 
akin to those of van Gogh. Lifestock production was 
increased through adoption of so-called factory 
farming techniques, changing, for example, chicken 
from a luxury to a convenience food but raising 
widespread community concerns about animal welfare 
and creating substantial environmental problems 
associated with effluent management. 

The changing face of the countryside, 
increasing awareness of the impacts on biodiversity, 
and concerns about potential impacts on human health 
has led to an upsurge of public disquiet (Shoard 1980, 
Harvey 1997, 2001, Humphrys 2001, Green 2002), 
but reform of the European agricultural system, 
although debated for several decades has been slow to 
eventuate. The idealized countryside of an urban 
population is often a bucolic dream, the product of a 
Romantic imagination, and ignores the earlier 
extensive changes wrought by the Agricultural 
Revolution and enclosures (Fox and Butlin 1979), but 
the issues raised by commentators such as Harvey 
(1997) are nevertheless well documented and cannot 
lightly be dismissed. 

However, the public concern over the 
consequences of agricultural policy runs contrary to 
the public expectation, also developed since 1945, of 
a never ending supply of cheap food. This expectation 
is well summarized by Gummer (2001) (John Gummer 
is a former UK Minister for Agriculture and Secretary 
of State for the Environment). 

“Many in the rich world, who do not blanch 
at forking out £30,000 for a more fashionable motor 
car, will refuse to expend threepence more on a fresher 
lettuce or a tastier loaf of bread. Food, which ought to 
rank highest among our spending priorities, has been 
relegated to the rank of necessity, and in this advanced 
civilization of ours, only luxuries deserve to be prized. 
We take necessaries as our right and expect them to 
be delivered at a discount. 

So it is that food prices demand a smaller and 
smaller proportion of a prosperous household’ s income 
and take less time than ever for the average worker to 
earn. What is more, now that packaging, distribution 
and preparation are necessary on-costs, the basic food 
content of what we buy represents an even smaller 
proportion of what we pay. With one in three meals 
eaten out of the home and most of the rest to a growing 


330 


extent pre-prepared, that proportion will continue to 
fall”. 

In demanding cheap food the public neglects 
to take into consideration the considerable subsidies 
paid out of the taxpayers’ pocket to farmers by 
governments. 

The trend to cheap food has also been assisted 
by the growth of supermarket chains. For many 
foodstuffs, prices to farmers are determined by a global 
oligopoly of retailers. With the availability of cheap 
airfreight a further consequence of the growth of the 
supermarket chains is the abolition of seasons, with 
the same range of produce being available globally 
year round. This means that the shelves of European 
supermarkets may be replete with sugar peas from 
Zambia, green beans from Kenya, salad greens from 
Tanzania and southern Africa, even brussel sprouts 
from Australia. When the costs of freight and 
packaging are considered the return to farmers must 
be very small, and does not take into account 
environmental degradation and disruption of traditional 
agricultural economies. Additionally agriculture in 
developing countries is adversely affected by dumping 
of surpluses from elsewhere — tomato producers in 
West Africa for example cannot compete even in their 
local market against heavily subsidized Italian canned 
tomatoes (Bradshaw 2003). 

There is little doubt that if global agriculture 
had remained as it was in 1945 it would have been 
impossible to support the current human population 
of about six billion. Currently global food production 
is capable of providing adequate nutrition for the world 
population (Waterlow ef al. 1998), famine and 
malnutrition are the consequences of failures of 
political systems, not of agriculture. What is less certain 
is whether, without extensive adoption of new 
molecular techniques, it will be possible to feed the 
predicted human population in 2050 of eight billion 
(Waterlow et al. 1998), particularly given the added 
impacts of global warming. 

The increased production is due in part to an 
increase in the land area devoted to agriculture but is 
largely the consequence of new technologies. The 
development of agricultural chemicals, mechanization 
and application of conventional plant and animal 
breeding can rightly be regarded as scientific and 
engineering triumphs. Nevertheless broader questions 
of ecological sustainability were not part of the agenda 
until recently, and the true costs of food production 
have rarely been calculated. 

How is this discussion relevant to any debate 
about use of genetically modified organisms in 
Australia? 

Firstly, despite talk about free trade and level 


Proc. Linn. Soc. N.S.W., 125, 2004 


P. ADAM 


playing fields, world agricultural markets are still 
heavily influenced by government subsidies and policy 
intervention. Whether the original motivation for these 
government programs can still be justified (and in the 
case of the much maligned European Common 
Agricultural Policy, maintenance of the social structure 
of rural communities was as important as ensuring self- 
sufficiency) it is unlikely that there is political will for 
change. Australia’s agricultural future, both in terms 
of access to overseas markets and ability to control 
imports, will be very much determined by what 
happens in Washington and Brussels. The US is 
currently seeking action by the World Trade 
Organisation to require the European Union to lift its 
restrictions on genetically modified food (Sanger 
2003). The outcome of these proceedings will have 
global implications, determining for example whether 
Australia could similarly impose controls on imports 
of genetically modified organisms. President Bush has 
argued that Europe’s approach has discouraged use of 
genetically modified foods in the third World and this 
contributed to continuing famine in Africa (Sanger 
2003). Most would argue that African famines have a 
number of causes, and that absence of genetically 
modified crops is not one of them. There will be 
continuing pressure for increased efficiency (as 
measured in production of cheap food) and this will 
have social and economic consequences. Secondly 
agribusiness is global, and increasingly vertically 
integrated, so that much plant breeding and 
development of genetically modified plants will be in 
the hands of a few multinationals, with the results 
imported into Australia. The growing hegemony of a 
few agribusiness company has considerable 
implications beyond the issue of genetic modification. 
Any choices that farmers might have in terms of crop 
varieties or chemicals are becoming increasingly 
illusory. President Eisenhower famously warned of the 
influence of the military industrial complex — his words 
would equally apply to agribusiness. 

To date, the introduction of genetically 
modified crops in Australia has been, as it also has 
been in Europe, a public relations disaster for 
agribusiness. However, information about the nature 
of the genetically modified crops has not been readily 
accessible to the public. 

David Murray’s “Seeds of Concern’ attempts 
to fill the gap, with the objective of promoting informed 
debate. The author is a distinguished plant scientist so 
his critique cannot be dismissed by the more 
enthusiastic proponents of the technology as well 
intentional but misinformed. However, while the book 
is aimed, in part, at the intelligent lay person, the 
assumed level of chemical/biochemical knowledge is 


Proc. Linn. Soc. N.S.W., 125, 2004 


high and may well deter the intended audience. 

The technology to achieve genetic 
manipulation exists, and the genie cannot be put back 
in the bottle. Murray recognizes this and discusses 
possibilities for using the technologies which could 
potentially be of considerable benefit to humankind. 
Unfortunately the benefits of the majority of 
genetically modified plants released to date accrue to 
agribusiness and farmers rather than the consumer. 
Genetic manipulation is possible in the whole range 
of organisms. Modification of microorganisms for 
industrial processes has not attracted much public 
attention, modification of domestic animals (including 
farmed fish) is still largely at the trial stage, but 
genetically modified plants are in the landscape and 
market place and very much in the public eye. 

Murray provides a very succinct introduction 
to plant cell biology and the techniques of genetic 
manipulation. In commercially released genetically 
modified plants the most frequent changes are the 
introduction of herbicide resistance or of genes which 
express insecticidal compounds. Murray explains the 
underlying basis for both changes, but also discusses 
actual and potential drawbacks with the use of plants 
modified in these ways in the field. It is far from clear 
that in the long term these approaches will have benefit. 
The number of potential applications of genetic 
modification which have been touted in the media is 
very large. Murray explains how in some cases, such 
as lowering caffeine levels in coffee or preventing 
expression of polyphenol oxidases, the proponents are 
ignorant of the biological function of these compounds 
or of consumer requirements. The yield costs of genetic 
modification are often, as Murray points out, given 
little consideration. What Murray regards as failings 
of current patenting regimes and of Australia’s Plant 
Breeders Rights legislation are discussed at some 
length. I would support the critique although 
recognizing that commercial interests would be 
expected to take a different view. These difficulties 
also arise with conventional plant breeding, but, with 
the heavy investment in genetic modification are likely 
to be more apparent as companies seek to protect what 
they regard as their intellectual property. 

The regulatory framework for release of 
genetically modified organisms currently applying in 
Australia is discussed in some detail. While this regime 
is important, Murray does not raise the issue of general 
lack of regulation of many agricultural activities. 
Farmers world wide argue that they are over regulated, 
and arguments in favour of less regulation abound (see 
Ridley 2001, Pennington 2001, and the continuing 
opposition by farmers in Eastern Australia to control 
of land clearing). While it is the case that some aspects 


331 


BOOK REVIEW - SEEDS OF CONCERN 


of agriculture are heavily regulated, key decisions are 
left to farmers. The cover of ‘Seeds of Concern’ shows 
a field of canola (rape); the decision of European 
farmers to adopt broad acre rape cultivation had a 
profound visual and ecological impact, but was not 
one in which the broader community participated. The 
change in the northern hemisphere from spring to 
winter cereal cultivation has had major effects on many 
bird species, but again was a decision of farmers and 
agribusiness. In the Australian context changes from 
pastoralism to cropping, or the spread of cotton 
growing (both changes which may be facilitated by 
the development of genetically modified plants) will 
have social, ecological and environmental 
consequences, but the decisions will be made by 
landholders with few avenues for external scrutiny, 
let alone any requirement for approval. Even if the 
specific concerns of the Gene Technology Regulator 
are met, the broader questions raised by agricultural 
change will remain unaddressed. These are clearly 
issues beyond those which Dr. Murray set out to 
address, but they do need to be placed on the policy 
agenda. 

Dr. Murray also manages within the compass 
of his admirably concise book to mount a defence of 
Mendel against claims that his results might have been 
‘polished’, express skepticism about proposals to clone 
thylacines, and have a few swipes against the 
pontifications of the great and the good. Despite in a 
few places being ill served in matters of layout by the 
publisher, this is an important contribution to the debate 
about genetically manipulated plants and of the future 
of agriculture. 

There are indeed ‘seeds of concern’, perhaps 
leading to seeds of doubt. 


REFERENCES 


Bradshaw, S. (2003) Unfair meal gives taste of global trade 
pitfalls. Broadcast on ABC ‘Landline’ 13 April 
2003. Transcript at http://www.abc.net.au/ 
landline/stories/s828402.htm 

Carson, R. (1962) Silent spring. (Houghton Miflin, Boston). 

Fox, H.S.A. & Butlin, R.A. (Eds) (1979) Change in the 
countryside. (Institute Of British Geographers, 
London). 

Green, B. (2002) The farmed landscape: the ecology and 
conservation of diversity. In Remaking the 
landscape. The changing face of Britain. Ed. J. 
Jenkins pp.183-210. (Profile Books, London). 

Gummer, J. (2001) Farming and the CAP. In A countryside 
for all. The future of rural Britain. (Ed. M. 
Sissons). pp.57-68. (Vintage, London). 

Harvey, G. (1997) The killing of the countryside. (Jonathan 
Cape, London). 


332 


Harvey, G. (2001) Reinventing agriculture. In A countryside 
for all. The future of rural Britain. (Ed. M. 
Sissons). pp.77-88. (Vintage, London). 

Humphrys, J. (2001) The great food gamble. (Hodder and 
Stoughton, London). 

Pennington, M. (2001) Deregulating the land: an alternative 
route to urban and rural regeneration. In A 
countryside for all. The future of rural Britain. 
(Ed. M. Sissons). pp.35-47. (Vintage, London). 

Ridley, M. (2001) Denationalising the land. In A countryside 
for all. The future of rural Britain. (Ed. M. 
Sissons). pp.25-33. (Vintage, London). 

Sanger, D. (2003) Bush blames European rules on GM food 
for hunger in Africa. The Sydney Morning Herald, 
May 23. p.12. 

Shoard, M. (1980) The theft of the countryside. (Temple 
Smith, London). 

Waterlow, J.C., Armstrong, D.G., Fowden, L. & Riley, R. 
(Eds) (1998) Feeding a world population of more 
than eight billion people. A Challenge to science. 
(Oxford University Press, Oxford). 


Proc. Linn. Soc. N.S.W., 125, 2004 


BOOK REVIEW 


M.L. Augee 
Wellington Caves Fossil Studies Center, 89 Caves Road, Wellington NSW 2820 


Duyker, Edward. 2003. Citizen Labillardiére: a naturalist’s life in revolution and exploration (1755-1834) 
Edward Duyker. The Miegunyah Press, Melbourne 2003. $59.95 


This book is a splendid example of the need 
for academic presses in Australia; Miegunyah Press is 
an imprint of Melbourne University Publishing. It is a 
scholarly, extremely well documented work covering 


a poorly known but important aspect of Australia’s ~ 


early history. As such it cannot be considered an “easy 
read” nor a likely undertaking for a commercial 
publisher. It is however a goldmine of information and 
will be a valuable reference source for anyone seeking 
information on the roots of natural history study in 
Australia or the contributions of French scientists to 
such study. The use of this book as a reference is greatly 
facilitated by three complete indexes (botanical, 
zoological and general) and a fully detailed 
bibliography. The amount of research into resources 
in several languages required to produce this book is 
amazing. Such detail insures its use for many years to 
come. 

“Citizen Labillardiére” can be read simply as 
a biography of a very interesting man who lived 
through very interesting times. He stands astride the 
“troubles” in France as the country shifted from the 
Ancien Régime to the National Assembly and 
constitutional monarchy; to a republic overtaken by 
the “Reign of Terror’; to an Empire (Napoleon); to 
the restoration of the monarchy; and then to the final 
overthrow of the Bourbons. The “citizen” in the book 
title indicates Labillardiére’s general political leaning, 
but it is also the story of a man of science trying for 
the middle road in turbulent politics. It is therefore a 
story of survival and adventure. Much of the adventure 
comes from travel, often arduous, over much of the 
globe. 

After graduating in medicine (1772), 
Labillardiére immediately begins his life of travel with 
ajourney to England. There he establishes connections 
with Joseph Banks and other British scientists that will 
serve him well in the future. He travels through the 
Alps and Lebanon, always collecting botanical 
specimens and possessing a remarkable enthusiasm for 
climbing mountains. Then comes his big break; he is 
appointed as a naturalist on the expedition to be lead 
by Bruny d’ Entrecasteaux in search of the missing La 
Perouse (last seen by the British at Botany Bay on 10 


March 1788). This small fleet, consisting of the 
Recherche and the Espérance, left France on 28 
September 1791. After stops at Tenerife and the Cape 
of Good Hope, where Labillardi¢ére made many 
collections, they landed in Van Diemen’s Land, at 
Recherché Bay, on 20 April 1792. There Labillardiére 
collected 5,000 specimens in five weeks. Sailing north 
in search of La Perouse, the expedition went to New 
Caledonia, New Guinea and the Solomons before 
returning to Recherche Bay. At that time Labillardiére 
records positive encounters with the native people; 
indeed he seems to have been a sympathetic and astute 
observer of the Tasmanian natives. 

Leaving Van Diemen’s Land, the expedition 
went to Tonga, passing close by the tip of New 
Zealand, and to New Caledonia. A return to Van 
Diemen’s Land, with the great likelihood of 
discovering that it was not part of the mainland and 
discovering the strait later to be discovered by Bass, 
was not to be, and d’Entrecasteaux died at sea on 19 
July 1793. Already falling apart with rifts between 
royalists and republicans, the expedition died when, 
upon reaching Java, the Dutch were found to be at 
war with France. The Dutch imprisoned Labillardiere 
and others, although not the royalists who collaborated 
with them. Labillardiére was finally released on 29 
March 1795 and, after spending six months at Ile-de- 
France enroute, was back in Paris in March 1796. 

Meanwhile most of his collection of botanical 
specimens had been taken to England and were claimed 
by the royalist French court in exile there. Banks 
however used his strong influence and the collection 
was returned to Labillardiére. 

There was an odd interlude in the life of 
Jacques-Julien Labillardiére when, in the summer of 
1796, he went to Italy as part of a commission to 
oversee seizure of Italian art and other treasures taken 
as tribute after General Napoleon Bonaparte conquered 
that country. In 1800 Labillardiére was made a full 
member of the Académie des sciences. 

Besides the political turmoils of his time, 
Labillardiére also straddled a revolution in biological 
science, during the change from strict application of 
Linnean principles (which saw taxonomy as a rigid 


BOOK REVIEW - CITIZEN LABILLARDIERE 


set of categories, determined in the case of plants by 
the structure of the sexual parts of flowers) to a more 
natural approach (systematics including as much 
evidence as possible, especially physiological and 
morphological). Labillardiére’s “Sertum austro- 
caledonicum” published in 1824 abandoned the strict 
Linnean approach. It should be noted, however, that 
well before that time zoologists (such as the towering 
figure of Baron Cuvier) had widely accepted the natural 
system. Not unlike today when about the only 
resistance to modern systematics, such as cladistics, 
comes from botanical taxonomists, many of whom one 
suspects strongly regret that upstart Antoine-Laurent 
de Jussieu who started tinkering with the Linnean 
system in 1789. 

Labillardiére died 8 January 1834 aged 79. 
He had a truly adventurous life and made a significant 
contribution to the advancement of science, although 
the claim by Duyker that he was “one of the founders 
of botany, zoology and ethnography in Australia” is 
overly generous. Labillardiére was basically a collector 
with a sometimes off-hand approach to record keeping. 
Nonetheless, he did record many “firsts” and many 
names he gave to plants are valid today. 


334 


Proc. Linn. Soc. N.S.W., 125, 2004 


INSTRUCTIONS FOR AUTHORS 


(this is an abbreviated form — the full instructions can be obtained from our web site or from the Secretary) 


1. The Proceedings of the Linnean Society of New South Wales publishes original research papers dealing 
with any topic of natural science, particularly biological and earth sciences. 


2. Manuscripts should be submitted to the Editor (M.L. Augee, PO Box 82, Kingsford NSW 2032). All 
manuscripts are sent to at least two referees and in the first instance three hard copies, including all figures 
and tables, must be supplied. Text must be set at one and a half or double spacing. 


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4. References are cited in the text by the authors’ last name and year of publication (Smith 1987). For three 
of more authors the citation is (Smith et al. 1988; Smith and Jones 2000). Notice that commas are not used 
between the authors’ names and the year. The format for the reference list is: 
Journal articles: 
Smith, B.S. (1987). A tale of extinction. Journal of Paleontological Fiction 23, 35-78. 
Smith, B.S., Wesson, R.I. and Luger, W.K. (1988). Levels of oxygen in the blood of dead Ringtail 
Possums. Australian Journal of Sleep 230, 23-53. 
Chapters or papers within an edited work: 
Ralp, P.H. (2001). The use of ethanol in field studies. In ‘Field techniques’ (Eds. K. Thurstle and P.J. 
Green) pp. 34-41. (Northwood Press, Sydney). 
Books: 
Young, V.H. (1998). “The story of the wombat’. (Wallaby Press, Brisbane). 


5. An abstract of no more than 200 words is required. Sections in the body of the paper usually include: 
INTRODUCTION, MATERIALS AND METHODS, RESULTS, DISCUSSION, 
ACKNOWLEDGEMENTS and REFERENCES. Some topics, especially taxonomic, may require variation. 


6. Subheadings within the above sections should be in the form: 
Bold heading set against left margin 
This is the form for the first level headings and the first line of text underneath is indented 
Underlined heading set against left margin 
This is the next level, and again the first line of text underneath is indented. 
Further subheadings should be avoided. 
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7. Up to 10 KEYWORDS are required. These are often used in computer search engines, so the more 
specific the terms the better. ‘Australian’ for example is useless. Please put in alphabetical order. 


8. Paragraphs are to be set off by a tab indentation without skipping a line. Do not auto-format the first line 
(i.e. by using the “first line” command in WORD). All auto-formatting can be fatal when transferring a 
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Use 12 point Times New Roman font. 
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10. Figures can be line drawings, photographs or computer-generated graphics. No figures will be accepted 


INSTRUCTIONS FOR AUTHORS 


larger than 15.5 X 23 cm. Width of lines and sizes of letters in figures must be large enough to allow 
reduction to half page size. 


If a scale is required it must be presented as a bar within the figure. It is the editor’s prerogative to reduce or 
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While there is no objection to full page size figures, it is journal policy to have the legend on the same page 
whenever possible and figures should not be so large as to exclude the legend. Figure legends should be 
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11. Tables must be provided on separate pages at the end of the manuscript and NOT placed within the text. 
It is essential that table legends are not set within the table but are supplied separately as with figure legends. 
It may be necessary to reduce or enlarge tables but the legends must remain in the same font as the text (10 
point Times New Roman in the final form). 

While the draft manuscript text is expected to be in 12 point type, it may be necessary to use smaller font size 
for large tables. Do not use vertical lines in tables unless absolutely necessary to demark data columns. Keep 
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WORD or EXCEL tables are acceptable. 


12. Details of punctuation, scientific nomenclature, etc. are to be found in the complete instructions. 


336 Proc. Linn. Soc. N.S.W., 125, 2004 


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Part 2 — monotreme papers 


PUNT 


CAT 


235 


243 


259 


273 


277 


279 


287 


301 


319 


Grant, T.R. 
Captures, capture mortality, age and sex ratios of platypuses, Ornithorhynchus anatinus, during studies 
over 30 years in the upper Shoalhaven River in New South Wales. 
Grant, T.R., Griffiths, M. and Temple-Smith, P.D. 
Breeding in a free-ranging population of platypuses, Ornithorhynchus anatinus, in the upper Shoalhaven 
River, New South Wales - a 27 year study. 
Grant, T.R. 
Depth and substrate selection by platypuses, Ornithorhynchus anatinus, in the lower Hastings River, New 
South Wales. 
Lunney, D., Grant, T. and Matthews, A. 
Distribution of the platypus in the Bellinger catchment from community knowledge and field survey and its 
relationship to river disturbance. 
Grant, T.R., Lowry, M.B., Pease, B., Walford, T.R. and Graham, K. 
Reducing the by-catch of platypuses (Ornithorhynchus anatinus) in commercial and recreational fishing 
gear in New South Wales. 
Bethge, P., Munks, S., Otley, H. and Nicol, S. 
Platypus burrow temperatures at a subalpine Tasmanian lake. 
Higgins, D.P. 
Ultrasonography of the reproductive tract of the short-beaked echidna (Tachyglossus aculeatus). 
Higgins, D.P., Tobias, G. and Stone, G.M. 
Excretion profiles of some reproductive steroids in the eeees of captive female short-beaked echidna 
(Tachyglossus aculeatus) and long-beaked echidna (Zaglossus sp.). 
Hassiotis, M., Paxinos, G. and Ashwell, K.W.S. 
Anatomy of the central nervous system of the Australian echidna. 
Rowe, M.J., Mahns, D.A. and Sahai, V. 
Monotreme tactile mechanisms: from sensory nerves to cerebral cortex. 
Proske, U. and Gregory, J.E. 
The role of push rods in platypus and echidna — some speculations. 


Part 3 — obituary, book reviews, instructions to authors 


327 
329 


333 


335 


Obituary: Mervyn Edward Griffiths 1914-2003. 

Adam, P. 
Admiral Doenitz’s legacy. A review of the book “Seeds of concern. The genetic manipulation of plants” by 
D.R. Murray (2003). 

Augee, M.L. A review of the book “Citizen Labillardiere: a naturalist’s life in revolution and exploration (1755- 
1834)” by Edward Duyker. 

Instructions for authors. 


PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W. 


VOLUME 125 "wil 


TITUTION LIBRARIES 


UL 


3 9088 01205 6412 


Issued 20 February 2004 
CONTENTS 


Part 1 — general contributions 


1 
"\ 43 


57 
67 
73 


97 


110 


145 
141 


165 


Moulds, T. 

Review of Australian cave guano ecosystems with a checklist of guano invertebrates. 

Young, G.C. 

A Devonian brachythoracid arthrodire skull (placoderm fish) from the Broken River area, Queensland. 

Cohn, J. 

Effects of slashing and burning on Thesium australe R. Brown (Santalaceae) in coastal grasslands of 
NSW. 

Mound, L. and Masumoto, M. 

Trichromothrips veversae sp.n. (Insecta, Thysanoptera), and the botanical significance of insects host 
specific to Austral bracken fern (Pteridium esculentum). 

Timms, B.V., Shepard, W.D. and Hill, R.E. 

Cyst shell morphology of the fairy shrimps (Crustacea: Anostraca) of Australia. 

Lindley, |.D. 

The Yule Island Fauna and the Origin of Tropical Northern Australian Echinoid (Echinodermata) 
Faunas. 

Errata from Lindley, !.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua 
New Guinea: Clypeasteroida. Proceedings of the Linnean Society of New South Wales 124, 125-136, 
and Lindley, 1.D. (2003). Echinoids of the Kairuku Formation (Lower Pliocene), Yule Island, Papua New 
Guinea: Spatangoida. Proceedings of the Linnean Society of New South Wales 124, 153-162. 

Lindley, |.D. 

Some living and fossil echinoderms from the Bismarck Archipelago, Papua New Guinea, and two new 
echinoid species. 

Zhen, Y.Y., Percival, |.G. and Webby, B.D. 

Conodont faunas from the Mid to Late Ordovician boundary interval of the Wahringa Limestone 
Member (Fairbridge Volcanics), central New South Wales. 
Wright, A.J. and Strusz, D.L. , 
Wenlock (Early Silurian) brachiopods from the Orange District of New South Wales. 
Rickards, R.B. and Wright, A.J. 
Early Silurian graptolites from Cadia, New South Wales. 
Edgecombe, G.D. and Wright, A.J. 
Silicified Early Devonian trilobites from Brogans Creek, New South Wales. 

Edgecombe, G.D. 

A new species of the henicopid centipede Dichelobius (Chilopoda: Lithobiomorpha) from southeastern 
Australia and Lord Howe Island. 

Weaver, H.J. and Aberton, J.G. 

Asurvey of ectoparasite species on small mammals during autumn and winter at Anglesea, Victoria. 

Allen, S., Marsh, H. and-Hodgson, A. 

Occurrence and conservation of the dugong (Sirenia: Dugongidae) in New South Wales. 


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VOLUME 126 
March 2005 


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The Middle Triassic Megafossil Flora of the Basin Creek 
Formation, Nymboida Coal Measures, New South Wales, 


Australia. Part 4. Umkomasiaceae. Dicroidium and Affiliated 


Fructifications 


W.B.KeitH Hotmes! AnD H.M.ANDERSON? 


'46 Kurrajong Street, Dorrigo, NSW 2453, Australia (Hon. Research Fellow, Geology Department, 
University of New England, Armidale, NSW 2351, Australia); 
*Hon. Palaeobotanist, National Botanical Institute, Pretoria 0001, South Africa 


Holmes, W.B.K. and Anderson, H.M. (2005). The Middle Triassic megafossil flora of the Basin 
Creek Formation, Nymboida Coal Measures, New South Wales, Australia. Part 4. Umkomasiaceae. 
Dicroidium and affiliated fructifications. Proceedings of the Linnean Society of New South Wales 126, 
1-37. 


The forked leaves of the morpho-genus Dicroidium are the most commonly occurring foliage in the 
collections from two quarries in the Middle Triassic Basin Creek Formation at Nymboida, N.S.W. The 
extensive Nymboida collections include leaves which, in gross morphology, range from simple with 
entire margins to pinnatifid, pinnate, bipinnatifid and bipinnate forms. The wide range of variation, which 
includes intergrading forms, creates problems in establishing species boundaries. For comparison with other 
Gondwanan material, the Nymboida leaves have been placed in five “species complexes’ distinguished 
as “Dicroidium coriaceum’, “D. odontopteroides’, “D. lineatum’, “D. dubium’ and ‘D. zuberi’. In each 
complex there is a continuum of variation of form and there are intergrading forms that link each complex. 
Illustrations of over eighty leaves demonstrate the range of variation present. A single leaf only of D. 
elongatum has been collected. An unusual leaf is described as ?D. nymboidense sp. nov. Fertile material 
affiliated with Dicroidium includes three species of female strobilus, Umkomasia distans, U. sessilis and 
U. sp. A. together with dispersed cupules and ovules. Microsporophylls are placed in Pteruchus sp. cf. P. 


matatamajor and a single specimen in P. sp. A. 


Manuscript received 17 February 2004, accepted for publication 18 August 2004. 


Keywords: Dicroidium, Middle Triassic flora, Nymboida Coal Measures, Pteruchus, Umkomasia. 


INTRODUCTION 


This is the fourth in a series of papers describing 
the rich and diverse Middle Triassic megafossil floras 
from two quarries located near the village of Nymboida 
in north-eastern New South Wales. A locality map 
showing the Coal Mine and Reserve Quarries and a 
summary of the geology of the Nymboida Sub-basin 
was included in Part 1 (Holmes 2000), which also 
dealt with the Thallophyta and Sphenophyta. Part 2 
(Holmes 2001) included descriptions of 14 taxa of 
the Filicophyta known from material preserved in a 
fertile state or remains of sterile material with known 
fern relationships. Twenty four morpho-taxa of fern- 
like leaves of unknown relationships were described 
in Part 3 (Holmes 2003). 

In this paper abundant material of the leaf 


morpho-genus Dicroidium and its affiliated male and 
female fructifications, Pteruchus and Umkomasia 
respectively, from the two Nymboida localities is 
described and illustrated. 


A history of Dicroidium 

Dicroidium appeared first in the Middle Early 
Triassic (Smithian), becoming more numerous and 
diverse in the Middle and Late Triassic (Ladinian to 
Carnian) of the Gondwana super-continent (Retallack 
1977, Anderson and Anderson 1983). There are no 
reliable Jurassic records (Anderson and Anderson 
2003, p.243) and the genus is presumed to have 
become extinct at the Triassic-Jurassic boundary. The 
success and diversification of the genus is recorded 
in numerous publications since the first description 
and illustration of three leaf fragments from Tasmania 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


by Morris (1845, as Pecopteris odontopteroides). The 
genus Dicroidium was erected by Gothan (1912), 
who recognised that the forked Gondwanan leaves, 
usually referred to Thinnfeldia, were distinct from 
the unforked leaves of the Northern Hemisphere. 
Walkom (1917) retained the name Thinnfeldia and 
later erected the genus Johnstonia (Walkom 1925b) 
for simple forked fronds from Tasmania. 

In an overview of genera attributed to the 
‘Thinnfeldia’ series, in which he accepted Dicroidium, 
Frenguelli (1943) also erected the new genera 
Dicroidiopsis, Diplasiophyllum, Xylopteris and 
Zuberia for Gondwana forked leaves. Townrow (1957) 
and Archangelsky (1968) synonymised Frenguelli’s 
new genera with Dicroidium. Retallack (1977), in a 
biostratigraphical review of Dicroidium and allied 
genera, retained Johnstonia and Xylopteris. Anderson 
and Anderson (1983), in their extensive study of 
Dicroidium in the Molteno Flora (South Africa), 
provided a detailed historical review, references and 
comprehensive lists to 1982 of all the illustrated 
material from Gondwana that they considered to fall 
within the genus. They also discussed synonyms and 
forking fronds distinct from Dicroidium. 

Gothan (1912) studied the cuticle of Dicroidium 
and showed that it was distinct from Thinnfeldia. Jacob 
and Jacob (1950) and Townrow (1957) added further 
evidence for the separation. Anderson and Anderson 
(1983) carried out a comprehensive cuticular 
study (light and scanning electron microscopy) 
on Dicroidium in the Molteno Formation. Based 
on permineralised material from Antarctica, Pigg 
(1990) and Boucher et al. (1993) have described the 
anatomy of Dicroidium leaves. The permineralised 
fructifications, Pteruchus and Umkomasia, have 
been described by Yao et al. (1995) and Slavins et al. 
(2002) respectively. 

While the forked leaves of Dicroidium are 
ubiquitous in Gondwana Triassic floras (Retallack 
1977; Anderson and Anderson 1983) their origin 
is unresolved (Boucher et al. 1993; Axsmith et al. 
2000). Archangelsky (1996) suggested that the 
Carboniferous pteridosperm Botrychiopsis may 
provide a plausible ancestral morphology for the 
Umkomasiaceae. Retallack (1980) noted that the 
multi-forked Lepidopteris callipteroides, which 
occurs immediately above the Late Permian coal 
seams in the Sydney Basin, probably belonged to 
the pteridosperm stock that gave rise to Dicroidium, 
most likely through intermediate forms similar to 
‘Dicroidium’ gopadense from Nidpur in India (Bose 
and Srivastava 1971). Palaeozoic pteridosperms from 
Siberian and Cathaysian sources that survived the 
end-Permian extinction in refugia as yet unidentified 


should also be considered. Since Thomas (1933) first 
suggested Umkomasia as a possible early angiosperm 
this has been much debated and remains unresolved 
(see Slavins et al. 2002 for a recent discussion). 
Thomas (1933) placed Dicroidium 
and its affiliated fertile organs in the Family 
Corystospermaceae. This family name has been 
widely used but according to ICBN rules a family 
must be based on the ovulate genus, in this case 
Umkomasia and therefore Umkomasiaceae. Meyen 
(1987) placed the Family Umkomasiaceae and the 
Class Umkomasiales in the gymnosperm Order 
Ginkgoopsida. Anderson and Anderson (2003), in 
their recent review of Umkomasia and Pteruchus 
from the Molteno Flora, followed this classification. 


The question of Dicroidium attachment 

While the leaves of Dicroidium are abundant 
and widespread throughout Triassic Gondwana, 
the form of a whole plant is still debated. Various 
authors have suggested that Dicroidium ranged from 
shrubs to tall trees (Retallack 1980; Anderson et al. 
1998; Anderson and Anderson 2003) or was a large 
tree (Taylor 1996). Petriella (1978) reconstructed 
Dicroidium as a palmiform tree to 10m high. The 
reconstruction by Retallack and Dilcher (1988, fig.10) 
showed a tall deciduous forest tree in a seasonally wet 
lowland. 

Despite the large numbers of Dicroidium leaves 
preserved in the Nymboida sediments, no leaves 
have as yet been found attached to a stem. The only 
convincing specimen elsewhere of leaves attached 
to a stem was illustrated by Anderson and Anderson 
(1983, Pl. 88, fig. 1). Another incomplete specimen 
(Anderson and Anderson 1983, P1.88, fig. 2) suggested 
that the leaves were attached in a fascicled manner 
to a stem. Anderson and Anderson (2003 p. 257, text 
fig.5) base their reconstruction on these specimens. 

In a paper describing Umkomasia uniramia, 
Axsmith et al. (2000, figs 6 and 8) illustrated 
a Dicroidium odontopteroides leaf apparently 
contiguous with, or overlain by, a stem of a plant 
with long and short shoot morphology. By analogy 
with extant plants bearing long and short shoot 
morphology (e.g. Gingko biloba) it would be 
unlikely in the extreme for a plant with this growth 
morphology to bear a leaf on the long shoot section of 
a stem subsequent to the formation of well-developed 
short shoots. On the illustrated Antarctic specimen 
even the short shoots are in a leafless state. The 
close association of the Dicroidium leaf and stems 
bearing short shoots suggests an affiliation but this 
is not exclusive as other leaves (e.g. Heidiphyllum, 
Taeniopteris) are present in the same deposit. We do 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


not accept the claim of Axsmith et al. (2000) that their 
specimen “demonstrates unequivocal evidence” of 
Dicroidium attachment. 

Archangelsky (1968) argued a case for the 
affiliation of silicified logs of Rhexoxylon and 
Dicroidium leaves. Meyer-Berthaud et al. (1993) 
described permineralised twigs from Antarctica 
as Kykloxylon and suggested that the stems bore 
Dicroidium fremouwensis Pigg (1990) leaves, but in 
neither of these cases have the leaves been found in 
organic connection. 


Dicroidium and its fertile organs 

Thomas (1933) described the female organ 
Umkomasia and the male Pteruchus. Based on close 
association and similarity of cuticles he regarded them 
as the fructifications of Dicroidium. This affiliation 
is now generally accepted (Townrow 1962; Holmes 
1982; Anderson and Anderson 1983, 2003; Crane 
1985; Retallack and Dilcher 1988; Yao et al. 1995 and 
Slavins et al. 2002). 

The fertile genera Karibacarpon (Lacey 1976, 
Holmes and Ash 1979) and Fanerotheca (Anderson 
and Anderson 2003) have also been linked to 
Dicroidium. The species described by Axsmith et al. 
(2000) as Umkomasia uniramia from Antarctica is an 
ovulate organ with single pedicillate cupules arranged 
in a terminal radial whorl on a peduncle attached to a 
leafless short shoot on a mature stem bearing further 
leafless short shoots. From the significant differences 
in the architectural arrangement of this fructification 
we believe it should be placed in a morpho-genus 
separate from Umkomasia. 


Problems with Dicroidium taxonomy 

In many early records (Walkom 1917, 
Frenguelli 1943) the forked leaves now known as 
Dicroidium (Gothan 1912) were placed in several 
genera and particularly in Thinnfeldia. Walkom 
(1917) considered his “Thinnfeldia” leaves to be ferns 
as they were closely associated with frond fragments 
bearing sori. Townrow (1957) separated these fertile 
fragments from Dicroidium as they were indeed ferns 
which Holmes (2001) has placed in the new genus 
Herbstopteris. Several genera of forked fronds have 
been described as separate from Dicroidium based 
mainly on morphological differences of pinna or 
pinnule shape (eg Johnstonia, Xylopteris, Zuberia 
and others). Townrow (1957), Archangelsky (1968), 
Holmes (1982) and Anderson and Anderson (1983, 
2003) have regarded most of these additional genera 
as synonyms of Dicroidium. Leaves illustrated by 
Anderson and Anderson (1983, P1. 74, figs 1-9) which 
showed typical D. elongatum (= Xylopteris elongata) 


Proc. Linn. Soc. N.S.W., 126, 2005 


pinnae and typical D. odontopteroides pinnae on 
the same leaf indicate how closely related these 
‘genera’ are. It is far more likely that those leaves 
are interspecific rather than intergeneric ‘hybrids’ or 
even one polymorphic species. Some South American 
workers continue to use a multiplicity of generic 
names for Dicroidium leaves (Artabe 1990; Artabe et 
al. 1998; Gnaedinger and Herbst 1998, 2001). 

Early authors appear to have worked only on 
limited museum collections or had very little field 
experience because they failed to recognise the 
variability within their ‘species’ and the intergrading 
forms that may have linked the ‘species’. Holmes 
(1982) discussed the problems created by the 
variation and intergrading forms of Dicroidium in 
the Middle Triassic Benolong Flora. He referred to 
the work of Meyen (1979), who had demonstrated 
from a large population of Permian pteridosperm 
leaves previously placed in several genera and many 
species that they all belonged in the single species 
Rhaphidopteris praecursoria. Rees and Cleal (1993) 
showed that the leaves of a Jurassic pteridosperm 
previously placed in six species and four genera all 
belonged to Archangelskya furcata. 

The revision by Retallack (1977) of the 
Dicroidium genus (plus Xylopteris, Johnstonia and 
Tetratilon, which he retained as allied but distinct) 
was a significant attempt to provide a taxonomic 
guide to the genus that would mainly be useful for 
stratigraphic purposes. However, his descriptions and 
stylised sketches of the taxa he recognised failed to 
demonstrate the range of variation within a single 
population of each taxon. The necessity under the 
IBCN Code (ICBN 2001) for typification has also 
exacerbated the problem of dealing with variable 
fossil populations. 

The photographic record by Anderson and 
Anderson (1983) of 1133 individual Dicroidium leaves 
arranged in ‘palaeodemes’ was a notable achievement 
and clearly demonstrated the diversity within single 
populations of Dicroidium leaves. They were the 
first to use palaeo-gamma taxonomy for Dicroidium, 
i.e. using the data of comprehensive collections 
from many localities. They defined a ‘palaeodeme’ 
as a “collection of specimens representing a single 
breeding population showing a normal distribution of 
variation and derived from a single fossil assemblage 
from a discrete lithological unit”. The same authors 
did not attempt to define precise boundaries between 
taxa as they considered “it would be unproductive 
considering the general morphological fluidity within 
the genus” and they had “little doubt that Dicroidium 
speciation was anything but simple and came closer 
to the reticulate speciation model of Sylvester- 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Bradley (1977)”. Based on the palaeodeme approach, 
Anderson and Anderson (1983, 1989) listed a total 
of 32 taxa comprising 10 species, 17 subspecies and 
15 formae. In their recent revision (Anderson and 
Anderson 2003) Dicroidium now comprises 21 taxa 
of specific rank plus 15 formae. 

The reasons listed by Boucher et al. (1993) 
provided a good summary of why Dicroidium is not 
a well-understood genus: there is a large amount of 
morphological variation; there is inconsistency in 
naming; variations in cuticular features are not well 
enough known; complete plants are not known; 
sample sizes are small especially from remote areas; 
and often only fragments are preserved. 


MATERIALS 


In the Holmes’ catalogued collection of over 
2600 slabs of Nymboida material, approximately one 
third of the slabs bear leaves that can be attributed 
to Dicroidium. The female and male fertile organs 
Umkomasia and Pteruchus affiliated with Dicroidium 
are rare and have been found at a ratio of one 
identifiable specimen to c. 70 leaves. This is closely 
similar to the ratio observed at the Benolong locality 
(Holmes 1982). In the Molteno assemblages Anderson 
and Anderson (2003, p. 240, 250) also noted that the 
fertile organs were rare and gave the return for each 
locality only in terms of man-hours spent chipping 
open fossiliferous slabs. 

The Nymboida fossil plant material is preserved 
mostly as carbonaceous compressions, but a tectonic 
heating event during the Cretaceous (Russell 1994) 
has destroyed the cuticle of otherwise beautifully- 
preserved specimens. 

In addition to the material held in the Holmes 
collection we have examined other Nymboida 
specimens housed in the University of New England 
Geology Department and the Australian Museum. 


OUR METHOD OF NOMENCLATURE 


Earlier attempts to classify Nymboida 
Dicroidium \eaves were made from limited and 
usually fragmentary specimens, i.e. using mainly 
palaeo-alpha taxonomy (De Jersey 1958, Flint and 
Gould 1975, Retallack et al. 1977). Retallack (1977 
Microfiche Frames G11-G13) listed 12 Dicroidium 
taxa plus one each of Johnstonia and Tetraptilon 
(both here included in Dicroidium) and two ovulate 
species from the Coal Mine Quarry. 

In this present revision we have used mainly 


palaeo-beta taxonomy and based our classification on 
our comprehensive collection of over 2600 selected 
and catalogued slabs that have been accumulated 
during many trips to the Nymboida quarries over a 
period of almost 40 years. The bulk of the collection 
was made from fallen blocks that had been blasted or 
bulldozed from the quarry faces. The exact source of 
the in situ material is mostly unknown. Except in rare 
cases such as the slabs containing large numbers of 
similar leaves (e.g. Figs 2 and 6A), we are unable to 
identify specific populations or palaeodemes as was 
achieved by Anderson and Anderson (1983, 1989, 
2003). Future work involving in situ collecting from 
specific assemblages (i.e. from discrete lithological 
units) would allow palaeo-gamma taxonomy and the 
material to be documented into palaeodemes. 

Dicroidium foliage is the most commonly 
preserved fossil at Nymboida andalso in the catalogued 
collection. The forked leaves occur in a diversity of 
forms ranging and intergrading from simple leaves 
with entire margins to pinnate to bipinnate. Faced 
with this continuum one could place all the leaves 
in one polymorphic species or, on the other hand, 
describe numerous species each based on a ‘single’ 
type specimen. Due to this range of variation and 
intergradation we are unable to determine reliable 
diagnostic features. This makes it difficult to identify 
the Nymboida Dicroidium material to a specific 
level. 

Although the sediments in the Nymboida 
quarries were deposited over a relatively short period 
of geological time (Holmes 2000) they do represent 
many different sedimentary facies, e.g. coal seams, 
shales, fossil soils, siltstones and sandstones. The 
enclosed fossils certainly would have been derived 
from a range of habitats so it is most likely that various 
species of Dicroidium plants had evolved or adapted 
to occupy particular environmental niches. Only one 
variable species of Preruchus plus a possible second 
species are recognised in the Nymboida collections 
while at least three separate species of the female 
Dicroidium fructification Umkomasia are present. It 
is therefore most likely that several discrete species 
of plants bearing Dicroidium leaves were growing in 
the region in the Middle Triassic. 

For the Nymboida Dicroidium leaves to be 
compared with collections from other geographical 
and stratigraphical localities we have separated 
the leaves into informal complexes based on leaf 
morphology, which range from simple to pinnate, 
bipinnatifid and bipinnate. Except for the single 
specimen placed in D. elongatum (Figs. 19, 20A) and 
the few rare and problematic specimens (Figs 20B-E) 
we have grouped the foliage material into five ‘species 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


complexes’ each named for a previously-described 
species that represents the core of the complex. 
Specimens in the Selected References lists also have 
a gross morphology identifiable with the core of the 
complex. Each complex includes a particular section 
of the range of diversity of the Nymboida Dicroidium 
leaf collections and is linked by intergrading forms 
with adjacent complexes. We illustrate over 80 
reasonably well-preserved leaves (Figs 1-18), that 
demonstrate the variation within each complex and 
the obvious forms intergrading between successive 
complexes. While we acknowledge that our form of 
classification is subjective, it is based on a large and 
representative collection of reasonably well-preserved 
material. 


Due to constraints imposed by the lack of . 


storage facilities at the Australian Museum only 
the specimens illustrated in this paper have been 
allocated AMF numbers. All other specimens remain 
in the Holmes Collection. 


SYSTEMATIC PALAEOBOTANY 


Order Ginkgoopsida Meyen 1987 
Class Umkomasiales Meyen 1984 
Family Umkomasiaceae Meyen 1984 


Genus Dicroidium Gothan 1912 


Type species 
Dicroidium odontopteroides (Morris 1845) 
Gothan 1912 


Synonymy [Synonymised by]: 

Johnstonia Walkom 1925b [Townrow 1957] 

Dicroidiopsis Frenguelli 1943 [Archangelsky 1968] 

Diplasiophyllum Frenguelli 1943 [Archangelsky 
1968] 

Zuberia Frenguelli 1943 [Townrow 1957] 

Xylopteris Frenguelli 1943 [Archangelsky 1968] 

Tetraptilon Frenguelli 1950 [Anderson and Anderson 
1983] 

Hoegia Townrow 1957 [Archangelsky 1968] 


‘Dicroidium coriaceum complex’ 
Figures 1A-C; 2A-E 


Selected References 

1925b Johnstonia coriacea, Walkom, figs 6,7 
1927 Johnstonia coriacea, Du Toit, fig. 12D, 13B 
1932 Johnstonia coriacea, Du Toit, text fig. 2A 
1967 Dicroidium coriacium, Jain and Delevoryas, 


Proc. Linn. Soc. N.S.W., 126, 2005 


Pl. 91, fig. 1 

1982 Dicroidium coriaceum, Holmes, figs 3C, D 

1983 Dicroidium coriaceum subsp. coriaceum, 
Anderson and Anderson, PI. 36, figs 3-6; P1.76, 
figs 1-6 

1983 Dicroidium coriaceum subsp. dutoitii, 
Anderson and Anderson, Pl. 41, figs 1-28; P1.76, 
figs 12-17 


Description 

Forked leaves of variable length and width, 
usually entire, sometimes lobed and grading to 
pinnatifid. 


Discussion 

D. coriaceum was first described from Tasmania 
by Johnston (1887) as Rhacophyllum coriaceum. 
Walkom (1925b) re-examined Johnston’s material 
and transferred it to the new genus Johnstonia. He 
noted that the margins were entire or slightly lobed. 
Specimens attained a length up to 100 mm above the 
dichotomy and the breadth of the larger specimens 
was 10 mm, though in general they were narrower. 
Antarctic material with leaves having a broad lamina 
and the fork closer to the apex were described by 
Townrow (1967) as D. dutoitii. He selected as the 
holotype a leaf illustrated by DuToit (1927, text fig. 
12D) from the Molteno Formation of South Africa, 
which differed quite significantly from the Antarctic 
specimens. Retallack (1977) placed the Antarctic 
material in Johnstonia coriacea var obesa with the 
South African material retaining the epithet dutoitii. 
Anderson and Anderson (1983) have separated D. 
dutoitii from D. coriaceum on palaeodeme evidence 
at localities in the Molteno Formation although their 
illustrations show some overlap in leaf dimensions. 
From South America, Jain and Delevoryas (1967, PI. 
91, fig. 1) illustrated a single slab that encompasses the 
size range of both above species. Holmes (1982) also 
noted a wide variation in size range of D. coriaceum 
in the Benolong flora of NSW. The Nymboida leaves 
illustrated in Figure | are a larger form, ranging 
from 120-140 mm long and 5-10 mm wide and from 
entire to lobed, and could be regarded as a form of D. 
dutoitii Townrow (1967). As only three slabs of this 
larger form are present in the Nymboida collections 
and their full range of variation is not known we place 
them in the D. coriaceum complex. 

The specimens illustrated in Figures 2A-E and 
Figure 4C were all collected from a single slab of 
whitish siltstone. We believe that, with the exception 
only of the pinnate fragment on the lower left of Fig. 
2A, all the other leaves represent a single population. 
They range in length from c. 40-80 mm long and 3- 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


7 mm wide. The leaf margins are mostly entire but 
lobed and pinnatifid forms (Fig. 2E and Fig. 4C) are 
present. The latter forms intergrade with, and are very 
close to, the pinnatifid and pinnule coalescing forms 
of the ‘D. odontopteroides complex’. 


‘Dicroidium odontopteroides complex’ 
Figures 3A-E; 4A-G; SA-D; 6A; 7A-C; 8A; 9A-C; 
10A,D,E 


Selected References 

1845 Pecopteris odontopteroides, Morris, Pl. 6, fig. 
3 

1890 Thinnfeldia odontopteroides, Feistmantel, Pl. 
26, figs 2, 2a 

1917 Thinnfeldia lancifolia, Walkom, P1. 7, fig. 2; 
Pl. 3, fig. 1 

1975 Dicroidium odontopteroides, Flint and Gould, 
Pl. 3, figs 10, 11 

1982 Dicroidium odontopteroides var. moltenense, 
Holmes, figs 4A, B 

1983 Dicroidium odontopteroides subsp. 
orbiculoides, Umk 111 palaeodeme, Anderson 
and Anderson, Pl. 42 

1983 Dicroidium odontopteroides forma 
odontopteroides, Umk111 palaeodeme, Anderson 
and Anderson, Pl. 43 

1992 Dicroidium odontopteroides, Taylor et al., fig. 
1 

2000 Dicroidium odontopteroides, Axsmith et al., 
fig. 3, leaf only 

2001 Dicroidium odontopteroides, Gnaedinger and 
Herbst, fig. 4E (in gross morphology but with 
denser venation) 


Description 

Usually once-forked pinnate frond to 160 mm 
long; pinnae to 40 mm long and 15 mm wide, not 
basally contracted, ranging from semi-orbicular to 
broadly triangular, to elongated rectangular or slightly 
tapering, with rounded or broadly obtuse apex; 
the longer pinnae with a midrib and alethopteroid 
venation; shorter rounded pinnae with odontopteroid 
venation as in Fig. 7C. The specimens illustrated in 
Figs 7A and B represent the core of this Nymboida 
complex. The leaf in Fig. 8A is the largest specimen 
from this complex in the collection. 


Discussion 

Taxonomic confusion has arisen since the 
original description and illustrations of Dicroidium 
odontopteroides (as Pecopteris odontopteroides) 
by Morris (1845). Only three frond fragments were 


illustrated and Morris noted that the specimen (PI. 
6, fig. 4) with more elongate pinnae was probably 
a variety of the species. When compared with 
the range of variation encompassed in our D. 
odontopteroides complex, Morris’s three specimens 
show a very limited range of variation as would be 
expected for this morpho-species. However, due to 
the lack of understanding of the range of variation 
that commonly occurs in a single population, leaves 
similar to Morris’s elongated pinna form were raised 
to specific rank i.e. Dicroidium lancifolium (Gothan 
1912; Walkom 1917; Frenguelli 1943; and others, see 
Hypodigm Lists of Anderson and Anderson 1983). 
The presence of a midrib (alethopteroid venation) 
in the more elongated pinnae has been regarded 
by some authors as a specific or varietal diagnostic 
feature. The form of venation is usually dependent 
on the length and shape of the pinnae, which varies 
according to the position on the frond and whether on 
the inside or outside of the fork. Both odontopteroid 
and alethopteroid forms of venation may be observed 
on a single frond. Leaves placed in D. lancifolium 
by several authors are best regarded as long-pinnaed 
forms of D. odontopteroides. 

The range of variation of fronds within this 
Nymboida complex encompasses forms that accord 
with several published species, subspecies, varieties 
or forms. On the classification of Retallack (1977, 
Microfiche Frames I1 to I7) these varieties include D. 
odontopteroides var. moltenense, D. odontopteroides 
var. obtusifolium and D. odontopteroides vat. 
odontopteroides. 

Specimens with short broad pinnae (Figs 3A- 
D; 4C, D, G; 7C) agree with leaves from the D. 
odontopteroides subsp. orbiculoides palaeodeme 
of Anderson and Anderson (1983, Pl. 42), which 
included D. crassinervis forma  obtusifolium, 
D. odontopteroides subsp. orbiculoides and D. 
crassinervis forma crassinervis. Boucher et al. (1993) 
separated compression and impression material from 
Mt Falla in the Beardmore Glacier area of Antarctica 
into D. odontopteroides, D. lancifolium and D. dubium 
based on minor morphological and cuticular features, 
which may have represented a normal range of 
variation within a single species complex. Frenguelli 
(1950) erected the genus Tetraptilon for Dicroidium 
odontopteroides-like leaves from Argentina in which 
the frond had a double fork. This feature has been 
recorded also in D. odontopteroides assemblages 
from Australia (Flint and Gould 1975, PI. 3, fig. 10; 
Retallack 1977) and South Africa (Anderson and 
Anderson 1983, Pl. 87, figs 1,2,4 and 6, Pl. 88, fig. 
1). Anderson and Anderson (1983) synonymised the 
genus Tetraptilon with Dicroidium. In our collections 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


these multiple forked fronds comprise c. 1% of the 
D. odontopteroides complex. The forking may result 
in three, four or five branches, with four branches 
being the most common (Figs 5A-D and Fig. 6A). 
On some bedding planes this is the only form of leaf 
preserved. This suggests that for some trees double- 
forked leaves were the normal frond and not aberrant 
as was surmised by Anderson and Anderson (1983, 
Pls 71, 87) because of their rare occurrence. The 
slab illustrated in Figure 6A is a good example of the 
range of variation in a single population of double- 
forked leaves. A single specimen of a double-forked 
D. dubium (Fig. 6B) has also been collected. 

In a recent paper, Gnaedinger and Herbst 
(2001) recognised the great morphological variability 
in Dicroidium leaves from three Upper Triassic 
formations innorthern Chile. They illustrated anumber 
of examples that indicated an intergrading ‘line’ from 
D. obtusifolium to D. odontopteroides (varieties 
moltenense and remotum) to D. odontopteroides var. 
odontopteroides to D. lancifolium var. lancifolium. 
This range of morphology is here included in our ‘D. 
odontopteroides complex’. 

Leaves illustrated in Figures 9A, B are forms 
intergrading with extremes from the D. lineatum 
complex. 


‘Dicroidium lineatum complex’ 
Figures 10B,C; 11A,B,D; 12A-C 


Selected References 

1883 Gleichenia lineata, Tenison-Woods, PI. 3, fig. 
6; Pl. 8, fig. 2 

1898 Thinnfeldia indica var. falcata, Shirley, P|. 7, 
fig. 2 

1917 Thinnfeldia acuta, Walkom, PI. 3, fig. 4 

1977 Dicroidium lancifolium vat. lineatum, 
Retallack, Microfiche Frame H17 

1983 Dicroidium odontopteroides subsp. lineatum, 
Anderson and Anderson, PI. 64, figs 12-29; Pl. 
65, figs 1-3; Pl. 79, figs 4, 6 

1985 Dicroidium lancifolium var. lineatum, Artabe, 
Pl. 3, fig. 5 

1992 Dicroidium lancifolium, Taylor et al., fig. 2 

2001 Dicroidium lancifolium var. lineatum, 
Gnaedinger and Herbst, fig. 2, A-E, fig. 3 L 


Description 
Forked pinnate leaf to 200 mm long; pinnae 
elongated-triangular to 35 mm long, tapering to 


acute apex, broad base, usually decurrent. 


Discussion 


Proc. Linn. Soc. N.S.W., 126, 2005 


This complex is closely allied to the D. 
odontopteroides complex but has been separated on 
the basis of the decurrent pinnae which taper to an 
acute apex. 

Retallack (1977, Microfiche Frames H17, 
H18) placed leaves of this complex as a variety of D. 
lancifolium, which we regard as belonging in the ‘D. 
odontopteroides complex’. Leaves comparable to our 
‘D. lineatum complex’ were described by Anderson 
and Anderson (1983) from the Molteno Formation of 
South Africa as D. odontopteroides subsp. lineatum 
but were later raised to specific rank (Anderson and 
Anderson 2003). 

In the Nymboida collection, forms with undulate 
margins or incipient lobes (Figs 11C; 12A,D) are 
linking forms with the ‘D. dubium complex’ below. 


‘Dicroidium dubium complex’ 
Figures 6B;.13A-C; 14A-E; 15A-D; 16A 


Selected References 

1890 Gleichenia dubia, Feistmantel, Pl. 26, fig. 3 

1908 Thinnfeldia odontopteroides, Seward, fig. 4 

1928 Thinnfeldia talbragarensis, Walkom, PI. 27, 
fig. 1 

1947 Thinnfeldia talbragarensis, Jones and 
deJersey, Pl. 1, fig. 5 

1965 ‘Thinnfeldia’ talbragarensis, Hill et al., Pl. T4, 
fig. 6 

1982 Dicroidium dubium var. dubium, Holmes, fig. 
7D 

1983 Dicroidium dubium subsp. dubium, Anderson 
and Anderson, Pl. 33, figs 21-31 

1983 Dicroidium dubium subsp. tasmaniense, 
Anderson and Anderson, PI. 44, figs 7-16; Pl. 53, 
figs 15-20; Pl. 59, figs 1-14; Pl. 60, figs 1-5 

1983 Dicroidium dubium subsp. switzifolium, 
Anderson and Anderson, Pl. 44, figs 17-20 


Description 

Forked bipinnatifid leaf; pinnae elongate, 
tapering to acute apex; margin variously lobed, 
proximal lobes sometimes separated to the pinna 
rachis but coalescing distally. Leaves variable in size 
from 70 - 200 mm long; pinnae in mid-portion of leaf 
from 15 - 90 mm long, 4 - 15mm wide. 


Discussion 

Fronds of the ‘D. dubium complex’ are the 
most commonly occurring Dicroidium fossils at 
the Nymboida quarries. Leaves that represent the 
core of this complex (Figs 13C, 14D,E, 15B,C) are 
bipinnatifid to partially bipinnate and agree well with 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


the Molteno palaeodemes of D. dubium subsp. dubium 
as recognised by Anderson and Anderson (1983, 
p. 106). Pinnae on the same frond may vary from 
entire to lobed and pinnate. Specimens with narrower 
pinnae and undulate margins are intergrading forms 
with the ‘D. lineatum complex’ while the bipinnate 
forms intergrade with those of the ‘D. zuwberi complex’ 
(Fig. 15A). The apical leaf fragment showing large 
bipinnatifid pinnae (Fig. 16A) may be compared 
with D. dubium var. australe (sensu Retallack 1977, 
Microfiche Frame H8). 


‘Dicroidium zuberi complex’ 
Figures 16B,C; 17A-D; 18A 


Selected References 

1890 Thinnfeldia odontopteroides, Feistmantel, P1. 
24, figs 1,2; Pl. 25, figs 1,2 

1917 Thinnfeldia feistmantelii, Walkom, P1. 2, figs 
12) 

1944 Zuberia zuberi, Frenguelli, Pl. 4 

1975 Hoegia papillata, Flint and Gould, PI. 2, figs 
4,5 

1977 Dicroidium zuberi, Retallack et al., Figs A-F 

1979 Dicroidium zuberi, Petriella, Pl. 2, fig. 5 

1983 Dicroidium zuberi, Anderson and Anderson, 
Pl. 61, figs 1-13; Pl. 62, figs 1-4; Pl. 81, figs 1-5 

1985 Dicroidium zuberi var. papillatum, Artabe, P1. 
4, fig. 2 


Description 

Frond bipinnate, small to large, 150 - 600 mm 
long; pinnules variable in shape, rounded to blunt 
rhomboid or falcate, inclined towards pinna apex, 
coalescing distally and apically. Basiscopic pinnules 
often decurrent on main rachis. 


Discussion 

Leaves of this complex are widespread but 
not common at the Nymboida localities. Solitary 
large leaves are sometimes preserved intact in beds 
of sandstone thus indicating they were tough and 
resistant to damage during transport. D. zuberi is 
differentiated from D. dubium by the presence of 
pinnules divided to the pinna rachis, but there are 
numerous intergrading forms with deeply bipinnatifid 
fronds (Fig. 17A,B). 

Retallack (1977, Microfiche Frame G12) listed 
three varieties of D. zuberi from Nymboida based on 
the pinnule shape of mostly fragmentary material. 
Our extensive collections clearly demonstrate 
intergrading forms that link the varieties. Indeed 
pinnules on the same leaf may include different 


varietal types (Figs 16B, 18A). The range of pinnule 
shapes in the Nymboida ‘D. zuberi complex’ supports 
the decision of Anderson and Anderson (1983) to 
recognise the previously described D. zuberi varieties 


feistmantelii, sahnii, barrealensis and papillatum as 


normal variations within a single species. 

We agree with Archangelsky (1968), Retallack 
(1977) and Anderson and Anderson (1983, 2003) 
that the continued placement of these leaves into the 
separate genus Zuberia (Artabe 1990; Gnaedinger 
and Herbst 2001) is not warranted. 


Dicroidium elongatum (Carruthers) Archangelsky 
1968 
Figures 19, 20A 


Holotype 
Designated by Retallack (1977) Sphenopteris 
elongata Carruthers 1872, Pl. 27, fig. 1 


Selected References 

1883 Trichomanides spinifolium, Tenison-Woods, 
Pl. 3, fig. 7 

1898 Trichomanides elongata vat. spinifolia, 
Shirley, Pl. 5, fig. 2 

1917 Stenopteris elongata, Walkom, PI. 6, figs 1, 3 

1965 Xylopteris spinifolia, Hill et al., Pl. TS, fig. 7 

1977 Xylopteris spinifolia, Retallack, Microfiche 
Frames J13, 14 

1982 Dicroidium spinifolium, Holmes, Fig. 5B,C 

1983 Dicroidium elongatum forma remotifolium, 
Anderson and Anderson, p.115, Pl. 48, figs 24-32 


Description 

Known only from a single incomplete specimen 
from the Coal Mine Quarry (AMF125082). Frond 
bipinnate, c. 120 mm long; pinnae linear at c. 45° 
to main rachis; pinnules well-spaced, elongated 
triangular to linear. 


Discussion 

This solitary specimen is preserved in siltstone 
together with other fragments that appear to have been 
subjected to long distance transport before burial. In 
the probable contemporary Benolong Flora (Holmes 
1982) leaves of a ‘Dicroidium elongatum complex’ 
comprised 20% of the preserved plant remains. 
Holmes reconstructed the Benolong environment as 
a dry sclerophyll woodland on low fertility sandy 
soils. The presence of a rich fern and fern-like leaf 
flora at Nymboida (Holmes 2001, 2003) suggests 
a vegetation growing in a moist environment with 
rich soils. This solitary specimen of D. elongatum 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


together with associated plant fragments was most 
likely transported from a drier less fertile upstream 
source. 

D. elongatum is a species complex of pinnate to 
tripinnate intergrading forms. In the past the various 
forms have been separated into species, subspecies 
and formae. Jones and de Jersey (1947) considered 
the pinnate to tripinnate forms to represent an 
evolutionary series but all forms are now known to 
occur together. Holmes (1982) noted that leaves with 
intergrading characters of most forms were present in 
the Benolong Flora. 


? Dicroidium nymboidense Holmes sp. nov. 
Figure 20B 

Diagnosis 

A small pinnate frond; pinnae few, large 
elongate-ovate, rounded, irregularly lobed; venation 
odontopteroid, two decurrent veins entering each 
pinna then dividing acutely up to five times to form 
parallel venation to the pinna apices. 


Description 

This new morpho-taxon is based on a single 
frond impression and counterpart. Frond pinnate, 
115 mm long with base missing; curving rachis with 
five opposite pairs of pinna and a terminal pinna, 
increasing in size apically; fifth pair of pinnae to 55 
mm long, 25 mm wide, elongate-ovate with two or 
three broad irregular obtuse lobes; texture of pinnae 
thin, venation conspicuous; two veins enter each 
decurrent pinna at c. 30°, arch and divide from three 
to five times then run parallel to each other to pinna 
apex; the upper of the two primary veins forming a 
short median vein. 


Holotype 
AMF125083 Australian Museum, Sydney. 


Type Locality 

Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle 
Triassic. 


Discussion 

This rare and problematical frond is placed 
with reservations in Dicroidium. The rachis curves 
sideways basally but the leaf base is missing and 
no fork is preserved. However, on the right margin 
of the specimen there is a pinnule fragment with 
venation that divides and runs parallel and is aligned 
at an angle appropriate for a pinna if it were a portion 
of a forked Dicroidium frond. The venation, while 


Proc. Linn. Soc. N.S.W., 126, 2005 


essentially odontopteroid, is somewhat similar to that 
in the decurrent, ovate, sometimes lobate pinnules 
of Nymboidiantum spp (Holmes 2003, Figs17- 
19). However the size and texture of the pinnae of 
?Dicroidium nymboidense is unlike that of any 
Nymboidiantum spp. While we are uncertain of its 
generic placement it is certainly a distinct species and 
is described and illustrated to draw attention to its 
presence in the Nymboida Flora. 


Dicroidium sp. A 
Figures 20C,D 

Description 

Apical portions of two fronds with extremely 
widely spaced pinnae; Fig. 20C is pinnate, pinnatifid 
with slightly lobed or entire strongly decurrent pinnae; 
Fig. 20D is bipinnate with broadly triangular pinnules 
and basiscopic pinnules decurrent on or attached 
directly to the main rachis. 


Discussion 

The wide spacing of the pinnae of these 
incomplete fronds sets them apart from all other more 
completely preserved fronds. They may possibly 
represent aberrant forms of D. dubium and D. zuberi 
respectively. D. pinnis-distantibus (Kurtz) Frenguelli 
(Retallack 1977, Microfiche Frame M 8) also has 
very widely spaced pinnae but with entire pinnae 
margins. 


Dicroidium sp. B 
Figure 20E 


Description 

The apical portion of a very large bipinnatifid 
Dicroidium-like frond with deeply dentate pinna 
margins. 


Discussion 

This single portion of an obviously very large 
frond differs by its size and shape from all other 
bipinnatifid material from Nymboida. D. sp. B differs 
from the large D. dubium var. australe (Retallack 
1977, Microfiche Frame H8) and D. dubium ssp. 
helvetifolium (Anderson and Anderson 1983, p. 105) 
by the acute triangular pinna lobes. A distal fragment 
of a somewhat similar-sized but pinnate frond from 
the Late Triassic Ipswich Basin was _ attributed 
by Walkom (1917) to Danaeopsis hughesi, now 
Dicroidium hughesi Lele (1962), a species described 
from India. 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Fertile organs associated with Dicroidium leaves 
Genus Umkomasia Thomas 1933 


Type species 
Umkomasia macleani Thomas 1933 p.203, figs 
1, 2, 56; Pl. 23, figs 1-4 


Umkomasia distans Holmes 1987 
Figures 21A; 22A 


Holotype 
AMF 63824, Holmes 1987, fig. 3, fig. 4, A-D 


Paratypes 
AMF 63825-28 


Description 

An Umkomasia strobilus with widely spaced 
alternate and spirally arranged branches on an 
elongate axis; branches with one or two opposite 
pairs of cupules and a terminal pair or a single cupule; 
cupules pedicillate, irregularly rounded, 3.5 to 5 mm 
in diameter. 


Discussion 

U. distans was described and _ illustrated 
by Holmes (1987) based on three specimens of 
reasonably preserved cupule bearing axes from the 
Coal Mine Quarry, Nymboida. Additional material 
has been collected (Fig. 21A) but this is fragmentary. 
For comparisons with other material see Holmes 
(1987). This species was regarded as distinct from the 
Molteno species by Anderson and Anderson (2003, 
Table 51). 


Umkomasia sessilis Holmes 1987 
Figures 21B,C; 22B 


Holotype 
AMF63829 and counterpart AMF63830, 
Holmes 1987, fig. 5, fig. 6 A-C 


Paratype 
AMF63831 


Description 

An elongate Umkomasia strobilus with 
alternate, spirally arranged branches bearing two 
pairs of opposite, sessile cupules 4 - 5 mm in 
diameter. 


Discussion 

U. sessilis differs from U. distans by the sessile 
cupules. U. decussata Anderson and Anderson (2003) 
has sessile cupules but differs by the more numerous 
pairs of cupules with a decussate arrangement. The 
partial strobilus located on the lower left hand side of 
Figure 10A probably belongs here. 


Umkomasia sp. A 
Figure 22C 


Description 

A portion of a single elongate Umkomasia 
strobilus with well-separated alternate branches, each 
branch with one or more (?) pairs of cupules borne on 
long expanding pedicels; cupules strongly decurved, 
8-10 mm long. 


Discussion 

U. sp. A differs from U. distans and U. sessilis 
by the larger decurved cupules and by the elongated 
and expanding pedicels. The cupules of U. gracilliaxis 
Anderson and Anderson (2003) from the Molteno 
Formation are decurved and of a size similar to U. sp. 
A. However, their cupules with four lobes each is a 
feature not preserved on this Nymboida specimen. 


Isolated Umkomasia cupules 
Figures 22D-F 


Specimen AMF125092 (Fig. 22D) shows a 
pair of adjacent and opposite dorsally compressed 
semicircular cupules each c. 16 mm in diameter, 
with a 4 mm wide peduncle attached to the proximal 
margin. These cupules are closely comparable with 
the cupule shown in a dorsal view of Karibacarpon 
(Umkomasia) feistmantelii (Holmes and Ash 1979, 
Fig. 6.2) from the Early Triassic Lorne Basin 
assemblage and with cupules from the Sydney Basin 
(Walkom 1925a, Pl. 31, fig. 9; Retallack 1980, fig. 
21.9E). No additional specimens have been found at 
Nymboida to show whether these cupules may have 
split on dehiscence into several lobes as occurs in U. 
feistmantelii fructifications (Holmes and Ash 1979, 
fig. 6. 3-5; Walkom 1925a, Pl. 29, fig. 9; 1932, Pl. 5, 
figs 3-5; Retallack 1980, Fig 21.9F). 

Specimen AMF125093 (Fig. 22E) is a thick 
rounded tapering mass, 15 mm wide, of carbonaceous 
material attached to a stout peduncle 2 mm wide. 
On opposite sides of the mass are decurved acute 
projections. This problematic specimen may represent 
a pair of large fleshy conjoined cupules, perhaps 
similar to Umkomasia sp. of Retallack (1980, Fig. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


21:9E): 

Specimen AMF 125094 (Fig. 22F) is an isolated 
dehisced cupule c. 7 mm wide, split into ?4 acute 
lobes. This cupule was thin textured in contrast to 
the thick fleshy appearance of the previous specimen. 
Preservation is too poor to allow for comparisons 
with known fructifications. 


Dispersed Umkomasia ovules 
Figure 21D 


Dispersed platyspermic ovules occur frequently 
on some horizons at both the Coal Mine and Reserve 
Quarries. The ovules illustrated are c. 5 mm wide and 
c. 7mm long, with an acute bifid apex. They are similar 
in form and shape to the ovules illustrated by Thomas 
(1933, fig. 33) and Anderson and Anderson (2003, PI. 
82, figs 11-16) and associated with Dicroidium leaves 
at the Umk111 locality of the Molteno Formation. In 
size they are closest to the ovules associated with U. 
quadripartita from the Mat 111 locality (Anderson 
and Anderson 2003, PI. 85, figs 7-10). 


Genus Pteruchus Thomas 1933 


Type species 
Pteruchus africanus Thomas 1933, fig. 34, Pl. 24, 
fig. 71 


Pteruchus sp. cf P. matatimajor Anderson and 
Anderson 2003 
Figures 23A-E 


Selected references 

1947 Pteruchus cf africana, Jones and de Jersey, text 
fig. 51 

2003 Pteruchus  matatimajor, 
Anderson, p. 254, Pls 92-94 


Anderson and 


Description 

This morpho-taxon is based on the five 
incomplete strobili illustrated and numerous other 
detached heads. The main axis is stout, to 2.5 mm 
wide, length not known; microsporophylls oval to 
linear-oblong, in pairs on a slender forked peduncle; 
heads 11-23 mm long, 5-9 mm wide; microsporangia 
spindle- or cigar-shaped, 2-3 mm long, 0.35-0.5 mm 
wide, covering whole abaxial surface. 


Discussion 


None of the Nymboida specimens is complete. 
Figure 23B clearly shows the paired nature of the 


Proc. Linn. Soc. N.S.W., 126, 2005 


microsporophylls. In this feature and size they are 
closely similar to the Preruchus strobilus illustrated 
by Jones and deJersey (1947, text fig. 51) from 
Ipswich, and also with P. matatimajor (Anderson and 
Anderson 2003) from the Matatiele assemblage in 
the Molteno Formation. We consider the Nymboida 
material as close to P. matatimajor and quite separate 
from P. africanus, which has smaller unpaired 
microsporophylls. The identification of P. dubius and 
P. johnstonii from the Coal Mine Quarry by Retallack 
(1977, Microfiche Frame G12) is doubtful as it was 
based on fragmentary material. The microsporophylls 
placed in Pteruchus johnstonii from the middle 
Triassic Benolong assemblage by Holmes (1982) 
differ from the Nymboida specimens by their shorter, 


~ more rounded unpaired heads. P. feistmantelii (Holmes 


1987 p.172) from the Lorne Basin (Holmes and Ash 
1979) and the Sydney Basin (Walkom 1925a, PI. 31, 
fig. 10; Retallack 1980) was very much longer and 
more gracile. 

It is interesting to note that in both the Molteno 
and Nymboida localities where P. matatimajor or a 
comparable form occurs, the leaves of the D. dubium 
complex are the most common Dicroidium. 


Pteruchus sp. 
Figure 23F 


Description 
A detached microsporophyll, 40 mm long and 
9 mm wide; forking distally. 


Discussion 

This microsporophyll is larger than all other 
Nymboida Prteruchus heads and the forking structure 
is unique. In size it approaches the long gracile 
microsporophyll of P. feistmantelii from the Lorne 
Basin, Australia (Holmes and Ash 1979). 


CONCLUSION 


Leaves of the Dicroidium morpho-genus are 
the most commonly occurring fossils in the large 
collections made from the Nymboida quarries. 
Collections have been made from differing 
sedimentary facies and may represent vegetation 
from various habitats depending on the degree of 
transport or dispersal before burial. The Nymboida 
leaves ascribed to Dicroidium display a wide range of 
variation and intergrading forms ranging from simple 
to pinnate to bipinnate.To identify individual leaves 
with previously described species ignores the presence 
of the intergrading forms. Therefore we have separated 


11 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


the leaves into five ‘species complexes’ each centred 
on a widely distributed and well-recognised morpho- 
species. The single leaf of D. e/ongatum present in the 
Nymboida flora is possibly a transported specimen 
from a source far beyond the Nymboida flood-plain 
vegetation which comprises the bulk of the preserved 
material. Future descriptions of all fossil floras should 
recognise the variability and range of intergrading 
forms due to ecological factors, the chance factors 
involved in the fossilisation process and discovery of 
the fossilised material. 


ACKNOWLEDGEMENTS 


W.B. Keith Holmes acknowledges with appreciation 
the enthusiastic help by his late wife Felicity and his 
daughters in collecting specimens from the Nymboida 
quarries over a period of almost 40 years. The operator of 
the quarries during that period, the late Mr Brian Foley, 
always provided valuable assistance during our visits by 
exposing a never-ending supply of material for examination. 


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354. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


elie 


eal 


i 


i. 


Figure 1. A-C. Dicroidium coriaceum complex. Larger leaves with mainly entire margins. A. 
AMF125016; B. AMF125017; C. AMF125018. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


15 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 2. A-E. Dicroidium coriaceum complex (excluding the leaf on lower left of Fig. 2A). Smaller leaves 
with margins ranging from entire to pinnatifid. All specimens from the same slab of siltstone and represent 

a single population. A. AMF125019; B. AMF125020; C. AMF125021; D. AMF125022; E. AMF125023, 
pinnatifid leaf on left hand side intergrading with shorter pinnuled forms of the D. odontopteroides complex. 
Scale bar = 1 cm. 


16 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


4S 


gaat 
Epa gots at 
wo iS 


Figure 3. A-E. Dicroidium odontopteroides complex. Pinnate leaves with short pinnules. A. AMF125024; B. 
AMF125025; C. AMF125026; D. AMF125027; E. AMF125028. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 . 7 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 4 A-G. Dicroidium odontopteroides complex. Pinnate leaves with short, variously rounded and 


inclined pinnae. A. AMF125029; B. AMF125030; C. AMF125031; D. AMF125032; E. AMF125033; F. 
AMEF125034; G. AMF125035. Scale bar = 1 cm. 


18 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


A 


3, 4 or 5 branches 


into 
=lcm 


ing 


Scale bar 


innate leaves fork 


IP 
D. AMF125039. 


dium odontopteroides complex 


1crol 
B. AMF125037 


D 


D 


AMF 125036 


Figure 5 A- 


AMF 125038; 


C 


> 


p! 


19 


Proc. Linn. Soc. N.S.W., 126, 2005 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 6. A. Dicroidium odontopteroides complex — an assemblage of double-forked leaves demonstrating 
size and shape variation. AMF 125040. B. Dicroidium dubium complex — a double forked leaf. AMF125041. 
Scale bar = 1 cm. 


20 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 7. A-C. Dicroidium odontopteroides complex. Larger leaves with variously elongated pinnae. A. 
AMF 125042; B. AMF125043. C. AMF125044, leaf showing ‘odontopteroid’ venation. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


21 


TRIASSIC FLORA FROM NY MBOIDA - UMKOMASIACEAE 


Figure 8. A. Dicroidium odontopteroides complex. The largest leaf in the complex. AMF 125045. Scale bar 
= | cm: 


22 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 9. A-C. Dicroidium odontopteroides complex. A, B. Forms approaching Dicroidium lineatum com- 
plex. A. AMF125046; B. AMF125047; C. AMF125048. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


23 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 10. A,D,E. Dicroidium odontopteroides complex, approaching Dicroidium lineatum complex. A. Note 
portion of Umkomasia sessilis strobilus on lower right. AMF125049; D. AMF125050; E. AMF125051. B,C. 
Dicroidium lineatum complex. B. AMF125052; C. AMF125053. Scale bar = 1 cm. 


24 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 11. A,B,D. Dicroidium lineatum complex. A. AMF 125054; B. AMF125055; D. AMF125056. C. Inter- 
grating form between Dicroidium lineatum complex and Dicroidium dubium complex. AMF 125057. Scale 
bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 25 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


st 


. 
4 


Figure 12. A,D. Forms intergrading between Dicroidium lineatum complex and Dicroidium dubium complex. 
A. AMF125058; D. AMF125059. B,C. Dicroidium lineatum complex. B. AMF125060; C. AMF125061. 
Scale bar = 1 cm. 


26 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 13. A-C. Dicroidium dubium complex showing range of pinna forms. A. AMF125062; B. 
AMF125063; C. AMF125064. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


Pal 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 14. A-E. Dicroidium dubium complex. A. AMF 125065; B. AMF125066; C. AMF125067; D. 
AMF 125068; E. AMF125069. Scale bar = 1 cm. 


28 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 15. A-D. Dicroidium dubium complex. A and D approaching Dicroidium zuberi complex. A. 
AMF 125070; B. AMF125071; C. AMF125072; D. AMF125073. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


29 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 16. A. Dicroidium dubium complex- large bipinnatifid form, AMF 125074. 
B, C. Dicroidium zuberi complex. B. AMF 125075; C. AMF125076. Scale bar = 1 cm. 


30 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 17. A-D. Dicroidium zuberi complex. A. AMF 125077; B. AMF 125078; C. AMF125079; D. 
AMF125080. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 31 


TRIASSIC FLORA FROM NYMBOIDA - UMKOMASIACEAE 


Figure 18. A. Dicroidium zuberi complex. AMF125081. Scale bar = 1 cm. 


32 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 19. Dicroidium elongatum AMF 125082. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


TRIASSIC FLORA FROM NY MBOIDA - UMKOMASIACEAE 


Figure 20. A. Dicroidium elongatum AMF 125082. B. ?Dicroidium nymboidense sp. nov. AMF 125083. C, D. 
Dicroidium sp. A. C. AMF125084; D. AMF125085. E. Dicroidium sp. B. AMF125086. Scale bar = | cm. 


34 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 21. A. Umkomasia distans. Holotype. AMF63824. B, C. Umkomasia sessilis. B. Holotype AMF63831. 
C. AMF 125087. D. Dispersed Umkomasia ovules showing bifid micropyle. AMF125088. Scale bar = | cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 35 


TRIASSIC FLORA FROM NY MBOIDA - UMKOMASIACEAE 


Figure 22. A-C. Umkomasia sp. strobili and cupules: A. Umkomasia distans. AMF 125089. B. Umkoma- 
sia sessilis. AMF125090. C. Umkomasia sp. AMF125091. D-F. Umkomasia cupules: D. AMF125092; E. 
AMF 125093; F. AMF125094. Scale bar = | cm. 


36 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 23. A-E. Pteruchus sp.cf. P. matatimajor microsporophylls. A. AMF125095; B. AMF125096; C. 
AMF125097; D. AMF125098; E. AMF125099; F. Preruchus sp. A. AMF125100. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


37 


f 


MC MORRIA At ROR RBM SOR REESE AE 


v 


The Middle Triassic Megafossil Flora of the Basin Creek 
Formation, Nymboida Coal Measures, New South Wales, 
Australia. Part 5. The Genera Lepidopteris, Kurtziana, 
Rochipteris and Walkomiopteris. 


W.B.KeITH HoLMes! AND HeErpI M. ANDERSON? 


'46 Kurrajong Street, Dorrigo NSW 2453, Australia (Hon. Research Fellow, University of New England, 
Armidale NSW 2351); 746 Kurrajong Street, Dorrigo NSW 2453, Australia (Hon. Palaeobotanist, National 


Botanical Institute, Pretoria 0001 South Africa). 


Holmes, W.B.K. and Anderson, H.M. (2005). The Middle Triassic megafossil flora of the Basin 
Creek Formation, Nymboida Coal Measures, New South Wales, Australia. Part 5. The genera 
Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris. Proceedings of the Linnean Society 
of New South Wales 126, 39-79. 


Leaves attributed to the gymnosperm genera Lepidopteris, Kurtziana and Rochipteris and the 
leaf sedis incertae Walkomiopteris eskensis (Walkom) Holmes and Anderson gen. et comb. nov. 
are described from two quarries in the Basin Creek Formation of the Middle Triassic Nymboida 
Coal Measures. Based on extensive collections of leaves, the morpho-genera Lepidopteris and 
Kurtziana each reveal a wide range of variation. Lepidopteris is divided into three “‘morpho- 
species complexes’ with intergrading forms based on L. madagascariensis, L. africana and 
L. stormbergensis and a new species L. dissitipinnula apparently without links to the three 
complexes. The Lepidopteris fertile organs, Peltaspermum and Antevsia are present. Kurtziana 
is separated into two ‘morpho-species complexes’ based on K. brandmayri and K. cacheutensis. 
Rochipteris is a diverse genus but of very rare occurrence. Six new species are described; R. 
obtriangulata and R. tubata display a close-spaced spiral or whorled arrangement. Seven leaves 
of R. incisa have been examined. R. sinuosa, R. pusilla and R. nymboidensis are represented by 
single dispersed leaves. Walkomiopteris eskensis (Walkom) gen. et comb. nov. is redescribed 


from additional Nymboida material. 


KEYWORDS: Kurtziana, Lepidopteris, Middle Triassic Flora, Nymboida Coal Measures, 


Rochipteris, Walkomiopteris. 


INTRODUCTION 


In this fifth part of a series describing the early 
middle Triassic Nymboida flora, leaves assigned to 
three gymnosperm genera: Lepidodopteris with some 
associated fertile organs, Kurtziana, Rochipteris and 
the new genus Walkomiopteris sedis incertae are 
illustrated and described. 

Part 1 (Holmes 2000) of this series dealt with the 
Bryophyta and Sphenophyta, Part 2 (Holmes 2001) 
with the Filicophyta, Part 3 (Holmes 2003) with 
fern-like foliage and Part 4 (Holmes and Anderson 
in press) with the genus Dicroidium and its fertile 
organs Umkomasia and Pteruchus. 

The material described below is based mainly 
on the collections made by the senior author and his 


family from two Nymboida quarries over a period of 
almost forty years. Details of the Coal Mine Quarry 
and the Reserve Quarry together with a summary 
of the geology of the Basin Creek Formation, the 
Nymboida Coal Measures and the Nymboida Sub- 
basin were provided in Holmes (2000). 


METHODS 


In living populations of extant plants there is often 
a large range of variation in leaf shape, e.g. juvenile, 
adult, shade, sun-leaves, etc. A preserved holotype in 
a herbarium seldom exhibits this range of variation. 
It is only through wide experience in the field that 
this variation can be recognised and appreciated. In 
the fossil record, leaves constitute the vast majority 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


of preserved plant organs. Most early palaeobotanical 
taxonomy was based on very limited material in the 
field or a few museum specimens. Original diagnoses 
rarely acknowledged the variation that could occur 
in a ‘natural’ species. To compound the problem 
of separating fossil leaves into ‘natural’ species, 
the assemblages from one locality may represent 
vegetation from a range of habitats and growing 
through an unknown period of time. As in Part 4 of this 
series, which dealt with the highly variable morpho- 
genus Dicroidium (Holmes and Anderson in press), 
we address the problem of variability observed in the 
large collections of Lepidopteris and Kurtziana leaves 
by creating ‘species complexes’. A ‘species complex’ 
includes leaves displaying a range of variation centred 
on a previously-described species. Intergrading forms 
often link the ‘complexes’. The Selected References 
are of specimens we consider to represent a mid- 
range for each ‘species complex’. Leaves illustrated 
in the Figures should enable comparisons to be made 
with material from other locations and horizons. 
Leaves in the genera Rochipteris and Walkomiopteris 
are represented respectively by ten and six specimens 
only, so are placed in morpho-species based on gross 
morphology of the material available. 

At the Nymboida quarries most specimens are 
preserved as carbonaceous compressions in which 
the gross morphology is usually well-preserved. 
However, spores and cuticles have been destroyed by 
a tectonic heating event during the Cretaceous Period 
(Russel 1994). 

Type and illustrated material is housed in the 
Australian Museum, Sydney. Some additional 
specimens are in the collections of the Geology 
Department, University of New England, Armidale, 
NSW, and the type of Walkomiopteris eskensis is 
housed in the collections of the Queensland Museum, 
Brisbane. 


SYSTEMATIC PALAEOBOTANY 


Ginkgoopsida S.V.Meyen 1987 
Peltaspermales F. Nemejc 1968 
Peltaspermaceae H.H. Thomas ex T.M. Harris 1937 
Genus Lepidopteris Schimper 1869 
Type species Lepidopteris stutgardiensis Schimper 
1869 


The proposal by Poort and Kerp (1990) to unite 
the leaves of Lepidopteris ‘natalensis’ with the ovulate 
organ Pel/taspermum thomasii, which occur together 
at the Waterfall locality in the Molteno Formation 
(locality Umk111 of Anderson and Anderson 1983), 


40 


in the ‘natural genus’ Meyenopteris is untenable. The 
leaf species Lepidopteris stormbergensis has priority 
over L. natalensis. An additional leaf and ovulate 
species have been described from the same Umk111 
locality (Anderson and Anderson 2003). Until fruit, 
leaves and stems are found in organic connection, it 
is premature to erect a ‘natural genus’. In accordance 
with ICBN (2001) Articles 1.2 the morpho-genus 
Lepidopteris should be retained for dispersed 
leaves and the morpho-genera Peltaspermum and 
Antevsia for the dispersed female and male organs 
respectively. 

Leaves of the Lepidopteris genus occur in the 
Permian of the Northern Hemisphere. The first 
Gondwanan record is of L. callipteroides, a branched 
leaf form from the basal Narrabeen Group (earliest 
Triassic), of the Sydney Basin (Retallack 2002). 
This species apparently did not persist through to the 
Middle Triassic.For many Gondwanan Lepidopteris 
leaves the application of specific names has at times 
been questionable. Some species are known only from 
impressions while some, with better preservation have 
been described with cuticle information (Carpentier 
1935; Townrow 1960, 1966; Baldoni 1972; Baldoni 
and de Cabrera 1977; Anderson and Anderson 1989). 
However, as noted by Townrow (1966), there are 
problems of identification of specimens both with or 
without cuticle. Rigby (1977) suggested reserving the 
name L. stormbergensis for all leaves lacking cuticle 
and L. natalensis and L. madagascariensis for those 
with preserved cuticle, while Retallack in Retallack et 
al. (1977) argued that the diverse range of leaves from 
the Cloughers Creek Formation in the Nymboida Coal 
Measures was best placed in L. madagascariensis on 
the basis of the thick leaf substance and mostly obtuse 
pinnules although no cuticle was present. We regard 
the Cloughers Creek leaves as being best placed in 
our ‘L. stormbergensis’ and ‘L. africana’ complexes. 

In the Nymboida collections, Lepidopteris leaves 
are preserved on c. 3% of the catalogued slabs. 
While individual leaves may be identified on gross 
morphology with the types of L. madagascariensis, 
L. africana or L. stormbergensis, there are numerous 
intergrading forms that link these three ‘species’. The 
same problem was noted by Holmes (1982) for the 
Benolong Flora where leaves of L. stormbergensis, 
L. africana and L. mortonii formed an intergrading 
series. Anderson and Anderson (1989) noted that 
at 11 of their 30 localities where both L. africana 
and L. stormbergensis were present, in most cases 
they formed an unbroken morphological range and 
were regarded as one palaeodeme. At the remaining 
19 localities only one or other of the species was 
present. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


As our collections of Lepidopteris have been 
made from the various sedimentary facies in the 
two Nymboida quarries they were probably sourced 
from differing vegetation types (Retallack 1977; 
Holmes 2000) and the Lepidopteris material may 
indeed belong to several true species each specific 
to one palaeodeme. However, on our present state 
of knowledge we must accept the collection as 
representing a single variable population sample. To 
enable the Basin Creek material to be compared with 
that from other localities, we place the Lepidopteris 
leaves in three ‘species complexes’ while noting 
the intergrading forms that link the complexes. A 
distinct leaf-type with widely-spaced pinnules and 
no intergrading links with the three ‘complexes’ is 


erected as a new morpho-species.A notable feature . 


of Lepidopteris leaves, with and without preserved 
cuticle, is the usual presence of ‘blisters’ or ‘lumps’ on 
the main and/or pinna rachis (Townrow 1956, 1960, 
1966; Holmes 1982; Anderson and Anderson 1989, 
2003) resulting from a proliferation of epidermal cells 
around trichome bases. Townrow (1960) noted that 
‘in a population of leaves attributed to Lepidopteris 
stormbergensis there was a series of leaves with 
rachises ranging from smooth to markedly blistered. 
Lumps or blisters are not apparent on the Nymboida 
Lepidopteris leaves but some do have punctate or 
striate rachises. Townrow (1966) described the main 
rachis of Lepidopteris as having a wing and with 
dorsally-attached pinnae. A rachis wing 1s not evident 
in the Nymboida material and the pinnae appear to be 
attached laterally. 


‘Lepidopteris madagascariensis complex’, based on 
L. madagascariensis Carpentier 1935 
Figures 1A,B; 2A—C 

Selected references 

1935 Lepidopteris madagascariensis, Carpentier, 
P1.3, figs 3,4 

1936 Lepidopteris madagascariensis, Carpentier, 
PL.5, fig.4 

1966 Lepidopteris madagascariensis, Townrow, 
Text fig. LE, P1.1, fig.1 

1975 Lepidopteris madagascariensis, Flint and 
Gould, P1.2, figs 1, 2 

1979 Lepidopteris madagascariensis, Holmes and 
Ash, Fig.5.6, 5.7 

1983 Lepidopteris madagascariensis, Retallack, 
Fig.5A 

1995 Lepidopteris madagascariensis, Retallack, 
Figs 2A, 3A 


Proc. Linn. Soc. N.S.W., 126, 2005 


Description 

Small broad-elliptic bipinnate leaves c. 50-150 
mm long, c. 40-70 mm wide, leaf base truncate, 
main rachis 2 mm wide and tapering to apex, 
sometimes punctate and/or longitudinally: striate; 
c. 20 pairs of well-separated opposite to alternate 
straight or arching pinnae, decreasing in length 
basally and apically, are attached at a high angle 
towards base, at c. 60° in mid-frond and more acute 
apically; pinnules not conjoined, oblong, truncate to 
obtusely rounded, attached by whole base to pinna 
rachis at c. 60°; first basiscopic pinnule decurrent; 
with one or more pinnules attached laterally on 
rachis between pinnae. These latter are generally 
referred to as ‘zwischerfiedern’. 


Material 
AMF126801—3, AMF126805 Coal Mine 
Quarry; AMF126804 Reserve Quarry 


Discussion. 

Typical leaves of this complex are not as 
numerous as those in the ‘L. africana complex’. It 
is distinguished from the two complexes below by 
the oblong pinnules with obtuse apices separated to 
the base on the mid-frond pinnae and by the higher 
angle of attachment of these pinnules. However, 
basal and apical pinnae often have coalescing to 
coherent pinnules. Figure 2A shows, on the same 
slab, portions of several leaves that obviously 
represent a single population (palaeodeme). One leaf 
is typically ‘madagascariensis’ while others show 
pinnules becoming coherent and grading into the 
form of the ‘L. africana complex’. 


‘Lepidopteris africana complex’, based on L. 
africana (Du Toit 1927) Holmes 1982 
Figures 2D; 3A,B; 4A,B; 5A,B 


Selected references. 

1927 Callipteridium africana, Du Toit, P1.27 

1944 Callipteridium argentinum, Frenguelli, P1.1, 
figs 1,2 

1965 Lepidopteris stormbergensis, Hill et al., P1.T6, 
fig. 1 

1977 Lepidopteris madagascariensis, Retallack et 
al., fig.9D 

1982 Lepidopteris africana, Holmes, Figs 8C, 8D 

1983 Lepidopteris africana, Anderson and 
Anderson, P1.13, fig.1 

1989 Lepidopteris africana, Anderson and 
Anderson, p.92, figs 1-3, P1.13, figs 1-10, 
P1.43, figs 1-16 


41 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


1998 Lepidopteris madagascariensis, Gnaedinger 
and Herbst, figs 14a—c 

2001 Lepidopteris madagascariensis, Gnaedinger 
and Herbst, fig. lla 

2003 Lepidopteris africana, Anderson and 
Anderson, p.157, fig.1 


Description 

Small to medium-sized bipinnatifid leaves 120— 
>170 mm long, 25—70 mm wide, with a truncate 
leaf-base 4 mm wide, tapering gradually to apex; 
pinnae closely spaced, longest at 2/3 of leaf length 
where attached at c. 45° to main rachis, apically the 
pinnae decrease in length and become more acute, 
basally the pinnae have a higher angle of attachment, 
become shorter, with rounded apices and entire to 
undulate margins; pinnae with coherent pinnules 
forming a serrate margin; the basiscopic base of the 
pinnae strongly decurrent along the main rachis to 
the acroscopic base of the pinna below, leaving no 
space for zwischerfiedern. 


Material 
AMF 126806 Reserve Quarry; AMF126807—12 
Coal Mine Quarry 


Discussion 

The leaves illustrated in Figures 3A,B and 
4A were exposed on one bedding plane and surely 
represent a single population (palaeodeme). Figure 
4B shows another bedding plane assemblage showing 
many ‘L. africana’ leaves of varying size together 
with a fragment of a leaf with larger separated 
pinnules that approaches ‘L. stormbergensis’ but 
with pinnules coalescing distally and apically. Large 
leaves with pinnules becoming less coherent form 
intergrading links between ‘L. africana complex’ and 
‘L. stormbergensis complex’ (Figs 5C; 6A,B; 7A). 


‘Lepidopteris stormbergensis complex’, based on L. 
stormbergensis (Seward 1903) Townrow 1956 
Figures 6C; 8A,B; 9A,B 


Selected references. 

1903 Callipteridium stormbergense, Seward, P1.7, 
fig.5 

1927 Lepidopteris stuttgardensis, Du Toit, P1.28 

1956 Lepidopteris stormbergensis, Townrow, figs 
1A, 1B 

1960 Lepidopteris stormbergensis, Townrow, text 
figs SC,F,G 

1965 Lepidopteris stormbergensis, Hill et al., P1.T6, 
fig.2 


42 


1975 Lepidopteris stormbergensis, Flint and Gould, 
P1.2, figs 1,2 

1977 Lepidopteris madagascariensis, Retallack et 
al. fig. 9A 

1982 Lepidopteris stormbergensis, Holmes, fig.8A 

1983 Lepidopteris stormbergensis, Anderson and 
Anderson, P1.13, figs 2,3 

1989 Lepidopteris stormbergensis, Anderson and 
Anderson, p.93, figs 1-4, P1.26, figs 2—S, Pl. 27, 
figs 1-4 

1998 Lepidopteris madagascariensis, Gnaedinger 
and Herbst, P1.3, fig.h, figs 14a,c 

2003 Lepidopteris stormbergensis, Anderson and 
Anderson, p.157, fig.4 


Description. 

Large bipinnate to tripinnatifid leaves, broad- 
oblanceolate, to 400 mm long and 180 mm wide. 
Rachis to 5 mm in mid-frond; pinnae opposite to 
alternate, longer pinnae at mid-frond attached at c. 
80°-45°, closely spaced to overlapping on larger 
and tripinnatifid fronds; pinnules on mid-portion of 
mid-pinnae 6—25 mm long, 3—6 mm wide, tapering 
to acute or narrow obtuse apex, margin entire to 
serrate. On the largest leaves (Figures 8B, 9B) 
the pinnules are deeply lobed to pinnatisect. First 
basiscopic pinnule attached to base of pinna rachis 
or strongly decurrent on main rachis; nil to three 
zwischerfiedern between pinnae on main rachis in 
mid-portion of leaf. 


Material 
AMF126816—21, AMF126851, all Coal Mine 


Quarry. 


Discussion 

The leaf assemblage preserved on AMF 126819, 
AMF 126821 and AMF126851 are parts and 
counterparts from the same bedding plane and show 
fronds ranging from bipinnatifid to tripinnatifid and 
include the largest in the collection (Figure 8B). 
Zwischerfiedern preserved on the tripinnatifid leaf 
(Figure 9B) are broad-elongate with a lobed margin. 
This assemblage demonstrates the large range of 
variation even within a single population. 


Lepidopteris dissitipinnula Holmes and Anderson 
sp. nov. 
Figures 1|OA—C 


Diagnosis 

A medium-sized Lepidopteris leaf with sub- 
opposite slightly arching to recurved pinnae; 
pinnules well-spaced, elongated-linear with obtuse 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


apices, margins parallel, entire to lobate. 


Description 

Based on two specimens, both with base and 
apices missing, length preserved to 110 mm; rachis 
at base of preserved section 2-3 mm wide; pinnae 
elongate-lanceolate, alternate, c. 12 mm apart, longest 
pinnae to 65 mm. Specimen AMF113528 (Figures 
10A,B) has slightly recurved pinnae attached at c. 
60°; AMF126823 (Figure 10C) has lower pinnae 
attached at right angles and arching slightly and the 
upper pinnae attached at c. 60°, but this may also be an 
artifact of preservation. Pinnules opposite, spaced c. 
one pinnule width apart, decurrent, straight or slightly 
arched, apex rounded-obtuse, margins parallel, entire 


to lobate. Pinnules are longest at mid-pinna, to 12 . 


mm long, 1—2 mm wide, decreasing in length basally 
and apically, basal basiscopic pinnule not decurrent 
on main rachis; one or two pairs of narrow, elongate 
zwischerfiedern on main rachis between pinnae. 


Holotype 
AMF 113528 and counterpart AMF113529 
Australian Museum, Sydney. 


Type Locality 
Coal Mine Quarry. Basin Creek Formation, 
Nymboida Coal Measures, Middle Triassic. 


Other material 
AMF 126823 Coal Mine Quarry. 


Name Derivation 
dissitus, (Lat.) well-spaced; referring to the 
well-separated pinnules. 


Discussion 

Lepidopteris dissitipinnula differs from all 
described Lepidopteris morpho-species by the 
elongated-linear well-spaced pinnules and perhaps 
is closest to L. madagascariensis. However, at 
Nymboida there are no intergrading forms to link 
L. dissitipinnula with the ‘L. madagascariensis 
complex’. Both specimens of L. dissitipinnula are 
preserved in a white sandstone matrix in contrast to 
all other Lepidopteris, material which is preserved in 
black to grey shales and mudstones, thus suggesting 
they were sourced from ecologically separated 
populations. 


Genus Peltaspermum Harris 1937 


Type species. Peltaspermum rotula Harris 
1937 


Proc. Linn. Soc. N.S.W., 126, 2005 


The ovulate organ Peltaspermum had a wide 
Laurasian and Gondwanan distribution. It is generally 
accepted as the female fructification of the plant that 
bore Lepidopteris leaves (Thomas 1933; Harris 1937, 
Townrow 1960, Anderson and Anderson 2003). 
Poort and Kerp (1990) revised the Peltaspermum - 
Lepidopteris association based on western and central 
European material. They proposed the creation of 
the ‘natural genus’ Pe/taspermum by emending the 
diagnosis of Peltaspermum to include Lepidopteris 
leaves. As Peltaspermum and Lepidopteris are both 
morpho-taxa under ICBN (2001) Article 1.2 a new 
name would be required for a ‘natural genus’. 

Despite the large number of Lepidopteris leaves 
in the Nymboida collection, Peltaspermum is known 
only from two incomplete strobili and two detached 
peltate discs. 


Peltaspermum cf monodiscum Anderson and 
Anderson 2003 
Figures 11A—E 


Description 

Based on two incomplete strobili. Axes as 
preserved c. 40 mm and 25 mm long, 2 mm wide; 
six discs c. 4 mm in diameter attached at 8-10 mm 
intervals singly or opposite each other by a peduncle 
c. 5 mm long. Each disc is c. 6 mm wide, pendant, 
showing four decurved, linear lobes 4 mm long, 
1 mm wide. As the fossils represent sideways- 
compressed discs, the number of lobes in life would 
be eight. A single detached disc and its counterpart 
(Figures 11D, E) show a radially symmetrical disc c. 
9 mm in diameter with ten linear lobes around the 
circumference, each c. 1 mm wide and 2—3 mm long. 
A possible peduncle protrudes from one side of the 
disc but its point of attachment is uncertain. 


Material 
AMF126824—6 Coal Mine Quarry. 


Discussion 

Anderson and Anderson (2003 pp. 152, 158, 159) 
described and illustrated from the Molteno Formation 
of South Africa some reasonably intact strobili with 
lobed receptacles attached singly to an axis. They 
refer to the receptacles as ‘discs’. Their specimens 
have 11 or 12 lobes. The Nymboida material with 
8-10 lobes is otherwise closely comparable. 


Peltaspermum sp A 
Figures 11F,G 


43 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Description 

One specimen and its counterpart shows a 
spherical disc 18 mm in diameter with c. 13-14 
broad obtuse lobes around the margin, each separated 
by an incision or ridge reaching from half to two 
thirds distance to the centre, which is marked with 
an irregular-shaped abcission scar c.1.8 mm _ in 
diameter. 


Material 
AMF 126852 and counterpart AMF 126853 Coal 
Mine Quarry. 


Discussion 

Peltaspermum sp. A differs from P cf 
monodiscum by the larger size and less incised lobes. 
Similar detached peltoid discs have a wide distribution 
and are associated in Gondwana deposits with the 
peltaspermaceous genera Lepidopteris (Harris 1937; 
Holmes and Ash 1979; Holmes 1982, Anderson 
and Anderson 2003) and Scytophyllum (Zamuner 
et al. 1999) and in the Northern Hemisphere with 
Lepidopteris, Tatarina, Comia, Pachydermophyllum 
and Scytophyllum (Meyen 1987). This isolated 
Nymboida disc has insufficient diagnostic features to 
place it in any known species. 


Genus Antevsia Harris 1937 
Type species Antevsia zeilleri (Nathorst) Harris 
1937 


Antevsia strobili have been recorded from 
Rhaetic localities in Sweden (Antevs 1914), 
Greenland (Harris 1932) and from the Upper Triassic 
Molteno Formation of South Africa (Anderson and 
Anderson 2003). Antevsia has been linked at those 
occurrences with Lepidopteris on the basis of similar 
cuticles (Antevs 1914; Harris 1932) and the same 
distinctive blistering on the strobilis axes as on the 
foliar rachises (Anderson and Anderson 2003). 


Antevsia sp A 
Figures 12A—C 


Description 

Two fragmentary specimens from Nymboida 
show clusters of sessile sporangia. AMF 126828 
(Figures 12A,B) is a portion of a strobilis overlain by 
a fragment of a Sphenobaiera leaf. The curved axis, 
which may be an almost complete branch, is c. 60 mm 
long, 1.4 mm wide at the base and tapering to 0.8 mm 
distally. Blisters are not apparent. Clusters of up to 
five irregularly elliptic microsporangia to 2 mm long 
are scattered along the branch axis. It is not certain 


44 


whether the sporangial sacs are sessile or shortly 
pedunculate. The second specimen, AMF126829 
(Figure 12C), is of two detached clusters and some 
scattered sporangial sacs to 5 mm long. Associated 
with the sporangia is a detached oval-shaped 
indeterminate ovule. 


Material 
AMF126828 and AMF126829 Coal Mine 


Quarry. 


Discussion 

Antevsia sp. A has some similarities to Antevsia 
mazenodensis Anderson and Anderson (2003) from 
the Molteno Formation but the preservation is not 
sufficient for specific determination. 


Order Matatiellales Anderson and Anderson 2003 
Family Matatiellaceae 
Genus Kurtziana Frenguelli 1942a 
Type species Kurtziana cacheutensis (Kurtz) 
Frenguelli 1942a 


In frond morphology and venation pattern 
Kurtziana differs from all other Gondwanan 
ginkgoopsid leaf genera. Based on mutual occurrence 
Anderson and Anderson (2003) have given a Grade 
2 affiliation of Kurtziana leaves with the female 
strobilis Matatiella and placed the leaf genus in the 
Order Matatiellales. 

The genus Kurtziana was erected by Frenguelli 
(1942a) for unforked pinnate leaves with pinnae 
having contracted pinna bases and attached laterally 
to the rachis. These leaves from Argentina were first 
illustrated by Kurtz (1921, Pl. 16, figs 198-199) as 
Danea cacheutensis. A second species, K. brandmayri 
Frenguelli (1944), was erected for leaves in which the 
pinnae were closely-spaced to imbricate and attached 
to the dorsal surface of the rachis. A very large 
leaf from Chile has recently been described as K. 
paipotensis Herbst and Gnaedinger (2002). Kurtziana 
leaves with preserved cuticle have been described by 
Petriella and Arondo (1982) and Artabe et al. (1991). 
Herbst and Gnaedinger (2002) have erected the new 
morpho-genus Alicurana for Kurtziana leaves with 
preserved cuticle. To date Kurtziana is best known 
from South American localities. 

Kurtziana leaves are also known from South 
Africa (Du Toit 1927; Anderson and Anderson 1983). 
In a recent publication, Anderson and Anderson 
(2003) illustrated five species and noted the presence 
of 16 species, generally of rare occurrence, from the 
Molteno Formation of South Africa. From Australia, 
the leaf ‘Thinnfeldia’ eskensis Walkom (1928) is here 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


transferred to Kurtziana cacheutensis. 

Kurtziana leaves are represented on c. 2% of 
catalogued slabs in the Nymboida collection. They 
are preserved as impressions and due to their probable 
coriaceous nature, only rare examples show clear 
details of the venation. One bedding plane in Coal 
Mine Quarry was covered with complete Kurtziana 
leaves, perhaps indicating an autumnal deposit of a 
deciduous plant. The Nymboida specimens exhibit a 
range of variation in frond and pinna size, in pinna 
shape and manner of attachment to the rachis. Some 
agree closely with the types of K. cacheutensis and 
K. brandmayri, others with K. cacheutensis sensu 
Herbst and Gnaedinger (2002) from collections that 
also exhibited a variation in leaf form. We have 
separated the leaves into two ‘species complexes’ 
based essentially on the perceived dorsal or lateral 
attachment of the pinnae to the rachis, a feature 
sometimes obscured by the manner of preservation. 


‘Kurtziana brandmayri complex’, based on 
Kurtziana brandmayri Frenguelli 1944 
Figures 13A,B 


Selected references. 

1944 Kurtziana brandmayri, Frenguelli, text fig. 2, 
Pl. 4, figs 1,2 

1965 “Thinnfeldia” eskensis, Hill et al., Pl. TS, figs 
3,4 

1991 Kurtziana brandmayri, Artabe et al., Fig. 1 

2002 Kurtziana brandmayri, Herbst and 
Gnaedinger, figs 2A—C; Pl. 4, figs D—F 


Description. 

Kurtziana leaves with elliptic lamina, to c. 
240 mm long, 100 mm wide, rachis to 4 mm wide, 
decreasing in width distally, striate and sometimes 
punctate, with expanded leaf base. Pinnae opposite to 
sub-opposite, closely spaced to overlapping, sessile, 
with contracted auriculate or caudate bases, attached 
to the dorsal surface of the rachis; oblong to linear- 
ovate or tapering to rounded-acute apex, 40—50 
mm long, 8-18 mm wide; basal pinnae broad-oval, 
increasing in length to mid-portion of leaf. Angle of 
attachment of pinnae to rachis, from 80° near base to 
75° in mid-leaf and becoming more acute apically. 


Material 
AMF 126830—1 Coal Mine Quarry. 


Discussion 

The Nymboida specimen on Figure 13A agrees 
closely with the type of K. brandmayri (Frenguelli 
1944, Pl. 4, figs 1,2) by the closely-spaced to 


Proc. Linn. Soc. N.S.W., 126, 2005 


overlapping oblong pinnae with obtuse apices and 
constricted auriculate bases attached at a high angle to 
the rachis. Other specimens with a dorsal attachment 
of the pinnae and contracted bases differ by the pinnae 
being not so closely-spaced, with a more acute angle 
of attachment and by the tapering of the pinnae to a 
narrower rounded-acute apex. 


‘Kurtziana cacheutensis complex’, based on 
Kurtziana cacheutensis (Kurtz) Frenguelli 1942a 


Figures 14A—D;15A; 16A,B; 17A 


Selected references 

1928 ‘Thinnfeldia’ eskensis, Walkom, P1.27, fig. 2, 
P1.28, fig. 1 

1942a Kurtziana cacheutensis (Kurtz) Frenguelli, 
Pl.1 

1975 Dicroidium eskense, Flint and Gould, Pl. 3, 
fig. 3 

1983 Kurtziana cacheutensis, Anderson and 
Anderson, PI. 9, fig. 5 

2002 Kurtziana cacheutensis, Herbst and 
Gnaedinger, Fig.l A-H 


Description 

Elliptic-ovate pinnate leaves, 100-200 mm 
long, 40-100 mm wide, pinnae attached laterally to 
a Striated rachis 2-3 mm wide, opposite to alternate, 
separated by c. one pinna width, linear-oblong, 
tapering slightly to acute rounded apex, to 45 mm 
long, 9 mm wide in midleaf, pinna acroscopic base 
contracting to midvein, basiscopic base contracted 
or variously decurrent. Pinnae inserted laterally on 
rachis at c. 60° becoming more acute apically. 


Material 
AMF126832-9 Coal Mine Quarry. 


Dicussion 

This is the most common form of Kurtziana at 
Nymboida, sometimes forming monotypic autumnal 
deposits on a bedding plane (Figure 15A) or associated 
with leaves of the conifer Rissikia (Figure 16B). This 
species complex is separated from the K. brandmayri 
complex by the lateral attachment of the pinnae to 
the rachis. Some specimens (Figures 15A and 16A) 
are closely comparable with the illustrated type of 
K. cacheutensis (Frenguelli 1942a). However, there 
is a wide range of variation in leaf size, pinna size 
and shape, spacing and inclination of pinnae to rachis 
and the degree of contraction or decurrence of the 
basiscopic base of the pinna. Figure 17A shows two 
leaves with extreme decurrent pinnae in the apical 


45 


PRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


half of the leaf and contacted pinnae in the basal half. 
Some leaves (e.g. Figure 14C) appear to have pinnae 
with asymmetrical laminae but this is probably an 
artifact of preservation caused by the inrolling of one 
edge of the pinna. 


Order Petriellales Taylor et al. 1994 
Family Kannaskoppiaceae Anderson and Anderson 
2003 
Genus Rochipteris Herbst, Troncoso and 
Gnaedinger 2001 
Type species Rochipteris lacerata (Arber) Herbst et 
al. 2001 


Flabellate leaves with anastomosing venation 
have long been known from the Triassic floras of 
Gondwana. Early records were from Queensland by 
Carruthers (1872, as Cyclopteris cuneata) and Shirley 
(1898, as Sagenopteris cuneata), from Tasmania 
by Johnston (1888 as Cyclopteris australis), from 
Victoria by Chapman (1927 as Psygmophyllum 
fergusoni), from New Zealand by Arber (1917 as 
Chiropteris lacerata), from Chile by Solms-Laubach 
(1899 as Chiropteris copiapensis) and South 
Africa by Seward (1903 as Chiropteris cuneata). 
Other records may be found in Etheridge (1895), 
Chapman and Cookson (1926), Frenguelli (1942b, 
1944) and Menendez (1951). The taxonomy of the 
group was confused. Retallack (1980) discussed the 
problems and subsequently recognised six species 
under an emended diagnosis for Ginkgophytopsis. 
Herbst et al. (2001) made a detailed analysis of 
the significant differences between the essentially 
Northern Hemisphere genus Ginkgophytopsis and 
the Gondwanan leaves. For the Gondwanan leaves 
they erected the new genus Rochipteris in which 
was included five species based mainly on material 
from Argentina and Chile. Barone-Nugent et al. 
(2003) redescribed leaves from the Leigh Creek Coal 
Measures of South Australia and the Ipswich Coal 
Measures of Queensland as Rochipteris etheridgei 
and R. ginkgoides respectively. 

In a recent significant publication on Gondwana 
Triassic gymnosperms, Anderson and Anderson 
(2003) described some remarkable material 
from the Molteno Formation of South Africa. 
Included were specimens of flabellate leaves with 
anastomosing venation and with either female or male 
fructifications attached to a stem. The female strobili 
were described as Kannaskoppia, the male strobili 
as Kannaskoppianthus and the attached leaves as 
Kannaskoppifolia. Under ICBN (2001), Article 1.2, 
detached leaves are regarded as morpho-taxa and 


46 


thus Rochipteris Herbst et al. (2001) has priority over 
Kannaskoppifolia. Kannaskoppifolia may be used as 
a genus for leaves attached to a stem. 

Anderson and Anderson (2003) regarded 
Kannaskoppiafolia (= Rochipteris) as “ubiquitous, 
diverse, long-lived, relatively frequent but generally 
lacking in abundance”. Barone-Nugent et al. (2003) 
noted that their Rochipteris species appeared to be 
distinct between separate basins and showed a strong 
degree of intra~-Gondwanic provincialism in marked 
contrast to Dicroidium species, which are widely 
distributed throughout Gondwana (Retallack 1977; 
Anderson and Anderson, 1983; Holmes and Anderson, 
in press). Forty years of collecting at Nymboida has 
yielded the six new species described below, but, with 
the exception of two species, the others are represented 
by a single specimen only. The species have been 
distinguished on leaf morphology, venation pattern 
and vein density. Two of the Nymboida species are 
important in demonstrating, for some species at least, 
that Rochipteris foliage is inserted on the stem either 
as a close spiral or a terminal whorl. 


Rochipteris obtriangulata Holmes and Anderson sp. 
nov. 
Figures 18A—C; 19A—D 


Diagnosis 

Obtriangular lamina, lateral and distal margins 
straight, entire; angle of divergence 20°—30°; 
veins sub-parallel, dichotomising c. five times and 
anastomosing twice in distal half of lamina; leaves 
attached in close spirals or whorls of eight to ten; 
venation density 20-25 per 10 mm. 


Description 

Based ona slab bearing impressions of one almost 
complete whorl and two incomplete whorls plus other 
isolated single leaves. The leaves are obtriangular, c. 
40 mm long and 12 mm wide at the truncate apex. 
Lamina diverging from the acute sessile base at c. 
20°-30°. Lateral margins straight, leaf apex truncate, 
straight and entire. Veins sub-parallel, dichotomising 
close to base and then four or five more times to leaf 
apex; from c. mid-lamina the veins converge and 
conjoin with adjacent veins, usually twice, to form 
irregular linear elliptical areoles; venation density at 
2/3 leaf length 20-25 per 10 mm. Foliage arranged in 
a close spiral or a terminal whorl of 8-10 leaves but 
leaf bases and stem not visible. 


Holotype 
AMF 126840 and counterpart AMF 126842; 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


paratype AMF126841, Australian Museum, Sydney. 


Type Locality 

Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle 
Triassic. 


Name Derivation 
obtriangulata (Lat.), obtriangular; referring to 
the reversed triangular leaf form. 


Discussion 

The straight lateral margins of the expanding 
laminae and the truncate entire apex differentiates R. 
obtriangulata from all other described Rochipteris 


species. They are close to the leaves from the . 


Molteno Formation locality Umk111 illustrated as 
Kannaskoppifolia sp. C by Anderson and Anderson 
(2003). 


Rochipteris tubata Holmes and Anderson sp. nov. 
Figures 20A—C 


Diagnosis 

Vase-shaped lamina, lateral margins concave, 
distal margin convex-rounded, entire; angle of 
divergence at base 15° increasing to 60°—90° 
apically; veins sub-parallel, dichotomising from near 
base, anastomosing in distal 2/3 of lamina. Foliage 
in a close spiral or whorl of c. 7 leaves; vein density 
20-25 per 10 mm. 


Description 

Based on one almost complete whorl of seven 
leaves. Lamina vase-shaped; leaves to 30 mm long, 
19 mm wide, rising from an attenuated base at c. 
15° and expanding distally in a curve to 60°—90°. 
Lateral margins concave, apical margin convex- 
rounded, slightly undulate. Venation sub-parallel, 
dichotomising c. five times from near base and 
anastomosing in distal 1/3 of lamina. Seven leaves 
apparently attached in a close spiral or terminal whorl 
but leaf bases and stem not visible. Density of veins 
c. 20-25 per 10 mm. 


Holotype 
AMF126843 Australian Museum, Sydney. 


Type Locality 

Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle 
Triassic. 


Proc. Linn. Soc. N.S.W., 126, 2005 


Name Derivation 
tubata (Lat.), trumpet; referring to the 
expanding outline of the leaf lamina. 


Discussion 

Rochipteris tubata is arranged in a whorl of 
leaves similar to R. obtriangularis but unfortunately 
in neither species is their attachment to the stem 
visible. The venation in both species is similar but 
R. tubata is separated on the basis of the expanding 
lamina and rounded leaf apex. The whorls of leaves 
in R. obtriangulata and R. tubata suggest a strong 
relationship to the stems bearing Kannaskoppia fruits 
(Anderson and Anderson 2003), which have small 
groups or whorls of leaves attached at intervals along 
the slender stem. 


Rochipteris incisa Holmes and Anderson sp. nov. 
Figures 21A—C; 22A 


Diagnosis 

Medium-sized cuneate leaf with arched apex; one 
to six deep incisions to below mid-lamina forming 
sub-parallel lobes; venation parallel, occasionally 
bifurcating or conjoining to form extremely elongated 
linear areoles. Venation density c. 18 per 10 mm. 


Description 

Based on three leaves from the Reserve Quarry 
and eight from Coal Mine Quarry. Leaf cuneate, to 
115 mm long and c. 70 mm wide; lateral margins 
straight or slightly concave, diverging from base at 
c. 45°-80°; apex semicircular, deeply incised to form 
a number of linear segments from 8-10 mm wide, 
incisions reaching to 1/3 distance from the lamina 
base, distal ends of segments entire or with a minor 
incision. Veins fine and parallel, dichotomising and 
occasionally conjoining to form extremely elongated 
areoles; density of veins c. 18 per 10 mm. 


Holotype 
AMF 126844 Australian Museum, Sydney. 


Type Locality 
Reserve Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures. 


Other Material 
AMF 126827, 126862, Reserve Quarry; 
AMF 126863—66, Coal Mine Quarry. 


Name Derivation 
incisa, (Lat.), incised, referring to the regularly 


47 


PRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


incised distal margin. 


Discussion 

Rochipteris incisa shows some similarities in 
shape and outline to the attached Molteno leaves 
Kannaskoppifolia vincularis (Anderson and Anderson 
2003). It differs from all other Nymboida Rochipteris 
spp in lamina shape and venation details. Rochipteris 
etheridgei (Arber) Barone-Nugent et al. is similar in 
outline to R. incisa but differs by its larger size, the 
less-deeply incised segments, the sinuate venation and 
by the presence of a broadly-flared leaf base (Barone- 
Nugent et al. 2003, fig. 3B, Pl. 1, figs 2-5). 

On the same slab and adjacent to the holotype of R. 
incisais asmaller spathulate leaf with venation similar 
to that of R. incisa (Figure 21B). This spathulate form 
is similar to leaves from Argentina that have been 
placed in Rochipteris cuneata (Carruthers) Herbst 
et al. 2001. That species was based on Cyclopteris 
cuneata Carruthers (1872), a poorly preserved leaf 
fragment with both apex and base missing. We 
believe that our spathulate leaf may belong to the 
same population as R. incisa and perhaps represents a 
juvenile or immature stage of development. 


Rochipteris sinuosa Holmes and Anderson sp. nov. 
Figures 23A—C 


Diagnosis 

A small flabellate leaf, diverging at c. 45° from 
short expanded leaf base; divided into two major 
segments by a deep incision; one segment again 
divided by a shallower incision; veins radiating 
from base, sinuous, approaching and diverging from 
each other, occasionally dichotomising but rarely 
anastomosing. Vein density c. 18 per 10 mm. 


Description 

Based on a single leaf with apical margin missing. 
Leaf as preserved, 42 mm long, 25 mm wide; flabellate, 
expanding at c. 45° from a short flared leaf base 4 
mm wide. Lamina divided into two major segments 
by a medial incision commencing at c. 12 mm from 
the leaf base; the left segment is again divided by a 
narrow incision commencing at 18 mm from the base. 
The lamina lateral margins are slightly concave, the 
apical margin is missing. Veins run in a sub-sinuous 
manner parallel to the lateral margins, dichotomising 
occasionally, approaching and diverging from each 
other but rarely forming a true anastomosis. The 
apparent areoles are elongate-elliptic. Vein density in 
the upper portion of the lamina c. 18 per 10 mm. 
Holotype 


48 


AMF 126845 Australian Museum, Sydney. 


Type Locality 

Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures, Middle 
Triassic. 


Name Derivation 
sinuosa (Lat.) sinuous; referring to the venation. 


Discussion 

The gross morphology, sub-sinuous venation and 
paucity of anastomoses distinguishes this leaf from 
other Nymboida Rochipteris species. Rochipteris 
sinuosa closely resembles the specimens from the 
Llantenes Formation of Argentina attributed to 
Chiropteris copiapensis Steinmann and Solms by 
Menendez (1951, P1.3, figs 1-4). However, those 
specimens have been synonymised with Rochipteris 
lacerata (Arber) as a new combination by Herbst 
et al. (2001). Rochipteris lacerata was described 
originally from New Zealand by Arber (1917) as 
Chiropteris lacerata and after detailed discussion by 
Retallack (1980, 1983) was transferred to the genus 
Ginkgophytopsis. Rochipteris lacerata sensu Herbst 
et al. (2001) is larger than R. sinuosa, is deeply incised 
into several parallel-sided segments and has straight 
parallel venation that dichotomises and anastomoses 
to form long areoles. Rochipteris copiapensis 
(Solms-Laubach) sensu Herbst et al. (2001) is a large 
leaf divided into two equal segments with straight, 
bifurcating and anastomosing venation. In outline 
and venation pattern, R. sinuosa differs from the 
ten illustrated but undescribed Kannaskoppifolia 
= Rochipteris leaves from the Molteno Formation 
(Anderson and Anderson 2003). 


Rochipteris nymboidensis Holmes and Anderson sp. 
nov. 
Figures 24A—D 


Diagnosis 

A small cuneate leaf, lateral margins concave, 
apical margin convex, entire; venation dense, straight 
and parallel, dichotomising, very rarely anastomosing; 
vein density c. 30-35 per 10 mm. 


Description 

Based on a single specimen. A cuneate leaf 63 
mm long, with leaf base missing, 50 mm wide at the 
entire to slightly undulate apex. Angle of divergence 
from base c. 25°, increasing to 90° at lamina apex. 
Venation very fine, parallel, straight, dichotomising 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


and very rarely anastomosing. Density of veins across 
mid-upper portion of lamina c. 30-35 per 10 mm. 


Holotype 
AMEF126846 Australian Museum, Sydney. 


Type Locality 
Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures. 


Name Derivation 
nymboidensis, referring to the Type Locality. 


Discussion 

The entire leaf with very dense venation with only 
rare cross connections distinguishes this leaf from all 
other described species of Rochipteris. Rochipteris 
nymboidensis is similar in outline but differs by the 
denser venation and fewer anastomoses from the 
undescribed Kannaskoppifolia sp. D of Anderson and 
Anderson (2003). 


Rochipteris pusilla Holmes and Anderson sp. nov. 
Figures 25A—C 


Diagnosis 

A very small cuneate leaf, lateral margins 
slightly convex, apex entire to undulate; venation 
dichotomising to five times from base, becoming more 
dense apically, conjoining to form linear areoles in 
apical half of lamina; in upper 2/3 of lamina venation 
density c. 21 per 10 mm. 


Description 

Based on a single almost complete leaf and its 
counterpart. Lamina narrow cuneate, 20 mm long, 14 
mm wide; leaf base truncate 1.5 mm wide, diverging 
at ca. 45°, lateral margins entire, slightly convex; 
apex entire to slightly undulate. Two veins enter the 
base of the lamina, each bifurcates five or six times 
to terminate at distal margin. In the distal 1/3 of the 
lamina adjacent veins sometimes conjoin to form 
elliptic areoles, which become shorter and narrower 
towards the leaf apex; vein density across the upper 
2/3 of the lamina is c. 21 per 10 mm. 


Holotype 
AMF 126854 and counterpart AMF 126855; 
Australian Museum, Sydney. 


Type Locality 


Coal Mine Quarry, Nymboida. Basin Creek 
Formation, Nymboida Coal Measures. 


Proc. Linn. Soc. N.S.W., 126, 2005 


Name Derivation 
pusilla, (Lat.), very small; this being the smallest 
Rochipteris species as yet described. 


Discussion 

The _previously-described smallest _ leaf, 
Rochipteris tasmanica (Walkom) comb. nov., differs 
from R. pusillia by its larger size, the lamina expanding 
more widely and with a sparser, more open network 
of veins with a density of c. 10 per 10 mm (Walkom 
1925) The venation of R. pusilla is somewhat similar 
to that in the small but less diverging leaves illustrated 
as Kannaskoppifolia sp. A and K. sp. B by Anderson 
and Anderson (2003) from the Molteno Formation of 


_ South Africa. 


Sedis Incertae 


Genus Walkomiopteris Holmes and Anderson gen. 
nov. 


Walkomiopteris eskensis (Walkom) gen. et comb. 
nov. 
Figures 26A—F 


Type species Sphenopteris eskensis Walkom 1928, 
P1.16,3; text fig.4 


Combined diagnosis 

Small wedge-shaped to semi-circular leaves 
apparently arranged in pairs, axis unknown; proximal 
portion of lamina contracted to petiole-like base; 
primary vein thick at base, dichotomising up to three 
times to form sparse radiating veins to lamina apical 
margin. 


Description 

Based on the type specimen of Walkom (1928) 
and four additional specimens from the Nymboida 
quarries. The individual leaves are conjoined into 
pairs; 17-30 mm long, 15-20 mm wide, wedge- 
shaped to semicircular, margin entire or variously 
shallowly-lobed; contracted into a petiole-like base 
to 5 mm long; a stout midvein enters the base of 
the lamina and soon dichotomises up to three times, 
with the fine venules radiating distally; c. 16 vein 
endings around lamina apical margin. Walkom’s 
specimen from the Esk Beds of Queensland (Figure 
26A) shows a cluster of eight irregularly-arranged 
but not connected leaves, the best preserved leaf 
is c. 20 mm long, 15 mm wide with fine radiating 
and dichotomising veins. The Nymboida material 
is larger, to 30 mm x 20 mm. On two specimens 


49 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


(Figures 26B,C,D) leaves are conjoined into pairs 
while AMF113490 (Figure 26F) suggests a whorled 
arrangement but no axis or stem is preserved. While 
cuticle is not preserved, specimen AMF113492 shows 
an impression of cellular structure of rounded thick- 
walled cells and elongated rectangular cells along the 
veins (Figure 27C). 


Material 

Type specimen F1729 Queensland Museum, 
Brisbane, from railway cutting near Ottaba 
railway station. AMF113440, Coal Mine Quarry; 
AMF113491—3, AMF126848-9, Reserve Quarry 
Nymboida. 


Name derivation 

Walkomiopteris — for the eminent palaeobotanist 
and mentorto WBKH, DrA.B. Walkom, who described 
the type material from the Esk Beds of Queensland. 


Discussion 

Walkom (1928) noted that this was a unique form 
of leaf in the Australian Mesozoic. In the mistaken 
belief that the leaves were pinnately connected to a 
rachis, he placed them in the genus Sphenopteris, a 
generalised leaf form with similar venation and which 
ranges from Devonian to Jurassic. Sphenopteris 
probably includes both ferns and pteridosperms 
(Boureau 1975). Anderson and Anderson (1983 P1.9 
figs 2,3)illustrated as foliage gen. B, sp.A, paired leaves 
with radiating venation similar to Walkomiopteris, 
but later collected material reveals that they belonged 
to a pinnate fern (Anderson and Anderson in press). 


CONCLUSION 


Leaves of the form-genera Lepidopteris and 
Kurtziana are preserved on 3% and 2% respectively of 
catalogued slabs in the Holmes Nymboida Collection. 
This has provided ample material to appreciate the 
range of variation existing in the genera. Recognising 
this variation we have placed the Lepidopteris 
leaves into three ‘species complexes’, each complex 
includes a range of variation with intergrading forms 
linking the complexes. One leaf form without links 
to the “species complexes’ is described as the new 
species Lepidopteris dissitipinnula. Kurtziana leaves 
are separated into two ‘species complexes’ based on 
the dorsal or lateral attachment of the pinnae; each 
complex includes leaves of variable morphology. 
Leaves of Rochipteris are rare, but on selected 
diagnostic features we have erected six new morpho- 
species. Due to the very limited material, the variation 


50 


that may exist within a ‘species’ or the possibility of 
intergrading forms between ‘species’ is unknown. 
The leaf morphology of Walkomiopteris eskensis 
is unique among Gondwanan Triassic plants. This 
morpho-species probably represents the foliage of a 
gymnospermous plant. 


ACKNOWLEDGEMENTS 


W.B.K.H. deeply appreciates the help by his late 
wife, Felicity, and family in collecting the Nymboida 
material over many years. The late Mr Brian Foley, quarry 
operator, was always available and willing to turn over 
more mountains of material for inspection. We thank Ros 
Coy for providing photographic equipment and the staff 
at the National Botanic Institute, Pretoria, for the use of 
facilities. W.B.K.H. is assisted by a grant from the Betty 
Main Research Fund. 


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- Holmes, W.B.K., 2003. The Middle Triassic flora of the 


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New South Wales. Part 3. Fern-like foliage. 
Proceedings of the Linnean Society of NSW 124, 53- 
108. 

Holmes, W.B.K. and Anderson, H.M., in press. The 
Middle Triassic flora of the Basin Creek Formation, 
Nymboida Coal Measures, New South Wales. Part 4. 
Dicroidium. Proceedings of the Linnean Society of 
NSW 125, 

Holmes, W.B.K. and Ash, S.R., 1979. An early Triassic 
megafossil flora from the Lorne Basin, New South 
Wales. Proceedings of the Linnean Society of NSW 
103, 47-70. 

ICBN. 2001. “International Code of Botanical 
Nomenclature, Saint Louis Code (2000)’. W. Greuter 
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Johnston, R.M., 1888. “Systematic account of the geology 
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Kurtz, F., 1921 Atlas de las plantas fosiles de la Republica 
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en Cordoba 7, 129-158. 

Menendez, C.A., 1951. La flora de la Formacion Llantenes 
(Provincia de Mendoza). Revista Instituto Nacionale 
de Investigaciones en Ciencias Naturales (Botanica) 
2, 147-261. 

Meyen, S.V., 1987. Fundamentals of palaeobotany. 
(Chapman and Hall, London). 

Petriella, B. and Arrondo, O.G., 1982. El genero 
Kurtziana Frenguelli: su morphologia y vinculaciones. 
Ameghiniana 19, 209-215. 

Poort, R.J. and Kerp, J.H.F., 1990. Aspects of Permian 
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and Central Europe and a reclassification of species 
formerly included in Peltaspermum, Harris. Review of 
Palaeobotany and Palynology 63, 197-225. 

Retallack, G.J., 1977. Reconstructing Triassic vegetation 
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biostratigraphy of Gondwanaland. Alcheringa 1, 247- 
278. Alcheringa-fiche 1, G1-J16. 

Retallack, G.J., 1980. Middle Triassic megafossil plants 


51 


TRIASSIC FLORA FROM NY MBOIDA - SOME GYMNOSPERM GENERA 


and trace fossils from Tank Gully, Canterbury, New 
Zealand. Journal of the Royal Society of New Zealand 
10, 31-63. 

Retallack, G.J., 1983. Middle Triassic megafossil marine 
algae and land plants from near Benmore Dam, 
southern Canterbury, New Zealand. Journal of the 
Royal Society of New Zealand 13, 129-154. 

Retallack, G.J., 1995. An early Triassic fossil flora from 
Culvida Soak, Canning Basin, Western Australia. 
Journal of the Royal Society of Western Australia 78, 
57-66. 

Retallack, G.J., 2002. Lepidopteris callipteroides, an 
earliest Triassic seed fern of the Sydney Basin, 
southeastern Australia. A/cheringa 26, 475-500. 

Retallack, G.J., Gould, R.E. and Runnegar, B., 1977. 
Isotopic dating of a middle Triassic megafossil flora 
from near Nymboida, north-eastern New South Wales. 
Proceedings of the Linnean Society of NSW 101, 77- 
113. 

Rigby, J.F., 1977. New collections of plants from the Esk 
Formation, south-eastern Queensland. Queensland 
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Russel, N.J., 1994. A palaeothermal study of the Clarence- 
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Seward, A.C., 1903. Fossil floras of Cape Colony. Annals 
of the South African Museum 4, 1-122. 

Shirley, J., 1898. Additions to the fossil flora of 
Queensland. Queensland Geological Survey Bulletin 


32 


7, 19-25. 

Solms-Laubach, H. and G. Steinmann, G., 1899. 

Das Auftreten und die Flora der Rhatischen 
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Thomas, H.H., 1933. On some pteridospermous 
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Philosophical Transactions of the Royal Society of 
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Townrow, J.A., 1956. The genus Lepidopteris and its 
southern hemisphere species. Avhandlinger utgitt 
av Det Norske Videnskaps-Akademi I Oslo. 1. mat.- 
Natury. Klasse 1956 2, 3-28. 

Townrow, J.A.,1960. The Peltaspermaceae, a 
pteridospermous family of Permian and Triassic age. 
Palaeontology 3, 333-361. 

Townrow, J.A., 1966. On Lepidopteris madagascariensis 
Carpentier (Peltaspermaceae). Journal and 
Proceedings of the Royal Society of NSW 98, 203-216. 

Walkom A.B., 1925. Notes on some Tasmanian Mesozoic 
plants.Pt.1. Papers and Proceedings of the Royal 
Society of Tasmania 1924, 73-89. 

Walkom A.B., 1928. Fossil plants from the Esk district, 
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Zamuner, A., Atrabe, A. and Ganuza, D., 1999. A new 
peltasperm (Gymnospermophyta) from the Middle 
Triassic of Argentina. A/cheringa 23, 185-191. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 1, A,B. ‘Lepidopteris madagascariensis complex’ A. AMF 126801; B. AMF126802. Both Coal Mine 
Quarry. Scale bar = 1 cm 


Proc. Linn. Soc. N.S.W., 126, 2005 53 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 2. A. Centre leaf ‘Lepidopteris madagascariensis complex’, other leaves intergrading with 
‘Lepidopteris africana complex’. AMF126803, Coal Mine Quarry. B,C. ‘Lepidopteris mada- 
gascariensis complex’. B. AMF126804, Reserve Quarry. C. AMF126805, Coal Mine Quar- 
ry. D. Leaves grading to ‘L. africana complex’. AMF126806, Reserve Quarry. Scale bar = 1 cm. 


54 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 3. A,B. ‘Lepidopteris africana complex’. A. AMF 126807. B. AMF126808. Both Coal Mine Quarry. 
Scale bar = 1 cm. : 


Proc. Linn. Soc. N.S.W., 126, 2005 


35 


TRIASSIC FLORA FROM NY MBOIDA - SOME GYMNOSPERM GENERA 


Figure 4. A. ‘Lepidopteris africana complex’. AMF 126809. B. ‘L. africana complex’, leaf at top right “L. 
stormbergensis complex’. AMF 12806810. Both Coal Mine Quarry. Scale bar = 1 cm. 


56 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 5. A,B. ‘Lepidopteris africana complex’. A. AMF126811. B. AMF126812. C. ‘L. stormbergensis 
complex’. AMF126813. All from Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


57 


TRIASSIC FLORA FROM NY MBOIDA - SOME GYMNOSPERM GENERA 


‘ } 


oe 


Figure 6. A,B. ‘Lepidopteris africana complex’ intergrading with ‘L. stormbergensis com- 
plex’. A. AMF126814, Reserve Quarry. B. AMF126815, Coal Mine Quarry. C. Small com- 
plete leaf of ‘Z. stormbergensis complex’. AMF126816, Coal Mine Quarry. Scale bar = 1 cm. 


58 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 7. A. Intergrading form between ‘L. africana complex’ and ‘L. stormbergensis complex’. 


AMF 126817, Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


59 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


> 


eo 


Figure 8. A,B. ‘Lepidopteris stormbergensis complex’. A. AMF126818. B. AMF126819. Both from Coal 
Mine Quarry. Scale bar = | cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


: 4 x = Y ~ ; : 
Bi CS gee 


Eine = 


Figure 9. A,B. ‘Lepidopteris stormbergensis complex’. A. AMF 126820. B. AMF126851, lob 
ern arrowed. Both from Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


a 


ed zwisherfied- 


61 


TRIASSIC FLORA FROM NY MBOIDA - SOME GYMNOSPERM GENERA 


Figure 10. A-C. Lepidopteris dissitipinnula sp. noy. A, B. AMF 113528. Holotype. C. AMF126823. Both from 
Coal Mine Quarry. Scale bar = 1 cm. 


62 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 11. A-E. Peltaspermum cf monodiscum. A,B. AMF 126824. C. AMF126825. D,E. AMF126826. F,G. 
Peltaspermum sp. A. AMF 126852. All from Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


63 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 12. A-C. Antevsia sp. A. A,B. AMF126828. C. AMF126829. Both from Coal Mine Quarry. Scale bar = 
lcm. 


64 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 13. A,B. ‘Kutrziana brandmayri complex’. A. AMF126830. B. AMF 126831. Both from Coal Mine 
Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


65 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 14. A-D. ‘Kurtziana cacheutensis complex’. A. AMF 126832, showing venation. B. AMF126833. C. 
AMF 126834. D. AMF126835. All from Coal Mine Quarry. Scale bar = | cm. 


66 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 15. A. Leaf assemblage of ‘Kurtziana cacheutensis complex’. A. AMF 126836. Coal Mine Quarry. 
Scale bar = | cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 67 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 16. A,B. ‘Kurtziana cacheutensis complex’. A. AMF 126837. B. AMF126838, leaf associated with 
Rissikia sp. Both from Coal Mine Quarry. Scale bar = 1 cm. 


68 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 17. A. ‘Kurtziana cacheutensis complex’. AMF 126839. Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


TRIASSIC FLORA FROM NY MBOIDA - SOME GYMNOSPERM GENERA 


Figure 18. A-C. Rochipteris obtriangulata sp. nov. A. Portions of two whorls of leaves, AMF 126840 holo- 
type on left and AMF126841. B. AMF126840 and C. AMF126841 enlarged to show venation. Coal Mine 
Quarry. Scale bar = | cm. 


70 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 19. A-D. Rochipteris obtriangulata sp. nov., enlarged to show venation. AMF 126842, counterpart of 
holotype. Coal Mine Quarry. Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 7\ 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 20. A-C. Rochipteris tubata sp. nov. Holotype. AMF 126843. Coal Mine Quarry. 
Scale bar = 1 cm. 


72 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 21. A. Rochipteris incisa sp. nov. Holotype on left, AMF126844, on right, AMF 126827, juvenile leaf? 
B. AMF 126827. C. AMF126844, to show venation. Reserve Quarry. Scale bar = | cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 73 


74 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 22. A. Rochipteris incisa sp. nov. Holotype. AMF 126844. Reserve Quarry. Scale bar = | cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 23. A-C. Rochipteris sinuosa sp. nov. Holotype. AMF 126845. Coal Mine Quarry. Scale bar 
= cin: 


Proc. Linn. Soc. N.S.W., 126, 2005 


ue) 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


scale 


> 


Figure 24. A-D. Rochipteris nymboidensis sp. nov. Holotype. AMF 126846. Coal Mine Quarry. A and B 


bar 


1 cm.; C and D, scale bar = 0.2 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


76 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 25. A-C. Rochipteris pusilla sp. nov. Holotype. AMF126854. Coal Mine Quarry. 
Scale bar = 1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 


77 


TRIASSIC FLORA FROM NYMBOIDA - SOME GYMNOSPERM GENERA 


Figure 26. A-F. Walkomiopteris eskensis gen. et comb. nov. A. Holotype, QMF1729. Cutting near Ottoba 
railway station, Queensland. B. AMF113491. C. Counterpart of B. D,E. AMF113493. F. AMF113440. B-E. 
Reserve Quarry; F. Coal Mine Quarry. Scale bar = 1 cm. 


78 Proc. Linn. Soc. N.S.W., 126, 2005 


W.B.K. HOLMES AND H.M. ANDERSON 


Figure 27. A-C. Walkomiopteris eskensis gen. et comb. nov. AMF113492. Reserve Quarry. A,B. Scale bar = 1 
cm. C. To show cell structure. Scale bar = 0.1 cm. 


Proc. Linn. Soc. N.S.W., 126, 2005 79 


(CRAKS DEH LO“A 2SMI0K MLW ERM GUNERA 


7 


aaa 


clway Mande, Oieenelad. 6 Avie an = 
'@ (Ouarry L Coal Mine Cheatr®. Scolr hat Tom. = =a 
 ~ and olno2 ELA \erimuO svrses HL. SOCKET TMA vou pron oe Mee eres 
| tie LO ae al 
; ay 


a ee 


Revision of Discomesites and Estaingia (Trilobita) from the 
Lower Cambrian Cymbric Vale Formation, Western New 
South Wales: Taxonomic, Biostratigraphic and Biogeographic 


Implications 


JOHN R. PATERSON 


Centre for Ecostratigraphy and Palaeobiology, Department of Earth and Planetary Sciences, Macquarie 


University, NSW 2109 (agnostid@hotmail.com) 


Paterson, J.R. (2005). Revision of Discomesites and Estaingia (Trilobita) from the Lower Cambrian 
Cymbric Vale Formation, western New South Wales: taxonomic, biostratigraphic and biogeographic 
implications. Proceedings of the Linnean Society of New South Wales 126, 81-93. 


The taxonomy of Discomesites and Estaingia from the Lower Cambrian Cymbric Vale Formation 
of western New South Wales is revised. Discomesites is regarded as a valid subgenus of Pagetides. 
Pagetides (Discomesites) fragum is considered a senior subjective synonym of P. (D.) lunatulus. Pagetides 
(Discomesites) spinosus from the Shackleton Limestone in the Holyoake Range, Transantarctic Mountains, 
is considered to be a junior subjective synonym of P. (D.) fragum. Estaingia cerastes from the Cymbric 
Vale Formation is considered to be synonymous with Hsuaspis cf. H. bilobata from the Shackleton 
Limestone. The Cymbric Vale Formation trilobite fauna is of late Early Cambrian (late Botoman) age, 
equivalent to the Pararaia janeae Zone of South Australia, based on correlation of the Syringocnema 
favus archaeocyathan fauna. Absolute ages of recently dated tuffs from the Cymbric Vale and Billy Creek 
Formations are questioned, based on new information regarding the stratigraphic position of the Cymbric 
Vale Formation tuff in relation to archaeocyathan and trilobite biostratigraphy. The co-occurrence of 
Pagetides (Discomesites) fragum and Estaingia cerastes in the upper part of the Cymbric Vale Formation 
and in the Shackleton Limestone represents the first species-level correlation between the Lower Cambrian 
of Australia and Antarctica using trilobites. The distribution of these trilobite species, in association with 
the Syringocnema favus archaeocyathan fauna, provides supporting evidence that Australia and Antarctica 
were connected by a continuous carbonate-detrital shelf during the late Early Cambrian (mid-late Botoman), 


allowing faunal exchange between these regions. 


Manuscript received 2 June 2004, accepted for publication 18 December 2004. 


KEYWORDS: Antarctica, Australia, biogeography, biostratigraphy, Botoman, Early Cambrian, 
geochronology, Gondwana, New South Wales, Trilobita. 


INTRODUCTION 


Since the publication of Opik’s (1975b) 
taxonomic study on the trilobites from the Lower 
Cambrian Cymbric Vale Formation, several important 
palaeontological and geochronological studies on the 
Early Cambrian of Australia and Antarctica, relevant 
to the Cymbric Vale Formation, have been published 
(Kruse 1978, 1982; Debrenne and Kruse 1986; 
Bengtson et al. 1990; Zhuravlev and Gravestock 1994; 
Palmer and Rowell 1995; Jago et al. 1997; Jenkins 
et al. 2002). The scope of this study is to revise the 
taxonomy of two key trilobite taxa from the Cymbric 
Vale Formation, Discomesites and Estaingia, as well 
as review the archaeocyathan biostratigraphy of the 


Cymbric Vale Formation and stratigraphic position 
of a recently dated tuff within the unit, allowing the 
Cymbric Vale trilobite fauna to be placed in both a 
biostratigraphic and biogeographic context. 

The Cymbric Vale Formation is part of the Lower 
to early Middle Cambrian Gnalta Group, which 
crops out in the Mt Wright area of western New 
South Wales (Fig. 1). The Gnalta Group comprises 
(in ascending order): the Mount Wright Volcanics, 
Cymbric Vale Formation and Coonigan Formation. 
The Lower Cambrian Mount Wright Volcanics and 
overlying Cymbric Vale Formation are considered to 
be conformable, based on common archaeocyathan 
faunas (Kruse 1982). The uppermost beds of the 
Cymbric Vale Formation are disconformably overlain 
by the early Middle Cambrian ‘first discovery 


REVISION OF TRILOBITE GENERA D/SCOMESITES AND ESTAINGIA 


SYDNEY 


ea Post-Palaeozoic cover 


Mt y Upper Cambrian- 
Arrowsmith Yt fener Ordovician 
= Lower-Middle Cambrian 


Proterozoic 


Figure 1. Generalised geological map of northwest- 
ernNewSouth Wales, showinglocationof{MtWright; 
modified from Shergold et al. (1982, text-fig. 1D). 


limestone’ of the Coonigan Formation (Roberts and 
Jell 1990). 

The Cymbric Vale Formation attains a maximum 
thickness of 1900 m and consists predominantly of 
interbedded blue, green and grey-white chert and green 
to brown tuff (Kruse 1982). Archaeocyath-bearing 
limestone lenses occur throughout the formation and 
contain two distinct archaeocyathan faunas (Fig. 2): 
Fauna | (L96-L99) occurs in the upper Mount Wright 
Volcanics and in the lower part of the Cymbric Vale 
Formation and is assigned an early Botoman age; 
Fauna 2 (L100-L101) occurs in the upper Cymbric 
Vale Formation and has been assigned a mid-late 
Botoman age (Kruse 1978, 1982; Zhuravlev and 
Gravestock 1994; Kruse and Shi in Brock et al. 2000). 
The uppermost beds of the Cymbric Vale Formation 
consist of well-bedded lithic and feldspathic siltstone 
and sandstone interbedded with impure iron-rich 


82 


carbonate rocks with abundant trilobites, molluscs, 
brachiopods, eocrinoids and sponge spicules (Opik 
1975b; Kruse 1982; Jago et al. 1997). 

The first formal taxonomic study of the Early 
Cambrian trilobites from the Cymbric Vale Formation 
was by Opik (1975b). He recorded the trilobites 
Dinesis aff. granulosus (Lermontova), Estaingia 
bilobata Pocock, Strenax cerastes Opik, Strenax 
(Sematiscus) fletcheri Opik, Serrodiscus daedalus 
Opik, Meniscuchus menetus Opik, Discomesites 
fragum Opik, Discomesites lunatulus Opik and 
Pagetia sp. nov. Opik (1975b) correlated the Cymbric 
Vale fauna with Daily’s (1956) South Australian 
faunal assemblages 9, 11 and 12, equivalent to the 
Pararaia janeae Zone of South Australia (Bengtson et 
al. 1990), and to the Botoman Sanashtyk’ gol Horizon 
of the Altay-Sayan region of Siberia. 

Jago et al. (1997) recorded a trilobite faunule from 
a new locality within the Cymbric Vale Formation, 
reassigning the species Estaingia bilobata Pocock 
and Strenax cerastes Opik, originally documented by 
Opik (1975b), to a single redefined species, Hsuaspis 
cerastes (Opik), and also described Redlichia cf. 
ziguiensis Lin, a taxon previously unknown from 
the Cymbric Vale Formation. Jago et al. (1997) also 
suggested that the Cymbric Vale trilobite fauna is of 
late Early Cambrian (late Botoman) age. 


BIOSTRATIGRAPHIC IMPLICATIONS 


The taxonomic revision herein of Pagetides 
(Discomesites) fragum Opik, 1975b and Estaingia 
cerastes (Opik, 1975b) from the Cymbric Vale 
Formation of western New South Wales is based on 
reexamination of Opik’s (1975b) type material plus 
additional collections housed at Geoscience Australia 
(Canberra) and provides the first direct interregional 
correlation between the Lower Cambrian of Australia 
and Antarctica using trilobites. Palmer (in Palmer and 
Rowell 1995) described a number of Early Cambrian 
trilobite assemblages, ranging in age from Atdabanian 
to Toyonian(?), from the Shackleton Limestone in 
the Holyoake Range of the central Transantarctic 
Mountains. One of the Botoman assemblages 
(‘Assemblage 3’) contains the taxa Pagetides 
(Discomesites) spinosus Palmer and Hsuaspis cf. H. 
bilobata (Pocock). The synonymy herein of Pagetides 
(Discomesites) spinosus with P. (D.) fragum, and 
Hsuaspis cf. H. bilobata with Estaingia cerastes 
permits direct correlation of the trilobite fauna from 
the uppermost Cymbric Vale Formation with trilobite 
‘Assemblage 3’ from the Shackleton Limestone. 
Unfortunately, further support for this correlation. 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. PATERSON 


South 

Australian Flinders Ranges 
trilobite (South Australia) 
zones 


Wirrealpa 
Limestone 


| TOYON. {Siberian stages 


Oraparinna 


P. 
bunyerooensis 


NO 
ZONATION 


BOTOMAN 


Ajax Limestone 


NO 
ZONATION 


HAWKER GROUP 


ATDABANIAN | 


Aroona Creek 
Limestone 


Billy Creek Formation 


Narina 
Greywacke 


Mernmerna 
Formation 


Woodendinna Dol. Limestone 


Parachilna Formation 


Mt Wright 
(western New 
South Wales) 


522.841.8 Ma 


(SL13) roi 
SN eh sti Lol cat =| : 
‘avus es g S 
beds = 
3 0 
Cymbric = > 
Vale g i 
i r} 
Formation SiG 
517.8+2.1 Ma Q 
Petuereli cat 
525+8 Ma ——| 2 
(SL13) 3 
S 
Mount o 
o 


Wright 
Voleanics 


Figure 2. Correlation diagram of the Lower Cambrian successions of Mt Wright (western New South 
Wales) and the Flinders Ranges (South Australia). L96-L99 represent Fauna 1 and L100-L101 represent 
Fauna 2 of Kruse (1982). Tuff ages: Billy Creek Formation, 522.8 + 1.8 Ma (Gravestock and Shergold 2001); 
Cymbric Vale Formation, 517.8 + 2.1 Ma (Jenkins et al. 2002) and 525 + 8 Ma (Zhou and Whitford 1994). 


based on similar archaeocyathan faunas from the 
Cymbric Vale Formation (Kruse 1978, 1982) and the 
Shackleton Limestone (Debrenne and Kruse 1986) is 
complicated. Trilobites and archaeocyaths described 
from the Shackleton Limestone were sampled from 
different localities in the Holyoake Range; see 
discussion by Palmer and Rowell (1995:4). Exposed 
sections of Shackleton Limestone in the Holyoake 
Range are considerably thick (up to 200 m) and seem 
to conform to a coherent stratigraphy; however, each 
section is bounded either by intensely disturbed zones 
of folding or faulting, or fields of ice or névé (Rowell 
et al. 1988:399; Rees et al. 1989:343). Sampling of 
trilobites and archaeocyaths from disparate localities 
in the Holyoake Range, coupled with the structural 
complexity of the region, makes correlation between 
faunas difficult. 

Documentation of archaeocyathan faunas 
from the Cymbric Vale Formation (Kruse 1978, 
1982), and the development of a preliminary Early 
Cambrian archaeocyathan biozonation for Australia 
(Zhuravlev and Gravestock 1994), has allowed for the 


Proc. Linn. Soc. N.S.W., 126, 2005 


correlation of the Cymbric Vale Formation with the 
Lower Cambrian succession in South Australia. As 
noted by Kruse and Shi (in Brock et al. 2000), Fauna 
2 from the upper Cymbric Vale Formation can be 
correlated with the mid-late Botoman Syringocnema 
favus beds of the Adelaide Geosyncline (Fig. 2), 
based on five co-occurring species. Zhuravlev and 
Gravestock (1994, Table 2) have recorded the S. 
favus beds occurring in the Koolywurtie Member 
of the Parara Limestone on Yorke Peninsula in the 
Stansbury Basin, and in the upper Ajax Limestone 
(Ajax Mine and Mount Scott Range) and upper 
Wilkawillina Limestone (Wilkawillina Gorge) in the 
Arrowie Basin. Trilobites from coeval beds in the 
Flinders Ranges (Arrowie Basin) are representatives 
of the Pararaia janeae Zone (Bengtson et al. 1990). 
While species-level correlation between the Cymbric 
Vale and Flinders Ranges trilobites is not possible, 
generic similarities are evident with the occurrence of 
Serrodiscus and Estaingia in both areas (Opik 1975b; 
Bengtson et al. 1990). Since the Cymbric Vale trilobite 
fauna occurs in the uppermost part of the formation, 


83 


REVISION OF TRILOBITE GENERA DISCOMESITES AND ESTAINGIA 


stratigraphically above archaeocyathan Fauna 2 (= S. 
/avus beds), it is equivalent to the Pararaia janeae 
Zone (sensu lato) or possibly even younger, thus 
supporting the late Botoman age suggested by Jago et 
al. (1997). 

A dated felsic tuff from the Cymbric Vale 
Formation was recently re-calculated by Jenkins et 
al. (2002) using the SL13 U-Pb SHRIMP method. 
The tuff was originally collected by Zhou (1992) and 
subsequently dated by Zhou and Whitford (1994), 
yielding an age of 525 + 8 Ma (SL13). However, 
Jenkins et al. (2002) produced an age of 517.8 + 2.1 
Ma. Unfortunately, neither publication citing the age 
of the Cymbric Vale tuff (Zhou and Whitford 1994; 
Jenkins et al. 2002) provided information about the 
stratigraphic position of the tuff horizon, especially 
in regard to the local biostratigraphy. However, 
Zhou’s (1992) unpublished PhD thesis does provide 
an Australian standard national grid reference for the 
dated sample (i.e., 6332E; 65498N). Based on this 
grid reference and mapping by Zhou (1992, map 7) 
and Kruse (1982, text-Fig. 2), the dated tuff appears 
to have been collected in the vicinity of, or possibly 
stratigraphically above, Kruse’s (1982) localities L98 
and L99 in the lower Cymbric Vale Formation (Fig. 
Dy 

In light of the known stratigraphic position 
of the dated Cymbric Vale tuff, there appears to be 
an age discrepancy of dated tuffs from the Cymbric 
Vale Formation and the lower Billy Creek Formation 
(Flinders Ranges), based on the correlation of the 
Syringocnema favus beds. Gravestock and Shergold 
(2001) reported a SHRIMP age of 522.8 + 1.8 Ma 
from a tuff within the lower Billy Creek Formation 
using the same standard (SL13) used by Jenkins et 
al. (2002) to re-calculate the Cymbric Vale tuff. 
This means that the “older” (522.8 Ma) Billy Creek 
Formation tuff occurs stratigraphically above the 
Syringocnema favus beds in the Flinders Ranges, and 
the “younger” (517.8 Ma) Cymbric Vale tuff occurs 
stratigraphically below archaeocyathan Fauna 2 (= 
S. favus beds) in western New South Wales (Fig. 
2). This discrepancy implies that the age of the tuff 
horizon in the Cymbric Vale Formation or Billy 
Creek Formation is erroneous, or perhaps both ages 
are incorrect. This age discrepancy may be related to 
the standard used, since the reliability of the SL13 
standard has been questioned in recent years (see 
Jago and Haines 1998 for detailed discussion). It is 
interesting to note that the original age of the Cymbric 
Vale tuff of 525 + 8 Ma, calculated using the SL13 
standard (Zhou and Whitford 1994; Jago and Haines 
1998), is more in accord with the archaeocyath and 
trilobite biostratigraphy and the age of the Billy 


84 


Creek Formation tuff. Furthermore, ages of the 
Cymbric Vale and Billy Creek Formation tuffs using 
the alternative QGNG standard yield dates of 531.8 
+ 8 Ma and 529.6 + 1.8 Ma respectively (Jago and 
Haines 1998), that better conform to archaeocyath 
and trilobite biostratigraphy. 


BIOGEOGRAPHIC IMPLICATIONS 


Jago (in Brock et al. 2000) noted that Early 
Cambrian trilobites from Australia have close faunal 
ties with other regions of East Gondwana, such as 
Antarctica, South and North China, Iran and India. 
However, Jago also demonstrated that faunal links 
with distantly separated palaeogeographic regions 
such as western Gondwana (e.g. Morocco), Laurentia 
and Siberia are not uncommon. In discussing the 
biogeographic patterns of Early Cambrian trilobite 
faunas from Antarctica, Palmer (in Palmer and 
Rowell 1995:5) observed that ‘Antarctic faunas do 
not show any consistent similarity to faunas from 
any one geographic region of the rest of Gondwana’. 
This appears to be true for any palaeogeographic 
region when treating Early Cambrian trilobite faunas 
as a whole, although by closely observing faunal 
assemblages from specific time intervals, whether 
they be zones or stages, distinct biogeographic 
patterns often emerge (for example, see Theokritoff 
1979, 1985; Burrett and Richardson 1980; Alvaro 
et al. 2003). Moreover, Alvaro et al. (2003:17) have 
commented that in the Lower Cambrian ‘the ideal of a 
global biostratigraphy and palaeobiogeography suffers 
from both a relatively limited diversity of trilobites 
and their pronounced endemism. Furthermore, the 
distribution of trilobites is...strongly controlled by 
facies, so that even a precise interregional correlation 
is difficult’. 

The occurrence of Pagetides (Discomesites) 
fragum and Estaingia cerastes, in addition to the 
Syringocnema favus archaeocyathan fauna, in the 
Cymbric Vale Formation (Gnalta Shelf) and the 
Shackleton Limestone (Holyoake Range) provides 
supporting evidence that a continuous continental 
shelf connected southeastern Australia and the 
Transantarctic Mountains during the Early Cambrian 
(Rowell and Rees 1991; Courjault-Radé et al. 1992; 
Wrona and Zhuravlev 1996; Veevers etal. 1997; Brock 
et al. 2000; Veevers 2000). The palaeogeographic 
distance between the Gnalta Shelf and the Holyoake 
Range is estimated to be around 2500 km (Brock 
et al. 2000, Fig. 8), although the shelf may have 
extended as far as King George Island, implying a 
length of over 6500 km. Courjault-Radé et al. (1992, 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. PATERSON 


Fig. 2) have suggested that throughout the Early 
Cambrian, and indeed the Middle and Late Cambrian, 
a continuous carbonate-detrital shelf connected 
Australia and Antarctica. The widespread dominance 
of the Syringocnema favus fauna in Australia and 
Antarctica during the mid-late Botoman coincides 
with a global high sea level, allowing faunal exchange 
between these regions (Zhuravlev and Gravestock 
1994; Wrona and Zhuravlev 1996; Gravestock and 
Shergold 2001). This strong faunal link between 
Australia and Antarctica during the Botoman is 
also supported by conspecific occurrences of small 
shelly fossils (SSF), such as Ejiffelia araniformis 
(Missarzhevskty), Chancelloria —_ racemifundis 
Bengtson, Halkieria parva Conway Morris, Dailyatia 


ajax Bischoff, Lapworthella fasciculata Conway - 


Morris and Bengtson, Hyolithellus micans (Billings), 
Hyolithellus filiformis Bengtson, Byronia? bifida 
Wrona, and Aetholicopalla adnata Conway Morris, 
from glacial erratics of King George Island, Antarctica 
(Wrona 2004) and the Parara, Wilkawillina and Ajax 
Limestones and Kulpara and Mernmerna Formations 
‘in South Australia (Bengtson et al. 1990; Gravestock 
et al. 2001). 

The carbonate-detrital shelf connecting Australia 
and Antarctica may have persisted until at least the 
early Late Cambrian (Idamean), based on other 
conspecific occurrences of benthic, shelf-dwelling 
polymerid trilobites and other biotas. For example, 
Holmer et al. (1996) described an Early Cambrian 
lingulate brachiopod faunule, containing Eoobolus 
aff. E. elatus (Pelman), Karathele napuru (Kruse) 
and Vandalotreta djagoran (Kruse), from the glacial 
erratics of King George Island, Antarctica. These 
same brachiopod species have been documented 
from the Toyonian (latest Early Cambrian) Wirrealpa 
and Ramsay Limestones in South Australia (Brock 
and Cooper 1993; Gravestock et al. 2001), and early 
Middle Cambrian units in the Northern Territory: 
Tindall Limestone, Daly Basin (Kruse 1990); 
Montejinni Limestone of the Wiso Basin and Gum 
Ridge Formation of the western Georgina Basin 
(Kruse 1998); and the Top Springs Limestone, 
northern Georgina Basin (Kruse 1991). The latest 
Middle Cambrian (Glyptagnostus stolidotus Zone) 
trilobite species Rhodonaspis longula Whitehouse 
has been recorded from the Georgina Limestone, 
Glenormiston, Queensland, Australia (Whitehouse 
1939; Opik 1963) and inthe Spurs Formation, Northern 
Victoria Land, Antarctica (Jago and Cooper 2001). 
Furthermore, the early Late Cambrian (Idamean) 
trilobite species Protemnites magnificans Shergold 
and Webers has been recorded from the Dolodrook 
River limestones, eastern Victoria, Australia (Paterson 


Proc. Linn. Soc. N.S.W., 126, 2005 


and Laurie 2004) and from the Minaret Formation, 
Ellsworth Mountains, West Antarctica (Shergold and 
Webers 1992). 

The position of Australia and Antarctica during 
the Cambrian, coupled with the persistence of 
carbonate deposition in Australia and Antarctica 
throughout the Cambrian, indicates that these regions 
remained within the tropical Carbonate Development 
Zone (30 + 5° north and south latitudes) (McKerrow 
et al. 1992; Courjault-Radé et al. 1992; Brock et 
al. 2000; Li and Powell 2001). Hence migration of 
trilobite species between Australia and Antarctica 
was not inhibited by a latitudinal thermocline during 
the Cambrian. However, this does not explain the 
lack of conspecific occurrences of Early Cambrian 
trilobites between Australia and Antarctica, although 
one may argue that a paucity of data from Antarctica 
could be responsible. Moreover, there is a distinct 
absence of conspecific occurrences between western 
New South Wales (Gnalta Shelf) and South Australia 
(Adelaide Geosyncline). One possible explanation 
for the absence of conspecific occurrences of Early 
Cambrian trilobites between South Australia and 
Antarctica is the presence of a tectonic barrier during 
this time, i.e., the Kanmantoo Trough (Haines and 
Flottmann 1998; Fléttmann et al. 1998). The Early 
Cambrian palaeogeographic relationship between 
the eastern Arrowie Basin (eastern South Australia) 
and the Gnalta Shelf (western New South Wales) is, 
however, poorly understood. Various studies (e.g., 
Cook 1988; Gravestock and Hibburt 1991; Scheibner 
and Basden 1998; Zang 2002) have inferred 
connection between the eastern Arrowie Basin 
and the Gnalta Shelf via a common seaway. While 
correlation of the Syringocnema favus beds provides 
supporting evidence for a connected seaway during 
the late Early Cambrian (mid-late Botoman), there 
may have been some form of geographic barrier that 
inhibited trilobite migration between these regions. 
Firstly, it is important to note that the Cymbric Vale 
Formation trilobites occur stratigraphically above 
the S. favus beds and that during their temporal 
separation the palaeogeography between the eastern 
Arrowie Basin and the Gnalta Shelf may have altered 
significantly, severing ties between these regions. 
There are two possible palaeogeographic barriers 
that may have hindered migration of Early Cambrian 
trilobites between the eastern Arrowie Basin and the 
Gnalta Shelf: (1) the Mount Wright Volcanic Arc 
situated immediately to the west of the Gnalta Shelf; 
and (2) an inferred trough situated to the west of the 
Mount Wright Volcanic Arc (Scheibner and Basden 
1998, Fig. 14.6). A small outcrop of the Lower- 
Middle Cambrian Teltawongee beds, considered by 


85 


REVISION OF TRILOBITE GENERA D/ISCOMESITES AND ESTAINGIA 


Mills (1992) to be coeval with the Gnalta Group, 
occurs at the northern end of the Mount Wright Block 
at Nundora. The Teltawongee beds are thought to 
have been deposited as a turbidite slope-trough facies 
(Mills 1992). Scheibner and Basden (1998) suggested 
that the Teltawongee beds might extend beneath the 
large thickness of Devonian sediments within the 
Bancannia Trough, situated to the west of the Gnalta 
Shelf. This inferred trough would have created an 
oceanic barrier between the eastern Arrowie Basin 
and the Gnalta Shelf. 

Wrona (2004) observed that Lower Cambrian 
horizons in Australia and Antarctica that contain 
very similar faunas correspond with transgressive 
episodes that occurred during the early Botoman, 
late Botoman and middle Toyonian (Gravestock 
and Shergold 2001; Gravestock et al. 2001). Wrona 
(2004:52) suggested that several isolated basins 
might have existed along the East Antarctic craton 
and that those basins ‘connected only during the 
most prominent transgressions, thus allowing faunal 
exchange’. Alvaro et al. (2003) suggested that species 
migration of Cambrian trilobites along the western 
Gondwanan margin coincides with transgressive 
episodes and subsequent connection of neighbouring 
platforms. This also appears to have been the case 
in influencing trilobite migration patterns along the 
eastern Gondwanan margin, especially between 
Australia and Antarctica. 


SYSTEMATIC PALAEONTOLOGY 


Specimens come ftom the Commonwealth 
Palaeontological Collection (prefix CPC) housed at 
Geoscience Australia, Canberra. 


Order AGNOSTIDA Salter, 1864 
Suborder EODISCINA Kobayashi, 1939 
Family EODISCIDAE Raymond, 1913 
Genus PAGETIDES Rasetti, 1945 


Type species 
Pagetides elegans Rasetti, 1945, Early 
Cambrian, Sillery Formation, Quebec, Canada. 


Subgenus DISCOMESITES Opik, 1975b 
Type species 
Discomesites fragum Opik, 1975b, Early 


Cambrian, Cymbric Vale Formation, Mt. Wright, 
western New South Wales, Australia. 


86 


Discussion 

Palmer’s (in Palmer and Rowell 1995) treatment 
and diagnosis of Discomesites as a subgenus of 
Pagetides is supported here. 


Pagetides (Discomesites) fragum Opik, 1975b 
Fig. 3A-K, Fig. 4A-J 


Discomesites fragum Opik, 1975b:32, Pl. 5, Figs 
1-8. 

Discomesites lunatulus Opik, 1975b:34, Pl. 6, Figs 
1-4. 

Neopagetina sp., Rowell et al. 1989, p. 14, Fig. C. 

Pagetides (Discomesites) spinosus Palmer in Palmer 
and Rowell, 1995:7, Fig. 7. 


Material 

13 cranidia, 8 pygidia; CPC13176-13179 [Type 
material of Discomesites fragum]|, CPC13180- 
13183 [Type material of Discomesites lunatulus] and 
CPC37623-37635 [New topotype material from Site 
A of Opik (1975b)]. 


Discussion 

The characters that Opik (1975b) used 
to differentiate Discomesites fragum from D. 
lunatulus can be explained by ontogenetic variation, 
deformation, preservation or misinterpretation. Opik’s 
differential characters include: ornamentation, length 
of the occipital spine, pygidial shape and number of 
pygidial axial rings. 

Opik (1975b) distinguished Discomesites 
fragum from D. lunatulus by its dense granulose 
ornament; however, this difference is the result of 
either ontogenetic variation and/or preservation. 
Firstly, it is important to note that specimens of D. 
lunatulus are considerably smaller than those of D. 
jJragum. The type series cranidia of D. lunatulus range 
between 2.6-2.9 mm (n=2) in length (sag.), whereas 
the type series cranidia of D. fragum range between 
3.4-4.1 mm (n=3) in length (sag.). This correlation 
between size and ornamentation can also be observed 
in pygidia of Discomesites. Smaller pygidia (<2.7 mm 
in length) appear to lack the granulose ornamentation 
(Fig. 4C-H, J), whereas larger pygidia clearly show 
dense granulation (Fig. 4A-B, I). It is also worth 
mentioning that some specimens of Discomesites 
that lack granulose ornamentation, including those of 
D. lunatulus, have a coating of ‘desert varnish’ (iron 
oxide), while others appear to have been indurated 
or partially silicified. It is therefore possible that the 
difference in ornamentation may have been caused by 
weathering or diagenesis. 

The differentiation of Discomesites species — 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. PATERSON 


Figure 3. Pagetides (Discomesites) fragum Opik, 1975b. All scale bars = 1 mm. A, CPC13177, holotype 
cranidium, dorsal view, stereo pair; B, CPC13176, cranidium, dorsal view; C, CPC13178, latex cast 
of partial cranidium, dorsal view; D, CPC13180, holotype cranidium of Discomesites lunatulus Opik, 
1975b, dorsal view; E, CPC13181, cranidium, dorsal view; F, CPC37623, cranidium, dorsal view; 
G, CPC37627, cranidium, dorsal view; H, CPC37628, cranidium, dorsal view; I, CPC37632, cra- 
nidium, dorsal view; J, CPC37635, cranidium, dorsal view; K, CPC37629, cranidium, dorsal view. 


Proc. Linn. Soc. N.S.W., 126, 2005 87 


REVISION OF TRILOBITE GENERA D/ISCOMESITES AND ESTAINGIA 


Figure 4. Pagetides (Discomesites) fragum Opik, 1975b. A, CPC13179, latex cast of pygid- 


ium, dorsal view, stereo pair, scale bar = 1 mm; B, CPC37631, pygidium, dorsal view, scale 
bar = 1 mm; C-D, CPC13182, pygidium, dorsal and posterior views, scale bars = 1 mm; E-F, 
CPC13183, pygidium, dorsal and posterior views, scale bars = 0.5 mm; G-H, CPC37624, py- 
gidium, dorsal and oblique posterolateral views, scale bars = 0.5 mm; I, CPC37630, pygidium, 
dorsal view, scale bar = 1 mm; J, CPC37626, partial pygidium, dorsal view, scale bar = 1 mm. 


based on occipital spine length appears to have 
been exaggerated by Opik (1975b). He described 
the occipital spine of D. /unatulus as ‘slender and 
relatively long’. However, specimens of Discomesites 
that have a preserved occipital spine, including the 
holotype of D. fragum, show no significant difference 
in size and shape to that of the holotype of D. 
lunatulus. Therefore occipital spine length does not 
appear to be a reliable diagnostic character. 

Pygidial characteristics used by Opik (1975b) 
to distinguish species of Discomesites can be attributed 
to preservation or misinterpretation. Opik (1975b:32) 
differentiated D. fragum from D. lunatulus by its 
‘relatively broad pygidium with six axial annulations’, 
and stated that the latter species possesses seven axial 
rings. Firstly, tectonic deformation of fossils from the 
Cymbric Vale Formation is relatively common; see 
for example, specimens illustrated by Opik (1975b, 
Pl. 1, Figs 4-5; Pl. 2, Fig. 1). The larger pygidium of 
D. lunatulus illustrated by Opik (1975b, Pl. 6, Fig. 
3) (see Fig. 4C-D for dorsal and posterior views) 
has clearly been laterally compressed, thus appears 
to be narrower (tr.). This was, in fact, noticed by 
Opik (1975b:35) in his comments on the illustrated 


88 


specimens. Opik illustrated only two pygidia of D. 
lunatulus, of which only one (CPC13183) has a 
complete axis preserved. Examination of pygidium 
CPC13183 reveals the presence of only six distinct 
axial rings and a terminal piece. It is likely that Opik 
misinterpreted the change in slope at the base of the 
axial node on the terminal piece as an axial furrow. 
Therefore, based on the evidence above, D. /unatulus 
is herein considered a junior subjective synonym of 
D. fragum. 

Palmer (in Palmer and Rowell 1995:7) 
distinguished the Antarctic species Discomesites 
spinosus from Australian species of Discomesites 
based on the ‘presence of axial spines on the thoracic 
and pygidial segments and of distinct nodes on the 
pygidial margin’. Examination of Opik’s (1975b) 
type material of Discomesites, in addition to new 
topotype material from the Cymbric Vale Formation, 
reveals that smaller pygidia of Discomesites (<2.3 
mm sagittal length) have axial rings bearing short, 
median axial nodes or spines (Fig. 4E-H, J). Larger 
pygidia have less conspicuous axial nodes; in some 
cases the nodes appear to be absent (Fig. 4A-D, 1), 
implying that axial nodes vary ontogenetically. The 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. PATERSON 


pygidia of Discomesites spinosus illustrated by 
Palmer and Rowell (1995, Fig. 7.2-7.3) conform to 
this ontogenetic pattern, being less than 2.5 mm in 
length (sag.). 

The presence of marginal nodes opposite 
the pleural furrows on the pygidium of Discomesites 
spinosus appears to be a rather dubious diagnostic 
character. Marginal nodes are not clearly delineated 
on the pygidia illustrated by Palmer and Rowell (1995, 
Fig. 7.2-7.3). Apygidium of D. /unatulus (CPC13183) 
and an associated unnumbered pygidium display what 
appear to be faint marginal nodes opposite the pleural 
furrows. However, this character does not seem to 
be consistent in all pygidia of Discomesites from the 
Cymbric Vale Formation. 


Palmer (in Palmer and Rowell 1995:7) noted . 


that the Australian species of Discomesites have ‘a 
slight posterior deflection of the inner margin of the 
[anterior cranidial] border on the axial line’. This is 
certainly true for the majority of Discomesites cranidia 
from the Cymbric Vale Formation, although there is 
a great deal of variation, and some cranidia do not 
display this deflection at all (Fig. 3E, I, J). Therefore, 
D. spinosus is herein considered a junior subjective 
synonym of D. fragum. 


Order REDLICHIIDA Richter, 1932 
Suborder REDLICHIINA Richter, 1932 
Superfamily ELLIPSOCEPHALOIDEA Matthew, 
1887 
Family ESTAINGIIDAE Opik, 1975a 
Genus ESTAINGIA Pocock, 1964 


Type species 
Estaingia bilobata Pocock, 1964, Early Cambrian, 
Emu Bay Shale, Kangaroo Island, South Australia. 


Discussion 

Jell (in Bengtson et al. 1990:310) originally 
regarded Estaingia and Zhuxiella as junior subjective 
synonyms of Hsuaspis. This synonymy was supported 
in subsequent studies by Palmer and Rowell (1995), 
Nedin (1995), Jago et al. (1997) and Nedin and 
Jenkins (1999). Jell (an Bengtson et al. 1990) also 
suggested that Strenax may be regarded as a junior 
subjective synonym of Pseudichangia. Jago et al. 
(1997) supported Jell in synonymising Strenax with 
Pseudichangia, but considered both genera to be 
junior synonyms of Hsuaspis. Recently, Jell (in Jell 
and Adrain 2003:334) discovered a nomenclatural 
error between the synonymous genera Estaingia 
and Hsuaspis. Jell noted that Estaingia should be 
considered the senior name because the publications 


Proc. Linn. Soc. N.S.W., 126, 2005 


in which the name Hswaspis was first mentioned 
(Zhang et al. 1957; Zhang 1957) did not satisfy the 
ICZN criteria for availability, thus Hsuaspis must 
be considered nomen nudum in both publications. 
Therefore Hsuaspis became available in Lu et al. 
(1965), a year after its synonym Estaingia was erected 
by Pocock (1964). 


Estaingia cerastes (Opik, 1975b) 
Fig. SA-K 


Strenax cerastes Opik, 1975b:14, Pl. 2, Figs 1-6. 

Strenax (Sematiscus) fletcheri Opik, 1975b:16, Pl. 3, 
Figs 1-2. 

Estaingia bilobata Pocock, Opik 1975b:11, Pl. 1, 
Figs 1-7. 

Bergeroniellus sp., Rowell et al. 1989, p. 14, Fig. A. 

Hsuaspis cf. H. bilobata Pocock, Palmer in Palmer 
and Rowell 1995:16, Fig. 12. 

Hsuaspis cerastes (Opik), Jago et al. 1997:71, Fig. 
2B-L, Fig. 3. 


Material 

11 cranidia, 3 librigenae, 5 pygidia; CPC13152- 
13158 [Opik’s (1975b) illustrated material of 
Estaingia bilobata|, CPC13159-13163 [Type material 
of Strenax cerastes|, CPC13164 [Holotype cranidium 
of Strenax (Sematiscus) fletcheri| and CPC37636- 
37641 [New material from Site A of Opik (1975b)]. 


Discussion 

The revision of Estaingia cerastes from 
the Cymbric Vale Formation has been previously 
documented by Jago et al. (1997) and will only be 
briefly discussed here. Jago et al. (1997) noted 
that specimens of E£. bilobata illustrated by Opik 
(1975b) do not belong to this species because Opik’s 
Specimens possess a longer (sag.) glabella (relative 
to cranidial length) and show a marked forward 
expansion of the glabella, whereas the type cranidia 
of E. bilobata illustrated by Pocock (1964) display a 
shorter (sag.) glabella that either tapers anteriorly or 
has a slight waist. Nedin and Jenkins (1999, Fig. 4) 
have documented the difference in glabella length (or 
preglabellar field) between specimens of E. bilobata 
and E. cerastes. Nedin and Jenkins (1999) have also 
demonstrated that EF. bilobata and E. cerastes can 
be differentiated based on cranidial length/width 
ratios. Jago et al. (1997) also suggested that Opik’s 
specimens of E. bilobata and Strenax cerastes are 
likely to represent the same species based on the 
variation displayed in Opik’s illustrated specimens 
and those illustrated by Jago et al. (1997). Therefore, 
in placing Strenax in synonymy with Hsuaspis, Jago 


89 


REVISION OF TRILOBITE GENERA D/SCOMESITES AND ESTAINGIA 


Figure 5. Estaingia cerastes (Opik, 1975b). A, CPC13159, latex cast of holotype cranidium of Stre- 
nax cerastes, dorsal view, scale bar = 5 mm; B, CPC13152, cranidium, dorsal view, scale bar = 5 
mm; C, CPC13156, cranidium, dorsal view, scale bar = 5 mm; D, CPC13157, pygidium, dorsal 
view, scale bar = 2.5 mm; E, CPC13158, latex cast of pygidium, dorsal view, scale bar = 2.5 mm; F, 
CPC37638, pygidium, dorsal view, scale bar = 2.5 mm; G, CPC13153, cranidium, dorsal view, scale 
bar = 5 mm; H, CPC37637, cranidium, dorsal view, scale bar = 2.5 mm; I, CPC37636, cranidium, 
dorsal view, scale bar = 2.5 mm; J, CPC13164, latex cast of holotype cranidium of Strenax (Sematis- 
cus) fletcheri, dorsal view, scale bar = 1 mm; K, CPC37641, librigena, dorsal view, scale bar = 2.5 mm. 


et al. (1997) reassigned E. bilobata and S. cerastes 
described by Opik (1975b) from the Cymbric Vale 
Formation to a single species, Hsuaspis cerastes, 
reassigned herein to Estaingia cerastes (Opik). Jago 
et al. (1997:71) also noted that the holotype cranidium 
of Strenax (Sematiscus) fletcheri Opik, 1975b ‘is 
clearly an immature specimen which should not be 
the basis of a new taxon’. The holotype of Strenax 
(Sematiscus) fletcheri is considered herein a juvenile 
specimen (sagittal length: 3.0 mm) and junior 
subjective synonym of Estaingia cerastes. 

Jago et al. (1997) suggested that specimens 
of Hsuaspis cf. H. bilobata described by Palmer 
(in Palmer and Rowell 1995) from the Shackleton 
Limestone in the Transantarctic Mountains may 
belong to Hsuaspis (=Estaingia) occipitospina, 
originally described by Jell (in Bengtson et al. 1990). 


90 


Jago et al. (1997:71) based this interpretation on the 
fact that the Antarctic specimens ‘have a glabella of 
similar length to H. occipitospina as well as a similar 
preglabellar median ridge [= plectrum]’. The Antarctic 
specimens do appear to have a shorter (sag.) glabella, 
similar to E. bilobata and E. occipitospina, compared 
to that of E. cerastes. However, it is important to note 
that the Antarctic cranidia are considerably smaller 
(sagittal length: 3.8-5.6 mm) than the large holaspid 
cranidia of E. cerastes (sagittal length: 11-18 mm) 
illustrated by Opik (1975b) and Jago et al. (1997). 
Smaller cranidia of E. cerastes (Opik 1975b, Pl. 3, 
Figs 1-2; Jago et al. 1997, Fig. 2K) of similar size to 
the Antarctic cranidia show similar glabella lengths. 
Therefore the glabella of E. cerastes appears to 
become longer (sag.) relative to the cranidial length 
during ontogeny; a trend also observed by Jago et al. 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. PATERSON 


(1997:73). This trend is reversed in E. occipitospina 
in that the glabella becomes shorter (sag.) relative 
to cranidial length during ontogeny (Bengtson et 
al. 1990, Fig. 200). Furthermore, the presence of a 
plectrum does not appear to be a defining character 
amongst species of Estaingia, since all species possess 
this feature, including E. cerastes (Opik 1975b, Pl. 1, 
Figo bler2 hon. Wels. ries 2lapo ct alal997, 
Figs 2D, E, G, 3B). Jago et al. (1997) also observed 
that the Antarctic specimens possess an occipital 
node rather than an occipital spine; however, Jago et 
al. (1997:71) demonstrated that the occipital spine in 
E. cerastes varies considerably, with some specimens 
having a small node, while others possess a long, 
slender spine. 


Estaingia occipitospina can be further . 


differentiated from E. cerastes in having a longer 
(sag.) anterior cranidial border that tapers laterally 
and the palpebral lobe and eye ridge can be clearly 
distinguished, with the eye ridge being considerably 
narrower and of low relief in relation to the palpebral 
lobe. Specimens of E. cerastes and H. cf. H. bilobata 
have an anterior cranidial border of approximately 
equal length (sag., exsag.), and display a palpebral 
lobe that is continuous with the eye ridge. 

Further evidence to suggest that Estaingia 
cerastes and Hsuaspis cf. H. bilobata are synonymous 
can be seen in the morphology of the glabella and 
pygidium. Although the anterior part of the glabella 
in the Antarctic specimens is not greatly expanded, 
the glabella does show a slight expansion anteriorly. 
This can also be observed in small cranidia, of similar 
size to the Antarctic specimens, of E. cerastes (Opik 
1975b, Pl. 3, Figs 1-2; Jago et al. 1997, Fig. 2K). Based 
on the description and illustration of the pygidium of 
H. cf. H. bilobata given by Palmer (in Palmer and 
Rowell 1995:16, Fig. 12.4), there seems to be no 
apparent difference in the pygidia of E. cerastes (Fig. 
5D-F; Opik 1975b, Pl. 1, Figs 6-7). This pygidial 
similarity was in fact observed by Palmer (in Palmer 
and Rowell 1995:16), noting that ‘all of the pygidial 
characteristics [of H. cf. H. bilobata] are shared with 
specimens from the Cymbric Vale fauna of New 
South Wales, Australia, assigned by Opik (1975b) to 
Estaingia bilobata |= Estaingia cerastes]’. 


ACKNOWLEDGEMENTS 


Thanks to John Laurie (Geoscience Australia) 
for the loan of specimens housed in the Commonwealth 
Palaeontological Collection (CPC) and for assistance with 
locating extra material of Discomesites and Estaingia. 
David Whitford (CSIRO) kindly allowed access to a 


Proc. Linn. Soc. N.S.W., 126, 2005 


copy of Bo Zhou’s PhD thesis. Glenn Brock (Macquarie 
University) gave valuable advice on an earlier draft of this 
paper. Thanks to the two anonymous referees for helpful 
suggestions. Funding for this study came from a Macquarie 
University Postgraduate Research Fund (PGRF) grant. 


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93 


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First Record of an Australian Fur Seal (Arctocephalus pusillus 
doriferus) Feeding on a Wobbegong Shark (Orectolobus ornatus) 


SIMON ALLEN AND CHARLIE HUVENEERS 


Graduate School of the Environment, Macquarie University, N.S.W. 2109, Australia 


Allen, S. and Huveneers, C. (2005). First record of an Australian fur seal (Arctocephalus pusillus 
doriferus) feeding on a wobbegong shark (Orectolobus ornatus). Proceedings of the Linnean Society of 
New South Wales 126, 95-97. 


The Australian fur seal (Arctocephalus pusillus doriferus) is listed as a ‘vulnerable’ species in New South 
Wales (NSW) under the Threatened Species Conservation Act, recovering from exploitation by commercial 
sealing activities around southeastern Australia, Recent dietary studies indicate they are generalist predators 
that feed on a wide variety of both vertebrates (fish and, occasionally, birds) and invertebrates (cephalopods 
and, occasionally, crustaceans). While a small number of elasmobranchs have been reported from the diets 
of a variety of fur seal species, no published evidence exists of either fur seals preying on wobbegongs 
(Orectolobus spp.), or of large wobbegongs as prey items in the diet of any predator. Here we describe an 
account of an Australian fur seal feeding on a large ornate wobbegong (Orectolobus ornatus). Wobbegongs 
are also listed as ‘vulnerable’ in NSW by the IUCN, with commercial fishing catch having dropped over 
50% from 1990-2000. Knowledge of relationships between high trophic level species is important for 
assessing interactions between marine mammals and fisheries and also presents interesting challenges for 


the conservation of commercially targeted species. 


Manuscript received 20 July 2004, accepted for publication 22 September 2004. 


KEYWORDS: Arctocephalus, Australian fur seal, diet, Orectolobus, threatened, wobbegong. 


INTRODUCTION 


The diets of Australian and New Zealand fur seals 
(Arctocephalus pusillus doriferus and A. forsteri, 
respectively) have been extensively studied around 
southeastern Australia and New Zealand in the last 
decade (e.g. Gales & Pemberton 1994; Fea et al. 1999; 
Littnan 2004). Diagnostic techniques have primarily 
involved faecal and regurgitate sampling, while 
more recent work has also included stable isotope 
and fatty acid analyses (Littnan 2004). These studies 
have indicated that fur seals target a large number 
of prey species, with a relatively limited number of 
cephalopods and fish species constituting the majority 
of their diet. There is evidence of some seasonal and 
spatial variation in Australian fur seal diet (Hume et 
al. 2004; Littnan 2004) and seasonal variation in New 
Zealand fur seal diet (Fea et al. 1999). A very small 
portion of fur seal diet is made up of crustaceans, birds 
and some small elasmobranchs (Gales & Pemberton 
1994; Fea et al. 1999; Hume et al. 2004). Here we 
describe the first account of an Australian fur seal 
feeding on a large ornate wobbegong (Orectolobus 
ornatus). 


INTERACTION ACCOUNT 


During a coastal survey of small cetaceans 
from Port Stephens to Sydney on December 28" 
2003, an Australian fur seal (distinguished from the 
sympatric New Zealand fur seal by facial profile and 
fur colouration) was witnessed carrying the body of 
a large ornate wobbegong (distinguished from the 
sympatric spotted wobbegong O. maculatus by skin 
pattern and colouration). The interaction occurred 
approximately 3.2 nautical miles north of Norah Head 
lighthouse on the central coast of New South Wales 
(33°13.3’S, 151°35.2’E). Excellent conditions (Sea 
State 1, no cloud cover, clear water and being able 
to approach to within 5m of the animals) facilitated 
reliable identification of both species, with video 
footage of the event used to confirm identification 
and behaviour after the voyage. The shark’s head 
had been removed and the fur seal was thrashing 
the body from side to side in an apparent attempt to 
separate manageable portions of the shark’s flesh. 
This behaviour is common for pinnipeds feeding on 
prey too large to swallow (Rand 1959; Reeves et al. 
IQ22), 


FUR SEAL FEEDING ON WOBBEGONG SHARK 


Female Australian fur seals grow to a maximum 
length of around 1.5m, while males can reach 2.0- 
2.25m (Warneke and Shaughnessy 1985). The fur seal 
was estimated to be approximately 1.5m in length and 
the presence of a light mane suggested it was a sub- 
adult male. The ornate wobbegong becomes sexually 
mature at around 1.8m in length and grows to 2.9m 
(Last and Stevens 1994). The wobbegong’s total body 
length was estimated to be around 1.4m (sex was not 
determined). 

Only post-capture manipulation was witnessed, 
with no predation event observed, so we cannot 
discount the possibility that the shark was found dead 
or was scavenged from a fishing line by the fur seal. 
Wobbegongs are, however, commercially targeted 
using set-lines in NSW; 89% are gut-hooked, 100% 
remain alive until retrieved and killed by fishermen, 
and no wobbegong fisherman in NSW have witnessed 
line depredation by fur seals (C. Huveneers unpub. 
data). It is unlikely that a carcass would be discarded 
by a fisherman or that the shark could have been 
removed from the hook by the fur seal without tearing 
the shark’s body cavity. Predation thus seems to be the 
most plausible explanation for the above observation 
of an Australian fur seal carrying the body of an 
ornate wobbegong. 


ELASMOBRANCHS IN FUR SEAL DIET 


The remains of two spiny dogfish (Squalus 
acanthias) were found in 357 faecal and regurgitate 
samples of Australian fur seals hauling out around 
Tasmania (Gales and Pemberton 1994), while a 
more recent study of the same colonies found no 
elasmobranch remains in 1044 samples (Hume et 
al. 2004). Similarly, no sharks or rays were found in 
the diet of Australian fur seals around Kanowna or 
the Skerries, Victoria (n=1008; Littnan 2004). The 
remains of one dogfish were found in 584 faeces 
and regurgitates from New Zealand fur seals at the 
Otago Peninsula (Fea et al. 1999) and elasmobranchs 
including the puffadder shyshark (Haploblepharus 
edwardsii) have been recorded in the diet of the Cape 
fur seal (A. p. pusillus) off South Africa (Rand 1959; 
Martin 2004). Adult wobbegongs might be considered 
potential prey items for numerous _pinnipeds, 
cetaceans and large shark species, but there has been 
no published account to date. 


96 


CONCLUSION 


Pinnipeds and elasmobranchs are high-level 
predators that occupy important niches in marine 
ecosystems (e.g. Cortes and Gruber 1990; Read and 
Brownstein 2003). Interactions between them can have 
both direct and indirect effects on marine mammals, 
fish and invertebrates at lower trophic levels. 
Quantifying the diet of high trophic level species 
is therefore important for modelling of interactions 
between marine mammals and fisheries and assessing 
the effects of stock depletion by commercial fishing 
(see Goldsworthy et al. 2003; Myers and Worm 2003; 
Hutching and Reynolds 2004; Littnan 2004). It also 
presents challenges for conservation and fisheries 
management when predator/prey relationships involve 
more than one threatened or vulnerable species. This 
note represents the first record of a large wobbegong 
being fed upon by a fur seal. 


ACKNOWLEDGEMENTS 


We would like to thank Owen and Linda Griffith of 
Advance II charters. Luciana Méller and Kerstin Bilgmann 
of the Graduate School of the Environment provided video 
footage of the interaction. Comments by Charles Littnan 
and an anonymous reviewer greatly improved the clarity 
of this note. 


REFERENCES 


Cortes, E. and Gruber, S.H. (1990). Diet, feeding habits 
and estimates of daily ration of young lemon sharks, 
Negaprion brevirostris. Copeia 1, 204-218. 

Fea, N. I., Harcourt, R. and Lalas, C. (1999). Seasonal 
variation in the diet of New Zealand fur seals 
(Arctocephalus forsteri) at Otago Peninsula, New 
Zealand. Wildlife Research 26, 147-160. 

Gales, R. and Pemberton, D. (1994). Diet of the Australian 
fur seal in Tasmania. Australian Journal of Marine 
and Freshwater Research 45, 663-664. 

Goldsworthy, S., Bulman, C., He, X., Larcome, J. and 
Littnan, C. (2003). Trophic interactions between 
marine mammals and Australian fisheries: An 
ecosystem approach. In ‘Marine mammals: Fisheries, 
tourism and management issues’ (Eds N. Gales, 

M. Hindell & R. Kirkwood) pp. 62-99. (CSIRO 
Publishing: Collingwood) 

Hume, F., Hindell, M., Pemberton, D. and Gales, R. 

(2004). Spatial and temporal variation in the diet of 


Proc. Linn. Soc. N.S.W., 126, 2005 


S. ALLEN AND C. HUVENEERS 


a high trophic level predator, the Australian fur seal 
(Arctocephalus pusillus doriferus). Marine Biology 
144, 407-415. 

Hutchings, J.A. & Reynolds, J.D. (2004). Marine fish 
population collapses: Consequences for recovery and 
extinction risk. BioScience 54(4), 297-309. 

Last, P.R. and Stevens, J.D. (1994). ‘Sharks and Rays of 
Australia’. (CSIRO Australia). 

Littnan, C.L. (2004). Foraging ecology of the Australian 
fur seal (Arctocephalus pusillus doriferus). PhD 
thesis, Macquarie University, Sydney. 

Martin, R. A. (2004). Natural mortality of the puffadder 
shyshark (Haploblepharus edwardsii) due to 
two species of marine tetrapod, the Cape fur seal 
(Arctocephalus pusillus pusillus) and black-backed 
kelp gull (Larus dominicanus vetula) at Seal Island, 
False Bay, South Africa. Journal of Fish Biology 64, 
1-6. 

Myers, R.A. & Worm, B. (2003). Rapid worldwide 
depletion of predatory fish communities. Nature 423, 
280-283. 

Rand, R.W. (1959). The Cape fur seal (Arctocephalus 
pusillus): distribution, abundance and feeding habits 
off the south western coast of the Cape Province. 
Investigational report, Division of Fisheries South 
Africa 34, 1-75. 

Read, A. J. and Brownstein, C.R. (2003). Considering 
other consumers: fisheries, predators and Atlantic 
herring in the Gulf of Maine. Conservation Ecology 
7(4), 2. 

Reeves, R.R., Stewart, B.S. and Leatherwood, S. (1992). 
“Seals and Sirenians’. (Sierra Club Bookstore: San 
Francisco). 

Warneke, R.M. and Shaughnessy, P.D. (1985). 
Arctocephalus pusillus, the South African and 
Australian fur seal: taxonomy, biogeography and 
life history. In “Studies of Sea Mammals in South 
Latitudes’ (Eds. J.K. Ling and M.M. Bryden) pp 53- 
77. (South Australian Museum: Adelaide) 


Proc. Linn. Soc. N.S.W., 126, 2005 


97 


ae hear’. 


> 4 Se 
(Po) QASSMEMHON RREOONA SHARK 
Ae 


r i 

vale Assetalior d. gee Bo merle ‘. lecag w/t radlergenr A pee Ee ba 
4 erourl s Gin vench 20. “Cohen Sanwa Caredivas nite, 
Grigio, 1098), Thee fir etal “Pp iweus Cea hind Af 

nin Senyth and eae: At) 


art 


4 Lit wau‘s xu 
1 t 7 ’ : ; afl : e “ 
enw a gyi icy ‘72 oe etl live itn =acl Te ines Pirie erie seers 
) The eohbegome F tondl bod omer ate igaey tM 
 t.s dt “en (ora wag oot © | centages yp Bie 
ure. Tioitpiletion wee witheased an a tc i 
— ' served, So “~we cannot the. oflertl oat yen fen 2M 
ae eg ee oe me pasry PRY oe 
mame ine by (he fur veal Hig 

eu aligg atl Pics? anton ti" 

as, however, eniumencaaly reeled POG Ae meee 
it We 89% pre wul Kou, TIRE ane semen when prodiaian’ pm 
fil ceinicved and billed y Maneripen,, 4 hee sg na 
yawobbepoany Ashodndn in. NSW fate witreseed sin araane IT oP. : 

im ty Tat seal ‘Cc Faris wore yunools bare ; 
1 Wnikes hat a < ca wih Ue | FUN 
by nce. thar dhwrk could 1 { 
i ‘ wiikeon : 4 Pal 
ie I : i > “ x . 3 7 cate se Aino? Reb 
st plaumble explanation for the-above observation amid iigetndl 


“any AisICaliany FUP: Seal CreTving thst Rasy. OF ay Ashi "intel A titi inka as > viaeoe 
‘ houaer of the interaitacn, Ser i coined 4 

ge pepper ge 
ELASMOBRANCHS IN FUR-SEAL DIET ~~ "hea 9 ea 


ictiatish: lak Rae este accaaaly) Kedeaniag 1 (2hO1) As ws 
| | | bra amcatitA ds 


were kn +3 cul } Pour Hal¢c ~o dann: : mag's 55 
: : st Aa ji ity Tu HA ij ny > Ti? i 
' wa. 4 i} yf) [ heriow ( | vinLe «@& 
we ToCenl study. of the Sime cealonicsy ft Pe 
nobranch Tem ies saropic Hire 
OA). Sinai no vh wks of rave “Wére lownd i 


re dict of AvAtraltan Tit scale around Ketan or 
toy Skerrigs, Vedrorea Tye (O08: Linen 2 Teo 
Teinoins G) one Gogh Were Tain) ay “RS Rees... ' 
ane revurgilates fro New Zealand tut attest Oe as: 
>Chago fobinsula{Fea ct al. 7099) and | penned erneix: / 
me Neding -the puftadder shyéhurk (7 figeleihearr 
dwundsi!) have bows recorded iy the aie ghee. 
furseal (4. p. pasius) oi) South Atviee (Rigel 1949. . bepes: 
Martin 2604). Aduli wobbegongs tight he Gaaniniteed ‘dae — ‘Mar 
polemtial ptey “hate fe ae, - ‘ella: aerials 
cetaceans (ed Tange Stark vag hs 
hi pubitisled meant te ile. 


Effects of Javan Rusa Deer (Cervus timorensis) on Native Plant 


Species in the Jibbon-Bundeena Area, Royal National Park, 
New South Wales 


Davip Kerry! AND BELINDA PELLOW? 


'NSW Department of Environment and Conservation, PO Box 1967, Hurstville 2220 NSW. 
*Janet Cosh Herbarium, University of Wollongong, Wollongong 2522 NSW. 


Keith, D. and Pellow, B. (2005). Effects of Javan rusa deer (Cervus timorensis) on native plant species 
in the Jibbon-Bundeena area, Royal National Park, New South Wales. Proceedings of the Linnean 
Society of New South Wales 126, 99-110. 


A reconnaissance survey and exclosure experiment were carried out to examine the effects of Javan rusa 
deer on native flora and vegetation in Royal National Park on the southern outskirts of Sydney, Australia. Of 
78 native plant species examined during the survey, only nine showed no evidence of vertebrate herbivory 
or physical damage and the majority of these plants were ferns and sedges. The other 69 species showed 
effects that included defoliation (young and/or old leaves), removal of shoots, bark-stripping, stem breakages 
and destruction or consumption of reproductive material. These effects varied in severity between species 
and from place to place, and were inferred to have been caused by deer based on the local abundance of deer 
droppings, footprints and the scarcity of other vertebrate herbivores in the area. The survey also revealed 
localised soil erosion associated with high densities of deer footprints and droppings. An unreplicated 
exclosure experiment showed that planted saplings of Syzygium paniculatum, a threatened rainforest tree, 
suffered major defoliation, bark stripping, stem breakages and some mortality when exposed to deer for 
several months. Many of the surviving plants showed signs of recovery when deer were subsequently 
excluded, although full recovery of their leaf canopies could take several seasons. The observed effects on 
vegetation and individual plant species are consistent with studies on several other deer species in a range 
of ecosystems overseas. A model of the effects of deer herbivory based on plant life-history suggests that 
curtailment of seed production and seedling recruitment are likely to be the major impacts of deer on plant 
population viability. Reductions in net growth and survival of established plants and possibly post-dispersal 
predation of seeds are less likely to be significant influences. 


Manuscript received 31 October 2003, accepted for publication 18 August 2004. 
KEYWORDS: browsing, Cervus timorensis, deer, endangered ecological communities, feral animals, 


fire, grazing, herbivory, plant demography, Syzygium paniculatum, threatened species, threatening 
process. 


INTRODUCTION 


Javan rusa deer (Cervus timorensis) is one of six 
deer species that have established wild populations 
in Australia (Moriarty 2004a). Rusa deer were 
introduced into Royal National Park in 1906 by the 
Park Trust for exhibition purposes (NSW National 
Parks and Wildlife Service 2002). The seven 
introduced animals were initially kept within a fenced 
enclosure at ‘Deer Park’ on the Hacking River near 
Warumbul. The deer soon escaped and established 
a wild population that has persisted in the Park and 
adjacent areas to the present day. Hamilton (1981) 
suggested that rusa deer essentially replaced fallow 


deer (Dama dama), which were introduced some 20 
years earlier and likely to be the species reported as 
widespread in the Park in 1914 (Anon. 1914). High 
densities of Rusa deer are now regularly observed 
in the vicinity of Bundeena, Grays Point, Garie-Era 
and various sites along the Hacking River valley. 
The deer population is likely to have fluctuated since 
its introduction, although densities appear to have 
increased markedly since bushfires in January 1994, 
which burnt more than 90% of the Royal National 
Park (pers. obs.). After the fires, the population within 
Royal National Park was estimated to include less 
than 500 individuals. Quantitative surveys carried out 
between 1999 and 2001 indicate that the population 


RUSA DEER IN ROYAL NATIONAL PARK, SYDNEY 


increased from 2500 to 2900 individuals during that 
time (Moriaty unpubl. data, NSW National Parks and 
Wildlife Service 2002). 

Rusa deer grow to | m tall at the shoulder and 
weigh 100-160 kg. Males develop large antlers but 
females do not. The deer are active nocturnally, 
resting in dense native vegetation by day. They are 
dietary opportunists, apparently preferring grass but 
browsing opportunistically on the buds, shoots and 
leaves of woody plants and herbs (Bentley 1979). 
Recent dietary data from Royal National Park indicate 
that food sources vary with season and location. In 
native vegetation remote from settled areas, 80% of 
their average summer diet consists of native browse, 
with grasses making up the remaining 20% (Moriaty 
2004b). In other seasons, the grass component 
becomes negligible in these areas. However, the grass 
component varies seasonally from 40% to 70% in 
animals living close to cleared grassy areas (Moriarty 
2004b). 

Concerns about impacts on native vegetation 
have resulted in a recent preliminary determination 
of feral deer as a Key Threatening Process under the 
NSW Threatened Species Conservation Act 1995. A 
recent survey of rangers employed by Rural Lands 
Protection Boards in NSW identified deer as the 
most important emerging pest animal threat (West 
and Saunders 2003). More than 40% of respondents 
indicated that there had been a moderate to high 
increase in the distribution and abundance of wild 
deer in their area. In another recent survey more than 
80% of land managers reported browsing of native 
plants and agricultural crops as an impact of feral 
deer (Moriarty 2004a). Despite numerous detailed 
studies on the impacts of deer on native vegetation 
in other countries (e.g. Okuda and Nakane 1990, 
Veblen et al. 1992, Kay 1993, Jane 1994, Anderson 
and Katz 1993, Mladenoff and Stearns 1993, Khan 
et al. 1994, Augustine and Frelich 1998, Akasi and 
Nakashisuka 1999, McShea and Rappole 2000, Fuller 
and Gill 2001, Coomes et al. 2003, Rooney et al. 
2004), there are relatively few published data from 
Australia on this subject. In Royal National Park and 
adjacent urban areas, the high densities of deer, their 
free movement in open areas and their exclusion from 
some properties provide opportunities to examine the 
effects of deer on native vegetation. The aims of this 
study were to document the various impacts of deer 
on native vegetation, compile a list of native plant 
species affected by deer and to quantify the effects 
of deer on a threatened plant species, Syzygium 
paniculatum. 


100 


METHODS 


Study area 

Jibbon-Bundeena is a 300 ha area located in the 
far north-east corner of Royal National Park on the 
shores of Port Hacking on the southern outskirts of 
Sydney (lat. 34° 06’S, long. 151° 09’E). The bedrock 
of Hawkesbury sandstone is overlain in some parts by 
podsolised marine and aeolian sand dunes that may 
exceed 10 m in depth. The dune crests and slopes are 
freely drained, although the swales and flats may be 
periodically waterlogged. The exposed sandstones 
carry shallow to skeletal yellow earths with variable 
drainage characteristics. 

The area includes a mosaic of rainforests, eucalypt 
forests, heathlands and wetlands. Littoral rainforest on 
the hind dunes of Jibbon and Bonnie Vale beaches is 
dominated by Cupaniosis anacardioides with Acmena 
smithii, Glochidion ferdinandi subsp. ferdinandi and 
Banksia integrifolia subsp. integrifolia. \t forms part 
of an Endangered Ecological Community (Littoral 
Rainforest in NSW) under the NSW Threatened 
Species Conservation Act 1995 and includes Syzygium 
paniculatum, which is currently listed as a Vulnerable 
species. The eucalypt forests are found principally on 
deep sands. On dune slopes they are dominated by 
Corymbia gummifera and Angophora costata with 
an understorey of sclerophyllous shrubs and bracken. 
On sandy flats, Eucalyptus botryoides and A. costata 
form a taller forest with an understorey that includes 
a mixture of mesophyllous and_ sclerophyllous 
shrubs and herbaceous plants. Heathlands occur on 
deep sands and on the exposed sandstone plateau. 
Floristic composition varies between these substrates, 
although both communities comprise a dense to 
open cover of sclerophyll shrubs interspersed with 
sclerophyll sedges. Wetlands are restricted to lagoons 
in dune swales and swampy sand flats. They are 
mainly herbaceous communities dominated by 
Baumea juncea with B. articulata, Leptocarpus 
tenax and other sedges. They are currently listed 
as an Endangered Ecological Community (Sydney 
Freshwater Wetlands) under the NSW Threatened 
Species Conservation Act 1995. 

The township of Bundeena occupies about 100 
ha within the study area, covering sand dunes, sand 
flats and sandstone ridges. The majority of the town 
precinct comprises suburban dwellings on blocks 
<0.1 ha, some of which are fenced to exclude deer 
from gardens. There are substantial areas planted 
with exotic grass on road verges and in parks and 
yards. Native plants, relics of the original vegetation, 
persist as solitary individuals or in clumps throughout 
the town. At least some of the deer in the area spend © 


Proc. Linn. Soc. N.S.W., 126, 2005 


D. KEITH AND B. PELLOW 


some of their nocturnal foraging time within the 
suburban limits of Bundeena. They graze on grassy 
areas and are fed vegetable matter by a few local 
residents. They also browse on a variety of forbs and 
woody plants, including various vegetables, exotic 
ornamental plants and local native plants. 


Survey 

Extensive reconnaissance was _ undertaken 
throughout the native vegetation and the suburban 
area to observe deer behaviour and movement, record 
plant species consumed or damaged by deer and 
effects of deer on the structure of vegetation and soils. 
Observations were made opportunistically between 
1999 and 2003. Observations on native plants were 
confined to bushland areas within 1 km of Bundeena 
township, but at least 20 m beyond the suburban/ 
bushland boundary. The effects were classified 
into the following categories: young foliage and 
shoots consumed; young and old foliage and shoots 
consumed; bark stripped; woody stems broken; 
seedlings uprooted; inflorescences damaged or 
consumed; and unaffected. Qualitative observations 
on damage to vegetation structure and soils were also 
recorded during the reconnaissance of bushland. 

To examine the effects on soils where deer 
activity was concentrated, three sites were identified 
where deer were fed by local residents. All three sites 
were on deep, unconsolidated sand dune soils on the 
interface between eucalypt woodland and the eastern 
edge of the suburban area. They were separated from 
one another by distances of 200 metres, and not 
included in the reconnaissance survey, which sampled 
bushland away from the suburban interface. Soils and 
vegetation at these sites were observed between 1999 
and 2003, and qualitative descriptions of vegetation 
cover and soil level were compared to those of 
equivalent sites on the bushland/suburban interface 
approximately 50 m distant from each feeding site. 
The equivalent sites were also on sand and had similar 
vegetation (disturbed heathy woodland) and a similar 
management history to the feeding sites. 


Exclosure experiment 

Ninety-three individuals of Syzygium 
paniculatum were planted in a 0.1 ha yard, grassed 
with Pennisetum clandestinum and enclosed within 
1 m high wire fencing. The plants were 1.0 — 1.4 m 
tall and were watered for three months after planting. 
After one month, all plants were in a healthy condition 
and growing new shoots when the yard was opened, 
allowing deer access. Initially relatively few deer 
entered the yard, but after several weeks, several 
animals gained regular access during nights. A fter three 


Proc. Linn. Soc. N.S.W., 126, 2005 


months, the gate was closed. Deer gained intermittent 
access to the yard after that time until the perimeter 
fence was raised to a height of 1.5 m. As a qualitative 
control treatment, five plants were observed in an 
adjacent yard that was maintained to exclude deer 
throughout the duration of the experiment. Brush- 
tailed possums, the only other vertebrate herbivore 
on the site, were observed in both the treatment and 
control yards. Effects on Syzygium paniculatum were 
recorded six months after deer initially gained access 
as follows: >75% foliage and branchlets consumed; 
<75% foliage and branchlets consumed; main stem 
broken within 30 cm of base; bark removed; or 
foliage and branches unaffected. One year after deer 
had been excluded, plants were recorded as either 


_ dead, alive and growing new shoots or alive and not 


growing new shoots. 


RESULTS 


Vegetation structure and soils 

Reconnaissance of bushland suggested that 
structural irregularities in the vegetation, including 
tracks and open areas with low densities of woody 
plants, were associated with deer access and activity. 
Footprints indicated that a number of tracks in the 
area were used by both humans and deer. However, 
numerous tracks in the area showed no evidence of 
regular human usage and generally dissipated after 
some distance or terminated in open areas in the forest 
understorey or heathland. These open areas varied 
from 2 — 50 m’ and had conspicuously lower densities 
of shrubs and groundcover than the surrounding 
vegetation (Fig la). Resting deer were disturbed 
from some of these sites during reconnaissance and 
the presence of droppings, footprints and/or dead 
remains of deer at most sites suggested that they 
function as deer encampments. There was little 
evidence of significant surface erosion on the sand 
dunes within heathlands or woodlands, except along 
the tracks (Fig. 1b), which were depressed below the 
general soil surface. However, the wetland soils were 
exposed, compacted and deformed on the surface 
by deer footprints. These areas of bare compacted 
soil were most commonly encountered around the 
margins of wetlands. 

The three sites where deer were fed by local 
residents on the bushland/suburban interface had a 
low open cover of grasses, herbs and shrubs when 
observations began in 1999. By 2003, all three sites 
were denuded of vegetation cover and had lost up to 
0. 6 m of topsoil (Fig. 2). At two of the sites, retaining 
walls supporting built-up residential yards had been 


101 


RUSA DEER IN ROYAL NATIONAL PARK, SYDNEY 


Figure 1. Changes in the structure of vegetation and soils related to deer activity: a (top) clearing approxi- 
mately 220 m east of Bundeena within a dry sclerophyll forest from which all woody understorey plants have 
been eliminated and groundcover plants have been thinned exposing bare soil; b (bottom) an area near the 
beginning ofthecoastwalk opened up by deeractivity approximately 6monthspriorto photophraph, showing 
deer footprints on bared soil surface and lignotubers of Lambertia formosa exposed by significant soil erosion. 


102 Proc. Linn. Soc. N.S.W., 126, 2005 


D. KEITH AND B. PELLOW 


Figure 2. Site on the bushland/suburban interface at the end of Scarborough Street, Bundeena, where 
concentrated deer activity resulted in substantial loss of soil. The relic shrub with exposed root burl 
is Leucopogon ericoides. Feeding of deer ceased at this site approximately two years prior to photo- 
graph, though the animals continue to pass through the area and maintain an exposed soil surface. 


substantially undermined by erosion of the dune. The 
denuded areas had high densities of deer droppings 
and footprints, and varied in area from 400 to 1075 
m*. The roots of trees and large shrubs (Corymbia 
gummifera, Banksia integrifolia subsp. integrifolia) 
had been exposed and broken in the eroded sites. 
Substantial volumes of sand had been transported 
downslope and deposited within the adjacent area 
of native vegetation. The equivalent sites on the 
bushland/suburban interface, approximately 50 m 
from each of the feeding sites, showed lower densities 
of deer prints and dung, comparatively little evidence 
of erosion and sedimentation, and retained an open 
continuous cover of grasses, herbs and scattered 
shrubs. 


Plant survey 

Seventy native plant species from 29 families 
and two introduced species showed evidence of 
damage by deer (Table 1). Young foliage and shoots 
were preferentially browsed on most of the plant 
species recorded. Older and tougher leaves were less 
affected, although in many species there appeared to 
be little distinction between consumption of young 
and old leaves (Fig.3). The effects of browsing 
were spatially variable because some species that 


Proc. Linn. Soc. N.S.W., 126, 2005 


were heavily browsed in some areas appeared to be 
relatively unaffected in others. The removal of shoots 
was particularly frequent and conspicuous in many 
leguminous species (family Fabaceae), with some 
individuals being completely defoliated. Rainforest 
species, notably Acmena smithii, Cissus antarctica, 
Clerodendrum tomentosum, Rapanea howitteana 
and Syzygium paniculatum, also suffered high levels 
of defoliation. The plant families Epacridaceae 
and Proteaceae also had large numbers of affected 
species. 

Bark was removed or damaged on large woody 
stems of six species (Table 1). In some cases, scattered 
remains of bark and the abraided appearance of the 
stem from which bark was removed suggested that 
damage was probably caused by antler rutting. In other 
cases, particularly the rainforest species, bark had 
been torn off in strips and may have been consumed 
as food. In several monocotyledonous species, 
including orchids, Doryanthes and Xanthorrhoea, 
inflorescences had been consumed or destroyed. 
Remains of destroyed Xanthorrhoea inflorescences 
indicated that this occurred during the bud stage. 
Only nine species of plant consistently showed no 
evidence of damage by deer. Three of these were 
ferns and three were sclerophyllous sedges. 


103 


Agavaceae 
Anacardiaceae 
Anthericaceae 
Apiaceae 
Arecaceae 
Asparagaceae 
Casuarinaceae 
Casuarinaceae 


Dennstadetiaceae 
Dennstadetiaceae 


Dilleniaceae 
Elaeocarpaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Epacridaceae 
Euphorbiaceae 
Euphorbiaceae 


Iridaceae 
Lomandraceae 
Luzuriagaceae 
Moraceae 
Myrsinaceae 
Myrtaceae 
Myrtaceae 
Myrtaceae 
Myrtaceae 
Myrtaceae 


Myrtaceae 
Myrtaceae 
Myrtaceae 
Myrtaceae 


104 


Species 


Doryanthes excelsa 
Euroschinus falcata 
Thysanotus virgatus 
Platysace linearifolia 
Livistona australis 
*Asparagus densiflorus 
Allocasuarina distyla 
Casuarina glauca 
Caustis pentandra 
Lepidosperma concava 
Schoenus brevifolius 
Pteridium esculentum 
Hypolepis muelleri 
Hibbertia scandens 
Elaeocarpus reticulatus 
Astroloma pinifolia 
Brachyloma daphnoides 
Epacris longiflora 
Leucopogon ericoides 
Leucopogon parviflorus 
Monotoca elliptica 
Monotoca scoparia 
Styphelia viridis 
Phyllanthus gunnii 
Ricinocarpos pinifolius 
Acacia implexa 


Acacia longifolia 


Acacia suaveolens 

Aotus ericoides 

Bossiaea ensata 

Bossiaea heterophylla 
Dillwynia floribunda 
Kennedia rubicunda 
Phyllota phyllicoides 
*Senna pendula var. 
glabrata 

Viminaria juncea 
Patersonia glabrata 
Lomandra longifolia 
Geitonoplesium cymosum 
Ficus rubiginosa 
Rapanea howitteana 
Acmena smithii 
Angophora costata 
Corymbia gummifera 
Leptospermum laevigatum 
Leptospermum 
polygalifolium 
Leptospermum squarrosum 
Leptospermum trinervium 
Melaleuca nodosa 


Syzygium paniculatum 


Proc. Linn. Soc. N.S.W., 126, 2005 


Species 


D. KEITH AND B. PELLOW 


Young young and 


Bark |Woody|Seedlings| Inflorescences} Not 


foliage and | old foliage |stripped| stems | uprooted} damaged or 


shoots and shoots 
consumed | consumed 


consumed 


Notelea longifolia 
Caladenia caerulea” 
Cyrtostylis reniformis® 
Glossodia minor X 
Caladenia caerulea hybrids” 


Oleaceae 

Orchidaceae 
Orchidaceae 
Orchidaceae 


Pterostylis sp. ~ 
Dianella caerulea 


Orchidaceae 
Phormiaceae 
Poaceae Austrostipa pubescens 
Proteaceae 


Proteaceae 


Banksia ericifolia 

Banksia integrifolia subsp. 
integrifolia 

Banksia marginata 
Banksia oblongifolia 
Banksia serrata 
Conospermum taxifolium 


Proteaceae 
Proteaceae 
Proteaceae 
Proteaceae 


Proteaceae Hakea laevipes subsp. laevipes 


Proteaceae 
Proteaceae 
Proteaceae 
Proteaceae 
Proteaceae 
Proteaceae 


Hakea propinqua 
Isopogon anemonifolius 
Lambertia formosa 
Persoonia levis 
Petrophile pulchella 
Telopea speciosissima 
Proteaceae Xylomelum pyriforme 
Restionaceae Hypolaena fastigata 
Restionaceae Leptocarpus tenax 
Acronychia oblongifolia 


Cupaniopsis anacardioides 


Rutaceae 
Sapindaceae 
Sinopteridaceae |Pellaea falcata var. falcata 
Solanaceae 
Ulmaceae 
Verbenaceae 
Vitaceae 
Vitaceae 
Xanthorrhoeaceae| Xanthorrhoea resinifera 


Solanum stelligerum 

Celtis paniculata 
Clerodendrum tomentosum 
Cissus antarctica 

Cissus hypoglauca 


* Introduced species 
* Margaret Bradhurst, unpubl. data. 


Table 1. (opposite page and above) List of plant species affected by deer. Nomenclature follows Harden 
(1990-2002) and recent updates (www.plantnet.rbgsyd.gov.au). 


Effects on Syzygium paniculatum 

All 93 individuals of Syzygium paniculatum 
suffered some loss of foliage when deer gained 
access to the yard (Table 2). For a large majority of 
individuals, the level of defoliation was severe, with 
less than 25% of foliage remaining on the plant. 
About 15% of plants were severely damaged, having 
their bark stripped off or their main stem broken near 
ground level. However, about 90% of plants began 


Proc. Linn. Soc. N.S.W., 126, 2005 


to recover when deer were excluded, shooting new 
foliage in the next growing season. The five plants in 
the adjacent yard, where deer remained excluded, did 
not suffer any appreciable loss of foliage. Scats and 
nocturnal observations indicated that deer regularly 
gained access to the open yard, but not to the closed 
control, while common brush-tailed possums gained 
access to both yards and were the only other vertebrate 
herbivore observed at the site. 


105 


RUSA DEER IN ROYAL NATIONAL PARK, SYDNEY 


Figure 3. Defoliation caused by deer herbivory on 
shrubs of (a - upper left) Styphelia viridis subsp. 
viridis; (b- upper right) Leptospermum squarrosum; 
(c - lower left) Banksia marginata; (d - lower right) 
Persoonia levis. Various heathland sites on sand ca. 
400-500 metres from the eastern fringe of Bundeena. 


106 Proc. Linn. Soc. N.S.W., 126, 2005 


D. KEITH AND B. PELLOW 


Control 


Effect class 


plants (N=5) 
>75% foliage and branchlets consumed 0 
<75% foliage and branchlets consumed 0 
main stem broken within 30 cm of base 0 
bark removed 0 
foliage and branches unaffected 5 (100) 


Treatment 6 months after 
deer gained access 


Number (%) of Number (%) of plants 


Treatment one year after deer gained 
access and were subsequently excluded 


Alive & sprouting Alive but with Dead 


(N=93) new foliage no new foliage 

78 (84) 72 5 1 
3 (3) 3 0 0 
9 (10) 7 1 1 
3 (3) 2 0 l 
0 (0) - - - 


Table 2. Effects of deer on planted Syzygium paniculatum in the exclosure experiment. 


DISCUSSION 


Deer consumed a wide variety of plant material 
including young and old foliage, branchlets, bark 
and reproductive material of a large number of 
plant species from a broad taxonomic spectrum. 
The generalisation that deer limit regeneration 
and reproduction in a wide variety of plant species 
appears to hold for different species of deer studied 
across a broad range of ecosystems all over the 
world, including North America (McShea and 
Rappole 1999, Opperman and Merenlender 2000, 
Rooney 2001, Rooney et al. 2004), Europe (Kay 
1993, Fuller and Gill 2001, Rackham 2003) and Asia 
(Okuda and Nakane 1990, Kahn et al. 1994, Akasi 
and Nakashisuka 1999), where deer are native, and 
Australia (Moriarty 2004a), New Zealand (Jane 1994, 
Coomes et al. 2003) and South America (Veblen et al. 
1992), where they have been introduced. In both cases, 
deer populations seem to have increased recently, 
either as a result of expansion into new habitats at 
previously uninhabited locations (Forsyth et al. 2004) 
or as a result of landscape changes within their natural 
range (Mladenoff and Stearns 1993, Fuller and Gill 
2001). The large number and diversity of native plant 
species affected by deer in the Bundeena-Jibbon area 
were also consistent with previous studies in Royal 
National Park that have shown deer to be adaptable 
dietary generalists. Hamilton (1981), for example, 
showed that the proportion of food types consumed 
by deer varied with season and habitat. Deer faeces 
generally contained higher proportions of shrub and 
herb fragments in winter and/or spring, and higher 
proportions of grass fragments in other seasons. 
Faeces recovered from rainforest had the highest 
content of shrub and herb material (ca. 80%), while 
faeces from grassland/wet sclerophyll habitats had 


Proc. Linn. Soc. N.S.W., 126, 2005 


the highest content of grasses (70-90%) and faeces 
from dry sclerophyl! forest and heathland had roughly 
equal proportions of broad-leaf and grass material. 
Moriarty (2004b) recorded similar dietary variation 
from rumen analyses, with grasses comprising the 
majority of food in the vicinity of cleared areas, while 
native plants other than grasses were the major food 
source in other areas. 

Our list of plant species affected by deer is 
substantially larger than previously reported lists 
(Hamilton 1981), but most unlikely to be exhaustive. 
Uncommon species and herbaceous species, which 
may be consumed in their entirety, are likely to be 
under-recorded in reconnaissance surveys such as 
ours. There is also a risk that signs of browsing could 
have been overlooked on some plants or that evidence 
of browsing was erroneously attributed to deer. The 
latter source of errors is unlikely to be significant 
because wallaby and possum scats were rarely seen 
within the study area, whereas deer scats were very 
common. Alternative techniques entail different 
sampling errors. Analyses of faeces and rumen 
samples, for example, face difficulties of identifying 
plant fragments, high variability between samples and 
limitations that labour-intensive laboratory analysis 
impose on sample size. Nevertheless, it would be 
possible to compile a more comprehensive list of 
plants consumed by deer with increased sampling 
effort and a combined sampling approach including 
exclosure experiments and analyses of faeces and 
rumen. 

In addition to the direct effects of herbivory, deer 
had substantial, though localised, impacts on soils 
and vegetation structure. Dune soils were severely 
eroded at parts of the urban interface where feeding by 
humans lead to intensive deer activity. The comparison 
with equivalent sites nearby indicated that the severe 
impacts diminish rapidly with distance from feeding 
sites. However, destabilisation of the dune may result 


107 


RUSA DEER IN ROYAL NATIONAL PARK, SYDNEY 


in the longer term if the denuded sites become the 
catalyst for more widespread mobilisation of sand. 
Within native vegetation, shrub cover was locally 
reduced along deer tracks and in encampment areas 
but erosion of sand was generally minimal. Impacts 
on wetland soils were more marked, with increased 
exposure, compaction and surface deformation 
evident in all three wetlands inspected. 

The exclosure experiment showed that deer may 
have very substantial impacts on the populations of 
at least some plant species over a relatively short 
time frame. The extent of foliage and shoot removal 
precluded any chance of reproduction in the plants 
exposed to browsing by deer. The level of mortality 
caused by bark stripping and stem breakages, 
while comparatively small over the duration of 
the experiment, would account for an appreciable 
reduction in a cohort of saplings over several years. 
The combined effects of foliage and shoot removal 
and cumulative mortality are likely to delay or 
prevent the growth of individual plant canopies above 
the browse height. Syzygium paniculatum appears 
to be one of the more palatable native plants in the 
study area, despite the essential oils in its foliage 
and the availability of copious grass in the vicinity. 
The observed effects on S. paniculatum probably 
represent the more severe of those to be observed 
among wild plant populations although, in bushland, 
severe defoliation was observed in species from a 
wide range of plant families including Fabaceae 
and Epacridaceae (Fig. 2). Effects are likely to be 
particularly severe in rainforest communities, such 
as the Littoral Rainforest Endangered Ecological 
Community. They also have major implications for 
revegetation projects, which may be prone to major 
losses if deer gain access to the plantings before they 
grow beyond browse height (e.g. Opperman and 
Merenlender 2000). 

A number of limitations in the design of 
the exclosure experiment impose constraints on 
interpretation of the results. The comparison was 
based on unreplicated treatments with unbalanced 
sample sizes. In the field, plants would be exposed 
to deer browsing at a much younger stage and would 
not be surrounded by grassy areas as they were in 
our experiment. Despite these differences between 
the experimental conditions and those in the field, 
the symptoms of deer browsing observed in the 
experimental population were substantial (relative to 
the control plants) and similar to those observed in 
other rainforest and sclerophyll forest plant species 
during reconnaissance of native vegetation. Data 
from replicated exclosures, which have recently been 
established in native vegetation (Moriarty 2004b), 


108 


will help to resolve uncertainties that arise when 
interpreting our experimental results in the context of 
wild plant populations. 

In Fig. 4, we propose a model of impacts based 
on plant life histories as a means of structuring future 
experimental investigations on the medium- to long- 
term impacts of deer herbivory on native vegetation. 
The model proposes that deer herbivory has its 
largest impact on population viability by interrupting 
two major plant life-cycle processes: seedling 
establishment and seed production. Compared with 
established plants, seedlings have less capacity to 
recover after defoliation and could be more palatable 
due to the lower content of fibre, tannins and phenolic 
compounds in their leaves. Bushfires expose more 
seedlings to browsing by deer because they release 
seeds of many species from dormancy or canopy 
storages (Keith 1996). Populations of plant species 
that only regenerate from seed are exposed to greater 
risks of decline than those in which a proportion 
of pre-fire established plants survive. Factors that 
influence the density of post-fire deer populations, 
such as fire size and patchiness and deer dispersal 
patterns, are potentially important in mediating the 
impact of deer on seedling recruitment. 

A second mechanism of deer impact on the 
viability of plant populations is through the reduction 
of fruit production (Fig. 4), as the resulting decline 
in seed banks reduces the capacity for seedling 
recruitment. These effects are likely to be most 
significant in species such as terrestrial orchids 
and lilies, in which all reproductive material may 
be consumed in a single visit, and those species 
whose reproductive effort is largely limited to the 
post-fire period (Keith 1996). While the magnitude 
of reductions in fecundity remain to be quantified 
under varying levels of browsing, our qualitative 
observations during reconnaissance suggest that the 
current densities of deer populations in the Jibbon/ 
Bundeena area could be causing substantial reductions 
in seed production in a wide range of plant species. 

A third life-cycle process susceptible to 
interruption by deer herbivory is the survival and 
growth of established plants. These effects are 
likely to be less significant in plant species that are 
capable of growth above the vertical reach of deer 
(c. 1.5-2.0 m). However, sustained herbivory could 
slow or block the transition from juvenile to mature 
growth forms and could also reduce survival in 
mature individuals if bark stripping occurs at levels 
observed in the exclosure experiment. These effects 
have resulted in failure of revegetation projects 
elsewhere in Royal National Park (e.g. Hacking River 
Valley) and in other parts of the world (Augustine 


Proc. Linn. Soc. N.S.W., 126, 2005 


D. KEITH AND B. PELLOW 


Consumption of 
shoots reduces seed 
production 


(Dee eee oe SES SSS ee a SSS SSS SS Se 


Consumption of shoots 
reduces survival and growth 
rates of established plants 


Possible predation of seeds after release 
: May reduce seedling emergence (?) 


Seedlings 


Consumption of 
seedlings reduces 
seedling establishment 


Figure 4. Simplified plant life cycle showing life history processes influenced by deer herbivory. Proc- 
esses marked by unbroken lines are predicted to exert a greater influence on population viability than 


those marked by broken lines. 


and Frelich 1998, Opperman and Merenlender 2000, 
Coomes et al. 2003), imposing substantial additional 
costs on the rehabilitation of degraded areas. Deer 
are also implicated in the predation of seeds after 
their release from plants. For example, the removal 
of Telopea speciosissima seeds from experimental 
caches coincided with deposition of deer faeces at the 
site (T. D. Auld and A. Denham, unpublished data). 
However, the magnitude of seed losses is unknown. 
In wild populations, the impacts on survival 
and growth of established plants are likely to be less 
significant than effects on seedling recruitment and 
fruit production, but all three processes will have 
cumulative impacts on the population viability of 
many plant species. The magnitude of impacts for any 
given plant species will depend upon its palatability 
relative to other available forage, density of the deer 
population, the accessibility of edible plant tissues and 
the propensity to replace leaf tissue. The model (Fig. 
4) nevertheless predicts the decline of many woody 
plant species and some herbaceous species because 
the recruitment of new seedlings fails to compensate 
for the attrition of established plants. This is supported 
by Moriarty’s (2004b) observation that 30-70% fewer 
plant species occurred in understorey vegetation 
exposed to high deer densities compared with that 


Proc. Linn. Soc. N.S.W., 126, 2005 


exposed to low deer densities. On the other hand, deer 
herbivory could favour the relatively few unpalatable 
plant species, particularly those ferns and sedges that 
may spread vegetatively to occupy space vacated by 
declines in densities of palatable shrubs and herbs. 
The abundance of Preridium esculentum and relative 
scarcity of shrubs in forest understories adjacent to 
cleared grassy areas would seem consistent with this 
prediction. The predicted transformation of forest 
understories from structurally complex, floristically 
diverse assemblages to simple assemblages dominated 
by ferns and sedges is likely to reduce the suitability 
of habitats for a range of vertebrate and invertebrate 
fauna (Catling 1991, York 1999). 

Reducing the density of deer populations may 
slow or reverse some effects of deer herbivory, as 
suggested by the observed recovery of surviving 
individuals of Syzygium paniculatum when deer were 
excluded from the treatment yard. Such effects have 
also been observed elsewhere (Anderson and Katz 
1993, McShea and Rappole 2000). However, it is not 
known what level of deer control in the wild would 
produce such a response. Furthermore, the recovery of 
native vegetation would be impeded by deer-related 
soil erosion and compaction, or if densities of shrubs 
or their seed banks had already declined to very low 


109 


RUSA DEER IN ROYAL NATIONAL PARK, SYDNEY 


levels. Our model suggests that fire management and 
the control of deer after fire are likely to be crucial 
in managing impacts of deer herbivory on native 
vegetation. 


ACKNOWLEDGEMENTS 


We thank residents of Bundeena for access to their 
yards. Paul Adam, Tony Auld and Andrew Moriarty 
commented on a draft manuscript. 


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Anderson, R. C. and Katz, A. J. (1993). Recovery of 
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Anonymous (1914). Offical guide to the National Park, 
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1981). 

Augustine, D. J. and Frelich, L. E. (1998). Effects of 
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Bentley (1979). An introduction to deer in Australia: 
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Hawthorn Press, Melbourne. 

Catling, P. (1991). Ecological effects of prescribed burning 
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Coomes, D. A., Allen, R. B., David, M., Forsyth, D.M. and 
Lee, W. G. (2003). Factors Preventing the Recovery 
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Invasive Deer. Conservation Biology 17, 450-459. 

Forsyth, D. M., Duncan, R. P., Bomford, M. and Moore, 
G. (2004). Climatic suitability, life history traits, 
introduction effort, and the establishment and spread 
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Fuller, R. J. and Gill, R, M. A. (2001). Ecological impacts 
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Hamilton, C. E. (1981). Rusa deer in the Royal National 
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movements. MSc thesis. University of Sydney. 

Harden, G. J. (1990-2002). Flora of New South Wales. 
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Jane, G. T. (1994). The impact of browsing animals on 
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Zealand. Australian Journal of Botany 42, 113-124. 

Kay, S. (1993). Factors affecting severity of deer browsing 
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Keith, D. A. (1996). Fire-driven mechanisms of 
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K. (1994). Tree and shrub mortality and debarking by 
sambar (Cervus unicolour) in Gir after a drought in 
Gujurat, India. Biological Conservation 68, 149-154. 

McShea, W. J. and Rappole, J. H. (2000). Managing 
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Populations through Manipulation of Deer 
Populations. Conservation Biology 14, 1161-1170. 

Mladenoff, D. J. and Stearns, F. (1993). Eastern hemlock 
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Moriarty, A. (2004a). The liberation, distribution, 
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Moriarty, A. (2004b). The environmental impacts of 
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Proc. Linn. Soc. N.S.W., 126, 2005 


A New Early Silurian Species of Trimerella (Brachiopoda: 
Craniata) from the Orange District, New South Wales 


IAN G. PercivAL!* AND A.J. WRIGHT2* 


‘Geological Survey of New South Wales, Department of Primary Industries, State Geoscience Centre, 947- 
953 Londonderry Road, Londonderry, NSW 2753; *School of Earth and Environmental Sciences, University 
of Wollongong, NSW 2522; *Honorary Research Associate, Centre for Ecostratigraphy and Palaeobiology, 


Macquarie University, NSW 2109. 


Percival, I.G. and Wright, A.J. (2005). A new Early Silurian species of Trimerella (Brachiopoda: Craniata) 
from the Orange district, New South Wales. Proceedings of the Linnean Society of New South Wales 
126, 111-120. 


Trimerella australis, a new species of craniate brachiopod, is described from silicified material extracted 
from Early Silurian limestone in the Bowan Park district, 22 km west of Orange in central New South Wales. 
Accompanying conodonts of the Distomodus staurognathoides Zone indicate this unnamed unit is mid 
Llandovery (latest Aeronian to earliest Telychian) in age, and support correlation with the Cobblers Creek 
Limestone at the base of the Waugoola Group. As with most other occurrences of trimerellide brachiopods 
in the Late Ordovician and Early Silurian of the Lachlan Orogen, 7 australis completely dominates 
its depauperate faunal associates of corals including Aphyllum? sp., cf. Axolasma sp. and cf. Halysites 
cratus Etheridge, 1904 , and very rare atrypide brachiopods. Although all specimens of T: australis are 
disarticulated, the community is interpreted as preserved essentially in situ, representing a very shallow 


water Benthic Assemblage 2 environment. 


Manuscript received 5 October 2004, accepted for publication 16 February 2005. 


KEYWORDS: Benthic Assemblage, biostratigraphy, Brachiopoda, Craniata, early Silurian, halysitid, 


tetracorals, trimerellide 


INTRODUCTION 


Silurian trimerellide brachiopods from the 
Lachlan Orogen of eastern Australia are poorly known 
compared to their Late Ordovician predecessors. Only 
Keteiodoros bellense from the Dripstone Formation 
(Early Silurian, Wenlock) of the Oakdale Anticline, 
southeast of Wellington, NSW (Strusz et al. 1998), 
has been fully documented. One other trimerellide 
specimen, identified by Strusz (1982) as Trimerella 
sp. from the Walker Volcanics of the Canberra area, 
has been illustrated from Silurian strata in NSW; the 
age of this occurrence is reported as either Wenlock 
(Strusz 1982) or early Ludlow (Talent et al. 2003). 
Unidentified trimerellide material, possibly 7rimerella, 
is known from the Manildra district (Savage 1968), in 
limestone (probably allochthonous) of the Greengrove 
Formation, the age of which has been interpreted as 
either lower mid Llandovery (Munson et al. 2000) or 
mid to late Llandovery (Talent et al. 2003). Here we 
document the new species Trimerella australis, from 
the Bowan Park district west of Orange, in limestone 


of mid Llandovery age (Distomodus staurognathoides 
Conodont Zone, equated to the upper convolutus to 
lower crispus Graptolite Zones). This new species 
qualifies as the biostratigraphically most precisely 
constrained Silurian representative of this order 
presently known from eastern Australia. 


STRATIGRAPHIC SETTING AND AGE 


Abundant silicified specimens of the new species 
of Trimerella occur in an unnamed limestone, situated 
at Grid Reference 672400 mE 6315070 mN (GDA94 
co-ordinates) in the Quarry Creek area, east of the 
Bowan Park district, about 22 km west of Orange 
(Fig. 1). The limestone was first mapped in detail by 
Packham and Stevens (1955, fig. 1), who depicted 
it as two outcrops offset by an east-west fault. 
Immediately west of the Silurian limestone lie Late 
Ordovician volcanics equated to the Malachis Hill 
Formation. To the east, the more northerly outcrop of 
limestone that yielded the abundant trimerellides abuts 


EARLY SILURIAN TRIMERELLA FROM ORANGE DISTRICT 


500 750 


209° y 


0°o 


fo} 


REFERENCE 
Holocene alluvium 
Miocene basalt 


Silurian 


Quarry Creek limestone 


Siltstone 

unamed limestone 
Late Ordovician 

Malachis Hill Formation 


Bowan Park limestone 


road 
creek 
fault 


geological boundary 


9. VL Mite 
A in “Alda fom" i" i,t "is ie" fi "fags "8 UPB, Cy 
NOVEM MOVEV! VEO Ve va iviet 
Ve VON VOM 
¥ VV VE SV VeV Oa wy, 
ve 
Move 
VV MV ON OM May 
ee Vaan Ve re ee 
VV MMV Me Yul Moll: IV Mo 
WO MAM VM eM: Mew ieee 
NP ONOV VIM IV MENON NA VAY VV 


MOLONG 33°18'S 


Mount 
Canobolas 


Figure 1. Map of the Quarry Creek area, east of Bowan Park district, 22 km west of Orange, 
NSW, showing the location of the limestone in the W.T. section of Bischoff (1987) that yielded the 
trimerellide brachiopods and associated fauna documented in this paper. Geology adapted from 
Packham and Stevens (1955) and mapping by Packham in Jenkins (1986) and Rickards et al. (1995). 


graptolite-bearing beds (g12 locality of Packham and 
Stevens), considered by those authors to be of late 
Llandovery age. These graptolitic beds are probably 
faulted against the limestone. Revised identifications 
in Rickards et al. (1995, table 1) of many of the late 
Llandovery graptolite faunas found by Packham and 
Stevens have indicated that these localities should 
now be assigned ages in the early to mid Wenlock. 
R.B. Rickards (pers. comm. 2004) informs us that 


112 


the graptolite fauna from g12 contains Testograptus 
testis and should be reassigned to the late Wenlock 
lundgreni-testis Biozone. Farther east is limestone of 
Late Ordovician age, termed the Barton Limestone by 
Packham and Stevens (1955). Current terminology 
has seen this name suppressed in favour of the 
Bowan Park Limestone Subgroup that is extensively 
developed in the Bowan Park area to the west. 
Bischoff (1987) sampled the Silurian limestone 


Proc. Linn. Soc. N.S.W., 126, 2005 


I.G. PERCIVAL AND A.J. WRIGHT 


(approximately 16 m thick) in his W.T. section, 
and first recognised its mid Llandovery age, 
based on conodonts. These were identified by 
Bischoff as Aulacognathus angulatus, Distomodus 
staurognathoides (alpha morphotype), Oulodus 
australis, O. planus planus, Ozarkodina excavata 
eosilurica and Oz. waugoolaensis; two additional 
species — Oulodus panuarensis and Pterospathodus 
cadiaensis — were very rare in Bischoff’s collections. 
Our sample GSNSW C1892 from the upper part of 
the limestone, where the trimerellides are particularly 
abundant, yielded Oulodus australis, O. planus planus, 
Oz. waugoolaensis, together with Panderodus sp. The 
additional conodont species recorded by Bischoff 
(1987, table 5), but not identified in our sample, were 
restricted to the lower part of his W.T. section. Bischoff 
assigned this conodont fauna to his Aulocognathus 
antiquus — Distomodus  staurognathoides alpha 
Assemblage Zone, which he correlated with the lower 
turriculatus Graptolite Zone, possibly extending into 
the upper part of the preceding sedgwickii Graptolite 
Zone. Simpson (1995) queried the veracity of A. 
antiquus, and argued that Bischoff’s assemblage 
zone bearing this name should be equated with the 
Distomodus staurognathoides Zone of global usage. 
Uncertainty in interpretation of the Awlocognathus 
lineage proposed by Bischoff only affects the lower 
limit of the Zone, and Strusz (1996) (following 
Simpson) aligned the local conodont zone with 
the interval represented by the convolutus to lower 
crispus graptolite zones. This range encompasses the 
full extent of the age of the W.T. section as interpreted 
by Bischoff (1987). 

The Quarry Creek Limestone is regarded as 
either late Llandovery or early Wenlock in age 
(Bischoff 1987). Rickards et al. (1995) discussed 
possible problems with the conflict in age between 
this limestone and the overlying Panuara Formation, 
but this appears to have been resolved with revision 
of graptolite identifications from the lower part of the 
latter unit supporting an early Wenlock age. Munson 
et al. (2000) placed the Quarry Creek Limestone 
straddling the Llandovery-Wenlock boundary. 
Although Talent et al. (2003, figure 6) depicted 
the Quarry Creek Limestone as occupying a lower 
Wenlock horizon, they admitted the possibility 
(p. 200) that this unit is entirely upper Llandovery. 
Whatever its precise age range, the Quarry Creek 
Limestone is substantially younger than the unnamed 
limestone of Bischoff’s (1987) W.T. section, and the 
two horizons cannot be equated as shown in Jenkins 
(1986, figure 34) and Rickards et al. (1995, figure 2). 

The unnamed limestone in the W.T. section is thus 
the oldest Silurian stratum in the Quarry Creek area, 


Proc. Linn. Soc. N.S.W., 126, 2005 


and is correlated using conodonts (Bischoff 1987) 
with the Cobblers Creek Limestone at the base of the 
Waugoola Group in the Angullong district, SSW of 
Orange. Recognition of this relationship is significant 
in constraining the upper limit of the Panuara Hiatus 
that separates the Ashburnia and Waugoola groups. 
Krynen and Pogson (in Pogson and Watkins 1998, 
p.109) interpreted the base of the Waugoola Group 
as diachronous, ranging from early late Llandovery 
in the Angullong Syncline succession, to terminal 
Llandovery (P. amorphognathoides Conodont Zone) 
in the Quarry Creek area. However, the presence of 
mid Llandovery strata in the W.T. section indicates 
that the base of the Waugoola Group in the Quarry 
Creek area is essentially the same age as elsewhere, 
and hence is isochronous rather than diachronous. 


FAUNAL ASSOCIATES AND DEPOSITIONAL 
ENVIRONMENT 


As with almost all other occurrences of 
trimerellide brachiopods known from the Late 
Ordovician and Early Silurian of the Lachlan Orogen, 
T: australis is numerically abundant and completely 
dominates an otherwise depauperate group of faunal 
associates. As extraction of fossils was by dissolution 
of bulk limestone samples in dilute acids, the 
silicified residues obtained are believed to be fairly 
representative of the preservable elements of the 
trimerellide community. In the W.T. section, faunal 
associates comprise the tetracorals cf. Axolasma sp. 
(Fig. 2a, b) and Aphyllum? sp. (Fig. 2d-f), the halysitid 
tabulate coral cf. Halysites cratus Etheridge, 1904 
(Fig. 2c), an indeterminate finely-ribbed atrypide 
brachiopod (Fig. 2g-1) and a smooth atrypide (Fig. 
2}). Each of these taxa is represented by at most a 
few specimens only, compared to many dozens of 
T. australis valves (although a high proportion of 
the latter are fragmentary, due either to post-mortem 
breakage or, more likely, incomplete silicification). 
One example of Aphyllum? is preserved on the 
exterior surface of a dorsal valve of 7. australis, with 
the calyx adjacent to the anterior margin (Fig. 3p). 
Possibly the coral not only employed the trimerellide 
as a substrate but could also have obtained nutrients 
from the inhalant or exhalant currents of the living 
brachiopod. All examples of TZ australis are 
disarticulated valves, probably resulting from storm 
or current activity disturbing in situ specimens or 
redistributing deceased individuals (cf Webby and 
Percival 1983). Orientation of shells is generally 
horizontal, and erratic rather than consistently either 
convex up or down. 


113 


EARLY SILURIAN 7TRIMERELLA FROM ORANGE DISTRICT 


Figure 2. Corals and articulate brachiopods associated with Trimerella australis. All specimens 
silicified; scale bars in black represent 5 mm. (a, b) cf. Axolasma sp., MMF 44114, lateral and calical 
views, displaying axial vortex. (c) cf. Halysites cratus Etheridge, 1904, MMF 44115, with calical 
views of partial colony. (d-f) Aphyllum? sp., oblique view of calyx, MMF 44116, longitudinal broken 
fragment, MMF 44117 showing septa and budding, and fragment of interior of calyx, MMF 44118 
displaying acanthine septa. (g, h) indeterminate ribbed atrypide, external and internal views of 
partial ventral valve and fragment of attached dorsal valve, MMF 44119. (i) external view of dorsal 
valve of conjoined specimen of ribbed atrypide, MMF 44120. Note circular pitting on surface, possibly 
caused by predatory sponge. (j) dorsal view of conjoined valves of smooth atrypide, MMF 44121. 


Comparable trimerellide communities include 
Late Ordovician shell beds consisting of Eodinobolus 
stevensi in the lower Fossil Hill Limestone at 
Cliefden Caves and Daylesford Limestone at Bowan 
Park (Webby and Percival 1983, Percival 1995), 
and Belubula spectacula in the Belubula Limestone 
at Cliefden Caves (Percival 1995). Keteiodoros 
bellense from the Dripstone Formation (Wenlock) 
of the Oakdale Anticline (Strusz et al. 1998) is the 
only locally-documented Silurian example of an in 
situ trimerellide community. The Late Ordovician 
examples from Cliefden Caves and Bowan Park were 
assigned to a Benthic Assemblage | or upper B.A. 2 
depositional setting, equivalent to a marine shoreline 
(intertidal to very shallow subtidal) environment 
(Percival and Webby 1996). Ordinarily this would be 
in protected waters such as a lagoon or embayment, 
as trimerellides — lacking a pedicle attachment 
~ relied on posterior gravity weighting for stability 
of orientation when alive. Such shallow waters are 
also highly susceptible to disturbance during storms, 


114 


accounting for death assemblages and _ stacked 
shell beds commonly encountered in trimerellide 
occurrences. Strusz et al. (1998) attributed to 
Keteiodoros bellense in the Dripstone Formation 
a B.A. 2 (shallow subtidal) setting, in quiet waters 
inshore of a protective Palaeophyllum wave barrier. 

We found no evidence of a protective wave 
barrier, such as might have been formed by coral 
thickets or reefs, associated with the ZT. australis shell 
beds. However, comparable halysitid tabulates and 
solitary tetracorals associated with this occurrence, 
and also with Keteiodoros bellense, suggest that 
the depositional environments of these Silurian 
trimerellides were similar. The stratigraphic position 
of the 7: australis shell beds, in limestone deposited 
following the erosional Panuara Hiatus, also argues 
for very shallow water, nearshore conditions. 
Therefore, a quiet water depositional environment no 
deeper than B.A. 2, and possibly as shallow as B.A. 
1, is interpreted for this unit. 


Proc. Linn. Soc. N.S.W., 126, 2005 


I.G. PERCIVAL AND A.J. WRIGHT 


SYSTEMATIC PALAEONTOLOGY 


Phylum BRACHIOPODA 
Class CRANIATA Williams, Carlson, Brunton, 
Holmer and Popov, 1996 

Order TRIMERELLIDA Goryansky and Popov, 
1985 

Superfamily TRIMERELLOIDEA Davidson and 

King, 1872 

Family TRIMERELLIDAE Davidson and King, 

1872 
Genus Trimerella Billings 1862 


Type species 
Trimerella grandis Billings, 1862, by subsequent 
designation of Dall (1870). 


Diagnosis 

Shell dorsibiconvex, elongate triangular; ventral 
valve flattened, ventral interarea high, triangular, 
apsacline, with deep concave homeodeltidium 
occupying more than half of interarea; dorsal valve 
strongly convex, beak incurved; ventral umbonal 
cavities small or vestigial; both valves with distinctly 
raised visceral platforms, extending anterior of centre; 
visceral platforms with deep vaults, separated by 
median partition extending anterior to platform; dorsal 
hinge plate high, strongly incurved; dorsal vascula 
lateralia broad, slightly divergent, lacking trace of 
bifurcation (Popov and Holmer 2000, p.185). 


Species included: 

?T. asiatica Li, 1984; T: jiangshanensis (Li, 1984) 
(formerly Prosoponella) and T. zhoujiashanensis 
(Li and Han, 1980) (formerly Machaerocolella) 
— both genera synonymised with Trimerella 
by Percival (1995) and Popov et al. (1997); all 
preceding species from Late Ordovician (early 
Ashgill) Huangnehkang Formation, Jiangshan 
county, W. Zhejiang, China (according to Rong 
and Li 1993). 

T. attentuata Goryansky, 1972 from Early Silurian 
(late Llandovery to early Wenlock) Donenzhal 
Formation, Kazakhstan (Popov et al. 1997). 

T. acuminata Billings, 1862 from Silurian (Wenlock- 
Ludlow) Guelph Limestone, Ontario (Popov 
and Holmer 2000, p. 186); Niagara Group, Ohio 
and Illinois; Gotland and Faro islands, Sweden; 
Gorno Altay of Russia (Kul’kov 1967). 

T. billingsi Dall, 1871 from Silurian (Wenlock- 
Ludlow) Guelph Limestone, Ontario. 

T. grandis Billings, 1862 from Silurian (Wenlock- 
Ludlow) Guelph Limestone, Ontario; Niagara 
Group, Ohio. 


Proc. Linn. Soc. N.S.W., 126, 2005 


T. lindstroemi (Dall, 1870) from Silurian (Wenlock) 
Hogklint beds, Gotland (Popov and Holmer 2000, 
p. 186) and (Ludlow) Klinteberg Limestone, 
Gotland (Cocks in Murray 1985, p. 55). 

T. ohioensis Meek, 1871 from Silurian (Wenlock- 
Ludlow) Guelph Limestone, Ontario; Niagara 
Group, Ohio and New York (Popov and Holmer 
2000, p. 186). 

T. wisbyensis Davidson and King, 1874 from Early 
Silurian (Wenlock) of Gotland and Estonia. 


Trimerella australis sp. nov. 
Fig. 3a-q, Fig. 4a-b 


Type material 

Holotype MMF 44100; paratypes MMF 44101- 
44113, from unnamed Early Silurian limestone on 
‘Coorombong’ property, east of the Bowan Park 
district, about 22 km west of Orange, NSW. All 
specimens curated in the NSW State Palaeontological 
Reference Collection, held at the Geological Survey 
of NSW Geoscience Centre, Londonderry. 


Diagnosis 

Broadly acuminate 7rimerella with vestigial to 
shallow ventral umbonal chambers; narrow ventral 
platform divided medially by parallel-sided furrow, 
and deeply excavated beneath by twin vaults; 
dorsal platform supported by median septum longer 
than rudimentary counterpart in ventral valve but 
terminating well short of anterior margin. 


Description 

Dorsibiconvex shell apparently lacking external 
ornament or pronounced growth lamellae; widest at, 
or slightly anterior to, midlength. As all specimens 
are fragmentary, overall dimensions are estimated 
from reconstructions (Fig. 4); maximum width 
approximates 45-50 mm, and maximum length is 
probably at least 60 mm for ventral valves, and about 
45 mm for brachial valves. Height of conjoined valves 
estimated at no more than 20 mm. 

Ventral valve broadly acuminate with 
posterolateral margins forming a right angle at 
slightly incurved beak; pseudointerarea apsacline to 
orthocline. Homeodeltidium shallowly depressed, 
marked by incised flattened chevrons and extending 
to about one-third width of pseudointerarea at 
anterior edge (Fig. 3a); homeodeltiditum delineated 
from elongate triangular propareas by incised groove; 
deltidial ridges lacking, except in one possibly 
immature individual where the lateral margins of 
the homeodeltidum are slightly raised (Fig. 3g). A 
depressed flattened area immediately in front of the 


115 


EARLY SILURIAN TRIMERELLA FROM ORANGE DISTRICT 


116 Proc. Linn. Soc. N.S.W., 126, 2005 


I.G. PERCIVAL AND A.J. WRIGHT 


Figure 3. (LEFT) Trimerella australis sp. nov. from unnamed Lower Silurian limestone near Quarry 
Creek. a-b is the holotype, all other figured specimens are paratypes. All specimens 1.5 times natural size. 
(a, b) internal view of posterior fragment of ventral valve showing pseudointerarea, and oblique view of 
specimen to display umbonal and platform chambers, MMF 44100. (c) internal view of finely preserved 
posterior fragment of ventral valve, showing vaulted platform, MMF 44101. (d) internal view of posterior 
fragment of ventral valve showing pseudointerarea, MMF 44102. (e) internal view of posterior fragment 
of ventral valve showing pseudointerarea and platform, MMF 44103. (f) internal view of posterior 
fragment of ventral valve showing pseudointerarea and eroded platform represented by lateral and 
medial walls of chambers, MMF 44104. (g) internal view of posterior fragment of ventral valve showing 
partial pseudointerarea, MMF 44105. (h, i) internal view of posterior fragment of dorsal valve, oriented 
obliquely to show platform chambers, h, and normal view showing platform, i, MMF 44106. (j) internal 
view of posterior fragment of dorsal valve, showing pseudointerarea and platform, MMF 44107. (k) 
external view of posterior part of dorsal valve, partially eroded anteriorly revealing interior of platform 
chambers, MMF 44108. (1) internal view of posterior fragment of dorsal valve, showing pseudointerarea 
and platform, MMF 44109. (m) internal view of posterior fragment of dorsal valve, showing platform and 
marginal area, MMF 44110. (n) external view of fragment of ventral? valve, partially eroded anteriorly 
revealing interior of platform chambers, MMF 44111. (0, p) internal and external views of anterior fragment 
of dorsal valve showing median septum, 0, and tetracoral Aphyl/um? sp. growing adjacent to valve margin, 
p, MMF 44112. (q) internal view of posterior fragment of dorsal valve, showing platform, MMF 44113. 


anterior edge of the pseudodeltidium may represent the 
site of a very weakly impressed umbonal muscle scar 
(Fig. 3a, c, f). Umbonal chambers variably developed 
even in fully grown specimens, where they may be 
rudimentary (Fig. 3c) or shallowly excavated (Fig. 
3a, b, d, e, f) beneath pseudointerareas, but are always 
less prominent than visceral platform chambers (Fig. 
3b, c, n). Cardinal buttress absent, instead replaced 
by distinct narrow parallel-sided median furrow 


slightly depressed below level of adjacent visceral 
platform (Fig. 3a, c), although this feature is not 
always apparent (Fig. 3e). Platform surface smooth, 
lacking traces of muscle scars, and narrowly rounded, 
reflecting conical vaulted chambers that extend to 
posterior end of cardinal buttress furrow. V-shaped 
anterior extremity of platform supported by very short 
median septum not extending beyond mid-length of 
valve. Lateral muscle scars, inserted along crescentic 


Figure 4. Reconstructions of (a) ventral and (b) dorsal valves of Trimerella australis. 


Proc. Linn. Soc. N.S.W., 126, 2005 117 


EARLY SILURIAN TRIMERELLA FROM ORANGE DISTRICT 


furrows parallel to valve margins, prominent in some 
specimens (Fig. 3c). Pallial canals not visible. 

Dorsal valve elongately ovate, with rounded to 
subangular beak that tends to be slightly to markedly 
asymmetric, and is gently to moderately incurved 
(Fig. 3j, 1). Brachidial plate curvilinear, prominent 
but relatively low, occupying all of very low 
pseudointerarea that expands and merges laterally 
with broad marginal area raised slightly above 
valve floor; incised boundary between marginal area 
and valve floor Fig. 3m) may equate to crescent of 
previous authors. Marginal area indistinguishable 
anterior to midlength. Umbonal chambers lacking; 
umbonal muscle scar not apparent. Visceral platform 
narrow, distinctly vaulted with long conical chambers 
beneath (Fig. 3h, i, k); ventral surface strongly convex, 
bisected longitudinally by broad shallow median 
depression (Fig. 3j, m, q). No trace of muscle scars on 
visceral platform, which does not extend into anterior 
half of valve. Anterior edge of wall separating vaults 
beneath platform is continuous with thin low median 
septum, much longer than its counterpart in ventral 
valve but ending well short of anterior valve margin; 
the median septum on the one specimen available 
(Fig. 30, Fig. 4b) is estimated to terminate between 
three-quarters and four-fifths valve length. Internal 
shell surface smooth; no pallial canals discernible. 


Remarks 

Trimerella attentuata Goryansky, 1972 (revised 
by Popov et al. 1997), from the late Llandovery to 
early Wenlock Donenzhal Formation of Kazakhstan, 
is closest in age and general appearance to the new 
species. It differs in being much smaller (attaining 
just half the dimensions of 7: australis), and in the 
relatively longer extension of the visceral platform 
and median septum in the dorsal valve. The ventral 
platform of 7: attenuata is relatively wide and in two 
figured specimens (Popov etal. 1997) bears prominent 
diagonal growth lines, whereas that of 7: australis is 
narrow and smooth. There does, however, appear to 
be a comparable narrow median furrow developed on 
the platform of both species, and neither displays any 
conspicuous extension of a median septum anterior 
to the ventral platform. All illustrated examples of 7. 
attenuata are internal moulds that do not adequately 
reveal details of the pseudointerareas. 

Comparisons with previously — established 
Wenlock to Ludlow species are also hindered by 
significant differences in preservation. These species 
of Trimerella were originally defined on the basis of 
natural internal moulds that frequently lacked details 
of pseudointerareas and visceral platform surfaces. All 
of the species depicted by Davidson and King (1874) 


118 


and Hall and Clarke (1892) have been reconstructed 
with elongate median septa, that in the case of the 
dorsal valve extend almost to the anterior margin of 
the valve, unlike 7 australis. The new species also 
seems to be compressed dorsoventrally compared 
with most Wenlock to Ludlow forms. Few of these 
have been photographically illustrated in the 130 
years since Davidson and King’s (1874) monograph, 
but figures in the Treatise (Popov and Holmer 2000, 
p. 187) confirm the differences discussed above 
between all these species and T. australis. 

From species of 7rimerella described from the 
Late Ordovician (early Ashgill) Huangnehkang 
Formation of Jiangshan county, W. Zhejiang, China 
(Li and Han 1980, Li 1984), 7. australis is readily 
distinguished by its relatively deeply excavated 
ventral platform chambers and absence of a well- 
developed median septum extending anterior to this 
platform. Dorsal valves of both ?T. asiatica and T. 
zhoujiashanensis are inadequately known and cannot 
be compared with that of 7. australis. The new 
species is furthest removed morphologically from 
T. jiangshanensis (characterised by the presence of 
stout median septa extending to the margins of both 
valves). 


ACKNOWLEDGMENTS 


The presence of trimerellides in the unnamed limestone 
of Bischoff’s W.T. section was discovered during field 
studies in 1998 by Jonathan Dunningham of Emmanuel 
College, Cambridge. We are grateful to the owners of 
“Coorombong’ for access to their property. Preparation of 
the fragile silicified specimens was skilfully carried out by 
Gary Dargan, who dissolved the enclosing limestone in dilute 
hydrochloric acid. David Barnes carefully photographed 
the specimens and compiled the digital plates. Reviews by 
two anonymous referees were helpful in improving the final 
version of this paper. Publication by Percival is authorised 
by the Director of the Geological Survey of NSW. 


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120 Proc. Linn. Soc. N.S.W., 126, 2005 


Electrophoretic Evidence for the Presence of Tandanus tandanus 
(Pisces: Plotosidae) Immediately North and South of the 
Hunter River, New South Wales 


D.R. JERRY 


School of Marine Biology and Aquaculture, James Cook University, Townsville QLD 4811 
Dean.Jerry@jcu.edu.au 


Jerry, D.R. (2005). Electrophoretic evidence for the presence of Tandanus tandanus (Pisces: Plotosidae) 
immediately north and south of the Hunter River, New South Wales. Proceedings of the Linnean 


Society of New South Wales 126, 121-124. 


Eel-tailed catfish from the genus Zandanus are morphologically conservative. Previous allozyme 
electrophoretic surveys have demonstrated that up to three species of Zandanus catfish occur in south- 
eastern Australian freshwater streams. Two of these species are previously undescribed cryptic species. 
However, the taxonomic status of catfish i many coastal river systems is yet to be examined using 
allozyme electrophoresis. In this study four diagnostic allozyme markers were used to determine the 
taxonomic status of eel-tailed catfish in four NSW coastal populations from the Wallamba, Coolongolook, 
Hawkesbury and Georges River systems. Electrophoretic analyses demonstrated that the species of catfish 
in these four populations is T. tandanus. These results extend the distribution of T. tandanus to the coastal 
rivers immediately north and south of the Hunter River, NSW. 


Manuscript received 12 July 2004, accepted for publication 15 December 2004. 


KEYWORDS: allozymes, catfish, cryptic species, Zandanus. 


INTRODUCTION 


The eel-tailed catfish, Zandanus tandanus 
(Mitchell 1838), had until recently been regarded 
as a single, broadly distributed species that inhabits 
freshwater streams throughout the Murray-Darling 
Basin and coastal drainages of eastern Australia (Allen 
1989). However, allozyme electrophoresis studies in 
the 1990’s demonstrated that what was originally 
thought to be one species of Zandanus was in fact a 
complex assemblage containing up to an additional 
two undescribed cryptic species (Musyl 1990, Musyl 
and Keenan 1996, Jerry and Woodland 1997). These 
studies highlighted that the taxonomy of 7: tandanus 
should be revised to recognise the presence of at least 
three species of Zandanus in south-eastern Australia; 
i) T. tandanus which occurs throughout the Murray- 
Darling River Basin and in the Mary, Brisbane and 
Hunter coastal rivers; 11) an undescribed species of 
Tandanus within the coastal river systems between 
and including the Bellinger River and Manning 
Rivers and; 111) an undescribed species of TZandanus 
restricted to the coastal basin of the Clarence River 
system (and possibly the Richmond and Tweed River 


systems) (Fig. 1) (Jerry and Woodland 1997). 

The taxonomic status of eel-tailed catfish in 
many other NSW coastal river systems, however, 
is unresolved. For example, it is not known what 
taxonomic variant occurs immediately north and 
south of the Hunter River population of 7: tandanus. 
Of particular interest is whether the distribution of 
the “Bellinger” variant of Tandanus extends south 
to the Hunter River, or whether ZT. tandanus extends 
north. The aim of the present study therefore was to 
use species diagnostic allozyme markers to determine 
whether 7. tandanus has a wider distribution in the 
coastal drainages immediately north and south of 
the Hunter River (the area designated “taxonomy 
uncertain” in Fig. 1). 


MATERIALS AND METHODS 


Catfish were sampled from four coastal river 
drainages north and south of the Hunter River, NSW. 
The populations sampled were the Wallamba and 
Coolongolook Rivers (north of the Hunter River) and 
the Hawkesbury and Georges Rivers (south of the 


ELL-TAILED CATFISH IN NSW RIVERS 


= T. tandanus 


eal T. sp. (Clarence) 
By T. sp. (Bellinger) 


E35 Taxonomy uncertain 


Figure 1. Distribution of Tandanus tandanus, Tandanus sp (Clarence) and Tandanus sp (Bell- 
inger) in eastern Australia. T. tandanus occurs in the Murray-Darling, Brisbane, Mary and Hunter 
River drainages, T. sp (Clarence) in the Clarence and possibly Richmond and Tweed Rivers, and 
T. sp (Bellinger) in the Bellinger, Macleay, Hastings and Manning Rivers. Note; Tandanus cat- 
fish also inhabit other coastal drainages throughout eastern Australia, however, the taxonomic sta- 
tus of these populations has not been confirmed using diagnostic allozyme markers and it is pos- 
sible that one or more cryptic species are present. Currently they are considered to be 7. tandanus. 


Hunter River) (Fig. 2). Catfish were opportunistically 
sampled by gill netting during biological surveys (K. 
Bishop, personal communication), with two adult 
specimens collected from each of the river drainages. 
Upon capture whole specimens were immediately 
frozen and shipped to the laboratory on dry ice where 
liver and muscle tissues were excised. Tissue samples 
were prepared for electrophoresis according to the 
methods described by Shaklee and Keenan (1986). 
Musyl (1990) and Musyl and Keenan (1996) 
identified four Zandanus species diagnostic allozyme 
markers (International Enzyme Commission Number 
in parentheses); Glucose-6-phosphate isomerase 
GPI* (5.3.1.9.), Esterase EST* (3.1.1.-), Umbelliferyl 
esterases UMB-1* and UMB-2* (3.1.1.-). These 
markers were used to delineate the taxonomic status 
of the catfish samples according to the running and 
scoring conditions described in Jerry and Woodland 
(1997). To confirm the mobility of diagnostic alleles, 
the test populations were run against reference 
specimens of 7: tandanus (Hunter River) and 7. sp 
“Bellinger” (Manning River) in line-up gels for all 


122 


enzyme systems. 


RESULTS 


Catfish sampled from the four riverine systems 
exhibited identical allele motilities at all enzyme loci 
to those of the 7: tandanus reference sample from the 
Hunter River (Table 1). More specifically, test catfish 
samples exhibited the slower EST*(100) and UMB- 
2*(100) and the faster GPI*(100) and UMB-1*(100) 
alleles compared to the mobility of alleles diagnostic 
to the “Bellinger” variant from the Manning River. 
Although sample sizes were very small, no genetic 
variation was observed at any of the allozyme loci. 
This is consistent with the loci being “fixed” and 
diagnostic for different alleles among the various 
species. 


DISCUSSION 


Proc. Linn. Soc. N.S.W., 126, 2005 


D.R. JERRY 


Great Dividing Range 


Manning River 


Wallamba River 
Hunter River 


Coolongolook River 


Hawkesbury River 


Georges River 


Shoalhaven River 


—— 


v, 


Figure 2. River populations of Tandanus sampled from coastal drainages of central NSW. Hatched 
area represents known distribution of Tandanus sp (Bellinger) (ie Bellinger River south to the Manning 


River). 


Fixed alleles at the allozyme markers GPI*, EST*, 
UMB-1* and UMB-2* have been shown by several 
authors to be diagnostic in discriminating between 
the three known species of Tandanus inhabiting rivers 
and streams of south-eastern Australia (Musyl 1990, 
Musyl and Keenan 1996, Jerry and Woodland 1997). 
Therefore, based on the electrophoretic evidence 
presented herein, catfish that inhabit the two major 
coastal river drainages both north and south of the 
Hunter River can be considered to be 7: tandanus. 


The known distributional range of 7! tandanus in 
coastal drainages of NSW can be extended to include 
the Wallamba, Coolongolook, Hawkesbury-Nepean 
and Georges River systems. A variant of Tandanus 
is also found in coastal drainages as far south as the 
Shoalhaven River in southern NSW and given the 
close geographical proximity of these drainages, is 
likely to be 7. tandanus. However, further studies 
will be required to verify the taxonomic status of this 
population. 


Table 1. Allele motility at four species diagnostic allozyme loci of Tandanus catfish sampled from 
four NSW coastal rivers. Allele mobility is calculated as the relative distance moved in the gel of the 
allele compared to that of the Hunter River population (designated a mobility of 100). The Manning 
River sample is a representative of the “Bellinger” species of Tandanus (Jerry and Woodland 1997). 


Hunter Wallamba 


look 


Coolongo- 


Hawkesbury | Georges 


100 100 100 


100 100 


100 100 100 


100 100 


100 100 100 


100 100 


100 100 100 


Proc. Linn. Soc. N.S.W., 126, 2005 


100 100 


123 


ELL-TAILED CATFISH IN NSW RIVERS 


ACKNOWLEDGEMENTS 


I would like to thank Mike Ramsey who ran some of 
the line-up allozyme gels and Keith Bishop for providing 
the test catfish samples. 


REFERENCES 


Allen, G.R. (1989). ‘Freshwater Fishes of Australia’. TFH 
Publications, Neptune City NJ 

Jerry, D.R. and Woodland, D.F (1997). Electroporetic 
evidence for the presence of the undescribed 
“Bellinger” catfish (Jandanus sp.) (Teleostei : 
Plotosidae) in four New South Wales mid-northern 
coastal rivers. Marine and Freshwater Research 48, 
235-40. 

Mitchell, T.L. (1838). “Three Expeditions into the Interior 
of Eastern Australia’. W. Boone, London. 

Musyl, M.K. (1990). Meristic, morphometric and 
electrophoretic studies of two native species 
of freshwater fishes, Macquaria ambigua 
(Percichthyidae) and Zandanus tandanus (Plotosidae), 
in south-eastern Australia. Ph.D Thesis, University of 
New England, Armidale, NSW. 

Musyl, M.K. and Keenan, C.P. (1996). Evidence for 
cryptic speciation in Australian freshwater eel-tailed 
catfish, Zandanus tandanus (Teleostei : Plotosidae). 
Copeia 1996 (3), 526-34. 

Shaklee, J.B. and Keenan, C.P. (1986). A practical 
laboratory guide to the techniques and methodology 
of electrophoresis and its application to fish fillet 
identification. CSIRO Marine Laboratories Report 
No. 177. 


124 Proc. Linn. Soc. N.S.W., 126, 2005 


Diversity and Biogeography of Subterranean Guano Arthropod 
Communities of the Flinders Ranges, South Australia. 


TimotHy Mou.tps 


Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, The 
University of Adelaide, South Australia 5005. 
timothy.moulds@adelaide.edu.au 


Moulds, T. (2005). Diversity and biogeography of subterranean guano Arthropod communities of the 
Flinders Ranges, South Australia. Proceedings of the Linnean Society of New South Wales 126, 125-132. 


This study documents the arthropod diversity and ecology of guano-associated species in 12 caves and 
mines of the Flinders Ranges, South Australia. Twenty two species from 12 orders and two classes are 
recorded. This represents a five-fold increase in species richness across the region from previously published 
and unpublished records. Eregunda Mine and Weetootla Gorge Mine 2 were the most diverse with five and 
six species recorded respectively. Specious groups included the tenebrionid beetle genus Brises and the 
emesine reduviid bug genus Armstrongula. Guano communities in the Flinders Ranges contain species 
in common with the Nullarbor Plain to the west and isolated arid karst areas to the north. There are few 
affinities with species found in the moist coastal regions of south-eastern Australia. 


Manuscript received 30 August 2004, accepted for publication 20 October 2004. 


KEYWORDS: Arthropoda, biospeleology, cave, food web, guano, invertebrate. 


INTRODUCTION 


Guano-associated invertebrate communities are 
poorly known for the vast majority of Australian 
caves. Animals in these communities are classified 
according to their ecological dependence on guano as 
either obligate guano-dwelling animals (guanobites), 
opportunistic guano-dwelling animals (guanophiles) 
or transitory guano-dwelling animals (guanoxenes) 
(Gnaspini and Trajano 2000; Humphreys 2000). The 
composition and evolution of guanophilic arthropod 
communities is dramatically different from resource- 
poor troglobitic (obligate cave-dwelling) communities 
(Gnaspini 1992; Gnaspini and Trajano 2000). The 
presence of virtually unlimited food resources 
enables a wide diversity of normally epigean (surface- 
dwelling) species to utilise the stable conditions found 
in caves. This has been previously demonstrated by 
studies in the south-east and Nullarbor Plain karst 
areas of South Australia (Richards 1971; Moulds 
2004). 

Several karst regions containing guano deposits 
are located in South Australia, ranging from high 
rainfall coastal areas such as Kangaroo Island and 
the lower south-east, to dry arid areas such as the 
Nullarbor Plain, Flinders Ranges and Davenport 


Ranges. The guanophilic arthropod assemblage in the 
maternal chamber of Bat Cave, Naracoorte, located in 
the upper south-east of South Australia, has received 
the most intensive study for guano invertebrates in 
Australia (Hamilton-Smith 2000; Sanderson 2001; 
Bellati et al. 2003; Moulds 2003). A major study of 
cavernicolous arthropod diversity and ecology on 
the Nullarbor Plain (Richards 1971) documented 
the subterranean communities, including those 
associated with guano. This was the first Australian 
study to document guanophilic arthropod ecology 
and provide possible geographic relationships with 
other Australian guanophilic communities. 

The Flinders Ranges, situated between the 
immense karst area of the Nullarbor Plain to the 
west, and the karst areas of south-eastern South 
Australia, contain a number of widely-scattered 
caves in horizontal or gently dipping Neoproterozoic 
crystalline limestone (Lewis 1976; Webb et al. 2003). 
Many of these caves support small (< 50) transient 
populations of cave-dwelling bats, dominated by the 
inland cave bat (Vespadelus findlaysoni Kitchener, 
Jones and Caputi), with the chocolate wattled bat 
(Chalinolobus morio Gray) occasionally recorded. 
The only previously published accounts of guanophilic 
arthropods for the Flinders Ranges are records of the 
beetle Brises acuticornis Pascoe from three caves 


SUBTERRANEAN GUANO ARTHROPODS 


(Hamilton-Smith 1967; Mathews 1986). In addition, 
several unpublished records of undetermined Acarina, 
Coleoptera (Carabidae) and Diptera (Nycteriibidae) 
were recorded by Elery Hamilton-Smith (Moulds 
2004). 

This study was undertaken due to the paucity of 
knowledge of cavernicolous invertebrates, specifically 
guano invertebrates, of this important biogeographic 
region that links the comparatively well-studied 
eastern and western karst regions of South Australia. 
The diversity of guano-associated arthropods in 12 
exemplar caves and mines of the Flinders Ranges is 
documented. Data collected through direct observation 
and relevant literature are combined to summarise 
species interactions as a food web. Biogeographic 
relationships of guanophilic arthropods are believed 
to be more closely related to nearby arid karst areas 


South Australia 


Lyndhurst 


| 
! 


Copley 


Lake Torrens 
(salt) 


Parachilna 


Brachina 
Gorge 


(e\ 


Bunyeroo |fe\ fe\ 
Gorge 


Hawker 


Cradock 


To Pt Augusta 


McKinleys Daughters Cave fe\ 


Eregunda Mine 


than to wetter coastal areas. These relationships are 
assessed and discussed. 


METHODS 


Nine caves and three mines were sampled during 
two field trips in April and September 2003 (Fig. 
1). Specimens were primarily collected individually 
using hand-held forceps due to the extremely 
localised guano deposits at most sites (Upton 1991). 
Guano samples were also taken from Weetootla 
Gorge Mines, Eregunda Mine, and Chambers Gorge 
caves for extraction of arthropods in Tullgren funnels 
as described by Upton (1991). Guano was also 
collected when available in sufficient quantities and 
measured for pH, allowing micro-habitat conditions 


Arkaroola 


Weetootla Gorge fe\ 
Balcanoona 


Moro Bat Cave fe\ 


Lake Frome 
(sait) 


fe\ 
Chambers 
Gorge 


fe\ 


Oraparinna 


Bat Cave 
fe\ Cave or mine sampled 


@ Major town 


{ ~~. _ Major road 


25km 50km 
ae AR ee eeemeeres eae | 


Figure 1. Localities of guano sites visited in the Flinders Ranges. Brachina, Bunyeroo, Chambers, and 


Weetootla gorges all contain two sites. 


126 


Proc. Linn. Soc. N.S.W., 126, 2005 


T. MOULDS 


to be assessed. The caves examined during this study 
do not include every guano-bearing cave in the area, 
but rather represent a cross-section of active bat caves 
found throughout the Flinders Ranges. A list of all 
Flinders Ranges caves historically known to contain 
guano can be found in Hamilton-Smith et al. (1997). 


Terminology 

Australian caves are referred to by a binomial 
alpha-numeric system according to Mathews (1985), 
with those of the Flinders Ranges using the prefix *F’. 
Mines are not included in this system and are referred 
to by name only. The division of caves into four 
environmental zones (entrance, twilight, transition, 
and deep zones, according to the amount of light and 
degree of interaction with surface climatic conditions) 
follows Humphreys (2000). 


Cave and mine site descriptions 

The majority of caves examined during this 
study are small, rarely extending into complete 
darkness or attaining a deep zone. Weetootla Gorge 
in the Gammon Ranges was the northern-most area 
examined. Weetootla Gorge Mines | and 2 (Fig. 1) 
are horizontal magnesite adits excavated prior to the 
early 1970s. The entrances were gated in the early 
1990s with 15 cm grids preventing access by large 
animals, although inland cave bats are still able to 
negotiate the entrances. Weetootla Gorge Mine 2 was 
found to contain 36 inland cave bats, counted using an 
infrared video of the flyout at dusk (T. Reardon, pers. 
comm. 2003). The adit is approximately 90 m long 
and breaks into a natural rift at its termination. Bat 
roosts are located near the end of the adit, 50-70 m 
from the entrance. Small guano piles 1 m wide by 1-2 
m in length lie directly on the solid magnesite floor. 
Weetootla Gorge Mine 1, located 200 m downstream 
of Weetootla Gorge Mine 2, contained only two 
widely separated guano piles with no evidence of 
fresh guano. No bats were sighted inside this adit, 
suggesting the roost is used infrequently. 

The remainder of mines and caves examined were 
situated in the northern and central Flinders Ranges 
and are described from north to south (Fig. 1). Moro 
Bat Cave (F47), located in Moro Gorge, is 50 m above 
a permanent stream and extends into the cliff face 
terminating in the transition zone. The cave contains 
several bat guano deposits and dung of the yellow- 
footed rock wallaby (Petrogale xanthopus Gray) 
which uses the entrance area as a daytime retreat. The 
vertical slot entrance to McKinleys Daughters Cave 
(F175) located near stream level leads to a narrow 
high aven (terminal roof hole). A thin veneer of dry 
guano and numerous small mammal bones sit on a fine 


Proc. Linn. Soc. N.S.W., 126, 2005 


silt floor in the twilight zone. Unidentified bats were 
present high in the aven. Eregunda Mine, north-east of 
Blinman, is a 25 m long adit containing several guano 
deposits under an active roost of inland cave bats (for 
further details see Moulds, in press). Two unnamed 
caves, on the southern side of Chambers Gorge in 
a high valley, contain several inland cave bats and 
substantial desiccated guano in the transition zone 
chambers. Two caves near the Teamster’s Campsite in 
Brachina Gorge contain minor guano deposits mixed 
with fine soil. One of these is located at river level 
and the other approximately 40 m above the river. 
Two caves at the western end of Bunyeroo Gorge, 
near river level, are small with only one reaching the 
transition zone and the other only containing a twilight 
zone. Oraparinna Bat Cave, located north of Wilpena 
Pound, contained extensive amounts of guano within 
the primarily horizontal joint controlled passages. 


Guano microhabitat conditions 

Guano caves in the Flinders Ranges often have 
extremely low relative humidities and are commonly 
characterised by dry, acidic, pellet-like guano, even 
under active bat roosting areas that normally have 
moist, basic conditions (Harris 1970; Decu 1986; 
Gnaspini and Trajano 2000). Weetootla Gorge Mines 
1 and 2 had relative humidities less than 20% during 
September 2003. This has consequently affected the 
water content of guano deposits, which have been 
historically recorded from 3.3% (Arcoota Creek 
Cave) to 12.7% (Clara St. Dora Cave) (Winton 
1922), comparable to the driest guano found in Bat 
Cave, Naracoorte (Moulds 2003). Several artificial 
entrances in Oraparinna Bat Cave opened for guano 
mining have been the cause of desiccated guano piles 
near these entrances, limiting the distribution of some 
arthropods. Despite an active bat roost in Weetootla 
Gorge Mine 2, virtually no fresh guano was found. 
The guano beneath an active bat roost in Weetootla 
Gorge Mine 2 had a pH of 5.5. 


RESULTS 


Species recorded 

Twenty two arthropod species were collected from 
12 orders and two classes (Table 1). This represents a 
substantial increase from the single species previously 
recorded in the literature (Brises acuticornis) and the 
four unpublished species records of Elery Hamilton- 
Smith. Two sites, Eregunda Mine and Weetootla 
Gorge Mine 2, were extremely diverse with five and 
six orders recorded respectively. Active arthropod 
communities were found at all 12 sites, but not in 


127 


128 


Location 
Brachina 


Gorge 


Bunyeroo 
Gorge 


Chambers 
Gorge 


Mount 
McKinley 


Moro 


Gorge 


Oraparinna 


Point Well 


Weetootla 
Gorge 


SUBTERRANEAN GUANO ARTHROPODS 


Cave 
Unnamed 
river cave 
Unnamed 
hillside cave 


Unnamed cave 


Unnamed cave 


Unmaned 
cave no. 1 
Unnamed 


bat cave 


McKinleys 
Daughters 
Cave (F175) 
Moro Bat 
Cave (F47) 


Oraparinna 
Bat Cave 
(F8) 
Eregunda 
Mine 


Mine | 


Mine 2 


Order 
Coleoptera 
Neuroptera 


Neuroptera 


Hemiptera 
Hemiptera 


Neuroptera 
Zygentoma 
Araneae 
Neuroptera 
Orthroptera 
Diptera 
Hemiptera 
Neuroptera 
Lepidoptera 
Lepidoptera 
Neuroptera 
Coleoptera 
Coleoptera 
Coleoptera 
Araneae 
Coleoptera 
Hymenoptera 
Pseudo-scor- 
pionida 
Psocoptera 
Araneae 
Coleoptera 
Araneae 
Blattodea 
Coleoptera 
Hemiptera 
Neuroptera 


Orthroptera 


Family 
Anobiidae 
Myrmeleontidae 


Myrmeleontidae 


Reduviidae 
Reduviidae 


Myrmeleontidae 


Nicoletiidae 


Myrmeleontidae 
Gryllidae 


Reduviidae 
Myrmeleontidae 
Noctuidae 
Pyralidae 
Myrmeleontidae 
Anobiidae 
Anobiidae 
Tenebrionidae 
Pholcidae 
Tenebrionidae 
Formicidae 
Cheliferidae 


Pholcidae 
?Dermestidae 
Pholcidae 


Tenebrionidae 
Reduviidae 
Myrmeleontidae 


Gryllidae 


Genus 


Aeropteryx 


Aeropteryx 


Armstrongula 


Armstrongula 


Aeropteryx 


Trinemura 


Aeropteryx 


Armstrongula 
Aeropteryx 
Dasypodia 


Aeropteryx 


Ptinus 


Brises 


Brises 
Iridomyrmex 


Protochelifer 


Brises 
Armstrongula 


Aeropteryx 


Species 
sp | 
sp | 
sp | 


sp | 
sp 2 


sp | 

sp | 

sp 1 

sp | 

sp | 

sp | 

sp | 

sp | 
selenophora 
sp 1 

sp 1 
exulans? 

sp | 
acuticornis 
sp 1 
undetermined 


purpureus 


sp | 


sp 1 
sp 2 
sp | 
sp 3 
sp | 
caraboides 
sp 3 
sp 1 
sp | 


Table 1. Arthropods collected from guano deposits in the caves and mines of the Flinders 
Ranges. Caves are listed alphabetically by the area in which they are found. Some speci- 
mens could only be identified to subfamily or genus. 


Proc. Linn. Soc. N.S.W., 126, 2005 


T. MOULDS 


Pseudoscorpionida 
Pholcid spiders 


Psocoptera 


Anobiid beetle adult 
Anobiid beetle larvae 


Dipteran adult 


Dipteran larvae 


Gryllidae adult 
Gryllidae nymph Tenebrionidae adult 


enebrionidae larvae 


Blattodea adult 


Blattodea nymph 


Epigean flora 


Iridomyrmex purpureus| 


Figure 2. Food web of a Flinders Ranges guano community. Arrows 
represent the direction of energy flow within the food web. Guano 
ecosystems are extremely variable, consisting of numerous micro- 
habitats differentiated by moisture, pH and temperature. Fungi and 
bacteria are an important basis for guano ecosystems, providing us- 
able nutrients for many species unable to consume guano directly. 


very old guano deposits that were extremely dry and 
powdered. The beetle B. acuticornis duboulayi Bates 
(Tenebrionidae) was found in Oraparinna Bat Cave 
and an unidentified Brises larvae in Eregunda Mine. 
The second record of B. caraboides, from Weetootla 
Gorge Mine 2, greatly increases the distribution of this 
species, previously known only from the type locality 
near Eucla on the Nullarbor Plain. The emesine 
reduviid genus Armstrongula (Hemiptera) (Table 1) 
has a wide distribution in the Flinders Ranges, with 
three undescribed species recorded. Previously known 
species of Armstrongula are recorded from under bark 
near the Bogan River, New South Wales (Wygodzinsky 
1950). The presence of an unidentified Protochelifer 
species (Pseudoscorpionida) in Eregunda Mine in 


Proc. Linn. Soc. N.S.W., 126, 2005 


the central Flinders Ranges is an 
important intermediate record 
for this widespread, and often 
cavernicolous genus, between 
the Nullarbor Plain and _ the 
south-east of South Australia. 
The cosmopolitan beetle Ptinus 
exulans. Erichson (Anobiidae), 
and an unidentified anobiid, were 
also recorded from Oraparinna 
Bat Cave. 


Neuroptera larvae 


Food web and species 


interactions 
A food web for the Flinders 
Ranges hypogean guano 


communities is shown in Figure 
2. This was constructed using 
numerous direct field observations 
of many taxa combined with 
previously documented feeding 
biology of taxa from _ the 
literature. 

The trophic basis of all the 
discrete ecosystems examined 
is bat guano. Tenebrionid beetle 
adult and larvae (B. caraboides) 
and nymphal cockroaches were 
observed directly scavenging 
on guano deposits in Weetootla 
Gorge Mine 2. Tenebrionid beetles 
also act as general scavengers 
of organic material. Meat ants 
(Iridomyrmex purpureus Smith) 
were observed in the twilight zone 
of Eregunda Mine removing fresh 
guano and carrying it to their nest, 
possibly as a food source (Moulds, 
in press). The presence of ants 
adds a unique element to the species interactions at 
this locality by providing a potentially rich external 
food source for many of the predatory species such as 
pholcid spiders, reduviid bugs and neuropteran larvae. 
More commonly, guano deposits form a direct energy 
source for a succession of bacteria, yeast and fungi 
that support the majority of arthropods found in these 
environments (Fletcher 1975). Anobiid beetles have 
been recorded feeding on fungi and insect remains 
in guano deposits and spider webs (Richards 1971; 
Hickman 1974). 

Gryllid crickets shelter in caves during the 
day and feed on plant material growing near cave 
entrances when conditions are favourable, similar to 
rhaphidopohorid cave crickets (Richards 1961, 1965, 


2) 


SUBTERRANEAN GUANO ARTHROPODS 


1966). An individual was observed in September 
2003 feeding upon arthropod remains in an unnamed 
bat cave in Chambers Gorge. 

Myrmeleontid neuropteran larvae are common 
inhabitants of sandy floors in entrance and twilight 
zones, capturing small arthropods including ants, 
small beetles, fly larvae, and Psocoptera that fall 
into their conical pits. The presence of guano near 
neuropteran pits attracts additional prey for these 
sedentary predators, making guano deposits a 
beneficial habitat. In the larval stage, Neuroptera are 
part of guano food webs, but the adults play little 
role other than foraging for both plant and animal 
food in the epigean environment (New 1991). Adult 
Neuroptera were, however, commonly found during 
the day, sheltering in many of the caves examined 
(Table 1). 

Reduviid bugs of the subfamily Emesinae are 
common predators in many subterranean guano 
deposits, stalking arthropods in small groups (Moulds, 
unpublished data). Individuals were observed in 
September 2003 on the guano surface in McKinleys 
Daughters Cave and caves in Bunyeroo Gorge. These 
bugs form the top predator within the Flinders Ranges 
guano deposits. Prey were generally consumed where 
captured, although sometimes were dragged away 
from guano deposits for later consumption. 

A single cheliferid pseudoscorpion individual 
was also found in guano deposits. Pseudoscorpions 
spend most of the time under rocks, only emerging to 
hunt micro-arthropods. 


DISCUSSION 


Environmental limitations of population size 

The most limiting factors against the development 
of large guanophilic arthropod communities in the 
Flinders Ranges are low humidity and transient bat 
populations that limit guano sites in volume and 
continuity. Guanophilic communities commonly 
inhabit environments of saturated humidity with 
many species preferring the strongly basic conditions 
associated with fresh guano (Moulds 2003). Low 
humidity, common in the Flinders Ranges, severely 
reduce fungal growth as many of the opportunistic 
phycomycetes found on fresh guano are susceptible 
to desiccation (Poulson 1992; Poulson and Lavoie 
2000). Reduced growth of fungi, the primary food 
source of guanophilic communities, results in lower 
species abundance and diversity when compared with 
guanophilic communities in more humid locations 
such as coastal south-eastern Australia (Yen and 
Milledge 1990; Eberhard and Spate 1995; Bellati 


130 


et al. 2003). Further, the paucity of moist substrates 
removes key refugia for the numerous moisture- 
dependant species commonly found in guano caves. 
Consequently, families such as Jacobsonidae, 
Sciaridae and Sphaeroceridae are notably absent 
from the Flinders Ranges, and have been replaced 
by arid-adapted species such as tenebrionid beetles. 
Arid-adapted species comprise a substantial part 
of the species richness for caves across the entire 
region. The often stochastic semi-permanent bat 
colonies in the Flinders Ranges can have catastrophic 
consequences for guanophilic arthropod communities 
reliant on fresh moist guano for survival. 


Biogeography and dispersal mechanisms 

The guanophilic arthropod fauna of the Flinders 
Ranges shows closest similarity in species diversity 
to that of the Nullarbor Plain and isolated karst areas 
to the north and east. The tenebrionid beetles Brises 
acuticornis, B. caraboides and the carabid Thenarotes 
speluncarius Moore are found in both regions 
(Moore 1967; Richards 1971; Mathews 1986). Brises 
acuticornis is recorded from epigean and hypogean 
habitats and may use rabbit or wombat burrows for 
shelter during the day, aiding in long range dispersal 
ability (Mathews 1986). The major extension to the 
known distribution of B. caraboides from near Eucla 
on the Nullarbor Plain to the Gammon Ranges in the 
northern Flinders Ranges is significant as it provides 
additional species similarities between these two 
regions. 

The isolation of three species of Armstrongula 
within the Flinders Ranges suggests that increasing 
aridity through the region may have prevented the 
movement of hydrophilic cavernicolous species 
between karst areas. The disjunct karst formations 
of the region are also likely to have restricted the 
dispersal ability of other species. The occurrence and 
distribution of emesine reduviid bugs in guano caves, 
including those of the Flinders Ranges, is presently 
poorly understood but no records are known for this 
subfamily from the wetter southern karst areas of 
Australia. 

The relationship of the single Protochelifer 
(Pseudoscorpionida) collected from the Flinders 
Ranges (Eregunda Mine) to P. naracoortensis Beier 
from south-east South Australia and to P. cavernarum 
Beier from the Nullarbor Plain is unknown at present. 
It is unclear if this single record from the Flinders 
Ranges is indicative of a paucity of pseudoscorpions 
in general or simply a result of minimal collecting. 

The guano mite Uroobovella  coprophila 
Womersley, ubiquitous in southern Australian bat 
caves, is notably absent from the Flinders Ranges and 


Proc. Linn. Soc. N.S.W., 126, 2005 


T. MOULDS 


other arid localities (Moulds 2004). This is probably 
due to rapid desiccation of fresh guano, even beneath 
active bat roosts, excluding this mite as it relies 
upon fresh, moist, highly basic guano and virtually 
disappears during periods when these conditions 
are not available (Harris 1973). This species shows 
a strong association with breeding colonies of the 
large bent-wing bat, Miniopterus schreibersii (Kuhl), 
which does not occur in the Flinders Ranges or other 
arid regions (Churchill 1998). The similar distribution 
pattern of U. coprophila and M. schreibersii suggest 
this mite may be phoretic on bats or guano-associated 
invertebrates such as carabid beetles, although no 
observations have been reported. Phoresy requires 
further investigation to determine its importance in 
the distribution of this, and other, guano-associated 
species. 

The change in fauna composition moving south 
from the Flinders Ranges into the wetter costal areas 
of south-eastern Australia is marked, with several taxa 
such as the guano mite U. coprophila, histerid beetles 
and phorid flies becoming dominant on fresh guano. 
This study represents only a first step in documenting 
the diversity of cavernicolous guanophilic arthropods 
in the Flinders Ranges. Many previously known 
caves remain to be investigated and new caves are 
still being discovered. The transient nature of bat 
colonies and their relatively small numbers make the 
study of guanophilic fauna difficult. However, the 
region warrants further attention as it represents an 
important interface between the cavernicolous fauna 
of the Nullarbor Plain and the wetter coastal areas of 
south-eastern Australia. 


ACKNOWLEDGEMENTS 


This project was made possible by grants from The 
Nature Conservation Society of South Australia, The 
Wildlife Conservation Fund of South Australia, Department 
of Environment and Heritage South Australia, and The 
University of Adelaide. Terry Reardon, Matilda Thomas and 
Chris Grant provided extensive field assistance and Eddie 
Rubessa provided cave location data. This project would 
not have been possible without access to the Nepabunna 
Aboriginal lands, Flinders Ranges. Marta Kasper, John 
Jennings and Andy Austin are thanked for editorial 
comments. The comments by the two anonymous referees 
greatly improved this paper. 


Proc. Linn. Soc. N.S.W., 126, 2005 


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Proc. Linn. Soc. N.S.W., 126, 2005 


Song Analyses of Cicadas of the Genera Aleeta Moulds 
and 7ryella Moulds (Hemiptera: Cicadidae) 


M.S. Moulds 
Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010 
maxm(@austmus.gov.au 


Moulds, M.S. (2005). Song analyses of cicadas of the genera Aleeta Moulds and Tryella Moulds 
(Hemiptera: Cicadidae). Proceedings of the Linnean Society of New South Wales 126, 133-142. 


The songs of Aleeta curvicosta, Tryella castanea, T. crassa, T. kaumaand T. rubra are analysed. Differences 
between the songs of the five species are discussed in addition to differences across the distribution of 
species and within populations. Timbal action of all four Tryella species was found to be similar to that 
of Aleeta curvicosta, that is a single muscle contraction produces multiple sound pulses as each rib of the 


timbal buckles. 


A corrigenda to a recent review of the systematics of these genera (Moulds 2003) is provided. 


Manuscript received 18 October 2004, accepted for publication 10 January 2005. 


KEYWORDS: Aleeta, cicada, song, timbal action, Tryella. 


INTRODUCTION 


In a recent review of the genus Abricta Stal 
(Moulds 2003) I showed that the Australian species 
in that genus are best placed in two new genera, 
Abricta curvicosta (Germar) to Aleeta Moulds and 
the remaining species to Tryella Moulds. Aleeta and 
Tryella are very distinct from other Australian genera, 
the nearest being Chrysocicada Boulard. Adults 
characteristically perch head downwards and also 
call in that position (Fig. 1). They sing both during 
the heat of the day and at dusk, the dusk call seeming 
more vigorous. Timbal structure of A/eeta and Tryella 
is also very similar, there being many long parallel 
ribs (9-13 in Aleeta, 9-11 in Tryella) interspersed by 
very short ribs (compare figs 20, 22-29 in Moulds 
2003). 

Below I compare the songs of Aleeta curvicosta 
and four species of Zryella. In particular I discuss 
differences between the five species as well as 
differences across the distribution of 7. castanea 
(Distant) and T. crassa Moulds, and differences within 
a single population of 7: crassa. I also investigate the 
timbal contraction mechanisms of the four 7ryella 
species and show how these are similar to that of 
Aleeta curvicosta. 

Only the song of Aleeta curvicosta has been 
studied previously. Young (1972a) showed that 


A. curvicosta has neurogenic timbal muscles that 
contract alternately. He also described and illustrated 
the timbal, which is strongly and evenly ribbed so 
that during the inward movement each rib buckles 
separately producing 7-9 separate pulses of sound with 
each muscle contraction. A similar mechanism has 
been described for the periodical cicada Magicicada 
cassinii (Fisher) (Young & Josephson 1983b). Young 
calculated the pulse repetition frequency for the 
natural song of A. curvicosta as 1050/sec and the 
sound frequency range from 7.5-10.5 kHz. In his 
second paper concerning sound production in cicadas 
(Young 1972b), a slow speed oscillogram of the 
song of A. curvicosta was included, which clearly 
shows the spaced introductory phases characteristic 
of this species. This paper also included a sonogram 
of the song not represented in his first paper (Young 
1972a). 

Young and Josephson (1983a) further investigated 
timbal muscle contraction and mb buckling in A. 
curvicosta confirming the interpretation of Young 
(1972a). They calculated the muscle contraction 
frequency for the timbals as 72 Hz. Some brief 
additional data concerning the song of curvicosta have 
been provided by Ewart (1995) and Moulds (1990), 
but no further study of the calling mechanisms has 
been published. 


CICADA SONG ANALYSIS 


Figure 1. A male of Tryella crassa in full song. The 
body is raised above the substrate and the wings are 
held clear to prevent damping. 7ryella and Aleeta spe- 
cies normally sing with the head facing downwards. 


MATERIALS AND METHODS 


Calling songs were recorded in the field under 
natural conditions using an Akai X V portable tape 
recorder at a tape speed of 9.525 cm/sec and an AKG 
D19C dynamic microphone. Sound analysis data 
and oscillograms were generated using a Kay DSP 
Sonagraph Model 5500. 

Twenty three recordings were obtained in all, 
from nine localities, covering a total of four species 
of Tryella across seven localities in addition to a 
recording from Aleeta curvicosta (Table 1). 

Acoustic terminology follows that of Greenfield 
(2002). In particular, a pulse is here defined as “. . 
a brief packet of sound or vibrational waves that 
generally corresponds with a single repetitive action”. 
Pulse repetition frequency is defined as the number of 
pulses produced by the timbals each second and has 
been calculated from that part of the call containing 
uninterrupted contractions. 


134 


RESULTS 


Note: oscillograms and sonograms (Figs 
2-27) are at the end of the paper 


Song structure by species 

Aleeta curvicosta (Germar) 
recorded at Waverley Creek, Central 
Queensland (Figs 6, 19), showed song 
characteristics that agreed closely with 
data obtained from specimens from New 
South Wales (Young 1972a, 1972b, and 
Young & Josephson 1983a, 1983b). Pulse 
repetition frequency was measured at 1025/ 
s, very close to the 1050/s as measured by 
Young. Pulses were arranged in syllables as 
described by Young (1972a), who found that 
the syllables represented a single buckling 
of a timbal and each pulse the buckling of 
a single rib (Fig. 19). In the introductory 
part of each complete sequence, syllables 
were grouped into short echemes in which 
the amplitude rises then suddenly cuts off, 
with the interval between these echemes 
gradually decreasing until they merge 
into the continuous phase; this structure is 
identical to that described by Young (1972a, 
1972b). Young (1972a) also found that 
within the continuous phase, syllables were 
grouped into clusters of four, representing 
four alternating timbal contractions between 
the right and left timbals. This differs from 
groupings of three found in an individual 
from Waverley Creek (Fig. 19). Fig. 19c 
also shows the coalescence of the syllables produced 
by the alternate contractions of left and right timbals. 
Song carrier frequency was 6-12 kHz, also close to the 
6.5-11.5 kHz range given by Young. Young (1972b) 
described the song as having a rasping quality that is 
clearly more pronounced in this species than in the 
four Tryella species studied. 

Tryella castanea (Distant) had a continuous 
regular call that showed little variation in internal 
structure. Amplitude modulation was remarkably 
even overall, but there was strong internal modulation 
(Figs 3, 7a, 8a, 21a) with the song switching 
alternately every 0.25 s or so between phrases with 
syllables compressed and phrases where the syllables 
were separated by very brief (up to ca. 4 ms) gaps. 
The syllables are believed to represent single timbal 
buckles of multiple pulses (see ‘Timbal buckling’ 
below). The pulse repetition frequency was measured 
at approximately 750/s for the dusk call and 1050/ 
s for the day call. The song carrier frequency was 


Proc. Linn. Soc. N.S.W., 126, 2005 


M.S. MOULDS 


Table 1. Summary of song recordings of Aleeta curvicosta and Tryella species. 


Species Locality No. Date 
recorded recorded 
Aleeta curvicosta (Germar) QLD: Waverley Ck 1 23-Jan-1992 
Tryella castanea (Distant) NT: Dingo Ck 3 1-Jan-1992 
T. crassa Moulds NT: Adels Grove 8) 18-Dec-1991 
NT: Mataranka 5 11-Jan-1992 
QLD: W of Georgetown 8 16-Jan-1992 
T. kauma Moulds QLD: Walkers Ck 1 15-Jan-1992 
T. rubra (Goding & WA: Kununurra 8) 1-Jan-1992 
Froggatt) NT: Timber Ck 3 25-Dec-1991 
NT: Top Springs 1 24-Dec-1991 


concentrated between 8-13 kHz (Figs 21b, 22b, 23b) 
with weak side bands extending as low as 2 kHz and 
as high as 14 kHz (Figs 7b, 8b). To the human ear, 
the high concentrated frequencies are dominated by 
the lower side bands which, combined with the rapid 
but regular pulse repetition frequency, gives the calla 
vigorous rhythmic buzzing sound. 

Tryella crassa Moulds had a continuous, even 
call that showed little amplitude modulation (Figs 4, 
9-14, 20a). Syllables were coalesced into echemes of 
14-18 that were interspersed by two, or sometimes 
three, syllables only slightly separated from each 
other and the coalesced echemes (Figs 9-14, 25-27). 
The syllables consisted of a train of pulses inferred 
to result from a single timbal buckle (see ‘Timbal 
bucking’ below) (Fig. 20). The pulse repetition 
frequency ranged from around 1100/s during the heat 
of the day to near 850/s during the dusk chorus, but 
even within single populations noticeable variation 
was encountered (Figs 12-14). The song carrier 
frequency was concentrated between 7-12 kHz (Fig. 
20b) with an extreme range of approximately 4-15 
kHz. This high sound frequency was audible only as 
a hiss and the ear hears mainly the regular grouped 
pulse frequencies, which give the song a slight 
buzzing quality. 

Tryella kauma Moulds had a continuous regular 
song with slight amplitude modulation (Figs 2, 
24). Syllables of sound (timbal contractions) were 
arranged evenly through the call in discrete echemes 
(approximately 25-70 ms long) with very distinct 
inter-echeme gaps. The pulse repetition frequency 
was near 1120/s taken from a recording made during 
midmorning. The dominant song carrier frequency 
was concentrated between 8-13 kHz with weak side 
bands reaching 6 and 14 kHz. These high frequencies 


Proc. Linn. Soc. N.S.W., 126, 2005 


and the small size of this species give the song a quiet 
hiss-like quality. 

Tryella rubra Goding & Froggatt had a song 
distinctly divided into echemes of usually 8-10 
syllables (timbal contractions) that were coalesced 
together, each of similar amplitude (Figs 5, 15-18, 
25, 26). The length of these echemes varied within 
single populations and apparently also between 
populations (Figs 15-18). Sometimes the interval 
between echemes contained isolated syllables. The 
pulse repetition frequency ranged from 1220/s at 
Kununurra to 1280/s at Top Springs and Kununurra; 
all specimens were recorded during the heat of the 
day. The dominant song carrier frequency for all 
individuals was concentrated between 6-11 kHz (Figs 
25b, 26b, 27b) with very weak side bands extending 
from 1.5 kHz to 12 kHz. These high frequencies and 
the short repetitive echemes give the song a hiss-like 
buzzing quality. 


Song comparisons 

For all five species examined song characteristics 
clearly differentiate each (Figs 2-6). By far the 
most distinctive call was that of Aleeta curvicosta, 
primarily because of its unique introductory phrasing 
of discrete echemes (Fig. 6). In fact, the arrangement 
of echemes was the main component of song 
structure to show consistent and easily recognisable 
differences between all species. While there were 
some differences in echeme pattern both within and 
between populations of conspecifics (e.g. in Tryella 
crassa, Figs 9-11 and 12-14), oscillograms clearly 
showed these differences never approached the degree 
of difference shown intraspecifically (Figs 2-6). 

The characteristic regular pulse pattern of 
T. castanea was identical for two very different 


135 


CICADA SONG ANALYSIS 


phenotypes of this species from Dingo Creek that 
otherwise were associated only on the basis of male 
genitalia and allozymes (Moulds 2003) (Figs 7- 
8). Tryella castanea is one of the most variable of 
Tryella species in both pigmentation and size (see 
Moulds 2003, Figs 54a-e). The song of 7. kauma 
(Fig. 2) is similar to that of 7 castanea but differs in 
being a more vigorous call and having a much more 
even amplitude. The songs of 7: crassa and T. rubra 
are also similar to each other in that each consists 
of a regular succession of distinct echemes (Figs 
4, 5) and, like Aleeta curvicosta, the syllables from 
individual timbal contractions coalesce together; 
however the echemes differ structurally (crassa 
with 14-18 syllables per echeme, rubra with 8-10) 
and the echemes are separated by different interval 
structures. 

Pulse repetition frequency was remarkably 
similar for all five species. Day calls fell within the 
range 1050-1280/s but were probably dependent to 
some extent upon temperature. Populations of 7° 
crassa had pulse repetitions for day calls ranging 
from 1220-1280/s. Dusk calls, on the other hand, 
showed much lower pulse repetition frequencies, as 
low as 850/s. Dusk calling by 7: castanea showed a 
similar low pulse repetition frequency rate of 750/ 
Ss compared with a day call rate of 1050/s. These 
low pulse repetition rates for dusk calls are almost 
certainly a consequence of lower temperatures; very 
hot day temperatures raise the body temperature of 
day-calling individuals. 

Similarly, song frequency showed little 
intraspecific variation, the ranges concentrated 
between 6-13 kHz, but with all species individually 
showing a broad range of at least 5 kHz, thus making 
species diagnosis by frequency alone unreliable. 
However, frequency distribution patterns from 
sonograms did suggest that further investigation may 
show features characterising species. For example, T. 
castanea was unique in having weak, but nevertheless 
distinct, side bands extending to as low as 2 kHz (Fig. 
8b). 


Timbal buckling 

Analyses of the calling songs of these species also 
provided an opportunity to compare timbal buckling 
actions. The high pulse repetition frequencies of 7. 
castanea, T. crassa, T: kauma and T: rubra suggest 
timbal action equivalent to that detailed for A. 
curvicosta by Young (1972a), where each inward 
buckle of a timbal produced a train of discrete pulses 
caused by the individual buckling of ribs. 

For all five species studied, oscillogram pulses, 
when aligned against sonograms comprising only 


136 


the strongest frequency distributions, showed trains 
of pulses (syllables) corresponding with clusters 
of descending frequency. Extrapolating from the 
work of Young (1972a) for A. curvicosta, these are 
interpreted as individual rib buckles from a single 
inward buckling of the timbal, the frequency of 
each rib buckle falling progressively as the timbal 
collapses. Aleeta curvicosta from Waverley Creek 
showed six such pulses for each timbal buckle (Fig. 
19), compared with the 7-9 rib pulses range recorded 
by Young (1972a). Tryella crassa showed 7-8 pulses 
(Fig. 20), 7. castanea 6-7 (Figs 21-23), T. kauma 9 
(Fig. 24) and T. rubra 6-7 (Figs 26-27). 

While it is unlikely that the numbers I have 
recorded reflect a full range of the number of pulses 
resulting from single timbal buckles for each species, 
evidence suggests that multiple pulsing from a single 
timbal buckle does occur in all these species. In other 
words, for each of these species a single inward timbal 
buckle produces a very rapid pulse train that in turn 
leads to very high pulse repetition frequencies. 


DISCUSSION 


The male calling songs of the five species 
recorded, Aleeta curvicosta, Tryella castanea, T. 
crassa, T: kauma and T. rubra, each showed unique 
characteristics enabling clear separation of each 
species by song alone. This separation held true for the 
distinct morphs of T: castanea: the songs of the two 
very different morphs proved identical, confirming 
the association of these morphs previously derived 
from male genitalia and allozymes. 

The arrangement of the introductory phrases in 
the song of Aleeta curvicosta differed significantly 
from those of the four 7ryella species examined, 
reflecting their generic separation. More subtle 
differences (e.g. a concentration of intense pulse 
frequency between 6 and 11 kHz) probably account 
for the small difference in the perception of the call 
detectable to the human ear when compared to songs 
of Tryella species. However, there is also an overall 
similarity in song structure between all five species. 
All have similar high frequency ranges, similar high 
pulse repetition frequencies and, at least in part, 
continuous, regular buzzing sequences to their songs. 
These characteristics appear to characterise the songs 
of the Australian Aleeta and Tryella species. Further, 
a single timbal buckle in each of the five species 
appears to give rise to a succession of individual 
pulses as each rib buckles in succession rather than all 
ribs buckling in unison as in many other cicadas. It is 
this multiple pulsing from individual rib buckling that | 


Proc. Linn. Soc. N.S.W., 126, 2005 


M.S. MOULDS 


produces the very high pulse repetition frequencies 
for Aleeta and Tryella species. 


Corrigenda to Moulds (2003) 

p. 245, column 2, line 1: delete ‘be’ 

p. 272, key to species, couplet 3: “16’ should read ‘6’ 
p. 272, key to species, couplet 4, 3rd line: ‘no’ 
should read ‘not’ 


ACKNOWLEDGEMENTS 


Dr A. Ewart and Dr D.C. Marshall provided extensive 
discussion and comments on the manuscript, my wife 
Barbara and son Timothy gave assistance with field work 
and Dr Shane McEvey helped with preparing electronic 
images; to all I am most grateful. 


REFERENCES 


Ewart, A. (1995). Cicadas. In Ryan, M. (ed.), Wildlife of 
Greater Brisbane. Queensland Museum, Brisbane. 
Pp. 79-88. 

Greenfield, M.D. (2002). Signalers and receivers: 
mechanisms and evolution of arthropod 
communication. Oxford University Press, Oxford. 
414 pages. 

Moulds, M.S. (1990). Australian Cicadas. New South 
Wales University Press, Kensington. 217 pages, 24 
plates. 

Moulds, M.S. (2003). An appraisal of the cicadas of the 
genus Abricta Stal and allied genera (Hemiptera: 
Auchenorrhyncha: Cicadidae). Records of the 
Australian Museum 55, 245-304. 

Young, D. (1972a). Neuromuscular mechanism of 
sound production in Australian cicadas. Journal of 
Comparative Physiology 79, 343-362. 

Young, D. (1972b). Analysis of songs of some Australian 
cicadas (Homoptera: Cicadidae). Journal of the 
Australian Entomological Society 11: 237-243. 

Young, D. and Josephson, R.K. (1983a). Mechanisms of 
sound-production and muscle contraction kinetics 
in cicadas. Journal of Comparative Physiology 152, 
183-195. 

Young, D. and Josephson, R.K. (1983b). Pure-tone 
songs in cicadas with special reference to the genus 
Magicicada. Journal of Comparative Physiology 152, 
197-207. 


Proc. Linn. Soc. N.S.W., 126, 2005 137 


CICADA SONG ANALYSIS 


2 Tryella kauma 


Ae 
( 


Vi 
|, A | 
| i hy 
Walle Hidde wT hit ahy 
} 


3 Tryella castanea , 025s __, 


1s 
rr 


6 Aleeta curvicosta 


Figs 2-6. Slow oscillograms of the free song (male 
calling song) of Aleeta curvicosta and four Tryella 
species: (2) 7. kauma, Walkers Creek near Normanton, 
N. Queensland; (3) 7: castanea, Dingo Creek, western 
NT; (4) 7) crassa, Mataranka, NT; (5) 7: rubra, Top 
Springs, NT; (6) A. curvicosta, after Young and Joseph- 
son, 1983a. Note that figure 6 is at a different time scale. 


138 


7b 


0.25s 


Tye Punppneay ocr ey ji 


oe | 


| 


Hid, 
iil | i r bin ® ii uli nual Uldal Hiatt a4 


Figs 7-8. Tryella castanea; (a) synchronised oscillo- 
grams and (b) sonograms for two males from Dingo 
Creek, NT: (7) dark individual; (8) pale individual. 
(a) Waveform and (b) spectrographic analyses. 


Proc. Linn. Soc. N.S.W., 126, 2005 


M.S. MOULDS 


Figs 9-11. Tryella crassa; slow speed oscil- Figs 12-14. Tryella crassa; slow speed oscillo- 
lograms at identical time scale of day call for grams recorded during a 10-minute interval, at 
3 individuals recorded during a 35-minute in- identical time scale, of dusk call for 3 individuals 
terval from 40km W of Georgetown, Qld. from Mataranka, NT: (12) echemes widely sepa- 
rated by intermediate syllables (i.e. single timbal 
contractions); (13) echemes with moderate sepa- 
ration; (14) echemes with minimum separation. 


Proc. Linn. Soc. N.S.W., 126, 2005 139 


CICADA SONG ANALYSIS 


15 lt 3 timbal contractions 
Top Springs 19a 31.2 ms 


16 


Kununurra 


3 coalesced timbal contractions 


17 
Timber Ck 


specimen 1 


Fig. 19. Aleeta curvicosta; slow speed oscillogram of 
the (a) free song, (b) synchronised high speed oscillo- 
grams, and (c) sonogram of day call from an individual 


18 


Timber Ck from Waverley Creek, Central Queensland. (b) and (c) 
specimen 2 0.25s show a sequence of three timbal contractions showing 


the coalescence ofthesyllables from these contractions. 


Figs 15-18. Tryella rubra; oscillograms of day calls at 
identical time scales for 4 individuals: (15)Top Springs, 
NT; (16) Kununurra, WA; (17-18) Timber Creek, NT 


140 Proc. Linn. Soc. N.S.W., 126, 2005 


M.S. MOULDS 


AA (DN Ud 


1a Lone timbal contraction 


—————~ | =] one timbal contraction” 


15.6ms 


21b 


22a L___Jone timbal contraction 
—~ man — = 14 KHz 


Fig. 20. Tryella crassa; (a) synchronised oscillo- 
gram and (b) sonogram of dusk call for an individ- 
ual from Adels Grove, Lawn Hill Stn, Queensland. 


6 KHz 


Figs 21-23. Tryella castanea; (a) synchronised os- 
cillograms and (b) sonograms of day call for a sin- 
gle individual from Dingo Creek, NT: (21) a train 
of pulses; (22) portion of the same train of pulses at 
greatly increased time scale showing individual tim- 
bal buckles within a series of timbal contractions; 
(23) a single timbal contraction showing sound pulses 
generated from the separate buckling of seven ribs. 


Proc. Linn. Soc. N.S.W., 126, 2005 141 


CICADA SONG ANALYSIS 


a es nae one timbal contraction 


Fig. 24. Tryella kauma; (a) synchronised oscil- 
logram and (b) sonogram of day call for an in- 
dividual from Walkers Creek, Queensland. 


Figs 25-27. Tryella rubra; (a) synchronised oscil- 
lograms and (b) sonograms of a day call for a sin- 
gle individual from Top Springs, Queensland: (25) 
a train of pulses; (26) portion of the same train of 
pulses at greatly increased time scale, the sonogram 
showing positions of individual timbal buckles; (27) 
a sequence of three timbal contractions showing the 
coalescence of the syllables from these contractions. 


142 Proc. Linn. Soc. N.S.W., 126, 2005 


Llandovery (Early Silurian) Graptolites from the Quidong 
Basin, NSW 


R.B. Rickarps', R.A. PARKES? and A.J. WriGHT*4 


‘Department of Earth Sciences, University of Cambridge, Cambridge, England CB2 3EQ; 7MUCEP, 
School of Earth and Planetary Sciences, Macquarie University, N.S.W. 2109; *School of Earth and 
Environmental Sciences, University of Wollongong, N.S.W. 2522, and * Honorary Research Associate, 
MUCEP, School of Earth and Planetary Sciences, Macquarie University, N.S.W. 2109. 


Rickards, R.B., Parkes, R.A. and Wright, A.J. (2004). Llandovery (Early Silurian) graptolites from the 
Quidong Basin, NSW. Proceedings of the Linnean Society of New South Wales 126, 143-152. 


Late Llandovery (Early Silurian) graptolites from several localities in the Merriangaah Siltstone, Quidong 
Basin, southern NSW, are described as Monograptus priodon (Bronn, 1835), Oktavites falx (Suess, 1851) 
and Oktavites bodentoeriensis Loydell, 2003. This is the first use of the generic name Oktavites in Australia. 
The firm age for this fauna to the spiralis graptolite Biozone and new age data from the Quidong Limestone 
place a maximum age for the unconformity between the Siltstone and the overlying Quidong Limestone, 
constraining the Quidongan Orogeny between the latest part of the Llandovery and the late or latest 


Wenlock. 


Manuscript received 6 September 2004, accepted for publication 12 January 2005. 


KEY WORDS: Early Silurian, graptolites, Llandovery, Merriangaah Siltstone, Quidong. 


INTRODUCTION 


Graptolites were first reported from the Quidong 
Basin in southeastern NSW by Crook et al. (1973, p. 
116), who listed “Retiolites geinitzianus angustidens, 
Monograptus cf. auduncus and Monograptus of the 
priodon group” from the Merriangaah Siltstone, 
and inferred an age of “late Llandoverian to early 
Wenlockian” (middle of the Early Silurian). This 
material was found at Quidong by M. Tuckson (see 
Sherwin 1972) and the identifications cited by Crook 
et al. (1973) were by G.H. Packham. The Quidong 
Basin, 20 km N of Delegate (Fig. 1), is a farming 
region, although sulphides (copper—lead—zinc) in 
the carbonates of the area were mined in the 1860s, 
and the locality continues to be targeted as a mineral 
prospect (McQueen 1989). 

Here we describe graptolites collected by us in 
May 2004 from three localities in the Quidong Basin 
(Fig. 2). Re-collection of the fauna was necessitated 
by the almost complete disappearance of the original 
collection; three poorly preserved, indeterminate 
specimens exist in the Mining Museum collections, 
presumably representing salvage from the collections 
of the University of Sydney. 


This graptolite fauna is important because Crook 
et al. (1973) recognised the Quidongan Orogeny to 
account for what they considered an unconformity 
between the Merriangaah Siltstone and the overlying 
Quidong Limestone (Fig. 2). Scheibner (1972) 
originally introduced the term without definition. 
Crook et al. (1973, p.116) inferred that the Quidong 
Limestone was Ludlow in age, based on comparison 
of brachiopods in the mudstone conformably 
overlying the Quidong Limestone (the Delegate River 
Mudstone) to those in the “Ludlovian Silverdale 
Formation at Yass”. This age was consistent with 
the late Wenlock-early Ludlow age assigned to the 
limestone by Hill (1943, p. 58) based on the similarity 
of the Quidong rugose coral fauna to the rugose fauna 
at Yass from the Bowspring and Hume limestones of 
the Silverdale Formation. Packham (1969, p. 121) also 
concluded the Quidong Limestone was “Wenlockian 
to early Ludlovian”, comparing its diverse fauna of 
brachiopods, trilobites and corals with faunas from 
Hattons Corner, Yass. On this basis the orogeny was 
placed by Crook et al. (1973) within the Wenlock. 
Our studies of the Quidong Basin graptolites allow 
refinement of the age of this graptolite fauna; along 
with new data on the age of the Quidong Limestone, the 


SILURIAN GRAPTOLITES FROM THE QUIDONG BASIN 


Queensland 


a eee 
| Brisbane 


ee eee 
Victoria ‘ 


Tasmania 


|/ ~~ DELEGATE \ 


Delegate River Mudstone Buckleys Lake Fault 


Quidong Limestone Adamellite 
g Delegate Inferred fault 
Adamellite 


mid toLate 


Early Silurian Silurian 


UNCONFORMITY —— 
Yalmy Grou Thrust 


Merriangaah Siltstone (undifferentlate 
bere] Tombong Formation sandstone 


siltstone, shale) Inferred thrust 


2UNCONFORMITY 
ss Akuna Mudstone 


MVGiRIECaISHale Inferred geological boundary 


‘Bombala Beds’ 

(Adaminaby Group - : : om 
undifferentiated sediments, Fold axis syncline, anticline 
turbidites; sandstone, mudstone, 

shale) 


Geological boundary 


Late 


Cambalong Metamorphic 
Complex 


Ordovician Ordovician 


Early 


Figure 1. Location and geological setting of the Quidong Basin in the Tombong Block in southeastern 
New South Wales (modified after Lewis and Glen 1995, and McQueen 1989). 


144 Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, R.A. PARKES AND A.J. WRIGHT 


i mee stan em 
a Se i 


9853 Bis 


5918000 


sa 


eames 0% 
sn a : 


Recent 
Alluvium *K Graptolite sample location — — Drainage lines 
Tertiary 
Quartz pebble/ — = Fault === Inferred fault bas Shaft or pit 
cobble alluvium 
mid to Late Silurian —— Geological boundary = _iTrack 
Delegate River Mudstone 
! 2 ------ Inferred geological boundary [J Dam 
Quidong Limestone 
“UT SsCUNCONFORMITY h x Fold axis syncline, anticline Necdias Quarry 
Early Silurian . ‘ 
: ; i Strike and dip direction 
Merriangaah Siltstone of bedding 


Tombong Formation 


Figure 2. Geology of part of the northern area of the Quidong Basin. Sites sampled for graptolites 


indicated by W1015, W1016 and W1017. 


age of the hiatus can be more closely constrained. 

It is not our purpose here to discuss critically 
the whole matter of Early and Middle Silurian 
diastrophism in NSW. Crook et al. (1973) discussed 
the relationship of the tectonic history of the Quidong 
Basin area in relation to that of the Canberra and 


Proc. Linn. Soc. N.S.W., 126, 2005 


Orange districts. As two ofus (RBR, AJW) are engaged 
in studies with Gordon Packham of the graptolite-rich 
sequences at the Spring-Quarry Creek area, at Four 
Mile Creek and the Angullong district near Orange, 
it is premature to comment on the nature of breaks 
in the sequence that have been recognised there by 


145 


SILURIAN GRAPTOLITES FROM THE QUIDONG BASIN 


Packham (1969) and Jenkins (1978, 1986). Further, 
the nature of the Merriangaah Siltstone — Quidong 
Limestone contact is a topic being investigated by 
RAP, and is only briefly discussed here. 


GEOLOGICAL SETTING 


The Quidong Basin is a structural entity of 
approximately 25 sq. km representing the preserved 
remnants of a sedimentary basin comprising Mid to 
Late Silurian sediments that unconformably overlie 
a 500 m-thick Early Silurian quartzose turbidite 
pile. Collectively, these Silurian sediments occupy 
a fault-bounded, triangular area defined as the 
Tombong Block (Lewis and Glen 1995) that sits 
within Ordovician turbiditic sandstones and shales 
(‘Bombala Beds’ — Adaminaby Group) (Fig. 1). 
The Tombong Block forms part of the southernmost 
section of the Hill End—Cooma Zone, a meridional 
structural zone situated in the east of the Lachlan Fold 
Belt (Lewis and Glen 1995). 

The Early Silurian component of the Tombong 
Block comprises two units: the lower Tombong 
Formation and the higher Merriangaah Siltstone. 
The Tombong Formation occupies the bulk of the 
Tombong Block and consists of approximately 
400 m of quartz-rich sandstones and siltstones and 
interbedded shales. The presence of chert and slate 
pebbles in the Tombong Formation indicates that it 
was possibly derived from the Adaminaby Group 
(Lewis et al. 1994). Beds in the unit are generally 30 
cm to 50 cm thick, but range up to | m in thickness, 
and have lateral continuity equivalent to outcrop 
exposure. No fossils have been recovered from the 
Tombong Formation. 

The relationship between the Tombong 
Formation and the overlying Merriangaah Siltstone 
is conformable. The transitional boundary linking 
these formations is well-exposed on the northwest 
margin of the Quidong Basin (Fig. 2). Where the 
Merriangaah Siltstone is absent on the western and 
southern margins of the Basin (Fig. 1), the contact 
between the Tombong Formation and the overlying 
Quidong Limestone is disconformable. 

In the north of the Quidong Basin (Fig. 2) are 
extensive exposures of the Merriangaah Siltstone, 
which is estimated to be at least 80 m thick (Lewis et 
al. 1994, p. 35), and is composed of laminated beds of 
fine to very fine quartz sand intercalated with coarse 
quartz siltstone. The graptolite specimens in this 
study were recovered from beds composed of sand- 
sized grains. Bed thicknesses are mostly between 5 
cm and 15 cm; cross-laminations and ripple marks 


146 


are common. The trace fossils Paleodictyon isp. and 
?Gordia isp. were described from the unit (Webby 
1969). Our material was collected at three localities 
(Fig. 2), as follows: 


W1015: the riverside location illustrated by Crook et 
al. (1973, plate 2, fig. B), approximately 3-4 m 
below the unconformity. This locality has yielded 
a monospecific graptolite fauna of M. priodon; 

W1016: most westerly locality sampled, about 10 m 
below unconformity. This locality has yielded 
Oktavites bodentoeriensis; and 

W1017: 1-2 m below the unconformity first 
recognised by Herbert (1965) and Woodhill 
(1965). The fauna from here is Oktavites falx and 
M. priodon. 

The mid to Late Silurian fill of the Quidong Basin 
consists of the highly fossiliferous Quidong Limestone 
and the conformably overlying, erosionally-truncated 
Delegate River Mudstone. Conodonts recovered from 
the Quidong Limestone by one of us (RAP) indicate 
that the unit ranges from the late or latest Wenlock. 
The unconformity separating the Quidong Limestone 
from the underlying Merriangaah Siltstone is 
angular. 

During Honours studies at the University 
of Sydney, Woodhill (1965) and Herbert (1965) 
described the unconformity based on lithologic 
relationships and the angular contact between the 
Merriangaah Siltstone and the Quidong Limestone. 
The deformation indicated because of the angularity 
of the unconformity has been attributed to “inferred 
periods of compression” (Gray 1997, p. 149) that were 
a feature of the stabilization of the Lachlan Fold Belt 
between the end of the Ordovician and the Middle 
Devonian (Collins and Vernon 1992). However, 
cleavage trends within the Merriangaah Siltstone 
and the overlying mid to Late Silurian sediments are 
similar (RAP, unpublished studies), suggesting that, 
if the cleavage resulted from horizontal shortening, 
such deformation was subsequent to the termination 
of the second round of basin fill and was therefore 
not responsible for the angularity. An alternative 
hypothesis indicating that extensional rifting produced 
the angular unconformity has been suggested by 
Pickett (1982, p. 10) and Glen (1992, p. 373), the 
angular discordance being a manifestation of block 
rotation on listric normal faults that formed part of a 
new, or renewed, round of basin extension. 

Significantly, the late Llandovery start of the hiatus 
in sedimentation represented by the unconformity 
in the Quidong Basin is approximately coeval with 
the metamorphism of turbidites forming the Cooma 
Complex, based on an age for the metamorphism. 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, R.A. PARKES AND A.J. WRIGHT 


(43343 Ma) derived from detrital zircon and 
monazite by Williams (2001). The Cooma Complex 
is one of five fault-bound metamorphic complexes in 
the Eastern Metamorphic Belt (EMB) located in the 
southeastern part of the Lachlan Fold Belt (Johnson, 
1999, fig. 2); the Cambalong Complex, 6 km to the 
east of the Quidong Basin (Fig. 1), is another in 
this narrow (<50 km wide), generally meridionally- 
trending Belt. If the complexes comprising the EMB 
are similar in age and the metamorphism, at least in 
part, is due to compressional deformation (Johnson, 
1999, p. 440), then the Quidong Basin and Cambalong 
Complex represent juxtaposed coeval elements with 
contrasting structural styles separated by a thrust 
boundary (Fig. 1). 


SYSTEMATIC PALAEONTOLOGY 


Figured material is lodged in the Australian 
Museum (AMF). Three specimens from Quidong 
are held by the Geological Survey of NSW (MMF 
-18915-7) and are now lodged in the repository at 
Londonderry, NSW. 


Class Graptolithina Bronn 1849 
Order Graptoloidea Lapworth, 1875 
Family Monograptidae Lapworth, 1873 
Genus Monograptus Geinitz, 1852 


Type species 
Lomatoceras priodon Bronn, 1835; 


subsequently designated by Bassler (1915). 


Monograptus priodon (Bronn, 1835) 


Figures 3A-B 
1835  Lomatoceras priodon Bronn, p. 56, pl. 1, fig. 
13% 
1842 Gr Priodon; Geinitz, pp. 699-700, pl. 10, 
figs 16A-B. 
1850  Grapt. priodon. Bronn; Barrande, pp. 38-40, 


pl. 1, figs 3-9, 14, (non 1-2, 10, 11-13). 
Monograptus priodon (Bronn, 1835); 
Loydell, pp. 107-112, pl. 5, figs 2, 12; text- 
fig. 20, figs 4-5, 11, 26. 

Monograptus priodon (Bronn, 1835); Storch 
& Serpagli, pp. 42-43, pl. 9, figs ?3, 4-5, 
text-fig. 123A, ?H. 

Loydell (1993) synonymised well over one hundred 
records of M. priodon; of these, however, M. 
rickardsi Hutt, 1975 seems to us specifically distinct 
from M. priodon. Loydell demonstrated the very 
widespread record of M. priodon in late Llandovery 


1993 


1993 


Proc. Linn. Soc. N.S.W., 126, 2005 


and Wenlock strata. 


Material 

Numerous adult rhabdosomes were collected 
from localities W1015 and W1017. Specimens 
from W1017 are associated with Oktavites falx, are 
preserved in low relief with minimal pyritisation and 
are current-aligned. Specimens from locality W1015 
are rather weathered, pyritised adult specimens in full 
relief and are also current-aligned. At locality W1015 
no associated species was found. 


Description 

Monograptus thabdosome, at least 15 cm long, 
with a distal dorsoventral width of 2.8-3.0 mm in 
three-dimensional specimens; proximal observed 
dorsoventral width of rhabdosome 1.0 mm and 
observed thecal spacing 11-14 in 10 mm; distal thecal 
spacing 8-11 in 10 mm; thecal overlap ca '2; thecal 
hooks strongly retroverted with lateral processes (but 
not spines); sicula not seen. 


Remarks 

This material is on slabs covered with current- 
oriented specimens, but no proximal ends have been 
found, although some specimens probably end within 
10 mm of the sicula. Specimens from W1017 are 
extremely well-preserved, agree with many other 
descriptions of the species and are typical of those 
found in the late Llandovery. 


Genus Oktavites Levina, 1928 


Type species 

Graptolithus spiralis Geinitz, 1842, subsequently 
designated by Obut (1964), from the Llandovery of 
Germany. 


Remarks 

Oktavites was not recognised in the 1970 edition 
of the graptolite Treatise, being considered a junior 
synonym of Monograptus (Bulman 1970, p. V132) 
for reasons given by Bulman and Rickards (in Bulman 
1970, p. V132). However, the thecal structure of 
Oktavites spiralis has long been quite well-known; 
Loydell (1993) effectively redefined Oktavites in 
modern terminology and, at the same time, drew a 
contrast with species of Spirograptus, including the 
type species S. turriculatus (Barrande, 1850). Thus 
Oktavites has broadly triangular thecae with the thecal 
apertures laterally expanded, whereas Spirograptus 
has 1 or 2 apertural spines, sometimes apertural 
symmetry, but usually little lateral expansion. The 


147 


SILURIAN GRAPTOLITES FROM THE QUIDONG BASIN 


Figure 3. (a-b) Monograptus priodon (Bronn), respectively AM F 123128, 123129 from locality 
W1017, preserved in moderate relief, specimen a being quite close to the proximal end, specimen b 
of mesial thecae; scale bar 1 mm. (c-l) Oktavites bodentoeriensis Loydell, respectively AM F123120, 
123114, 123116, 123119, 123121, 123118, 123115, 123113, 123124 and 123117, all from locality 
W1016; scale bars of c-h, k-l are 0.10 mm; scale bars of i-j are 1 mm. (m) Oktavites falx (Suess), 
AM  F123125, from locality W1017; heavy bar indicates a lineation on the bedding plane that 
may be soft sediment deformation rather than tectonic deformation; scale bar of m is 1 mm. 


148 Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, R.A. PARKES AND A.J. WRIGHT 


generic name Oktavites has not been previously 
used for Australian graptolites, although O. spiralis 
was recorded from the Melbourne Trough, Victoria 
as Monograptus spiralis (see Rickards and Sandford 
1998). Neither Oktavites falx nor O. bodentoeriensis 
(see below) has previously been recorded from 
Australia. 


Oktavites falx (Suess, 1851) 


Figure 3m 

1851  Graptolithus falx n. sp.; Suess, p. 119, figs 
10a-b. 

1945  Spirograptus falx (Suess, 1851); Pribyl, pp. 
11-32, pl. 5, figs 1-6. 

21990 Oktavites falx (Suess); Ge, pp. 152-153, pl. 
64, figs 3, 6, 9. 

1993 Oktavites? falx (Suess, 1851); Loydell and 
Cave, figs 8k-n. 

21994 Monograptus aff. falx (Suess, 1851); 
Zalasiewicz and Tunnicliffe, text-fig. 8A-B. 

1998  Oktavites? falx (Suess, 1851); Storch, pp. 

124-5, text-fig. 3, figs 11-13. 

2003 Oktavites falx (Suess, 1851); Loydell, pp. 
59-60, text-fig. 1, figs 11-12. 

Lectotype 


The specimen figured by Suess (1851, pl. 9, 
fig. 10a) from the spiralis zone of the Litohlavy 
Formation, Praha-Mala Chuchle, Bohemia, was 
designated by Pribyl (1945). 


Material 

Two specimens from locality W1017, AM 
F123125a-b and AM F123126a-b; the former is on a 
bedding plane with many well-preserved specimens 
of M. priodon. Specimens are more or less flattened 
except for the proximal end, which is in low relief and 
well preserved. 


Description 

Oktavites with low-angled triangular thecae, 
reaching 13-9 in 10 mm; proximal dorsoventral width 
0.4-0.5 mm, distally about 1 mm; thecal apertures 
with small lateral expansion and suggestions in places 
of tiny spines; dorsal wall strongly recurved; thecal 
height at thl (hence dorsoventral width at same point) 
0.4-0.5 mm; distal thecae inclined at a lower angle 
and less triangular than proximal ones (20° down to 
10°); thecal overlap slight; rhabdosome with some 
spiral coiling beginning around th7; sicula 1.2 mm 
long, apex to just above level of hook of thl; origin 
of thl halfway from sicular aperture; sicular aperture 
simple; virgella short, slim spine. 


Proc. Linn. Soc. N.S.W., 126, 2005 


Remarks 

The proximal thecae of O. falx are similar to 
those of O. bodentoeriensis from Quidong but are 
essentially smaller, much more closely spaced (13-12 
in 10 mm compared with 7% -8 in 10 mm) and with 
a lower metathecal height. Storch’s (1998) specimens 
from Spain are very close to the Quidong specimens, 
perhaps beginning their spiral coiling a little later 
(ca thl0-15, rather than ca th7), but otherwise 
having the same dimensions and measurement. The 
stratigraphically earlier forms illustrated by Loydell 
(2003) have stronger rhabdosomal coiling but have 
exactly the same proximal end as the Quidong 
specimens. 


Oktavites bodentoeriensis Loydell, 2003 


Figs 3C-L 
2003 Oktavites bodentoeriensis sp. nov.; Loydell, 
p. 60, text-fig. 1, figs 14-17; text-fig. 3. 
Holotype 


Specimen figured by Loydell (2003, text-fig. 
1, fig 15) from the lower spiralis Biozone of the 
Rauchkofel Bodentéri section, Carnic Alps, Austria. 


Material 

Two adult specimens, AM F123113 and 123115, 
and eight early growth stages, AM F123114, 123116- 
121, 123124 all from locality 1016. Two further, 
poorly-preserved possible early growth stages from 
the same locality, AMF 123122-3. 


Diagnosis 

Oktavites with some rhabdosomal coiling 
beginning ca thl2; prosicula 0.16-0.28 mm long; 
sicula 1.2-1.5 mm long; virgella short and fine. 
Proximal dorsoventral width of rhabdosome 0.6-0.8; 
distal dorsoventral width 0.75-0.85 mm. Proximal 
thecal spacing 7 in 10 mm; distal thecal spacing 7.5-8 
in 10 mm; thecal overlap very low (diagnosis modified 
after Loydell on the basis of our new material). 


Description 

Of the two adult rhabdosomes AM F23113 (Fig. 
3J) shows no twisting of the stipe after th7, whereas 
AM F123115 (Fig. 31) begins to twist at around th10- 
11, so an open spiral coiling of rhabdosome can be 
predicted. 

Prosicula well seen on several specimens, 
occuring as isolated specimens on bedding planes. 
Several longitudinal spiral strengthening threads 
visible (Fig. 3C) and these may coalesce to form 
nema, a fine thread up to 0.7 mm long and commonly 


149 


SILURIAN GRAPTOLITES FROM THE QUIDONG BASIN 


seen on early growth stages. There seems to be a slight 
constriction at the origin of the metasicula (Figs 3d, 
g, k). When complete, the sicula is 1.2-1.5 mm long 
and its apex is invariably above the level of the hook 
of thl. 

Origin of thl very low on metasicula, perhaps 0. 1- 
0.15 mm above sicular aperture, which has a diameter 
of up to 0.12 mm. Th1 completed before protheca of 
th2 begins (Figs 3e, k). Protheca of th2 very slim 
(0.1 mm), and expands only slightly as metatheca 
is approached. A marked change occurs in angle of 
free ventral wall with onset of metatheca (a change 
from 5° to 20-40°) and metatheca is quite high (giving 
the full dorsoventral width). Thecal hook occupies 
about 1/3 of metathecal height. Overall thecal profile 
axially elongate-triangular, with prominent hook 
showing no sign of torsion; there is some indication 
of apertural expansion and there are tiny thecal spines 
(Fig 31, thS and th6 of visible thecae). Central part 
of metathecal hook strongly retroverted, facing dorsal 
side of rhabdosome. 


Remarks 

The nature of the thecal hook confirms Loydell’s 
(2003) attribution of this species to Oktavites 
rather than Spirograptus (which has less transverse 
expansion of the thecal aperture) or Jorquigraptus 
(which shows thecal torsion of the metathecal axis). 
The Quidong specimens are very close to Loydell’s 
originals from the Carnic Alps and differ only in 
having slightly more widely-spaced thecae (7.5-8 in 
10 mm compared with 8-10 in 10 mm). The Quidong 
specimens give a fuller idea of the early development, 
which is not well known in species of Oktavites 
other than the type species O. spiralis. Oktavites 
bodentoeriensis differs from O. fa/x in the same part 
of the rhabdosome, in having a more robust proximal 
end and different thecal spacing. 


BIOZONAL AGE OF THE QUIDONG BASIN 
GRAPTOLITES 


The age indicated for the assemblage is probably 
early spiralis Biozone (in old terminology early to 
mid crenulata Biozone: Llandovery, Early Silurian). 
Monograptus priodon has a long time range, 
possibly appearing (Loydell 1993) in the upper part 
of the turriculatus Biozone (earliest Telychian=late 
Llandovery), but certainly is common from the 
griestoniensis Biozone (Telychian) to the early middle 
Wenlock. However, both O. falx and O. bodentoeriensis 
are restricted to the Llandovery; the former appears 
in the early spiralis Biozone (more coiled forms) 


150 


and ranges into the upper spiralis Biozone (almost 
uncoiled specimens), whereas the latter was recorded 
from the early spiralis Biozone of the Carnic Alps 
by Loydell (2003). This seems the most likely level 
in the spiralis Biozone for the Quidong material. 

Crook et al. (1973) listed the following graptolites 
from Quidong: Retiolites geinitzianus angustidens 
Elles and Wood, Monograptus aduncus Bouéek and 
Monograptus ex gr. priodon (Bronn), stating that they 
were identified by G.H. Packham. It was suggested 
that the last of these named forms was similar to 
M. parapriodon Bouéek because of the narrow 
thabdosome and high thecal spacing; if so, it is 
different from the specimens of M. priodon described 
here but has broadly the same age implication, as 
Monograptus parapriodon occurs in the crenulata 
Biozone. Retiolites g. angustidens ranges from the 
crispus Biozone to the early Wenlock. Monograptus 
aduncus is now referred to Monoclimacis and is 
from the early Wenlock rather the late Llandovery; 
however, we would need to re-examine this 
material given the improvements in recent years 
of our understanding of Monoclimacis. We have 
been unable to locate these specimens but the ages 
indicated are not in dispute with our more precise age 
of early spiralis Biozone, except for the identification 
of Monoclimacis aduncus. With the help of Dr Ian 
Percival and Dr Lawrence Sherwin we were able 
to examine three specimens (MMF 18915-7) in the 
collections of the Geological Survey of NSW (now 
held in the NSW State Palaeontological Reference 
Collections at the Geological Survey of NSW 
Geoscience Centre, Londonderry); as no retiolitids 
were present, part of the original collection is missing. 

We have also been unable to check the 
Pickett (1982) record of M. sedgwickii Portlock 
but, as we remark in the following description 
of M. priodon, there is a preservational view of 
M. priodon that can appear superficially like M. 
sedgwickii; however, even under these circumstances 
the two have a quite different thecal overlap. 


ACKNOWLEDGEMENTS 


We are grateful to property owners James and Marlene 
Adams of ‘Canigou’ and Buddy Stevenson of ‘Quidong’ 
for allowing us access to their properties for specimen 
collection at localities in the Quidong Basin. Mandy Lyne 
kindly assisted with collecting. Dr Jan Percival and Dr 
Lawrence Sherwin helped us to locate early collections 
of Quidong graptolites. RBR acknowledges financial 
support from the Royal Society. We thanks two anonymous 
reviewers for their helpful comments, including the point 
about the Sardine Graben. 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, R.A. PARKES AND A.J. WRIGHT 


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152 Proc. Linn. Soc. N.S.W., 126, 2005 


Silurian graptolites from the Barnby Hills Shale and Hanover 
Formation, New South Wales 


R.B. Rickarps!, J.R. FARRELL?, A.J. WricuTt? and E.J. MorGcaAn* 


‘Department of Earth Sciences, University of Cambridge, Cambridge, England CB2 3EQ; *School of 
Education, Macquarie University, N.S.W. 2109; *School of Earth and Environmental Sciences, University 
of Wollongong, N.S.W. 2522; and Research Associate, MUCEP, School of Earth and Planetary Sciences, 

Macquarie University, N.S.W. 2109; and *Arundell Geoscience, P.O. Box 170, Belmont 6984, Western 
Australia (formerly Geological Survey of N.S.W., Department of Mineral Resources, P.O. Box 53, Orange 


N.S.W. 2800). 


Rickards, R.B., Farrell, J.R., Wright, A.J. and Arundell, E.J. (2005). Silurian graptolites from the Barnby 
Hills Shale and Hanover Formation, New South Wales. Proceedings of the Linnean Society of New 
South Wales 126, 153-169. 


Additional collections of graptolites from the Barnby Hills Shale and new collections of graptolites 
from the Hanover Formation in the Lachlan Fold Belt of central western NSW are documented. The Late 
Silurian Hanover Formation is shown to range from the spineus Biozone (late Ludlow) to the parultimus 
Biozone (Pridoli). A fauna containing Monograptus ludensis is recorded from the Barnby Hills Shale, 
which is now known to range from the /udensis Biozone (late Wenlock) to the inexspectatus or kozlowskii 
Biozone (late Ludlow). New dendroid graptolite taxa described here include Dendrograptus typhlops sp. 
nov. from the Barnby Hills Shale and Dictyonema paululum hanoverense subsp. nov. from the Hanover 
Formation. Monograptus spineus, from the Hanover Formation, is reported for the first time outside Europe. 
The new data confirm that strata assigned to the cornutus and praecornutus biozones (late Ludlow) are 
widely distributed in central NSW, and confirm previous suggestions for a latest Ludlow sea level highstand 


followed by a shallowing. 


Manuscript received 9 September 2004, accepted for publication 10 January 2005. 


KEY WORDS: Barnby Hill Shale, graptolites, Hanover Formation, Ludlow, Pridoli, Silurian. 


INTRODUCTION 


A diverse Late Silurian (late Ludlow: 
inexspectatus or kozlowskii Biozones) graptolite 
fauna was described by Rickards and Wright (1997a) 
from the Barnby Hills Shale at Neurea, NSW (Fig. 
1). In this paper we document further biostratigraphic 
control for this region of NSW by describing 
additional Silurian graptolites from the Barnby Hills 
Shale from various localities, and documenting the 
first Silurian graptolites from the Hanover Formation 
in the vicinity of Cumnock in central western NSW 
(Fig. 1). These new collections provide definitive age 
controls for the latter formation, in particular. 


GEOLOGICAL BACKGROUND 


The fossils described here are from two largely 
Late Silurian formations located in the northern part 


of the Lachlan Fold Belt, New South Wales, in the 
region from east of Orange to south of Wellington 
(Fig. 1). The Barnby Hills Shale is part of the Early 
Silurian to earliest Devonian Mumbil Group, a 
carbonate - volcanic - fine-grained clastic sequence, 
which was deposited on the Mumbil Shelf. The 
Hanover Formation was deposited in the Cowra 
Trough, a marine basin to the west of the Mumbil 
Shelf, and forms part of the Early Silurian to earliest 
Devonian Cudal Group (Meakin and Morgan 1999). 


Barnby Hills Shale 

Strusz (1960) introduced the term Barnby Hills 
Shale Member for the upper unit of his “‘Mumbil 
Formation’, and Vandyke and Bymes (1976) 
subsequently raised the unit to formation status; the 
Mumbil Group was established by Pickett (1982). 
Discussions of the formation were given by Morgan 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


Key for Figs 1 & 2 


volcanilitharenite and volcanilithrudite 
and limestone blocks (L). 


- fe] Soil cover 
3 Hill wash, derived from 
§ Catombal Group 
° 
= Ferruginized deposits 
& 
a2 
5 § Catombal Group 
ray 
i a | Dgg_ Garra Limestone 
Gees ?Mungallala Member - shale and 
§ Ea conglomerate 
o : 
3 Deg Cuga Burga Voicanics - 
= 
oO 


Camelford Limestone 


BABY on 
er 


Barnby Hills Shale - shale, chert, 
meta-dolerite, rhyolite, Ordovician 
limestone blocks (OL) and isolated 
blocks of Camelford Limestone (CL). 


Narragal Limestone 


Fairbridge Volcanics 


Geological boundary 
aia ee ss Inferred geological boundary 


eae Fault 
— Thrust fault 

— Inferred fault line 

Sie Dip, strike and facing 
<R- Ephemeral creek with dam, creek 
= Road and track 

68KiL Graptolite locality 

& Quarry 


Figure 1. Simplified geology of the area between Wellington and Cumnock (NSW) (modified after 
Talent and Mawson 1999) showing major Silurian and Early Devonian carbonate units and associated 
shale sequences along the Mumbil Shelf, and showing location of Fig. 2. 


(in Pogson and Wyborn 1994; in Meakin and Morgan 
1999). 

Talent and Mawson (1999) and Cockle (1999) 
advocated the relegation of this unit toa junior synonym 
of the Wallace Shale, a formation erected for strata in 
the Four Mile Creek area, SSE of Orange (Stevens and 
Packham 1953) but which also occurs in the Spring- 
Quarry Creek area (Packham and Stevens 1955) west 
of Orange. The Wallace Shale exhibits distinctive red, 
green and yellow banding, and is easily distinguished 
lithologically from the Barnby Hills Shale. The age 


154 


of the Barnby Hills Shale is late Wenlock to late 
Ludlow, based on earlier data (Rickards and Wright 
1997a) as well as data presented here. The only age- 
diagnostic fossils described from the Wallace Shale 
are the Pridoli graptolites described by Sherwin and 
Rickards (2002) and, probably, the Pridoli graptolite 
fauna from near Cadia described by Rickards et al. 
(2001); the deeply weathered graptolitic strata at the 
latter locality cannot be assigned with certainty to the 
Wallace Shale on the basis of their lithology. There 
is insufficient reason, we believe, to synonymise and 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


discard this very distinctive stratigraphic unit (see 
also Zhen et al. 2003). 

The formation crops out mainly in four meridional 
fault slices (Morgan et al. 1999) and extends from the 
town of Molong in the south to north of Wellington 
in the north, a strike distance of almost 80 km. The 
bulk of the fauna described here from the Barnby 
Hills Shale is from the western belt that extends from 
‘The Gap’ to north of Eurimbla (Figs 1-2). Farrell 
(1992; in Talent 1995) recognised its Late Silurian 
age in the western belt, and also identified a number 
of interbedded limestones of Silurian and Ordovician 
ages, which had been earlier noted by Byrnes (in 
Pickett 1982, fig. 19). Dating of graptolites from 
shales enclosing Ordovician limestone outcrops in 
the northern section by Sherwin (1994a, 1997) as 
Silurian proved that the limestone is allochthonous 
and that the shales correlate with the Barnby Hills 
Shale (Zhen et al. 2003). 

The western belt is faulted at its base against 
the Late Devonian Catombal Group (Fig. 2). In the 
south, the formation is conformably overlain by the 
- Siluro-Devonian Camelford Limestone but, to the 
north, it is faulted against the early Devonian Cuga 
Burga Volcanics. The Silurian limestone outcrops in 
the southern section are Pridoli and are thought to 
be remnants of the lower horizons of the Camelford 
Limestone ‘grounded’ along the Curra Creek 
Thrust (Farrell 2001). To the east, the Barnby Hills 
Shale forms part of a continuous Siluro-Devonian 
sequence. 

In the vicinity of the original road cutting type 
section just east of the Mitchell Highway at Neurea 
(see Rickards and Wright 1997a, and Fig. | herein) 
the formation conformably overlies the Narragal 
Limestone. This road cutting, which yielded the 
material described by Rickards and Wright (1997a), 
has been, regrettably, over-collected and the now 
almost barren exposures have been also severely 
damaged by large-scale removal of rock for other, 
non-geological purposes. Morgan (in Meakin and 
Morgan 1999) nominated a new type section for the 
formation along a railway cutting at Dripstone (Fig. 1) 
where exposures are good and formation boundaries 
are exposed. Unfortunately graptolites are extremely 
rare in this section: on our first visit we found rare 
graptolites identified in the field as Bohemograptus, 
Linograptus and Dictyonema, but these specimens 
have been mislaid; no more material was collected on 
a second visit to the section. In this type section the 
formation is 290 m thick; in the Eurimbla area, the 
western belt appears to reach a thickness of over 700 
m but the unit may be internally folded (Morgan in 
Meakin and Morgan 1999). 


Proc. Linn. Soc. N.S.W., 126, 2005 


We also describe and illustrate poorly-preserved 
graptolites from this formation near Lewis Ponds; this 
locality, 15 km northeast of Orange, was described 
as the Mullions Reserve by Meakin in Pogson and 
Wyborn (1994). This material was first reported by 
Sherwin (1993) as “Monograptus bohemicus subsp., 
with siculae of hercynicus type”, and the occurrence 
was further documented by Meakin (in Pogson and 
Wyborn 1994). All the abundant graptolites (Fig. 
5D-E) belong to a monospecific assemblage of 
Bohemograptus paracornutus Rickards and Wright, 
1999b, are tectonically deformed, and occur in black 
cleaved pyritic slates; this is the southernmost known 
fossiliferous development of the formation. 


Hanover Formation 

The term Hanover Formation was introduced by 
Maggs (1963) and first published by Offenberg et al. 
(1971). Bradley (in Pickett 1982, fig. 14) included the 
formation in the Cudal Group. Morgan (in Meakin 
and Morgan 1999) found that the unit was much more 
widespread than shown by previous workers and 
provided a thorough discussion of the formation. The 
Hanover Formation crops out in a number of complex 
meridional fold and fault repetitions from Cumnock 
to Geurie (Morgan et al. 1999). The fossils described 
here were found in the easternmost block of Hanover 
Formation, in a rail cutting east of Cumnock (Fig. 1). 
Previous workers considered the rocks in the cutting 
to be part of a Devonian package (the ‘Carinya Shale’ 
of Maggs [1963] and Offenberg et al. [1971]), but 
Morgan (in Meakin and Morgan 1999) determined a 
Silurian age and correlated the rocks with the Hanover 
Formation. 

The Hanover Formation has few diagnostic 
features and closely resembles the Barnby Hills 
Shale in lithology, as well as a number of other shale- 
siltstone dominated facies in the area. The formation 
conformably overlies the Wenlock Canowindra 
Volcanics and is overlain either by the Early Devonian 
Cuga Burga Volcanics or their lateral equivalent, the 
Berkley Formation. In the study area the formation 
is faulted at its base against the Cudal Fault, and is 
internally complexly folded and faulted. No one 
section has yielded both top and base of the Hanover 
Formation, so its total thickness is not known; Bradley 
(in Pickett 1982) estimated a thickness of 300 m. 


Lithological Description 

The Barnby Hills Shale and the Hanover 
Formation consist dominantly of poorly-outcropping 
siltstone and shale, with subordinate interbedded fine- 
to coarse-grained volcaniclastic sandstone, radiolarian 
chert, marly siltstone and detrital calcareous horizons 


155 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


Figure 2. Simplified geological map showing graptolite localities in the area between The Gap and 


Larras Lee. 


(Morgan in Meakin and Morgan 1999; Morgan 
in Meakin and Morgan 1999). The formations are 
characterised by poorly- to well-bedded and finely- 
laminated buff siltstone and shale, and vary from 


156 


brown to grey to green to red. Beds are in the order 
of 1 mm to 2 cm in thickness. Laminations exhibit 
slump, scour, grading, flame and other sedimentary 
structures, and bioturbation is common (Morgan in 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


Meakin and Morgan 1999; Morgan et al. in Meakin 
and Morgan 1999). 

Numerous detrital limestone beds are present 
within both formations, including allochthonous 
limestone blocks up to 180 m long. The beds are 
commonly highly fossiliferous and are of varying 
ages. In the Hanover Formation, a massive to well- 
bedded fossiliferous detrital limestone bed is present 
100 macross strike to the west of the graptolite locality, 
for which Percival (1998) determined a probable late 
Llandovery age. In the Barnby Hills Shale, large 
allochthonous limestone blocks containing Late 
Ordovician fossils (Zhen et al. 2003, Webby 1969) lie 
adjacent to one of the graptolite localities (68 KIL on 
Figure 2A). In contrast, a large limestone block at the 
southern end of the western belt of the Barnby Hills 
Shale contains Late Silurian fossils (Farrell 2001). 


PALAEONTOLOGY AND AGE 


Barnby Hills Shale 

Graptolites from the Barnby Hills Shale were 
first recorded by Strusz (1960) as Monograptus 
bohemicus. Sherwin (1993, 1994b) identified the 
common species in the fauna as an undetermined 
subspecies of Bohemograptus bohemicus, indicating 
a middle to late Ludlow age; a late Ludlow age 
was suggested by Sherwin (1997). Sherwin (1994a, 
1997) identified Saetograptus colonus from shale 
in the KIL study area in the western belt, indicating 
an early Ludlow age (basal ni/ssoni Biozone to part 
way through the scanicus Biozone). No graptolites 
were described or illustrated until Rickards and 
Wright (1997a) described the substantial late 
Ludlow (inexspectatus or kozlowskii Biozones) 
fauna of dendroids and graptoloids including the 
age-diagnostic Bohemograptus bohemicus tenuis 
(Boucek 1936) from the belt at Neurea (Fig.1). The 
Barnby Hills Shale is locally conformably overlain 
by the Pridoli to Lochkovian Camelford Limestone 
(Farrell 1992, 2003). 

Meakin (in Pogson and Wyborn 1994, p. 105) 
drew attention to the occurrence of this formation 
at South Mullion Reserve, ENE of Orange. The 
occurrence at this locality (W859) of poorly preserved 
Bohemograptus paracornutus (Figs 5D-E) indicates 
the cornutus Biozone. 

The graptolite faunas described here from this 
formation are thus from four localities (Figs 1-2): 


68 KIL (Fig. 2): ‘Kildara’ property; /udensis Biozone, 


late Wenlock, Barnby Hills Shale; Monograptus 
ludensis (Murchison, 1839) and indeterminate 


Proc. Linn. Soc. N.S.W., 126, 2005 


retiolitids. 

Blind Gully (WEEM 13b: see Fig. 2): praecornutus 
Biozone, late Ludlow, Barnby Hills Shale; 
Bohemograptus praecornutus Urbanek, 
1970; Linograptus posthumus Richter, 1875; 
Dendrograptus sp. nov.; Acanthograptus 
aculeatus aculeatus (Poéta, 1894); Thallograptus 
acanthicus vandenbergi Rickards and Wright, 
1997a; and Dictyonema sp. 

W 825: Dripstone railway cutting, type section of 
Barnby Hills Shale, late Ludlow; Bohemograptus, 
Dictyonema and Linograptus (not described or 
figured); this collection has been mislaid. 

W 859: roadside quarry at South Mullion Reserve, 
lower Lewis Ponds Road (Bathurst 1:250 000 
sheet, grid reference 705420E, 6320760N). 
Poorly-preserved and highly-deformed 
graptolites occur in cleaved, pyritic black slate. 
The only species identified is the late Ludlow 
Bohemograptus paracornutus Rickards and 
Wright, 1999b. 


Hanover Formation 

The Hanover Formation has yielded graptolites 
including Saetograptus chimaera and brachiopods 
(Sherwin 1997), and plant fragments (Morgan in 
Meakin and Morgan 1999). Maggs (1963) identified 
Monograptus colonus, M. salweyi, M. bohemicus 
and ?Spinograptus spinosus from a locality 3 km 
roughly along strike north of the graptolite locality 
described here; his identifications suggest the 
nilssoni or scanicus Biozone, early Ludlow. Sherwin 
(1996) identified the graptolites Agastograptus spp., 
?Paraplectograptus sp. and Monograptus ludensis, 
indicating a latest Wenlock age (/udensis Biozone). 
The Hanover Formation thus probably extends from 
late Wenlock throughout the Ludlow and possibly to 
the end of the Pridoli, considering that the formation 
appears to be conformably overlain by the early 
Devonian (Lochkovian) Cuga Burga Volcanics and 
Berkley Formation (Morgan et al. in Meakin and 
Morgan 1999). Graptolite data reported here indicate 
an age range of spineus Biozone (late Ludlow) to 
parultimus Biozone (Pridoli). 

The fauna described here from this formation is 
from one locality: 


WEEM 409 (a, c): spineus (late Ludlow) and 
parultimus (early Pridoli) Biozones, Cumnock 
railway cutting, Hanover Formation; Dictyonema 
elegans Bulman, 1928; Dictyonema paululum 
hanoverense subsp. nov.; Pristiograptus shearsbyi 
Rickards and Wright, 1999b; Monograptus 
spineus (Tsegelnuk, 1976); Neocolonograptus 


SW 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


parultimus (Jaeger, 1975). Three collections have 
been made from different levels in this cutting, 
as follows: WEEM 409 has Neocolonograptus 
parultimus and Dictyonema elegans and D. 
paululum; WEEM 409a has ?Neocolonograptus 
parultimus, Linograptus posthumus, P. shearsbyi 
and WN? wmitchelli, and WEEM 409c_ has 
Monograptus spineus and P. shearsbyi. 


DEPOSITIONAL ENVIRONMENTS 


The Barnby Hills Shale and Hanover Formation 
were deposited in a quiet, deepwater environment, 
reflecting the progressive subsidence of the Cowra 
Trough and Mumbil Shelf during the Middle to Late 
Silurian (Byrnes 1976; Morgan, Colquhoun and 
Meakin in Meakin and Morgan 1999), which allowed 
widespread distribution of fine-grained sediments. 
The time of initiation of subsidence varied, beginning 
in the Cowra Trough in the Wenlock, and on the 
western margin of the Mumbil Shelf in the early 
Ludlow. Ordovician basement rocks were still 
locally exposed, shedding large limestone blocks 
downslope into deeper water (Morgan, Colquhoun 
and Meakin in Meakin and Morgan 1999; Zhen et 
al. 2003; Zhen and Percival 2004). By middle or late 
Ludlow the Mumbil Shelf was totally submerged. 
Fine-grained sedimentation continued in the Cowra 
Trough and on the margins of the Mumbil Shelf into 
the earliest Devonian. Parts of the Shelf underwent 
shallowing during the late Ludlow or early Pridoli, 
permitting local deposition of the Camelford 
Limestone. Conditions changed substantially in the 
early Devonian, with the onset of dominantly mafic 
to intermediate volcanism in the Cowra Trough and 
on the Mumbil Shelf (Cuga Burga Volcanics and 
Berkley Formation), thus terminating quiet mud 
sedimentation (Morgan, Colquhoun and Meakin in 
Meakin and Morgan 1999). 

The late Ludlow praecornutus Biozone, and the 
slightly younger cornutus Biozone characterised by 
B. paracornutus and B. b. tenuis, are now known to 
range widely within central N.S.W. Both characteristic 
taxa have been recorded from the Yass area by 
Rickards and Wright (1999b); we have also seen B. 
paracornutus in collections made by G.H. Packham 
just south of Neurea, and probably the same species 
collected by him from the Manildra area (Savage 
1968). Bohemograptus b. tenuis was described from 
Neurea by Rickards and Wright (1997b). 

In both the Yass (Rickards and Wright 1999b) 
and Neurea (Rickards and Wright 1997a) areas, 


158 


Ludlow graptolitic beds are succeeded sharply by 
shallow water strata (the Rainbow Hill Marl at Yass 
and the Camelford Limestone near Wellington), 
providing evidence for a sea-level highstand followed 
by a latest Ludlow shallowing. Various authors have 
drawn attention to the regression at the end of the 
Ludlow (Talent 1989, Johnson et al. 1991, Johnson 
and McKerrow 1991, Kaljo et al. 1995) and Pickett 
(1982) noted the widespread deposition in NSW 
during the period dominated by bohemograptid 
graptolites; the NSW setting appears to conform to 
the global sea-level pattern at this time. 


SYSTEMATIC DESCRIPTIONS 


Material described here is deposited in the Australian 
Museum, Sydney (AM F). 


Class Graptolithina Bronn, 1849 
Order Dendroidea Nicholson, 1872 


Genus Dendrograptus J. Hall, 1858 


Type species. 
Graptolithus hallianus Prout, 1851; subsequently 
designated by J. Hall (1862). 


Dendrograptus typhlos sp. nov. 
Fig. 3A-B 


21995 Dendrograptus sp.; Rickards et al., p. 18, fig. 
12B. 

21997a Dendrograptus sp.; Rickards and Wright, p. 
214, fig. 6A. 

21999b Dendrograptus sp.; Rickards and Wright, p. 
191, fig. 3A. 


Type Material 

Holotype AM F114618; paratypes AM F114620- 
1; all well-preserved though not showing the 
autothecae too well; all from Blind Gully, locality 
WEEM 13B; praecornutus Biozone, late Ludlow, 
Barnby Hills Shale. 


Etymology 
The species name is the Greek word for blind, 
after the Blind Gully locality. 


Diagnosis 

Dendrograptus branching every 1-3 mm, more or 
less regularly, at angles from 40-90°; younger parts 
of colony have a lateral stipe width of 0.2-0.4 mm _ 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


“rj 


oe 


Figure 3A-G. A-B, Dendrograptus typhlos sp. nov.; respectively AM F114618 (holotype) and 114620; 
question mark indicates possible holdfast in holotype. Blind Gully, praecornutus Biozone, Ludlow. 
C-D, Dictyonema elegans Bulman, 1928; respectively AM F 92382 and 92381, locality WEEM 409, 
parultimus Biozone, Pridoli; E, Dictyonema paululum hanoverense subsp. nov., AM F92380 (holotype), 
WEEM 409, parultimus Biozone, Pridoli. F, Acanthograptus aculeatus aculeatus (Pocta, 1894), AM 
F114616, Blind Gully, praecornutus Biozone, Ludlow. G, Thallograptus acanthicus vandenbergi 
Rickards and Wright, 1997a, AM F14619, Blind Gully, praecornutus Biozone, Ludlow. Scale bars 1 mm 


Proc. Linn. Soc. N.S.W., 126, 2005 159 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


but, nearer the proximal end, lateral widths of 0.3-0.5 
mm occur. Possible compound stipes proximally but 
no indication of them distally. Autothecae difficult 
to see, possibly spaced at ca 30 per 10 mm. Largest 
specimen 25 mm x 20 mm. 


Remarks 

Perusal of the studies of Rickards et al. (1995) and 
Rickards and Wright (1997a, 1999b) demonstrates that 
identification of Dendrograptus specimens is not easy. 
In Australian rocks the genus seems to be relatively 
rare, occurring sporadically throughout the Silurian 
and the earliest Devonian (Rickards and Wright 2001). 
The specimen figured by Rickards et al. (1995, fig. 
12B) as Dendrograptus sp. has similar dimensions 
to D. typhlos and may be an earlier occurrence of it: 
the Quarry Creek specimen is from the /undgreni/ 
testis Biozone of the Wenlock. Dendrograptus sp. (of 
Rickards and Wright 1997a, fig. 6A) from the Ludlow 
Barnby Hills Shale at Neurea has similar dimensions 
but may have a lower autothecal spacing (15-20 in 
10 mm). Dendrograptus sp. (of Rickards and Wright 
1999b, fig. 3A) from the praecornutus Biozone of 
the Yass district also has an apparently lower thecal 
spacing (20 in 10 mm), although is otherwise similar 
and occurs only a little lower stratigraphically than 
the Blind Gully specimens. 


Genus Dictyonems J. Hall, 1851 


Type species 
Gorgonia retiformis J. Hall, 1843; subsequently 
designated by Miller (1889). 


Dictyonema elegans Bulman, 1928 
Fig. 3C-D 


1928 Dictyonema elegans sp. nov; Bulman, p. 52, 
text-fig. 26; pl. 6, figs 22-3. 
1997a Dictyonema elegans Bulman; Rickards and 
Wright, p. 214, figs SC, 8D. 
A fuller synonymy was given by Rickards and Wright 
(1999b). 
Material i 
Several fragmentary specimens (AM F 92381-2, 
AM F114613 a-b, AM F114614) from locality WEEM 
409, parultimus Biozone, Hanover Formation, near 
Cumnock. 


Description 


Dorsoventral width (excluding ventral apertural 
processes) 0.4-0.5 mm. Ventral apertural processes 


160 


robust and relatively short: however, their distal 
extremities may be broken. Autothecal spacing 20 
in 10 mm. One specimen (Figure 3D) may be close 
to the holdfast position, or may enclose the holdfast 
between 3 or 4 diverging stipes; the latter are seen 
partly in profile and partly in dorsoventral view. No 
obvious signs of bithecae. 


Remarks 

Rickards and Wright (1999b) gave a summary of 
the variation seen in Australian records of the species, 
which is now known to range through the Ludlow 
and Pridoli (the type British material being Wenlock). 
The present specimens agree with previous records 
(Rickards and Wright 1997a) from the Barnby Hills 
Shale in having an autothecal spacing of only 20 in 
10 mm, which may be contrasted with the types and 
with Quarry Creek specimens from the Wenlock and 
Ludlow respectively, which have 25 in 10 mm, and 
with the Yass Ludlow and Pridoli specimens, which 
have 30 in 10 mm. Thus, whilst it is certain that these 
are all very similar forms, it cannot be argued that 
the range of variation is as yet constrained in any 
stratigraphic sense. Even so, D. elegans has great 
potential in this regard. 


Dictyonema paululum Bulman, 1928 


1928 Dictyonema paululum, n. sp.; Bulman, p. 58, 
pl. 5, figs 9-11; ?pl. 4, fig. 13. 


Dictyonema paululum hanoverense subsp. nov. 
Fig. 3E 


Material 

Holotype, a single fragment of well-preserved 
stipe in profile view, AM F 92380, locality WEEM 409, 
parultimus Biozone, Pridoli; Hanover Formation. 


Diagnosis 

A subspecies of Dictyonema_ paululum 
characterised by dorsoventral width of 0.50 mm, 
crowding of autothecae (28-30 in 10 mm) and 
dissepiments. 


Description 

Five autothecae seen on the specimen, spaced 
at ca 28-30 in 10 mm. Autothecal apertures slightly 
isolated and, whilst there is no dorsal apertural spine, 
there is a long ventral spine approaching 0.5 mm in 
length. Fine dissepiments seen at dorsal edge of stipe; 
spacing rather close at 20-30 per cm, possibly one to 
each autotheca. One possible bithecal tube visible on 
free ventral wall of penultimate theca. No indications 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


of internal structure save traces of some autothecal 
overlap. 


Remarks 

This specimen differs from Bulman’s late 
Llandovery originals in that the dorsoventral width 
is greater (0.50 mm compared with 0.15-0.25 mm), 
as is the crowding of autothecae (28-30 in 10 mm 
compared with 19-20 in 10 mm in Bulman’s originals) 
and dissepiments. In overall appearance they are very 
similar, however, and especially resemble Bulman’s 
(1928, pl. 5, fig. 11) specimen. Dictyonema elegans 
Bulman, 1928 (see also Rickards and Wright 1997a) 
is another similar form, although all the biocharacters 
have differentmeasurements; clearly they belong to the 
same dictyonemid group typified by slightly isolated 
autothecal apertures, ventral spines, no dorsal process, 
and external bithecae. Dictvonema paululum australis 
Rickards and Jell, 2002, from the griestoniensis 
Biozone of the Graveyard Creek Subprovince of 
Queensland, has very similar dimensions overall but 
differs in having a low dissepimental spacing (12-14 
in 10 mm) and a higher stipe spacing (20+). 

Dictyonema paululum hanoverense subsp. nov. is 
not associated with dendroids other than D. elegans, 
but the Blind Gully locality has yielded several 
rhabdosomal fragments we refer to Dictyonema 
spp. indet.; further material is needed for a more 
satisfactory identification. 


Genus Acanthograptus Spencer, 1878 


Type species 
Acanthograptus granti Spencer, 1878; by original 
designation. 


Acanthograptus aculeatus aculeatus (Poéta, 1894) 
Fig. 3F 


1894 Jnocaulis aculeatus n. sp.; Poéta, p. 199, pl. 
7, figs 22-25. 

1894 Inocaulis demetosa n. sp.; Poéta, p. 200, pl. 
7, figs 7, 8a. 

21909 Inocaulis diffusus (Spencer); Gurley in 
Bassler, p. 53, fig. 68. 

1957  Acanthograptus aculeatus (Poéta); Boucek, 
p. 88, pl. 15, figs 1-9, text-figs 37 a-g. 

1995  Acanthograptus aculeatus (Poéta); Rickards 
et al., p. 24, figs 11F, 15C-F. 


Material 

One well-preserved rhabdosome, AM F114616a, 
b, from Blind Gully locality WEEM 13B, Barnby 
Hills Shale, praecornutus Biozone. 


Proc. Linn. Soc. N.S.W., 126, 2005 


Description 

Acanthograptus with a lateral stipe width of 1.5 
mm, parallel-sided; 10-12 twigs per 10 mm of stipe, 
on each side; twigs alternating on opposite sides (in 
dorsal or ventral view); stipe diversions infrequent 
and irregular; twigs may have two autothecae in each 
termination; broad bases to twigs suggest presence 
of bithecal openings there; stipe may be compound; 
central part of stipe ca 0.4 mm in diameter. 


Remarks 

Previous records of A. a. aculeatus range from 
the late Wenlock to early Ludlow, but the Blind Gully 
specimens are late Ludlow (praecornutus Biozone). 
Thus the species seems to have a long range with 
little morphological change. However, Rickards and 
Wright (1997a) described a more slender subspecies, 
A. a. neureaensis, from the inexspectatus or kozlowskii 
biozonal level in the Barnby Hills Shale at Neurea. 


Genus Thallograptus Ruedemann, 1925 


Type species 
Dendrograptus? succulentus Ruedemann, 1904; 
by original designation. 


Thallograptus acanthicus Bouéek, 1957 


Thallograptus acanthicus vandenbergi Rickards and 
Wright, 1997a 
Fig. 3G 


1997a_ Thallograptus acanthicus vandenbergi 
subsp. nov.; Rickards and Wright, pp. 219-20, 
fig. 6. 


Material 

Four specimens AM F114619, 114622-4, and one 
possible specimen AM F114617a-b, all from Blind 
Gully locality WEEM 13B, praecornutus Biozone, 
late Ludlow, Barnby Hills Shale. 


Description 

Thallograptus reaching 20 mm x 15 mm, with 
robust, spiky stipes which branch and diverge 
more or less at right angles; stipes compound, with 
bundles of long, narrow tubes; main stipe width 
1.5 mm proximally, down to 0.5 mm most distally; 
progressive dichotomies become narrower, and all 
stipe termination is by single, conspicuous autothecal 
tubes. 


161 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


Remarks 

No bithecae have been detected and the 
autothecal spacing cannot be given until more 
autothecal openings can be seen on the ventral stipe 
surface. This subspecies resembles Acanthograptus 
a. aculeatus but the latter is more slender and has a 
very regular association of twigs opening alternately 
on one side of the stipe and the other. Thallograptus a. 
vandenbergi is a smaller and more slender subspecies 
than the type subspecies. The species is very rare, 
with one specimen only recorded from Bohemia (the 
holotype of Boucek’s 1957 nominate subspecies) and 
five definite Australian specimens. 


Order Graptoloidea Lapworth, 1875 
Genus Pristiograptus Jaekel, 1889 


Type species 
Pristiograptus frequens Jackel, 1889, by original 
designation. 


Pristiograptus shearsbyi Rickards and Wright, 
1999b 
Figs 4D-E 


1999b_ Pristiograptus shearsbyi n. sp.; Rickards and 
Wright, p. 194, figs 3J-P, 11A-B, 13B-E. 


Material 

AM F 92371, 92376, 114574-81 from localities 
WEEM 409 and 409c, respectively the parultimus 
and spineus biozones, Hanover Formation. 


Description 

Straight, thin Pristiograptus with dorsoventral 
width proximally 0.5 mm, distally 0.8-1.2 mm; sicula 
1.2-2.1 mm long, reaching midway between apertures 
of thl and th2 in case of shorter siculae, and up to 
the level of th2 aperture in rhabdosomes with long 
siculae; )) = 1.2-1.4 mm; thecal overlap ca /%; thecal 
inclination 20-30°; proximal thecal spacing 11 in 10 
mm; distal thecal spacing 10 in 10 mm; sicula with 
short, dorsal tongue. 


Remarks 

This material is very close to the Yass material; 
the species ranges from the praecornutus Biozone into 
the Pridoli. Some Hanover specimens have a slightly 
greater dorsoventral width (up to 1.2 mm compared 
with 0.85 mm) and some have a longer sicula (up to 
2.1 mm compared with 1.65 mm). 


162 


Genus Monograptus Geinitz, 1852 


Type species 
Lomatoceras priodon Bronn, 1835; subsequently 
designated by Bassler (1915). 


Monograptus spineus (Tsegelnuk, 1976) 
Fig. 4A 


1976 Acanthograptus spineus sp. n.; Tsegelnuk, p. 
113, pl. 34, figs 6-9. 

1983 Bugograptus spineus (Tsegelnuk); Tsegelnuk, 
p. 145, fig. 34. 

1988 | Monograptus spineus (Tsegelnuk); Koren’ et 
al., p. 17, fig. 8. 
1995 Monograptus (Uncinatograptus) spineus 
(Tsegelnuk); Urbanek, p. 3, figs 1D, 2, 7C-E. 
1997 Monograptus (Uncinatograptus) spineus 
(Tsegelnuk); Urbanek, pp. 149-154, pl. 11, figs 
3-6; pl 12; pl. 13, figs 13, 35-41. 

1997 | Monograptus spineus (Tsegelnuk); Koren’ 
and Sujarkova, p. 80, pl. 6, figs 1-2, text-figs 
14A-I. 


Material 

A single, well-preserved proximal end, AM 
F 92370, from locality WEEM 409c, Hanover 
Formation, spineus Biozone, Ludlow. 


Description 

The well-preserved proximal end has 3.5 thecae, 
the sicula, virgella, virgula and sicula thickening 
bands all clearly preserved. Thecal overlap low; thecal 
hooks well-developed, with a pair of lateral spines and 
a central retroflexed portion showing fuselli on th1. 
Upper sicular band coincides with base of th2, but it 
may or may not represent pro-metasicular boundary. 
Second band halfway along sicula, which is 1.4 mm 
long, its apex reaching midway between th! and th2. 
Thecal spacing 12.5 in 10 mm and dorsoventral width 
1.1-1.2 mm. } = 0.9 mm. 


Remarks 

This specimen is the first record of the species 
outside Podolia, Poland, and Russia. Urbanek 
(1997) considered the species indicative of the late 
Ludfordian (Ludlow) spineus Biozone. It overlaps 
with the lower (Ludlow) part of the range of the 
more common species M. formosus Boucek, 1931, 
previously recorded from Yass, N.S.W. by Packham 
(1968) and Rickards and Wright (1999b). 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


Figure 4A-L. A, Monograptus spineus (Ysegelnuk, 1976), AM F92370, WEEM 409c, spineus 
Biozone, Ludlow. B-C, Monograptus ludensis (Murchison, 1839), AM F114572-3, 68 KIL, 
ludensis Biozone, Wenlock. D-E, Pristiograptus shearsbyi Rickards and Wright, 1999b, AM 
F92376 and 923371, WEEM 409c, spineus Biozone, Ludlow. F-J, Neocolonograptus parultimus 
(Jaeger, 1975), respectively AM F92379, 92377, 92378, 92374 and 92375, WEEM 409, parultimus 
Biozone, Pridoli. K-L, Bohemograptus praecornutus Urbanek, 1970, AM F102918 from Blind 
Gully; and MMF 33611, WEEM 409a, both praecornutus Biozone, Ludlow. Scale bars 1 mm. 


Proc. Linn. Soc. N.S.W., 126, 2005 163 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


Monograptus ludensis (Murchison, 1839) 
Fig. 4B-C 


1839  Graptolithus ludensis n. sp.; Murchison, p. 
694, pl. 26, fig. 2 (non fig. 1). 

1996 Colonograptus ludensis (Murchison); Lenz 
et al., p. 1396, figs 3P, D, 4M-R. 

1997b Monograptus ludensis (Murchison); Rickards 
and Wright, pp. 236-7, figs 2D-H, 4A-E, SJ. 

For further references see Rickards et al. (1995) and 

Rickards and Wright (1997b). 


Material 

Well-preserved specimens: AM F114572-3, 
114582-600, from locality KIL 68, /udensis Biozone, 
Barnby Hills Shale. 


Description 

Robust pristiograptid-like rhabdosomes up to 30 
mm long and 2 mm broad, but with aperture of thl 
clearly rounded; proximal end and sicula often slightly 
curved ventrally; proximal dorsoventral width 0.8-1.0 
mm; proximal thecal spacing 14-10 in 10 mm; sicula 
1.8-2.2 mm reaching to level of aperture of th2; thecal 
overlap ca % - %4; thecal inclination 45-50°. 


Remarks 

These forms are very close to many previous 
descriptions and would correspond with the sort 
of M. ludensis that has been sometimes called M. 
praedeubeli Barca and Jaeger, 1990; this matter was 
discussed by Rickards and Wright (1997b). 


Genus Neocolonograptus Urbanek, 1997 


Type species 
Monograptus lochkovensis Pribyl, 1940; by 
original designation (Urbanek 1997, p. 128). 


Neocolonograptus parultimus (Jaeger, 1975) 
Fig. 4F-J 


1899  Monograptus ultimus n. sp.; Perner, p. 13, pl. 
10, figs 4, 5 (non fig. 14 a, b =Neocolonograptus 
ultimus). 

1975 Monograptus parultimus n. sp. Jaeger, p. 
119, pl. 2, figs 4, 8; text-fig. 4. 

1997 Neocolonograptus  parultimus (Jaeger); 
Urbanek, pp. 166-7, pl. 21, figs 1-7; fig. 48. 

1999b Monograptus parultimus Jaeger, 
Rickards and Wright, p. 165, figs 3Q-S, U. 


LOWS; 


More detailed synonymies are given by Urbanek 
(1997), Koren’ and Sujarkova (1997) and Rickards 


164 


and Wright (1999b). 


Material 

Well-preserved specimens from localities WEEM 
409 and 409a: AM F 92374-5, 92377-9, 114601-11, 
parultimus Biozone, Hanover Formation. 


Description 

Neocolonograptus thabdosomes up to 12 mm 
long with distal dorsoventral width up to 1.6 mm, 
proximally 0.6-0.8 mm and proximal end often with 
slight ventral curvature; sicula 1.6-2.0 mm long, with 
occasionally-preserved thickening bands, apex below 
level of aperture of th2; dorsal tongue conspicuous; 
virgella short; slight rounding on first theca or some 
higher thecae; thecal overlap ca '2; thecal spacing 
10.5-13 in 10 mm; >» = 1.3-1.4, 1.6 in one specimen. 


Remarks 

This material is close to that described from 
Yass by Rickards and Wright (1999b), which in turn 
was shown to be close to Jaeger’s types from Kosov 
Quarry; however, the Barnby Hills Shale specimens 
have a > value closer to that of the material from south 
Tien Shan described by Koren’ and Sujarkova (1997). 
There is also a suggestion in our present collection 
that the apertural undulations of the proximal thecae 
are slightly less than in the specimens from Yass. It is 
possible that they are slightly earlier, perhaps near the 
base of the parultimus Biozone. 


Genus Bohemograptus Pribyl, 1967 
Type species 


Graptolithus bohemicus Barrande, 
original designation. 


1850; by 


Bohemograptus praecornutus Urbanek, 1970 
Fig. 4K-L 


1970 Bohemograptus  praecornutus 0. Sp.; 
Urbanek, pp. 301-10, text-fig. 16, pl. 20C, pls 23, 
24 

1999b Bohemograpius  praecornutus  Urbanek; 
Rickards and Wright, pp. 200-202, figs 5C-L, 
13K 

A full synonymy is given in Rickards and Wright 

(1999b) 


Material 

Two specimens, AM F102918, and MMF 33611 
from Blind Gully locality WEEM 13B near Cumnock; 
and a less well-preserved specimen, AM F114612 
from the same locality; praecornutus Biozone; 


Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


Barnby Hills Shale. 


Description 

Robust Bohemograptus with tight ventral 
curvature. Dorsoventral width at thl 0.6-0.7 mm, 
distal dorsoventral width not seen; proximal thecal 
spacing 14 in 10 mm (distal thecal spacing not seen); 
thecal overlap ca 1/2; thecal inclination 40-50°; > 
1.0-1.4 mm. Conspicuous sicula with pronounced 
dorsal tongue, 1.4-1.8 mm long, reaching to a little 
above aperture of thl. 


Remarks 
These specimens differ from the Yass material 
only in having a slightly longer sicula in one 


specimen: all the other measurements agree closely. _ 


They are, therefore, very close indeed to Urbanek’s 
(1970) originals from Poland. As in the Yass district 
the presence of B. praecornutus can be taken as an 
indicator of the praecornutus Biozone. 


- Bohemograptus paracornutus Rickards and Wright, 
1999b 
Fig. 5D-E 


1999b Bohemograptus  paracornutus nn.  sp., 
Rickards and Wright, p. 202, figs SM-Q, 6A-N, 
7, 9A-B, 10A-E. 


Material 

AM F 114767-8 and at least 50 other specimens 
from the Mullions Reserve locality, all highly deformed 
tectonically, and in a monotypic assemblage. 


Remarks 

The highly deformed nature of the specimens 
precludes a useful description. There is, however, no 
doubt about the nature of the strikingly flared sicula, 
which is identical to that of the type material from 
Yass, nor in the nature of the thecae with their slightly 
raised lateral apertural rim and gently concave free 
ventral wall. Some specimens (e.g. Fig. 5E) are 
abnormal in thecal spacing and nature of the sicula, 
but this may be caused by tectonic deformation. 


Genus Enigmagraptus Rickards and Wright, 2004 
Type species 
Neocucullograptus? yassensis Rickards and 


Wright, 1999; by original designation. 


Species recognised 
Enigmagraptus yassensis (Rickards and Wright, 


Proc. Linn. Soc. N.S.W., 126, 2005 


1999b); E. sp. cf. yassensis (Rickards and Wright, 
1999b); E. mitchelli (Rickards and Wright, 1999b); 
E. pennyae Rickards and Wright, 2004. 


Diagnosis 

One of the tiniest known graptolites, with a 
dorsoventral width up to 0.25 mm; widely spaced 
thecae; axially elongate protheca usually developed 
from thread-like origin; tiny metatheca up to half 
thabdosome width, consisting of hood derived 
from dorsal metathecal wall, and variously enrolled 
ventrally to enclose simple ventral thecal margin; 
small sicula with virgella and dorsal apertural process 
in type species (corrected after Rickards and Wright 
2004). 


Remarks 

Since the description of the Yass material 
of E. yassensis and E. mitchelli by Rickards and 
Wright (1999b), E. yassensis has also been found 
at Cumnock and E. sp. cf. yassensis has been found 
at the ‘borrow pit locality (W910) near Cadia mine 
(Rickards et al. 2001). Enigmagraptus mitchelli 
was, until now, known only from the type locality 
at Yass. Enigmagraptus pennyae was described by 
Rickards and Wright (2004) from the same locality 
(W910) as E. sp. cf. yassensis. All these localities are 
Pridoli; W910 is probably late Pridoli, whereas the 
other occurrences are best assigned to the parultimus 
Biozone, early Pridoli. 


Enigmagraptus mitchelli (Rickards and Wright, 
1999b) 
Fig. SA-C 


1999b Neocucullograptus? mitchellin. sp.; Rickards 
and Wright, p. 200, figs 4S, T. 


Material 

Three specimens, AM F 92372-3 and 114571, 
all from locality WEEM 409a; parultimus Biozone; 
Hanover Formation. 


Description 

There is one proximal end with a partially 
preserved sicula and 2 % thecae, with virgella and 
virgula also preserved. Sicula may be ca 0.5 mm 
long, although apex is missing; it reaches only part 
way along thl. Thecal spacing of this proximal part, 
and a fragment possibly also close to proximal end 
(Fig. SA), ca 7.5-8 in 10 mm but there is some soft 
sediment deformation along specimen shown in Fig. 
SC so thecal spacing of 7 in 10 mm might be more 
likely in undeformed material. A more distal fragment 


165 


SILURIAN GRAPTOLITES FROM THE LACHLAN FOLD BELT 


a 


Figure 5A-E. A-C, Enigmagraptus mitchelli (Rickards and Wright, 1999b), respectively AM 
F92373, 92372 and 114571, WEEM 409a, ?parultimus Biozone, Pridoli; D-E, Bohemograptus 
paracornutus Rickards and Wright, 1999, AM _  F114768 and_ 114767 _ respectively; 
approximately cornutus Biozone, late Ludlow, W859, Mullions Reserve. AM F114767 appears 
to have abnormal dimensions: heavy bar = tectonic stretching direction. Scale bars 1 mm. 


(Fig. SA) shows 7 in 10 mm. Maximum dorsoventral only a few degrees at most. 

width, including hook, of most distal fragment is 0.4 

mm, but proximal end is only 0.2 mm at thl. Late Remarks 

metathecal part has a dorsoventral width of 0.15 mm These rare specimens are similar to those described 
on th2 and 0.2 mm on most distal thecae seen. Thecal from the same stratigraphic level (parultimus 
overlap low and thecal angle (of free ventral wall) | Biozone) fromYass by Rickards and Wright (1999b). 


166 Proc. Linn. Soc. N.S.W., 126, 2005 


R.B. RICKARDS, J.R. FARRELL, A.J. WRIGHT AND E.J. MORGAN 


They differ in being more slender and in having a 
higher thecal spacing (7 in 10 mm compared with 4.5 
in 10 mm). However, it is possible that these represent 
the proximal ends of the same species: certainly the 
profile of the hook is very similar as are most other 
features such as overlap and inclination. Only four 
specimens are known (including one from Yass) and 
the nature of the thecal hook is far from certain. 


ACKNOWLEDGMENTS 


Support for a visit to Australia by RBR was supported 
by The Royal Society; Emmanuel College, Cambridge; 
the Department of Earth Sciences, Cambridge; and the 
University of Cambridge. Field work by AJW was supported 
by the University of Wollongong. EJM publishes with the 
permission of the Deputy Director-General of the NSW 
Department of Primary Industry and Mineral Resources. 
We are grateful to two anonymous reviewers who greatly 
improved an initial draft of the manuscript. 


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Altitude, Frost and the Distribution of White Box (Eucalyptus 
albens) on the Central Tablelands and Adjacent Slopes of NSW 


W.S. Sempce! AND T.B. KOEN? 


"NSW Department of Infrastructure, Planning and Natural Resources, PO Box 53, Orange NSW 2800 
bill.semple@dipnr.nsw.gov.au 
“NSW Department of Infrastructure, Planning and Natural Resources, PO Box 445, Cowra NSW 2794 


Semple, W.S. and Koen, T.B. (2005). Altitude, frost and the distribution of white box (Eucalyptus albens) 
on the Central Tablelands and adjacent slopes of NSW. Proceedings of the Linnean Society of New South 


Wales 126, 171-180. 


The apparent rarity of white box (Eucalyptus albens) at high altitude has been explained by assuming an 
intolerance to low temperatures. These propositions were assessed by (a) a field survey of the occurrence 
of white box at high altitude on part of the NSW Central Tablelands and adjacent slopes and (b) a pot 
trial study of the response of seedlings of white box and yellow box (E. melliodora) to low temperatures 
during winter 1997. The field survey confirmed that, unlike yellow box, white box was absent from a large 
part of the Central Tablelands. However, it was recorded at altitudes up to 925 m a.s.l. near Orange but at 
lower altitudes further south. Aspect was not limiting at high altitude though low slopes appeared to be. 
Survival and growth of white and yellow box seedlings were low at the high altitude site but there was 
little difference between species regardless of whether frosted seedlings were subjected to early or delayed 
exposure to direct morning sunlight. Hence, alternative explanations for the local distribution pattern of 
white box on the Central Tablelands and adjacent slopes need to be examined. 


Manuscript received 15 November 2004, accepted for publication 2 February 2005. 


KEYWORDS: altitude, aspect, Eucalyptus albens, Eucalyptus melliodora, frost, lignotuber, 


photoinhibition. 


INTRODUCTION 


White box (Eucalyptus albens Benth.) extends 
from south-east Queensland along the slopes of NSW 
to central Victoria. Disjunct populations occur in 
eastern Victoria, western Victoria and the Southern 
Flinders Ranges of SA. The most recent description 
of the habitat of E. albens was prepared by Prober 
and Thiele (1993) and draws heavily on previous 
accounts. They reported that it generally occurs on 
’ fertile soils derived from a wide variety of parent 
materials. Within its area of occurrence, mean annual 
rainfall ranges from 500 - 800 mm, mean maximum 
temperature in the hottest month 27 - 32 °C, mean 
minimum temperature in the coldest month -1 - 5 °C 
and frost frequency 5 - 70 per year. 

Cambage (1902) and Beadle (1981) suggested 
that its absence from higher parts of the tablelands 
was due to ‘coldness’ or intolerance of heavy frosts. 
It has been recorded at high altitude but its upper 
altitudinal limit is difficult to determine as altitude is 
rarely recorded on locality details accompanying 


herbarium specimens. Actual and apparent altitude 
records in CSIRO’s (since discontinued) Eucalist, 
Victorian Flora Information System and NSW 
National Herbarium databases (Table 1) indicated a 
record at ~1060 m above sea level (a.s.l.). Boland 
et al. (1984), however, reported an occurrence from 
an unspecified locality, possibly in northern NSW, at 
1200 m a.s.l., some 600 m higher than in an earlier 
edition of this publication (Hall et al. 1970). 

Though frosts are common at high altitudes, they 
are also common in ‘frost hollows’ produced by cold 
air drainage at lower altitudes. Hence, if E. albens is 
intolerant of heavy frosts it would not be expected 
to occur in depressions subject to frequent frosts. 
The relative occurrence of E. melliodora Cunn. ex 
Schauer (yellow box) and E. al/bens in the Central 
Western Slopes and Central Tablelands botanic 
subdivisions of NSW (explained and mapped in 
various editions of Anderson, e.g. 1968) suggests that 
this may be the case. For example, around Bathurst 
~670 m a.s.l.), which is located in a basin subject to 


DISTRIBUTION OF WHITE BOX 


Table 1. Some high altitude records (metres above sea level) for E. albens across its natural 
range from north to south. Sources: CSIRO’s ‘Eucalist’ (since discontinued), Victorian Flora 
Information System (VFIS), National Herbarium of NSW (NSW). 


Specified Possible maximum Location Source(1997) 
altitude altitude 

900 - Glenn Innes - Emmaville, NSW Eucalist 
- 850 Top of Mt. Wallaby, Woolomin NSW 
: 1100 Hanging Rock, Nundle, NSW NSW 

780 - WSW of Quirindi. NSW Eucalist 
- 10604 Pinnacle Lookout [Coolah Tops] NSW 

731 - Wattle Flat - Sofala Eucalist 
- 9608 Mt. Remarkable, S.A. Eucalist 
- 1204°¢ Mt. Mcquarie, Blayney, NSW NSW 

900 - South of Suggan Buggan, Vic. VFIS 

850 - Southeast of Suggan Buggan, Vic. VFIS 
= 704? The Paps, Mansfield, Vic. NSW 
- 1167® Mt. William, Grampians, Vic. Eucalist 


“Confirmed by M. Sharp (pers. comm., 2001) who also noted that E. albens occurs elsewhere in Coolah Tops 


National Park at elevations up to ~1090 m a.s.l. 


8 Occurs on the southern foothills but is unlikely to extend to summit ridge (W. Semple, personal observations). 
©. Most likely a record from the slopes of Mt. Macquarie, where E. albens occurs up to at least 870 m a.s.l. (this 
paper). A summit location cannot be confirmed due to its conversion to exotic pine forest. 

D._ Not observed above 550 m a.s.1. on this hill (J. Lawrence, pers. comm., 2001). 

E_ Probably a mis-identification as E. albens is not mentioned in Elliot et al. (1984). 


cold air drainage, E. melliodora occurs in both the 
basin and at higher altitudes, whereas E. albens is 
restricted to the latter (e.g. towards Hill End). Around 
Orange (800 - 900 m a.s.l.), which is located on a 
plateau, E. melliodora is relatively common up to 
~900 m a.s.l. but E. albens is rare. However, within 
the Central Western Slopes, E. albens commonly 
occurs on hills and slopes and E. melliodora on flats 
and lower slopes. These patterns could be explained, 
at least partially, by assuming that E. melliodora has 
a higher tolerance to frost than does E. albens. This 
explanation is at variance with other authorities and 
local observations. For example, Bower et al. (2002) 
reported that E. melliodora is more common on more 
fertile, though perhaps less well-drained, soils than E. 
albens. Though E. albens is not typically associated 
with poor soils, its local distribution pattern is often 
explained in terms of differences in soil properties 
(e.g. Prober 1996). 

If frost does affect the local distributions of E. 
melliodora and E. albens, it would be expected to 
operate at a sensitive stage of the life cycle, viz. the 
seedling, especially those that emerged in autumn or 


172 


winter. According to Cremer (1990), frost can affect 
young seedlings in three main ways: (a) ‘wilting’ 
where leaves become flabby and darkened with a 
waterlogged appearance, which is followed within 
days by drying; (b) damage to roots which are more 
sensitive than shoots; and (c) ‘frost heave’ where 
the stem of the seedling is gripped by a frozen 
soil crust and forced upwards by underlying ice 
crystals resulting in roots being detached from the 
soil. A further possibility is that of ‘stress-induced 
photoinhibition’ where photosynthetic capacity is 
reduced in stressed seedlings and that under high - 
light conditions, more light energy is absorbed than 
can be used or dissipated. The visual effects of this 
would presumably be leaf death similar to ‘wilting’. 
Based on work by Ball et al. (1991), Egerton (1996) 
proposed chronic cold-induced photoinhibition as a 
major reason for the absence of eucalypt seedlings 
growing to immediate north of E. pauciflora Sieber ex 
Sprengel trees near Canberra. However, earlier work 
with ~30 cm tall, frost-hardened, subalpine eucalypt 
seedlings in a radiation frost room by Harwood (1980) 
indicated no difference in leaf damage between 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.S. SEMPLE AND T.B. KOEN 


seedlings that were exposed, on one occasion only, to 
bright sunlight v. darkness following frosting. 

As part of research into the factors affecting the 
recruitment of E. albens, its local distribution pattern 
around Mt. Canobolas on the Central Tablelands 
of NSW was investigated in the mid 1990s. 
Preliminary results (Semple 1997) indicated that E. 
albens mainly occurred on slopes with non-easterly 
aspects at altitudes above 780 m a.s.l. On the basis 
of these results and Egerton’s (1996) hypothesis, the 
survey was extended to a wider area and the effects 
of low temperatures on young eucalypt seedlings 
were investigated in a pot trial during winter 1997. 
The hypotheses tested in the pot trial were that (1) 
seedlings of E. albens are less frost tolerant (as 
assessed by survival and indices of growth) than 
those of E. melliodora and (2) frosted seedlings of 
E. albens (and possibly those of E. melliodora) are 
adversely affected by exposure to direct sunlight. 


METHODS 


Upper altitudinal limits of E. albens on the 
tablelands and adjacent slopes 

Over a number of years, roads radiating from two 
local high points, Mt. Canobolas (1397 m a.s.1.) and 
Mt. Macquarie (1204 m a.s.].) were examined for the 
highest occurrences of E. albens. Each location was 
plotted on a map and altitude and aspect recorded. 
Trees, which could be confidently identified in 
adjacent paddocks, were also included in the survey. 
All 11 sites in Semple’s (1997) earlier survey were 
revisited. 

It was appreciated that the reduced likelihood of 
roads traversing the highest points in the landscape, 
the absence of public roads in some areas, and 
selective tree removal on roadsides were potential 
sources of error in this technique. 


Effect of frost + early morning sunlight on 
seedlings of E. melliodora and E. albens 

Seedlings were raised in 30 cm diameter x 27 cm 
deep black plastic pots containing sandy loam topsoil 
from an E. albens site at Cowra overlain by ~2 cm 
of seed-raising mixture. A slow-release fertiliser 
was mixed with the soil in all pots in an attempt to 
overcome any nutritional problems that may have 
adversely affected E. melliodora, which usually 
occurs on fertile soils. Half of the 16 pots used in 
the experiment were randomly allocated to seed 
(collected from around Molong on the upper Central 
Western Slopes) of E. albens and the other half to 
E. melliodora in early March 1997. Seedlings and 


Proc. Linn. Soc. N.S.W., 126, 2005 


weeds were progressively removed until 30 similar- 
sized seedlings of either species were present in each 
pot. Due to settling of the soil, the rim of each pot 
was later cut down to ~2’% cm above the soil surface. 
At the commencement of the experiment on 1 May 
1997, seedlings were at the four to six leaf stage with 
those of E. albens generally being more advanced due 
to earlier emergence. 

Two pots of each species were randomly 
allocated to four treatments, consisting of the factorial 
combination of light (Delayed or Early Light) and 
location (Orange, 870 m a.s.l. or Cowra, 380 m 
a.s.l.). The Delayed Light treatment was realised 
by placing pots ~1 m to the west of an existing or 
constructed north-south opaque steel fence, thereby 
delaying exposure to direct morning sunlight until 
1035 hours (early July) at both locations. The Early 
Light treatment was achieved by placing the pots in 
an open area exposed to direct sunlight at 0740 hours 
(early July) at Cowra and 0805 hours at Orange where 
‘sunrise’ was delayed by topography and vegetation. 
Pots were watered as required, avoiding prolonged 
waterlogging. At 1 to 2 weekly intervals, the positions 
of the four pots in each location by light treatment 
were rotated. Each pot was also rotated through 180° 
in an attempt to evenly distribute shading from the 
rim of the pot. At the same time, seedlings were 
counted, dead plants removed and apparent cause of 
death noted. 

On 15 September 1997, each seedling was 
assessed for height of main stem (from the cotyledons 
to the upper-most live leaf) and the number of live 
pairs (or part pairs) of seedling leaves on the main 
stem. Pots with high numbers of healthy plants (viz. all 
those at Cowra) were systematically thinned to about 
15 plants per pot. On 13 October when the likelihood 
of further frosts was low, all pots were relocated to a 
concrete apron with an automated watering facility at 
Cowra to evaluate subsequent growth under uniform 
conditions. Seedling numbers were recounted in 
early November and, together with measurements of 
heights, in early January 1998. 

Data (survival, mean numbers of leaves and 
mean heights) were analysed as a 2? factorial design 
of Location (2) x Species (2) x Light treatment (2), 
replicated twice, using analysis of variance methods. 
Visual examination of residual diagnostic graphs 
indicated a non-normal distribution in the September 
seedling height data and a transformation to natural 
logarithms was carried out. Treatment means were 
examined for significant differences (P = 0.05) 
using the least significant difference (lsd) multiple 
comparison procedure (Steel and Torrie 1960). 


173 


DISTRIBUTION OF WHITE BOX 


"YY Uy 

My Yj Yy Uj 
———— 5 Y 
< 


Y 
‘Uy 


Figure 1. Part of the Central Tablelands and adjacent slopes of NSW showing areas where Eucalyp- 
tus albens is likely (hatched area, including the small area south of Carcoar) or unlikely (unhatched 
area) to occur. Numbered +s indicate locations of E. albens at the highest elevations on roads (thick 
lines). Thin lines indicate the 1000 m (near Mts. Canobolas and Macquarie and Hobbys Yards in the 


south-east) and 700 m contours. 


RESULTS 


Upper altitudinal limits of FE. albens on the 
tablelands and adjacent slopes 

Eucalyptus albens was absent from most roadsides 
above 900 m in the vicinity of Orange but to the south- 
west, its highest occurrence was rarely above 750 m 
a.s.l. It was absent from many roadsides, particularly 
those to the south-east of Orange. The occurrence or 
non-occurrence of E. albens, as determined in the 
roadside survey, is shown in Fig. 1. All occurrences 
were of mature trees though regeneration was evident 
at some sites. 


174 


Elevation 

Thirty main sites (Fig. 1) and nine nearby 
subsidiary sites were identified as being the highest 
elevation occurrences on the roads travelled. The 
highest elevations were recorded near Orange: to 
the east on the fall of the Central Tablelands to the 
Macquarie River valley (925 m a.s.l., site 2) and to 
the north (890 m a.s.l., site 9a) on the western fall of 
the tablelands. A disjunct population at high elevation 
(up to 870 m a.s.l., site 28) was also recorded near 
Carcoar on the western and north-western slopes of Mt. 
Macquarie. The site with the lowest elevation (660 m 
a.s.l., site 23) occurred west of Lyndhurst on the Mid 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.S. SEMPLE AND T.B. KOEN 


Table 2. Mean monthly terrestrial minima and frost frequencies at the Orange Agricultural 
Institute* (1975-96) and the Department of Infrastructure, Planning and Natural Resources 
Research Centre at Cowra® (1943-97), together with monthly data for 1997. 


Terrestrial minima (°C) Frost© frequency (days/month) 


Cowra Cowra Orange Orange Cowra Cowra Orange Orange 


Gnacan) 997 (mean) e897 (mean) 1997 (Gnean) | 1 1997 
May 373 4.6 DD, Dy 6.3 3 Te 8 
June 1.3) -1.2 -0.2 -2.5 10.9 19 13.6 24 
July 0.1 -1.8 -1.2 -4.1 (39) 20 18.1 28 
Aug 0.8 -0.9 -0.8 -2.7 1-42 19 16.3 25 
Sept 2.6 3.4 0.8 1.4 6.8 | eS: 5) 
Oct 3) 4.3 3.0 21) 1.9 4 a2 9 


“ 890 m a.s.l and 3.4 km from the Orange experimental site. 
5 381 maz.s.l. and c.100 m from the Cowra experimental site. 
© A frost was considered to have occurred when a minimum of < -0.9 °C was recorded at 2.5 cm above grass. 


Western Highway. Many other sites in the south-west 

.were of relatively low elevation, e.g. sites 17, 19, 
21, 22, 24 and 26 were at elevations below 750 m 
a.s.1. The situation was similar but less marked in the 
north-east (sites 3, 4, 6, 7 and 8). 


Aspect 
Approximately 25 % of the 39 occurrences 


occurred on crests and hence could not be 
allocated a single aspect though averaging was 
attempted. Virtually all occurrences, especially at 
higher altitudes, were on sloping land. No sites 
occurred in drainage lines but at one relatively low 
elevation site (720 m a.s.l., site 3a), E. albens 
extended down-slope to a drainage line. When each 
site was allocated an aspect (N, E, S or W quadrants), 
the numbers of sites in each quadrant declined from 
W=N>E>S. Of the 12 highest elevation sites, 1.e. 
=800 m ASL, numbers in each quadrant were W = E 
=N>S. 


Effect of frost + early morning sunlight on 
seedlings of E. melliodora and E. albens 

Monthly terrestrial minima were lower and frost 
frequencies were higher than average at both sites 
from June to August 1997 (Table 2). During the main 
period of the experiment, | May to 12 October, 97 
frosts were recorded at Orange and 63 at Cowra. Five 
or more days of consecutive frosts occurred on six 
occasions at Orange and seven at Cowra. The lowest 
terrestrial minima recorded were -8 °C at Cowra and 
-9 °C at Orange, both on 21 July. Frosts persisted for 
longer at Orange than at Cowra — particularly in the 


Proc. Linn. Soc. N.S.W., 126, 2005 


Delayed Light treatment. The soil in the pots was often 
frozen at or just below the surface during midwinter 
in Orange. 


Effect of frosts on seedlings 

At Orange a progressive decline in numbers of 
seedlings of both species commenced in June (Fig. 2). 
Seedlings in the Early Light treatment were adversely 
affected initially but by mid August, mean numbers of 
survivors were similar in all treatments. Most deaths 
at Orange were frost-related, i.e. wilting and/or frost 
heave with the latter particularly affecting small (<6 
leaves) seedlings, which were more common in £. 
melliodora than in E. albens populations. None of the 
deaths at Cowra (20 by mid October) showed definite 
frost effects (Table 3). 

In July, it was noted that some seedlings of both 
species had produced shoots in leaf axils as well from 
the cotyledon area, 1.e. the site where the lignotuber 
would subsequently develop. When assessed in mid 
September, axillary shoots were present in 95 % of all 
seedlings and ‘lignotuberous shoots’ in 78 %. The early 
development of lignotuberous shoots apparently had 
no adverse effect on the development of lignotubers, 
which were present in 89 % of surviving seedlings in 
early January 1998. 


Differences between treatments 

Mean seedling survival (in October), main 
stem height and numbers of leaves (in September) 
were significantly higher at Cowra than at Orange 
(Table 4). Apart from mean numbers of leaves on E. 
melliodora seedlings being greater than on E. albens, 


IIS) 


DISTRIBUTION OF WHITE BOX 


Table 3. Probable cause of death of seedlings (120 of each species initially) in both Early and 
Delayed Light treatments at (a) Orange and (b) Cowra between 1 May and 11 October 1997. 


Unknown Frost heave ‘Wilt’ Frost heave All causes 
and ‘wilt’ 

(a) Orange 

Eucalyptus albens 2 12 67 6 87 
E. melliodora 5) 32 3] 2 70 
(b) Cowra 

Eucalyptus albens 6 0 0 0 6 
E. melliodora 14 0 14 


there were no significant differences between the 
species across the two sites. Significant interactions 
suggested that mean height and number of leaves 
were significantly higher for all seedlings in the 
Delayed than in the Early Light treatment at Cowra 
but not at Orange; and that across both sites, EL. albens 
seedlings were significantly taller in September in the 
Delayed than the Early Light treatment (Table 4). 
Most of the seedlings present in mid October 
1997 survived until early January 1998 at which 
time the mean height of seedlings raised at Cowra 
was significantly higher than those raised at Orange 
but the difference was less marked than previously. 
There were no other differences between species and 
treatments and their interactions (Table 4). 


DISCUSSION 


Natural occurrence of E. albens at high altitudes 

The roadside survey data indicated that E. albens 
did not have a consistent upper altitudinal limit 
around Mts. Canobolas and Macquarie on the Central 
Tablelands of NSW. In the south-west of the study 
area it did not occur above 750 m whereas nearer to 
Orange it extended to elevations above 800 m. At the 
latitude of about Millthorpe (~33° 30’ S), E. albens 
does not occur further east than is shown in Fig. 1 
(apart from a recently-discovered, small disjunct 
population at 750 m a.s.l. on the northern footslopes 
of Mt. Panorama at Bathurst) despite the availability 
of other low elevation sites in the Bathurst Basin. It 
extends further east to the north of Orange (as shown) 
and to the south of the study area in the vicinity of 
Abercrombie, south of Blayney. Clearly, the Central 
Tablelands represent a barrier to the distribution of 
E. albens but on its own, altitude (and presumably 
frost severity) does not appear to be a limiting factor 
except at very high altitudes. 

Other potential limiting factors include higher 


176 


rainfall and waterlogging, which may favour other 
eucalypt species on the tablelands, and soil differences. 
High altitude occurrences of E. albens were sometimes 
associated with certain soil landscapes (composites of 
soils, topography and lithology) as mapped by Kovac 
et al. (1990). The disjunct population near Carcoar 
was associated with ‘Razorback’ soil landscape (steep 
to rolling topography with shallow well-drained soils 
derived from the Sofala Volcanics) and occurrences 
north-east of Orange were loosely associated with 
‘Panorama’ soil landscape (steep to level-crest 
topgraphy with moderately fertile soils derived 
from Tertiary basalt) and similar areas too small to 
be shown on Kovac et al.’s map. A close association 
with basic igneous material was reported from the 
Macquarie region north of Dubbo by Biddiscombe 
(1963, p. 20), who suggested that “soil nutrient status 
may be more decisive to E. albens than is moisture 
status’. However, further south in the South-eastern 
Riverina (Moore 1953) and Monaro (Costin 1954) 
regions, E. albens was reported to occur on a wide 
variety of soils and parent materials but generally on 
steep to undulating topography. 

The earlier supposition that E. albens did not occur 
on easterly aspects (i.e. those likely to be exposed 
to early morning sunlight) at high altitude was not 
supported by the expanded roadside survey. This was 
clearly indicated by its presence on easterly aspects at 
the two highest altitude sites. The preponderance of 
northerly and westerly aspects on the western part of 
the Central Tablelands was probably responsible for 
the earlier supposition. As all of the higher altitude 
occurrences were on slopes, it is likely that low 
slope rather than aspect may be a factor that limits its 
occurrence at high altitude. 


Differences between seedlings of E. melliodora 
and E. albens in the pot trial 

Seedlings at Cowra performed significantly 
‘better’ than those at Orange in all attributes measured - 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.S. SEMPLE AND T.B. KOEN 


Mean number 
of surviving seedlings 


Terrestial 
minima (°C) 


May/97 Jun/97 


Mean number 
of surviving seedlings 


@ 
eon? 2 


a etna! 


agseg $ 
8 @ go ie 


Terrestial 
minima (°C) 


May/97 Jun/97 


Jul/97 Aug/97 Sep/97 


Jul/97 =Aug/97 Sep/97 


—@— E£. albens ‘early’ 
—O— E. albens ‘delayed’ 
—v— E melliodora ‘early’ 
—V7— E. melliodora ‘delayed' 


Oct/97 Nov/97 Dec/97 Jan/98 


—@— E albens ‘early’ 

—O— E. albens ‘delayed! 
—v— E. melliodora ‘early’ 
—¥— E. melliodora ‘delayed’ 


Oct/97 Nov/97 Dec/97 Jan/98 


Figure 2. Survival of E. melliodora and E. albens seedlings under two morning light treat- 
ments, ‘delayed’ and ‘early’, during winter 1997 at Orange (870 m a.s.l.) and Cowra (380 m 
a.s.l.). Seedlings at Cowra were thinned on 15 September and numbers presented have been ad- 
justed for this. All seedlings were relocated to an early light area at Cowra on 13 October 1997. 
Also shown are daily terrestrial minimum temperatures at each site. Open circles indicate frosts. 


(Table 4). Conditions at Orange were particularly 
harsh during the 1997 winter and this was exaggerated 
by seedlings being in uninsulated above-ground pots 
where freezing of the topsoil occurred in midwinter. 
This was probably uncommon under normal conditions 


Proc. Linn. Soc. N.S.W., 126, 2005 


at Orange and seedlings may have experienced 
conditions more typical of an altitude that was well 
above the site — a suggestion that would also apply to 
the Cowra seedlings. Seedling deaths at Orange were 
mainly attributed to frost heave and ‘wilt’. Other 


al 


DISTRIBUTION OF WHITE BOX 


Table 4. Differences in mean survival (at October 1997), numbers of leaf-pairs (Sep- 
tember 1997) and mean stem height (September 1997 and January 1998) between 

seedlings of two eucalypt species exposed to two light treatments at two sites. Within 
columns in each section, values followed by the same lower case letter are not signifi- 


cantly different (P = 0.05). 


Survival Leaf-pairs log (Height+1) Height (mm) 
% per seedling Sept 1997 Jan 1998 
SITE 
Orange (ca 870 m a.s.1.) 34.6a 3.15a 2.89 a (17.0) “ 407 a 
Cowra (ca 380 m a.s.1) 91.7b TA1b 4.39 b (80.0) 476 b 
Isd (5%) 19.9 0.73 0.19 61 
SPEGIES 
E. albens =e 4.53 a - - 
E. melliodora - 6.03 b - - 
Isd (5%) 0.73 
SITE x LIGHT 
Orange Early - 3.55a 2.97 a (18.5) - 
Orange Delayed - 2.75 a 2.81 a (16.7) - 
Cowra Early - 6.75 b 4.18 b (64.1) - 
Cowra Delayed - 8.07 ¢ 4.61 c (99.5) - 
Isd (5%) 1.03 0.27 
SPECIES x LIGHT 
E. albens Early - 4.08 a 3.50a (32.0) - 
E. albens Delayed - 4.97 a 3.82 b (44.8) - 
E. melliodora Early - 6.22 b 3.65 ab (37.5) - 
E. melliodora Delayed - 5.85 b 3.60 ab (35.6) - 
Isd (5%) 1.03 0.27 


“Back transformed means (mm) in parentheses. 


B Only statistically significant main order and interaction effects have been tabulated. 


presumed effects of frost at both Cowra and Orange 
were the production of shoots from leaf axils and the 
cotyledon or “proto-lignotuber’ area. 

Though seedling densities in the pots were 
initially higher than would be expected in cases of 
natural regeneration, it was unlikely that the results 
were confounded by the effects of competition, 
which if operative, would have been more likely to 
affect growth than survival. At Orange, pots were 
not crowded due to the many deaths. Also, young 
woodland eucalypts, including E. melliodora, grow 
little if at all during winters at Orange (Semple and 
Koen 2001). At Cowra, where seedlings did grow and 


178 


deaths were few, densities in all pots were similar, 
including after thinning in September 1997. 
Competition effects would have been constant across 
treatments and hence, would not have confounded the 
relativity of the results. 

There was no significant difference between the 
survival rates of the two species but it was possible 
that the susceptibility of the smaller E melliodora 
seedlings to the unusual occurrence of frost heave at 
Orange (Table 3) may have masked differences at that 
site and possibly across sites in the analysis. The only 
significant difference detected between the two species 
was higher numbers of leaves in E. melliodora. The 


Proc. Linn. Soc. N.S.W., 126, 2005 


W.S. SEMPLE AND T.B. KOEN 


results therefore suggest that although E. melliodora 
seedlings may be slightly more frost tolerant than 
those of E. albens, the difference is unlikely to explain 
the rarity of E. albens, relative to E. melliodora, at 
altitudes up to ~900 m a.s.l. or in ‘frost hollows’ at 
lower altitudes. 

The fact that some E. albens seedlings survived 
in uninsulated pots at an altitude of 870 m a.s.l. 
at Orange, together with a natural occurrence of 
mature trees at 925 m a.s.l., is further evidence that 
alternative explanations need to be sought for the 
rarity of E. albens on the Central Tablelands. Though 
it is possible that it may have been more common 
before European settlement in the mid 1800s and 
since selectively removed, Cambage (1902), who 
travelled widely in this area, did not record it at any 
site where it is currently absent. 


Differences between delayed and early light 
treatments in the pot trial 

Pots in the Delayed Light treatment were 
exposed, at least at the colder Orange site, to a longer 

period of frosting, which may have enhanced the 
formation of ice crystals in the soil. Hence, seedlings 
in this treatment had increased likelihood to damage 
by frost heave but a lower likelihood of cold- 
induced photoinhibition. As any one or more of these 
factors may have affected the results, they cannot be 
considered separately. 

Despite deaths being more common at Orange 
in the Early Light treatment during early winter (Fig. 
2), differences in survival rates between the two 
treatments were not evident by spring. Significant 
differences in growth indices were evident at Cowra 
and across sites for E. al/bens such that seedlings in the 
Delayed Light treatment were taller and/or had more 
leaves than those exposed to Early Light. However 
by January, following a period of enhanced growing 
conditions, these effects were far less pronounced 
(Table 4). 

The pot trial results, together with natural 
occurrences on high altitude sites with easterly 
aspects, suggested that early morning sunlight’s 
supposed adverse effect on frosted seedlings 
(photoinhibition) was not a useful hypothesis for 
explaining the presence/absence of E. albens on the 
slopes and tablelands. 


CONCLUSIONS 
It was hypothesised that the local distributions of 


E. albens and E. melliodora on the Central Tablelands 
(where E. melliodora is relatively common and E. 


Proc. Linn. Soc. N.S.W., 126, 2005 


albens is rare) and adjacent Central Western Slopes 
(where E. albens tends to occur on upper slopes and 
E. melliodora on lower slopes) could be explained 
by different responses to frost. Pot trials at high and 
low altitude sites during the 1997 winter suggested 
that seedlings of both species were relatively frost 
tolerant. Though seedlings exposed to delayed 
morning sunlight were taller and/or had more leaves 
than those exposed to early morning sunlight, the 
differences were marginal and relatively short-lived 
when conditions for growth improved. Neither light 
treatment significantly affected seedling survival. 
Difference in frost tolerance between the two species 
was therefore an unlikely explanation for their 
distribution pattern on the slopes and tablelands. 

Observations of the occurrence of E. albens at 
high altitude in the field also supported its apparent 
tolerance to low temperatures (at least up to 925 m 
a.s.l.) regardless of aspect. Differences in geology, 
slope, drainage and/or soils are probably more 
important factors than exposure to frost in explaining 
the localised occurrence of E. al/bens on the Central 
Tablelands and adjacent slopes. 


ACKNOWLEGEMENTS 


We are indebted to numerous people for assisting with 
this project. Ifeanna Tooth and Jean Metcalfe maintained 
the seedlings and carried out many of the measurements 
at Cowra. Madeleine Rankin and Pat Farrelly accessed 
meteorological records at Cowra and Orange respectively. 
John Lawrence and Michael Sharp checked some of the 
high altitude records reported in Table 1. Kristina McColl, 
Phillip Wierzbowski and the late Doug Boland accessed E. 
albens databases (National Herbarium of NSW, Victorian 
Flora Information System and Eucalist respectively). 
Ashley Leatherland prepared the E. albens distribution map 
and Des Lang provided constructive criticism of an earlier 
manuscript. Thanks to you all. 


REFERENCES 


Anderson, R.H. (1968). “The Trees of New South Wales’ 
(fourth edition). (NSW Department of Agriculture / 
Government Printer: Sydney). 

Ball, M.C., Hodges, V.S. and Laughlin, G.P. (1991). Cold- 
induced photoinhibition limits regeneration of snow 
gum at tree-line. Functional Ecology 5, 663-668. 

Beadle, N.C.W. (1981). “The Vegetation of Australia’. 
(Cambridge University Press: Cambridge). 

Biddiscombe, E.F. (1963). A vegetation survey in the 
Macquarie region New South Wales. Division 
of Plant Industry Technical Paper 18 (CSIRO: 
Melbourne). 


179 


DISTRIBUTION OF WHITE BOX 


Boland, D.J., Brooker, M.I.H., Chippendale, G.M., Hall, 
N., Hyland, B.P.M., Johnston, R.D., Kleinig, D.A.and 
Turner, J.D. (1984). ‘Forest Trees of Australia’ (fourth 
edition). (Thomas Nelson / CSIRO: Melbourne). 

Bower, C., Semple, B. and Harcombe, L. (2002). 
‘Eucalypts of the Central West of NSW’(second 
edition). (Department of Land and Water 
Conservation: Orange). 

Cambage, R.H. (1902). Notes on the botany of the interior 
of New South Wales. Part VII — from Forbes to 
Bathurst. Proceedings of the Linnean Society of NSW 
27, 561-591. 

Costin, A.B. (1954). ‘A Study of the Ecosystems of 
the Monaro Region of New South Wales’. (NSW 
Government Printer: Sydney) 

Cremer, K.W. (1990). Frost. In ‘Trees for Rural Australia’ 
(Ed. K.W. Cremer) pp. 217-224. (Inkata Press: 
Melbourne). 

Egerton, J. (1996). Tree planting in cold climates — lessons 
from fundamental research. Australian Journal of Soil 
and Water Conservation 9(1), 37-42. 

Elliot, R., Blake, T. and Brownlie, J. (1984). ‘A Field 
Guide to the Grampians Flora’ (revised edition). 
(Algona Publications: Northcote). 

Hall, N., Johnstone, R.D. and Chippendale, G.M. (1970). 
‘Forest Trees of Australia’ (third edition). (Australian 
Government Publishing Service: Canberra). 

Harwood, C.E. (1980). Frost resistance of subalpine 
Eucalyptus species I. Experiments using a radiation 
frost room. Australian Journal of Botany 28, 587-599. 

Kovac, M., Murphy, B.W. and Lawrie, J.W. (1990). ‘Soil 
Landscapes of the Bathurst 1:250 000 Sheet’. (Soil 
Conservation Service of NSW: Sydney). 

Moore, C.W.E. (1953). The vegetation of the South- 
eastern Riverina, New South Wales I. The climax 
communities. Australian Journal of Botany 1, 485- 
547. 

Prober, S. (1996). Conservation of the grassy white 
box woodland: rangewide floristic variation and 
implications for reserve design. Australian Journal of 
Botany 44, 57-77. 

Prober, S.M. and Thiele, K. (1993). The ecology and 
genetics of remnant grassy white box woodlands in 
relation to their conservation. Victorian Naturalist 
110, 30-36. 

Semple, W.S. (1997). Eucalypt regeneration in white box 
(Eucalyptus albens Benth.) communities. M. Litt. 
thesis, University of New England, Armidale. 

Semple, W.S. and Koen, T.B. (2001). Growth rate and 
effect of sheep browsing on young eucalypts in an 
anthropogenic Themeda grassland. The Rangeland 
Journal 23, 182-193. 

Steel, R.G.D and Torrie, J.H. (1960). ‘Principles and 
Procedures of Statistics’. (McGraw Hill Book 
Company Inc.: New York). 


180 


Proc. Linn. Soc. N.S.W., 126, 2005 


Collections of Galerina (Agaricales, Fungi) Made by J.B.Cleland 
and Housed in the State Herbarium of South Australia 


ALEC Woop 


School of Biological, Earth and Environmental Sciences, 
University of New South Wales, Sydney, 2052, NSW, Australia 


Wood, A. (2005). Collections of Galerina (Agaricales, Fungi) made by J.B.Cleland and housed in the 
State Herbarium of South Australia. Proceedings of the Linnean Society of New South Wales 126, 181- 


196. 


Twenty-five collections by J.B. Cleland of Galerina (or which have been regarded as possibly belonging 
to Galerina) have been studied and their true status has been determined. Details are provided of the size 
and state of the collections and results are given of microscopic analysis of the material. The results have 
been evaluated in the light of recent taxonomic studies and suggestions are provided about the taxonomic 
position of each of the collections, together with discussions of the reasons for the conclusions. Ten of the 
collections have been shown to belong to genera other than Galerina. All the remaining collections have 
been assigned to previously described species of Galerina - G. lurida, G. marginata, G. muscolignosa, G. 


unicolor, G. vittiformis. 


Manuscript received 16 December 2003, accepted for publication 20 October 2004. 


KEYWORDS: Agaricales, Australia, Cleland, Galerina, herbarium, mushrooms. 


INTRODUCTION 


J.B.Cleland (1878-1971) was one of the great 
collectors of Australian larger fungi over a long period, 
first in New South Wales and later and particularly 
in South Australia. His work on fungi was largely 
summarised in his ‘Mushrooms and Toadstools and 
other Larger Fungi of South Australia’ (1934-1935), 
though he continued collecting until very late in his 
life. The basic material of his book has been revised by 
Grgurinovic and published as ‘Larger Fungi of South 
Australia’ (1997). The collections of Galerina were 
re-examined as part of the work for a forthcoming 
volume by Australian Biological Resources Study 
on some genera of the Family Cortinariaceae in 
Australia (in press 2005). A summary of these 
results is published there, but this paper in addition 
provides a full documentation of the status of each 
of the collections, notes on the size and state of each 
collection together with details of the original Cleland 
collecting notes. Some of these collections were in 
part discussed by Cleland in his first paper, Australian 
Fungi: Notes and Descriptions. No. | (Cleland and 
Cheel 1918). Cleland’s understanding of concepts of 
European species was guided particularly by the work 


of Rea (1922). The earlier work of Massee (1892- 
1895) was important for Rea and the illustrations of 
Cooke (1881-1891) were also a major influence. The 
use of European names should be interpreted in this 
light, with due consideration for later European use 
of these same names, particularly by Watling and 
Gregory (1993). 


MATERIALS AND METHODS 


Material was examined in 5% KOH and stained 
with Congo Red. Spores were examined and drawn 
at x2000. Cystidia were drawn at x1000. Spore 
shapes were named following the nomenclature of 
Bas (1969). Spore details are reported as a range 
of spore sizes, mean length and width (X) and 
mean ratio of length:breadth (Q). Ornamentation of 
the spores was recorded in terms of height (high, 
medium or low), width (coarse, fine) and shape 
of the tip of the ornamentation (blunt, pointed). 

Cystidia shapes were described following 
the categories of Vellinga (1988). In many cases 
cystidia could not be recovered. Measurements 
of the cystidia, when the form is lageniform, are 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


recorded as length, width of the basal portion, width 
of the neck and width of the apex if it is inflated. 
In most cases, the collections were originally 
described as being species of Galera. As it was later 
determined that this was not a valid generic name, 
the name Galerina became the universally accepted 
generic name for these species (see Donk 1962). 

The species are arranged alphabetically according 


to the species names on the packets. The collections 
were first located in the Waite Agricultural Research 
Institute (ADW) and later the whole of the Cleland 
material was transferred to the State Herbarium of 
South Australia (AD). For completeness, both the 
earlier and the current numbers are cited. However 
the AD numbers are the current valid numbers. 


COLLECTIONS EXAMINED 


1. Galerina (Galera) campanulata 


AD-C 42538 


(ADW 13793) 


Microscopic details: Spores 15.0—18.0 x 9.6—10.5 um, X= 16.3x 10.3 um, Q = 1.59, strongly ferruginous, 
oval to elliptic, wall thick, smooth, with no visible perispore, with broad very evident apical germ-pore. 
Cystidia sparse, scattered, fusiform to lageniform, never capitate. Basidia clavate to pyriform, two-spored. 
Pileal surface mostly collapsed, but seems clearly to be a distinct thin layer of thin-walled globose cells. 


Packet label: 
Galerina (Galera) campanulata 
Milson Is. 10/11/14 J.B.Cleland 


+Pencil annotation 
( = Galerina) 
on rich soil Milson Is (Sydney) NSW 


Cleland Notes: (sparse) 


Gills narrow, ascending, adnate. Cap pallid brown. Stem hollow, almost white (brownish tint). 


On rich soil. 

Milson Island 

10/11/14 

Galera 

Spores 13.6—15.5 x 8.5 um. 


Collection: The collection consists of six fruit-bodies, each about 1 cm in diameter, cap convex to conical, 


with a long thin stipe. 


Notes: This species was named by Cleland as Galera campanulata Massee, a species which in the British 
Fungus Flora (Dennis, Orton and Hora, 1963) is regarded as a doubtful species, and is not recorded by 

Watling and Gregory (1993) or Moser (1983). Cleland, in the discussion of this species in the 1918 paper, 
compares it to Galera silignea and also discusses differences between this species and Galera tener. Since 


these are now regarded as Conocybe species, this suggests that Cleland regarded this collection as belonging 
to what can now be regarded as Conocybe. The description of this species by Rea (1922) probably represents 
current interpretation of this species at the time: pileus deep cinnamon, persistently campanulate; stipe pallid, 
base darker; gills tawny cinnamon; spores 12 x 7 um; smell strong; by roadsides. 

The microscopic features detailed above, smooth spores with germ-pore and cellular cuticle, seem to 
clearly indicate a species of Conocybe. The cystidia of the related species, C. tener, are lecythiform with a 
distinct globular apex. Hence if the rarely found cystidia are representative of the collection, it is not close to 
Conocybe tener, but belongs in a quite different section of the genus. 

If the few cystidia found above are regularly present, this collection could still be a Conocybe but of 
a different section. At this stage the identity of the collection remains in doubt, but the genus is clear, the 
smooth spores with apical germ-pore could not be Galerina, particularly with a cellular cuticle. The other 
less likely possibility would be a species of the genus Pholiotina section Piliferae, but this genus is less often 
found in Australia than Conocybe. 


182 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


2. Galerina (Galera) campanulata AD-€ 42539 (ADW 13794) 

Microscopic details: Spores 10.2—11.1 x 7.2-8.4 um, X = 10.6 x 8.0 um, Q = 1.34, pallid dull ferruginous, 
blunt ovoid, apex depressed with fairly indistinct germ-pore, smooth, with no visible perispore, wall visibly 
thickened. No cystidia could be recovered. Basidia clavate to pyriform, clearly four-spored. 


Packet label: 

Galera campanulata 

Neutral Bay 18/12/14 J.B.Cleland 
+ Pencil annotation 

Galerina 

on wet ground in lawn (Sydney) NSW 


Cleland Notes: Collection envelope has note on outside — 
Cap conico-campanulate, yellowish brown, darker towards summit, edge slightly striate. Gills pale fawn. 
Stem white, hollow, brittle, shining. 
On wet ground in lawn. Neutral Bay. 18/12/14. 
Spores yellow brown, oval, with several small vacuoles, 13—13.8 x 7.7—8.5 um. 
A single paper slip inside the envelope has the same details with the following variation 
Cap about 1/2” diameter. Conico-campanulate. Slightly sticky. Tawny brown, darker towards summit. 
Slightly striate. 
Gills pale cream. 
Among grass 
Neutral bay 
19/2/14 


Collection: The collection consists of a single fruit-body. 


Notes: This collection seems clearly related to the previous collection, though the spores seem different. 
Possibly they represent two-spored and four-spored variants of the same species. The absence of cystidia 
makes more certain results almost impossible. See under the previous collection (AD-C 42538) for more 
discussion of the other possibilities. 


3. Galerina (Galera) lateritia AD-C 42540 (ADW 13788) 

Microscopic details: Spores 10.8-12.9 x 7.5-8.4 um, X = 12.2 x 8.1 pm, Q = 1.51, clear ferruginous, ovoid, 
wall smooth, apex a little thin, with an indistinct germ-pore, with no visible perispore. Basidia clavate to 
pyriform. No cystidia could be located. Pileal surface collapsed and difficult to reconstruct, but seems to be a 
thin complete layer of globose thin-walled cells. 


Packet label: 

Galera lateritia Adelaide 22/9/13 J.B.Cleland 
+ Pencil annotation 

Galerina 

Amongst grass S. Aust. 


Cleland Notes: 

Pileus conical 3/4 x 3/4”, very pale fawnish white, ? slightly striate. Stem white 2”, slightly bulbous, 
attenuated up, finely striate, no ring, hollow. Gills fairly close, narrow, pale fawn, just adnexed, hymen... 
mec: 

Among moss. Adelaide. 22/9/13. 


Collection: The collection consisted of three good substantial fruit-bodies, each about lcm in diameter. 


Notes: This, and the next two collections probably should be regarded as the same species as the characters 
are very similar but this would require cystidia to decide the issue. Clearly they are not Galerina as they have 


Proc. Linn. Soc. N.S.W., 126, 2005 183 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


a cellular cuticle and smooth spores. Clearly on the characters available, this is a species of Conocybe, or 
possibly a Pholiotina Section Piliferae, but cystidia and other features would be needed to decide the issue. 
Conocybe seems to be the most likely genus. 

Galerina lateritia is now regarded as being the same as Conocybe lactea, which is a very pale species. 


4. Galerina (Galera) lateritia AD-C 42541 (ADW 13789) 

Microscopic details: Spores 13.5—14.1 x 7.5—8.7 um, X = 13.6 x 8.2 um, Q = 1.70, deep brown to chocolate 
in mass, ovoid, thick-walled, smooth, with clear narrow apical germ-pore. Basidia pyriform, mostly partly 
collapsed. No cystidia could be recovered, even though the material appeared to be in good condition. 


Packet label: 
Galera lateritia 


Sydney 20/3/14 J.B.Cleland 
+ Pencil annotation 

Galerina 

Amongst grass, Sydney, NSW (+ formalin specimen) 
Cleland Notes: 


Pale brownish fawn, apex particularly conical, about 3/8 x 3/8”, apex acute to obtuse. Gills reddish 
brown, narrow, crowded, just free. Stem whitish, silky. Hollow, attenuated up, thin, | 3/4”. 
Amongst grass; Sydney 20/3/14 

Galera ? lateritia. Formalin specimen. 

Other collection like this has spores brown, 12.5 x 7—7.6 pm. 


Collection: The collection is of 6-7 fruit-bodies which are in good condition; the dried fruit-bodies are up to | 
cm in diameter. 


Notes: This is clearly not a Galerina. This collection and the next one clearly have a cellular cuticle and 
either belong to the genus Conocybe or to Pholiotina Section Piliferae. These three collections may represent 
variants of the same species or they may represent two close but different species. They clearly do not 
represent Conocybe /ateritia as it is now understood, as this is a very pale species. At the moment it seems 
best to regard all three collections as belonging to the same species. 


5. Galerina (Galera) lateritia AD-C 42542 (ADW 13790) 

Microscopic details: Spores 12.0—15.0 x 8.49.6 um, X = 12.8 x 8.7 um, Q = 1.47, deep ferruginous, oval, 
thick-walled, smooth, with no visible perispore, with distinct apical germ-pore; basidia large, pyriform, often 
collapsing. No cystidia of any kind could be recovered. 


Packet label: 
Galera lateritia 
Milson Is. 29/11/14 J.B.Cleland 
+ Pencil annotation 
Galerina 
Amongst grass Milson Island, NSW 
Kew No 6 see also Formalin Specimen No 27 


Cleland Notes: 

Conical then expanded to become broadly conical with pointed umbo, umbo dark tan, rest pale tan, 
densely striate. Gills narrow, very crowded, adnate, yellowish brown. Stem | 3/4” white, finely streaked, 
attenuated upwards, hollow, base slightly bulbous. 

Amongst grass Milson Island 29/11/14 
Spores yellow brown, oval slightly oblique, 12 x 8 pm. 


Collection: The collection has two fruit-bodies, together with three fruit-bodies glued to the back of the 


184 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


collecting slip, each about | cm diameter. 


Notes: This is clearly not a Galerina species, because of the smooth spores with germ- pore. Obviously it is 
close to the previous collections. In the absence of any cystidia, it should be regarded as another collection of 
the previous species, 1.e. a Conocybe or Pholiotina Section Piliferae species. The former should be regarded 
as the more probable in terms of the known frequency of the two genera in Eastern Australia. 


6. Galerina mycenoides AD-C 42543, — (ADW 13715) 

Microscopic details: Spores 7.5—8.1 x 4.5—5.1 um, X = 7.9 x 4.9 um, Q = 1.62, pale golden, broadly ovoid, 
thin-walled, smooth, without apical germ-pore, with no visible perispore. Basidia narrowly clavate, four- 
spored. Cheilocystidia abundant 60-80 x 3—8 um, cylindrical to fusiform or narrowly lageniform, apex often 
slightly enlarged 4—7 um; no pleurocystidia could be recovered. 


Packet label: 

Pholiota mycenoides 

Orange NSW 10/10/16 J.B.Cleland 
+Pencil annotation 

Galerina mycenoides 

on ground 


Cleland Notes: 

Pholiota. Cap convex then expanded, centre dimpled when moist, somewhat chestnut and striate, drying 
to pallid tawny white. Gills adnate, cinnamony brownish, moderately close. Stem 1”, often wavy, 
brownish tan, white down often near base, base a trifle swollen, slightly hollow. Ring as superior whitish 
fibres, often obscure, sometimes very definite. 

On ground. Orange 10/10/16. 

Spores 7—8.5 x 4.24.5 um. 


Collection: The collection is of numerous fruit bodies singly in granules of clay soil, all dirty and granular. 


Notes: This is not a species of Galerina as the spores were thin-walled and smooth, without a germ-pore. 
Nor was this a species of Pholiota Section Aporini, as the spores are thin-walled and too pale. The collection 
probably represents a species of Tubaria even though the spores were not easily collapsing, nor were they 
reniform. It probably comes closest to Tubaria rufofulva which also has similar cheilocystidia, though the 
cap colours seem somewhat different. See Grgurinovic (1997) and Moser (1983) for other related species. 
Galerina mycenoides has larger, finely rough spores, and other different features. See also Rea (1922), who 
places Galerina mycenoides in Pholiota, and reports it as growing among moss. 


7. Galera hypnorum AD-CS5506 (ADW 13787) _ 

Microscopic details (from packet a): Spores 12.3—13.2 x 6.6—7.9 um, X = 12.6 x 6.9 um, Q = 1.80, well 
coloured, elliptic to amygdaliform, plage usually obvious, flat, smooth and usually with a distinct rim, 
perispore thin, mostly obvious, not swelling or loosening, ornamentation moderately low to low, coarse, 
blunt. Cheilocystidia fairly sparse, broadly lageniform; pleurocystidia absent. . 


Packet label: 

Galera hypnorum 

Greenhill Rd., 27/6/21 J.B.Cleland 
+ Pencil annotation 

Holotype Galerina nyula 

Amongst moss Adelaide 


Cleland Notes: 


Galera hypnorum 
Up to 1/4”, campanulate, umbonate, striate, watery cinnamon, paler when dry. Gills tawny cinnamon, 


Proc. Linn. Soc. N.S.W., 126, 2005 185 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


tending to be distant, adnate. Stem up to 1”, brownish cinnamon, slender. Amongst moss. 
Greenhill Road 27/6/21 
Spores oblique, 11 to over 12.8 x 7.5, yellow brown. 


Collection: There are two sub-packets — 
a. with fragments of 2-3 fruit-bodies, with the label “winged spores”. 
b. with fragments of 4-5 fruit-bodies, with the label “ellipsoid spores”. 
There are no separate collecting notes for these individual fruit-bodies. 
Packet b. is clearly the collection referred to in Grgurinovic as the other collection. From it the following 
details were found: a 
Microscopic details: Spores 11.7—12.6 x 6.0—-8.7 um, X = 12.03 x 8.28 um, Q = 1.45, well coloured, 
ovoid to slightly elliptic, plage not obvious, a vague flat area above the apiculus, perispore not visible, 
ornamentation low to moderately low, fairly coarse, blunt. Cystidia could not be recovered. 


Notes: Sub packet a. Material clearly corresponds to the description of Galerina nyula in all the details 
published by Grgurinovic (Grgurinovic 1997), and doubtless corresponds to some of the records of Galerina 
hypnorum in Australia. However it is quite distinct from Galerina muscolignosa (see Wood 2001), which has 
distinctly calyptrate spores and which seems to be the common species in much of Eastern Australia. Also 
Galerina oreophila may also be confused with Galerina nyula, but Galerina oreophila has more distinctly 
lageniform cystidia, slightly broader spores, a mixture of two-spored and four-spored basidia and an alpine or 
sub-alpine habitat. 

Sub packet b. Material has microscopic features that suggest it may be a collection of a species of 
Cortinarius because of the lack of cystidia and the spores without a plage and without visible perispore. It 
probably represents a species of the subgenus Telamonia, but further identification will await more work on 
that sub-genus, and it would be made more difficult by the lack of any macroscopic field details. 


8. Galerina nyula AD-C5507, |. (ADW 13785) 

Microscopic details: Spores 7.8—9.3 x 4.8—-5.4 um, X = 8.9 x 5.2 um, Q = 1.72, fairly well coloured, elliptic 
to vaguely amygdaliform, plage flat, without rim, not obvious, appears smooth, perispore thin, often not 
obvious, often somewhat loosening but not fully calyptrate, ornamentation low to very low, somewhat coarse, 
blunt. The material was too fragmentary for cystidia to be recovered. 


Packet label: 

Galera hypnorum 

Lisarow 5/8/16 J.B.Cleland 
+ Pencil annotation : 

Galerina nyula 

Lisarow NSW 


Cleland Notes: No field notes were present. 


Collection: The collection consists of fragmentary parts of about three fruit-bodies, among moss, with conical 
mycenoid caps, small and conical to convex with long thin stems. Clearly among moss. 


Notes: The distinctive characteristics of the spores indicate that this collection does not represent Galerina 
nyula. Rather it should be regarded as belonging to Galerina muscolignosa despite a smaller degree of 
loosening of the perispore on the spores, because of the spore size and the level of ornamentation on the 
spores. 


9. Galera hypnorum AD-C 5508 — (ADW 137867) 

Microscopic details: Spores 7.5—9.6 x 5.4-6.3 um, X = 9.0 x 5.3 um, Q = 1.69, well coloured, elliptic to 
slightly amygdaliform, plage slightly flatter, with no distinct rim or smooth patch, perispore thin, often a little 
loosening or slightly calyptrate, ornamentation low, a little coarse, blunt. No cystidia could be recovered from 
the limited material. 


186 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


Packet label: 

Galera hypnorum 

Mosman 23/7/16 J.B.Cleland 
+ Pencil annotation 

Galerina nyula 


amongst moss Mosman (Sydney) NSW 
Miss Clarke (Watercolour) No 133 Formalin specimen 229 
Cleland Notes: 


Galera. Cap conico-campanulate 1/4”, base 5/16” high, ?without definite umbo, dark yellow brown, 
striate, drying pallid tan. Gills moderately distant, ascending, adnate, yellow brown, not ventricose . Stem 
up to 1”, slender, yellow brown. 

Amongst moss. Mosman 


Collection: The collection consisted only of fragmentary material. 


Notes: Comparisons with Collection AD-C 5507 above seem to indicate that it is the same species, and the 
same remarks apply. This collection also represents Galerina muscolignosa. 


10. Galera hypnorum AD-C5509— _ (ADW 13783) 

Microscopic details: Spores 8.4—9.6 x 5.4-6.3 um, X = 9.0 x 5.6 um, Q = 1.60, well coloured, elliptic, 
slightly amygdaliform in profile, plage smooth, flat, with a slight rim, ornamentation very low, moderate, 
rounded, perispore clearly present, thin, regularly loosening, sometimes variously in bubbles. No cystidia 
could be recovered from the limited material. 


Packet label: 

Galera hypnorum 

J.B.Cleland, no locality; no date 
+ Pencil annotation 

Galerina nyula 


Cleland notes: 
Only torn fragments in packet; only a few scraps can be partially reconstructed - Galera (new sp. ?) 
...apex of stipe mealy... 
...MOSS... 


Collection: The collection was very fragmentary, with no intact fruit-bodies remaining; one partial cap 
remained and produced the spores described above; the material was too fragmentary for cystidia to be 
recovered. 


Notes: Comparison with Collections AD-C 5507 and AD-C 5508 above seems to indicate that they represent 
the same species. Clearly they do not represent Galerina nyula for the reasons given above. It fits best within 
Galerina muscolignosa. 


11. Galera hypnorum AD-C 5510 (ADW 13784) 

Microscopic details: Spores 10.5—12.6 x 6.3—7.5 um, X = 11.7 x 7.1 um, Q = 1.66, well coloured, ovoid, 
some slightly pointed at apiculus, only rarely slightly amygdaliform in profile, plage not visible or distinct, 
without a rim, no visible perispore, ornamentation low to very low, a little coarse, rounded. Basidia often or 
mostly, two-spored. No cystidia could be recovered. 


Packet label: 


Galera hypnorum 
J.B.Cleland no locality or date 


Proc. Linn. Soc. N.S.W., 126, 2005 187 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


+Pencil annotation 
‘Galerina nyula’ 
vide Miss Clarke Watercolour 133 
Formalin spec 229 

(Perhaps same coll as AD-C 5508 ? G.Bell 02) 


Cleland notes 
No macroscopic details, a single slip has, in pencil (JBC) - spores 8—8.5 x 6 um, oval, peculiar double 
outline, ?wing at one end, yellow brown, edge a little turned in when young, so as to be globular (with a 
sketch of a globular head and two spores, clearly calyptrate). 

And around it, in the same hand (JBC), in fine ink 
Galera hypnorum Batsch 
Vide Miss Clarke Picture 133 Formalin specimen 229 
Rec. in Trans. Roy. Soc. SA XLII, 1918 p 119 


Collection: The collection consists of about seven fruit bodies in fair condition. The collection was clearly 
made from soil with moss. 


Notes: Cleland’s second set of notes (above) presumably may mean that he thought that it was the same 
species as the illustration he cited. He does not necessarily mean that this was a comment about this 
collection or that this one was the one that was painted by Miss Clarke. 

As the spores figured in Cleland’s notes are clearly calyptrate, and the spores of the current specimen are 
clearly not calyptrate, one suspects that the written slip in the packet does not correspond with the specimen 
and has been misplaced from elsewhere. This is confirmed by the spore sizes cited by Cleland (8—8.5 x 6 um) 
while the present specimens have much larger spores (10.5—12.6 x 6.3—7.4 um). 

From the details available from the specimens, as the spores are mostly produced on two-spored basidia, 
the spore size and morphology suggest this represents a collection of Galerina vittiformis possibly var. 
pachyspora. Final certainty could not be produced from the details that could be gained from the specimens. 
However, it is clear that the specimens do not represent Galerina nyula. 


12. Pholiota pumila AD-C 42544 (ADW 13720) 

Microscopic details: Spores 7.5—9.0 x 4.24.8 um, X = 8.2 x 4.8 um, Q = 1.71, golden, fairly thin-walled, 
elliptic to slightly amygdaliform, plage usually not marked, flat, smooth, sometimes with a small rim, 
perispore thin, sometimes a little swollen and occasionally slightly loosening, ornamentation low, fine, a little 
blunt. A few narrow lageniform non-capitate cystidia were recovered, both cheilocystidia and pleurocystidia 
present and of similar morphology. 


Packet label : 

Pholiota pumila 

Spit, Sydney 9/7/16 Amongstmoss  J.B.Cleland 
+Pencil annotation 

Galerina 

formalin specimen No 216 


Cleland notes: 

Pholiota. Cap 3/16”, broadly conical, faintly striate, apex rather pointed, yellow brown, finely granular, 
with less hygrophanous appearance. Gills adnate, very pallid brown, rather distant, with short ..!. Stem 
5/8”, attenuating up, moderately stout, pallid brown, somewhat silvery mealy, slightly hollow, ring 
superior ?definite. 

Amongst moss. Spit. 


Collection: The collection consists of only a few small fruit-bodies, clearly among moss. Dried fruit-bodies 
small and mycenoid. 


188 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


Notes: The finely rough spores clearly indicate a species of Galerina. If Cleland’s notes are accurately 
interpreted, with a ‘ring’, then the specimens almost certainly fit Galerina lurida. Though the spores seem 
more finely rough, all the other features fit Galerina lurida well. It was not possible to detect remains of a 
ring on the stipe of the dried specimens, but this would not be unusual with this species, where the texture of 
the ring is variable and hence it persists in differing degrees in mature specimens. On balance, this collection 
should be regarded as being of Galerina lurida. 


13. Pholiota pumila AD-C 42545 (ADW 13721) 

Microscopic details: Spores 7.8-9.0 x 4.5—5.4 um, X = 8.4 x 5.0 um, Q = 1.69, Golden ferruginous, 

elliptic to amygdaliform, plage large, flat, smooth, sometimes with a distinct rim, perispore thin, obvious, 
sometimes swelling irregularly and a little loosening, but not calyptrate, ornamentation low, fairly fine, blunt. 
Cheilocystidia fairly frequent, narrowly lageniform, apex distinctly rounded to slightly capitate, 40-50 x 
8-12 x 3-5 x 6-9 um, pleurocystidia similar, fairly frequent. 


Packet label: 

Pholiota pumila : 

Amongst moss Mosman 13/8//16 J.B.Cleland 
+Pencil annotation 

Galerina 


Cleland notes: 
Pholiota.....moss 
Cap 3/8”, convex then nearly plane, trace fibres, dark reddish brown, striate. 
Gills reddish brown, adnate, moderately close. Stem 1”, dark brown, slightly striate, solid 
? Film of rather dirty white ring. 13/8/16 


Collection: The collection consists of three fruit-bodies, in good condition. The specimens are clearly more 
substantial than those for collection 42544. 


Notes: Details of this collection are similar to those for collection AD-C 42544. Despite the differences in 
habit, they both should be regarded as specimens of Galerina lurida. Clearly, with rough spores, this is not a 
Pholiota. These collections may indicate that Galerina lurida is a somewhat variable species. 


14. Galerina subifinosa AD-C 42546 (ADW 13792) 

Microscopic details: Spores 10.5—12.0 x 7.5—8.7 um, X = 11.4 x 8.2 um, Q = 1.40, ferruginous, sometimes 

a little pale, ovoid or slightly elliptic to slightly amygdaliform, plage flat, smooth, with a low rim, not very 
strongly developed, perispore not visible, ornamentation moderate, coarse, blunt, apex not mucronate. 
Basidia mostly collapsed, clavate, mostly four-spored, with a few two-spored. Cheilocystidia fairly common, 
narrowly lageniform, apex rounded, not capitate, 50-75 x 7-14 x 2-7 x 5—7 um. Similar pleurocystidia also 
present. 


Packet label: Galerina subifinosa 

Mosman 30/7/16 J.B.Cleland among moss 
+Pencil annotation 

? G. rubiginosa 

? misreading of Cleland handwriting (G.Bell 02) 


Cleland notes: 
Spores 10.5—11 x 7.5 um, oblique, oval, finely rough, yellow brown. 
Cap 3/8", base to high, conico-campanulate, obtuse umbonate, dark chestnut, coarsely ribbed to umbo. 
Gills dark reddish brown, adnate, broad, moderately distant. 
Stem to 1 1/4”, slender, dark brown, slightly, hollow. 
Among moss. Mosman 30/7/16 
Collection: The collection consists of a single fruit-body only. 


Proc. Linn. Soc. N.S.W., 126, 2005 189 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


Notes: This collection seems clearly to belong to Section Galerina of Galerina because of the presence 

of clear pleurocystidia. In this group it seems to be part of the Galerina vittiformis complex. It does not 
correspond with Galerina vittiformis var. pachyspora because of the four-spored basidia, darker cap colours 
and slightly different spore shape. The name Galerina subifinosa appears to be an unpublished manuscript 
name and hence has no status. It seems probable that it represents a misreading of Cleland’s label, which was 
originally intended to be Galerina rubiginosa. This is given added support by the fact that apart from these 
two collections, no Galerina rubiginosa collections are found in the Cleland collections, when that species 
was recorded in the 1918 paper. Galerina rubiginosa, as it is now understood, is one of the species within 
the Galerina vittiformis complex, and it has been split between several species. The Cleland collections, with 
darker cap colours, slightly smaller spores and four-spored basidia do not clearly fit any of the current species 
or varieties. The nearest would be Galerina vittiformis, possibly as a new form or variety. It may represent 
Galerina vittiformis var. vittiformis f. tetraspora (see Singer and Smith 1964, and Breitenbach and Kranzlin 
2000). This collection seems to correspond to the one quoted by Cleland in the 1918 paper, with the note 
“Miss Clarke Watercolour No. 132”. 


15. Galerina subifinosa AD-C 42547 (ADW 13791) 

Microscopic details: Spores 10.5—12.6 x 6.6—7.8 um, X = 11.6 x 7.4 um, Q = 1.58, ferruginous, sometimes 
a little pale, ovoid or a little elliptic to slightly amygdaliform in profile, with distinct flat smooth plage, with 
slight rim, with no visible perispore, ornamentation moderate to low, coarse, blunt. Basidia clavate, much 
collapsed and reviving poorly, four-spored, with only a few two-spored. Cheilocystidia mostly collapsed, 
narrow lageniform, longish, not capitate. Similar pleurocystidia also clearly present. 


Packet label: 

Galerina subifinosa 

Mosman NSW amongst moss 30/7/16 J.B.Cleland 
+Pencil annotation 

(Galerina rubiginosa) 

(probably a misreading of J .B. Cleland handwriting G. Bell 02) 


Cleland notes: 
Pileus campanulate, 3/16” high, 3/8” broad, almost chestnut, striate, striae darker, not definitely viscid. 
Gills adnate, reddish brown, moderately distant. 
Stem slender 1”, reddish brown, slightly mealy, trace of being hollow. 
Amongst moss Mosman 30/7/16 


Collection: The collection consists of three fruit-bodies, in fairly good condition, fairly small, with some 
sandy soil and moss. 


Notes: This collection is clearly a Galerina species, and probably represents another collection of the species 
found above in collection AD-C 42546. Its identity is discussed fully there. 


16. Galerina (Pholiota) subpumila AD-C 11883 (ADW 12930) 

Microscopic details: Spores 9.3—-10.2 x 6.0-6.6 tum, X = 9.5 x 6.4 um, Q = 1.49, well coloured, golden, wall 
slightly thick, quite smooth, apical germ-pore small or narrow, but clearly distinctly present. Cheilocystidia 
narrowly lageniform, not capitate, 30-40 x 4-8 x 10-13 pm, a few similar pleurocystidia also present. 


Packet label: 

Pholiota subpumila 

Greenhill Rd., 12/6/26 J.B.Cleland 
+Pencil annotation 

Holotype 

(Greenhill Rd. runs between SE corner of Adelaide and summit of Mt. Lofty) 
Cleland notes: 


190 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


Pileus up to 5/8” to 1 1/8”, convex then flattened or a little depressed, umbonate when young, sometimes 
a little wavy, shining waxy looking, dark tan. 

Gills rather close, adnate or slightly decurrent, watery brown, rather triangular. 

Stem equal or slightly attenuated upwards, 1 1/2”, whitish, fibrillose with a slight tinge ? of cap colour. 
Spores yellow brown 8—9.5 x 5.5 um, oblique. 


Collection: The collection consists of five fruit-bodies, in good condition, with moss. 


Notes: With smooth spores, which also have a distinct germ-pore, the collection clearly represents a Pholiota, 
not a Galerina. It has been well re-described by Grgurinovic (1997) as Pholiota subpumila and the current 
study has confirmed the details given there. 


17. Galerina (Pholiota) subpumila AD =Ci2393 (ADW12931) 

Microscopic details: Spores 8.7—9.9 x 6.0—7.2 um, X = 9.09 x 6.45 um, Q = 1.41, golden, wall distinctly 
thickened, completely smooth, with narrow apical germ-pore, usually narrow but always distinctly present. 
Cheilocystidia narrow lageniform to lageniform, clearly present and fairly frequent, less frequent similar 
pleurocystidia also present. 


Packet label: 

Pholiota subpumila 

Greenhill Rd., 11/6/27 J.B.Cleland 
+Pencil annotation 

On moss 

(Runs between SE corner of Adelaide and summit of Mt. Lofty) 


Cleland Notes: 
No notes of macroscopic details. 
Spores yellow brown, 9 x 6.5—7 um 
On moss 


Collection: The collection consists of five fruit-bodies in good condition, with some soil and debris. 


Notes: This collection clearly matches all the features of Pholiota subpumila - see discussion under the 
previous collection and Grgurinovic (1997). 


18. Galerina (Pholiota) subpumila’ AD-C 12104 (ADW 12929) 

Microscopic details: Spores 7.8-10.5 x 5.7-6.6 um, X = 8.9 x 6.2 um, Q = 1.44, well coloured, golden, 
ovoid, wall moderately thickened, smooth, with small distinct apical germ-pore, constantly and clearly 
present. Cheilocystidia fairly frequent, fusiform to narrowly lageniform, pleurocystidia less common, but 
clearly present, of similar morphology. 


Packet label: 

Pholiota subpumila 

Waterfall Gully, SA 27/6/21 J.B.Cleland 
+Pencil annotation: 

amongst moss 

Miss Fiveash watercolour 24 

(34° 58’ S; 136° 41’ E ) 


Cleland notes: 
Cap 5/8” convex, umbonate (obtuse), pallid yellow brown, edge rather mealy. Gills dingy greyish brown, 
decurrent (slightly), moderately close, watery cinnamon. Stem up to | 1/2” pallid whitish, with a superior 
well-marked whitish ring, solid. Flesh watery. Amongst moss. Waterfall Gully 27/6/21 


Proc. Linn. Soc. N.S.W., 126, 2005 191 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 
Base of stem ?occasionally swollen. 
Spores rather a dull dark brown, thin-walled, ellipsoid but a little irregular, thick-walled 8 x 6.4 um. 
Collection: The collection consists of numerous fruit-bodies, in good condition, with soil and debris. 


Notes: Clearly this collection is Pholiota subpumila, as are the previous collections. For a discussion of this 
species, see there and Grgurinovic (1997). 


19. Galerina (Pholiota) subpumila AD-C 22424 (ADW 12928) 

Microscopic details: Spores 8.4—9.6 x 5.1—6.0 um, X = 9.1 x 5.6 um, Q = 1.62, strongly and deeply coloured, 
elliptic, profile distinctly amygdaliform, without apical callus and apex not less ornamented, plage large flat, 
smooth, obvious, with small rim, ornamentation high, coarse, blunt, perispore obvious, thick, swelling, but 
only occasionally slightly loosening, never calyptrate. Cheilocystidia and pleurocystidia both clearly present, 
similar, narrowly lageniform, apex slightly capitate but never abruptly so, 45-50 x 3—7 x 10-13 um. 


Packet label: 


Pholiota subpumila 

Eagle on the Hill 6/6/32 J.B.Cleland 
+Pencil annotation 

(34° 59” S ; 138° 40” E) near moss 

Galerina 


Cleland notes: 

Pileus ochraceous tawny XV '2” slightly convex, slightly umbilicate, substriate. Gills ochraceous tawny, 
adnate to decurrent, rather distant. Stem 3/4”, same colours, slender, slightly fibrillose. Ring indefinite, 
rather distant, stem cartilaginous. Flesh same colour. Near moss. 

Spores golden brown, 8 x 4.5 um, obliquely elliptic, perhaps slightly rough. 


Collection: This collection consists of three small fruit-bodies on a twig of wood, with some soil and debris. 
No velar remains are now visible on the stipe. Clearly the original fruit bodies were quite small. 


Notes: Because of the rough spores, with plage, without germ-pore, this collection is clearly not of Pholiota 
subpumila, but clearly represents a species of Galerina. The species that it might represent are G. marginata 
or G. lurida. It is probably best regarded as a small specimen of Galerina marginata, since Galerina lurida 1s 
clearly not lignicolous and it has spores and cystidia that are slightly different from the present collection. 


20. Galerina (Pholiota ) subpumila AD-C 22425 (ADW 12932) 

Microscopic details: Spores 8.4—9.3 x 5.1-6.0 um, X = 8.9 x 5.6 um, Q = 1.59, well coloured, ovoid to 
elliptic, amygdaliform in profile, plage obvious, smooth, mostly with a distinct rim, ornamentation obvious, 
moderately low, coarse, blunt, perispore marked, swollen, with some slight loosening but never calyptrate. 
Cheilocystidia and pleurocystidia clearly present, but not abundant, of similar morphology. Narrowly 
lageniform, apex rounded or sometimes vaguely capitate, 40-50 x 12-16 x 4-7 x 4-9 um. 


Packet label: 

Pholiota subpumila 

Coromandel Valley 26/6/27 J.B.Cleland 
+Pencil annotations 

Galerina 

amongst moss 

Clarendon, Coromandel Valley, SA. 


Cleland notes: 


Pholiota. Moss. Cap near Sudan Brown III. Stem paler cap, drying to Tawny Olive XXIX. Stem up to 3”, 
equal, or attenuating up, ring, pallid, subsuperior, not striate. Not hygrophanous. Spores 7.5 x 5.2 um, 


192 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


oblique, rather ovate, dark yellow brown. 


Collection: This collection consists of three fruit-bodies, each fairly substantial, clearly from among soil and 
moss and clearly not on wood. One of the dried fruit-bodies shows traces of a fine fibrillose ring. 


Notes: Because of the rough spores with a plage and without a germ-pore, this collection is clearly a species 
of Galerina and not of Pholiota. Hence the identification as Pholiota subpumila is incorrect. The substantial 
fleshy habit and habitat on soil, not on wood, clearly point to a good collection of Galerina unicolor if that 
species is recognised as being separate from Galerina marginata (for discussion of this point, see Wood 
2001). 


21. Galerina unicolor AD-C 42548 — (ADW 13728) 

Microscopic details: Spores 9.3—10.5 x 5.7—-6.3 um, X = 10.0 x 6.0 um, Q = 1.68, elliptic to amygdaliform, 
plage smooth, often marked with abrupt margin, ornamentation moderately low, coarse, blunt, perispore 
obvious, thin, not swelling or loosening at all. Cheilocystidia and pleurocystidia not in good condition, sparse 
but clearly present, similar, narrowly lageniform, clearly not bifurcate. 


Packet label: 

Pholiota unicolor 

Lawn 6/16 J.B.Cleland 
+ Pencil annotation: 

Galera 

(probably Sydney NSW) 


Cleland notes: 
No macroscopic details 
Spores yellow from 8.5 x 5.2 um ?over 10.5 x 5.2 um, ?oblique .... swollen hyphae 


Collection: The collection consists of numerous fruit-bodies, in good condition, clearly fleshy, with moss 
and debris, and one fruit-body clearly on heavy bark, but substrate connection not clear for any of the other 
specimens. 


Notes: On balance, this collection should be retained as Galerina unicolor since there is no certainty that 
the substrate was wood. But note that if consensus on species limits changes, this would become Galerina 
marginata. 


22. Pholiota unicolor AD-C 42549 (ADW 13729) 

Microscopic details: Spores 8.4—9.6 x 5.4-6.3 um, X = 9.0 x 5.9 um, Q = 1.54, elliptic to amygdaliform, 
plage distinct, flat, smooth, with distinct rim, ornamentation low to very low, coarse, blunt, perispore mostly 
obvious, usually thick and somewhat swollen, some slightly loosening, but never calyptrate. Cheilocystidia 
and pleurocystidia sparse and difficult to find, but both clearly present and similar, narrowly lageniform, 
always simple, never bifurcate. 


Packet label: 

Pholiota unicolor 

Mt. Wilson 6/6/15 J.B.Cleland 
+Pencil annotation 

Galerina 

Mt. Wilson (NSW) 

Kew No 24 Miss Clarke Watercolor No 85 


Cleland notes: 


Pholiota. Pileus at first deeply reddish tan, drying to pale brown, smooth, convex, umbonate, 1”. Gills 
adnate, reddish brown, moderately close. Ring ?moderately distant, marked. Stem | 1/2”, slightly 


Proc. Linn. Soc. N.S.W., 126, 2005 193 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


attenuated upwards, base a little swollen, covered with whitish mealy fibrils, brownish below, solid. 
On a separate piece of paper: Spores 8.5 to 10.4 x 5.2 1m, oblique. Chrysalis-like. Brown. 


Collection: A good collection of five fruit-bodies in good condition, all growing on old wood. 


Notes: On balance, because of the substrate, this collection should be regarded as Galerina marginata, 
because the spores have a thick perispore and low wall ornamentation. The wood substrate is quite clear and 
underlines the species identification. 


23. Pholiota unicolor AD-C 42550 — (ADW 13730) 

Microscopic details: Spores 8.7—9.9 x 5.1-6.6 um, X = 9.1 x 5.6 um, Q = 1.63, ovoid to elliptic or 
amygdaliform, mostly with clear large smooth flat plage, with distinct rim, ornamentation low, a little narrow, 
blunt, perispore sometimes not obvious, often fairly thick and swollen and some with a slight degree of 
loosening. Cheilocystidia and pleurocystidia, sparse and difficult to recover, but both similar and clearly 
present, narrowly lageniform, always simple and never bifurcate. 


Packet label: 
Pholiota unicolor 


Lisarow 5/8/16 J.B.Cleland 
+Pencil annotations 

Galerina 

on trunks (fallen) Lisarow (NSW) 


Cleland notes: 
(Pileus) 3/4”, nearly plane, dingy darkish brown and finely striate, drying to pallid brownish. Gills adnate, 
dingy cinnamon. Stipe dirty brown, fibrillose, ?streaked. 
On trunks (fallen). Lisarow 5/8/16 
Spores dull brown, oblique 8.5—9.0 x 4—5 ym. 


Collection: The collection consists of eight small fruit-bodies in good condition, clearly attached to wood 
fragments. There is no visible annulus on the dried material. Habit naucorioid, but much smaller than the 
previous collections. 


Notes: Since Cleland called this collection Pholiota unicolor one assumes the presence of some kind of ring 
and some degree of robust stature. If this is so, there is no reason for this collection not to be regarded as 
being Galerina marginata since it 1s clearly on wood. 


24. Pholiota unicolor AD-C4A2S55 iy Oe(AD WHS 731) 

Microscopic details: Spores 8.1—9.0 x 4.5—5.1 um, X = 8.4 x 4.8 um, Q = 1.76, elliptic to amygdaliform, 
plage large, flat, smooth, mostly without a rim, ornamentation low, fairly fine, blunt, perispore thin, not very 
obvious, sometimes distinctly swollen and occasionally a little loosened. Cheilocystidia and pleurocystidia 
sparse, difficult to recover, but both clearly present and with similar morphology, narrowly lageniform, apex 
simple, never bifurcate. 


Packet label: 

Pholiota unicolor 

Lisarow 5/8/16 J.B.Cleland 
+Pencil annotations 

Galerina 

Scattered on fallen trunks Lisarow NSW 


Cleland notes: 


Pileus | 1/4 “, convex acutely umbonate, watery yellow brown and edge finely striate, drying pallid 
brown, smooth. Gills moderately close, adnate with a decurrent tooth, pale cinnamon, becoming dingy 


194 Proc. Linn. Soc. N.S.W., 126, 2005 


A. WOOD 


cinnamon. Stipe up to 1 1/2” ?slender, base somewhat swollen, pallid to brownish, fibrillosely streaked. 


Ring superior, often slight. 


Differs from P. unicolor in being larger, umbo marked, gills not triangular. 


Collection: The collection consists of six fruit-bodies on wood or heavy bark. 


Notes: The collections were made clearly from wood and the specimens are clearly somewhat fleshy. Though 
they appear larger than the previous collection, they probably are still smaller than many current collections. 
However, it should be regarded as a good collection of Galerina marginata. 


25. Psilocybe foenisecii AD-C 5608 


(ADW 13155) 


Microscopic details: Spores 10.5—12.3 x 7.2-8.7 um, X = 11.6 x 7.7 um, Q = 1.51, ovoid, wall distinctly 
thick, with small distinct germ-pore, distinctly and clearly smooth. Cheilocystidia present, ventricose, 


pleurocystidia absent. 


Packet label: 
Psilocybe foenisecii 
Ryde 

+Pencil annotations: 
Galerina sp. 
On roadsides 


NSW 27/5/16 


‘Cleland notes: 
No macroscopic description present. 


J.B.Cleland 


P. foenisecii ?. Spores dull dark brown not definitely purple, 8.5—10.5 , 


occasionally 12 x 5.2 to7 um 


(Cap) brown and striate when moist, pallid white when dry. 


On roadsides. Ryde 27/5/16 


Collection: The collection consists of several small fruit-bodies in fairly good condition, on soil debris with 


some small plant material. 


Note — AMY 1986 ‘not P. foenisecii, spores smooth, probably a Galerina.’ 


Notes: Clearly this collection matches the description of Psilocybe korra Grg. (See Grgurinovic 1997) There 
this collection is cited as the only other collection in addition to the type (AD-C 5609) from Adelaide. Other 
collections under the same name are correctly named Panaeolus foenisecii, which has verrucose spores, a 
cellular cap cuticle and different cystidia. That species is now usually regarded as Panaeolina foenisecii. 


ACKNOWLEDGMENTS 


The invaluable assistance of Graham Bell of the South 
Australian Herbarium (AD) in locating the specimens 
and providing important documentation is gratefully 
acknowledged. The assistaance of Pam Catcheside in 
deciphering the handwriting of Cleland is also gratefully 
acknowledged. Financial assistance for this work by 
the Australian Biological Resources Study is gratefully 
acknowledged. Facilities for the later part of this work were 
kindly provided by Professor Anne Ashford. 


Proc. Linn. Soc. N.S.W., 126, 2005 


REFERENCES 


Bas, C. (1969). Morphology and Subdivision of Amanita 
and a Monograph of the Section Lepidella. Persoonia 
5, 285-579. 

Breitenbach, J., and Kranzlin, F. (2000). “Fungi of 
Switzerland. Vol. 5. Agarics Part 3.’ (Mykologia: 
Lucerne). 

Cleland, J. B. (1934). ‘Toadstools and Mushrooms and 
Other Larger Fungi of South Australia.’ (Government 
Printer: Adelaide). 


195 


THE CLELAND COLLECTIONS OF GALERINA (FUNGI) 


Cleland, J. B. and Cheel, E. (1918). Australian Fungi: 
Notes and Descriptions. No. |. Transactions and 
Proceedings of the Royal Society of South Australia 
42, 88-138. 

Cooke, M.C. (1881-1891). ‘Illustrations of British Fungi.’ 
8 vols. (Williams and Norgate:London.) 

Dennis, R.W.G., Orton, P.D., and Hora, F.B. (1960). New 
Check list of British agarics and boleti. Transactions 
of the British Mycological Society 43, Supplement: 
1-225. 

Donk, M.A. (1962). The Generic Names Proposed for 
Agaricaceae. Beihefte zur Nova Hedwigia. Heft 5. 
(Cramer: Weinheim). 

Grgurinovic, C. A. (1997). ‘Larger Fungi of South 
Australia.’ (Botanic Gardens of Adelaide and State 
Herbarium and Flora and Fauna of South Australia 
Handbooks Committee: Adelaide). 

Massee, G.E. (1892-1895). “British fungus flora.’ 4 vols. 
(George Bell and Sons:London). 

May, T. W. and Wood, A. E. (1997). “Catalogue 
and Bibliography of Australian Macrofungi 1. 
Basidiomycota p.p..’ (Australian Biological 
Resources Study:Canberra). 

Moser, M. (1983). “Keys to Agarics and Boleti 
(Polyporales, Boletales, Agaricales, Russulales).’ 
(Roger Phillips:London). 

Rea, C.R. (1922). “British Basidiomycetae.’ (University 
Press:Cambridge). 

Smith, A. H. and Singer, R. (1964). ‘ A Monograph of the 
genus Galerina Earle.’ (Hafner:New York). 

Vellinga, E. C. (1988). Glossary. In ‘Flora Agaricina 
Neerlandica. Vol. 1.’ (Eds Bas, C., Kuyper, Th., 
Noordeloos, M. E. and Vellinga, E. C.) pp. 51-64. 
(Balkema:Amsterdam). 

Watling, R. and Gregory, N. M. (1993). ‘British Fungus 
Flora, Agarics and Boleti. Part 7. Cortinariaceae p.p..’ 
(Royal Botanic Garden:Edinburgh). 

Wood, A.E. (2001). Studies in the Genus Galerina 
(Agaricales) in Australia. Australian Systematic 


Botany 14, 615-676. 


196 Proc. Linn. Soc. N.S.W., 126, 2005 


A Recent Expansion of its Queensland Range by Eupristina 


verticillata, Waterston (Hymenoptera, Agaonidae, Agaoninae), 


the Pollinator of Ficus microcarpa 1.f. (Moraceae). 


J.R. MCPHERSON 


School of Tropical Biology, James Cook University, Townsville Q 4811. 
Address for correspondence: 74 Sunbury Street Geebung 4034. 
E-mail: John _R_McPherson@ourbrisbane.com 


McPherson, J.R. (2005). A recent expansion of its Queensland range by Eupristina verticillata, Waterston 
(Hymenoptera, Agaonidae, Agaoninae), the pollinator of ficus microcarpa 1.f. (Moraceae). Proceedings of 
the Linnean Society of New South Wales 126, 197-201. 


In 2004, the first Ficus microcarpa seedlings were observed self-establishing in Brisbane, Queensland. 
Prior to this the only F microcarpa in the city were cultivated specimens. This self-establishment is a 
certain indicator of the presence of Eupristina verticillata, the obligate pollinator wasp of F. microcarpa. 
A Ficus species must exceed a critical population size (CPS) for its pollinator, a species of agaonid wasp, 
and other non-pollinator symbionts to colonize a new area and then maintain their new population. This 
CPS has often been estimated to be approximately 300 mature trees. The CPS for F’ microcarpa has been 
exceeded in Brisbane for some time. Brisbane has been colonized by the wasps Odontofroggatia galili (a 
gall species), since at least 1975, and E. verticillata, since at least 2004. Previously, the southern extreme 
of the range of E. verticillata was central Queensland, approximately 600 km north of Brisbane. This was 
probably achieved through a stationary, inland, trough system drawing tropical air to the southeast over a 
few days. Brisbane’s new E. verticillata population may not persist, as it must compete for short-styled F- 
microcarpa flowers with a long established O. galili population. Further, it must contend with a winter of 


greater duration and lower mean temperature than its tropical origins may allow. 


Manuscript received 24 September 2004, accepted for publication 16 February 2005. 


KEYWORDS: dispersal, Eupristina verticillata, Ficus microcarpa, Odontofroggatia galili, persistence, 


population, Queensland. 


INTRODUCTION 


On March 26, 2004, four Ficus microcarpa L.f. 
seedlings were found growing on a monument and 
road reserve infrastructure at North Quay, Brisbane, 
Queensland. The largest of these was approximately 
100 mm in height and diameter (Figure 1). Prior to 
this the only F’ microcarpa occurring in Brisbane 
were deliberately cultivated specimens. The Settler’s 
Monument, which identifies the nascent colony of 
Moreton Bay’s first graveyard, is located under the 
canopy of a F’ microcarpa (Figure 2) that is part of an 
ongoing phenology study begun in September 1996. 
The presence of these seedlings indicates the presence 
in Brisbane of the obligate pollinator of F: microcarpa, 
identified by Wiebes (1994) as being Eupristina 
verticillata Waterston, whose former southern limits 
were in central Queensland. No F microcarpa 


seedlings were noted in Brisbane’s CBD prior to 
March 2004, and none were located in a major 1996 
Ficus hemi-epiphyte audit of 21 Brisbane parks that 
involved checking 3,580 trees and palms (McPherson 
1999). Ficus microcarpa syconia investigated by the 
author at various times prior to March 2004 contained 
only the non-pollinator wasp Odontofroggatia galili 
Wiebes, although E. verticillata would have arrived 
prior to then but went undetected. 

As a genus Ficus has two distinct characteristics: 
the floral receptacle or syconium (the ‘fig’) containing 
large numbers of male and/or female flowers; and 
apart from rare exceptions (Kendelhue and Hockberg 
1997) each Ficus spp. being exclusively pollinated 
by its own unique species of agaonid wasp. Further, 
a ‘typical’ individual Ficus exhibits strict synchrony 
of syconial initiation and development, but does so 
asynchronously relative to local conspecifics. While 


RANGE OF THE POLLINATOR WASP EUPRISTINA VERTICILLATA 


a Ficus population may exhibit seasonal highs 
and lows in syconial production, at any time in a 
sufficiently large population some individuals bear 
syconia at various developmental stages. Departures 
from this ‘typical’ pattern occur, as syconia in all 
stages of development within a single tree have been 
reported for Ficus aurea Nutt. (Bronstein and Patel 
1992), Ficus benjamina L. (Corlett 1984), Ficus 
macrophylla Desf. ex Pers. (Gardner and Early 1996), 
and F. microcarpa (Corlett 1984; Bronstein 1989). 

The syconia of monoecious Ficus spp. such as 
F. microcarpa are functionally protogynous. Female 
pollinator wasps depart maturing syconia when 
male flowers are shedding pollen. Entering a young 
syconium when female flowers are receptive, they 
gall the flowers with short styles and pollinate those 
with long styles. Adult pollinator wasps live only two 
to three days, so rapid location of receptive ‘female 
phase’ syconia year round is critical to their persistence 
in an area (Ware and Compton 1994; Gardner and 
Early 1996). Various estimates have been made 
regarding the Critical Population Size (CPS) of Ficus 
spp. that allows pollinator persistence. There is a 
degree of “in principle’ consensus that approximately 
100 Ficus individuals are required for establishment 
of wasps and approximately 300 individuals needed 
for long-term wasp persistence (McKey 1989; 
Thompson, ef a/. 1991). Different Ficus spp. seem to 
exhibit longer or shorter mean periods between crops. 
This would affect CPS, as would asynchronous intra- 
tree production of syconia. More study is required to 
ascertain the CPS for different Ficus spp. and even of 
a single Ficus species growing in different climates. 

Many non-pollinator agaonid wasp species share 
syconia with pollinators. In the case of F. microcarpa, 
O. galili is a common ‘cuckoo’ species in syconia 
of F. microcarpa. It has been present in Brisbane 
since at least 1975 (Boucek 1988) and probably 
considerably longer. The long-term presence of 
O. galili in Brisbane indicates that F) microcarpa 
exceeds minimum CPS for the wasp population’s 
persistence. Survival of most non-pollinator agaonids 
depends on cohabitation of syconia with pollinators 
as unpollinated syconia usually abscise (Ramirez 
and Montero, 1988). However, abscission can be 
prevented by the presence of galls resulting from a 
few non-pollinator agaonids such as Odontofroggatia 
Ishii spp. (Boucek, 1988), allowing development of 
seedless, mature syconia. 

It is likely that most agaonid wasps disperse only 
short distances. However, reports of wasps covering 
long distances exist. Several Australian agaonid wasp 
species have become naturalized in New Zealand, 
presumably after being caught in a major air current 


198 


(Gardner and Early 1996). Stowing away in aircraft or 
vehicles also allows long dispersal distances (McKey 
1989). 

Ficus microcarpa occurs naturally in suitable 
habitats throughout the tropics of south and east Asia 
and Australia, with central Queensland as the ‘natural’ 
southern limit for F) microcarpa in Australia (Chew 
1989). Due to its hardiness, beauty and excellent 
shade it has been widely planted in the tropics and 
subtropics of the world. It isa common park and street 
tree in Brisbane, thriving in the moist-subtropical 
climate. Huge specimens dating from the 19th and 
early 20th Centuries can be found in Brisbane’s CBD 
and inner suburbs. Innumerable small to medium 
sized specimens, planted in the late 20th to early 
21st Centuries are growing in parks and along roads 
throughout the city and frost-free areas of southeast 
Queensland. 

Ficus microcarpa is an invasive urban ‘weed’ 
in the Americas, Hawaii and elsewhere, beginning 
its invasion after the introduction of its pollinator 
(McKey, 1989). Finding a vacant ecological niche, 
both plant and pollinator rapidly established and 
spread. Given its reputation as an invader of urban 
space, and since it is autochthonous to much of coastal 
Queensland, the question arises as to why it has not 
reached and survived in Brisbane prior to this recent 
occurrence. 


MATERIALS AND METHODS 


Monthly surveillance of 39 mature Ficus of 
various species, including eight F’ microcarpa, has 
been undertaken since September 1996 for Brisbane’s 


Figure 1.A Ficus microcarpaseedling atleftand a Fi- 
cus benjamina seedling at right, growing in the base 
of the Settler’s Monument, North Quay, Brisbane, 
26/03/04. The F. microcarpa has an erect habit with 
mostly obovate, flat leaves, while FE. benjamina has a 
pendulous habit with mostly ovate, curling leaves. 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. McPHERSON 


Figure 2. The Settler’s Monument, North Quay, 
Brisbane, located beneath a medium-sized Ficus 
microcarpa, 26/03/04. 


CBD. Data on the phenology of these Ficus spp. and 
any other interesting observations have been recorded. 
These data included notes on any seedlings appearing 
near mature trees. 

On March 27, 2004 “male phase’ syconia that had 
no wasp exit holes were collected from an immense 
F. microcarpa in the Brisbane City Botanic Gardens 
and incubated in glass jars covered by paper towels 
and sealed with rubber bands. After two days, wasps 
that had emerged were killed and O. galili identified 


by using the key of Boucek (1988) and E. verticillata 
using the Wiebes (1994) key. During each subsequent 
month, syconia exhibiting exit holes were collected 
from each of the eight study trees, dissected, and 
investigated for seeds and galls. Seeds and galls were 
easily differentiated as galls were either hollow or 
contained pre-emergent wasps. 

Using mean monthly temperature data supplied 
by the Regional Observations Database (Accessed 
July 16, 2004) of the Brisbane Office of the Bureau of 
Meteorology, mean monthly minimum temperatures 
for Brisbane Aerodrome during the months from May 
to September during the years in the periods 1950-99 
and 2000-03 were compared using Students t test. 


RESULTS 


The four seedling F) microcarpa were noted 
during the phenology audit for March 26, 2004. 
Subsequent to the initial discovery, two new F- 
microcarpa seedlings germinated on the Settler’s 
Monument or nearby infrastructure. As yet they have 
not been found establishing in other areas of the 
CBD. 

Syconia that lacked exit holes released both 
E. verticillata and O. galili during incubation. If 
possible, specimens of both species will be lodged 
with the Queensland Museum should later verification 
of either taxon be thought necessary. Syconia from 
which wasps had departed contained a mix of seeds 
and galls until August 2004, when only galls could be 
found. Since germination was occurring naturally in 
the CBD the viability of the seeds was not tested. 

Only July returned a significant difference 
between monthly mean minimum temperatures in 
the blocks 1950-99 and 2000-03 (Table 1). The July 
mean of 7.8°C for 2000-03 was less than the 9.5°C 
mean of for 1950-99. 


Table 1. Comparisons of mean monthly minimum temperatures at Brisbane Aerodrome, Eagle Farm, 


for the years in the periods 1950-99 and 2000-03. 


May June 
Mean minimum 13.8°C 11.0°C 
temp. 1950-99 
Mean minimum 12.6°C 10.2°C 
temp. 2000-03 
Student’s t 1.862 22) 
Significance P=0.680 P= 0.228 


Proc. Linn. Soc. N.S.W., 126, 2005 


July August September 
DIE 10.1°C IDSC 
7.8°C OBC 12.4°C 
2.249 1.299 0.346 
P=0:029 P=0.200 P=0.731 


199 


RANGE OF THE POLLINATOR WASP EUPRISTINA VERTICILLATA 


DISCUSSION 


At some time between 1999 and 2004, E. 
verticillata reached Brisbane, found and pollinated 
a number of F. microcarpa. Seed from one of these 
trees has been germinating in cracks in the mortar 
of the stone-block Settler’s Monument and in the 
surrounding footpath on North Quay. The precise 
date of the arrival is unknown but, from the size of 
the seedlings initially found, it would be closer to five 
years. 

As per the dispersal of Australian agaonids to 
New Zealand (Gardner and Early 1996) it is probable 
that FE. verticillata travelled south in a rapidly 
moving, surface air mass. If this was the case, it is 
likely that quite a few F- microcarpa were pollinated 
in subtropical coastal Queensland and as far south 
into New South Wales as the air mass penetrated. 

It would require a northwesterly wind blowing 
steadily at over 12.5 kmh' to move wasps the 
approximately 600 km from the Rockhampton area 
to Brisbane in less than two days. While unusual, 
this can happen when trough systems sit for a few 
days over inland Queensland. Tropical air is pulled 
southeast, resulting in hot northwesterly winds 
blowing over southeast Queensland and northeast 
New South Wales. Clouds of fig wasps emerging from 
natal syconia could be caught up in this air mass. 

Will E. verticillata persist in Brisbane? Possibly. 
A likely pattern is occasional population irruptions 
followed by local extinction. Wasps were still present 
in good numbers in June 2004 and successfully 
dispersing between trees, but seed could not be located 
during August, indicating a population decline. 
Hypothetically, if E. verticillata had dispersed to 
Brisbane at some time prior to 1999 it may have failed 
to establish due to strong competition for short-styled 
flowers from O. galili. Gardner and Early (1996) 
noted a decline in the numbers of pollinator wasps per 
syconia after the dispersal of non-pollinator agaonids 
to New Zealand. Also, Brisbane’s winter may be too 
long and its temperatures too low for the survival of 
E. verticillata. 

Ultimately, E. verticillata has had over a century 
in which to establish a permanent population in 
Brisbane but has yet to succeed. Any earlier failure 
to establish a permanent population can not be 
attributed to climatic factors, as Brisbane’s winters 
in the early 21*' Century were marginally cooler than 
winters in the second half of the 20" Century. It may 
be successful if the steadily expanding F! microcarpa 
population has reached a size where a residue E. 
verticillata population could survive competition and 


winter in suitable refugia, and then expand rapidly 
as the season warmed. A suitable refugium would 
have a warmer microclimate and a large number of 
F. microcarpa growing within it. This would ensure 
short pollinator dispersal flights, less stress during 
cold, adverse weather and consistently available 
‘female’ phase syconia. 

If E. verticillata does become established, then 
Ficus microcarpa would begin to establish itself as 
a hemi-epiphyte and lithophyte throughout Brisbane. 
While a boon for local frugivores, this would be a 
disaster for the local authority and other property 
managers. Ficus spp. have tremendous potential to 
damage trees and infrastructure. At least six Ficus 
spp. are already establishing in Brisbane as hemi- 
epiphytes or lithophytes (McPherson 1999), causing 
various amounts of damage and requiring ongoing 
attention to minimize their impact. None of these 
species has a literature reputation for invasiveness 
to compare with F! microcarpa (Starr 2003). Already 
hard-pressed and under-funded vegetation control 
services would be presented with yet another 
problem plant to manage should F: microcarpa, the 
acknowledged “Big Kahuna’ of invasive Ficus spp., 
begin establishing in numbers in Brisbane. 


ACKNOWLEDGEMENTS 


A/Prof Betsy Jackes, School of Tropical Biology, 
James Cook University and Mr James Harper, Brisbane 
City Council’s Community Development Officer — 
Communication, kindly commented on an early draft of the 
article. 


REFERENCES 


Boucek, Z. (1988). “Australasian Chalcidoidea 
(Hymenoptera): A Biosystematic Review of Genera 
of Fourteen Families, With a Reclassification of 
Species’ (C.A.B. International, Wallingford, UK) 

Bronstein, J.L. (1989). A mutualism at the edge of its 
range. Experientia 45, 622—637. 

Bronstein, J.L. and Patel, A. (1992). Causes and 
consequences of within tree phenological patterns in 
the Florida strangling fig, Ficus aurea (Moraceae), 
American Journal of Botany 79, 1, 41-48. 

Chew, W.L. (1989). Moraceae. In ‘Flora of Australia’, 
3, pp. 15-68. (Australian Government Publishing 
Service, Canberra, ACT). 

Corlett, R.T. (1984). The phenology of Ficus benjamina 
and Ficus microcarpa in Singapore. Journal of the 
Singapore National Academy of Science 13, 30-31. 


Proc. Linn. Soc. N.S.W., 126, 2005 


J.R. McPHERSON 


Gardner, R.O. and Early, J.W. (1996). The naturalization 
of banyan figs (Ficus spp., Moraceae) and their 
pollinating wasps (Hymenoptera: Agaonidae) in 
New Zealand. New Zealand Journal of Botany 34, 
103-110. 

Kendelhue, C. and Hockberg, M.E. (1997). Active 
pollination of Ficus sur by two fig wasp species in 
west Africa, Bio, Vol. 29, No. 1, pp. 69-75. 

McKey, D. (1989). Population biology of figs: applications 
for conservation. Experientia 45, 661-672. 

McPherson, J.R., 1999. “Studies in urban ecology: 
strangler figs in the urban parklands of Brisbane, 
Queensland, Australia’, Australian Geographical 
Studies, Vol. 37, No. 3, pp. 214-229. 

Ramirez, W.B. and Montero, J.S. (1988). Ficus 
microcarpa L., F: benjamina L. and other species 
introduced in the New World, their pollinators 
(Agaonidae) and other fig wasps. Revista Biologia 
Tropica 36, 2B, 441-446. 

Regional Observations Database, “Daily Data for 
Brisbane’, (No publication date), Commonwealth 
Bureau of Meteorology (Accessed July 16th 2004) 

Starr, F., Starr, K. and Loope, L., 2003. ‘Ficus microcarpa, 
Chinese banyan, Moraceae.’ United States Geological 

; Survey, Biological Resources Division. Hawaii 

Thompson, J.D., Herre, E.A., Hamrick, J.L. and Stone, 
J.L. (1991). Genetic mosaics in strangler fig trees: 
Implications for tropical conservation. Science 254, 
1214-1216. 

Ware, A.B. and Compton, S.G. (1994). Responses of fig 
wasps to host plant volatile cues. Journal of Chemical 
Ecology 20, 3, 785-802. 

Wiebes, J.T. (1994), ‘The Indo-Australian Agaoninae 
(Pollinators of Figs).’ (Koninklijke Nederlandse 
Akademie van Wetenschappen, Amsterdam). 


Proc. Linn. Soc. N.S.W., 126, 2005 


201 


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A New Phyllolepid Placoderm Occurrence (Devonian Fish) from 


the Dulcie Sandstone, Georgina Basin, Central Australia 


GAVIN C. YOUNG 


Department of Earth and Marine Sciences, Australian National University, Canberra ACT 0200 
gyoung@ems.anu.edu.au 


Young, G.C. (2005). A new phyllolepid placoderm occurrence (Devonian fish) from the Dulcie 
Sandstone, Georgina Basin, central Australia. Proceedings of the Linnean Society of New South 
Wales 126, 203-213. 


A new phyllolepid placoderm occurrence from a low level in the Dulcie Sandstone, Georgina 
Basin, Northern Territory, lies about 200 metres stratigraphically above an older fish assemblage 
containing Wuttagoonaspis. A new species Austrophyllolepis dulciensis 1s characterised by an 
unusually broad anterior ventrolateral plate. The stratigraphic range of other species in this genus 
from southeastern Australia and Antarctica suggest a Givetian-Frasnian age. Early members of 
the order Phyllolepida are endemic to east Gondwana, and other phyllolepids of similar age 
occur in Turkey and Venezuela. In the Northern Hemisphere (Europe, Russia, Greenland, North 
America) phyllolepids are restricted to the latest Devonian stage (Famennian). This disjunct 
space-time distribution for the group supports a Gondwanan origin for the Phyllolepida, and 
later access to northern landmasses resulting from closure of the ocean between Gondwana and 
Laurussia at or near the Frasnian-Famennian boundary. 


Manuscript received 25 June 2004; accepted 15 September 2004. 


KEYWORDS: Devonian, Georgina Basin, Phyllolepida, Placoderm fishes. 


INTRODUCTION 


Devonian fossil fish remains from central Australia 
were first documented by Hills (1959), who identified 
the placoderm genera Bothriolepis and Phyllolepis 
and concluded a Late Devonian age for the Dulcie 
Sandstone in the Dulcie Range, N.T. (Georgina Basin; 
Fig. 1). An older fish assemblage from the basal part 
of the Dulcie Sandstone at the northwestern end of 
the Dulcie Syncline was discovered during geological 
mapping by the Bureau of Mineral Resources in 1961. 
Further material was collected by the author from 
these and many new localities during two field trips 
to the Georgina Basin (1974, 1977). The entire fauna 
of the lower assemblage, including a new species of 
the genus Wuttagoonaspis Ritchie (1973), has been 
described by Young and Goujet (2003). Ten new 
localities in the lower part of the Dulcie Sandstone 
were documented, of which six (localities GY74/8- 
13) were collected in 1974 along the southern flank 
of the Dulcie Range on the southeastern edge of the 


Barrow Creek 1:250 000 sheet (Fig. 1B). There were 
no taxa in common with the earlier descriptions of 
Hills (1959) and Young (1985, 1988), which dealt 
with younger fish assemblages from the upper part 
of the Dulcie Sandstone (localities GY1-7 of Young 
1988: fig. 4). A diagnostic group in these younger 
assemblages are the phyllolepid placoderms. 

The specimens described in this paper were 
collected from one of the localities (GY74/8) 
documented by Young and Goujet (2003). Initially 
they were put aside as unidentified smooth mudclast 
impressions in sandstone. However, the shape of bones 
now clearly demonstrates that the sample represents 
impressions of bones belonging to a phyllolepid 
placoderm. Some 19 genera representing 13 taxa 
of family or higher rank have been documented by 
Young and Goujet (2003) in the Wuttagoonaspis 
Assemblage, which is known from localities over 
about one million square kilometres of the Australian 
continent. However, phyllolepid placoderm remains 
have never been confirmed in that assemblage. This 


DEVONIAN PLACODERM FROM GEORGINA BASIN 


[Early Palaeozoic 


hae syncline 


Figure 1. A, location of the Dulcie Range, on the south- 
western flank of the Georgina Basin (GB) in northern 
Australia. B, Fossil fish localities in the northwestern 
part of the Dulcie Syncline, with geology generalised 
from the Barrow Creek and Alcoota 1:250 000 sheets 
(modified from Young and Goujet 2003: fig. 1). The 
samples described herein come from locality GY74/8. 


204 


new phyllolepid occurrence is_ significant 
in coming from an intermediate level in the 
Dulcie Sandstone, all previous examples of 
phyllolepids (Hills 1959; Young 1988) coming 
from near the top of the sequence. 

Phyllolepids, like Wuttagoonaspis, have 
dermal bones with ridged ornament. Before 
Wuttagoonaspis was described by Ritchie 
(1973), all fish remains with ridged ornament 
from the Australian Devonian were referred 
to Phyllolepis without question (e.g. Rade 
1964). In Europe Phyllolepis is only known 
from the youngest stage of the Late Devonian 
(Famennian), and on this basis all such 
occurrences inAustralia were originally assigned 
to the Famennian (e.g. Hills 1929, 1931). 
But Hills (1958) also noted that the temporal 
significance of different placoderm genera in 
the Upper Devonian of Europe did not apply 
in Australia, and Young (1974) demonstrated 
that some phyllolepid occurrences were older 
(Frasnian) than in Europe. Some authors (e.g. 
Ritchie 1973) considered the phyllolepids and 
Wuttagoonaspis to be only distantly related, 
and recently Dupret (2004) has regarded 
the ridged ornament as a non-homologous 
character. The alternative hypothesis (e.g. 
Miles 1971; Young 1980; Long 1984; Young 
and Goujet 2003) is that ridged ornament is a 
shared derived feature, which with some others 
indicates that Wuttagoonaspis and phyllolepids 
are sister groups. Apart from ornament, the 
shape of bones is quite different in the two 
taxa, so impressions with ridged ornament can 
be readily assigned to one or the other group if 
bone margins are complete. Thus, examination 
of the ridged fragment called Phyllolepis by 
Gilbert-Tomlinson (1968) showed clearly that it 
belongs to Wuttagoonaspis (Young and Goujet 
2003, fig. 3E). Similarly, impressions of the 
inner unornamented surface described below 
can unequivocally be assigned to a phyllolepid 
on the basis of bone shape. 


Locality 

Fossil fish localities GY74/8-13 along the 
southern flank of the Dulcie Range (Fig. 1B) 
supposedly came from the basal 10 m of the 
Dulcie Sandstone (Haines et al. 1991: 32), but 
poor outcrop, and some material collected from 
scree, make the actual interval of occurrence 
difficult to establish (Young and Goujet 2003). 
The boundary with the underlying Tomahawk 


Proc. Linn. Soc. N.S.W., 126, 2005 


G.C. YOUNG 


beds (Cambro-Ordovician) is variously interpreted, as 
a faulted contact on the second edition of the Barrow 
Creek 1:250 000 geological map (Haines et al. 1991) 
but as a disconformity on the adjacent Huckitta map 
sheet (Freeman 1986). 

The most easterly of these fish occurrences 
(GY74/8, Figure 1B) is recorded in field notes 
as being immediately above the contact with the 
underlying Tomahawk beds and was the first at which 
fish remains were discovered by the author in the 
lower Dulcie (on 7 July 1974). This is the locality 
yielding the specimen described below. Locality 
74/8 was about 800 m west of the field campsite at 
Lurapulla Waterhole. 


Stratigraphic level 
The samples were labelled ‘top of Dulcie; 
collected 7/7/74’. According to field diary records 
only locality 74/8 was visited on that day, when the 
adjacent section through the Dulcie Sandstone was 
also examined. Stereo air photos of locality 74/8 
(Barrow Creek 17-10-70, run 9, photos 0025, 26) 
- show the highest elevation in the vicinity of two ridges 
about | and 1.8 km to the NW, where beds have a very 
shallow dip (5° on the Barrow Creek geological sheet, 
first edition). Lower beds are more inclined, with a 
dip of 16° marked on the second edition geological 
map just above Lurapulla Waterhole. Estimates 
based on air photo interpretation suggest that the 
samples described below came from an interval 
about 200+ m above the fish horizon at the base of 
the sequence. At locality 74/8 the basal level yielded 
only indeterminate phlyctaenioid arthrodire remains 
(impressions with tuberculate ornament). However, 
another locality along strike about 3 km to the NW 
(74/11) produced some 50 samples belonging to seven 
taxa, including Wuttagoonaspis (Young and Goujet 
2003: table 1). At none of the 26 localities covered 
in that study were phyllolepid remains identified. It 
was concluded that younger bothriolepid-phyllolepid 
placoderm assemblages did not occur in the lower 
part of the Dulcie Sandstone, nor in the Cravens Peak 
Beds (contra Gilbert-Tomlinson 1968, and Draper 
1976). However, they are recorded from the upper 
part of the Dulcie Sandstone at the southeastern end 
of the Dulcie Range (about 490 and 600 m above 
base; Young 1988), where the type section records a 
thickness of 621 m. These horizons occur in sandstone 
mesas in the core of the syncline, representing the 
upper subdivision of the formation and are separated 
by valleys of recessive strata from more prominent 
outcrops of the lower Dulcie Sandstone (e.g. Freeman 
1986, pl. 20). 

Freeman (1986) identified the upper and lower 


Proc. Linn. Soc. N.S.W., 126, 2005 


facies in the vicinity of the type section but dismissed 
the suggestion (Gilbert-Tomlinson 1968) of a possible 
paraconformity within the lower unit. He noted a 
decrease in thickness to an estimated 250 m in the 
northwestern exposures on the Huckitta sheet. No 
sections were measured by Haines et al. (1991) on 
the Barrow Creek sheet, but the Dulcie Sandstone is 
much thinner (30-40 m) on the Elkedra sheet to the 
north-west (Stidolph et al. 1988). 

A question arises as to whether the thinning 
of the Dulcie is general throughout the sequence 
or whether the lower or upper part is missing from 
the thinner stratigraphic sections. As first noted by 
Gilbert-Tomlinson (1968), the lower and upper fish 
assemblages in the Dulcie generally do not occur 
together. The occurrence reported here is the only 
one known so far that represents two distinct fish 
faunas within the same section. Previously (Young 
1985: 251) a stratigraphic thickness of about 430 m 
was estimated between the lower Wuttagoonaspis and 
upper Bothriolepis — Phyllolepis fish assemblages, but 
the new fossil sample described below comes from 
some 200 m above the lower fish horizon. Decrease 
in thickness to the north-west from the type section 
would place this new phyllolepid occurrence near 
the top of the formation as exposed on the Barrow 
Creek sheet. If it correlates with those from the SE 
it could be assumed that the middle recessive part 
of the Dulcie Sandstone in the type section has been 
lost, retaining two distinct fish assemblages with no 
taxa in common within a section some 200-300 m 
thick. It is considered more likely that it represents 
a lower phyllolepid assemblage, given that a 
rather diverse fauna of unknown biostratigraphic 
relationship is indicated from fragmentary remains 
already described (Young 1988). Comparison can be 
made with the Pertnjara Group in the Amadeus Basin, 
where thickness extrapolations suggest a separation 
between the assumed level for the Wuttagoonaspis 
fauna in the base of the Deering Siltstone Member 
and the Bothriolepis assemblage within the Harajica 
Sandstone Member to be about 280 m at Stokes Pass, 
and possibly as low as 100 m at Dare Plain (Young 
1985: 251, 252). A new species of the phyllolepid 
Placolepis has recently been described from the 
Harajica fish assemblage (Young in press a). 


ABBREVIATIONS 


Measurements of total length (L), breadth (B), 
length of the anterior division of the AVL (L,,), 
and level of the lateral corner (Ic) of the PVL 
(as a percentage of total length) are summarised 


205 


DEVONIAN PLACODERM FROM GEORGINA BASIN 


in Figure 3 (below). Bone proportions are given 
as a ratio of breadth to length expressed as a 
percentage (abbreviated as ‘B/L index’). Standard 
abbreviations for placoderm dermal bones and other 
structures are used in the text and figures as follows: 
ADL, anterior dorsolateral plate; 

AL, anterior lateral plate; 

AMV, anterior median ventral plate; 

AVL, anterior ventrolateral plate; 

cf.IL, contact face overlapping interolateral plate; 
cf.SP, contact face overlapping spinal plate; 

IL, interolateral plate; 

Ic, lateral corner; 

m.AMV, margin abutting anterior median ventral 
plate; 

m.PMV, margin abutting posterior median ventral 
plate; 

0a.AVL, area overlapped by AVL plate; 

pect, pectoral embayment (margin) of AVL plate; 
PMV, posterior median ventral plate; 

PNu, paranuchal plate; 

PVL, posterior ventrolateral plate; 

ppec, prepectoral corner; 

ptpec, postpectoral corner; 

SP, spinal plate. 


SYSTEMATIC PALAEONTOLOGY 


Class PLACODERMI McCoy, 1848 
Order PHYLLOLEPIDA Stensi6, 1934 


Diagnosis 

Placoderms in which the nuchal plate is much 
enlarged, as broad or broader than long, and surrounded 
by five smaller paired bones including paranuchal, 
marginal, postorbital and preorbital plates. Paranuchal 
plate with well-developed postnuchal process, and 
rostral, pineal, and central plates absent from skull 
roof. Trunk armour relatively broad; median dorsal 
plate lacks an inner keel; anterior dorsolateral 
plate with narrow elongate exposed area; anterior 
ventral and posterior lateral plates absent; posterior 
dorsolateral and anterior and posterior median ventral 
plates reduced or absent. Anterior ventrolateral 
plates short and broad, and posterior ventrolaterals 
triangular, with ossification centres near anteromesial 
corners; both ventrolateral plates relatively flat, 
lacking a lateral lamina, and meeting in part or all 
of the midline in non-overlapping sutures. Dermal 
ornament mainly of smooth concentric ridges, with 
some tubercles and tubercle rows. 


Remarks 
The diagnosis provided by Ritchie (1984: 344) 


206 


was slightly reworded from that of Denison (1978: 
41), and included reference to the absence of rostral 
and pineal plates from the skull, used by Denison 
to separate his two suborders ‘Antarctaspina’ and 
‘Phyllolepina’. Antarctaspis is now regarded as an 
actinolepid related to Toombalepis and Yujiangolepis, 
which also have converging sensory grooves on 
the nuchal plate (Young and Goujet 2003: fig. 16), 
a primitive feature and not a criterion for indicating 
close relationship to phyllolepids. Ritchie (1984: 
346) gave an additional diagnosis for the family 
Phyllolepidae that combined features of Denison’s 
subordinal and family diagnoses, but there are no 
good criteria for grouping genera within the order at 
this stage. 

The above diagnosis excludes characters that 
are evidently primitive by outgroup comparison 
(e.g. converging sensory canal grooves on the skull, 
median dorsal plate short and broad and lacking inner 
keel, sliding dermal neck-joint). New characters 
include features of the trunk armour bones typical 
of the group. The much reduced external part of 
the anterior dorsolateral (ADL) plate is a condition 
approached in Bryantolepis, Kujdanowiaspis (Denison 
1958: fig. 108) and Wuttagoonaspis. The anterior 
ventrolateral (AVL) is relatively broad compared 
to that of other primitive arthrodires, in which the 
posterior ventrolateral (PVL) has complex midline 
overlaps, and a prominent lateral lamina (Denison 
1958: figs. 112, 114). The ossification centre of the 
PVL is normally placed laterally near the posterior 
end of the lateral lamina (e.g. Actinolepis; Mark- 
Kurik 1973: fig. 13), but its anteromesial position 
in phyllolepids is well shown by the concentric 
ornament of Austrophyllolepis (Long 1984: fig. 11D). 
New phyllolepids from southeastern Australia show 
that the posterior dorsolateral (PDL) plate is retained 
in some members of the order (Young, in press b), 
although it is lost in Phyllolepis, Austrophyllolepis, 
and probably Placolepis (Long 1984; Ritchie 1984). 

Genera included in the family and order are 
Phyllolepis, Austrophyllolepis, and Placolepis, plus 
two new phyllolepid taxa from southeastern Australia 
(Young in press b). The genus Pentagonolepis 
Lohest, 1888, from the Famennian of Belgium, was 
synonymised with Phyllolepis by Leriche (1931), 
Stensié (1939) and Denison (1978). However, the 
ridged ornament on the nuchal plate from the skull roof, 
on which the species Phyllolepis (Pentagonolepis) 
konincki is based (Stensid 1939, fig. 6B), suggests 
a rounded anterior margin (incomplete), somewhat 
similar to the corresponding bone of Placolepis 
described by Ritchie (1984). The Belgian specimen 
differs from Placolepis in that the middle pitline is — 


Proc. Linn. Soc. N.S.W., 126, 2005 


G.C. YOUNG 


directed towards the lateral corner of the plate, as in 
Phyllolepis, rather than well behind that corner. It 
seems likely that the paranuchal plate from the same 
skull had extensive contact with the postorbital plate. 
Restudy of this material, and any new specimens from 
Belgium, is needed to confirm the shape of the nuchal, 
but ‘Pentagonolepis ’konincki Loheste, 1888 could be 
a valid taxon, morphologically intermediate between 
Placolepis and Phyllolepis in the configuration 
of skull bones. It is noted that the first Devonian 
tetrapods from continental Europe have recently 
been described from these localities, in the Evieux 
Formation of Belgium (Clément et al. 2004). 


Austrophyllolepis Long, 1984 


Diagnosis 

Phyllolepids in which the sensory groove passes 
off the paranuchal plate in the anterior third of plate 
length, and the external surface of the marginal plate 
is similar in breadth and length, with the postmarginal 
sensory canal junction in about the middle of plate 
‘length. A small suborbital plate is articulated to an 
ossified process below the postorbital plate, and the 
submarginal plate may be ossified. The trunk armour 
has a posterior median ventral plate that forms a 
distinct notch in the mesial margins of anterior and 
posterior ventrolateral plates. The ridged ornament 
includes extensive areas of tuberculation and some 
ridge duplication. 


Austrophyllolepis dulciensis sp. nov. 


Name 

Shortened from “Dulcie’, after the type formation 
and locality (Dulcie Sandstone and Range, Georgina 
Basin, Northern Territory). 


Diagnosis 

A species of Austrophyllolepis in which the 
anterior ventrolateral plate is as broad as long 
(breadth/length index about 100), with anterior and 
posterior divisions about 20% and 30% of total 
length and a slightly concave spinal margin; posterior 
median ventral plate elongate and free part of spinal 
plate relatively short. 


Remarks 

The generic diagnosis from Young and Long 
(in prep.) is updated from Long (1984). Most of the 
characters are not observable in the material described 
below, but the evidence of a posterior median ventral 
(PMV) plate in the trunk armour is the criterion for 
referring the species to Austrophyllolepis (see further 


Proc. Linn. Soc. N.S.W., 126, 2005 


comments below). Characters in the specific diagnosis 
separate the new species from previous species of 
Austrophyllolepis. The unusually broad anterior 
ventrolateral (AVL) plate distinguishes A. dulciensis 
sp. nov. from all previously described phyllolepids in 
which this plate is known. 


Material 

ANU V3064 (holotype; associated AVL, PVL 
and SP plates); ANU V3065 (incomplete PVL plate), 
both preserved as impressions in hard sandstone. 


Locality and Horizon 

Locality GY74/8, southern flank of Dulcie 
Range (Fig. 1B), from a horizon about 200 m above 
the exposed base of the Dulcie Sandstone (see Young 
and Goujet 2003: 5 for locality details). 


Description 

The impressions of closely associated phyllolepid 
trunk armour plates comprise an almost complete 
right AVL plate, overlain by a SP plate and a right 
PVL plate (Fig. 2A). Another AVL is separated by 45 
mm from the first (Fig. 2B), and an incomplete left 
PVL impression is preserved on the second smaller 
sandstone sample (ANU V3065). The two AVL plates 
(termed AVL#1, AVL#2) were first assumed to be 
internal impressions of left and right plates from one 
individual, because both appeared to lack ornament. 
Closer examination showed that both are right plates. 
The second specimen (AVL#2; Fig. 2B) is slightly 
convex rostrocaudally, and must have been an external 
surface impression, even though it is smooth. A new 
phyllolepid genus with smooth dermal bones has 
recently been documented from southeastern Australia 
(Young, in press b). However, in ANU V3065 absence 
of ornament may be a post-mortem effect due to 
‘sand-blasting’ in a river current removing the ridged 
ornament from most of the external surface. Short 
sections of relatively coarse ridges are retained on the 
downturned posteromesial and spinal margins of the 
bone (ri, Fig. 2B), presumably protected from abrasion 
beneath the sediment-water interface. The relatively 
flat ventral plates of the phyllolepid trunk armour 
would have made them hydrodynamically stable 
with either external or internal surfaces uppermost, in 
contrast to some other common placoderm remains, 
for example the strongly angled trunk armour bones 
of the widespread antiarch Bothriolepis. 

The anterolateral and prepectoral corners of 
AVL#2 are incomplete, and the posterolateral part 
is missing off the edge of the sample. This and the 
first AVL clearly came from two individuals of about 
the same size, and shape differences are assumed 


207 


DEVONIAN PLACODERM FROM GEORGINA BASIN 


pect 


Figure 2. ANU V 3064 showing associated AVL, PVL and 
SP plates of Austrophyllolepis dulciensis sp. nov. (A), 
the second AVL plate (B). Both are latex casts of sand- 
ammonium 


stone impressions whitened with 


to be intraspecific variation, given the very similar 
proportions of the two restored plates (see below). 
AVL#1 (Fig. 2A) exposes the visceral surface, which 
is gently concave rostrocaudally, with the anterior 
margin curved up, forming a slight rim mesially. 
Around the anterolateral corner a second ridge 
inside the margin defines a narrow contact face for 
the IL and SP plates (cf.IL, cf.SP, Fig. 2A, 3A). A 
diagonal groove crosses the visceral surface of the 
AVL in Placolepis (Ritchie 1984: fig. 11A-B) and is 
a distinctive feature of that genus. It is not developed 
in the Victorian material of Austrophyllolepis (Long 
1984, 1989) and is absent in this new specimen. The 


208 


spinal margin, which in all other 
phyllolepid taxa is convex, in this 
AVL is gently concave anteriorly 
and straight posteriorly (Fig. 3A). 
The mesial margin, normally 
straight in phyllolepids, is slightly 
convex anteriorly and concave 
posteriorly, again showing the 
raised rim typical of phyllolepid 
AVL plates where they form a non- 
overlapping suture in the midline. 
In that case the margin must be 
straight, but AVL#1 indicates a 
gap in the midline for a small PMV 
plate, as in Austrophyllolepis from 
Victoria (Long 1984: fig. 11D). 
In AVL#2 the preserved spinal 
margin and the posterior part of 
the mesial margin are also gently 
concave (Fig. 2B), as in AVL#1. 
Reconstruction (Fig. 4), based on 
AVL#2, suggests that the Dulcie 
Range species must have had a 
larger PMV than in the Victorian 
species. Of these, A. ritchiei had 
the longer PMV (Long 1984: 
figs. 7B, 13B) but with a concave 
external margin (convex in the 
new specimens). There is a distinct 
short posteromesial margin 
on AVL#1 (m.PMV, Fig. 3A), 
occasionally seen in other species 
; (e.g. interpreted as a broken edge 
r in Phyllolepis concentrica by 
Stensi6d 1939: fig. 1). 

AVL#2 differs in lacking a 
posteromesial margin, but has an 
anteromesial margin defined by 
distinct corners (m.AMYV, Fig. 4). 
Ritchie (1984: 342-3) assumed 
the AMV to be absent in European 
Phyllolepis and in Placolepis, as 
did Long (1984) for Austrophyllolepis. One specimen 
assigned to A. youngi does suggest an anterior median 
element (Long 1984: fig. 18C), as in Phyllolepis 
woodwardi from Europe (Stensid 1939: fig. 2). 
An AMV plate is also present in a new phyllolepid 
taxon from Merriganowry, N.S.W. (A. Ritchie, pers. 
comm.). Since both AMV and PMV plates were 
primitively present they would be expected to be 
variably developed in basal phyllolepid taxa (for 
example the two PMV plates in Victorian material; 
Long 1984: fig. 6). 

Both AVLs as restored are as broad as long (B/L > 


and 


chloride. 


Proc. Linn. Soc. N.S.W., 126, 2005 


G.C. YOUNG 


Figure 3. Austrophyllolepis dulciensis sp. nov. A, AVL plate 


of Fig. 2A (AVL#1) slightly restored. B, PVL plate 


of Fig. 2A, restoration of external surface. ANU V3064. 


index 100), witha similar length of the anterior division 
(19-20%). In contrast, in both Austrophyllolepis and 
Placolepis budawangensis (Long 1984; Ritchie 1984: 
fig. 11), and all species of Phyllolepis from Europe 
(Stensid 1939), the AVL is consistently longer than 
broad. Since the precise measurements for the AVL 
as summarised in table form by Stensié (1939: 7) 
were not clearly defined, measurements used here are 
indicated in Fig. 3. 

The main difference in the two AVLs described 
here is the orientation of the posterior margin, giving 
a greater posterior angle (~78°) in AVL#1 than in 


Proc. Linn. Soc. N.S.W., 126, 2005 


AVL#2 (~60°), the latter approximating to 
the anteromesial angle of the PVL (~65°). 
Distortion in the specimens is discounted 
because the anteromesial angle of the AVL 
is much the same in both (75-80°). 

The complete PVL impression shows 
the visceral surface of a left plate (Fig. 2A), 
with a raised inner rim along its straight 
mesial margin (L 33 mm) which abutted 
against the right PVL. A thickened zone 
inside the anterior margin probably reflects 
the position of the overlap area for the AVL 
on the external surface (0a.AVL, Fig. 3B). 
The B/L index (73) places this PVL at the 
broader end of the variation for Placolepis 
budawangensis, which is broader than in 
Austrophyllolepis edwini, A. youngi and 
three European species (Phyllolepis orvini, 
Ph. nielseni, Ph. tolli), but more narrow than 
in A. ritchiei (Young 1988: table 1), with 
Austrophyllolepis differing from the other 
taxa in the more posterior level of the lateral 
corner (Long 1984: fig. 7; 1989: fig. 4). The 
shape and proportions of the PVL described 
here are also notably different from the two 
PVLs from the Amadeus Basin (Young 
1988: table 1). The second incomplete PVL 
(ANU V3065) shows the mesial margin of a 
right plate impression, also 33 mm long, and 
thus perhaps from the same individual. 

The complete PVL of Fig. 2A does not 
fit properly against the underlying AVL, but 
makes a good fit with AVL#2, so these were 
used for the reconstruction (Fig. 4). The 
associated SP plate impression is missing 
both ends, but restored length is about 
33 mm. The distal 4 mm of the preserved 
part has a closed posterior margin, with 
the bases of about 4 denticles preserved on 
what is assumed to be the ventral margin. 
This margin shows a slight angle in the 
middle of the preserved part, assumed to fit 
into the concave margin on the AVL. The opposite 
lamina (presumed to be dorsal) is more rounded. This 
SP plate cannot be restored with the free part of the 
spine as long as in Placolepis, or in the Victorian 
Austrophyllolepis (33% and 25% of total length 
respectively; Ritchie 1984: 342; Long 1984: 272). 
However, in Phyllolepis orvini the free part of the SP 
plate is much shorter (Stensi6 1936: fig. 20). 


209 


DEVONIAN PLACODERM FROM GEORGINA BASIN 


Figure 4. Austrophyllolepis dulciensis sp. nov. Attempted restora- 
tion of ventral trunk armour; outline shape, and plates of the right 
side based on the AVL plate of Fig. 2B (AVL#2), and the PVL plate of 
Fig. 2A; left side indicates the shape of AVL#1 (Fig. 2A); gap left 
for the PMV plate suggests a variable shape in this species (PMV). 


DISCUSSION 


These few phyllolepid remains are apparently 
‘primitive’ compared to Phyllolepis in Europe by 
showing evidence of a PMV plate of appreciable 
size, a character by which they can be referred to 
Austrophyllolepis Long, 1984. However, they differ 
from all three species of the genus so far described 
(Long 1984, 1989). It is unclear how reliable the 
criterion of PMV presence will prove as a generic 
character, since both AMV and PMV were presumably 
present in phyllolepid ancestors and their reduction 
and loss may have occurred independently in different 
lineages. For the present this criterion is retained to 
allocate this species to the genus Austrophyllolepis. 
Although the ridged ornament is not preserved, the 
distinctive triangular PVL plate clearly places these 
impressions within the Phyllolepida as defined 
above. In other arthrodires, and in Wuttagoonaspis 
(e.g. Young and Goujet 2003: fig. 8G), the PVL has 
a distinct lateral lamina, the primitive condition. The 
loss of this lamina, giving an essentially flat bone, may 
account for the fact that impressions of the phyllolepid 
PVL are more commonly found than other bones of 
the trunk armour, because its hydrodynamic qualities 
would have resembled those of a flat mud clast. 


210 


The fact that this 
phyllolepid evidently came 
from a horizon some 200+ 
m_ stratigraphically above 
a completely different fish 
assemblage, which lacks 
phyllolepids but contains 
the possible primitive sister 


group Wuttagoonaspis, 
might suggest that this 
is a_ relatively early 


phyllolepid occurrence. Ina 
comparison of stratigraphic 
thicknesses between fish 
horizons across __ central 
Australia, Young (1985: 
fig. 10) concluded that the 
previous Dulcie Range 
phyllolepid occurrences 
could be somewhat younger 
than the phyllolepid — 
bothriolepid occurrences 
in the Harajica Sandstone 
of the Pertnjara Group in 
the Amadeus Basin. This 
new description of probable 
Austrophyllolepis sp. lower 
in the Dulcie sequence 
suggests that the diversity of phyllolepids in central 
Australia is greater than previously thought, as is now 
demonstrated in southeastern Australia (Young, in 
press b) and Antarctica (Young and Long submitted). 
By comparison with species of both Austrophyllolepis 
and Placolepis in southeastern Australia, the age of 
the lower phyllolepid occurrences in central Australia 
can be assumed to lie within the Givetian-Frasnian 
interval. Recent work on the age of the Mount Howitt 
fish assemblage within the Middle-Late Devonian 
volcanics of eastern Victoria, the type locality for 
Austrophyllolepis, supports a late Middle Devonian 
age on both palaeontological and isotopic evidence 
(Long 1999; Compston 2004). 

The diversity of phyllolepid taxa in East 
Gondwana, and their earlier occurrence than in the 
Devonian of the Northern Hemisphere, supports the 
hypothesis of a Gondwanan origin for the group, 
followed by dispersal into Laurussia in the Famennian 
resulting from palaeogeographic change at or near the 
Frasnian-Famennian boundary (Young 1989, 1990, 
1993b, 2003). A summary of biostratigraphic ranges 
(Fig. 5) shows diverse phyllolepids in the Givetian- 
Frasnian of East Gondwana, with related forms in the 
largely endemic Wuttagoonaspis assemblage (Young 


Proc. Linn. Soc. N.S.W., 126, 2005 


G.C. YOUNG 


rachytera 
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LATE DEVONIAN 


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BIOSTRATIGRAPHIC RANGE OF PHYLLOLEPID TAXA 
ZONATIONS 
[E. GONDWANA] GONDWANA LAURUSSIA 


rhenana ae 
jamieae a Se 


Ph. undulata 
Ph. concentrica 
Ph. woodwardi 

Ph. orvini 
Ph. nielseni 
Ph. delicatula 


Figure 5. Summary of known stratigraphic ranges (?Pragian-Famennian) for wutta- 
goonaspids-phyllolepids in East Gondwana (Australia, Antarctica), and the genus Phyl- 
lolepis in Laurussia (ANT= Antarctica; AMAD= Amadeus Basin). Approximate interval 
for Austrophyllolepis dulciensis sp. nov. from the Dulcie Sandstone indicated, based on 
the range of other species of the genus. Australian stratigraphic range data updated 
from Young (1993a: fig. 9.3) and Young (1999: fig. 5). Conodont zonation after Talent et 
al. (2000). Approximate alignment of macrovertebrate (MAV), miospore (GH, GF, VCo, 
LN, LV), and conodont zones modified from Young (1996) and Young and Turner (2000). 


and Goujet 2003) probably extending down into the 
Early Devonian, some 22 conodont zones before 
species of the genus Phyllolepis are documented in the 
Northern Hemisphere (in the Famennian rhomboidea 
conodont zone). This disjunction in time and space is 
an outstanding feature of global distribution patterns 
amongst Devonian fishes. 


Proc. Linn. Soc. N.S.W., 126, 2005 


ACKNOWLEDGMENTS 


Peter Davis provided assistance with the 1974 field 
work, and Lindsay and Joan Johannsen (Baikal Station), 
and John and Audrey Turner (Jinka Station), are thanked 
for hospitality. Information on field localities and Georgina 
Basin geology was provided by J.H. Shergold and E.C. 
Druce. P. Kruse provided copies of recent geological maps. 
H.M. Doyle prepared and curated the sample. A. Ritchie 
(Australian Museum, Sydney) and J.A. Long (Museum of 
Victoria, Melbourne) provided unpublished information 


211 


DEVONIAN PLACODERM FROM GEORGINA BASIN 


and casts of phyllolepid material for comparison. E. 
Resiak and J. Laurie (Geoscience Australia) provided 
access to the Georgina Basin palacontological collection, 
and Margaret Drury helped with early air photos in the 
Geoscience Australia library. Ben Young assisted with 
sorting of latex casts and matching to original specimens, 
and Val Elder contributed to collection management. This 
research was a contribution to IGCP Projects 328, 406, 410 
and 491. Financial support was provided by ANU Faculties 
Research Fund Grant F00108, and Professor P. De Deckker 


is thanked for provision of facilities in the Department of 
Earth & Marine Sciences, ANU. Comparative study of 


phyllolepid placoderms from East Gondwana was begun in 
Berlin during the tenure of a Humboldt Award (2000-2001), 
and the support of the Alexander von Humboldt Foundation, 
and provision of facilities in the Museum ftir Naturkunde, 
Berlin (Prof. H.-P. Schultze) are gratefully acknowledged. 


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Mark-Kurik, E. (1973). Actinolepis (Arthrodira) from the 
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Western Australian Museum, Supplement 65, 85 pp., 
42 figures. 

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213 


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Cowralepis, a new genus of phyllolepid fish (Pisces, Placodermi) 
from the Late Middle Devonian of New South Wales, Australia 


ALEXANDER RITCHIE 


Research Fellow, Palaeontology, Australian Museum, 6 College St., Sydney, New South Wales 2010 
alexr@austmus.gov.au 


Ritchie, A. (2005). Cowralepis, a new genus of phyllolepid fish (Pisces, Placodermi) from the Late 
Middle Devonian of New South Wales, Australia. Proceedings of the Linnean Society of New South 
Wales 126, 215-259. 


Cowralepis mclachlani, a new genus and species of phyllolepid placoderm (Pisces, Placodermi), is 
described from numerous articulated specimens discovered near Cowra, New South Wales, Australia. 
Cowralepis, represented by a growth series, illustrates ontogenetic changes from juvenile to adult and 
throws new light on the dermal, endocranial, visceral and axial skeleton of phyllolepids and on placoderm 
interrelationships. The head shield is longer than the trunk shield, the reverse of the situation in other 
phyllolepid genera. The presence of two pairs of upper tooth plates, plus a posterior dorsolateral plate, 
an anterior median ventral plate and one or more posterior median ventral plates in the trunk shield is 
confirmed. The phyllolepid jaw apparatus and associated structures, first reported in Austrophyllolepis, are 
reinterpreted. The branchial skeleton, an occipital ossification and a fused synarcual, previously unknown 
in phyllolepids, are described. Cowralepis had an ossified vertebral column, a large epicercal caudal fin 
and small pelvic fins but lacked a dorsal fin. The Cowra/lepis material has suffered regional tectonism and 
illustrates why tectonic deformation must be taken into account in the interpretation of fossils from ancient 


fold belts. 


Manuscript received 5 October 2004, accepted for publication 16 February 2005. 


KEYWORDS: Cowralepis, Placoderm, phyllolepid, tectonism. 


INTRODUCTION 


In January1993, during the preliminary 
excavation of a major Late Devonian fish site near 
Canowindra, New South Wales (Ritchie in press), 
the late Mr Reg Dumbrell, a Canowindra resident, 
showed the writer some Devonian fish specimens he 
had collected from another site, about 15 km south, 
in Cowra Shire. Dumbrell’s best find, on a piece of 
black shale 10 cm square, was a small, articulated 
armoured fish, a phyllolepid, with the tail of a second 
individual lying beside it. Articulated phyllolepids 
had previously been recorded from only two sites in 
the world — Dura Den, Fife, Scotland, and Mt Howitt, 
Victoria, Australia. 

The source of Dumbrell’s find was unexpected; 
the margins of the approach roads to Merriganowry 
Bridge, a causeway over the Lachlan River, 20 km 
northwest of Cowra, NSW, and about 2 km from any 
natural rock outcrop. In March 1993 the writer and 


colleagues visited the site, located more fish fossils 
and solved the mystery. The fish-bearing black shale 
was not in situ. It had been quarried elsewhere, 
trucked in and crushed for road-base material - the 
local roads were literally paved with fossil fish! 

Cowra Shire Council confirmed the source to be 
a small quarry 2 km from Merriganowry Bridge, just 
north of Merriganowry Hall and only 50 metres east 
of the main road from Cowra to Forbes (Young 1999, 
fig.1B, locality 6). This quarry, on private land, had 
been worked on and off for 10-15 years, apparently 
without anyone noticing, or at least reporting, the 
presence of abundant, well-preserved fossil fishes in 
the shale. 

With permission of the new landowner, Mr Alex 
McLachlan, the writer first visited Merriganowry in 
June 1993 and uncovered a fossil treasure trove. Some 
sixty phyllolepid specimens, large and small, many 
complete with tails, were recovered by the writer in 
three days digging! At a conservative estimate, tens 


A NEW GENUS OF DEVONIAN PLACODERMS 


or hundreds of thousands of fossil fish specimens 
had been quarried from this site and crushed for 
road material before its scientific importance was 
recognised. 

When informed ofits unique scientific importance, 
Mr McLachlan immediately arranged for the quarry 
to be secured and fenced, at considerable cost, and 
made it available to the Australian Museum for 
long-term, systematic investigation under scientific 
supervision. Since September 1993 the Merriganowry 
site has been excavated by paying volunteer groups 
supervised by the writer and Dr Zerina Johanson from 
the Australian Museum and organised by an eco-tour 
operator, Monica Yeung, Gondwana Dreaming Inc., 
of Canberra. 

All of the material collected by these groups 
has been retained for scientific study and deposited 
with the Australian Museum, providing a unique 
scientific research resource. Several hundred fish 
specimens have been recovered from Merriganowry, 
many of them articulated individuals complete with 
tails, and they represent a continuous growth series 
from juveniles 5 cm long to adults over 35 cm long. 
Strangest of all, after 11 years of supervised group 
digs, every fish fossil recovered from Merriganowry 
(with the exception of dissociated scales in associated 
coprolites) appears to belong to a new genus of 
phyllolepid placoderm, Cowralepis. 

Phyllolepids were dorsoventrally flattened 
placoderms with a distinctive, sub-concentric ridged 
ornament, hence their name meaning ‘leaf-scale’. 
Most phyllolepid finds consist of dissociated or 
fragmentary dermal bony plates; articulated specimens 
are extremely rare, which is why the Merriganowry 
site is So important. 

Most of our previous knowledge of phyllolepids 
comes from three genera: Phyllolepis from the 
Late Devonian of the Northern Hemisphere, and 
Austrophyllolepis Long, 1984 and Placolepis Ritchie, 
1984 from the Middle-Late Devonian of Australia; for 
reviews of earlier phyllolepid discoveries see Long 
(1984 263-4) and Ritchie (1984 321-3). Many of the 
new phyllolepid discoveries since 1984 have come 
from Gondwanan sites. In Australia these include 
the Amadeus and Georgina Basins, central Australia 
(Young 1985, 1988, in press a), southeastern New 
South Wales (Young in press b), Victoria (Long 1989) 
and Queensland (Turner et al. 2000). Phyllolepids have 
also been described from Antarctica (Young 1989, 
1991; Young and Long in press), Venezuela, South 
America (Young et al. 2000; Young and Moody 2002) 
and Turkey (Janvier 1983). Articulated phyllolepids 
have also recently been reported from Pennsylvania, 


216 


U.S.A. (Daeschler et al. 2003). 

Phyllolepid fishes have not yet been recorded 
either from Asia, where diverse fossil fish assemblages 
of appropriate age are extensively documented, or 
from Africa. 


GEOLOGICAL SETTING 


Horizon and age. 

The phyllolepids described here all come from 
the Merriganowry Shale Member that was uncovered 
by quarrying for road-base material in the 1980’s and 
early 1990’s. The more resistant outcrops (rhyolitic 
volcanics) forming the east margin of the quarry 
had initially been mapped as part of the Canowindra 
Volcanics, of possible Silurian age. The quarry 
exposed a conformable relationship between the 
fish-bearing black shales and underlying volcanics, 
showing that the latter must also be Devonian in age, 
not Silurian. 

Young (1994, 1999, figs 2, 3) correlated the 
volcanic sequence at Merriganowry with the much 
younger Dulladerry Volcanics, which cropout over 
a large area of NSW between Narromine, Dubbo, 
Forbes and Bathurst and underly the Late Devonian 
Hervey Group on the western edge of the Hervey 
Syncline. The youngest Dulladerry Volcanics are 
now estimated to be Late Middle Devonian in age 
(Givetian), which would date the Merriganowry 
black shales as either Late Givetian or Early Frasnian 
(Young 1999, fig. 5B). This is consistent with the fish 
and associated palaeobotanical evidence. 

No direct contact has been identified between 
the west-dipping shale and volcanic sequence at 
Merriganowry and the overlying Hervey Group, but 
sandstone ridges | km north of Merriganowry quarry, 
with the same strike and dip, have been mapped as 
Peaks Sandstone, the basal unit of the thick Hervey 
Group, suggesting a conformable sequence between 
them. 

Although the very limited exposure of the 
Merriganowry Shale Member precludes a total 
thickness estimate, its exposed lower section, directly 
overlying the volcanics in Merriganowry quarry, 
dips at 45° to the WSW (245°), and the fish-bearing 
sequence is estimated to be at least 25 metres thick. 
Excavations over the past 12 years have confirmed 
the presence of fish remains on hundreds of different 
bedding planes throughout the sequence, from just 
above the underlying volcanics to the top of the 
section exposed in the quarry. 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Environment of Deposition. 

Young (1999, 144; P. O’Brien, pers. comm. 1994) 
reported that “the locality is a black shale deposit 
closely associated with the underlying volcaniclastics 
including rhyolite blocks and possible flows. The 
shale includes slumped beds and graded interbeds of 
rhyolitic sand up to 30 cm thick, suggesting deposition 
in a lake of sufficient size and depth for turbidity 
currents to be generated, with water depths of tens 
to hundreds of metres, perhaps an elongate deep lake 
such as are associated with strike slip faulting.” 

Krynen (in Pogson and Watkins 1998, 224-5), in 
formally naming the Merriganowry Shale Member, 
noted “this sequence suggests initial subaerial 
outwash fan deposition in a volcanic rift setting 
passing upwards to turbiditic deposition in a deep 
lake environment following at least local cessation of 
volcanism. The formation of the lake may have been 
associated with strike-slip faulting within the volcanic 
rift.” 


Associated fauna and flora 

The Merriganowry fauna is extremely restricted. 
The only identifiable fish remains, recovered after 11 
years of systematic collecting with the assistance of 
many hundreds of volunteers, almost all appear to 
belong to one genus of phyllolepid, Cowralepis gen. 
nov., described here. A few fragments of eurypterid 
integument have been recovered from one layer high 
in the section. Apart from these, the only other fossils 
recovered are fragmentary plant remains including 
lycopods (‘Protolepidodendron’) and _ branching 
stems (cf. Prearamunculus) and what appears to be 
an egg sac (Fig. 18D). 

What the abundant phyllolepids ate remains a 
mystery - probably plant debris and small invertebrates 
not preserved as fossils. Coprolites up to 20 mm long, 
presumably derived from the dominant phyllolepids, 
are common throughout the Merriganowry section 
but are invariably deeply weathered. When cleaned 
out, these may reveal traces of their contents, but 
preserved only as natural moulds. They include 
comminuted bone fragments, some of which appear 
to be of phyllolepid origin, and several coprolites 
contained abundant minute scales, about the size and 
shape of thelodont denticles, but not well enough 
preserved to be identifiable. 

It appears unlikely that the phyllolepids lived 
on the bottom of the lake because the shale layers 
are generally smooth, flat and undisturbed, free of 
burrowing organisms and with very few trace fossils. 
The abundance of articulated phyllolepid specimens 
complete with delicate tails indicates an absence of 


Proc. Linn. Soc. N.S.W., 126, 2005 


scavengers and suggests that the bottom conditions 
were probably anoxic. 

The observation that about half of the 
dorsoventrally flattened phyllolepids found in situ at 
Merriganowry were buried upside down suggests that 
they probably lived in the upper, better-oxygenated 
water levels and only after death fell randomly 
to the bottom where they lay undisturbed until 
covered. The same conditions must have persisted 
for many thousands of years as the shale deposit 
slowly accumulated, during which time the fauna 
was dominated by phyllolepid fishes to the virtual 
exclusion of all other groups. 

The Merriganowry fauna, and the conditions 
under which it lived and was buried, thus presents a 
sharp contrast with the Late Devonian mass-kill fish 
site discovered only a few kilometres to the north, 
near Canowindra, New South Wales (Ritchie in 
press). The Canowindra assemblage of placoderms 
and sarcopterygians, dominated by two antiarch 
genera, Bothriolepis and Remigolepis, comes from 
a single bedding plane on which many thousands 
of fishes lie crowded tightly together, almost all of 
them (about 95%) buried right way up. Canowindra 
represents a unique time capsule, a sample of a single 
population killed and buried quickly as the result of 
a local environmental disaster, probably an extreme 
drought. 


MATERIAL AND METHODS 


Note: to facilitate cross-referencing of specimens all 
figures are located at the end of the paper 
from page 240 


The original bone was often preserved but 
deeply weathered and was removed by washing, by 
airbrasives, mechanically or with dilute hydrochloric 
acid. The resulting detailed natural moulds (negatives) 
were cast using black-pigmented latex rubber. The 
latex casts were whitened with ammonium chloride 
sublimate and photographed with a Nikon Coolpix 
digital camera. Most of the images used here show the 
original uncorrected dimensions, with the exception 
of several specimens (Figs 6a-D, 9F, 10H, 11B, 16A- 
C) in which these have been digitally modified, as 
indicated, using Adobe Photoshop LE, to correct for 
the effects of regional tectonic deformation. 

Silicone rubber moulds were prepared from the 
latex casts (which have a limited shelf life) and most 
of the figured specimens have been replicated as resin 
casts (positives) for collection and study purposes 
and for scientific exchange with other institutions. 


DG 


A NEW GENUS OF DEVONIAN PLACODERMS 


Interpreting tectonically deformed fossils — the 
long and the short of it. 

Not only do the Merriganowry phyllolepids 
represent a very wide growth range, from juveniles 
to adults, but they also display considerable variation 
both in relative proportions and in ornament (Fig. 2). 
Some specimens are deceptively symmetrical (Fig. 
5), but most show visible evidence of deformation 
and/or asymmetry. 

They confirm a) that the Merriganowry Shale 
Member had undergone significant shear strain, b) that 
the fish fossils provide a means of directly measuring 
this and c) that much of the morphological variation 
and deformation at Merriganowry can be attributed 
to post-mortem tectonic deformation, which must be 
taken into account in interpreting such material. 

Many of the fossils found in areas subjected to 
major regional tectonism are distorted, a phenomenon 
documented for many types of fossils since the 
mid-19th century. Ramsay and Huber (1983, 127- 
149) illustrated how tectonically deformed fossils 
(trilobites, brachiopods, corals, graptolites, ammonites 
etc.) could provide structural geologists with a useful 
method of strain measurement. Graphical techniques 
developed for restoring deformed fossils have mostly 
been used for invertebrates and plants (Cooper 1990) 
but less often for vertebrates. 

Tectonic strain is most readily detected in 
fossil organisms that were originally bilaterally 
symmetrical, where the original axis of symmetry 
can only lie in two directions, at right angles on the 
bedding plane without any loss of symmetry. Such 
organisms (the Welsh Ordovician trilobites, Angelina 
and Bathyuriscus are classic examples) provide 
the simplest subjects for study (Ramsay and Huber 
1983, figs 8.7, 8.8). Where the original shape of these 
fossils is already known it can be used to determine 
the shear strain in deformed specimens and restore 
them to their original shape. Dorsoventrally flattened 
(and bilaterally symmetrical) armoured fishes such as 
phyllolepids provide ideal subjects especially where, 
as at Merriganowry, innumerable specimens were 
buried articulated and undisturbed post mortem. 

Despite the widespread use of length/breadth 
ratios and angular measurements in the description 
and classification of fossil vertebrates, such 
correction techniques have rarely been applied to 
fossil vertebrates. Failure to recognise, and correct 
for, tectonic deformation has undoubtedly led to 
misidentification of fossil species and proliferation of 
invalid taxa. 

Because most of the Cowralepis specimens were 
discovered only after blocks had been excavated 
for splitting by volunteers, their original orientation 


218 


in the ground had been lost. A small number of 
Cowralepis specimens were found in situ (<S%) as 
the outcrop was excavated and their orientation could 
be measured before removal. 

The extremes of deformation displayed by 
Cowralepis specimens from Merriganowry are 
illustrated by two types of specimens displaying 
bilateral symmetry - broad symmetric and narrow 
symmetric - depending on whether the broad 
symmetry axis is parallel to the long or to the short 
axis of the strain ellipse (Fig. SA-D). Such specimens 
cannot be used to determine shear strain. Most of 
the phyllolepids from Merriganowry are visibly 
asymmetric, or skewed, to various degrees, and 
appropriate correction techniques need to be applied. 


Digital reconstruction of deformed fossils. 

The first step is to determine the direction of 
greatest principal extension (GPE) and smallest 
principal extension (SPE) of the fossil. At 
Merriganowry the GPE coincides approximately 
with the strike direction of the steeply dipping shale 
layers and the SPE lies at right angles to it, up-dip 
and down-dip. Cowralepis specimens discovered in 
situ are marked with strike and dip directions before 
removal (Fig. 3B). 

In specimens where the original orientation of 
the specimen has been lost the GPE orientation may 
be determined from prominent directional wrinkling 
of the smooth bony plate surfaces. This marks the 
intersection of the axial plane cleavage with the 
bedding plane (Figs 1A, 4A, 5A-D, 11A, 14A), and its 
orientation can be estimated to within a few degrees. 

Where such surface wrinkling is not well 
developed, the GPE can be identified using 
standard techniques used by structural geologists 
for determining first generation cleavage and 
incipient cleavage in clastic rocks of low to very 
low metamorphic grade (Durney and Kisch, 1994). 
Dr David Durney, Macquarie University, visited 
Merriganowry with the writer to advise on their use 
in determining the GPE of Cowralepis specimens. 

Using graphical methods described by Ramsay 
and Huber (1983, fig. 8.5, Breddin curve) and Cooper 
(1990, fig. 5, Wellman’s method - deformed right 
angles) the degree of shear strain at Merriganowry 
was estimated. It was then tested on several examples 
in which two Cowralepis specimens were recovered 
lying close together on the same bedding plane but 
in different orientations. The availability of modern 
computers and digital image processing programmes 
makes this process much easier. After trial and error, 
the approximate degree of correction required to 
bring both specimens back to symmetry, and to the — 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


same proportions, was determined and this has since 
been applied to many single Cowralepis specimens 
(Figs 6A-D, 9E, F, 10G, H, 11A, B). 

When the GPE had been determined, a digital 
image of the fossil (or cast) was rotated until the GPE 
was horizontal. Using Adobe Photoshop the image 
was then reduced by 10% in the direction of GPE and 
increased by 10% in the direction of SPE. Reducing 
the GPE and increasing the SPE by the same amount 
produces a corrected image with approximately the 
same surface area as the deformed original (Figs 9F, 
10H, 118; strain directions indicated by arrows with 
percent correction indicated). 


How many species? 

The remarkable range of morphological variation 
in phyllolepids from Merriganowry raises the question 
— how many species are present at this site? If the 
specimens illustrated here (cf. Figs. 2, 5) had been 
recovered from several different sites, how many 
‘species’ or even ‘genera’ of phyllolepid might have 
been erected? 

Many factors are involved: the presence of 
unique characters and morphological variation 
(breadth/length and angular measurements); 
variation in ornament; extent and direction of sensory 
canal grooves; differential changes during growth 
(allometry); differential changes through time 
(heterochrony); differences in preservation (selective 
diagenesis); sexual dimorphism; sample size, and 
whether the sample comes from one horizon or 
from different levels ; and, where relevant, tectonic 
deformation. 

This is illustrated by the problems encountered 
in interpreting phyllolepid material from three widely 
separated Devonian sites in southeastern Australia 
— Mount Howitt, Victoria (Long 1984), Nettleton’s 
Creek, New South Wales (Ritchie 1984) and 
Merriganowry, New South Wales (this paper). 

The Mount Howitt fauna, excavated by Monash 
University teams in the 1960’s and 70’s, comprises 
a diverse assemblage of late Middle Devonian 
fishes (placoderms, acanthodians, sarcopterygians, 
actinopterygians) represented by hundreds of 
articulated specimens in all stages of growth. These 
include a small number of placoderm specimens 
described by Long (1984) as a new genus of 
phyllolepid, Awstrophyllepis, represented by two 
species, A. ritchiei and A. youngi. (1984, figs 8 and 
13 respectively) 

Long (1984, 274) differentiated A. ritchiei (fig. 7) 
from A. youngi(fig 13)“.. only by the proportions of the 
dermal armour (figs 8, 25), specifically the preorbital, 
paranuchal, nuchal, median dorsal, anterior and 


Proc. Linn. Soc. N.S.W., 126, 2005 


posterior ventrolateral plates. Although the ornament 
appears to be more finely developed in A. youngi 
relative to A. ritchiei it is not a distinguishing feature 
for the species as a whole.” A. ritchiei was shorter 
and wider and A. youngi was longer and narrower, but 
otherwise there was very little to separate them. 

Long (1984, fig. 8) also analysed proportional 
differences in the breadth/length index of the nuchal 
and median dorsal plates of Austrophyllolepis, based 
on a small sample of 23 specimens. These appeared 
to fall into two distinct clusters for each plate, with 
no overlap, and were interpreted as support for two 
species of Austrophyllolepis at Mt Howitt. 

The writer suspected this interpretation might 
be incorrect after examining several specimens of 
Austrophyllolepis from Mt Howitt in the Australian 
Museum collection (not included in Long’s sample), 
some of which, when measured, fell between the 
ranges shown for A. ritchiei and A. youngi. Casts of 
the figured specimens of A. ritchiei (Long 1984, figs 
3, 4) and A. youngi (1984 fig. 9) showed the same 
signs of deformation as those seen in phyllolepids 
from Merriganowy (asymmetry, surface wrinkling 
aligned in the direction of GPE) indicating that the 
Mt Howitt phyllolepids had also experienced tectonic 
deformation. 

The computer image correction techniques 
used on Merriganowry phyllolepids were applied 
to the Mt Howitt specimens with similar results. 
The Austrophyllolepis ritchiei specimens became 
longer, narrower and symmetrical and the A. youngi 
specimens became shorter, broader and symmetrical. 

Long later admitted (1999, 36) “. . that there may 
only be one species present, as deformation of the strata 
can account for the distortion of plate measurements 
that separate the two originally named species.” The 
hypothesis that only one species of Austrophyllolepis 
is present at Mt Howitt can be readily tested by the 
recovery and analysis of additional Austrophyllolepis 
material from the Mount Howitt site, which is still 
accessible, and checking whether the deformation 
present in the fossils is directly related to regional 
tectonism. 

Asimilar wide range in length/breadth proportions 
was also observed in the head and trunk plates of 
another phyllolepid, Placolepis budawangensis 
Ritchie (1984), from Nettleton’s Creek, NSW 
described at the same time as Austrophyllolepis. This 
was especially noticeable in the nuchal plates (1984, 
figs 4, 5). Several ventral trunk shield plates, found 
closely associated on the same bedding plane (but 
lying in different orientations), probably came from 
one individual. They displayed similar ornament but 
differed markedly in shape and proportions (1984, fig. 


219 


A NEW GENUS OF DEVONIAN PLACODERMS 


11). This was attributed to post-mortem deformation 
caused by regional tectonism. All of the material came 
from one small excavation about two metres square 
in a creek bank, and from the same siltstone unit. The 
writer concluded that only one species of Placolepis 
was represented in the Nettleton’s Creek fauna. 

The same applied to the associated material of 
Bothriolepis longi Johanson and Young (1999, figs 3, 
5, 6). The presence of long/narrow and short/broad 
variants of Bothriolepis initially raised the possibility 
of two species being present (1999, 66) but this 
was dismissed. The morphological deformation at 
Nettleton’s Creek is largely the result of regional 
tectonism. 

In the analysis of tectonically deformed material, 
standard length/breadth ratios and angle measurements 
are of limited value. The writer applied the same 
length/breadth index method used by Long (1984, fig. 
8) to Cowralepis, but based on a much larger sample 
- 143 nuchal plates and 124 median dorsal plates. 
The length/breadth distribution of both samples (not 
shown here) were similarly very widely scattered 
but not meaningfully clustered. It is clear that such 
graphs cannot be used to demonstrate the presence 
of one or more species in a fauna, especially where 
the specimens come from different layers and where 
tectonic deformation is involved, as at Merriganowry, 
Mount Howitt and Nettleton’s Creek. 

The articulated phyllolepid specimens from 
Merriganowry, covering a wide size range, made 
possible another approach. Unlike length/breadth 
ratios, plate ratios along the same axis are not altered 
by distortion, so the most useful plates for analysis are 
the large median plates in the head and trunk shield, 
the nuchal and median dorsal plates. 

Eighty specimens (including many of the figured 
specimens) were selected in which both the nuchal 
and median dorsal plates remained in close association 
and lay in the same orientation. The length of each Nu 
and MD was measured separately along the midline 
and combined to provide data for the horizontal axis; 
the vertical axis illustrates the relative lengths of the 
nuchal and median dorsal plates (Fig. 3A.) 

The eighty specimen points (Fig. 3A) are widely 
scattered, with little obvious clustering (other than an 
abundance of smaller individuals), and are of little 
use in determining whether one or more species of 
phyllolepid are present at Merriganowry. What they 
do appear to illustrate is a dramatic allometric change 
in the relative lengths of the Cowralepis head and 
trunk shields during ontogeny. This is discussed more 
fully in the description of the dermal skeleton. 

Based on the material available for study, only 
one genus of phyllolepid placoderm appears to be 


220 


present at Merriganowry. Cowralepis gen. nov. 
displays a combination of characters not found in 
other phyllolepid genera. These include: head shield 
longer than trunk shield; only one pair of sensory canal 
grooves (central sensory line) crosses from circum- 
nuchal plates onto the Nu plate; lateral line canal 
groove passes off PNu onto Mg plate between 78- 
92% of PNu length (intermediate between Placolepis 
and Austrophyllolepis); a fenestra is present between 
the PtO, Mg and PNu plates and the Nu plates (most 
noticeably in juvenile specimens); a small posterior 
dorsolateral plate is present under the lateral margins 
of the median dorsal plate. 

The phyllolepid specimens from Merriganowry 
cover a wide size range and display a remarkable 
degree of morphological variation; they also come 
from many different levels within the Merriganowry 
Shale Member and thus represent a large and mixed 
sample covering a wide time span. Most importantly, 
every phyllolepid specimen from Merriganowry 
has been deformed, limiting the use of standard 
proportional length/breadth ratios for taxonomic 
determination. 

While it is certainly possible that the material 
recovered from Merriganowry may well include more 
than one species of Cowralepis, it is not possible to 
confirm or refute this at present. 

The course proposed here is to designate a single 
species, Cowralepis mclachlani sp. nov. to establish 
the new taxon in the realisation that it may include 
material from more than one species. It will take 
many more years of excavation, including the use of 
heavy equipment, to sample throughout the whole 
shale section at Merriganowry before the question of 
how many species are present can be tested. Twelve 
years of digging using many hundreds of volunteers 
has barely made a dent in the Merriganowry quarry, 
but this unique site has the potential for a long-term 
systematic research program, with educational and 
eco-tourism benefits as a bonus. 


Abbreviations. 

ADL, anterior dorsolateral plate; AL, anterior 
lateral plate; AMV, anterior median ventral plate; ant 
ridge, inner anterior ridge on AVL plate; Art, articular; 
AVL, anterior ventrolateral plate; Bhy, basihyal; 
buttr AL, buttress on inner surface of ADL plate, 
below AL plate; Chy, ceratohyal; esl, central sensory 
line; d.end, ductus endolymphaticus; d.pr, dorsal 
process; d.sp, dorsal spine; f.bhy, buccohypophysial 
foramen on parasphenoid; fen fenestra; GPE, 
direction of Greatest Principal Extension; h.arch, 
haemal arch; h.sp., haemal spine; Ign, inferognathal; 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


IL, interolateral plate; ioc, infraorbital canal groove; 
l.ind, lateral indentation in parasphenoid; Ile, main 
lateral line sensory canal groove; MD, median dorsal 
plate; Mg, marginal plate; mp.PNu, mesial process on 
paranuchal plate; n.arch, neural arch; n.can., neural 
canal; not., notochord; Nu, nuchal plate; oa.AVL, 
overlap area for AVL plate; oa.Nu, overlap area for 
nuchal plate; Occ oss, occipital ossification; Ot, otolith; 
p-pao, paraotic plate; PDL, posterior dorsolateral 
plate; pect, pectoral embayment (margin) of AVL 
plate; pel.b, basal plate of pelvic fin; pel.d, distal 
element of pelvic fin; PMg, posterior marginal plate; 
PMV, posterior median ventral plate; PN, postnasal 
plate; pnpr, postnuchal process of paranuchal plate; 
PNu, paranuchal plate; ppl, posterior pitline; PrO, 


preorbital plate; Psp, parasphenoid; PtO, postorbital — 


plate; Sgn ant, anterior superognathal; Sgn post, 
posterior superognathal; SMg, submarginal; soc, 
supraorbital canal groove; Sp, spinal plate; SPE, 
direction of Shortest Principal Extension; sub-h.sp., 
sub-haemal spine; Syn, synarcual; vpl, ventral pitline; 
Zyg., zygopophosis. 


SYSTEMATIC PALAEONTOLOGY 


Class PLACODERMI McCoy, 1848 
Order PHYLLOLEPIDA Stensi6, 1934 


Diagnosis 

Placoderms with greatly enlarged nuchal plate, 
slightly wider than long, surrounded by five pairs of 
smaller bones — paranuchals, marginals, postorbitals, 
post nasals and preorbitals. Paranuchal plate with 
well-developed postnuchal process. Rostral, pineal 
and central plates absent from skull roof. Anterior 
and posterior superognathals; anterior superognathal 
large and wide, posterior superognathal very 
small; inferognathal long, narrow anteriorly, wider 
posteriorly. Trunk armour broad and dorsoventrally 
flattened; median dorsal plate lacks an inner keel; 
anterior dorsolateral plate with long, narrow exposed 
area; posterior dorsolateral plate reduced or absent; 
anterior lateral plate small, rhomboid; posterior 
lateral plate absent. Anterior ventrolateral plate short 
and broad, with ossification centre near posteromesial 
comer; posterior ventrolateral plate triangular, with 
ossification centre near anteromesial corner; both 
plates relatively flat with straight, non-overlapping 
mesial suture. Interolateral plate long, narrow, with 
denticulate postbranchial lamina; anterior median 
ventral plate short, broad, separating interolaterals; 
posterior median ventral plate(s) small, narrow, 


Proc. Linn. Soc. N.S.W., 126, 2005 


variable in number (0-2). Dermal ornament consists 
of smooth, slightly undulating, subconcentric ridges, 
locally of rows of tubercles. Pelvic fins small; dorsal 
fin absent; caudal fin large, epicercal. 


COWRALEPIS MCLACHLANI 
new genus and species 
Figs 1-20 


‘a new genus and species of phyllolepid’: Krynen, 
1998, 225. 
‘a new phyllolepid’: Young, 1999, 144. 


Name 

The genus is named after Cowra (Aboriginal for 
‘place of the rocks’), the nearest town in central west 
New South Wales; and /epis (Gk), ‘scale’. 

The species name acknowledges the contribution 
of Mr Alex McLachlan, owner of ‘Mooroonbin’, 
near Cowra, NSW (which includes Merriganowry), 
in enclosing the quarry site and making it available 
to the Australian Museum for scientific investigation; 
also after the Lachlan River near where the first 
discoveries were made. Only one species, Cowralepis 
mclachlani sp. nov. is recognized at present from the 
Merriganowry site. 


Repository 

All of the described, figured and mentioned 
specimens in this paper are lodged in the palaeontology 
collections of the Australian Museum, Sydney, as 
indicated by the prefix AMF. 


Holotype 

AMEF103767a, b, medium sized individual 
complete with tail, seen in dorsal and ventral view 
(Fig. 1A-D) 


Figured material 
AMF90003a(Fig.4A,6A);AMF90004 (Fig. 16C); 
AMF90007b (Figs 12A, 14A); AMF9001 1a, b (Figs 
18E, F; Fig. 19); AMF90012 (Fig. 7H): AMF90018 
(Fig. 5D); AMF90027 (Fig. 2C); AMF90029a (Fig. 
7G); AMF90034a, b (Fig. 2D); AMF90044b (Fig. 
21); AMF90048a (Fig. 17A, B); AMF90048b (Figs 
17C, D); AMF90051 (Fig. 5C); AMF90053 (Fig. 
2B); AMF90054a, b (Fig. 7D, E); AMF96747a (Fig. 
2G); AMF96750 (Fig. 2E); AMF96751a (Fig. 13A); 
AMF96755Sa, b (Fig. 13C); AMF96762 (Figs 4B, 6B, 
7C); AMF96764 (Fig. 16A); AMF96765 (Fig. 16B); 
AMF96779 (Figs 7A, B, LOE, 12E); AMF96780 (Fig. 
10F); AMF96781 (Fig. 9A, B); AMF96783 (Figs 9E- 
G, 12F); AMF96784 (Fig. 7F); AMF96785 (Fig. 13D, 


221 


A NEW GENUS OF DEVONIAN PLACODERMS 


E); AMF96786 (Fig. 10G, H); AMF100018 (Fig. 5D); 
AMF103753a, b (Figs 5A, B, 6C, D); AMF103754 
(Fig. 71); AMF103755 (Fig. 1OA); AMF103756 (Fig. 
2A); AMF103763 (Fig. 13B); AMF103768 (Fig. 9D); 
AMF103770 (Fig. 8C, D); AMF103776 (Fig. 14B); 
AMF103778a (Fig. 2J); AMF103784 (Fig. 2H); 
AMF103787 (Fig. 10D); AMF104154b (Figs 8A, 
B, 9C); AMF 104155 (Fig. 10C); AMF104157a (Fig. 
2F); AMF104160 (Fig. 8F); AMF104164 (Fig. 10B); 
AMF12715la (Fig. 18D); AMF127152 (Fig. 8E); 
AMF127154b (Fig. 18B); AMF127156 (Figs 11A, B, 
12B-D); AMF127159 (Fig. 18A); AMF127162 (Fig. 
8G). 


Locality 

All of the figured material comes from a small 
quarry, the type locality (and only known exposure) 
of the Merriganowry Shale Member. The quarry 
(GR642750 6373750) is on a small rise on the east side 
of the Cowra to Forbes road on Mooroonbin property, 
1.5 km northwest of Merriganowry homestead and 20 
km northwest of Cowra, New South Wales. 


Diagnosis 

Moderately large phyllolepid, reaching ca. 35 cm 
in length. Widest part of nuchal plate lies just posterior 
to centre of ossification; anterior nuchal margin 
more angular than in Placolepis, less angular than 
in Austrophyllolepis and Phyllolepis. Nuchal plate 
same length as or longer than median dorsal plate in 
juveniles; nuchal plate 20-60% longer than median 
dorsal plate in adults. Supraorbital canal, infraorbital 
canal and pit-line grooves not developed on nuchal 
plate. Lateral line canal groove on paranuchal plate 
diverges anteriorly from nuchal margin and canal 
groove crosses anterolateral margin between 78-92% 
PNu length. Marginal plate separates postorbital 
and paranuchal plates. Marginal plate and posterior 
division of postorbital plate lack contact with the 
nuchal plate in juveniles, leaving fenestra between 
them; marginal plate and posterior division of 
postorbital plate meet lateral margin of nuchal in 
adults, fenestra reduced. Posterior dorsolateral plate 
present; small, subtriangular, hidden under lateral 
margin of median dorsal plate; posterior lateral 
plate absent. Anterior and posterior median ventral 
plates both present. Occipital region of endocranium 
ossified; vertebral column fused anteriorly, forming 
long narrow synarcual under median dorsal plate. 


Remarks 

In addition to Cowralepis gen. nov. described here, 
there are now five other named genera of phyllolepids 
- Phyllolepis, Austrophyllolepis, Placolepis, plus two 


222 


new genera recently described by Young (in press) 
from the Middle Late Devonian of the Pambula area, 
southeastern New South Wales. The later are known 
only from a handful of dissociated, and mostly 
incomplete, head and trunk plates. 

Cowralepis differs from all of these in that the 
head shield is longer than the trunk shield; it is also 
the only phyllolepid known in which a fenestra is 
developed between the nuchal (Nu) and lateral cranial 
plates (PtO, Mg, PNu). Cowralepis differs from 
Phyllolepis, Austrophyllolepis and Placolepis in the 
presence of a posterior dorsolateral plate (PDL) in the 
trunk shield; a feature it apparently shares with the 
two new genera from near Pambula, NSW (Young, in 
press) discussed beow. 


DESCRIPTION 


The three main developmental divisions of the skull 
and skeleton in vertebrates are 
a) dermal skeleton - includes the skull roof, 
cheek and operculum, denticulated or tooth- 
bearing bones of the palate and inside and 
outside of the lower jaw, and small dental 
plates of the buccal cavity and inside visceral 
arches. 
b) endocranium. 
c) visceral skeleton - Meckel’s cartilage plus 
palatoquadrate, forming the core of the upper 
and lower jaws, plus branchial arches. 


Dermal skeleton 

The head shield of phyllolepids consisted of a 
very large nuchal plate (Nu) bordered anteriorly and 
laterally by five pairs of smaller plates, interpreted 
here as the preorbitals (PrO), postnasals (PN), 
postorbitals (PtO), marginals (Mg) and paranuchals 
(PNu), the last of which articulate with the trunk 
shield. Another pair of small plates, the submarginals 
(SMg) flanked the Mg plates laterally, but were only 
loosely associated with them. 


Nuchal 
In Cowralepis the Nu was slightly wider than long 
and sub-polygonal in shape (Fig. 6A, C, 15A) with the 
greatest width posterior to the centre of ossification. 
In this it is closest to Placolepis and differs from 
Phyllolepis and Austrophyllolepis, in which the Nu 
is always widest anterior to the centre of ossification 
(Fig. 20B). The anterior margin of the Nu was more 
angular than in Placolepis and less angular than in 
Phyllolepis and Austrophyllolepis. 
Cowralepis differs most 


noticeably from — 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Placolepis, Austrophyllolepis and Phyllolepis in the 
relative lengths of its nuchal and median dorsal plates. 
In the other three genera the Nu plate is always shorter 
than the MD. In all but a few Cowralepis specimens it 
is the Nu that is longer, sometimes much longer, than 
the MD. Eighty Cowralepis specimens were selected 
in which the dorsal shield was well preserved and the 
Nu and MD plates were closely associated and lying 
in line. Unlike length/breadth measurements, the 
relative lengths of such plates should be unaffected 
by the tectonic deformation. 

The results, illustrated graphically in Fig. 3A, 
show the relative length of the Nu and MD plates in a 
wide range of growth stages of Cowralepis and appear 
to indicate a dramatic increase in length of the head 
shield relative to the trunk shield during ontogeny. 


Length Nu+MD samplenumber Nu/MD x 100 
30 —50 mm 20 individuals av. 110.7% 
50 —70 mm 33 individuals av. 107.8% 
70 —90 mm 10 individuals av. 108.8% 
90 — 110 mm 11 mdividuals av. 123.2 
-110—130mm _ 2 individuals av. 113.5% 
130—150mm _ 2 individuals av. 134.0% 
150-170 mm _ 2 individuals av. 139.0% 


In only a few specimens of Cowralepis was the 
Nu plate slightly shorter than the MD. In sixty- three 
individuals with a combined Nu+ MD length between 
30 mm and 90 mm, the Nu plate was on average 8- 
10% longer than the MD. The main increase in the 
relative length of the head shield appears to have 
taken place from 90 mm (Nu + MD) upwards, with 
many specimens known in which the Nu plate is 20- 
40% longer than the MD. 

In the holotype, AMF103767 (Figs 1A, 3A), the 
Nu is 32% longer than the MD; in AMF90003A (Figs 
4A, 6A) the Nu is 39% longer. The most spectacular 
example is also the largest known specimen of 
Cowralepis, AMF103754a (Fig. 71) in which the Nu 
plate is 60% longer than the MD. 

These results indicate that, in at least one 
phyllolepid, Cowralepis, the head shield grew 
relatively larger than the trunk shield during later 
stages of growth and the head shield/trunk shield 
length index was not fixed but varied with age. 

Cowralepis differs from all other phyllolepid 
genera in that only one pair of sensory canal grooves 
cross onto the Nu plate from the surrounding circum- 
nuchal plates; this is the central sensory line (Figs 2, 
5A, C, 15A; csl) which is also visible on the ventral 
surface of the Nu (Figs 4B, 9A, 11). In contrast, three 
pairs of sensory canal grooves cross onto the Nu plate 
in Austrophyllolepis and four pairs in Placolepis and 


Proc. Linn. Soc. N.S.W., 126, 2005 


Phyllolepis (Fig. 20B), as discussed below. 


Circum-nuchal plates 

The two anterior pairs of plates on the head shield 
were interpreted differently in Placolepis Ritchie 
(1984, fig. 2) and Austrophyllolepis Long (1984, 264, 
fig. 1). The writer followed Denison (1978, fig. 29) 
in identifying the median pair as the postnasal plates 
(PN) flanked by preorbital plates (PrO) plates; Long 
interpreted the median pair as the PrO plates flanked 
by the PN plates. The general consensus now is that 
the latter version is preferred, with the qualification 
that some uncertainty remains about the homology of 
the plates identified as the postnasals (Fig. 15A). 


Preorbital plate 

The PrO plates that form most of the anterior 
margin of the head shield are subrectangular, wider 
than long, and meet mesially in a sinuous suture. They 
are subdivided longitudinally by the supraorbital 
sensory canal groove (Fig. 15, soc) into a smaller 
mesial portion and a larger lateral division. The 
anterior margin of the head shield in Cowralepis (Figs 
1, 2, 4-6, 13A, 14B) is straighter than in Placolepis 
and Austrophyllolepis and was not indented medially 
as in Phyllolepis (Fig. 20B). 

In Cowralepis the suborbital canal (soc) 
terminates at the posterior margin of the PrO plate 
and does not cross onto the nuchal plate as in 
Placolepis, Austrophyllolepis, and Phyllolepis, in all 
of which the soc crosses onto the Nu plate, maintains 
the same course and converges towards the centre of 
ossification of the Nu (Fig. 20B). 

An unusually preserved specimen of Cowralepis 
(Fig. 14B, soc) reveals why the supraorbital canal 
grooves appear to terminate suddenly at the posterior 
dermal margin of the PrO plates. In this specimen, 
seen in dorsal view, the circum-nuchal plates are in 
place but the nuchal plate has been lost, exposing 
the normally hidden overlap areas of the PrO and 
PN plates. The supraorbital canal grooves on the left 
and right PrO plates converge posteriorly towards 
the posterior overlap area where they turn sharply 
towards the midline. Just before reaching the midline 
they again turn sharply posteriorly to resume their 
original course, but would have been hidden under 
the anterior margin of the Nu plate. 


Postnasal plate 

The postnasal plates flank the PrO plates laterally 
to form the anterolateral corners of the head shield. 
The PN plates are relatively larger in Cowralepis 
than in Placolepis, Austrophyllolepis and Phyllolepis 
(Fig. 20B). The V-shaped canal groove on the PN 


225 


A NEW GENUS OF DEVONIAN PLACODERMS 


plate is interpreted as a loop of the infraorbital canal 
(Fig. ISA, ioc). It is also visible as a prominent 
ridge on the visceral surface of the plate (Figs 1B, 
4B, 5B, 9A, E, 10B, C, 11, 13A). In Cowralepis 
and Austrophyllolepis the infraorbital canal does not 
cross onto the nuchal plate, unlike in Placolepis and 
Phyllolepis (Fig. 20B). 


Postorbital plate 

The postorbital plate is relatively larger in 
Cowralepis than in Placolepis, Austrophyllolepis 
and Phyllolepis, especially its posterior division. The 
infraorbital sensory canal groove (Figs 15A, 20B, ioc) 
continues over the PtO onto the nuchal plate as the 
central sensory line (csl), as in the other three genera. 
In small to medium-sized specimens of Cowralepis 
the posterior division of the PtO is separated from the 
Nu plates by a crescentic fenestra (Fig. 2G, fen). In 
larger specimens (Figs 1A, 5A, 6A, 11, 15A, B) the 
posterior division of the PtO contacts the Nu margin 
along its full length (cf. below). 


Marginal plate 

TheMgplatein Cowralepisisasmall, subtriangular 
plate, longer than wide, tapering posteriorly and with 
a transverse anterior margin. It was closer in shape 
to the Mg plate of Austrophyllolepis than that of 
Placolepis (Long 1984, fig. 2B). The Mg is often 
partly obscured by the plates around it (Figs 1A, 2, 
5A, B, 6A, B) and its shape is most clearly seen in 
ventral views of the head shield (Figs 4B, 6B, 14A) 
or in specimens that are dissociated (Fig. 7F). A small 
projection on the mesial margin of the Mg fitted into 
a notch in the PNu plate where the lateral line canal 
groove crossed over (Fig. 7F). 

The Mg separated, and was overlapped anteriorly 
and posteriorly by, the PtO and PNu plates (as in 
Placolepis). In smaller Cowralepis specimens (Fig. 
2), the mesial margin of the Mg plate fell far short 
of the Nu margin, leaving a fenestra (shared with the 
PtO) between them. In larger individuals, the Mg plate 
enlarged mesially until it just met the nuchal margin, 
although it is not clear if an overlap relationship had 
developed between them. 

In Cowralepis, the relationship between the Mg 
plate and the plates surrounding it (PNu, PtO and 
Nu) changes during growth and appears to represent 
a morphological condition intermediate between 
Placolepis and Austrophyllolepis, as discussed 
below. 


Paranuchal plate 


The posterolateral corner of the head shield in 
phyllolepids was formed by a rather large paranuchal 


224 


plate with a long, broad lateral division and a narrow, 
tapering, postnuchal process (pnpr) that extended 
mesially almost to the midline (Figs 1, 2, 3A, C, 6A, 
ISA, B). The shape and overlap areas of the PNu in 
Cowralepis are best seen in isolated examples (Fig. 
7F, G) or where the nuchal plate has been dislodged 
(Fig. 14B). 

The lateral division of the PNu was traversed 
by a long curving groove that housed the lateral line 
sensory canal (IIc) and crossed the Mg/PNu boundary 
at different levels in different phyllolepid taxa. This 
character, used by Long (1984, fig. 2A, 266) to 
differentiate Austrophyllolepis from Placolepis and 
Phyllolepis, is one of the most useful characters in 
separating and diagnosing phyllolepid taxa. 

The length of the PNu, measured from its posterior 
margin to where the lateral line intersects its lateral 
margin, is expressed as a percentage of the total PNu 
length (cf. Young in press b, fig. 2A). Because both 
of these dimensions lie in the same direction their 
ratios are not altered by tectonic deformation, unlike 
breadth/length ratios. 

In Austrophyllolepis the lateral line canal crossed 
the lateral margin of the PNu at around 68-72% of 
total PNu length. It is thus intermediate between 
Phyllolepis orvini Heintz (1930) from Greenland 
(Stensid 1934, pl. 5, figs 1, 2) in which the lateral 
canal crossed at 48-56% PNu length and Placolepis 
(Ritchie 1984, fig.6A-C) where the canal crossed onto 
the Mg at the apex of the PNu, effectively 100%. In 
a new genus from southeastern NSW described by 
Young (in press b, figs 3A, C, 4A) the PNu llc index 
is 71%, placing it within the Austrophyllolepis range. 

The same analysis was applied to the 
Merriganowry phyllolepid material. Thirty-six 
examples of Cowralepis PNu plates were measured, 
many of them representing left and right PNu plates 
from the same individual. In Cowralepis the lateral 
line groove crossed the PNu margin at between 
78.5% and 92% of the PNu length, with an average of 
86%. This range therefore falls intermediate between 
that of Placolepis and Austrophyllolepis but does not 
overlap with either of them. 

The lateral line canal groove on the PNu plate 
of all phyllolepids turns sharply mesially towards 
the posterolateral corner of the nuchal plate. In 
Placolepis, Austrophyllolepis and Phyllolepis it 
continues onto the nuchal surface for a short distance 
as the posterior pit-line (ppl), but no trace of a pit-line 
has been detected in Cowralepis. 

The mesial margin of the PNu plate (Fig. 7F, 
G) is formed by a well-developed overlap area for 
the nuchal plate (oa.Nu) and carries a short blunt 
process that lay under the posterolateral corner of — 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


the nuchal plate. On the posterior margin of this 
process lies a deep groove directed anteromesially. It 
is suggested here that this groove (Fig. 7F, G, d.end) 
may have housed the endolymphatic duct that in most 
placoderms reached the surface via a small foramen on 
either the nuchal or paranuchal plate. In Cowralepis 
there is no visible surface opening on either plate but 
it is possible that the endolymphatic duct may have 
opened on the margin between the plates. 

Gavin Young (pers.comm.) has recently described 
anew phyllolepid from the south coast of New South 
Wales in which the paranuchal plate bears a strongly 
developed mesial process on the inner margin of the 
PNu (Young in press b, figs 3A-C, 4A, B, mp.PNu) that 
would have projected under the corner of the nuchal 
plate. This mesial process is clearly comparable to, 
but much larger than, the small blunt process seen in 
Cowralepis paranuchal plates (Fig. 7F, G, mp.PNu). 


Interrelationships of nuchal, paranuchal, 
postorbital and marginal plates. 

The Nu, PNu, PtO and Mg plates in Cowralepis 
illustrate a condition morphologically intermediate 
between that in Placolepis and Austrophyllolepis 
(Figs 15A, 20B). They also document a remarkable 
change in the relationships of these plates throughout 
ontogeny. 

Placolepis and Phyllolepis (Fig. 20B) illustrate 
the extremes of the range in the known phyllolepid 
genera. Placolepis differed significantly from 
Phyllolepis in the size and shape of its PNu plate (much 
shorter than in Phyllolepis) and 1n the relationships of 
the PNu with the Mg, PtO and Nu plates. 

In Placolepis (Ritchie 1984, fig. 8E) the lateral 
line canal groove closely followed the PNu/Nu margin 
to the anterior corner of the PNu where it crossed over 
the Mg onto the PtO. The Mg plate in Placolepis thus 
separated the PNu and PtO plates and was overlapped 
mesially by the lateral margin of the Nu. 

In Phyllolepis (Ritchie 1984, fig.8F) the lateral 
line canal groove on the PNu plate crossed from the 
much smaller Mg plate attached about midway along 
the PNu lateral margin. A large triangular anterior 
extension of the PNu (mesial to the lateral line canal) 
separated both the marginal plate and posterior part 
of the postorbital plate from any contact with the 
nuchal. 

Austrophyllolepis (Long 1984, figs 2, 7, 10, 13, 
19B) displayed a condition intermediate between 
that of Placolepis and Phyllolepis. The Mg plate 
was similar in size and shape to that of Phyllolepis 
but was attached more anteriorly to the PNu margin 
than in Phyllolepis. Austrophyllolepis also resembles 
Phyllolepis, but differs from Placolepis, in its large 


Proc. Linn. Soc. N.S.W., 126, 2005 


anterior extension of the PNu (1984, fig. 2A), which 
separated the Mg and Nu plates and contacted the 
posterior mesial margin of the PtO (Long 1984, figs 
2B, 19B). 

The new evidence from Cowralepis complements 
that of Placolepis, Austrophyllolepis and Phyllolepis. 
Cowralepis resembled Placolepis in that the Mg 
plate separated the PtO and PNu plates, and the 
anterior division of the PNu did not contact the 
PtO. Cowralepis resembled Austrophyllolepis and 
Phyllolepis in that neither the Mg plate nor the 
posterior margin of the PtO plate contact the lateral 
margin of the Nu (leaving a fenestra between them) 
— but only in juveniles and smaller individuals (Fig. 
2). In adult Cowralepis specimens the PtO meets the 
Nu along its full length while the Mg barely contacts 
the Nu, or just falls short, but it always separates the 
Pto and PNu plates (Fig. 15). 

The distinctive shape of the nuchal plate in 
Phyllolepis appears to be the derived condition in 
phyllolepids, compared to that in Placolepis. It could 
have arisen by either: 

a) aposterior migration of the lateral line canal 
groove on the PNu plate - since the angle of 
the lateral canal on the PNu does not appear 
to have changed significantly in the different 
phyllolepid genera, nor have its relationships 
to the marginal and postorbital plates (Long 
1984, fig. 2A; this paper Figs 15, 20B). 

b) OR, the alternative, preferred here, 
progressive enlargement_of the anterior 


division of the PNu plate, mesial to the 


lateral line, first separating the Mg plate 
and, later, the posterior part of the PtO, from 


the Nu - i.e. that the distinctive, parallel- 
sided shape of the nuchal plate in later 
phyllolepids (Phyllolepis) is largely the 
result of differential anterior enlargement of 
the paranuchal plate mesial to the lateral line 
canal groove (Fig. 20B). 

OR by a combination of both processes 


Submarginal plate 

One more dermal bone in the cheek area 
remains to be accounted for. Long described a small 
unornamented bone lying lateral to the marginal 
plate in Austrophyllolepis. It was only seen in two 
specimens, leaving its nature and identification 
uncertain. It lacked a laterosensory groove, appeared 
to be loosely attached to the cheek and was identified 
as the postmarginal plate (Long 1984, 284, fig.19B, 
C; PMg). 

A similar, but much larger, plate is present in most 
of the articulated specimens of Cowralepis (Figs 1A, 


225 


A NEW GENUS OF DEVONIAN PLACODERMS 


2B, C, E-G, I, J, 4A, B, 5D, 6A, B, 7F, 14A, B, 19). Its 
size, shape and relationship to the Mg are best seen in 
AMF96784 (Fig. 7F). The plate was long, narrow and 
slightly curved, with a shallow convex dermal surface. 
A low ridge crossed its dermal surface obliquely from 
the posteromesial corner to the anterolateral corner. It 
lay against the lateral margin of the Mg, but was only 
loosely attached to it and easily dislodged. From its 
position lateral to the Mg this plate appears to have 
formed part of a moveable cheek or gill cover and 
must therefore be the submarginal plate (SMg), not 
the postmarginal. 

Asubmarginal plate is present in many placoderms, 
loosely attached in the cheek area and usually closely 
associated with the hyomandibula (Janvier 1996 
fig. 4.42.14). The long, narrow submarginal of the 
dorsoventrally flattened Cowralepis resembles that 
found in deep-bodied ptyctodonts such as Ctenurella 
(Moy-Thomas and Miles 1971, fig. 8.13; Miles and 
Young 1977, figs 15, 19; Long 1997, figs 25, 28, 29) 
and Campbellodus and Austroptyctodus, both from 
Gogo, W.A. (Long 1997, figs 5, 6 and 28, 29, 35 
respectively) and there can be little doubt that they 
are homologous. 


Trunk shield 

The dorsal and ventral trunk armour of 
Cowralepis (Fig. 15A-C) is basically similar to that 
of other phyllolepid genera, with a few differences 
detailed below. The wide variation in length/breadth 
proportions of the trunk plates in Cowralepis (most 
of which can be attributed to regional tectonism) 
precludes the use of such measurements for diagnosis. 
The trunk (and cranial) plates of Cowralepis also 
display a wide variety of dermal ornament, ranging 
from almost smooth surfaces to dense subconcentric 
ridges, but whether this variation is inter-specific or 
intra-specific remains to be determined. 


Dorsal trunk shield 

The dorsal trunk shield of Cowralepis (Figs 1A, 
2, 4A, 5A, C, 6A, D, 15A, B) consisted of a large, 
subpentagonal median dorsal plate (MD) flanked 
laterally by an extremely narrow anterior dorsolateral 
plate (ADL) carrying the lateral line canal groove. 
A broad anterior flange on the ADL fitted under 
the posterior margin of the PNu plate on the head 
shield. Supported laterally by a similar flange, 
the postbranchial lamina, on the adjacent anterior 
lateral plate (AL), this provided phyllolepids with 
a sliding neck joint like that present in actinolepids 
and wuttagoonaspids. The spinal plate (Sp), linking 
the dorsal and ventral shields laterally, was relatively 
small, with a short posterior spine. 


The dermal surface of the anterior dorsolateral 
plate (ADL) was long and very narrow and often 
obscured by the adjacent AL and MD plates (Figs 1A, 
5A, C). Much more informative are specimens of the 
ADL seen in ventral view and showing its full shape, 
including the extensive mesial and lateral overlap 
areas that contacted the ventral margins of the MD 
and AL plates respectively (Figs 7A, B, 13A, D, 
15B). 

In AMF96779 (Fig. 7A, B) numerous cranial 
and trunk plates from one individual had become 
dissociated before burial. They included ventral views 
of the left (Fig. 7A) and right (Fig. 7B) ADL plates; an 
otolith from the same individual is figured elsewhere 
(Fig. 12E). A strongly developed, smoothly rounded 
ridge (ant ridge) extends the full length of the ventral 
anterior margin of each ADL plate and presumably 
supported the anterior flange on the ADL that fitted 
under the PNu plate in the head shield to form the 
sliding neck joint. 

The inner surface of both ADL plates display 
an unusual feature not previously recorded in 
phyllolepids. The course of the lateral line canal is 
visible as a narrow ridge (llc) on the inner surface, 
running the length of the plate. Where this meets the 
anterior margin another, more strongly developed, 
ridge emerges from the margin ventral to the lateral 
line canal ridge. 

The second ridge diverges posteriorly from the 
lateral line canal at 20-25°, becomes broader distally 
and bifurcates near the rear margin. Along most of 
its length this inner ridge on the ADL plate would 
have underlain the AL plate, acting as a strengthening 
buttress (buttr AL) for the latter. The same features are 
also visible in two figured specimens of Cowralepis 
in which the ADL plate is preserved in situ (Fig. 13A, 
D). 

No trace of either posterior dorsolateral (PDL) 
or posterior lateral (PL) plates has been reported in 
the dorsal shields of Phyllolepis, Austrophyllolepis 
and Placolepis, raising the question whether this was 
a primary or a secondary condition. Both PDL and 
PL plates are present in a wide range of arthrodires; 
separate PL plates are also found in early antiarchs and 
PDL plates in palaeacanthaspids and petalichthyids. 
This suggests that PL and PDL plates were primitive 
components of the placoderm trunk shield. 

The presence of a rather large PDL plate has 
been deduced in two new phyllolepid genera from 
Pambula, NSW, described by Young (in press b), 
although the PDL itself has not been recovered in 
either. In one taxon the median dorsal plate displays 
a short straight posterolateral margin between the 
lateral and posterolateral corners (Young, in press b, — 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


fig. 5A, B, Ic, plc), interpreted as the attachment point 
for a PDL plate. In the second new taxon, the ventral 
posterolateral margin of the MD plate displays a long, 
narrow contact face for a PDL plate (figs 7B, H, 8B, 
cf. PDL). In both of these new genera the PDL plate 
probably extended well beyond the lateral margins of 
the MD. 

Cowralepis provides another example of a 
phyllolepid retaining a PDL plate in its trunk armour 
and illustrates an interesting intermediate condition 
in the reduction and secondary loss of PDL plates 
from the trunk shield. PDL plates are preserved in 
many Cowralepis specimens at all stages of growth, 
including the holotype (Figs 1A, B, 4B, 5B, 6B, D, 
16A, 19), but they are very unusual dermal plates in 
that they were not visible at the surface (Fig. 15A, 
B). 

The explanation is simple: in Cowralepis, the 
PDL plate was reduced to a mere remnant tucked 
neatly under, and attached to, the lateral margin of 
the median dorsal and did not extend beyond it. This 
is best illustrated in AMF96762 (Figs 4B, 6B, 7C), 
seen in ventral view, where the right PDL is still in 
place but the left PDL had become detached and 
drifted sideways. This exposes a contact area for the 
PDL on the ventral side of the MD plate consisting 
of several shallow pits. The PDL was crescentic to 
sub-triangular, with a rounded and thickened lateral 
margin. The anterior extension of the PDL was still in 
contact with the posterior end of the ADL plate (Fig. 
16A). 


Ventral trunk shield 

The ventral trunk shield (Figs 1B, 4B, 5B, D, 6B, 
D, 8F, 15C, 16A,B, 19) consisted mainly of a pair of 
large, wide, flat anterior ventrolateral (AVL) plates, 
bordered anteriorly by long, narrow interolateral 
(IL) plates, and posteriorly by a pair of smaller, 
subtriangular posterior ventrolateral (PVL) plates. 
Whether phyllolepids ever possessed, or have lost, 
paired anterior ventral (AV) plates, like those present 
in actinolepids, wuttagoonaspids and the Chinese 


petalichthyid Eucaryaspis (Liu 1991), remains 
unknown. 
The interolateral (IL) plate was already 


known from Phyllolepis orvini (Stensi6 1936, fig. 
21), Phyllolepis woodwardi Stensié (1939, text- 
fig.2); Austrophyllolepis (Long 1984, figs 18, 20) 
and Placolepis (Ritchie 1984 fig. 11E-D). The IL in 
Cowralepis mclachlani is almost identical (Fig. 8 
A-E), a long, narrow, slightly curved plate tapering 
mesially to a sharp point. The post-branchial lamina 
was strongly convex and ormamented with 9-10 
parallel rows of very fine denticles (Fig. 8D). 


Proc. Linn. Soc. N.S.W., 126, 2005 


Although an anterior median ventral (AMV) 
plate was reconstructed in Phyllolepis orvini (Stensi6 
1969 figs 199A, B) the best evidence for an AMV in 
phyllolepids was provided by an articulated specimen 
of Phyllolepis woodwardi from Dura Den, Scotland 
(Stensi6 1934, text fig. 2D; 1936 text fig. 5; 1939 text 
fig. 2) in which a reasonably large, well-developed 
AMV is preserved in situ, flanked laterally by the 
interolateral plates (IL). 

Long (1984) was unable to identify an AMV in 
the Austrophyllolepis material from Mt Howitt and 
omitted it from the reconstructions of A. ritchiei and 
A. youngi (1984, figs 7, 13). In several specimens 
of Austrophyllolepis (1984, figs 5, 6, 11D, 18C) 
the anteriomesial corners of the AVL are distinctly 


_rounded and part of what may be a small AMV is 


visible in one specimen (fig. 18C). The writer also 
was unable to locate or identify an AMV plate in the 
dissociated material of Placolepis and omitted it from 
his reconstruction (Ritchie 1984, figs 2D, 8C), but 
again the anteromesial corners of the AVL plates are 
slightly rounded (figs 10J, K, 11F, G, 12A, B). 

Cowralepis reveals that the apparent absence of 
an AMV plate in Austrophyllolepis and in Placolepis 
is most probably due to the vagaries of preservation 
rather than original absence. As noted above, well- 
developed and robust IL plates are present in several 
specimens of Austrophyllolepis, but all of them were 
slightly displaced. The AMV was a much smaller 
plate, easily dislodged and lost post mortem. 

In many of the apparently complete, articulated 
specimens of Cowralepis from Merriganowry it is 
not uncommon to find both the AMV and IL plates 
missing. The explanation appears to be that, during 
burial and compaction of the wide but slightly 
convex, ventral trunk shield, the narrow, curved IL 
plates, which supported and presumably strengthened 
the anterior margin of the AVL plates, were displaced 
by flattening and swung forwards, especially in the 
midline, and were often lost. In the process, the 
associated AMV was also detached and, being much 
smaller, is easily overlooked. 

Every stage of this process is illustrated by 
Cowralepis. Many specimens show the AMV and IL 
plates still in situ and in association (Figs 1B, 4B, 5B, 
D, 14A, 15C). In most of the specimens where the 
IL plates have sprung forward, the AMV is missing. 
Other specimens show the AMV almost in place but 
slightly detached (Figs 8A, B, E) and reveal its size, 
shape and overlap areas. The AMV was three times 
wider than long. Its anterior margin was slightly 
concave and the posterior margin was bluntly pointed 
medially, matching the rounded corners of the AVL 
plates against which it fitted. In AMF 127152 (Fig. 8E) 


ay 


A NEW GENUS OF DEVONIAN PLACODERMS 


the AMV displays two pairs of overlap areas; laterally 
to accommodate the sharp pointed mesial ends of the 
IL plates and posteriorly for the anteromesial corners 
of the AVL plates. 

Rounded anteromesial corners are well displayed 
in most AVL plates of Austrophyllolepis (Long 1984, 
figs 4-7, 11-13) and are also present in AVL plates of 
Placolepis budawangensis Ritchie (1984, figs 10-12). 
On the new evidence from Cowralepis mclachlani, 
the writer would now have no hesitation in restoring 
an AMV plate in Placolepis even though none has yet 
been found. On balance it is suggested that most, if 
not all, phyllolepids probably retained an AMV plate 
in their ventral trunk armour. 

The first evidence for a posterior median ventral 
plate (PMV) came from a specimen of Phyllolepis 
woodwardi from Dura Den, Scotland (Stensié 1939, 
text-fig. 2) in which “fragments of bone, not nearer 
determinable” were noted and labelled (px) in the 
mid-ventral line at the junction of the AVL and PVL 
plates. Long (1984, 267) suggested that these may be 
“_. fragments of the axial skeleton”, exposed through 
a gap in the ventral trunk shield. He figured a rather 
large, narrow PMV plate in several specimens of 
Austrophyllolepis (1984, figs 7, 9B, 11D, 13, 18C) 
plus an example of “. . an abnormal development” in 
a juvenile specimen of Austrophyllolepis displaying 
two PMV plates (figs 4C, 6). 

The Cowralepis material indicates that this 
condition may not, in fact, be so abnormal in 
phyllolepids. In the holotype of Cowralepis 
mclachlani (AMF103767B, Fig. 1B) and in several 
other specimens (AMF127162, Fig. 8G) a single 
slender PMV is visible. More commonly two PMV 
plates are clearly present, the anterior PMV being 
normally longer than the posterior one (Figs 5D, 
16B). But other specimens from Merriganowry with 
apparently intact ventral shield show no trace of a 
PMV at the intersection of the AVL and PVL plates 
(Figs 4B, 6B, 8F). 

The new evidence from Cowralepis suggests that 
the number of PMV plates in phyllolepids was not 
fixed but could vary within a genus and, possibly, 
even within a species, from none to at least two. 


Sensory canals on the trunk shield 

The lateral line system continued posteriorly 
from the head shield over the narrow ADL plate on the 
dorsal trunk shield. In Placolepis, Austrophyllolepis 
and Phyllolepis an anterior mesial branch of this 
crossed onto the corner of the MD plate as the dorsal 
sensory canal groove (Long 1984, figs 7, 13; Ritchie 
1984 fig. 2A, C) but no trace of this canal groove has 
been detected in Cowralepis. 


228 


In the ventral trunk shield, the only trace of sensory 
canal grooves consists of a short shallow stretch of the 
ventral pit line (vpl) parallel to the posterior margin 
of the AVL plate in a few specimens of Cowralepis 
in which the ornament is well developed (Figs 5B, 
6D, 8F, vpl). The ventral pit line is also present in 
Placolepis, Austrophyllolepis and Phyllolepis. 


Tooth plates and parasphenoid 

Although lying deep inside the head, the tooth 
plates and parasphenoid originated as part of the 
dermal skeleton. Amongst the placoderms, only the 
arthrodires and the acanthothoracids had two pairs 
of gnathal plates in their upper jaw, the anterior and 
posterior superognathals (Sgn) that occluded against 
a single inferognathal plate (Ign) in the lower jaw. 
Where two pairs of superognathals are present the 
posterior pair are normally the larger. 

The only previous evidence for tooth plates 
in phyllolepids comes from Austrophyllolepis, in 
which large, broad superognathals and narrower 
inferognathals were preserved in several specimens 
(Long 1984, figs 5, 6, 12C, 16, 17, 18B), although 
usually slightly displaced. 

The upper and lower tooth plates are preserved 
in many specimens of Cowralepis, both in situ and in 
association (Figs 9, 10, 11, 14, 19) and have also been 
found as isolated elements. They confirm the presence 
of two pairs of supragnathal elements in Cowralepis: 
a large anterior supragnathal, corresponding closely 
to that of Austrophyllolepis, and a very small posterior 
supragnathal, nestling against its posterolateral 
margin. The small posterior element has only been 
identified in about a dozen of the many hundreds of 
phyllolepid specimens from Merriganowry, and it is 
not surprising, given its size, that it was not located 
in the much smaller sample of phyllolepid specimens 
available from Mt Howitt. 

The anterior superognathals (Figs 4B, 5B, 6B, 
D, 9A, E-G, 10A-C, G, H, 15C, D) were long, broad 
and flat, subtriangular in shape with a concave mesial 
margin. In the largest individuals of Cowralepis 
the anterior Sgn plates reached over 3 cm in length 
and 1 cm wide. They were broadest posteriorly and 
tapered anteromesially to a narrow apex near the 
anterior margin of the head shield where they met 
in the midline under the preorbital plates at an angle 
of 90-100° (Figs 14A, 19). The occlusal surface was 
densely covered with rows of small, sharp conical 
teeth radiating from the ossification centre of the plate 
and increasing in size towards the margins, but with a 
very sharp line of demarcation running up the midline 
towards the anterior apex (Fig. 10G, H); a similar 
linear feature can be seen in the superognathals of 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Austrophyllolepis (Long 1984, figs 16, 18B). 

The anterior Sgn in Cowralepis closely resembles 
the example depicted in Austrophyllolepis ritchiei 
(Long 1984, fig. 17), except that the teeth in the latter 
did not extend to the margins, leaving an edentulous 
rim all round the plate. The shape of this plate does 
not match that in the accompanying reconstruction 
(Long 1984; fig. 14B) where its posterior margin was 
shown as sharply truncated, straight and attached to a 
rounded plate. 

In sharp contrast to the anterior plate, the 
posterior superognathal was minute, only 2-4 mm in 
diameter (Figs 9E-G, 10A, B, C). It was semicircular, 
with one straight or slightly concave margin. Because 
of its size it is often obscured by larger ventral plates 
in the fossil specimens. The original relationships 
of the three gnathal plates are best demonstrated in 
AMF96783 (Fig. 9E-G) where the inferognathal is 
in occlusion against the anterior superognathal and 
the curved margin of the posterior Sgn neatly fits 
into the posterior margin of the larger superognathal; 
its straight margin is therefore the posterior face. In 
~ three other examples (Fig. 10A-C) the posterior Sgn 
is completely exposed but slightly displaced. 

Given the small size of this element, and the fact 
that in placoderms with two pairs of superognathals 
(arthrodires and acanthothoracids) the posterior 
element is normally larger than the anterior, Gavin 
Young (pers. comm.) has suggested another 
interpretation for this small plate in Cowralepis. 
Young et al. (2001, 673, fig. SA, sc) noted a third 
small denticulate plate, also ca. 2mm across, at the 
posterior end of the posterior superognathal plate in a 
buchanosteid arthrodire from Taemas, NSW 

The inferognathal (Ign) comprises two laminae 
meeting at right angles; a tooth-covered dorsal 
(horizontal) occlusal lamina and a smooth, vertical 
anterolateral lamina. In most Cowralepis specimens 
the Ign is preserved in occlusion against the anterior 
superognathal and shows only the smooth groove that 
housed Meckel’s cartilage (Figs 9A, E-G, 10A, E, 
11). The Ign was very narrow anteriorly, broadening 
out only in the posterior half; its occlusal area (Fig, 
10D) was much smaller and narrower than that of 
the anterior Sgn and a row of larger teeth extended 
along the outer ridge to the anterior apex (not visible 
here because of the rotation) as in Austrophyllolepis 
(Long 1984, figs 16, 17). The posterior margin of the 
inferognathal was smooth, tooth-free and serrated or 
sharply pointed (Fig. 10D, E). 


Parasphenoid 


In many placoderms the palate bore a small 
bone, the parasphenoid, with a median foramen 


Proc. Linn. Soc. N.S.W., 126, 2005 


for the buccohypophysial canal; whether this is the 
homologue of the parasphenoid in osteichthyan fishes 
is still uncertain. 

The parasphenoid is well preserved in situ in many 
specimens of Cowralepis and was apparently firmly 
attached to the palatal surface of the endocranium. It 
is best displayed in specimens where the branchial 
ossifications are either not preserved or have been lost 
(Figs 5B, 6D, 8A, 9A-F, 10G, H, 15C). 

Based on the new evidence from Cowralepis, 
the reconstruction of the subcranial ossifications of 
Austrophyllolepis (Long 1984, fig, 14A, B) needs 
to be modified. The parasphenoid is shown in the 
correct position but the superognathals lay farther 
forward, almost meeting under the anterior margin. 
The otoliths (which would have been covered by the 
parasphenoid in the position depicted) were situated 
more posterolaterally, lying between the parasphenoid 
and the paranuchal. 

The Cowralepis parasphenoid was very large 
(up to 1/3 the length of the head shield), flat and thin 
and was centrally situated under the nuchal plate. 
Its overall shape is best displayed in AMF96781 
(Fig. 9B), and it consisted of a slightly raised, 
lightly tuberculated central area which was widest 
posteriorly and tapered anteriorly. A single median 
buccohyphophysial foramen (f.bhy) is situated level 
with the widest part of the plate. 

The central raised area was surrounded by a 
wide, thin radiating flange with scalloped edges. 
It was deeply indented laterally (l.ind) opposite 
the buccohypophysial foramen as far as the central 
raised area and continues as a shallow tapering 
groove onto the central raised area. The Cowralepis 
parasphenoid does not differ significantly from that in 
Austrophyllolepis (Long 1984, fig. 17), known only 
from a few specimens. 

In a review of parasphenoids in the Placoderm1, 
Dennis-Bryan (1995) noted that parasphenoids had 
been recorded from 36 placoderm genera, of which 
all but three were arthrodires; one of the three 
exceptions was Austrophyllolepis (Long 1984). It was 
observed that placoderm parasphenoids appeared to 
be species specific, but were of limited use at higher 
taxonomic levels. In overall shape the Cowralepis 
parasphenoid most closely resembles those of the 
Gogo brachythoracid arthrodires, Goujetosteus and 
Eastmanosteus (Dennis-Bryan 1995, fig. 2 B, D), but 
these were much more robust structures and easily 
detached from the endocranium. 

Placoderm parasphenoids were divided into two 
main types - ‘primitive’ and ‘advanced’. ‘Primitive’ 
parasphenoids were flat, lacked lateral notches or 
grooves, had a central tuberculated area and a large 


229 


A NEW GENUS OF DEVONIAN PLACODERMS 


median buccohypophysial foramen, single or paired. 
‘Advanced’ parasphenoids were thicker, with lateral 
notches and grooves and buccal foramina reduced; 
tubercles were reduced or absent, and a _ well- 
developed median ventral crest was developed. 

In a prescient comment Dennis-Bryan (1995, 
136) cited two notable exceptions, Pholidosteus and 
Austrophyllolepis; “The former is considered to be an 
advanced arthrodire but has the so-called primitive 
parasphenoid, and in the latter case the reverse is true. 
It is perhaps possible that the dorsoventrally flattened 
phyllolepids are more advanced among placoderms 
than was first thought.” 


Visceral skeleton 

In many placoderms both the dermal skeleton 
and endocranium are known from a wealth of 
well-preserved material. The visceral skeleton of 
placoderms, however, is rarely preserved and is 
poorly known (Denison, 1978, 5-7; Janvier, 1996, 
153). In most cases it was probably unossified and, 
even where perichondral ossification was present, it 
was often delicate and easily destroyed by taphonomic 
processes. 

In all gnathostomes the principal paired visceral 
structures are the mandibular and hyoid arches 
and several branchial arches. The mandibular arch 
comprises the palatoquadrate and mandibular 
(Meckel’s) cartilages, both of which may be 
perichondrally ossified and, in the hyoid arch, the main 
elements (when preserved) are the hyomandibula, 
ceratohyal and basihyal elements. 

Prior to the discovery of Cowralepis the only 
direct evidence on the visceral skeleton in phyllolepids 
came from several Austrophyllolepis specimens in 
which various paired and unpaired ossifications were 
preserved lying against the ventral surface of the head 
shield, but displaced (Long 1984, figs 6, 16, 17, 18B, 
20). Long based his interpretation (fig. 17) on the 
only example in which these ossifications appeared 
to be in association and reconstructed them as three 
pairs of bones plus a large median parasphenoid and 
a pair of otoliths (Long 1984, fig. 14A). 

Long interpreted the three pairs of elements 
as the superognathals, metapterygoids (the middle 
division of the palatoquadrate) and the quadrates, and 
he reconstructed them as connected in series (Long 
1984, fig. 14B). He also suggested (Long 1984, 281) 
that this series might only have developed at maturity. 
Many of the articulated Cowralepis specimens, 
juvenile and adult, show some or all of the same 
visceral ossifications (and many others) in situ and 
in association and reveal that Long’s interpretation is 
invalid. 


230 


Immediately posterior to the gnathal plates, and 
in direct line with the inferognathal, lay a robust bone 
with a narrow posterior stem and a wide anterior 
margin. This bone was strongly perichondrally 
ossified but open at both ends (Figs 10F, 11, 12C, 
D, F). A prominent ventral ridge ran obliquely 
from its anteromesial to its posterolateral margin. 
Anterolaterally it flared out into a wide triangular 
flange, with a rounded anterior margin. Posteriorly, 
on the mesial side of the ventral ridge, there was a 
deep smooth-sided pit (Figs 10F, 12F). 

Long interpreted this plate as the quadrate and 
reconstructed it as attached anteromesially to a large, 
flat plate which he labelled the metapterygoid (Long 
1984, figs 14B, 17). 

Cowralepis suggests a different interpretation. 
The same two plates are preserved in a large number of 
specimens, in place and closely associated (Figs 9A, 
E, F, 11, 14A). The robust bone (Long’s ‘quadrate’) 
was not originally attached distally to the rounded 
plate (Long’s ‘metapterygoid’) but lay lateral to it 
(Fig. 15C, D). Although they have sometimes drifted 
slightly apart post mortem they appear to have been 
closely associated throughout growth. 

The anterior margins of both the rounded plate, 
and the robust plate lateral to it, covered (ie. lay 
below) the posterior ends of both the superognathals 
and the inferognathal. They therefore lay ventral to 
Meckel’s cartilage. This would suggest that the latter 
cannot be the quadrate, as originally proposed, and 
it is interpreted here as the articular, the posterior 
ossification of the lower jaw. 

Miles and Dennis (1979, fig.11B-E) illustrated a 
small perichondrally ossified articular plate attached 
to the rear of the inferognathal in the brachythoracid 
Harrytoombsia from Gogo, Australia. More recently 
Johanson (2003, fig. 4) has figured large, well-ossified 
articular bones attached to the inferognathal in other 
Gogo brachythoracids (Eastmanosteus, Gogopiscis 
and Jncisoscutum). 


Metapterygoid or ceratohyal? 
If the robust bone is the articular this raises the 


question — what was the rounded plate lying along its 
mesial margin? This was very thin and flatand normally 
seen in ventral view (Figs 9A, F, 11, 12C, 14A, 18F, 
19, Chy) but occasionally in dorsal view (Fig. 14B, 
Chy). It consisted of two layers of perichondral bone 
separated by a thin sheet of cartilage and continued to 
grow throughout life, as indicated by the prominent 
semicircular growth lines on the flat plate, which 
radiate from a centre of ossification midway along 
the straight margin. 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Since it lay ventral to the inferognathal (and to 
Meckel’s cartilage) it cannot be the metapterygoid 
(a palatoquadrate ossification) as suggested by Long 
(1984) and it is interpreted here as the ceratohyal 
(Fig. 15C, Chy), a ventral element of the hyoid arch. 

In smaller Cowralepis individuals (Figs 5B, 6D, 
9A, E, F) the ceratohyal was slightly longer than the 
articular but fell a long way short of the midline. In 
larger individuals (Figs 11A, B, 14A) the ceratohyal 
was almost twice as long as the articular and had 
expanded anteromesially to make contact with, and fit 
neatly against, the concave lateral margins of a large 
robust median plate, discussed below. 


Basihyal 
Almost every Cowralepis specimen in which 


the gnathal, articular and ceratohyal elements are 
preserved also features a large, robust, median 
perichondral ossification lying anteromesial to the 
ceratohyals and a short distance posterior to the 
symphysis of the upper and lower tooth plates (Figs 
1, 4B, 9A, B, E, F, 11, 14A, 18F). The distinctive 
features of this robust plate are illustrated here by 
stereopair images (Fig. 12A, B). 

The plate is about three times as long as wide, 
with a flat basal platform, rounded anteriorly and 
posteriorly and with concave lateral margins. A 
prominent, narrow ventral keel runs most of the 
length of the plate. The keel is shallowest anteriorly 
where it divides over the basal platform into numerous 
radiating buttresses and rises gradually towards the 
posterior margin where it terminates in a deep pit that 
probably housed a ligament operating to lower the 
jaw. Midway along the basal platform, on either side 
of the keel, there is a small round pit with a raised 
rim, most clearly seen in AMF96783 (Fig. 9E,F; cf. 
also 12A). 

In smaller individuals the ceratohyals are well- 
separated from this median plate (Figs 6D, 9A, F) 
but, during growth, they gradually increase in size 
until they meet the lateral margins of the median 
plate, extending almost to its anterior margin (Figs 
11, 14A). 

In virtually every specimen of Cowralepis in 
which it is preserved in situ this median bone lies 
posterior to, and is quite separate from, both the 
gnathal plates and the mandibular arch. The close 
lateral association with the ceratohyals confirms 
that it also lies ventral to the inferognathal and it is 
interpreted here as the anterior ossification of the 
hyoid arch, a median basihyal. 

Long (1984, fig. 17) figured a rather shapeless, 
unnamed median bone in Austrophyllepis overlapping 
the anterior margin of the parasphenoid but this was not 


Proc. Linn. Soc. N.S.W., 126, 2005 


identified or included in the reconstruction (fig. 14B). 
Given its close association with the parasphenoid 
this plate may be the homologue of the basihyal in 
Cowralepis (Figs 9A, B, 15C), but this can only be 
settled by the recovery of more phyllolepid material 
from Mt Howitt. 


Basibranchials or hypobranchials? 
In several medium-sized to large Cowralepis 


specimens in which the visceral skeleton is preserved, 
the large median plate interpreted above as the 
basihyal is closely followed posteriorly by up to four 
pairs of flat, rounded to irregular bony plates that 
meet in the midline (Figs 1B, 4B, 6D, 11, 14A, 18F, 
19). The anterior pair sometimes even wrap around 
the rounded posterior margin of the basihyal (Fig. 
14A). If the above interpretation of the mandibular 
and hyoid arch elements is correct, these four pairs of 
plates must be associated with the third to sixth visceral 
arches. This is very similar to Orvig’s reconstruction 
of the visceral skeleton in the ptyctodont, Crenurella 
(Miles 1971, fig. 8.13). 

Ossified basal visceral elements have only been 
recorded from Pseudopetalichthys (Stensid 1969, 
Figs 168A, B), Zapinosteus and now in Cowralepis. 
Stensi6’s reconstruction of the branchial skeleton 
in Tapinosteus (1963, figs 83A, B; 1969, fig. 166A, 
B), derived from serially sectioning the holotype, 
shows a median basihyal followed by three pairs 
of hypobranchials, (each pair subdivided with 
dotted lines) and depicted as separated by unpaired 
basibranchials. The reconstruction was based on a 
chondrichthyan model (e.g. Chimaera monstrosa; 
Stensi6 1969, fig. 169), consistent with Stensi6’s 
proposed grouping of chondrichthyans, placoderms 
and acanthodians into his Elasmobranchiomorphi, an 
association no longer considered to be valid. 

This raises the question — are the four pairs of 
visceral elements in Cowralepis the homologues of 
the basibranchials (which are median and unpaired 
in chondrichthyans) or the hypobranchials (which 
are paired)? Goodrich (1930) and others have 
suggested, from embryological evidence, that the 
basal visceral elements in gnathostomes may have 
been primitively paired. There is no trace of any 
median elements between the pairs of basal branchial 
elements in Cowralepis that met in the midline and 
the paired elements present here are interpreted as 
hypobranchials (Figs 6B, 11B, 14A, B, 15D, Hbr). 

The whole complex of basal visceral ossifications 
under the head of Cowralepis (large disc-shaped 
ceratohyals, a robust basihyal and several pairs of 
hypobranchials) would appear to have provided a 
firm floor to the buccal cavity, and the deep pit on the 


Pen 


A NEW GENUS OF DEVONIAN PLACODERMS 


posterior face of the basihyal (Fig. 12B) may have 
housed a strong ligament that depressed the lower 
jaw. 

There are many more specimens of Cowralepis 
displaying the visceral skeleton than those figured 
here and a fuller account, tracing its ontogenetic 
development, will be presented elsewhere. 


‘Suborbital’ 

Long (1984, figs 4A, 5, 14A, 16, 19) figured a small 
curved bone in the cheek area of Austrophyllolepis 
but found it difficult to homologise this with other 
placoderms because it lacked ornament and was 
folded into a “. . double lamina with a large valley in 
between” (Long 1984, 283). The plate was interpreted 
as a greatly reduced and modified suborbital plate 
(SO) from its situation below, or lateral to, the PtO 
plate, just where the infraorbital sensory line of 
most placoderms divided to send a supraoral line 
ventrally. 

A similar small curved bone is present in several 
specimens of Cowralepis (Figs 5B, 6D, 14B, 15C, 
hae), but is best preserved in AMF127156 (Fig. 11A, 
B), details of which are shown as stereopairs (Fig. 12C, 
D, hae). In this rather large individual of Cowralepis 
the bone is 6 mm long and 3 mm wide, bean-shaped 
and strongly convex. Its surface is very smooth and 
deeply creased longitudinally by a curving groove 
close to, and following, the concave margin. 

In most of the Cowralepis specimens in which 
this element is present it does not lie near the lateral 
margin of the PtO (as depicted in Austrophyllolepis) 
but farther in, under the inner margin of the PtO plate 
and almost touching the posterior end of the bone 
interpreted above as the articular. The new evidence 
from Cowralepis does not support the earlier 
identification of this bone as a suborbital plate. 

The very strongly convex shape, complete lack 
of ornament and its position rule out a dermal origin. 
Gavin Young (pers.comm.) has suggested it may be a 
small perichondral ossification of the visceral skeleton 
and, from its position near the mandibular joint, 
possibly a hyoid arch element, and this interpretation 
is provisionally accepted here (Fig. 15C, hae). 


Endocranium 

In many Cowralepis specimens two very robust, 
median ossifications lying entirely within the trunk 
armour are often clearly visible through the dorsal 
and ventral armour, which has been impressed onto 
them (Figs 4B, 6B). They are most clearly seen in 
situ in a few specimens where the ventral plates have 
either been lost, or are preserved as isolated elements 
(Fig. 13). 


232 


The shorter, wider, structure is the occipital 
ossification, the longer, narrower ossification is the 
synarcual, the anterior section of the axial skeleton 
(discussed below) and the junction between them is 
the craniothoracic articulation. 

The Occipital ossification lies anterior to the neck 
joint and represents the ossified posterior (occipital) 
part of the endocranium (Fig. 13A, B, D, E; Occ oss). 
It does not include the otic region and extends from 
the anterior margin of the ventral trunk shield to the 
posterior margin of the head shield. 

There is a discrepancy in shape between the 
occipital ossification in small individuals (Fig. 13D, 
E) and in larger specimens (Fig. 13A, B). In juveniles 
it is broad and flat, narrowest posteriorly with concave 
sides and a straight posterior margin (the neck joint). 
Anteriorly it flares into two round, flat processes 
meeting mesially, with a median notch on the anterior 
margin. 

In larger individuals (Figs 4B, 6B, 13A, B) the 
occipital ossification is much longer, like two spools 
attached side by side. The lateral margins are still 
concave and the anterior margin is wider than the 
posterior margin, but the central shaft of each half 
of the ossification is smoothly rounded with one or 
more rows of regularly spaced deep pits, separated by 
narrow ridges, developed around the circumference 
at either end. 

While this may indicate the presence of two 
species, differential development of the occipital 
ossification between small and large individuals, as 
noted here, has been described in pyctodontids by 
Miles and Young (1977, 166) 


Otoliths and paraotic plates 

Almost all articulated specimens of Cowralepis 
display two solid, bean-like structures lying against 
the ventral surface of the nuchal plate midway between 
the midline and the anterior process of the paranuchal » 
plate (PNu). Long (1984, figs 6, 9, 11, 12, 17, 18C) 
identified similar structures in Austrophyllolepis as 
otoliths and the Cowralepis material supports this. 

The otoliths were dense calcareous bodies, 
oriented anterolaterally (Figs 1B, 4B, 5B, D, 8A, 9A, 
11A, B, 13D, 14B). They were very solid bodies, not 
hollow, as demonstrated by the fact that the nuchal 
plate has often been moulded over them, and they 
were proportionally larger in smaller individuals than 
in more mature ones. One specimen of an isolated 
otolith displays its aggregate crystalline nature and 
surface structure in fine detail (Fig. 12E) 

In life, the otoliths were internal structures located 
within the fluid-filled sacculus of the left and right 
otic labyrinth and were deeply embedded within the 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


endocranium, the anterior part of which, in Cowralepis, 
was apparently cartilaginous. As noted by Long (1984, 
281) the position of the otoliths in Austrophyllolepis 
(and now also in Cowralepis) corresponds closely 
to the site of the saccular cavities in Buchanosteus 
(Young 1979), Kujdanowiaspis (Stensi6 1963) and 
other euarthrodires. Calcified otoliths have not been 
recorded from other placoderms, but their presence 
in both Cowralepis and Austrophyllollepis suggests 
that they were probably present in all phyllolepids. 
Otoliths were also present in the extinct acanthodians 
and are found in extant osteichthyans, but in both of 
these groups there are three separate otoliths in each 
otic capsule, not one. 

The otoliths in Cowralepis display an interesting 
feature not observed in Austrophyllolepis. In smaller 
individuals the posterior half of each otolith is often 
overlain or, more correctly, underlain by a subcircular 
to irregular bony disc with concentric growth rings 
(Fig. 8B). In slightly larger individuals this disc had 
grown to cover two thirds of the otolith (Fig. 11A, 
B), and in the largest individuals it was large enough 
- to completely cover the otolith, onto which it was 
moulded from below (Fig. 14A). Because these sub- 
otic plates appear to be underlain ventrally by the 
hypobranchial ossifications, and therefore lay dorsal 
to them, it is suggested that these plates were attached 
to the palate immediately under each otic capsule as 
paraotic plates (Fig. 15D, p.pao). 

The nearest equivalent in placoderms appears 
to be a large pair of denticle-covered paraotic plates 
developed on the palate in Nefudina qalibahensis, 
a thenanid placoderm from the Early Devonian of 
Saudi Arabia (Lelievre et al. 1995, 111, pl.1, fig.1, 
p.pao). The paraotics lay immediately under the otic 
capsules and were attached to the palatal surface of 
a well-ossified endocranium, just posterior to a large 
denticulate parasphenoid. Their robust nature and 
denticulate surface suggests they may have been 
used in food processing. In contrast, the paraotics 
in Cowralepis were thin, smooth-surfaced discs, 
lacking ornament and may have served to protect the 
undersurface of the otic capsules. Gavin Young (pers. 
comm.) has suggested that they may represent areas 
of ossification on the floor of the endocranium. 


Axial skeleton and fins 

In many placoderms the anterior vertebral 
elements, housed within the trunk shield, were 
fused into a synarcual ossification on which rested 
the median dorsal plate. Stensi6 reconstructed the 
posterior of the endocranium and a long narrow 
synarcual in Paraleiosteus (1969, fig. 34) in lateral 
view, and that of Cowralepis must have looked rather 


Proc. Linn. Soc. N.S.W., 126, 2005 


similar. 

The synarcual in Cowralepis was long and narrow 
and its overall shape is displayed in two individuals 
seen in ventral view, AMF96762 (Figs 4B, 6B) 
where it is overlain by the ventral trunk plates and 
AMF96751 (Fig. 13A) where the ventral plates have 
been lost, fully exposing both the synarcual and 
occipital ossifications. AMF96753 (Fig. 13C), an 
isolated synarcual, preserved in counterpart, displays 
both its dorsal (left) and ventral (right) surfaces 
revealing that, like the occipital ossification, 1t was 
clearly divided into left and right sections. The ventral 
surface displays a deep longitudinal median groove 
that widens anteriorly into an elongate diamond- 
shaped foramen. 

Long’s reconstruction of the trunk and fins in 
Austrophyllolepis (1984 Fig. 23), based on the few 
incomplete specimens available at the time (Long 
1984, figs 21, 22), can be revised from Cowralepis. 
There is no evidence for the presence of a dorsal fin in 
Cowralepis (or in Austrophyllolepis), the pelvic fins 
were much smaller than shown and were tucked under 
the body close to the ventral shield, and the caudal 
fin was epicercal (= heterocercal) with the notochord/ 
vertebral column supporting the upper lobe. 

The vertebral column is fully known in only 
a few placoderm genera, the best examples of 
which are Ctenurella (Stensid 1969, fig. 178) and 
Coccosteus (Miles and Westoll 1968, text fig. 48) in 
both of which a persistent unrestricted notochord was 
enclosed dorsally and ventrally by neural and haemal 
arches bifurcating around it. 

Many specimens of Cowralepis, small and large, 
have the vertebral column and tail fin attached, often 
almost complete and preserved in part and counterpart 
in fine detail (Figs 1, 4, 16-18). Even the smallest 
specimens display the beginnings of a well-ossified 
backbone (Figs 2D, 18A). The complete exposure 
of the vertebral elements indicates that Cowralepis 
lacked any scale-covering, a feature also noted in 
Austrophyllolepis by Long (1984, 297). 

In Cowralepis, the tail was relatively long and 
powerful, and the relative length of the dermal shield 
to the body and tail change during growth. In small to 
medium-sized individuals the body and tail forms 65- 
70% of the total length; in the largest specimens this 
drops to just over 50% of the total length (Fig. 16 B, 
©): 

Cowralepis had at least 60 vertebrae posterior 
to the trunk shield and pelvic fin. Anteriorly these 
consist only of dorsal (neural) and ventral (haemal) 
arcualia but posteriorly they deepen to support the 
caudal fin and were supplemented ventrally by an 
additional row of sub-haemal elements (Fig. 18C, 


233 


A NEW GENUS OF DEVONIAN PLACODERMS 


sub-h.sp) extending to the tip of the sharply pointed 
tail. 

The exquisite preservation of some Cowralepis 
specimens (Fig. 17A-D) allows reconstruction of the 
vertebral elements (Fig. 18C) and comparison with 
those of Coccosteus and with Incisoscutum ritchiei in 
which uncrushed vertebral elements were recovered 
by acid preparation (Dennis and Miles, 1981, 248-50, 
figs 21, 22). 

In Jncisoscutum the neural arch (housing the nerve 
cord) was separated from an overlying triangular 
opening (housing the dorsal ligament) by horizontal 
bony flanges that met mesially. In Coccosteus a 
similar transverse flange carried an anterior median 
bony projection that Miles and Westoll (1968, text. 
fig. 46) called the zygopophysis; by contrast, in 
Incisoscutum, the same flange carried a pair of well- 
developed zygopophyses. 

The condition in Cowralepis was simpler; the 
neural arch was an open V-shaped incision with no 
transverse flange separating the nerve cord and dorsal 
ligament, and with a single prominent zygopophysis 
at the apex of the neural arch (Fig. 18C, zyg). The 
dorsal spines were long, narrow and angled sharply 
backwards. 

Where the neural arch sat on the notochord it 
broadened out into large curved basal pads, as in 
Incisoscutum (Dennis and Miles 1981, fig 22). In the 
haemal arches the paired dorsal contact processes 
were oval with deep pits (Fig. 17D). The haemal 
spines were much longer than the dorsal spines and 
strongly sinuous in shape, especially in the caudal 
fin. An unusual feature in Cowralepis, not recorded 
in any other placoderm, was the development of an 
extra row of sub-haemal radial elements, considerably 
deepening the hypochordal lobe of the fin and 
increasing the propulsive force of the tail. 


Pectoral fin 

None of the remarkably complete Cowralepis 
specimens from Merriganowry display any trace 
of the pectoral fin which clearly lacked any scale 
covering, nor is there any trace of the scapulocoracoid 
or of fin basals, suggesting that these were probably 
cartilaginous and not perichondrally ossified. The size 
and shape of the pectoral fin in Cowralepis remains 
conjectural (Fig. 20A). 


Pelvic girdle and fin 
The pelvic girdle and fin skeleton is preserved in 


situ in many specimens of Cowralepis, of all sizes, 
including the holotype, AMF103767 (Figs 1A-D, 4B, 
6B, 7H, I, 16A-C, pel.b, pel.d). It lay immediately 
behind, or slightly overlapped, the posterior margin of 


234 


the ventral shield but it has also been found detached 
and isolated (Fig. 7D, E). 

The pelvic fin was first identified in 
Austrophyllolepis, based on two specimens (only 
one of which was figured) where it consisted of two 
perichondral ossifications, a larger anterior element, 
and a shorter, narrower, distal element (Long 1984, 
figs 21, 22, pel.b, pro). 

Based on the many well-preserved specimens of 
the same element in Cowralepis, the anterior element 
(basal pelvic plate) preserved in Austrophyllolepis is 
probably incomplete. In Cowralepis the basal pelvic 
plate was longer and narrower, with slightly concave 
margins (Fig. 7D, E, H, pel.b). The left and right 
pelvic plates met in the mid-ventral line and diverged 
posterolaterally at about 45° to the body axis. 

Long noted that part of the lateral margin of the 
basal plate appeared to bear “short grooves denoting 
serial divisions for articulation of cartilaginous pelvic 
fin ray elements” (1984, fig. 20, art; 285). No evidence 
of such grooves has been detected in Cowralepis and 
I suspect that this feature in Austrophyllolepis was 
probably an artefact of preservation. 

The shorter distal element (Figs 4B, 6B, D, 7D, E, 
H, I, pel.d) was interpreted by Long as a propterygium 
(1984, figs 21, 22, pro), the anterior of three principal 
cartilages in the paired fins of elasmobranch fishes, 
and thus a pre-axial element. In contrast, the smaller 
bone in the Cowralepis pelvic fin is normally attached 
distally to the basal plate, and in line with it, unlike 
in Austrophyllolepis where it appears to have been 
displaced. This suggests that it was an axial element. 

Janvier (1996, 244) noted that although one 
group of placoderms (ptyctodonts) may have 
possessed claspers on the pelvic fin, “. .the problem 
with pelvic claspers is that they are apparently 
lacking in all other placoderms, although very few 
pelvic fins have actually been preserved. Long (1984) 
has described a peculiar club-shaped endoskeletal 
(probably metapterygial) element in the pelvic fin of 
the phyllolepid Austrophyllolepis; this may suggest 
the presence of a pterygopodium that supported the 
clasper.” 

The distal element of the pelvic fin in Cowralepis 
(Figs 1C, D; 6B, D; 7D, E, H, I; 16A, pel.d) is 
interpreted as the homologue of the metapterygium in 
the elasmobranch fin. 


Sexual dimorphism 
Long (1984, 275, 286) discussed the possibility 


that the ‘propterygial’ element in Austrophyllolepis 
may have been a clasping organ, rather than just an 
extension of the pelvic fin. Zangerl (1981) pointed 
out that “. . the pelvic girdle of Austrophyllolepis 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


showed some resemblance to the male clasping 
organs of primitive chondrichthyans, particularly 
Cobelodus”. Judgement was suspended because of 
insufficient material; the only two specimens of the 
Austrophyllolepis pelvic skeleton were similar, with 
no evidence of dimorphism to indicate male and 
female versions. 

This can now be re-examined using the numerous 
examples of pelvic fins in Cowralepis. Because of the 
vagaries of preservation, and its relatively small size, 
the pelvic fin is not always visible even in individuals 
in which the body and tail is well preserved. However, 
where it is preserved, the pelvic fin skeleton is always 
approximately the same length in similar-sized 
individuals, with no evidence for larger and smaller 


variants, or for the presence of claspers, that might | 


indicate sexual dimorphism. 

Sexual dimorphism could still be present in the 
Merriganowry phyllollepid material, but it 1s difficult 
to see how evidence for it might be extracted from 
fossil specimens that have undergone strong tectonic 
deformation. 


The cannibal from Cowra 

If further confirmation was required concerning 
the topographic relationships of all the gnathal and 
visceral skeletal elements under the Cowralepis head 
shield it is provided by an unusual example of one 
individual that literally bit off more than it could 
chew! 

AMF9001 1a, b is a moderately large Cowralepis 
specimen in which the vertebral column of a second, 
smaller Cowralepis is seen disappearing under the 
anterior margin of the head shield in the midline 
(Fig. 18E). That it is not just a case of accidental 
superposition of one fish on another is revealed by 
the counterpart showing the ventral surface (Fig. 
18F) and by combining information from the part and 
counterpart (Fig. 19). In the larger individual most of 
the gnathal and branchial skeletal elements are present 
in situ and they even include the small left posterior 
superognathal (Sgn post.) exposed by separation of 
the articular from the ceratohyal. 

The smaller fish skeleton is shown here in black 
(Fig. 19). The vertebrae continue posteriorly under 
the ventral margin of the head shield and over the 
right superognathal. They pass between the basihyal 
and the right ceratohyal into the buccal cavity where 
they terminate against the posterior margin of the 
ventral trunk plates of the smaller individual (PVL + 
AVL plates; shown in black), still in association. The 
size of the prey demonstrates that the mouth opening 
in Cowralepis was at least as wide as the anterior 
margin of the head shield. 


Proc. Linn. Soc. N.S.W., 126, 2005 


The neural arches and their dorsal spines lie 
against the occlusal surface of the right superognathal 
and are overlain by the posterior part of the right 
inferognathal. Behind this the dorsal spines disappear 
under the anteromesial corner of the right ceratohyal 
that has been moulded over them. The vertebrae and 
associated trunk plates are partly overlain by several 
hypobranchial plates and the trunk plates disappear 
under the leading edge of the host’s ventral shield 
(AVL plates); the left and right interolateral plates 
have swung forwards and the AMV is missing, as 
commonly occurs in phyllolepids. 

This unique specimen confirms that the skeletal 
elements identified as  basihyal, ceratohyals, 
articulars and hypobranchials, together with the 
inferognathals, all lay ventral to the palatoquadrate 
and superognathals. 


An egg sac? 

One of the most intriguing finds from 
Merriganowry, AMF127151, preserved in counterpart 
(but only as a natural mould) appears to be an egg 
sac (Fig. 18D). It is 3 cm long, 1 cm wide and 
contains perhaps 200 tightly packed, uniform, oval 
bodies, each 3-4 mm long and oriented parallel to, or 
slightly oblique to the long axis of the whole mass. 
The surface material of each oval body is finely 
granular. The well-defined margins suggest that these 
tightly packed oval bodies were originally enclosed 
in a membrane or sac. There is obviously no way to 
investigate what they once contained. The only animal 
remains recorded from Merriganowry are phyllolepid 
fishes, possible thelodont scales (in coprolites), and 
rare eurypterid fragments. It is at least possible that 
they represent unhatched fish eggs and are presented 
here for the record. 


Relationships of phyllolepids 

Since their first discovery in Scotland in the early 
19 century (Agassiz 1844) the nature, origins and 
relationships of phyllolepids have been in dispute. At 
one time they were even thought to be ostracoderm 
agnathans (Woodward 1915, 1920) until Stensié 
(1934, 1936, 1939) confirmed that they were rather 
aberrant placoderms. 

Until the 1980s, phyllolepids were considered 
to be the sister group to the Arthrodira (Goujet 
1984) and several workers have suggested a close 
relationship with a distinctive endemic Australian 
arthrodire, Wuttagoonaspis Ritchie, 1973 (cf. 
Janvier 1996, figs 4.44, 4.49). Phyllolepids and 
wuttagoonaspids, which have now been recorded 
over a wide area in Australia (Young and Goujet 
2003), had a rather similar ridged ornament that has 


235 


A NEW GENUS OF DEVONIAN PLACODERMS 


been interpreted as a shared derived character of the 
two groups. In contrast, I (Ritchie 1973) suggested 
that phyllolepids and wuttagoonaspids were only 
distantly related, a view supported by Dupret (2004, 
48) who does not consider the ornament ridges in 
phyllolepids and wuttagoonaspids to be homologous 
and proposes Wuttagoonaspis as a sister group to all 
other arthrodires. Goujet and Young (1995, fig. 2) and 
Goujet and Young (2004, fig. 1) included both the 
Phyllolepida and Wuttagoonaspis in the Arthrodira, 
but as a polytomy, leaving their interrelationships 
unresolved. 

Dupret (2004) reviewed the phylogenetic 
relationships between actinolepids (which, like 
phyllolepids, had a sliding neck joint,) and other 
arthrodires, e.g. phlyctaeniids and brachythoracids 
(which had a rotating neck joint with articular 
condyles on the anterior dorsolateral plates). He 
grouped Phyllolepida with “Actinolepids” in 
“Actinolepidoidei”’, a paraphyletic assemblage, and 
suggested that Phyllolepida are the sister group to the 
Phlyctaenioidei (Phlyctaenii plus Brachythoraci). 


Palaeogeographic distribution of phyllolepids 

Of the ca 300 placoderm genera and eight 
placoderm orders that dominated the Devonian lakes, 
rivers and seas (Carr 1995), the phyllolepids display 
perhaps the most unusual stratigraphic distribution. In 
the former supercontinent of Laurussia (= Euramerica), 
where they were first found, phyllolepids are known 
only from the latest Devonian (Famennian). They are 
the only placoderm group with no Early or Middle 
Devonian fossil record in the Northern Hemisphere. 

Fossil discoveries from Gondwana since the 
early 1980s provide a possible explanation for the 
sudden appearance of phyllolepids in the Northern 
Hemisphere and a solution to the mystery of their 
origins (Young 1986, 1987, 1990, 1993a, 1993b, 
2003, in press a, b). With the exception of Venezuela 
and Antarctica, most of the phyllolepid finds from 
Gondwana have come from Australia, from much 
older deposits - Late Middle Devonian (Givetian) to 
Early Late Devonian (Frasnian) - than those in the 
Northern Hemisphere. They are also usually found, 
like their northern relatives, in freshwater sediments. 

This disjunct distribution of phyllolepids, in time 
and space, provides strong support for a dispersal 
episode in the mid-Late Devonian (Frasnian- 
Famennian boundary) at which time northern 
Gondwana and Laurussia came close enough to 
each other to allow the exchange of predominantly 
freshwater fish faunas. Phyllolepid fishes survived in 
Laurussia until the end of the Devonian but have left 
no record in Asia. 


236 


It is clear that much more remains to be uncovered 
about the early history of phyllolepid placoderms and 
that Gondwana, and especially Australia, displays 
the greatest potential for discoveries illustrating the 
origins and radiation of the Phyllolepida. 

Cowralepis mclachlani— gen. et sp. nov. 
makes a major contribution to our knowledge 
of the Phyllolepida. The Merriganowry site in 
central west New South Wales provides an ideal 
base for a long-term, multi-disciplinary scientific 
research programme into the palaeontological, 
palaeoecological and sedimentological history of a 
Late Middle Devonian lake and its fauna covering 
tens, or hundreds, of thousands of years. Under 
proper scientific supervision, this can be combined 
with, and financially supported by, a unique, hands- 
on, educational facility and eco-tourism experience, 
with long-term economic benefits for rural New 
South Wales. 


ACKNOWLEDGMENTS 


I wish to thank the Director and Trust of the Australian 
Museum for appointing me as a Museum Research Fellow 
following my retirement as Museum palaeontologist in 
1995 and for providing facilities and support enabling me 
to continue active research work on fossil fishes, especially 
in central west New South Wales. 

Without the support of Mr Alex McLachlan, owner 
of the property on which Merriganowry is situated, this 
study would not have been possible. His generous action 
in fencing and securing the Merriganowry site for scientific 
investigation by the Australian Museum has preserved a 
unique, world-class fossil assemblage for posterity, the full 
scientific, educational and eco-tourism potential of which 
has yet to be realised. 

The discovery of the Merriganowry site owes much to 
the late Mr Reg Dumbrell of Canowindra who discovered 
the remains of phyllolepid fossil fish near the Lachlan River 
and reported this to the Australian Museum. 

The remarkable assemblage of fossil specimens 
illustrated here represents only part of the material 
excavated from Merriganowry between 1993 and 2004 by 
many hundreds of paying volunteers, supervised by myself 
and Dr Zerina Johanson from the Australian Museum. 
These groups have been organised by Ms Monica Yeung 
of Canberra and Mr Bruce Loomes of Canowindra, through 
Gondwana Dreaming Inc. of Canberra. My deepest thanks 
to them and to all the dig participants who (mostly willingly) 
relinquished their fossil finds to the Australian Museum for 
this project and whose financial contributions helped to 
fund our research. 

Mr James Fairfax, of Bowral, New South Wales, 
generously funded our Canowindra Research Project (of 
which the Merriganowry study is part) for many years. 

I thank my palaeoichthyological colleagues, most 


Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


especially Dr Zerina Johanson, who has jointly supervised 
more than 60 weekend digs at Merriganowry; also Dr Gavin 
Young (Canberra), Dr John Long (Melbourne), Dr Robert 
Carr (Ohio), Dr Daniel Goujet (Paris), Dr Vincent Dupret 
(Paris) and many others for constructive discussions on 
phyllolepid anatomy and placoderm relationships. I would 
like to express particular gratitude to the referees, Dr Gavin 
Young and Dr Robert Carr, whose extensive and useful 
comments made this a much better paper. However, in the 
final analysis, the opinions expressed here are my own. 


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Stensi6, E. A. (1939). On the Placodermi of the Upper 
Devonian of East Greenland. Second supplement to 
Part I. Meddelelser om Gronland 97(3), 1-33, 6 pl 

Stensi6, E. A. (1963). Anatomical studies on the 
arthrodiran head. Part1. preface, geological and 
geographical distribution, the organization of the 
head in the Dolichothoraci, Coccosteomorphi and 
Pachyosteomorphi. Taxxonomic appendix. Kunglika 
Svenska Vetenskapsakademiens Handlingar. 9, 1- 
419. 


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Stensi6, E. A. (1969). Elasmobranchiomorphi 
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Masson). 

Turner, S., Basden, A. and Burrow. C. J. (2000). Devonian 
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Woodward, A. S. (1915). Preliminary report on the 
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Advancement of Science (1914), 84 Rpt. 

Woodward, A. S. (1920). Presidential address. 
Proceedings of the Linnean Society of London 1920: 
25-34. 

Young, G. C. (1979). New information on the structure 
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Young, G. C. (1980). A new Early Devonian piacoderm 
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Young. G. C. (1985). New discoveries of Devonian 
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Young. G.C. 1986. The relationships of placoderm fishes. 
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Young, G. C. (1987). Devonian palaeontological data 
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Young, G. C. (1988). New occurrences of phyllolepid 
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Young, G. C. (1989). The Aztec fish fauna of southern 
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Young, G. C. (1990). Devonian vertebrate distribution 
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Proc. Linn. Soc. N.S.W., 126, 2005 


239 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 1A-D. Cowralepis mclachlani n. gen. and sp. AMF 103767a, b. holotype, complete individual in part 
and counterpart. A) dorsal view; B) ventral view, C-D) detail, pelvic fins and vertebrae, in dorsal and ventral 
view respectively. Latex casts whitened with ammonium chloride sublimate. 


240 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 2A-J. Cowralepis mclachlani n. gen. and sp. Small to medium-sized individuals to the same 
scale illustrating the range of deformation in Cowralepis specimens from Merriganowry. A-C), short, 
broad symmetrical specimens; D-G) skewed specimens; (H-J) long, narrow symmetrical specimens. A) 
AMF 103756; B) AMF90053; C) AMF90027; D) AMF90034a, b; E) AMF96750; F) AMF104157a; G) 
AMF96747a; H) AMF103784; I) AMF90044b; J) AMF103778a 


Proc. Linn. Soc. N.S.W., 126, 2005 241 


A NEW GENUS OF DEVONIAN PLACODERMS 


160% 


Nu length 


MD length ne 


140% e ° 
! holotype 
e a AMF103767A 
e 


Figure 3A, B Cowralepis mclachlani n. gen. and sp. Tectonic deformation. A) relative lengths of dorsal plates 
in eighty Cowralepis specimens. Horizontal axis shows combined length of Nu+MD; vertical axis shows 
Nu/MD x 100. B) All Cowralepis specimens from Merriganowry have been tectonically deformed. The 
central figure depicts a hypothetical undeformed individual of Cowralepis. Surrounding figures, in different 
orientations, were enlarged by 10% in the direction of largest principal extension (GPE) and reduced by 10% 
in the direction of smallest principal extension (SPE), as indicated by accompanying strain ellipses. The 
strike anddip at Merriganowry are in register with the GPE and SPE of the strain ellipse, but this is probably 


coincidental. 


242 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


B. Cowralepis mclachlani n. gen. and sp. Two large individuals with tail. A) AMF90003a, dorsal 


2 


Figure 4A 


ventral view (cf. Fig. 6 for digitally modified versions of both). Latex casts whitened 


with ammonium chloride sublimate. 


>) 


view; B) AMF96762 


243 


Proc. Linn. Soc. N.S.W., 126, 2005 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure SA-D. Cowralepis mclachlani n. gen. and sp. Three symmetrical individuals illustrating short/broad 
and long/narrow examples of tectonic deformation. A, B) AMF103753a, b, dorsal and ventral (cf. Fig. 6C, 
D for same specimen after digital modification); C) AMF90051 dorsal view; D) AMF100018, ventral view. 
Latex casts whitened with ammonium chloride sublimate. 


244 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Ba abate tA ihe oe 
Caer 


i} 

ey 
oats 
eu 


f 


3 
Da’, 


Figure 6A-D. Cowralepis mclachlani n. gen. and sp. Digitally modified versions of A,B) long/narrow and 
C,D) short/broad Cowralepis specimens restored approximately to original proportions. A) AMF.90003a, 
dorsal surface (cf. Fig. 4A); B) AMF96762, ventral surface (cf. Fig. 4B); AMF103753, dorsal and ventral 
views (cf. Fig. 5A, B). 


Proc. Linn. Soc. N.S.W., 126, 2005 245 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 7A-I. Cowralepis mclachlani n. gen. and sp. A, B) AMF96779, left and right ADLs from same 
individual, in ventral view; C) AMF96762, left and right PDLs (cf. Figs 4B, 6B); D, E) AMF90054a, b, 
isolated pelvic fin skeleton, in counterpart; F) AMF96784, PNu with associated Mg, PMg and part of PtO, 
dermal view; G) AMF90029a, right paranuchal, dermal view; H) AMF90012, pelvic fins, dorsal view; I) 
AMF 103754, pelvic fins, dorsal view. Latex casts whitened with ammonium chloride sublimate. 


246 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 8A-G. Cowralepis mclachlani n. gen. and sp. Ventral trunk plates. A) AMF104154, partial head 

and trunk, ventral view; B) IL, AMV and otolith; C) AMF103770, right IL; D) detail of ornament; E) 

AMF 127152, anterior ventral margin with AMV and both IL plates slightly dislodged; F) AMF104160, 
AVL and PVL plates; no PMV visible; G) AMF127162, AVL and PVL plates; one PMV present. Latex casts 
whitened with ammonium chloride sublimate. 


Proc. Linn. Soc. N.S.W., 126, 2005 247 


A NEW GENUS OF DEVONIAN PLACODERMS 


a 
*, 
i 


"Sk Soa 


Figure 9A-G. Cowralepis mclachlani n. gen. and sp. A) AMF96781, ventral view; and B) AMF96781, detail 
of A); C) AMF104154, parasphenoid, ventral surface (cf. Fig. 8A); D) AMF103768, parasphenoid, dorsal 
surface; E) AMF96783, before digital modification; GPE and SPE indicate directions of Greatest and Shortest 
Principal Extension; F) AMF96783, after digital modification; arrows indicate direction and % of correction 
applied; G) AMF96783, right inferognathal and both superognathals in association. Latex casts whitened 
with ammonium chloride sublimate. 


248 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 10A-G. Cowralepis mclachlani n. gen. and sp. Gnathal elements. A) AMF103755S, left superognathals 
and inferognathal; B) AMF104164, left superognathals, inferognathal lost; C) AMF104155, left 
superognathals, inferognathal lost; D) AMF103787, right inferognathal and articular, Ign rotated to show 
occlusal surface; E) AMF96779, left inferognathal, ventral view; F) AMF96780, right articular, ventral 

view; G) AMF96786, before digital modification; GPE and SPE indicate directions of Greatest and Shortest 
Principal Extension; H) AMF96786, after digital modification; arrows indicate direction and % of correction 
applied. Latex casts whitened with ammonium chloride sublimate. 


Proc. Linn. Soc. N.S.W., 126, 2005 249 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 11A, B. Cowralepis mclachlani n. gen. and sp. AMF 127156, ventral surface of head shield and 
anterior margin of ventral trunk shield, A) before digital modification; GPE and SPE indicate directions of 
Greatest and Shortest Principal Extension; B) after digital modification; arrows indicate direction and % of 
correction applied. (cf. Fig. 12B-D for stereo pairs of basihyal, articulars and ?hyoid arch element). Latex 
cast whitened with ammonium chloride sublimate. 


250 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 12A-F. Cowralepis mclachlani n. gen. and sp. Stereopair images of visceral skeletal elements and 
otolith. A) AMF90007b, basihyal (cf. Fig. 14A); B) AMF127156, basihyal (cf. Fig. 11A); C) right articular, 
ceratohyal and hyoid arch element? (cf. Fig.9); D) AMF127156, left articular and hyoid arch element? (cf. 
Fig. 9); E) AMF96779, left otolith, ventral view; F) AMF96783, left articular. Latex casts whitened with 
ammonium chloride sublimate. 


Proc. Linn. Soc. N.S.W., 126, 2005 251 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 13A-E. Cowralepis mclachlani n. gen. and sp. Occipital and synarcual ossifications. A) AMF96751, 
head and trunk shields, ventral view; ventral shield lost, exposing axial skeleton; B) AMF103763, occipital 
ossification and synarcual, ventral view; C) AMF96753, isolated synarcual; dorsal (left) and ventral (right); 
D) AMF96785, juvenile dorsal shield, ventral view showing occipital ossification; synarcual lost. E) detail, 
ventral view. Latex casts whitened with ammonium chloride sublimate. 


O52 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 14A, B. Cowralepis mclachlani n. gen. and sp. A) AMF90007b, head shield of 

large individual, ventral view, showing all gnathal and visceral elements in association; B) 
AMF 103776, smaller individual, head in dorsal view, Nu lost, exposing visceral branchial 
skeleton and otoliths in dorsal view, surrounded by circum-nuchal plates . Latex casts whitened 
with ammonium chloride sublimate. 


Proc. Linn. Soc. N.S.W., 126, 2005 253 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 15A-D. Cowralepis mclachlani n. gen. and sp. A) reconstruction of craniothoracic armour, dorsal 
view; B) reconstruction of dorsal shield, ventral view (ventral shield omitted) showing overlap relationships 
of dorsal plates; C) reconstruction of craniothoracic armour, ventral view, showing gnathal elements, 
parasphenoid, otoliths, etc; D) reconstruction of dorsal shield, ventral view (ventral shield omitted) showing 
hypobranchials, occipital ossification and synarcual. 


254 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 16A-C. Cowralepis mclachlani n. gen. and sp. A) AMF96764, dorsal view; Nu and MD plates lost, 
exposing ventral elements; B) AMF96765, counterpart showing ventral shield, pelvic fins and complete 
vertebral column; C) AMF90004, dorsal view. Both specimens digitally modified (to correct for tectonic 
deformation) and drawn to same length to illustrate the relative lengths of dermal armour/tail in smaller and 
larger individuals.of Cowralepis. 


Proc. Linn. Soc. N.S.W., 126, 2005 255 


A NEW GENUS OF DEVONIAN PLACODERMS 


Wi, 


iG is Y, Yi 


Figure 17A-D. Cowralepis mclachlani n. gen. and sp. Caudal fin. A. B) AMF90048a, caudal fin seen from 
right; B) detail of anterior region; C, D) AMF90048b, same caudal fin seen from left; D) detail of anterior 
region. Latex casts whitened with ammonium chloride sublimate. 


256 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Figure 18A-F. Cowralepis mclachlani n. gen. and sp. A) AMF 127159, juvenile vertebral column; B) 
AMF127154b, isolated neural arcualia: C) neural and haemal arcualia reconstructed from AMF90048 (cf Fig. 
17); D) AMF127151a, egg sac from Merriganowry; E) AMF9001 1a, dorsal shield with smaller Cowralepis 
disappearing under anterior margin; F) AMF90011b, head shield in ventral view showing smaller Cowralepis 
inside buccal cavity (cf. Fig. 19 for interpretation). Latex casts whitened with ammonium chloride sublimate. 


Proc. Linn. Soc. N.S.W., 126, 2005 257 


A NEW GENUS OF DEVONIAN PLACODERMS 


Figure 19. Cowralepis mclachlani n. gen. and sp. Combined sketch of the dorsal and ventral surfaces in 
AMF9001 la, b. (cf. Fig. 18E, F). This individual appears to have died in the act of swallowing another 
Cowralepis individual that was too large, fortuitously providing invaluable information on the spatial 
relationships of the gnathal and branchial skeletal elements in Cowralepis mclachlani. 


258 Proc. Linn. Soc. N.S.W., 126, 2005 


A. RITCHIE 


Cowralepis mclachlani 
Late Middle Devonian (Givetian) 
Merriganowry, near Cowra, 
New South Wales 
(length - up to 35 cm) 


Figure 20. Cowralepis 
mclachlani n. gen. and sp. 
A) Reconstruction of 
Cowralepis with soft tissues of 
head margin restored and prob- 
able position of eye indicated. 
B) Four named phyllolepid genera 
showing the interrelationships of the 
postorbital (PtO), marginal (Mg), para- 
nuchal (PNu) and nuchal (Nu) plates of the 
head shield. 
[1] Placolepis budawangensis Ritchie, from the early 
Late Devonian (Frasnian) of southeastern New South 
Wales (Ritchie 1984, fig. 14B); 
[2] Cowralepis mclachlani new genus and species (this paper 
Fig. 15A) from the late Middle Devonian (Givetian) or early Fras- 
nian of central west New South Wales; 

[3] Austrophyllolepis ritchiei Long from Mt Howitt, Victoria, originally 
dated as Frasnian, now thought to be Givetian (Young, pers. comm.). Long’s reconstruction of A. ritchiei 
(1984, fig. 7A); digitally modified to correct for tectonic deformation at Mt Howitt. 

[4] Phyllolepis orvini Heintz, from the Late Devonian (Famennian) of East Greenland (after Denison, 1978, 
fig. 29). 


Placolepis budawangensis 


Proc. Linn. Soc. N.S.W., 126, 2005 259 


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<<) 7 77 
: - 


Larval Development and Autogeny in Ochlerotatus 


camptorhynchus (Thomson) (Diptera: Culicidae) from Southern 


Victoria 


Puitie S. BARTON AND JOHN G. ABERTON 


School of Ecology and Environment, Deakin University, Geelong, VIC 3217 


Barton, P.S. and Aberton, J.G. (2005). Larval development and autogeny in Ochlerotatus camptorhynchus 
(Thomson) (Diptera: Culicidae) from southern Victoria. Proceedings of the Linnean Society of New 
South Wales 126, 261-267. 


Larval development and autogeny was examined in the mosquito Ochlerotatus camptorhynchus 
(Thomson) from southern Victoria. Larvae of Oc. camptorhynchus were reared in the laboratory at 5 
constant temperatures (15, 20, 25, 30 and 35°C) and three constant salinities (0, 18 and 36 ppK). Of the five 
temperatures, survival ranged from 35.6% at 35°C to 84.4% at 20°C, and development times ranged from 
12.1 to 37.1 days at 35°C and 15°C respectively. The minimum threshold temperature for development was 
7.3°C, and the thermal constant was 324.0 + 12.8 SE degree-days. No differences in development times 
or survival were detected for the three salinities. Adult mosquitoes reared from field-collected pupae and 
larvae reared on a high-nutrition diet displayed no autogenous egg development. A positive relationship 
was found between adult body size (wing length) and fecundity in blood-fed adults, with fecundity ranging 


from 40 to 112 eggs per female. 


Manuscript received 14 October 2004, accepted for publication 16 February 2005. 


Key Words Autogeny, culicidae, larval development, fecundity, Ochlerotatus camptorhynchus 


INTRODUCTION 


The southern saltmarsh mosquito, Ochlerotatus 
camptorhynchus (Thomson), is an abundant mosquito 
Species in southern coastal Australia from southwest 
Western Australia around to the southern coast of New 
South Wales, including Tasmania (Dobrotworsky 
1965; Russell 1993). It is a confirmed vector of 
Ross River virus (Ballard and Marshall, 1986) and 
suspected vector of Barmah Forest virus (Aldred et al. 
1990; Russell 1995). Ochlerotatus camptorhynchus 
was also detected on the North Island of New Zealand 
during 1998 (Hearnden et al. 1999), and a large 
eradication program is now under way. 

Larvae of Oc. camptorhynchus are typically 
found in brackish to fresh ground pools associated 
with coastal swamps and bushland (Dobrotworsky 
1965; Lee et al. 1984), but also in some salinity- 
affected areas inland (Wishart 2002). No studies 
have previously investigated the responses of 
immature stages of Oc. camptorhynchus to different 
temperatures or salinities. 

Autogeny, or the ability to develop an initial batch 


of eggs without a blood meal, has been described 
to varying levels in several Australian mosquito 
species including Anopheles hilli Woodhill and Lee 
(Sweeney et al. 1973), Ochlerotatus vigilax (Skuse) 
(Sinclair 1976; Hugo et al. 2003), Culex annulirostris 
Skuse (Kay et al. 1986), Culex sitiens Wiedemann 
(Fanning et al. 1992), Ochlerotatus australis 
(Erichson) (Brust 1997) and Culex molestus Foérskal 
(Dobroworsky 1954). Of these mosquitoes, arguably 
the most closely related ecologically is Oc. vigilax, 
a saltmarsh mosquito replacing Oc. camptorhynchus 
as the dominant mosquito in northern coastal areas 
from southern New South Wales around to southern 
Western Australia (Lee et al. 1984; Russell 1993). 
Autogeny in Oc. camptorhynchus has not previously 
been investigated. Production of an autogenous egg 
batch may delay the first mosquito-host contact (Kay 
et al. 1986), and is thought to accelerate the rate of 
natural increase of mosquito populations (Tsuji et al. 
1990). 

The purpose of this study is to increase our 
understanding of the development and survival of 
larval Oc. camptorhynchus, which may provide for 
greater accuracy of timing of larvicide applications, as 


LARVAL DEVELOPMENT AND AUTOGENY IN A MOSQUITO 


well as a better understanding of the basic ecological 
parameters of this important mosquito species. 


METHODS 


Larval development 

Thirty newly hatched larvae (1“ instar) of Oc. 
camptorhynchus (<12hrs old) from a saltmarsh at 
Breamlea (144°35’ East, 38°13’ South), near Geelong 
in southern Victoria, were put into each of six 
translucent plastic containers with 300 ml of water 
(salinity = 18 ppK). Sets of six containers (total = 180 
larvae) were then placed in a water bath for each of 
15, 20, 25, 30 and 35°C. Larvae were fed ground K9® 
Gold Fish Food (Go-Pet Petcare Solutions) at rates 
of: Ist and 2nd instars = 0.12 mg/larva/day, 3rd instar 
= 0.48 mg/larva/day, 4th instar = 0.96 mg/larva/day. 
Any excess food was removed daily. Subsamples of 
20 adult females were taken from each temperature 
treatment, and those had a single wing removed 
and measurement with a graticule eyepiece at 10x 
magnification from the wing tip (excluding the fringe) 
to the arculus (Harbach and Knight 1980). 

To evaluate salinity tolerance, seawater (36 
ppK) was collected from near Breamlea and used as 
stock solution, diluted equally with distilled water to 
produce a concentration of 18 ppK, and with distilled 
water being used for the 0 ppK concentration. Thirty 
newly hatched larvae, again collected from the 
Breamlea saltmarsh, were added to 300 ml of each 
concentration (2 replicates each). Concentrations 
were kept constant by adding distilled water daily to 
compensate for evaporation. A constant temperature 
of 25°C was maintained with the use of a water bath. 
Larvae were fed as described above. 

Average development time and survival for all 
immature stages were calculated using frequency- 
weighted means, based on daily counts. Analysis 
of Variance with Student Newman-Keuls post-hoc 
tests (SPSS v11) was used to compare differences in 
total development times and survival to adulthood 
between treatments. The relationship between larval 
development times and temperature, plus wing length 
and temperature, was examined using least squares 
linear regression. The day-degrees (K) needed for 
development at each experimental temperature (T°C) 
and duration of development (t days) was calculated 
from K = t (T-C), where C is an extrapolation of 
minimum temperature for development. 


Autogeny 
Autogeny was assessed in two categories of 
mosquito: 1) adults derived from field-collected 


pupae, and 2) adults derived from larvae reared 
on a high nutrition diet. All larvae and pupae were 
taken from a field site at Breamlea flora reserve near 
Geelong in southern Victoria (144°35’ East, 38°13’ 
South, Fig 3.1). 

The field-sourced pupae were collected from the 
same site at Breamlea during September 2002 and 
October 2003 and were allowed to emerge over a 24- 
hour period in a cage of approx 0.5m’. The laboratory- 
reared larvae were collected during October 2003 as 
first instar and reared with a high nutritional feeding 
regime at a daily rate of: 1“ and 2" instar = 0.16 mg/ 
larva, 3 instar = 0.64 mg/larva, and 4" instar = 1.28 
mg/larva. The field-collected pupae and laboratory- 
reared larvae were kept in water taken from the field 
(36ppK) with salinity kept constant by topping up 
containers with distilled water. All emerged adult 
mosquitoes were given immediate access to 10% 
sucrose solution and males were not removed from 
the cages. Immatures and adults were kept at ambient 
laboratory temperatures (approx 20 + 5°C). Ten days 
after emergence, all female mosquitoes were removed 
and cold anaesthetised before dissection. Ovaries were 
dissected and placed on slides with a saline solution 
for inspection of follicles at 200x magnification. 
Recording of stages of ovarian development was done 
with reference to Clements and Boocock (1984). 

A separate sample of mosquitoes derived from 
pupae at Breamlea flora reserve (October 2003) was 
allowed to blood-feed three days after emergence. 
These mosquitoes were dissected seven days after the 
blood meal and their fecundity (number of eggs per 
female) recorded, as well as with wing length. 


RESULTS 


Larval development 

At all temperatures tested, the first and fourth 
instars had the shortest and longest development 
times, respectively (Table 1). A significant difference 
between total development time and temperature 
was obtained (d.f. = 4, M.S. = 1313.06, F = 66.65, 
P < 0.01). Mean development was 12.1 + 0.9 days 
at 35°C, and 37.1 + 1.3 days at 15°C. Survival to 
adulthood was significantly different between the 
temperature treatments (d.f. = 4, M.S. = 205.55, F = 
12.42, P < 0.01), with 84.4% survival at 20°C and 
35.6% survival at 35°C. No difference was obtained 
for total immature development in different salinity 
treatments (d.f. = 2, M.S. = 2.99, F = 0.22, P= 0.80). 
No difference was obtained for immature survival 
between the salinity treatments (d.f. = 2, M.S. = 4.50, 
F = 1.50, P=0.35). 


Proc. Linn. Soc. N.S.W., 126, 2005 


P.S. BARTON AND J.G. ABERTON 


Table 1 Duration, development rate and survival of immature stages of Oc. camptorhynchus at constant 
water temperatures and salinities. * Values followed by different letters are significantly different at the 
5% level (Student-Newman-Keuls test). “Proportion of immature development per day. 


Days (mean + SE) in each stage 


Tempera- | I II Il IV P Total* Develop- %Sut- 
ture °© ment rate’ —_ vival* 
15 ARO Seenoer- Om ooe-- leo lt Si Oe 7 0.9 37-1 3a, 01027 68.9 a,b 
20 SulheeOS o¢kvee il SOs 1D TSsb iS SAEED” Wosyes oy OB y3 84.4a 
2S a Om On 420 OMe 4 eae ale IO! 20:24 1:06 0105 76.1a 
30 Se aOlop eeSe- Ol 2-6-2 0.0 S25 OF a1 £09) 1334109d 01075 52.8 b 
35 Ee Ooms = 0 HO mee 10) 30-2 0:86 512 14090 0!083 3515 © 
Salinity (ppK) at 25°C 
0 D5#OES BRO” Ailes Sse I Aue) eC) Seals ee OOS 98.5 a 
18 PASI Sm eo eeH OOF 942-0) 2 = ll S010" 22002 Me0a 0:05 88.3a 
36 25208 S210 AIO SS sete SoS ee 0) OO SSI KOE OES) 93.3a 


Table 2. Percentage ovarian development in two categories of adult female Oc. camptorhynchus, 10 


-days post emergence. 


Percent ovarian stage 


Source of adults la Ib lia lib 


Pupae taken from field 0 19 


Larvae reared on high diet 0 12.4 42.8 


A significant linear relationship (d.f. = 1, M.S. = 
0.01, F = 109.01, P < 0.01) was obtained between 
water temperature and rate of development with a 
coefficient of determination of R? = 0.971 (Fig 1). The 
lower threshold for development was extrapolated to 
7.3°C, and the thermal constant required for complete 
development was calculated as 324.0 + 12.8 SE 
degree-days. A significant negative linear relationship 
Git 1 MES: = 12. — 638.183, P< 0:01) was 
obtained for wing length and temperature (Fig 2.) 


Autogeny 

No autogeny was observed in either category of 
Oc. camptorhynchus assessed in this study. The most 
common stage of follicular development in the two 
categories examined was stage Ila (field = 51.2%, 
laboratory = 42.8%) (Table 2). The next highest 
percentage of follicular development was stage IIb 
(field = 26.2%, laboratory = 32.4%). All blood fed 
mosquitoes developed eggs to stage V, seven days 
after the blood meal. A significant linear relationship 


Proc. Linn. Soc. N.S.W., 126, 2005 


iia ib “iva Ivbi  V Total 


No. 


Sil VRS) 06. 0 0 0 168 
324 9 34-0 0 0 145 


was found between body size (wing length) and the 
number of mature follicles per mosquito (d.f. = 1, MS 
= 5,668.00, F = 41.64, P < 0.01) (Fig 3). Fecundity 
of blood-fed females ranged from 40 to 112 eggs per 
mosquito (72.41 + 2.99 SE, n= 34). 


DISCUSSION 


This study demonstrates that Oc. camptorhynchus 
is well adapted to cooler temperatures and is widely 
tolerant of different salinities. In contrast to Oc. vigilax, 
the Breamlea population of Oc. camptorhynchus 
exhibits no autogeny, indicating a different survival 
strategy to its more northern congener. 

The development of larval Oc. camptorhynchus 
responded to temperature and was linear between 15 
and 35°C. None of the temperatures tested were lethal 
to the Victorian strain Oc. camptorhynchus. The 
development threshold temperature of 7.3°C for Oc. 
camptorhynchus suggests development of immatures 


263 


LARVAL DEVELOPMENT AND AUTOGENY IN A MOSQUITO 


0.10 


y =0.003x - 0.020 


2 
= 0.97 
0.08 R =0.971 


Figure 1. Relationship be- 
tween development rate of 
immature stages of Oc. camp- 
torhynchus and five con- 
stant water temperatures. 


0.06 


Development rate (1/days) 


0.00 
 pcamead || ul ie aati liar alae | ante fe gs ag 
W ater temperature (°C) 
48 
4.6 t = -0.067x + 5.493 
fa} R?= 0.957 
wn 
+H 
Figure 2. Relationship between & 
wing length of adult female & 
Oc. camptorhynchus and five S 
constant water temperatures. a 
"po 
“4 
= 
5 
a 


10 15 20 25 30 35 40 


Water temperature (°C) 


264 Proc. Linn. Soc. N.S.W., 126, 2005 


P.S. BARTON AND J.G. ABERTON 


140 


120 


100 


80 


60 


40 


Fecundity (number of eggs) 


20 


y = 40.452x - 103.791 
R? =0.565 


50 pe es oi 0 
Wing length (mm) 


4.5 5.0 S20, ow 


Figure 3. Relationship between wing length and fecundity for blood-fed 


Oc. camptorhynchus (n = 34). 


will continue in winter in southern Victoria, where 
minimum temperatures of approximately 8°C 
commonly occur (Bureau of Meteorology 1993). The 
larvae of Oc. camptorhynchus at 15°C had higher 
survival (69%) and a lower development threshold 
(7.3°C) compared to more northern species reared at 
15°C, such as Cx. sitiens from southeast Queensland 
(10% survival, 11.9°C threshold) (Mottram et al. 
1994), Cx. annulirostris from northern Victoria (3 % 
survival, 9.7°C threshold) (McDonald et al. 1980) and 
Ae. aegypti from north Queensland (24% survival, 
8.3°C threshold) (Tun-Lin et al. 2000). 

At 25°C, the absence of any affect of salinity 
on the development or survival of larval Oc. 
camptorhynchus suggests a high level of adaptation to 
saline conditions. The mechanism for survival in high 
salinities is likely to be a capacity to produce non- 
toxic osmolytes (Bradley 1987), which can neutralise 
the harmful effects of high salt concentrations. 
Possible interaction between salinity and temperature 
was not examined and differences in survival and/or 
development may be observed at varying salinities at 
higher or lower temperatures. 


Proc. Linn. Soc. N.S.W., 126, 2005 


During expected spring or summer temperatures 
of 15-25°C, and following inundation of larval 
habitat, survival of Oc. camptorhynchus will be 
high, depending on predation, and development 
may take 20-37 days. No threshold salinity, above 
which survival is curtailed, is apparent for Oc. 
camptorhynchus between OppK and 36ppK. Given 
the optimal stage for treatment with the control 
agent Bacillus thuringiensis israelensis are 2"! to 34 
instars, this suggests a large treatment window of 
approximately 7-15 days, and if s-methoprene were to 
be used, the time before treatment could be extended 
to 4 instar if necessary. However, in situations 
where prolonged temperatures of above 30°C occur, 
survival of Oc. camptorhynchus might be expected 
to be lower, and control may not be warranted. The 
operational criteria for control, therefore, should be 
based on larval densities, size of breeding site and on 
proximity to residential areas. 

Adult Oc. camptorhynchus derived from field- 
collected pupae and larvae reared with high nutrition 
in the laboratory both failed to exhibit autogeny. 
This differs remarkably compared to the saltmarsh 


265 


LARVAL DEVELOPMENT AND AUTOGENY IN A MOSQUITO 


mosquitoes Oc. vigilax from Australia (Sinclair 1976; 
Hugo et al. 2003), and Oc. taeniorhynchus from the 
USA (O’Meara and Edman 1975), where autogeny 
rates of up to 100% and 94.4% were observed in these 
studies, respectively. 

Of all the adults assessed for autogeny, the 
majority of displayed ovarian development to stages 
Ila or IIb, which is the previtellogenic resting stage 
described by Clements and Boocock (1984). At this 
developmental stage, it is thought that a blood meal 
is required to facilitate further follicular development 
and oviposition. That a greater percentage of adults 
reared on a high diet had follicles over stage IIb 
(12.4%) compared to adults derived from field- 
collected pupae (3.6%) suggests that nutrition may be 
important in influencing ovarian development in Oc. 
camptorhynchus. 

The fecundity of blood-fed Oc. camptorhynchus 
increased with body size, a well-known relationship 
documented with other mosquitoes (Nayar and 
Sauerman 1975; Armbruster and Hutchinson 2002). 
Larger mosquitoes might therefore be expected to 
produce larger egg batches 

The anautogeny apparent from the sampled Oc. 
camptorhynchus suggests this species may require 
blood meals for survival and egg development and 
therefore appetential dispersal after emergence 
might occur earlier, in relative terms, than for Oc. 
vigilax, as time for egg development and oviposition 
is not required. This in turn suggests that the role 
of Oc. camptorhynchus in biological transmission 
would commence earlier, as this species is unlikely 
to mature an autogenous egg batch. From an 
ecological standpoint, dispersal of anautogenous 
Oc. camptorhynchus would seem to be a risky 
survival strategy when suitable saltmarsh habitats 
are discontinuous, as on the Bellarine Peninsula 
where this study was performed. This result might be 
expected to change geographically, where different 
genetic and environmental factors may influence 
autogenous expression (Sota and Mogi 1995; Hugo 
et al. 2003), and is worthy of further study. 


ACKNOWLEDGEMENTS 


We thank Steve Sodomaco (Greater Geelong City 
Council) for information on larval habitat on the Bellarine 
Peninsula and Haylee Weaver (Central Queensland 
University) for assisting with the collection of mosquito 
larvae. Prof Brian Kay (Queensland Institute of Medical 
Research) provided helpful advice on the project and 
the manuscript. Dr David Slaney (Wellington School of 
Medicine and Health Sciences) provided helpful information 


266 


on development rates of larval mosquitoes. This study was 
supported by a Deakin University Postgraduate Research 
Scholarship. 


REFERENCES 


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Brust, R.A. (1997). Ecology of Aedes australis (Erichson) 
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48. 


Proc. Linn. Soc. N.S.W., 126, 2005 


267 


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BOOK REVIEW 


Bibliography of Australian Entomology 1687-2000 


by G. Daniels, 2004. Privately published by the author, P.O. Box 828, Mt Ommaney, Qld 4074. Email: 
g.daniels@uqconnect.net. Two volumes. viii, 1776 pages. A$275. 


This very impressive work, in two large 
hard-bound volumes, documents the Australian 
entomological literature from the earliest published 
record to the year 2000 and partly beyond. It lists 
over 52,000 references and attempts to include every 
publication that mentions an Australian insect. Some 
three quarters of the work (1322 pages) is occupied 
in documenting the references while the remaining 
quarter (450 pages) comprises a very comprehensive 
index and a list of journal titles. The work begins 
with a Contents listing and a short Introduction. 
The two volumes are case bound for heavy use. 

References are listed alphabetically by 
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format. Multi-authored works are cross referenced to 
each author. Taxonomic references are accompanied 
by a list of all new taxa described; these lists are 
sometimes extensive. Author’s given-names are 
included where known and biographical summaries 
are included for some authors. The bibliography also 
includes references on Collembola and Protura, groups 
which until recently were included in the class Insecta. 

While a serious attempt has been made 
to list every publication mentioning an Australian 
insect the author is aware that it is inevitable he has 
not achieved perfection. However, it is clear that 
this is by far the most comprehensive bibliography 
of Australian entomology ever published. Further, 
it is the only bibliography on all Australian 
entomological literature for the period 1931-2000. 

In addition to author listings in the References 
there are entries for topics such as Australian 
Museum, ANIC, Coleopterorum Catalogus, Genera 
Insectorum, Royal Entomological Society of London, 
School of Public Health & Tropical Medicine, 
Zoological Catalogue of Australia and names of ships 
of famous voyagers cross-referenced to relevant 
literature. References with complex or controversial 
publication details are annotated with notes on dates 
of publication, pagination of parts, authors, etc. 

The Index is an important part of this 
Bibliography. It is exceptionally detailed and 
comprehensive. References are arranged under 242 
major subject areas such as Behaviour, Christmas 


Island, Distribution, Insecticides, Forensic 
Entomology, Fungal Diseases, Keys, Malaria, 
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and Zoogeography. These are further divided by Order, 
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which in turn are further divided into family headings 
and in some cases species headings, the total entry 
running to over 7 pages of small print. This enables 
the user to locate literature on almost any subject. 

Works such as this require exceptional 
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is evident that the author has these attributes in 
abundance. Typographical errors are exceedingly 
rare in the reference list and index, although a few 
have crept into the Introduction. The author is to be 
congratulated on a super- human effort that provides 
a basic tool for Australian entomologists. It is clear 
that anyone wanting to get into the literature on any 
subject concerning Australian entomology should 
consult this impressive work. It is not available 
electronically but this is of little consequence 
because the Index is so comprehensive. At least 
as hard copy it will always be readily accessible. 


M.S. Moulds 
Australian Museum 


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CONTENTS - CONTINUED 


203 Young, G. 
A new phyllolepid placoderm occurrence (Devonian fish) from the Dulcie Sandstone, Georgina 
Basin, central Australia. 

IIS) Ritchie, A. 
Cowralepis, a new genus of phyllolepid (Pisces, Placodermi) from the Late Middle Devonian of 
New South Wales, Australia. 

261 Barton, P.S. and Aberton, J.G. 
Larval development and autogeny in Ochlerotatus camptorhynchus (Thomson) (Diptera: 
Culicidae) from southern Victoria. 

269 Book review - Bibliography of Australian Entomology 1687-2000 by G. Daniels. 

270 Instructions to authors 


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PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W. 
VOLUME 126 


vn 


TS EPO: [RT 
Issued 20 March 2005 
CONTENTS 


1 Holmes, W.B.K. and Anderson, H.M. 
The Middle Triassic megafossil flora of the Basin Creek Formation, Nymboida Coal Measures, New 
South Wales. Part 4. Umkomasiaceae. Dicroidium and affiliated fructifications. 

39 Holmes, W.B.K. and Anderson, H.M. 
The Middle Triassic megafossil flora of the Basin Creek Formation, Nymboida Coal Measures, New 
South Wales. Part 5. The genera Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris: 

81 Paterson, J.R. 
Revision of Discomesites and Estaingia (Trilobita) from the Lower Cambrian ehabde Vale Formation, 
western New South Wales: taxonomic, biostratigraphic and biogeographic implications. 

95 Allen, S. and Huveneers, C. 
First record of an Australian fur seal (Arctocephalus pusillus doiiferus) feeding on a wobbegong shark 
(Orectolobus ornatus). 

99 Keith, D. and Pellow, B. 
Effects of Javan rusa deer (Cervus timorensis) on native plant species in the Jibbon-Bundeena area, 
Royal National Park, New South Wales. 


111 Percival, |.G. and Wright, A.J. 
Anew Early Silurian species of Trimerella (Brachiopoda: Craniata) from the Orange district, New South 
Wales. 


121 Jerry, D.R. 
Electrophoretic evidence for the presence of Tandanus tandanus (Pisces: Plotosidae) immediately north 
and south of the Hunter River, New South Wales. 

125 ~~ ~Moulds, T. 
Diversity and biogeography of subterranean guano arthropod communities of the Flinders Ranges, South 
Australia. 

sees) Moulds, M. 
Song analyses of cicadas of the genera A/eeta Moulds and Tryel/a Moulds (Hemiptera: Cicadidae). 

143 Rickards, B., Parkes, R. and Wright, A.J. 
Llandovery (Early Silurian) graptolites from the Quidong Basin, NSW. 

153 Rickards, B., Farrell, J.R., Wright, A.J. and Morgan, E.J. 
Silurian graptolites from the Barnby Hills Shale and Hanover Formation, New South Wales. 

171 Semple, W.S. and Koen, T.B. 
Altitude, frost and the distribution of white box (Eucalyptus albens) on the central tablelands and adjacent 
slopes of NSW. 

181 Wood, A. 
Collections of Galerina (Agaricales, Fungi) made by J.B. Cleland and housed in the State Herbarium of 
South Australia. 

197 McPherson, J.R. 
A recent expansion of its Queensland range by Eupristina verticillata Waterston (Hymenopera, 
Agaonidae, Agaoninae), the pollinator of Ficus microcarpa |.f. (Moracea). 


CONTINUED INSIDE BACK COVER 
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