Memoirs of Museum Victoria 61(2): 121-127 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

Mitochondrial 12S rRNA sequences support the existence of a third species of

freshwater blackfish (Percicthyidae: Gadopsis ) from south-eastern Australia

Adam D. Miller, Gretchen Waggy 1 , Stephen G. Ryan and Christopher M. Austin 2

School of Ecology and Environment, Deakin University, PO Box 423, Warmambool, Vic. 3280, Australia

1 Current address: Department of Coastal Science, College of Marine Science, The University of Southern Mississippi,

703 East Beach Drive, Ocean Springs, MS, USA, gretchen.waggy@usm.edu

2 Author for correspondence: cherax@deakin.edu.au

Abstract Miller, A.D., Waggy, G., Ryan, S.G., and Austin, C.M. 2004. Mitochondrial 12S rRNA sequences support the existence

of a third species of freshwater blackfish (Percicthyidae: Gadopsis ) from south-eastern Australia. Memoirs of Museum

Victoria 61(2): 121-127.

Fish of the genus Gadopsis are a distinctive component of the freshwater fish fauna of south-eastern Australia.

Gadopsis marmoratus and G. bispinosus are the only two species recognised within the genus, with the former of uncer-

tain taxonomic status, as it is thought to be composed of at least two distinct geographical forms based on morphologi-

cal and allozyme data. The objective of this study was to investigate DNA sequence divergence in Gadopsis, especially

in the western portion of its distribution, using an approximately 400 base pair fragment of the mitochondrial small sub-

unit 12S rRNA gene region in order to reassess the taxonomy of the genus. Individuals from 1 1 locations were sequenced

and confirm that G. marmoratus and G. bispinosus are genetically distinct, and further that the G. marmoratus complex

consists of two divergent clades representing the previously identified northern and southern forms. The degree of diver-

gence between the three Gadopsis clades was similar (5-6% nucleotide substitutions), suggesting that they diverged from

a common ancestor at approximately the same period in geological time. These results are consistent with previous

allozyme studies and highlight the usefulness of mitochondrial DNA data coupled with allozyme information for clari-

fying taxonomic boundaries in morphologically conservative aquatic organisms.

Keywords Mitochondrial rRNA, taxonomy, blackfish, Percicthyidae, Gadopsis, Australia

Introduction

Fish of the genus Gadopsis, commonly known as the river or

freshwater blackfish, are endemic to south-eastern Australia

(including Tasmania), and carry out their entire life cycle in

freshwater (Jackson et al., 1996). The genus is phylogenetic-

ally distinct and its evolutionary origins remain uncertain as it

may have either evolved from a marine ancestor some 15

million years ago or had a more ancient Gondwanan fresh-

water origin (Sanger, 1984; Jerry et al., 2001).

Gadopsis belongs to the family Percicthyidae and contains

two currently recognised species, G. marmoratus (Richardson,

1848) and G. bispinosus (Sanger, 1984). Sanger (1986)

suggested, based on morphological and allozyme evidence, that

G. marmoratus potentially consists of a northern and a south-

ern species. However, Sanger (1986) did not formally recog-

nise the northern and southern forms of G. marmoratus as these

putative species were not found in sympatry and because the

taxonomic significance of the genetic and morphological

divergence between the two forms was uncertain.

Gadopsis marmoratus has a large geographic range that

includes tributaries of the Murray-Darling river system, as far

north as the Condamine River in southern Queensland. The

species is also found in Tasmania with endemic populations in

the north and translocated populations in the Huon River and

elsewhere in the south. Sanger (1986), assuming that G. mar-

moratus is in fact a complex of two species, suggested that

these taxa evolved in allopatry following the isolation of

Tasmania from mainland Australia. According to this scenario

ancestral gadopsids are thought to have been originally wide-

spread throughout Victoria and northern Tasmania and that the

formation of the two species may have occurred during periods

of raised sea levels which isolated Tasmanian from mainland

populations. Subsequently, when sea levels were lower during

the Pleistocene glaciation and land connections re-established

with the mainland, the Tasmanian form of G. marmoratus

invaded southern Victoria, consequently displacing the north-

ern G. marmoratus (Ovenden et al., 1988).

Sanger’s (1986) biogeographical hypothesis assumes that

the two forms of G. marmoratus behave as independent

122

A. D. Miller, G. Waggy, S. G. Ryan and C. M. Austin

species. This hypothesis also suggests that the two forms have

come into contact in the past and therefore it may still be pos-

sible to find locations where both the northern and southern

forms coexist in Victoria (Koehn and O’Connor, 1990). Sanger

(1986) suggested that the two forms may occur in sympatry in

the state’s southwest, an area encompassing the Gellibrand and

Glenelg river systems. An allozyme study by Ryan et al. (in

press) was unsuccessful in finding evidence supporting the

existence of sympatric populations of northern and southern

G. marmoratus in south-western Victoria. However, their find-

ings were consistent with Sanger’s (1986) results indicating

genetic divergence between the two forms inhabiting adjacent

river systems in this region.

In addition to Sanger’s and Ryan’s allozyme studies there

have been several studies of blackfish using DNA-based tech-

niques (Ovenden et al., 1988; Waters et al., 1994; Jerry et al.,

2001). These studies, however, are limited by minimal sam-

pling of the northern form of G. marmoratus, especially in the

western portion of its distribution. This study therefore extends

these studies by using Polymerase Chain Reaction (PCR)

amplification of an approximately 400 base pair fragment of

the mitochondrial 12S rRNA gene region coupled with direct

DNA sequencing, in order to further evaluate the taxonomic

status of the northern and southern forms of G. marmoratus

with emphasis on its western distributions. This approach was

chosen because mitochondrial DNA (mtDNA) has been found

to be very useful for inferring phylogenetic and taxonomic

relationships in groups of organisms where the protein-based

techniques of allozyme electrophoresis have lacked resolution

or produced ambiguous results (Hillis et al., 1996).

Methods and materials

Gadopsis samples. Tissue samples were obtained from specimens pre-

viously collected by Ryan et al. (in press). Sample selection was based

on the results of Ryan et al. (in press) together with three reference

sites based on previous studies by Sanger (1984) and Ovenden et al.

(1988). These three sites consisted of the MacDonald River

(Murray-Darling catchment, northern G. marmoratus ), the Gellibrand

River (south-west Victoria, southern G. marmoratus) and Cudgewa

Creek (north-east Victoria, G. bispinosus). The other samples obtained

by Ryan et al. (in press) included specimens of the G. marmoratus

complex from eight additional sites. These sites included Darlot Creek,

Bracknell Creek, the Wimmera River, and the Wannon River. In addi-

tion DNA sequences were provided by D. Jerry, James Cook

University, Queensland, for samples from Stony Creek, Victoria, rep-

resenting both G. bispinosus (GenBank accession number: AF294459)

and G. marmoratus and Little Forester Creek (Tasmania) also repre-

senting G. marmoratus (AF294452). The remaining specimens were

from Eight Mile Creek and Mosquito Creek (South Australia) (Fig. 1).

Samples of Maccullochella peeli peeli (Murray cod) and Bostockia

porosa (Western Australia nightfish) were included as outgroups

(sequences derived from GenBank, accession numbers: AF295060 and

AF295048)

DNA extraction and amplification of mtDNA. Total DNA was

extracted from muscle tissue using an extraction protocol developed by

Crandall et al. (1999). A fragment of the 12S mtDNA gene region

(approximately 400 bp) was amplified via PCR using the 12S c/d

primers described in Jerry et al. (2001). Double stranded PCR ampli-

fications were performed in 50 ml volumes consisting of: lx PCR

buffer, 2.0 pM MgC12, 0.2 mM dNTPs, 1 mM of each primer and 2

units Tag DNA polymerase. PCR amplifications were performed in a

Corbett PC-960 Microplate Thermal Sequencer and consisted of 30

cycles of denaturation at 94°C for 30 sec, annealing at 55 °C for 30 sec,

and extension at 72°C for 30 sec. An initial denaturation cycle of 3 min

at 94°C was used and the program ter-minated with a 5 min cycle at

72 °C. The PCR products were visualised in 1% agarose / TAE gels

stained with ethidium bromide under UV light.

Purification and sequencing. PCR products were purified using a

QIAquick PCR purification kit (QIAGEN) according to the manufac-

turers instructions. Purified DNA was quantified via direct comparison

with DNA marker (Promega DNA / HAE III marker) of kn own con-

centration, again visualized under UV light in a 1% agarose / TAE gel

containing ethidium bromide. Purified DNA was then sequenced

according to Australian Genome Research Facility (AGRF), University

of Queensland, protocols.

Data analysis. Sequence chromatograms were viewed using EditView

and edited using SeqPup software (Gilbert, 1997). Sequences were

aligned using the Clustal X program (Thomson et al., 1997) with

alignment-ambiguous regions excised prior to phylogenetic recon-

struction (Gatesy et al., 1993). Phylogenetic analyes were conducted

using a range of approaches implemented by the PAUP* software

package (Swofford, 1998). Phylogenetic signal within the data set was

assessed using the gl statistic from the random tree length-frequency

distribution (Hillis and Huelsenbeck, 1992). Phylogenetic relationships

were estimated using maximum parsimony, neighbour-joining and

maximum likelihood approaches. The most parsimonious tree was

identified using a full exhaustive search with support for branches

evaluated by 1,000 bootstrap replicates. Distance analysis was per-

formed using the Tajima-Nei model of evolution and the neighbour-

joining option with the number of bootstrap replicates set at 1,000. The

most appropriate model of evolution for the maximum likelihood (ML)

analyses was obtained via testing alternative modes of evolution using

Modeltest (Posada and Crandall, 1998).

Results

Approximately 400 bp of the mitochondrial 12S rRNA coding

region were sequenced for 12 individuals of Gadopsis from 11

locations. After sequence editing, 290 bp were used for sub-

sequent analysis (GenBank accession numbers: AF505866 -

AF505872). The random tree distribution based on the entire

data set including both the outgroup taxa is significantly

skewed to the left with gl= -0.899, P<0.01, indicating signifi-

cant phylogenetic information (Hillis and Huelsenbeck, 1992).

The random tree distribution within the ingroup taxa was also

significantly skewed (gl= -0.686, PcO.Ol).

Percentage sequence divergences and the number of

nucleotide substitutions among individuals (Table 1) indicate

the existence of three equally distinct groups within Gadopsis.

The individuals representing southern G. marmoratus (samples

9-12, Table 1) differ at 14-18 base positions (5-6% sequence

divergence) in comparison with northern G. marmoratus

individuals (samples 3-8), and 15-16 base positions (5-6%

sequence divergence) compared to G. bispinosus (samples 1

and 2). Gadopsis marmoratus (northern) and G. bispinosus

differed at 15-20 base positions (5-7% sequence divergence).

In contrast, comparison between samples within each of

these groups revealed much lower levels of divergence with

G. marmoratus (southern), G. marmoratus (northern) and

Mitochondrial sequences of freshwater blackfish

123

Figure 1. Sample locations: 1. Mosquito Creek, 2. Eight Mile Creek, 3. Wimmera River, 4. Wannon River, 5. Darlot Creek, 6. Bracknell Creek,

7. Gellibrand River, 8. Stony Creek, 9. Cudgewa Creek (G. bispinosus ), 10. Little Forester Creek (Tasmania), 11. MacDonald River (New South

Wales).

G. bispinosus showing differences at 0-1 bp positions

(0.00-0.03%), 0-8 bp positions (0-3% sequence divergence)

and 4 bp positions (1% sequence divergence) respectively.

Nevertheless, geographic variation was apparent within the

northern form of G. marmoratus. Blackfish samples from west-

ern Victoria and south-eastern South Australia (sites 1-4) differ

by 7-8 base positions from the two samples from the

Murray-Darling River system (sites 8-11). Variation within

each of these groups was minimal with haplotypes either being

identical or differing at only a single base position.

The degree of divergence between the outgroup taxa,

B. porosa and M. peeli peeli, and blackfish samples was sub-

stantial ranging between 12 and 16% (34-45 bp differences).

This was also similar to the difference between the two out-

group taxa (10% sequence divergence) (Fig. 2). The maximum

parsimony, distance and maximum likelihood methods gave

similar tree topologies. The Tamura-Nei model was chosen for

the maximum likelihood analysis, involving a full heuristic

search with support for branches evaluated by 100 bootstrap

replicates and the application of a gamma distribution shape

parameter value equal to 0.2293 and calculated base frequency

and substitutional rate matrix values. The significant feature of

the trees is the clustering of Gadopsis into three distinct clades,

representing the two putative northern and southern species of

G. marmoratus and G. bispinosus. These three clades are sup-

ported by high confidence values in each method of analysis

(66-100% bootstrap). Especially noteworthy is that the analy-

ses do not necessarily indicate that the northern and southern

forms are each other’s closest relative. While maximum parsi-

mony and maximum likelihood methods indicate an unresolved

trichotomy for the relationship between the three clades, the

distance approach suggests the northern G. marmoratus may in

fact be more closely related to G. bispinosus than to the south-

ern G. marmoratus , although the bootstrap support for this

relationship is low. The maximum likelihood and distance

analyses also highlight phylogenetic patterns within the

124

A. D. Miller, G. Waggy, S. G. Ryan and C. M. Austin

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Mitochondrial sequences of freshwater blackfish

125

Wimmera R (3)

Eight Mile Ck (2)

Mosquito Ck (1)

W arm on R (4)

MacDonald R ( 1 1)

Stony Ck (8)

Gellihrand R (7)

Darlot Ck (5)

L. Forester Ck (10)

Bracknell Ck (6)

Stony Ck (bispinosus) (8)

Cudgewa Ck (bispinosus) (9)

B. porosa

M. peeli peeli

Wimmera R (3)

Wnnnon R (4)

Mosquito Ck (1)

Eight Mile Ck (2)

MacDonald R (it)

Stohy Ck (8)

Stony Ck (bispinosus) (8)

Cudgewa Ck (bispinosus) (9)

L. Forester Ck ( 10)

Gellihrand R (7)

Darlot Ck (5)

Bracknell Ck (6)

B. porosa

M. peeli peeli

MacDonald R (11)

Stony Ck (8)

Wimmera R (3)

Eight Mile Ck (2)

Mosquito Ck (1)

Wannon R (4)

Gellihrand R (7)

Darlot Ck (5)

Bracknell Ck (6)

L, Forester Ck (10)

Stony Ck (bispinosus) (8)

Cudgewa Ck (bispinosus) (9)

M. peeli peeli

B. porosa

Figure 2. Phylogenetic trees, using the PAUP software package (Swofford, 1998). A, Maximum parsimony, using a full exhaustive search with

1000 bootstrap replicates. B, Distance analysis, using the neighbour-joining option, bootstrap replicates set at 1000. C, Maximum likelihood, using

the Tamura-Nei model with 100 bootstrap replicates.

northern form of G. marmoratus. The samples from south-

eastern South Australia and western Victoria (sites 1-4) form a

well supported clade (78-100% bootstrap support) as do the

individuals from Stony Creek and the MacDonald River from

the Murray-Darling River system (99-100% bootstrap

support) (Fig 2.)

Discussion

The taxonomic value of molecular genetic data is widely appre-

ciated (Hillis et al., 1996) and they can be used in essentially

the same way as other data to address issues concerning the

identification of taxonomic boundaries. Thus, finding that

the degree of nucleotide divergence between G. bispinosus and

the northern form of G. marmoratus is similar to that between

G. bispinosus and the southern form (5-7%), and substantially

greater than that observed within these groupings (0-3%),

strongly suggests that each represents a distinct taxonomic

entity.

The degree of divergence between the northern and southern

forms of G. marmoratus and G. bispinosus is very similar to

those reported by Jerry et al. (2001). These authors examined a

126

A. D. Miller, G. Waggy, S. G. Ryan and C. M. Austin

560 bp fragment of the 12S gene region for a single individual

representing each of the three genetic forms of blackfish as part

of phylogenetic study on Australian members of the family

Percichthyidae. It is also noteworthy that Jerry et al. (2001)

reported divergence levels of a similar or smaller magnitude for

a number of congeneric percichthyid species that are con-

sidered good biological species. This together with the fact that

G. bispinosus and G. marmoratus are known to behave as good

biological species based on their maintenance of genetic differ-

ences in sympatry (Sanger 1986), suggests that the northern

and southern forms of G. marmoratus also represent distinct

biological species.

Additional support for the validity of northern and southern

forms of G. marmoratus as discrete species derives from an

examination of intra- and interspecific levels of divergence.

Overall, the intraspecific comparisons average was 1.2%, com-

pared with an average of 5.2% divergence for interspecific

comparisons. The largest intraspecific comparison is 2.8%

divergence between samples of northern G. marmoratus from

Eight-Mile Creek in South Australia and the MacDonald River

in New South Wales. Finding this degree of intraspecific diver-

gence is not surprising given that the samples come from

independent drainages over 1000 km apart. Conversely, finding

that the Wannon River sample differs from the Darlot creek

sample by 14 base positions (6%) and are less than 100 km

apart strongly suggests that either a biological or geographical

barrier has limited or completely impeded migration of black-

fish between the two adjacent drainages. Decoupling of

genetic divergence and geographic separation between samples

indicates that the two forms represent good biological species

and suggests that if they do come into contact they are

unlikely to interbreed.

The application of the biological species concept to

allopatric populations has, however, been widely criticized and

is considered a persistent problem for taxonomic studies of

freshwater fish (McDowell, 1972). Some authors have called

for the abandonment of the biological species concept and its

replacement with lineage or genealogically-based concepts

such as the phylogenetic species concept (Claridge et al., 1997;

Avise and Walker, 1999; Shaw, 2001). An advantage of the

phylogenetic and related species concepts is that they allow

recognition of species in sympatry or allopatry because

genealogical relationships can be determined independently of

geographical status (Shaw, 2001). While there are also oper-

ational difficulties in the application of lineage-based species

concepts (Avise and Wollenberg, 1997; Sites and Crandall,

1997), the three distinct clades of blackfish identified in this

study, via all three methods of phylogenetic analysis, would

qualify for recognition as distinct species when applying a

lineage-based species concept.

It is unwise to base the determination of species boundaries

on a single source of information such as mitochondrial

sequences from a single gene region. Support for the taxo-

nomic conclusions of this study comes from studies of

allozyme and morphological variation (Sanger, 1986; Ryan et

al., in press) and restriction digests of the whole mitochondrial

genome (Ovenden et al., 1988). Allozyme data indicated sub-

stantial differences between G. marmoratus and G. bispinosus

(22% fixed differences) and between the northern and the

southern forms of G. marmoratus (11% fixed differences)

(Ryan et al., in press). It is noteworthy that while the allozyme

variation between the northern and southern forms is low

relative to that between G. marmoratus and G. bispinosus, the

extent of these differences are far greater than that detected

between samples within the northern and southern groupings.

Significantly, finding the same pattern of geographically abrupt

genetic discontinuity between samples of blackfish from west-

ern Victoria in both mitochondrial DNA and allozymes (Ryan

et al., in press), provides substantial support for the recognition

of distinct northern and southern species. Further, results

reported by Ovenden et al. (1988) are entirely consistent with

the findings of this study based upon restriction digest of the

whole mitochondrial genome.

An outcome of this study, which was not apparent from

allozyme analyses or Ovenden’s study (1988), is the finding of

geographic variation in mitochondrial 12S rRNA sequences

within the northern form of G. marmoratus. Specifically, based

upon the six samples analysed, it appears that Gadopsis from

western Victoria and south-eastern South Australia (sites 1-4)

form a monophyletic group distinct from those of the

Murray-Darling drainage system (sites 8 and 11). While the

degree of divergence among these two groups is considerably

less than that seen among the northern and southern G. maro-

moratus and G. bispinous, they nevertheless represent distinct

diagnosable lineages. It is noteworthy that Jackson et al.

(1996), without going into any great detail expressed the view

that blackfish from south-eastern South Australia may possibly

be taxonomically distinct, therefore it will be important to

investigate this pattern of variation in greater detail and deter-

mine if blackfish from this region may deserve taxonomic

recognition.

Independent of the consideration of taxonomic status of

Gadopsis spp., it is apparent that four evolutionary significant

units (Waples, 1995) can be recognised in Victoria. If sup-

ported by additional sampling, each of these units will require

the development of appropriate management strategies if the

blackfish biodiversity is to be conserved and protected. In addi-

tion to loss of populations due to habitat deterioration, trans-

locations associated with aquaculture, stocking of private water

bodies and the use of Gadopsis as live bait, are factors that

could threaten the integrity of local blackfish stocks. The

genetic hazards of local translocations are well illustrated by

the rapid genetic displacement of a freshwater crayfish species

in the south-west of Western Australia as a result of an

inadvertent translocation of a closely related species (Austin

and Ryan, 2002).

The major difference between the results of this and previ-

ous allozyme studies is that the allozyme data appear to under-

estimate the degree of divergence between the northern and

southern forms. In fact, the relationships among the three puta-

tive Gadopsis species remain an open question. The parsimony

and maximum likelihood analyses suggest an unresolved tri-

chotomy (see also Ovenden et al., 1988), the distance analysis

suggests that the northern form of G. marmoratus is more

closely related to G. bispinosus, although with poor bootstrap

support, Jerry et al. (2001) supports a closer relationship

Mitochondrial sequences of freshwater blackfish

127

between the southern form and G. bispinosus, and the allozyme

data suggests that the northern and southern G. marmoratus

species are the most closely related. These inconsistencies

leave the phylogenetic relationships among these species

unresolved, and therefore also their possible evolution and

biogeographic history (Sanger, 1986).

Given the vulnerability of Gadopsis from a conservation

perspective, the relatively high degree of genetic diversity

found in this study, and the unresolved phylogenetic relation-

ships among the three major Gadopsis lineages, it becomes

apparent that further research is important. Given the relatively

slow evolutionary rate of the mitochondrial 12S rRNA gene

region, it is suggested that genetic variation within and between

Gadopsis populations using more rapidly evolving gene

regions is determined to fully resolve geographical patterns of

genetic diversity in these taxonomically distinct groups and the

phylogenetic relationships among them. Further, the geo-

graphic sampling of Gadopsis for taxonomic and population

genetic analysis needs to be expanded, especially with respect

to populations in the eastern part of Victoria for which our

genetic knowledge is limited.

Acknowledgments

We thank all those who contributed to this study. Many thanks

to Dr Dean Jerry from James Cook University, Townsville,

Queensland, for supplying G. bispinosus samples, and Daniel

Ierodiaconou from the School of Ecology and Environment,

Deakin University, Warmambool, for GIS support. We would

also like to thank Dr Christopher P. Burridge from the

Molecular Ecology and Biodiversity Laboratory, Deakin

University, Warmambool, for his contribution to the revision of

this manuscript, and to the team at the Molecular Ecology and

Biodiversity Laboratory for their assistance in the field and the

laboratory.

References

Austin, C.M., and Ryan, S.G. 2002. Allozyme evidence for a new

species of freshwater crayfish of the genus Cherax Erichson

(Decapoda: Parastacidae) from the south-west of Western Australia.

Invertebrate Systematics 16: 357-367.

Avise, J.C., and Walker, D. 1999. Species realities and numbers in sex-

ual vertebrates: perspectives from an asexually transmitted genome.

Proceedings of the National Academy of Science 96: 992-995.

Avise, J.C., and Wollenberg, K. 1997. Phylogenetics and the origin of

species. Proceedings of the National Academy of Sciences 94:

7748-7755.

Claridge, M.F., Dawah, H.A., and Wilson, M.R. 1997. Practical

approaches to species concepts for living organisms. Pp 1-13 in:

Species: The Units of Biodiversity. Chapman and Hall Publishers:

New York.

Crandall, K.A., Fetzner, J.W., Lawler, S.H., Kinnersley, M., and

Austin, C.M. 1999. Phylogenetic relationships among the

Australian and New Zealand genera of freshwater crayfishes

(Decapoda: Parastacidae). Australian Journal of Zoology 47:

199-214

Gatesy, J., DeSalle, R., and Wheeler, W. 1993. Alignment- ambiguous

nucleotide sites and the exclusion of systematic data. Molecular

Phylogenetics and Evolution 2(2): 152-157

Gilbert, D.G. 1997. SeqPup software. Indiana University.

Hillis, D.M., and Huelsenbeck, J. P. 1992. Signal, noise, and reliabili-

ty in molecular phylogenetic analyses. Journal of Heredity 83:

189-195

Hillis, D.M., Mable, B.K., Larson. A., Davis, S.K., and Zimmer, E.A.

1996. Nucleic acids IV: sequencing and cloning. Pp. 336-339 in:

Hillis, D.M., Moritz, C., and Mable, B.K. (eds), Molecular system-

atics (2nd edn). Sinauer Associates: Sunderland.

Jackson, P.D., Koehn, J.D., Lintermans, M., and Sanger, A.C. 1996.

Family Gadopsidae, freshwater blackfishes. Pp. 186-190 in:

McDowall, R.M. (ed.), Freshwater fishes of south-eastern

Australia. A.H. and A.W. Reed Publishing: Sydney.

Jerry, D.R., Elphinstone, M.S., and Baverstock, P.R. 2001.

Phylogenetic relationships of Australian members of the family

Percichthydae in ferred from mitochondrial 12S rRNA sequence

data. Molecular Phylogenetics and Evolution 18(3): 335-347

Koehn, J.D., and O’Connor, W.G. 1990. Biological information for the

management of native freshwater fish in Victoria. Victorian

Government Printer: Melbourne. 165 pp.

McDowall, R.M. 1972. The species problem in freshwater fishes and

the taxonomy of diadromous and lacustrine populations of

Galaxiias maculatus (Jenyns). Journal of the Royal Society of New

Zealand 2(3): 325-367.

Ovenden, J.R., White, R.W.G., and Sanger, A.C. 1988. Evolutionary

relationships of Gadopsis spp. inferred from restriction enzyme

analysis of their mitochondrial DNA. Journal of Fish Biology 32:

137-148.

Posada, D., and Crandall, K.A. 1998. MODELTEST: testing the model

of DNA substitution. Bioinformatics 14: 817-818.

Ryan, S.G., Miller, A.D., and Austin, C.M. in press. Allozyme variation

and taxonomy of the river blackfish, Gadopsis marmoratus

Richardson, in western Victoria. Proceedings of the Royal Society

of Victoria.

Richardson, J. 1848. Ichthyology. In: Zoology of the Voyage ofH.M.S

Erebus and Terror, Under the Command of Capt. Sir J. C. Ross,

R.N. During the Years 1839 to 1843. Longman, Brown and

Longmans: London.

Sanger, A.C. 1984. Description of a new species of Gadopsis (Pisces:

Gadopsidae) from Victoria. Proceedings of the Royal Society of

Victoria 96: 93-97.

Sanger, A.C. 1986. The evolution and ecology of the Gadopsis mar-

moratus complex. Ph.D. thesis. The University of Melbourne,

Melbourne.

Shaw, K.L. 2001. The genealogical view of speciation. Journal of

Evolutionary Biology 14: 880-882.

Sites, J.W., and Crandall, K.A. 1997. Testing species boundaries in bio-

diversity studies. Conservation Biology 11(6): 1289-1297.

S wofford, D.L. 1998. PAUP*. Phylogenetic analysis using parsimony

(*and other methods ). Version 4. Sinauer Associates: Sunderland.

128 pp.

Thomson, J.D., Gibson, T.J., PlewniakF., Jeanmougin, F., and Higgins,

D.G. 1997. The Clustal X windows interface: flexible strategies for

multiple sequence alignment aided by quality analysis tools.

Nucleic Acids Research 24: 4876-4882.

Waples, R.S. 1995. Evolutionary significant units and the conservation

of biological diversity under the Endangered Species Act. American

Fisheries Society Symposium 17: 8-27.

Waters, J.M., Lintermans, M., and White, R.W.G. 1994.

Mitochondrial-DNA variation suggests river capture as a source of

vicariance in Gadopsis bispinosus (Pisces: Gadopsidae). Journal of

Fish Biology 44(3): 549-551.

Memoirs of Museum Victoria 61(2): 129-133 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

Dimorphic brooding zooids in the genus Adeona Lamouroux from Australia

(Bryozoa: Cheilostomata)

Philip E. Bock 1 2 and Patricia L. Cook 2

1 School of Ecology and Environment, Deakin University, Melbourne Campus, Burwood Highway, Burwood, Vic. 3125

(pbock @ deakin.edu. au)

2 Honorary Associate, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

Abstract Bock, P.E., and Cook, PL. 2004. Dimorphic brooding zooids in the genus Adeona Lamouroux from Australia (Bryozoa:

Cheilostomata). Memoirs of Museum Victoria 61(2): 129-133.

The genus Adeona is a characteristic and common part of the Australian shelf fauna, extending to the tropical Indo-

West Pacific. The genus first appears in the fossil record of the Miocene of south-eastern Australia. Zooid dimorphism

has been recognised initially from subtle differences in the external appearance, which have not been described

previously. Detailed examination has shown enlarged brooding zooids with marked differences from autozooids in the

internal stmcture of the peristomes and in the occurrence of a primary calcified orifice.

Keywords Bryozoa, bryozoans, Cheilostomata, Adeonidae, Recent, Australia, brooding, dimorphism

Introduction

The Family Adeonidae includes genera with colonies which are

mainly erect and bilaminar, and some which are encrusting.

Frontal wall development is umbonuloid as demonstrated in a

study of a range of Recent material (Cook, 1973). Further

analysis of skeletal structures in a number of Australian and

New Zealand examples was done by Lidgard (1996). Enlarged

brooding zooids are well-known in the genera Adeonellopsis

MacGillivray, 1886, Reptadeonella Busk, 1884 (Hayward and

Ryland, 1999), and Dimorphocella Maplestone, 1903.

Brooding dimorphs are also characteristic in Adeonella

Maplestone, 1903 and Laminopora Michelin, 1842, which dis-

play schizoporelloid frontal wall development. For that reason

these latter genera are considered to belong to a distinct family,

the Adeonellidae (although there is disagreement on this issue,

see Lidgard, 1996). In the genus Adeona Lamouroux, 1812,

enlarged brooding zooids (“ooecial cells”) were first identified

by Busk (1884: 181) in the species A. appendiculata Busk,

1884 from Twofold Bay, off Eden, NSW. His illustration (pi.

33, fig. 6) shows two zooids with slightly larger secondary ori-

fice dimensions. He also figured the opercula of brooding and

non-brooding dimorphs (fig. 47). Cook (1973: 249, 250)

inferred the presence of brooding zooids in unidentified

material from the collection of the Natural History Museum,

but these were not described or illustrated. A study of skeletal

regeneration in a specimen of Adeona (Wass, 1983), included

illustrations of early ontogenetic stages of regenerating zooids.

These show dimorphism in the size of the secondary calcified

orifices but no comment was made about this dimorphism.

Wass (1991) illustrated zooidal variation in Adeona, and

remarked on the larger size of inferred brooding zooids, and the

porous distal plate in the calcified orifice of these zooids.

At least 15 species of Adeona have been described, mainly

from the Recent of Australia, with some from Indonesia, Japan

and South Africa, and one from Brazil. A thorough revision of

this group is badly needed; it is expected that some of these

species will be better placed in other genera, while it is also

believed that detailed study will reveal more undescribed

species from Australia. The type species, Adeona grisea

Lamouroux, 1812 was collected from Australia by the Baudin

expedition: the exact locality is not known. It is assumed that

the type material of Lamouroux was largely destroyed but

material in the Nice Museum may be relevant (Tillier, 1977).

Most of the specimens collected from southern Australia are

of folded and branching bilaminate fenestrate sheets, often

forming colonies 150-250 mm wide and high. These large

colonies are attached to the sea floor by a complex articulated

stem built of porous calcareous segments joined by cuticular

tubes, forming a stout trunk (Bock and Cook, 2000). The orig-

inal illustration of Lamouroux (1816) shows a single fenestrate

sheet but it is uncertain if this is a specific character or if it rep-

resents an early stage of colony development. Colonies with a

lanceolate, non-fenestrate form are known from Indonesia and

Western Australia as Adeona foliif era Lamarck, 1816 ( =Adeona

foliacea Lamouroux, 1816).

130

P. E. Bock and P. L. Cook

Material and methods. The collection of Museum Victoria includes a

large amount of material of the family Adeonidae from southern

Australia, including very large colonies collected from near Port

Phillip heads in the late nineteeth and early twentieth centuries. This

has been supplemented by colonies dredged from Bass Strait in the

1980s, and from the Great Australian Bight in 1995. The latter

material was sampled using the CSIRO Marine Laboratories vessel,

RV Franklin. The taxonomic revision of this collection is a major proj-

ect which has been barely commenced.

The material examined is from three samples: stations GAB-033,

GAB- 113, BSS-119 (locality details given in explanation of figures).

Four colonies were used, apparently belonging to four distinct species,

although species identification has not been attempted in this pre-

liminary study. These species appear to be neither Adeona cellulosa

(MacGillivray, 1869) nor A. wilsoni (MacGillivray, 1881). It is pos-

sible that the material may be identified as one or more of the four

species and one variety defined by Kirchenpauer, 1880 {Adeona

albida, A. arborescens, A. intermedia, A. macrothyris and A. foliacea

var. fascialis ) but the type material of these has not been seen.

In order to prepare the interior of the frontal shield for examination,

dry colony fragments were cemented to a glass slide with domestic

cyanoacrylate adhesive ('Superglue'). After polymerization, the entire

upper layer of zooids, together with the basal wall of the lower layer,

was removed by abrasion. After dissolving the adhesive with acetone,

the remaining material was cleaned of organic tissue using bleach. The

resulting preparation clearly shows details of the internal skeletal

structures, as revealed in the illustrations.

Observations

The genus Adeona includes several species, with the most

common colony form composed of a boxwork of branching bil-

aminar fenestrate sheets. Some species form simple sheets, or

branches without fenestrae. Colonies of the complex types are

up to 30 cm in diameter; the age of these colonies is unknown

but is suspected to be of the order of some tens of years.

Colonies are attached, usually to solid substrates, by a slightly

flexible stem composed of calcareous stem joints with cuticu-

lar connecting tubes (Bock and Cook, 2000).

The systematically important characters of Adeona are con-

sidered to include gross colony morphology, fenestra size, type

of heterozooids on the fenestral rim and the zooidal skeletal

characters. Zooidal appearance changes greatly and rapidly

with ontogeny, so that young zooids at the growing edge of the

colony should be more reliable for identification than older

zooids with thick secondary calcification. Internal characters of

the zooid skeleton should also reveal useful characters, partic-

ularly in the shape of the orifice, although no information on

this has been published previously.

Specimens of a number of species of the genus have been

examined externally after removal of tissue using bleach,

revealing that zooid dimorphism is common. The four exam-

ples illustrated here differ in the zooid and orifice proportions,

the relative position of frontal avicularia and foramen opening,

and the presence or absence of avicularia on brooding zooids.

It is inferred that the dimorphs with larger secondary orifices

are brooding zooids. The calcified external opening of the ori-

fice is generally larger in the brooding zooids. This is best seen

in zooids near the growing edge of the colony where secondary

calcification is less well-developed (Figs 1, 4). Exa mi nation of

the interior of the frontal shield has revealed considerable

differences between brooding and non-brooding zooids (Fig.

2). The interior opening of the brooding zooid peristome is

much larger and leads to a distal peristomial chamber with

smooth calcification. This space contained large single

embryos which were observed during the process of sectioning

the colony. The smooth calcification of the brood chamber ter-

minates proximally at a line that is continuous with the ring

scar of the body chamber of the zooid (Fig. 3). The lateral and

distal walls peripheral to the ring scar are perforated with

numerous pores leading to tubes communicating with the

septular pores on the exterior surface of the calcified skeleton.

The porous distal plate observed by Wass (1991) is the frontal

surface of the peristomial brood chamber (Figs 5, 8).

Observations on the fairly small sample which has been

examined in detail shows that brooding zooids may occur in

clusters of up to at least 20 zooids (Fig. 6). These clusters do

not appear to be closely related to colony margins or to be prox-

imal to colony fenestrae. However, in other colonies brooding

zooids appear to be scattered with no obvious clustering.

Other differences between brooding zooids and non-

brooding zooids are relatively subtle. Brooding zooids tend to

be larger (Fig. 4) but not greatly (Fig. 1). In one example the

brooding zooids lack avicularia (Fig. 5) but the significance of

this character is not known. In many examples, a thin laminar

plate is seen on each side of the orifice at the position where the

hinge of the operculum would be articulated (Fig. 3). These are

considered to be paired condyles, which have not been noted in

this genus previously. These are also present in non-brooding

zooids (Fig. 2).

The illustrations show the complexity of the frontal shield in

Adeona. The wall over the epistegal space is clearly multi-

layered (Figs 4, 8). A thin layer forms immediately above the

epistegal space and an overlying porous layer develops by

secondary thickening, beneath the hypostegal coelom (Fig. 7).

The rapid thickening of the outer layer of secondary calcifi-

cation on the colony surface obscures such relationships except

at the growing margin. The pores seen in the outer layer (Fig.

8) are conventionally termed areolae; they are assumed to be

for communication between the hypostegal coelom and the

main zooid. However, the complexity of these pores suggests

that the complete account of their function is yet to be eluci-

dated. Additional work needs to be done using thin sections,

together with methods of reconstructing the paths of com-

munication pores. Further investigation should also exa mi ne

the relationship between the tubes communicating between

the zooid interior and the hypostegal coelom and those

communicating with adjacent zooids.

Previous accounts, subsequent to Cook (1973), have

described the external opening to the epistegal space as a spira-

men rather than as an ascopore as this opening does not lead to

a true compensation sac in the interior of the zooid. However,

this terminology is ambiguous as the term spiramen normally

applies to an opening leading from the frontal surface into a

peristome, as in the genus Porina. The frontal openings in

Adeona and Adeonellopsis are more closely analogous to

frontal wall foramina in the families Arachnopusiidae and

Exechonellidae, and preferably should be termed foramina. In

the material examined, the foramen initially develops in the

Dimorphic brooding zooids in bryozoans

131

Figure 1. Growing edge of colony of Adeona species 1 (stn GAB- 113,

Great Australian Bight, depth 106 m, 34°36'S, 119°55'E). Starred

zooids are inferred brooding dimorphs. Arrow indicates zooid also

arrowed in Fig. 2. Scale = 500 pm.

Figure 3. Detail of the distalmost of the two brooding zooids in Fig. 2

(not reversed). Shows ring scar at margin of epistegal space (arrow),

plate-like condyle (arrow), and numerous communication pores later-

al to orifice and brood chamber, leading to hypostegal coelom. Scale =

100 pm.

Figure 2. Interior of part of material shown in Fig. 1 . Image is reversed

left to right for comparison of zooid positions in the fragment. Arrow

indicates zooid also arrowed in Figure 1 . Two brooding dimorphs show-

ing enlarged orifice and peristomial brood chamber. Scale = 200 pm.

Figure 4. Growing edge of colony of Adeona species 2 (stn GAB- 113

as in Fig. 1). Ontogenetic thickening of secondary calcification and

development of avicularia. Distalmost three zooids are brooding

dimorphs, as well as orifice in left proximal margin. Arrow marks cal-

cification around foramen. Scale = 500 pm.

132

P. E. Bock and P. L. Cook

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^ •v 4 ^ / -

^ ^/v '^svfi

V ■- Vr

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: -rs *3tlL ■ r.,: ,.-‘4 ^*7

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Figure 5. Same colony as Fig. 4. Proximal zooid and left zooid are

brooding dimorphs, with well-defined ‘distal plate’ in orifice, and with

no avicularium, having communication pores in its place. Distal and

right zooids are non-brooding zooids. Scale = 200 pm.

Figure 6. Adeona species 3 (stn BSS-119, Bass Strait, depth 92 m,

39°6.7'S, 143°28.7'E). Interior of colony section between three

fenestrae, showing clusters of brooding dimorphs at top left, bottom

left, and centre right. Scale = 1 mm.

Figure 7. Same specimen as Fig. 6. Detail of single non-brooding

zooid. Showing multi-layered frontal calcification, with communica-

tion pores in outer layer. Scale = 200 pm.

Figure 8. Adeona species 4 (stn GAB-033, Great Australian Bight,

depth 106 m, 34°36'S, 119°55'E). Two brooding zooids with partly bro-

ken frontal walls, showing distal plate and condyle in calcified orifice.

Scale = 200 pm.

Dimorphic brooding zooids in bryozoans

133

lower layer of calcification, and the secondary layer

progressively thickens above it (Fig. 4).

Acknowledgments

We should like to thank Dr Yvonne Bone (University of

Adelaide) and the Master and crew of RV Franklin for the

opportunity for one of us (PEB) to participate in the survey

which collected material from the Great Australian Bight in

July 1995.

References

Bock, P. E. and Cook, P. L. 2000. Early astogeny of Adeona colonies.

Pp. 161-167 in: Herrera Cubilla, A., and Jackson, J.B.C. (eds),

Proceedings of the 11th International Bryozoology Association

Conference. Smithsonian Tropical Research Institute: Balboa.

Cook, P.L. 1973. Preliminary notes on the ontogeny of the frontal body

wall in the Adeonidae and Adeonellidae (Bryozoa, Cheilostomata).

Bulletin of the British Museum of Natural History ( Zoology ) 25:

243-263.

Busk, G. 1884. Report on the Polyzoa collected by H.M.S. Challenger

during the years 1873-1876. Part 1. The Cheilostomata. Report on

the Scientific Results of the Voyage of the H.M.S. “Challenger”,

Zoology 10: 1-216.

Hayward, P.J., and Ryland, J.S. 1999. Cheilostomatous Bryozoa. Part

2. Hippothoidea - Celleporoidea. (2nd edn) In: Barnes, R.S.K., and

Crothers, J.H. (eds), Synopses of the British Fauna (New Series) 14.

The Linnean Society of London and Estuarine and Coastal Sciences

Association, Field Studies Council: Shrewsbury.

Kirchenpauer, G. H. 1880. Uber die Bryozoen-Gattung Adeona.

Abhandlungen aus dem Gebiete der Naturwissenschaften,

Herausgegeben von dem naturwissenschaftlichen Verein in

Hamburg 7: 1-24.

Lamarck, J.B.P.A. de M. de 1816. Histoire naturelle des Animaux sans

Vertebres ... precedee d'une introduction offrant la determination

des caracteres essentiels de l' animal, sa distinction du vegetal

et des autres coorps naturels, enfin, exposition des principes

fondamentaux de la zoologie. Verdiere: Paris. 568 pp.

Lamouroux, J.V.F. 1812. Extrait d'un memoire sur la classification des

Polypiers coralligenes non entierement pierreux. Nouveau Bulletin

Scientifique de la Societe Philosophique 3: 181-188.

Lamouroux, J.V.F. 1816., Histoire des polypiers Coralligenes

Flexibles, vulgairement nommes Zoophytes. F. Poisson: Caen. 559

pp.

Lidgard, S. 1996. Zooidal skeletal morphogenesis of some Australian

and New Zealand Adeonellopsis (Cheilostomatida). Pp. 167-177

in: Gordon, D.P, Smith, A.M., and Grant-Mackie, J.A. (eds),

Bryozoans in Space and Time. NIWA: Wellington.

MacGillivray, P.H. 1869. Descriptions of some new genera and species

of Australian Polyzoa; to which is added a list of species found in

Victoria. Transactions and Proceedings of the Royal Society of

Victoria 9: 126-148.

MacGillivray, P.H. 1881. On some new species of Catenicella and

Dictyopora', and on Urceolipora, a new genus of Polyzoa.

Transactions and Proceedings of the Royal Society of Victoria 17:

84-87.

Maplestone, C. M. 1903. Further descriptions of the Tertiary Polyzoa

of Victoria. 9. Proceedings of the Royal Society of Victoria (new

series) 16: 140-147.

Michelin, J. H. L. 1840-1847. Iconographie zoophytologique, descrip-

tion par localites et terrains des Polypiers fossiles de France et pays

environnants etc. P.Bertrand: Paris. 348 pp. (1842, pp. 41-72).

Tillier, S. 1977. Les types de bryozoaires de la collection Risso.

Annales du Museum d'Histoire Naturelle de Nice 5: 153-154.

Wass, R.E. 1983. Regeneration of calcification in the Adeonidae

(Bryozoa: Cheilostomata). Memoir of the Association of

Australasian Palaeontologists 1: 305-310.

Wass, R.E. 1991. Intracolonial variation in the cheilostome genera,

Adeona and Adeonellopsis. In: Bigey, F.P., and d'Hondt, J.-L. (eds),

Bryozoaires Actuels et Fossiles: Bryozoa Living and Fossil.

Bulletin de la Societe des Sciences Naturelles de I'Ouest de la

France, Memoire hors serie 1: 523-529.

Memoirs of Museum Victoria 61(2): 135-182 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A review of Australian Conescharellinidae (Bryozoa: Cheilostomata)

Philip E. Bock 1 2 and Patricia L. Cook 2

1 School of Ecology and Environment, Deakin University, Melbourne Campus, Burwood Highway, Burwood, Vic. 3125.

(pbock @ deakin.edu. au)

2 Honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

Abstract Bock, P.E. and Cook, PL. 2004. A review of Australian Conescharellinidae (Bryozoa: Cheilostomata). Memoirs of

Museum Victoria 61(2): 135-182.

The family Conescharellinidae Levinsen is defined and is regarded as comprising seven cheilostome genera

( Conescharellina , Bipora, Trochosodon, Flabellopora, Zeuglopora, Crucescharellina and Ptoboroa). The astogeny of

colonies, that consists of frontally budded zooids with “reversed” orientation, is briefly described and compared between

genera. The morphology of zooids and heterozooids is defined and keys to genera and Australian species are provided.

Taxa that were first described from Australia or from reliable subsequent records are redescribed and illustrated where

possible. Australian specimens that have been identified as non- Australian species, have generally been found to be dis-

tinct and are here redescribed as new species. Some non-Australian records of specimens previously assigned to

Australian species have also been re-examined. These are described and sometimes referred to other taxa. Altogether,

eight previously described species that have not been found in the present material are discussed and 27 taxa are

described from collections, principally from the eastern and southern coasts of Australia and from the Tertiary of Victoria.

Eighteen of these are considered to be new species. Where possible, type or at least topotype material of previously

described species has been examined. Colonies from the collections of Museum Victoria (NMV) and the Natural History

Museum, London (BMNH), have been examined. New species from Australia described here are: Conescharellina cog-

nata, C. ecstasis, C. diffusa, C. obscura, C. stellata, C. plana, C. perculta, C. pustulosa, C. ocellata, C. macgillivrayi, C.

humerus; Trochosodon fecundus, T. asymmetricus, T. diommatus, T. aster, T. anomalus, T. praecox and Crucescharellina

australis. In addition, the New Zealand bryozoan Trochosodon multiarmatus (Gordon, 1989) (not Bipora multiarmata

Maplestone, 1909) is described as Trochosodon gordoni sp. nov.

Keywords Bryozoa, bryozoans, Cheilostomata, Conescharellinidae, fossil, Recent, Australia, new taxa

Introduction

The Bryozoa sorted from dredge samples offshore from south-

eastern and south-western Australia in the past 25 years have

revealed a wide diversity of species, with many apparently

undescribed. The present study is of the family Cones-

charellinidae. The principal collection programs were the Bass

Strait Survey by the Victorian Institute of Marine Science and

the National Museum of Victoria (now Museum Victoria)

(stations with BSS prefix), Museum Victoria’s South-eastern

Australian Slope Survey (SLOPE prefix), and the RV Franklin

1995 shelf survey of the Great Australian Bight including areas

to the west by Dr Y. Bone (University of Adelaide) (GAB pre-

fix). Further collections were made by Gary C.B. Poore on an

expedition with the Western Australian Museum to the

Dampier Archipelago, north-western Australia in 1999 (DA-02

prefix). All these surveys used epibenthic sleds and grabs to

collect sediments. Sampling of sandy sea-floor sediments, fol-

lowed by careful sorting, yields examples from a wide range of

groups adapted to loose sediments (Hayward and Cook 1979,

Bock and Cook, in press). In view of the unexpected diversity

from the scattered survey stations, it is to be expected that yet

more species remain undiscovered.

In addition, an interesting series of partially sorted speci-

mens, labelled in C.M. Maplestone’s hand, from the NMV col-

lection, includes some boxes labelled “S.A.” (i.e. South

Australia) and others with no locality. These last are labelled

with the names of Maplestone’s species from New South

Wales, described by him in 1909 and include specimens of

species that have not been reported again. They do not occur in

any other collections except as “types” in the Australian

Museum, and as “cotypes” that are held in the Natural History

Museum (London) (BMNH), that were originally sent to

London by Maplestone and were registered in 1909. Among

others, these include examples of five species of

Conescharellinidae, labelled as Bipora biarmata, B. multiar-

mata, B. magniarmata (all now referred to Conescharellina ),

Bipora ( ^Trochosodon ) ampulla and Zeuglopora lanceolata.

136

P. E. Bock and P. L. Cook

The Appendix includes full data on station locations and

species occurrences.

A further collection from the Natural History Museum

(London) was originally one of the sediment samples collected

by H.M.S. Challenger. These were stored in the Mineralogy

Department and remained uninvestigated until the 1970s. One

sample, from Challenger stn 185 (11°25'35"S, 144°2'0"E,

249-286 m. near Raine Island, on the outer rim of the Great

Barrier Reef, Cape York, Queensland), was first examined in

1972-73. This sample included foraminiferans and minute

bryozoan colonies, some of which were figured by Cook and

Lagaaij (1976). Cook (1981) later emphasised and illustrated

the striking similarities in size and general appearance of these

two different components of the sample. Further examination

of the numerous bryozoan colonies has revealed that three

species of Trochosodon and one of Crucescharellina are

present. Busk (1884) did not include stn 185 in his Report as its

bryozoan component was undiscovered. Similarly, a specimen

of Crucescharellina sp. from Challenger stn 169 (37°34'0"S,

179°22'0"E, 1295 m, off New Zealand) also remained unre-

ported although a preparation of the single colony is preserved

in the BMNH collection.

Colonies of fossils Conescharellina from the Miocene of

Victoria are also included in this study (see Appendix).

Colonial development. The group of conescharellinids dis-

cussed below construct small colonies that are anchored into

the soft-sediment substratum by one or several cuticular roots.

The colony may develop and grow below the water-sediment

interface or live slightly above the sea-floor. Colonies are

conical or lenticular except in the genus Crucescharellina

which branches into several horizontal arms.

Notes on astogeny of colonies. The astogeny of

“conescharellinids”, like that of “batoporids” (Batoporidae),

has been the subject of a considerable amount of theoretical

discussion that was reviewed by Waters (1919) and Harmer

(1957: 722). Full explanation had to await the description of

concepts of frontal budding (Banta, 1972) and reversed frontal

budding (Cook and Lagaaij, 1976). The type of astogeny gen-

erally known as “reversed frontal budding” occurs in all genera

of Conescharellinidae and Batoporidae but is not unique to

these families. A closely similar form of budding occurs in the

orbicular, flattened colonies of Orbituliporidae. In addition the

rounded and lenticular colonies of the numerous species of the

genus Sphaeropora have a similar type of budding. This genus,

however, is closely related to Celleporaria ; both genera are

referable to the family Lepraliellidae.

Frontal budding was first described by Banta (1972) in

encrusting colonies of Schizoporella ; different sequences were

also illustrated by Cook (1985). Essentially, a frontal bud is

formed by enlargement of an existing hypostegal coelom,

bounded frontally by an intussusceptive expansion of frontal

cuticle. The nutrients necessary to support the growth of the

bud are derived from the pre-existing zooid or zooids, via the

frontal septular pores in the calcified frontal shield. Frontal

buds often have an orientation closely similar to that of the

“parent” zooid but in some mammilliform growths where buds

are derived from more than one “parent” zooid, the orientation

may be random, the orifices occurring with no reference to the

position or direction of the originating zooids.

These forms of frontal budding occur frequently in

ascophoran cheilostomes, particularly among “schizoporellid”

and “celleporid” genera. However, different types of frontal

budding may occur among “anascan” and “cribrimorph”

genera. For example, new branches in the erect “anascan”

Rhabdozoum develop from an elongated frontal bud that arises

from extended calcification surrounding the opesia of a single

zooid (Cook and Bock, 1994). In the cribrimorph Anaskopora,

interzooidal frontal buds arise from uncalcified “windows” in

the chambered pores surrounding each zooid and the resultant

colonies may resemble those of conescharellinids in organisa-

tion (Arnold and Cook, 1997). In Corbulipora, buds arise from

the uncalcified pelmatidia in the spines of the frontal shield

(Bock and Cook, 2001). Encrusting colonies of Trematooecia

and Fatkullina exhibit a reversal of polarity of orifice within

zooids but new buds arise from vertical interior walls

(Grischenko et al, 1998 (1999)). In the Conescharellinidae all

zooid orifices are reversed with respect to the direction of

growth and all zooids are interzooidal frontal buds.

In “reversed frontal budding” the buds arise regularly

between or among the series of frontal septular pores of two or

more neighbouring zooids. The orientation of the primary

orifice is with the “distal” border directed towards the

ancestrular or adapical region. In nearly all the colonies con-

sidered here, most of the frontal shield of a zooid is over-

grown and concealed by the next generation of zooids at the

growing edge (see Cook and Lagaaij, 1976; Pizzaferri and

Braga, 2000). The remaining frontal regions surrounding

the orifices (exposed frontal shields) form the exterior sur-

face of the colony except for the proliferal region. An analo-

gous arrangement occurs in the leaf-like colonies of

Flabellopora and Zeuglopora where the zooids of either

surface interdigitate, forming a superficially “bilaminate” erect

colony (see p. 175).

Mode of life. All living colonies of Conescharellinidae are

known or inferred to be anchored to a substratum by one or

more cuticular roots or extrazooidal rhizoid systems. Generally,

the majority of roots, or the greater part of rhizoid systems, is

located at or near the adapical region of earliest astogeny. There

is evidence from living specimens that metamorphosis of the

larva produces a binary complex consisting of a pair of ances-

trular and root elements (Cook and Chimonides, 1985). Roots

were first described in living colonies of Conescharellina by

Whitelegge (1887); they have also been illustrated by Silen

(1947), Harmer (1957) and Cook (1979, 1981). The mode of

life of small, conescharelliniform and flabelloporiform

colonies, especially early in astogeny, appears to be interstitial,

almost without exception. The minute colonies exist within the

upper centimetres of the sediment surrounded by sand grains

and shell fragments. The colonies are anchored randomly to

minute particles with no particular orientation with regard to

gravity. Colonies are robust and are preserved in the sediment

samples after death. These samples include associations of sev-

eral species, with each species showing colonies at different

growth stages. The function of the roots seems to be purely one

Australian Conescharellinidae (Bryozoa)

137

of anchorage in most genera, not of support, in contrast to the

turgid, extrazooidal rhizoid systems of Sphaeropora and

Parmularia (Cook and Chimonides, 1981, 1985; Brown et al.,

2002). In some species of Flabellopora, however, the more

numerous and larger roots may have a supportive function.

Roots may extend up to 10 mm or more from the colony

surface (Silen, 1947). They are usually thin and delicate, as

illustrated by Cook (1981, pi. A fig. 1) in Trochosodon optatus

Harmer, 1957. They arise from special pores that are formed in

the outer walls of frontally budded, interzooidal kenozooids.

These are quite small and are in communication with the sur-

rounding zooids and kenozooids through small septular pores,

that were described by Levinsen (1909), Livingstone (1925)

and Cook and Lagaaij (1976). In the Conescharellinidae, many

of the root pores that have been reported have a lunate shape,

although others are circular. Both types have been reported to

occur in a single colony; it has been suggested that the circular

pores may be an early ontogenetic stage of the lunate pores

(Harmer, 1957). No colony has been observed here to develop

both kinds of root pore. The lunate shape has given rise to a ter-

minology that has included “lunooecia”, “semilunar pores” and

“semilunar slits”. Root pores are frequent in the earlier stages

of astogeny, occurring amongst both the autozooid orifices and

the avicularian series. Lunate pores often possess a pair of

lateral avicularia, whereas circular pores may be surrounded by

a circlet of avicularia.

The association of a solitary coral, Dunocyathus parasiticus

T. Woods, with colonies of Conescharellina was documented

by Maplestone (1910) in specimens from New South Wales and

South Australia. He considered that the position of the coral,

that usually occupies the entire antapical region of the bryozoan

colony, was evidence of the orientation in life of

Conescharellina , because “the delicate tentacles of the coral

would be crushed” if they rested on the substratum. Harmer

(1957: 724, text-fig. 69) examined a specimen from

Maplestone in the collections of Cambridge Museum. He con-

cluded that Maplestone’s theoretical orientation was probably

correct, as the adapical region of the bryozoan colony was

usually without feeding zooids but was the origin of roots. Of

course, as the actual, interstitial mode of life does not involve a

hard substratum, and as the anchoring, not supportive, nature of

roots, together with the minute size of colonies, is unaffected

by gravity, these theories are of historical interest only. It

appears possible that the coral component of the association

did not live interstitially. A total of 22 bryozoan-coral associ-

ations has been found among the specimens examined here.

Two of these involve Conescharellina multiarmata, seven

C. magniarmata, ten C. cognata, and three C. species (Figs ID,

2F). Although the majority of coral specimens grow from the

antapical surface of the bryozoan colony, three are asymmetri-

cally developed and one occurs at the adapical end of a small

colony. The adjustment of the growth of both organisms seems

to be mutually advantageous. There is no evidence of the

bryozoan occluding the coral, although calcification has devel-

oped laterally, that appears to originate from the bryozoan (Figs

ID). The large avicularian mandibles of C. magniarmata

probably discouraged settlement on any other but the antapical

region but C. multiarmata has only very small avicularia. One

significant correlation may be that all the colonies showing

the association have a “high” conical shape and few or no

antapical cancelli.

Abundance and diversity. The very strong correlation between

the occurrence of minute colonies and fine-grained sediments

was noted by Harmer (1957) and was also emphasised by Cook

(1981). The paucity of earlier records and of numbers of spec-

imens from each sample is almost certainly an effect of collec-

tion bias. Strikingly different observations have resulted where

samples of the sediments themselves have been examined

(Hayward and Cook, 1979; Cook, 1981). The Australian speci-

mens described by Maplestone (1909), from a single dredge

haul in 146 m off New South Wales, also illustrate this differ-

ence, as no fewer than 145 specimens were found, that

belonged to eight nominal species, now known to be referable

to four genera. A total of 79 specimens of Conescharellinidae

were reported by Harmer (1957) from 16 Siboga stations from

the East Indies. These were described as belonging to 18 nom-

inal species and five genera. Silen (1947) also listed 79 speci-

mens, that he referred to nine species and three genera, from

eight stations that overlapped both the Siboga area and the

“Philippines” region reported by Canu and Bassler (1929).

Canu and Bassler included 25 stations with conescharellinids,

identifying 32 nominal species belonging to four genera.

Analysis of sediments from south-eastern Africa revealed 31

specimens belonging to two genera from six stations (Hayward

and Cook 1979). In contrast, Gordon (1985) listed only eight

colonies, belonging to two species, from five stations in the

Kermadec region. Unfortunately, other reports on collections

have not always included consistently the total number of spec-

imens of species from each locality. Gordon (1989) described

six species from 41 stations from southern New Zealand;

Gordon and d’Hondt (1997) also reported six species from 18

New Caledonian stations but gave no estimate of abundance.

Lu (1991) described 24 species referred to Conescharellinidae

from the South China Sea and tabulated estimates of colony

abundance from each of 27 stations. As noted above, Harmer

(1957) was the first to remark on the correlation of sediment

type with the presence of minute, rooted colony forms. Apart

from Gordon and d’Hondt (1997), all the above-mentioned

authors give some indication of sediment type at each collect-

ing station. With hardly any exception, these are of sand, mud,

or ooze, depending on the depths at which they occurred. Cone-

scharelliniform colonies belonging to the Conescharellinidae

are often associated with slope (200 to 1000 m), or even abyssal

depths. Several records given by Harmer (1957), Gordon

(1989) and Gordon and d’Hondt (1997) are from depths in

excess of 1000 m or even 4000 m.

Morphology of structures with characters used in specific

determination

Colony shape and structure. The genera of Conescharellinidae

are characterised to a large extent by shape, that reflects the

arrangement and proportion of autozooids, kenozooids and

avicularia. The principal axis of most colonies extends from the

ancestrular or adapical region to the proliferal or antapical

region. In Conescharellina , autozooids are arranged with their

138

P. E. Bock and P. L. Cook

orifices in apparent radial or in quincuncial series and alternate

frequently with series of avicularia. They often surround a core

of small kenozooids (cancelli). These are budded centrally from

the frontal septular pores on the inner edge of the autozooid

walls and occupy a variable area on the antapical surface. The

successive whorls of autozooids, in fact, always alternate

radially in the antapical direction (quincuncial). The distance

between whorls varies, so that the orifices may appear to form

almost continuous radial chains in colonies with a “high” con-

ical shape, but are obviously quincuncially arranged in colonies

with a “low” cone. In Trochosodon, the conical autozooid

arrangement is very similar but there is little or no central keno-

zooidal core resulting in a more obvious quincuncial arrange-

ment. In Ptoboroa, that does not occur from Australia, the

colonies are stellate with a prominent central root kenozooid.

In Bipora, the radial axes occurring in Conescharellina are

greatly reduced in one dimension; the kenozooidal core is flat-

tened producing an intervening layer of cancelli and a fan-

shaped colony. In Flabellopora and Zeuglopora, the reduction

of all but two of the radial axes is complete. The autozooids are

budded in alternating and interdigitating series with no inter-

vening kenozooids. Colonies are elongated and leaf-like or

occasionally trilobate. In Crucescharellina, it is the adapical to

antapical axis that is completely reduced and the radial axes

elongated, discrete and often branched. This produces a cruci-

form colony with only one series of zooid orifices on one face

and an antapical, “non-zooidal” series on the other (see also

Silen 1947). The colonies of Crucescharellina and trilobate

Flabellopora have the potential to grow far larger than those of

the more conical genera such as Conescharellina, Trochosodon

and Bipora. In Conescharellina, the shape of the cone appears

to be decided early in astogeny and is often apparently

species-specific. For example, the cones of C. biarmata,

C. multiarmata and C. diffusa are usually higher than wide,

whereas those of C. ebumea and C. obscura are wider than

high. The angle of the frontal surface to the vertical axis also

affects the extent and nature of the kenozooidal core. This

forms an interior cone, or cylinder, completely filling the

antapical surface, or lines a shallow concavity (see C. cognata.

Figs 3F, G). Most colonies of the conical genera have a mature

growth stage antapically in that there is no further budding of

autozooids but in that the “cancellated” kenozooidal core is

itself covered by a smooth extrazooidal lamina with small,

intervening avicularia (C. ebumea. Fig. 1G; C. plana. Fig.

10D). These are often derived from the frontal septular pores of

the exposed shields of the most proliferal of the antapical

whorls. Later development of cancelli may include alternating

series of kenozooids and small avicularia.

For some species, examination of large samples has shown

that they may exhibit a wide range of colony shape and of

avicularian size, although in other species variation appears

minor. Particularly in early astogenetic stages, orifices tend to

be quincuncial and the small colonies dome-shaped. In later

astogeny, the orifices may appear radially arranged and the

colonies conical (see C. ecstasis. Figs 5 A, B). Ontogenetic

changes affect both the adapical and antapical regions, with the

development of secondary calcification that obscures zooidal

characteristics. In all colonies, zooid orifice and avicularian

dimensions increase with astogenetic age and there is no

distinct zone of astogenetic repetition. Usually, root pores and

other kenozooids remain almost constant in size, although they

may become surrounded by extrazooidal calcification or by

groups of secondary avicularia, forming specific patterns.

Variation in colony shape and in the astogenetic timing of

“mature” characteristics often reduce the value of past taxo-

nomic descriptions, such as those of Canu and Bassler (1929).

The earliest astogenetic stages have not been observed in

any genus but may be inferred from analogous structures in

other “sand fauna” colonies and from study of minute stages

that infrequently occur in samples. It is inferred that the ances-

trula is anchored to a sand grain or similar object within the

upper layers of sediment, as has been observed in

Conescharellina, Sphaeropora and Parmularia (Cook and

Chimonides, 1981, 1985). The position and orientation of the

first zooidal buds relative to the ancestrula indicate the

eventual mode of growth and structure of the subsequent

colony. For example, Harmer (1957) illustrated very young

colonies of Trochosodon linearis and T. optatus and analysed

their budding patterns. The ancestrula and paired primary buds

formed a radially directed triad, followed by “cycles” (whorls)

of alternating zooids, increasing in size and number.

Kenozooids and small avicularia were budded on the adapical

surface. Almost exactly the same series of astogenetic changes

may be traced in very young colonies of Conescharellina. Cook

(1981: pi. A fig. 6) illustrated a young colony of Cruces-

charellina (as Agalmatozoum sp.) showing a central adapical

area of rhizoid pores (probably overlying the ancestrular

region), with four autozooids forming the earliest stages of a

cruciform colony. Gordon and d’Hondt (1997) illustrated a

slightly older colony with five arms and a central, adapical area

of rhizoid pores and avicularia, very similar in appearance.

Primary orifice. The primary orifice is invariably sinuate, the

sinus defined by a pair of condyles, that may be prominent or

minute. The dimensions of all primary orifices increase with

astogeny but the proportions appear to remain virtually the

same within species. Although the differences among species

are minute and are usually only observable in scanning electron

micrographs, they are constant and correlated and therefore

taxonomically valid. The shape of the sinus varies from round-

ed to subtriangular and is species- specific but it may vary

slightly among populations.

Secondary orifice. Secondary orifices are variable, usually

being confined to raised lappets of lateral peristome. In appar-

ently radial series, these produce an appearance described as

“costulate” (Canu and Bassler, 1929). Sometimes peristomes

are tubular and very prominent marginally (e.g. in Trochosodon

and Ptoboroa) but may be elongated without being prominent

at the colony surface (e.g. Conescharellina plana).

Ovicell. Ovicells are known in two of the genera described

here, Conescharellina and Trochosodon. They are globular,

hyperstomial and often extremely delicately calcified, with an

exposed, frontal entooecium. Ovicells apparently originate

from the small, adapical pore placed close to the border of the

maternal zooid orifice. Gordon (1985) clearly illustrated the

Australian Conescharellinidae (Bryozoa)

139

early stages of ovicell ontogeny in Conescharellina, showing

the ectooecial and entooecial calcified layers developing from

the adapical and antapical sides of this pore respectively.

Colonies of C. diffusa and T. fecundus also show traces of both

layers, associated with the adapical pore (Figs 6B, 17C).

Harmer (1957) described ovicells as peristomial but, although

they are closely associated with the adapical edge of the peris-

tome, they are not derived from it nor do they normally include

any part of it (see Trochosodon praecox ). Ovicells have also

been illustrated by Maplestone (1910) and by Livingstone

(1925b). Silen (1947) illustrated asymmetrically placed ovi-

cells; these may be inferred to occur in T. asymmetricus, from

the asymmetric position of the adapical pore, although ovicells

have not been found (Fig. 19A). Harmer (1957) also described

asymmetrically placed ovicells, and noted their fragility in

some species. Lu (1991, pi. 17 fig. 5C) illustrated part of an

ovicell, in a species he called “ Conescharellina radicata ” from

the South China Sea. No ovicells were mentioned in the

description, that apparently refers to C. radiata Canu and

Bassler (1929: 493, pi. 67 figs 1-3). Although the majority of

recorded ovicells is from the later astogenetic stages of growth,

ovicells have been found very early in astogeny in

Conescharellina africana (see Cook 1966, 1981; Hayward and

Cook, 1979) and in Trochosodon praecox sp. nov. (as

Trochosodon sp. in Cook and Lagaaij, 1976; Cook, 1981).

Colonies of Conescharellina with embryos in their ovicells

were found within a rhizoid and sediment mass belonging to

Parmularia from off Townsville, Queensland, in 1982. These

too, were very fragile, and often became detached when

colonies were moved. The mature colonies occurred together

with very young specimens that were anchored by means of a

minute, turgid ancestrular rhizoid element. Embryos were

released from the mature colonies, that apparently spent their

entire life interstitially (Cook and Chimonides, 1985). The

roots of adult colonies were not turgid or supportive but

anchored the colonies with random orientations with regard to

gravity, within the rhizoid mass of Parmularia. Ovicells, some-

times with embryos in situ, have been found in the present

collection in colonies of Conescharellina plana (stn BSS-167),

C. diffusa (Dampier, N.W. Australia), C. stellata (stn GAB-

019), C. obscura (stn GAB-048), Trochosodon fecundus

(Dampier, N.W. Australia), and T. praecox (Cape York,

Queensland). Study of these examples strongly suggests that in

many cases the basal wall of the ovicell, that is formed by

ectooecium developing from the adapical side of the pore, is

covered by cuticle that is in contact with the frontal shield only

at the point of origin. This explains the ease of detachment of

ovicells in many specimens (see Figs 9H-I). The entooecium

may be ridged and occasionally is porous. The ridges appear to

form pores marginally where the entooecial layer meets the

ectooecium. Ovicells are distinctive as there is no contribution

to their structure by any zooid other than the maternal zooid.

This is a result of the reversed nature of the frontal astogeny.

Avicularia. Avicularia vary considerably in size, distribution

and orientation but may provide some distinguishing character

states among species. The great majority has small, rounded

rostra, that may be minute ( Conescharellina multiarmata ).

Some species have large avicularia; in Crucescharellina and

Zeuglopora these may be spathulate. Those near the orifices

appear to be adventitious in most cases, derived from frontal

septular pores of zooids at the proliferal region, becoming

slightly immersed as the next whorl of zooids is budded. Larger

avicularia appear to be budded interzooidally (Fig. 3D).

Avicularia on the antapical surface are often derived from can-

celli and may alternate with them. Cancelli are kenozooids

originating from septular pores in the antapical part of the

frontal shield in the proliferal region. In some colonies of

Conescharellina that have only just reached a mature asto-

genetic stage, the avicularia follow the radial rows of frontal

septular pores of the last-budded, proliferal whorl, and may not

be accompanied by any cancelli (see Fig. 3F). Avicularia have

a bar that often bears one or more calcareous spinous pro-

cesses (ligulae) on the palatal side. In one species, C. diffusa ,

small spinous processes are present on the non-palatal side of

the bar (Figs 6A, B); in another, C. stellata, the non-palatal area

is sometimes occupied by a thin lamina that may be perforate

(Figs 9B, E). Generally, the palates are without any expanded

cryptocystal margin but this occurs in C. magniarmata (see Fig.

3B). Large, acute avicularia also occur in C. ecstasis (Fig. 4)

and were seen in the living specimens of Conescharellina sp.

from Queensland mentioned above. These last were very

active, snapping shut and holding any surrounding objects in

the sediment. Whether their function is one of stabilization

or defence, or perhaps both, is unknown, as is that of small

avicularia.

Superfamily Conescharellinoidea Levinsen, 1909

D’Hondt (1985: 11) stated that the diagnosis was “confondue

avec celle des Conescharellinidae publiee par Ryland (1982)”.

He included only the family Conescharellinidae Levinsen,

1909.

Family Conescharellinidae Levinsen, 1909

Type genus. Conescharellina d’Orbigny, 1852.

Description. Free-living Cheilostomata attached to small par-

ticles by cuticular roots originating from kenozooidal pores. All

autozooids with reversed frontal budding; ancestrular region

adapical, with root pores and avicularia. Zooids elongated;

frontal wall composed of two parts; “exposed”, surrounding the

primary orifice, and “concealed”, only visible completely in the

antapical region of the colony. Primary orifice usually sinuate,

often with paired condyles, almost terminal, in the centre of the

exposed frontal wall. Avicularia adventitious or interzooidal,

often in patterns among autozooids. Antapical regions often

occupied by kenozooids (cancelli), originally budded from the

septular pores of the frontal walls of zooids of the proliferal

region. Extrazooidal calcification and/or secondary kenozooids

and avicularia often budded from the primary cancelli. Other

avicularia budded directly from the proliferal zooids. Ovicells

hyperstomial, originating from an adapical pore, globular, not

closed by the operculum, entooecium and ectooecium often

delicate and fragile; usually distinct from the peristome but

occasionally associated with it through a foramen.

140

P. E. Bock and P. L. Cook

Remarks. The family includes closely related groups of species

with distinctive colony forms that define genera. The typical

growth pattern of each genus restricts variation of the colony

form; minor differences in astogenetic pattern and zooid mor-

phology may be important in distinguishing species. All known

species have in common: small size (usually less than 10 mm

in maximum dimension), anchorage to sediment particles by

roots arising from special kenozooids, and a strong association

with fine-particle sediments, often from continental slope and

lower slope depths. All species have sinuate primary orifices,

often with condyles. Most species have interzooidal or adven-

titious, frontally budded avicularia, that form patterns among

the autozooid orifices. Special pores, derived from kenozooids,

are the origin of roots. These may be generally distributed or

confined to the regions of earliest astogeny. The family

resembles the Batoporidae and the Orbituliporidae in its

reversed frontal budding pattern but differs in the structure

of the primary orifice and the few known ovicells. It also

resembles the Lekythoporidae, another group including several

closely related genera that have erect branching colonies with a

type of reversed frontal budding (Bock and Cook, 2000). These

last genera, however, have zooidal and ovicellular characters

that show stronger links with the family Celleporidae. Gordon

(1989) has suggested that the Conescharellinidae and

Orbituliporidae should be included with the Lekythoporidae in

a single superfamily This view was not accepted by Bock and

Cook (2000) who noted that Sphaeropora Haswell, 1881, that

also has globular to lenticular colonies formed by reversed

frontal budding, anchored by supportive, turgid, extrazooidal

rhizoids, is closely related to Celleporaria, and is therefore

assignable to the family Lepraliellidae. Reversed frontal

budding itself may not therefore reflect any close systematic

relationships.

Six of the seven genera of Conescharellinidae are repre-

sented in Australia, often by several species. However, the type

species of Conescharellina, C. angustata d’Orbigny, 1852, was

described from the Philippines, and that of Flabellopora,

F. elegans d’Orbigny, 1852, from the China Sea. The type

species of Trochosodon, T. linearis Canu and Bassler, 1927,

occurred from Borneo, and the type species of

Crucescharellina, C. japonica Silen, 1947, from Japan. Two of

the remaining genera, Bipora and Zeuglopora, have Australian

type species: Bipora flabellaris Levinsen, 1909, and

Zeuglopora lanceolata Maplestone, 1909 respectively. The

type species of Ptoboroa, Trochosodon pulchrior Gordon,

1989, occurs from New Zealand.

No attempt has been made here to review or revise the

numerous species of Conescharellina, Trochosodon and

Flabellopora introduced and described by Canu and Bassler

(1929) from the Philippine region. Similarly, the synonymies of

these species, and of further new taxa introduced from the same

region by Silen (1947), from the East Indies by Harmer (1957),

and the South China Sea by Lu (1991), cannot be assessed

without examination of all relevant material. It is possible that

some of the species from eastern and south-eastern Australia

described by Tenison Woods (1880), Whitelegge (1887) and

Maplestone (1909), may be synonymous with some, or part, of

the nominal species described by later authors from the west

Pacific region. Similarly, it is possible that some taxa, intro-

duced here as new, may have been described earlier by these

authors, or even later by Gordon (1989), or Gordon and

d’Hondt (1997) from the New Zealand and New Caledonian

regions.

There is no unequivocal record of a member of the

Conescharellinidae, as defined here, from the European

Tertiary. Conescharellinopsis Labracherie, 1975, described

from the Lower Eocene of Aquitaine, has the type species C.

vigneauxi Labracherie, 1975 (p. 151, pi. 4 figs 4-11). This

species appears to be similar to species of Atactoporidra, as

described by Cook and Lagaaij (1976), with which it was

associated and is not referred to the Conescharellinidae here.

Conescharellina perfecta Accordi (1947), from the Upper

Eocene of northern Italy, has been demonstrated to belong to

the genus Lacrimula (Batoporidae) by Cook and Lagaaij (1976)

and more recently by Zagorsek and Kazmer (2001) who gave a

full synonymy. Lacrimula perfecta also appears to be con-

generic with another north Italian Eocene species,

Conescharellina eocoena Neviani, 1895. Cook and Lagaaij

(1976) suggested that it seems possible that all fossil records of

Conescharellina from western Europe may “prove to belong to

one species complex, attributable to Lacrimula” The

Conescharellinidae therefore seems to have an Indo-

west-Pacific and Australasian distribution only, perhaps

extending from the Eocene (Labracherie and Sigal, 1975), to

the present day.

Notes on the use of the name “Biporidae”. Zagorsek (2001:

558) and Zagorsek and Kazmer (2001: 73) introduced a super-

family “Biporidae Gregory, 1893” but no mention was made of

the genus Bipora Whitelegge, 1887. The superfamily was

described to include the family “Batoporoidea” (sic) Neviani,

1901 and the genera Lacrimula Cook, 1966 and Orbitulipora

Stoliczka, 1862. Neviani (1901) had, however, included in his

family “Batoporideae” [sic] only the genera Batopora Reuss

(for B. rosula) and Conescharellina d’Orbigny (for C. conica, a

manuscript name, almost certainly referable to Lacrimula

perfecta ; see Cook and Lagaaij, 1976 for discussion).

Gregory (1893: 223) suggested a classification of

Cheilostomata that included five Suborders. Two of these

included the “ascophorine” forms and consisted of the

Suborders Schizothyriata and Holothyriata. Gregory’s treat-

ment of families and subfamilies was not consistent but among

the Schizothyriata the family Schizoporellidae and subfamily

Schizoporellinae were provided (p. 239) with an informal des-

ignation of Schizoporella as type genus, and a reference to its

diagnosis by Hincks (1880). In a similar manner, the type genus

Schizoretepora was designated in a footnote for the family

Schizoreteporinae. The type genus Schismoporina was also

designated for the subfamily Schismoporineae in another foot-

note. The treatment of the subfamily Biporineae was com-

pletely different. No generic names were included but the

description given was “Schizoporellidae with a patelliform

unilaminate zoarium, with vibracularia systematically

arranged”. This is a parallel of the description of a subfamily of

Microporidae included in Gregory’s suborder Athyriata, called

the Selenarinae, similarly described as “Microporidae with

Australian Conescharellinidae (Bryozoa)

141

patelliform zoaria and vibracularia systematically arranged”.

The Biporineae may even have been introduced to provide a

form of symmetrical concept between the Athyriata and

Schizothyriata. Presumably, Gregory had in mind some lunu-

litiform ascophoran genus or genera that would be included in

his Biporineae but he did not mention the subfamily again, or

describe Bipora, or any other genus as belonging to it. In

addition, although Whitelegge’s (1887) paper and its reprint

(1888) were both listed in Gregory’s bibliography (on p. 274),

no mention of either was made anywhere in his text. Both

Whitelegge and Jelly (1889), whose Synonymic Catalogue was

also listed by Gregory in his bibliography, gave

Conescharellina in the synonymies of several species assigned

to Bipora. In fact, Jelly (1889: 20) referred to Whitelegge’s

paper under her entry for B. umbonata (Has well), and again (on

p. 64) under Conescharellina cancellata and C. elegans, where

Bipora was given in synonymy.

Gregory must therefore have been aware that other, earlier

authors had described a relationship between the two genera.

Any Conescharellinidae were, however, tacitly excluded from

the subfamily Biporineae by Gregory (1893: 225, 251), as the

genus Conescharellina was listed as belonging to the family

Celleporidae, a member of his suborder Holothyriata. Gregory

regarded Conescharellina as a senior synonym of Batopora and

described one species from the British Eocene, Cones-

charellina clithridiata, that is, in fact, referable to the

Batoporidae. This species was illustrated as Batopora by Cook

and Lagaaij (1976, pi. 2 fig. 1, pi. 5 fig. 5) and by Cook (1981,

pi. B fig. 4). One other species, B. glandiformis, was

erroneously referred to the cyclostome genus Heteropora by

Gregory (1893) but was briefly discussed and assigned to

Batopora by Cheetham (1966) and subsequently was assigned

to Atactoporidra by Cook and Lagaaij (1976). Waters (1904:

96) made the illuminating remark, with reference to Gregory’s

“undoubted abilities” that “sometimes angel visits stir up all

that has been done without establishing order” and “classifi-

cation has been left in a much more hopeless condition than it

was before . . . made by those who have swooped down on the

Bryozoa for a short visit”.

It seems that Biporineae is not a synonym of

Conescharellinoidea and there is no necessity to use any

emended suprafamilial name such as “Biporidae Gregory” to

include the “Batoporoidea” as used by Zagorsek and Kazmer

(2001), or the Conescharellinidae as used by Levinsen (1909).

“Biporinae” Maplestone (1910) was an informal usage of a

name and is a junior “synonym” of Levinsen’s (1909) name

Conescharellinidae. As Conescharellinidae has been in

common usage, the rule of priority can be ignored, as in ICZN

Rule 35.5.

Key to genera of Conescharellinidae

1. Colonies conical with circular cross section, or stellate 2

— Colonies not as above 4

2. Autozooids and avicularia frequently in antapically direct-

ed, alternating series. Autozooids not very prominent mar-

ginally; kenozooids forming a central core or as antapical

layers Conescharellina

— Colonies stellate, without central core of kenozooids, auto-

zooid orifices often arranged quincuncially, marginal

zooids prominent; avicularia often absent 3

3. Colonies with elongated peripheral peristomes; antapical

avicularia and cancelli rare Trochosodon

— Colonies with prominent central root kenozooid Ptoboroa

4. Colonies compressed laterally in one plane 5

— Colonies compressed antapically in one plane, often

branching Crucescharellina

5. Colonies with a laterally compressed cone, becoming fan-

shaped, zooids on each face separated by a narrow band of

kenozooids Bipora

— Colonies leaf-like, with no intervening kenozooids

between two interdigitating, frontally budded series of

zooids 6

6. Lateral margins of colonies serrated, often with groups of

prominent zooids or enlarged avicularia Zeuglopora

— Lateral margins not serrated, colonies sometimes trilobate

Flabellopora

Key to Australian species of Conescharellina

1. Avicularian rostra acute, longer than orifice 2

— Avicularian rostra rounded, smaller than orifice 6

2. Colonies domed, height and / or width > 3mm 3

— Colonies small, distinctly conical, higher than wide.

Avicularia paired, lateral oral, orientated laterally

C. biarmata

3. Avicularia lateral oral, single 4

— Avicularia lateral oral paired, orientated laterally and

adapically C. ecstasis

4. Avicularia randomly orientated C. angulopora

— Avicularia orientated laterally and adapically 5

5. Colonies domed, width and height subequal. Solid antapic-

ally. Avicularia with lateral cryptocyst lamina, and 3 large

ligulae C. magniarmata

— Colonies becoming very large, wider than high, diameter

10 mm. Antapical surface hollow, covered by cancelli . . .

C. cognata

6. Colonies large, height and / or width > 4mm 7

— Colonies small, height and / or width < 3mm 10

7. Root pores circular, surrounded by avicularia 8

— Root pores lunate, with paired avicularia 9

8. Colonies with patent orifices; avicularia lateral and ada-

pical, visible on antapical surface of marginal zooids ....

C. eburnea

— Orifices at the base of a long peristome, that is not raised

at the colony surface; avicularia minute, scattered and

paired, adapical C. plana

9. Colonies conical, higher than wide, height up to 5 mm;

avicularia and root pores in series alternating with orifices;

root pores without small avicularia; rostra with non-palatal

spinules C. diffusa

— Colonies flat, up to 13 mm diameter, or small and

discoidal; avicularia scattered or single, antapical and

peristomial; root pores with paired avicularia

C. obscura

10. Colonies with lunate root pores with paired avicularia 11

142

P. E. Bock and P. L. Cook

— Colonies with circular root pores surrounded by avicularia

11. Colonies domed, orifices in radial series alternating with

avicularia, root pores adapical (fossil) C. aff. diffusa

— Colonies conical, higher than wide, orifices with minute

lateral and antapical avicularia C. multiarmata

— Colonies stellate, marginal peristomes bilabiate or spout-

like; avicularia with non-palatal lamina C. stellata

— Colonies with pustular calcification adapically and anta-

pically; avicularia minute, one peristomial and antapical,

others scattered C. pustulosa

12. Colonies with prominent spout-like marginal peristomes

and numerous pairs of avicularia, some visible on the

antapical surface marginally C. perculta

— Colonies not as above 13

13. Colonies slightly domed , or raised centrally; orifices with

a long subtriangular sinus, peristomes raised laterally;

avicularia paired lateral-oral, visible on the antapical sur-

face of marginal zooids; cancelli absent . ... C. ocellata

— Colonies fairly flat, with bilabiate marginal peristomes;

orifice with a small rounded sinus; avicularia rare, anta-

pical surface with large cancelli C. macgillivrayi

— Colonies slightly raised centrally; orifices with a rounded

sinus and laterally raised peristome, with paired lateral

avicularia that form prominent “shoulders” on marginal

zooids C. humerus

Key to Australian species of Trochosodon

1. Colonies large, diameter 3-4.7 mm 2

— Colonies smaller 3

2. Colonies fairly flat, domed centrally, with numerous

tubular marginal peristomes T. ampulla

— Colonies lenticular, with prominent radial rows of peris-

tomes with paired avicularia on the antapical surface; root

pores lunate T. diommatus

3. Colony diameter 2-3 mm 4

— Colony diameter <2 mm 5

4. Colonies with bilabiate marginal peristomes, orifices quin-

cuncial with wide, shallow sinus; ovicells symmetrical,

root pores lunate T. fecundus

— Colonies with short, tubular peristomes, orifices radial,

sinus rounded; adapical pores asymmetric, root pores

circular T. asymmetricus

5. Colonies conical, higher than wide; zooid peristomes

prominent and curved; root pores rare, lunate T. anomalus

— Colonies as wide as high, or wider 6

6. Colonies very small, fairly flat, stellate; peristomes tubular,

with paired lateral avicularia; root pores lunate . T. aster

— Colonies minute, with an antapical dome of mamillate

calcification; peristomes tubular, with paired lateral avic-

ularia; ovicells small, robust, symmetrical; root pores

rounded T. praecox

Conescharellina d’Orbigny 1852

Conescarellina [sic] d’Orbigny 1852: 447.

Conescharellina. — Canu and Bassler, 1917. — Waters, 1919: 93. —

Canu and Bassler, 1929: 480. — Silen, 1947: 33. — Harmer, 1957:

726.— Gordon, 1989: 81.

Type species. Conescharellina angustata d’Orbigny, 1852, sub-

sequent designation by Waters, 1919: 93. [Canu and Bassler

(1917) had earlier incorrectly indicated C. cancellata (Busk,

1884), see Harmer (1957: 726)]. The mis-spelling of the name

as Conescarellina occurs only in the genus heading of

d’Orbigny (1852: 447): all other spellings of the name are as

Conescharellina. Conescharellina angustata was included in

Batopora by Reuss (1867: 224).

Description. Colony conical, with autozooids appearing to be

in radial series, either placed in rows alternating with avicular-

ia, or in quincunx with intervening avicularia. Cuticular roots

arise from circular or crescentic skeletal pores, concentrated in

the adapical region in some species. Orifices with an antapical

sinus, often with raised lateral peristomes. Avicularia adventi-

tious and interzooidal, usually budded in distinct patterns, with

acute or rounded mandibles, slung on a bar, that often has one

or more palatal ligulae. Ovicells hyperstomial, prominent,

derived from an adapical pore, with thinly calcified ectooecium

and entooecium. Central part of colony cone occupied by a

core of small kenozooids (cancelli), often accompanied by

avicularia, that may cover the antapical surface late in as

togeny.

Remarks. C. angustata was described by d’Orbigny (1852: 447,

pi. 714 figs 14-16) from the Philippine island of Basilan

(approx. 6°50'N, 122°E, in the Celebes Sea). The figured

colony (fig. 15) was an elongated cone with 8-9 apparently

radial series of zooids forming costules. The orifices are raised,

circular-to-oval, each with an asymmetrically arranged pair of

pores adapically, and a single series of “special” pores alternat-

ing with the zooid orifices in a radial depression. D’Orbigny

noted that the orifices were in quincunx, and figured the anta-

pical surface (fig. 16) showing five alternating series of prolif-

eral and subproliferal zooids, with no central cancelli.

D’Orbigny noted this particularly, comparing it with the antapi-

cal side of C. dilatata (see below). In view of the relatively

large size and possible maturity of the type colony (height

approximately 2.5 mm), it is unusual in Conescharellina for

cancelli to be absent. In fact, this is characteristic of

Trochosodon.

A scanning electron micrograph of the putative type speci-

men, from the Museum Nationale d’Histoire Naturelle, Paris,

has been provided by Drs D.P. Gordon and P.D. Taylor. It

resembles d’Orbigny’s figure 15 in its elongated conical shape

and radial series of zooid orifices. The adapical region is less

regular than the figure, and there are fewer zooid series but this

may be the result of damage.The rounded secondary orifices,

almost all of which have an adapical pore, are similar to those

figured but the additional pores shown near the orifices are not

present. D’Orbigny figured a radial series of pores in the

depression between zooid series, that were lateral to the ada-

pical edge of the adjacent orifices. Avicularia occupy a similar

position in the micrograph of the specimen but are far larger

and twice as frequent. These avicularia are small and rounded

with a delicate, simple bar. The orifices of fig. 15 are second-

ary and show a slightly raised peristomial rim; a few of those in

the micrograph also show a sunken primary orifice with a

rounded sinus. The specimen of C. angustata resembles some

Australian Conescharellinidae (Bryozoa)

143

of the more elongated colonies of C. diffusa. These differ in

their proportionally larger avicularia and the presence of

numerous lunate root pores, that are absent from C. angustata.

Waters (1905: 9, pi. 1 fig. 7) examined the type material of

C. angustata and gave a figure of the specimen from Basilan.

This does not show the entire colony but only a formalised rep-

resentation of four oval orifices and a single antapical avicular-

ium. Later, Waters (1919: 93) indicated C. angustata as type

species of Conescharellina, without comment. He also (1921:

419, pi. 30 fig. 18) figured but did not describe a speci-

men “from China, sent to me thus named by Jullien” as

C. angustata. This colony was also conical and very elongated,

with raised “costules” of radial rows of oval secondary orifices

separated by adapically placed pores. One elongated sinuate,

perhaps primary, orifice was figured, and small scattered pores

among the orifices may have represented avicularia. Unlike the

type specimen, the adapical region was occupied by extra-

zooidal or kenozooidal calcification. The figure is otherwise

similar to that of d’Orbigny’s C. angustata, with “costules” of

secondary orifices that are more elongated and with “pores”

less regularly spaced. Harmer (1957) was doubtful that the

three Siboga collection specimens from East Java, that he

nevertheless assigned to C. angustata, were identical with

d’Orbigny’s species. These colonies were not elongated; the

primary orifices were patent, with little or no peristome, and

were relatively wide with a rounded sinus. These specimens do

not appear to be conspecific with the type specimen.

D’Orbigny (1852: 447) also introduced but did not figure

Conescharellina dilatata from “Manille et detroit de Malacca”

[sic]. It differed from C. angustata in its greater width (“ensem-

ble plus large”) and in the presence of “un espace poreux” (pre-

sumably of cancelli) on the antapical surface. Waters (1905: 9,

pi. 1 fig. 6) gave a semidiagrammatical figure of two zooid ori-

fices from a specimen of C. dilatata from d’Orbigny’s collec-

tion from Manila. There were “two species in the tube” but he

did not indicate which of these he regarded as C. dilatata. As

before, only examination of the type material can elucidate

fully the characters and relationships of this species. However,

it is obvious that d’Orbigny’s C. angustata is closely similar to,

and congeneric with, many of the other taxa subsequently

referred to Conescharellina but description of its specific

characters must await examination of the type specimen.

Species recorded from Australia but not recognised in the

material examined here

Conescharellina philippinensis (Busk, 18540 and C. can-

cellata (Busk, 1854)

Lunulites philippinensis Busk, 1854 and L. cancellata Busk,

1854 were described and figured by Busk (1854: 101, pi. 113

figs 1-3 and 4-7 respectively) from the Philippines. They are

obviously species of Conescharellina but the characters

described and figured are not sufficiently clear to allow their

recognition and identification with other material with any cer-

tainty. It has been possible to examine specimens from the

“type suites” of L. philippinensis and L. cancellata but it must

be emphasised that until all Busk’s specimen suites have been

revised, little may be concluded as to the nature and the

identity of specimens later reported under these names.

According to Waters (1921: 419), Busk’s specimens in the

British Museum collection confused both species and included

at least two additional species. Harmer (1957: 742) did not,

however, agree with all Waters’ conclusions. The “type” slide

of L. philippinensis (BMNH 1854.11.15. 150) originally

included five colonies. Two of these have been lost in the past;

one was remounted as an additional slide and labelled in

Kirkpatrick’s hand. This very worn, separated colony may be

the original of Busk’s figure (1854: pi. 113 fig. 2). The other

specimens do not appear to have been figured, although all

three seem to be conspecific. The specimens are all worn and

show little detail. Two are flat and are less than 2 mm in

diameter. They include approximately five quincuncial genera-

tions of zooids and each whorl has nine to ten zooid orifices.

The marginal peristomes are slightly prominent and tubular; the

primary orifices cannot be seen. Small rounded pores, inferred

to have been avicularia, are interspersed randomly among the

zooid orifices and the antapical surface has a central cancellate

area. In both the larger colonies, the centre of the adapical sur-

face has two prominent rounded “bosses”, that are illustrated in

Busk’s pi. 113, fig. 2. It is not possible to recognise this species,

either among those described from the Philippines by Canu and

Bassler (1929) or in the Australian material examined here. The

“type” slide of L. cancellata (BMNH 1854.11.15.151) includes

four specimens that are in a better state of preservation. The

originals of Busk’s pi. 113, figs 4-7 are recognisable; an addi-

tional large, worn, unfigured colony, that does not seem to be

conspecific, is present (Brown, 1958: 82). The figured colonies

are distinctly domed; the largest, that is less than 2 mm in

diameter, includes approximately seven quincuncial zooid gen-

erations and nine to ten zooids per whorl. The peristomes are

only slightly raised and circular; the primary orifices are visible

and are rounded with a short, wide, almost semicircular sinus.

Traces of an adapical pore are present in a few zooids. Small

oval avicularia, with a delicate, simple bar, occur some-

what irregularly among the zooid orifices. No root pores are

visible; the antapical surface has a central cancellate area. This

species was apparently not among the other Philippine forms

described by Canu and Bassler (1929) and has certainly

not been recognised among the Australian specimens

examined here.

Waters’ (1921) account of L. cancellata is not at all clear. He

remarked “specimens from Busk’s own collection so named are

C. angustata d’Orb.” Harmer (1957: 742), when discussing

C. crassa, seems to have mistaken Waters’ (1921) reference to

C. angustata, as describing part of the type material of

L. cancellata. A specimen in Busk’s collection from the Sea of

Japan (BMNH 1899.7.1.1276 labelled Lunularia cancellata ) is

narrowly conical, with seven to eight radial series of orifices

and five to six zooid whorls. Zig-zag series of small oval avic-

ularia alternate with the orifice series; these also occur on the

antapical surface. The colony somewhat resembles d’Orbigny’s

C. angustata and may be the one mentioned by Waters. Waters

(1921) also stated that the specimens he described “from New

South Wales” (i.e. in 1887) “then called cancellata by me are

seen to be philippinensis .” Both names have been used for

144

P. E. Bock and P. L. Cook

several Australian records; references to Recent material

assigned to these species are discussed below under C. diffusa

and C. obscura.

It is unfortunate that little of the previously described

Tertiary material is extant. Various combinations and spellings

of C. cancellata having been quoted, particularly for specimens

from the Tertiary, by Waters (1881; 1882a; 1882b) and by

MacGillivray (1895). Maplestone (1904) tabulated several

additional fossil localities, including Campbells Point, Mitchell

River and Lake Gnotuk, together with his own observations of

material from Mornington. Unfortunately, Maplestone’s

specimens are not extant, and therefore his concept of fossil

B. cancellata and B. philippinensis must remain unknown. He

also listed B. elegans Waters ( -Bipora flabellaris ), from

Jimmys Point, that has otherwise not been reported as a fossil,

and therefore seems unlikely to be this species. MacGillivray

(1895: 89, pi. 12 fig.2) reported “ Bipora philippinensis''' from

the Tertiary of Schnapper Point and Muddy Creek, Victoria. His

specimen from Muddy Creek is extant (NMV P27728). It is a

fairly flat colony, with quincuncial zooid orifices with a small

sinus and scattered avicularia. The antapical surface has a large

cancellated area. This specimen appears to be referable to the

fossil species described here as Conescharellina macgillivrayi

sp. nov. Waters (1881) mentioned Recent specimens of

B. cancellata from Torres Strait but no fossil examples.

However, he appears to have believed that he had specimens

from the “Curdies Creek” locality, as he mentioned them

(Waters, 1882a) in connection with the “better preserved”

material he had from Bairnsdale, Victoria (Waters, 1882b: 512,

pi. 22 figs 10, 11, as Lunulites cancellatus), that he figured

showing the orifice and surrounding avicularia. These illustra-

tions suggest that the species may also have been

Conescharellina macgillivrayi . Whitelegge (1887: 341) listed

C. cancellata, remarking that he had several fossil examples

from Muddy Creek, Victoria, that might be identical with the

species recorded by Waters (1882b) but that in C. cancellata

and C. philippinensis “the identity can only be definitely settled

by comparison with the types”. Bipora cancellata was

recorded by MacGillivray (1895: 89, pi. 12 fig. 1) from

Bairnsdale; he noted that it was often difficult to distinguish it

from B. philippinensis. His specimen (NMV P22727) is a

conical colony with orifices arranged in radial series. The pri-

mary orifice has a fairly wide, rounded sinus and is flanked

antapically by a pair of small, rounded avicularia. The anta-

pical surface has very few cancelli. His specimen resembles

others from Bairnsdale, and is discussed here under

Maplestone’s Recent colonies of C. diffusa. Colonies from the

Miocene of Victoria and South Australia are numerous and

diverse; four species, C. ocellata, C. macgillivrayi , C. humerus

and C. aff. diffusa are described below.

The ovicells of C. cancellata were mentioned, in passing, by

Levinsen (1909: 310, pi. 23 figs 8a, b), who illustrated small,

globular ovicells with marginal pores and an oval zooid orifice

with an adapical pore. Three small rounded avicularia sur-

rounded the ovicelled zooid orifice. Levinsen did not give any

details of the provenance of the specimens illustrated and the

information given is insufficient for identification of the

species.

Conescharellina angulop ora (Tenison Woods, 1880)

Lunulites angulopora Tenison Woods, 1880: 7, pi. 1 figs 3a-c.

? Lunulites conica Haswell, 1881: 42, pi. 3 figs 7, 8.

1 Conescharellina incisa Hincks, 1881: 127 (sep. p. 68), pi. 4 figs

1-3.

? Bipora angulopora. — Whitelegge, 1887 (1888): 18.

not ? Lunulites angulopora. — MacGillivray, 1895: 46, pi. 8 fig. 1

(= Selenariopsis macgillivrayi Bock and Cook, 1996).

? Conescharellina angulopora. — Levinsen, 1909: 311, pi. 23 figs

7a-f.

not Conescharellina angulopora. — Gordon, 1985: 173, figs 20-23;

Gordon, 1989: 81, pi. 48B (see C. cognata).

Remarks. Search for type material of Tenison Woods has been

unsuccessful; consequently the characters of this species

remain somewhat doubtful. The colony was figured as a

distinct cone and the autozooids and avicularia occurred in

apparent alternating radial series. However, the description of

the orifice as “divided into two portions; one half triangular

constricted in the middle; the other semicircular”, taken

together with the illustration, indicates that Tenison Woods had

confused the avicularia with the secondary orifices. The illus-

tration shows at least one triangular avicularium accompanied

by a typical lunate root pore that he did not recognise as distinct

structures. His later remark “the cells are obliquely placed;

sometimes in contrary directions alternately”, also appears to

refer to avicularia, that have been described in other material

assigned to this species as having alternating orientations. The

description of “the vibracular pores” as “long and narrow, and

in a depressed area” and the illustration, showing irregularly

ovoid openings, apparently refers, in fact, to the secondary

autozooid orifices. Waters (1887: 199), describing specimens

he assigned to C. incisa (Hincks), remarked “This may be

Lunulites angulopora T. Woods, but apparently the avicularia

were mistaken for zooecial cells, and the zooecia for

vibracula”. Tenison Woods had only two specimens from

Port Stephens, New South Wales, that he noted were “worn”;

his type material has not been found. It seems unlikely that his

species is recognisable. Livingstone (1924) regarded C. conica

Haswell (1881), Lunulites incisa Hincks (1881, 1892), Bipora

biarmata, and B. magniarmata Maplestone (1909), all as

junior synonyms of L. angulopora Tenison Woods (1880). Both

Livingstone (1928) and Hageman et al. (1996) reported

C. angulopora from South Australia, and specimens labelled

Bipora angulopora occur in Maplestone’s collection from this

area. These specimens belong to at least two other taxa (see

C. cognata and C. diffusa ) but specimens in Maplestone’s

collection (NMV), inferred to have been from New South

Wales, are described below as C. species (C. angulopora

sensu Maplestone not T. Woods). Haswell (1881) gave an

illustration of his C. conica showing the orifices “upside

down”, so that the apparent antapical primary sinuses are in

fact, adapical parts of the peristome. He did not label

his types or conserve entire specimens (Livingstone, 1924).

Hincks’ (1881) type specimens of L. incisa are not avail-

able, so that the identity of these species and their pos-

sible synonyms remains in doubt, in spite of the superficial

similarity of his figure of L. incisa with that of C. conica (see

below).

Australian Conescharellinidae (Bryozoa)

145

Conescharellina crassa (Tenison Woods, 1880)

Lunulites ( Cupularia ) crassa Tenison Woods, 1880: 5, pi. 1 figs

la-c.

Bipora crassa. — Whitelegge, 1887: 343 [reprinted 1888: 18].

Conescharellina crassa. — Livingstone, 1924: 212. — Livingstone,

1925: 301, pi. 46 figs 1-5, text-fig. 1.

Description, (modified in part from Livingstone’s 1925

account). Colony a large, shallow cone, maximum diameter 10

mm, height 5 mm. Zooids arranged in quincunx. Primary ori-

fice elongated, with a fairly narrow but rounded sinus; lateral

peristomes raised, marginal peristomes prominent. Adapical

pore (“special pore”) large, on the edge of the peristome, form-

ing a tube. Root pores rounded, not lunate. Avicularia small and

rounded; with a bar and one ligula; one (possibly the “vibracu-

lar pore”) placed adapically to the orifice; others minute, some-

times paired, antapical and lateral, or irregularly scattered

among orifices, rounded. Antapical surface “spongy”, (inferred

to have consisted of cancelli), and “solid”.

Remarks. Tenison Woods (1880) mentioned “about a dozen

specimens” from Cape Three Points and Port Stephens, New

South Wales. They were collected from depths of approxi-

mately 130-150 metres. Whitelegge (1887, 1888) examined

these, the type specimens of C. crassa, that were then in the

Macleay Museum, Sydney. He remarked on the raised lateral

peristomes, the primary orifice, the subcircular avicularian

mandibles and the large, antapically placed pore (inferred by

Harmer (1957) to have been an avicularium) but did not

mention the antapical surface or the form of root pore.

Whitelegge (1887) noted that Tenison Woods’ figure was “the

first published figure which exhibits the form of the true

operculum-bearing aperture”. This was narrow and elongated,

with a rounded sinus. Livingstone (1925) also examined the

type specimens, and other colonies from New South Wales. He

redescribed C. crassa, noting that some of the “vibracular

pores” were “filament pores”, i.e. root pores. These were

rounded, not lunate. The raised lateral peristomes obscured the

orifice, with its fairly elongated, narrow sinus. The adapical

pore (“special pore”) was figured on the edge of the peristome,

forming a tube, very similar to the pore illustrated here in

C. multiarmata (Fig. 2D). Livingstone (1925) was the first to

suggest that “lunoecia” and “filament pores” had the same

function.

Harmer (1957: 740, pi. 48 figs 1-6, text-figs 70, I, 73)

described specimens from West Timor, the Arafura Sea and

Holothuria Bank (north-west Australia) as C. crassa. The

colonies resembled those reported from eastern Australia in

size and shape, having a concave antapical surface lined by

cancelli, and bordered by prominent zooids; they were, how-

ever, not solid antapically. The orifices had an elongated sinus

but were arranged in quincunx, not in apparently radial series.

Harmer noted that both the “vibracular pore” of Tenison Woods

(1880) and the “filament pore” of Livingstone (1925) might

have been avicularia. A small adapical pore (“proximal pore”)

was sometimes present in his material but the circular root

pores were found scattered among the orifices, not directly

associated with the peristomes. Three of Harmer’s preparations

have been examined (BMNH Siboga stn 59, West Timor, 390

m, 1964.3.2.8 part, and from Murray Island, Torres Strait, from

Haddon, 1890.3.24.17). The latter was mentioned by

Kirkpatrick (1890) who described the operculum as “broadly

pyriform”. They are large colonies, ranging from 10 to 12 mm

in diameter but are all very worn. Only one primary orifice is

clearly visible: it is wide, with a rounded sinus, unlike Tenison

Woods’ figure. Otherwise, Harmer’s C. crassa resembles the

original description but only examination of Tenison Woods’

type material, and comparison with that seen by Haswell from

Queensland, can decide if any of them are conspecific.

Livingstone (1925: fig. 1) described ovicells in the “small-

est specimen” of a group of colonies of C. crassa from north-

east of Port Jackson, at 137-146 m. These were “bean-shaped”,

wider than long, flattened frontally. They appear to have had an

ectooecial rim bordered by “a row of elongated pores”. The

figure of the ovicells depicts these pores as minute and cer-

tainly not elongated. Curiously, Livingstone (1925: 303) noted

the absence of “special pores” in the smallest colony that bore

the ovicells. His illustration (pi. 46 fig. 3) leaves no doubt that

the adapical pore is depicted. As its presence is a necessary part

of ovicell development, his observation requires explanation.

The illustration of the ovicells in C. crassa given by

Livingstone (1925: Fig. 1) is remarkably similar to that of the

ovicells of “Batopora pulchrior ” Gordon (1989: 81, pis 47F, G,

48A) from very deep water (914-3347 m) off New Zealand.

The ovicells of B. pulchrior lack marginal pores. B. pulchrior

is the type species of Ptoboroa Gordon and d’Hondt (1997), a

genus that appears to have closer links with Trochosodon than

with Batopora (see below).

Although specimens of C. crassa should be recognisable

from the descriptions of authors mentioned above, no colony in

the collections examined here appears to be assignable to this

species. Two species described here with large, relatively

flattened colonies are C. cognata and C. obscura.

Conescharellina depressa Haswell, 1881

Conescharellina ? depressa Haswell, 1881: 41, pi. 3 fig. 4.

Conescharellina depressa. — Livingstone, 1924: 212.

Description. Colony forming a low cone, concave antapically,

with prominent marginal zooids. Orifices with raised peris-

tomes, arranged in apparent radial series, alternating with large

avicularia with elongated, rounded, or semicircular mandibles.

Lunate root pores occur among the avicularia. Antapical

cancelli lining the concave surface (based on Livingstone,

1924).

Remarks. C. depressa was originally described from Port

Denison, Queensland. Whitelegge (1887) mentioned “5 or 6

specimens” but these do not seem to have been part of the type

material, although he mentioned no locality other than

Haswell’s. Livingstone (1924: 205) noted that Whitelegge had

informed him that Haswell did not label his type specimens,

and that he himself had seen only small fragments of each

species. The orifice was described with a sinus “about half the

diameter of the mouth; or ovate with a sub-triangular denticle

on each side near the base”. Harmer (1957: 743) regarded

C. depressa it as “nearly allied” to his C. crassa but the wide

146

P. E. Bock and P. L. Cook

primary orifice and lunate root pores suggest that it may be dis-

tinct. Until type material can be examined, the characters of this

species must remain uncertain; no colony resembling it has

been found in the present material.

Conescharellina conica Haswell, 1881

Conescharellina conica Haswell, 1881: 43, pi. 3 figs 7, 8.

ILunulites incisa Hincks, 1881: 68 (sep. p. 127), pi. 4 figs 1-3.

Description. Colony conical, distinctly higher than wide. Zooid

orifices apparently in radial rows, peristome raised laterally.

Primary orifice elongated, with a minute sinus. Avicularia

large, in raised rows, mandibles acute, orientated laterodistally,

in both directions; bar with well developed ligula.

Remarks. Haswell’s material was from Holborn Island,

Queensland, from 37 m, and Hincks’ specimens were from

Bass Strait (from less than 73 m) but their illustrations of the

primary orifices and avicularia appear to be closely similar.

Haswell’s figure, however, does not illustrate the primary ori-

fice, as the antapical side is uppermost. Hincks’ figure has a

reversed orientation and the apparent similarity in orifice shape

is accidental. Hincks (1892: 331, sep. p. 194), however, regard-

ed his species as synonymous with C. conica Whitelegge

(1887). Livingstone (1924) placed both species in the syn-

onymy of Conescharellina angulopora, see above. Without

examination of type material, the relationships of these three

nominal taxa remains uncertain.

Descriptions of species present in the material examined

Conescharellina sp.

Figures 1A-D.

Bipora angulopora. — Maplestone 1909: 268 (not Conescharellina

angulopora Tenison Woods).

Specimens examined. NMVF98977, F98978, F101878. 13 specimens,

all somewhat worn, labelled “ Bipora angulopora ” are present in the

Maplestone material, inferred to be from New South Wales.

Description. Colonies conical, slightly wider than high; zooid

orifices in apparent radial series, elongated, with prominent

condyles and a subtriangular sinus. Peristomes deep, worn, not

raised, with an adapical pore present. Avicularia very elongated

and narrow, usually occurring antapically and laterally, some-

times paired, usually alternating with zooid orifices, orientation

lateral or random, bar with a ligula. Small lunate root pores

adapical, with paired avicularia. Antapical surface solid, with

rows of small avicularia but no cancelli.

Height of colony up to 2.1 mm, width 2.6 mm, number of

whorls 9-10, number of zooids per whorl 10-12.

Remarks. Maplestone (1909: 268) listed “ Bipora angulopora ”

from the “Miner” dredgings from New South Wales but did not

describe his specimens. His labelled material agrees in part

with Livingstone’s (1924) description of some colonies of

C. angulopora. These were conical and had elongated avicular-

ia orientated in several directions. The colonies examined are

very worn, have adapical lunate root pores; the avicularia have

a ligulate bar. Generally, the orientation of the avicularia is

random but some colonies tend to have a pair of laterally

directed avicularia, somewhat similar to those of C. ecstasis

(compare Fig. IB with 5B), from which they are distinguished

by their orifice shape and the presence of a ligulate bar. The

peristomes are worn but a few show an adapical pore on their

outer edge. The antapical surface is smooth and thickly calci-

fied, with no sign of cancelli but with radial series of

minute avicularia. These colonies may be assigned to a general

category of “C. angulopora'' but, without examination of type

material of that species, it is not possible to be certain of

their identification, other than that they represent Maple-

stone’s (1909) concept of the species. Three of the colonies

have associated antapical solitary corals, one of which is

figured (Fig. ID).

Specimens from South Australia in the NMV Collection,

identified and labelled as “ Bipora angulopora ” by Maplestone

but never described, belong to two additional species,

C. cognata and C. diffusa (see below).

Conescharellina eburnea (Maplestone, 1909)

Figures 1E-H

Bipora ( Conescharellina ?) eburnea Maplestone, 1909: 270, pi. 72

figs 6a, b.

Conescharellina eburnea. — Livingstone, 1924: 212.

Specimens examined. BMNH 2000.2.23.3, New South Wales (2

colonies, part of material sent by Maplestone to the BMNH, labelled

“cotypes”); NMV F101879, stn SLOPE-2 (3 colonies).

Description. Colonies discoid, wider than high; slightly raised

centrally, marginal zooids prominent. Calcification finely

mamillate. Orifices in quincunx, patent, wide, with a rounded

sinus and distinct condyles. Peristome not raised, with a

rounded adapical pore on its edge. Root pores round, with a

circlet of 3—4 minute avicularia, tending to occur near the

adapical region only. Other avicularia paired, small, rounded,

closely distolateral to the adapical margin of the orifice, orien-

tated distolaterally, with a minute ligula. At the proliferal

margin, these avicularia appear on the antapical surface of

the zooids and closely resemble the paired avicularia of

C. ocellata, C. perculta, and Trochosodon diommatus (see also

Figs 11B, 13C, 21B). There are only a few cancelli on the solid

antapical surface but a regularly distributed series of small,

rounded avicularia.

Colony diameter 4-5 mm, height 1 mm, number of whorls

8, number of zooids per whorl 10.

Remarks. Maplestone’s specimens were from 22 miles east

of Port Jackson, from 146 m; he did not give details of the

number of colonies. There are few records of this species, all

from New South Wales. The distinctive features are the patent

orifices, that possess hardly any peristomial rim, the rounded

sinus, and the paired, antapical peristomial avicularia. The cir-

cular root pores, each surrounded by small avicularia, resemble

those of several other species of Conescharellina (see C. plana

sp. nov., C. ocellata sp. nov.) and of Crucescharellina australis

sp. nov.

Australian Conescharellinidae (Bryozoa)

147

Figure 1. A-D, Conescharellina sp. A-C, NMV F98977, colony; direction of growth arrowed; scale=l mm, B, detail of orifices and avicularia,

scale = 500 pm, C, detail of orifice with adapical pore (arrow), scale=100 pm. D, NMV F98978, colony from antapical surface with commensal

coral, scale = 1 mm. E-H, Conescharellina eburnea (Maplestone, 1909). E, BMNH 2000.2.3.33, adapical view of colony, scale=l mm. F-H,

NMV F98979. F, adapical view of colony showing root pores, scale = 500 pm. G, antapical view of colony with small, central and paired mar-

ginal peristomial avicularia (arrow), scale=500 pm. H, detail of orifices and avicularia, adapical pore arrowed, scale= 100pm.

148

P. E. Bock and P. L. Cook

Conescharellina biarmata (Maplestone, 1909).

Figures 2A, B

Bipora biarmata Maplestone, 1909: 268, pi. 75 figs la, b.

Conescharellina biarmata . — Harmer, 1957: 729.

Specimens examined. BMNH 2000.2.23.2, (4 colonies, part of

material sent by Maplestone to BMNH, labelled “cotypes”); NMV

F98980, no locality (89 colonies, labelled by Maplestone, probably

part of type material); NMV F101880, South Australia, (9 colonies,

from Maplestone, with “S.A.” on box); NMV F101881, stn SLOPE-19

(2 colonies); NMVF101882, stn BSS-170 (1 colony); NMV F101883,

no locality, slide labelled E3195, suspected material from “Endeavour”

(New South Wales) but no other information (2 colonies).

Description. Colonies very small, conical, distinctly higher

than wide. Calcification smooth. Orifices in radial series,

patent, with little raised peristome, elongated oval, with a very

small sinus formed by paired condyles. Frontal septular pores

slit-like. Adapical pore placed just outside the edge of the peri-

stome. Avicularia regularly paired, proximolateral to the orifi-

cial sinus, small, elongated, narrow, triangular, with a ligulate

bar, orientated laterally and slightly antapically. Lunate root

pores adapical, uncommon. Antapical surface with small

avicularia in mature specimens, adapical surface solid with

kenozooidal calcification and scattered avicularia.

Specimens 2.0-2.3 mm high, 1.4-1. 6 mm wide, and com-

prise approximately 10-13 astogenetic generations arranged in

radial rows. The number of zooids in each whorl, 6-8.

Remarks. Livingstone (1924) placed C. biarmata in the

synonymy of C. angulopora. Maplestone’s material differs

from Livingstone’s concept of this species in the consistently

small colonies, and the characters of orifices and avicularian

orientation. Harmer (1957) did not describe C. biarmata but

treated it as a distinct species. Maplestone’s specimens are

numerous and very consistent in characters that are shared by

the specimens from stn SLOPE- 19 and the slide E3195. Other

specimens with elongated, triangular, paired avicularia from

the SLOPE stations differ in several respects and are here

described as Conescharellina ecstasis (see below).

Conescharellina multiarmata (Maplestone, 1909)

Figures 2C-F

Bipora multiarmata Maplestone, 1909: 268, pi. 75 figs 2a, b.

Conescharellina multiarmata . — Livingstone, 1924: 212.

not Trochosodon multiarmatus. — Gordon, 1989: 83, pi. 49 D-F (=

Trochosodon gordoni sp. nov., see below).

Specimens examined. BMNH 2000.2.23.4; (3 colonies, part of

material sent by Maplestone to the BMNH, labelled “cotypes”); NMV

F98981, probably NSW (31 small colonies from two boxes labelled by

Maplestone, probably part of type material); NMV F101884, stn BSS-

169 (1 colony); NMV F101885, stn BSS-170 (1 colony); NMV

F98982, F98983, stn SLOPE-2 (20 colonies, 4 with roots, 2 with corals

on antapical side); NMV F101886, stn SLOPE-7 (1 colony); NMV

F101887 stn SLOPE-39 (2 colonies); NMV F101888, stn SLOPE-40

(47 colonies); NMV F101889. stn SLOPE-48 (1 colony); NMV

F101890, stn GAB-030 (1 colony).

Description. Colony conical, higher than wide. Calcification

smooth to finely mamillate. Orifices in radial series, small,

elongated, with a small sinus flanked by distinct condyles, and

a raised lateral peristome. Adapical pore obviously tubular,

opening on the inside of the peristome edge. Avicularia very

small, rounded, 4 placed laterally and 1 proximolaterally near

the orifice sinus; bar with a minute ligula. Frontal pores minute,

forming a pattern among the orifices. Antapical surface solid

with pores and small avicularia at maturity; marginal peristome

with small paired avicularia. Lunate root pores in radial series

with the zooid orifices, present towards the adapical region of

the colony.

Colonies 1. 9-2.2 mm in height and 1. 8-2.2 mms wide. They

comprise approximately 4-12 or more astogenetic generations,

arranged radially, and include 8-10 zooids per whorl. The

numerous colonies from the SLOPE stations are wider than

those from Maplestone’s collection, inferred to be from New

South Wales. Mature colonies may show a small central area of

cancelli on the antapical surface.

Remarks. C. multiarmata differs from C. biarmata, that also

has very small colonies, in the details of the primary orifice and

character and distribution of the minute avicularia. These alter-

nate with minute frontal pores and have a rounded rostrum; the

bar has a single small ligula. Paired avicularia are visible on

the antapical surface of marginal zooids but are not as promi-

nent as those of C. eburnea. In some zooids, the adapical pore

is prominent and a tubular extension of its calcification can be

seen to descend into the peristome, like that of Trochosodon

asymmetricus (Fig. 2D). The colonies from the SLOPE stations

are generally larger than those from the Maplestone collection.

A colony from stn SLOPE-2 has an incorporated antapical

solitary coral present (Fig. 2F).

Gordon (1989) identified specimens from New Zealand as

Maplestone’s species and assigned them all to Trochosodon.

The New Zealand material appears to be referable to

Trochosodon but certainly not to Conescharellina multiarmata :

it is here renamed Trochosodon gordoni sp. nov.

Conescharellina magniarmata (Maplestone, 1909)

Figures 3A, B

Bipora magniarmata Maplestone, 1909: 269, pi. 75 figs 3a, b.

Conescharellina magniarmata . — Harmer, 1957: 729.

Specimens examined. BMNH 2000.2.23.5 (2 colonies, part of material

sent by Maplestone to the BMNH, labelled “cotypes”); NMV F98984

(12 colonies, labelled in Maplestone’s hand, inferred to be from NSW);

NMV F101891, South Australia (1 colony, Maplestone’s specimens,

no other information); NMV F101892, stn BSS-167 (4 colonies with

roots and antapical corals); NMV F101893, stn BSS-170 (2 colonies

with antapical corals); NMV F101894, stn BSS-171 (1 colony); NMV

F101895, stn GAB-015 (1 colony); NMV F101896, stn GAB-020 (1

colony); NMV F101897, stn GAB-056 (1 colony).

Description. Colony conical, becoming large, broader than pre-

vious species, with adapical extrazooidal, kenozooidal and

avicularian growth forming a prominent “lump” quite early in

astogeny. Orifices in 6-8 radial series, slightly elongate, with

an extended sinus; secondary orifices with laterally raised

peristomes, forming “costules”. Adapical pore outside the

peristome. Avicularia very large, originating beside the orifice,

Australian Conescharellinidae (Bryozoa)

149

Figure 2. A-B, Conescharellina biarmata (Maplestone, 1909). NMV F98980. A, lateral view of colony, growth direction arrowed, scale=l mm.

B, detail of orifices with adapical pore, avicularia and slit-like frontal septular pores, scale= 100 pm. C-D, Conescharellina multiarmata

Maplestone, NMV F98981. C, Lateral view of colony, growth direction arrowed, scale=500 pm, D, detail of orifices from slightly antapical view,

showing tubular adapical pores, avicularia and minute frontal septular pores, scale = 200 pm. E-F, Conescharellina multiarmata (Maplestone,

1909). E, NMV F98983, orifices and avicularia; note pattern of septular pores, scale= 200 pm. F, NMV F98982, Antapical view of colony show-

ing small solitary coral. Note paired, marginal peristomial avicularia, scale= 500 pm.

acute, orientated distolaterally, with a wide, curved, palatal

flange surrounding an opesia, and a bar with at least 3 large

ligulae. Lunate root pores present, each with a pair of small

lateral avicularia. Mature colonies with solid antapical

extrazooidal calcification and avicularia.

Height of colonies 3.0 mm, diameter 3.0 mm, number of

zooid whorls 8-9, number of zooids per whorl 8.

Remarks. Livingstone (1924) regarded C. magniarmata as a

synonym of his “C. angulopord ’ but Harmer (1957) was

doubtful of the identity of the two forms. The avicularia cer-

tainly distinguish C. magniarmata. Few of the colonies

assigned here to C. cognata show any intermediate characters,

although the two forms are obviously very closely related. All

the specimens from eastern Australia have elongated conical

colonies but one from south-western Australia (stn GAB-015)

is very small and domed. However, it has typical “ magniar-

mata ”- type avicularia, with a wide palatal flange and is there-

fore referred to this species.

150

P. E. Bock and P. L. Cook

Fig. 3. A-B, C. magniarmata Maplestone, NMV F98984. A, Lateral view of colony, growth direction arrowed, note adapical secondary calcifica-

tion, scale= 1 mm. B, detail of orifice with adapical pore and avicularium, scale= 100 pm. C-G, Conescharellina cognata sp. nov. C-E. NMV

F98985, holotype. C, lateral view of large colony, growth direction arrowed, scale = 2 mm. D, margin of proliferal region in antapical view, show-

ing zooid orifices with adapical pores and “concealed” frontal shields with marginal septular pores. Chambers of alternating developing inter-

zooidal avicularia arrowed, scale = 500 pm. E, detail of orifices with adapical pores, one with entooecial and ectooecial laminae of developing

ovicell (arrow); large, interzooidal avicularia and lunate root pore with small paired avicularia, scale = 200 pm. F, NMV F98986, antapical view

of small colony, showing central area of cancelli and alternating “concealed” zooids with marginal septular pores, scale = 500 pm. G. NMV

F98985, holotype, detail of antapical surface of large colony, showing alternating series of cancelli, minute avicularia and extensions of septular

pores, scale = 200 pm.

Australian Conescharellinidae (Bryozoa)

151

Conescharellina cognata sp. nov.

Figures 3C-G

Holotype. NMV F98985, Maplestone collection, Kangaroo Island,

South Australia, labelled “ Bipora angulopora” .

Paratypes. NMV F101898, locality as above (23 colonies).

Other specimens. NMV F101899, Maplestone collection, South

Australia (no details) (23 colonies); NMV F101900, stn BSS-55

(1 colony); NMV F101901, stn BSS-65 (1 colony); NMV F101902, stn

BSS-117 (4 colonies with roots); NMV F101903, stn BSS-127

(1 colony); NMV F101904, stn BSS-130 (4 colonies); NMV F101905,

stn BSS-132 (1 colony); NMV F101906, stn BSS-138 (1 colony);

NMV F98986, stn BSS-155 (34 colonies); NMV F101907, stn BSS-

158 (12 colonies); NMV F101908, stn BSS-159 (12 colonies); NMV

F101909, stn BSS-161 (11 colonies); NMV F101910, stn BSS-162

(3 colonies); NMV F101911, stn BSS-170 (2 colonies); NMV

F101912, stn BSS-171 (8 colonies); NMV F101913, stn BSS-176

(6 colonies); NMV F101914, stn BSS-194 (3 colonies); NMV

F101915, stn GAB-019 (1 colony); NMV F101916, stn GAB-020

(5 colonies); NMV F101917, stn GAB-030 (3 colonies); NMV

F101918, stn GAB-045 (2 colonies, one with root); NMV F101919, stn

GAB-049 (1 colony with root and antapical coral); NMV F101920,

stn GAB-067 (1 colony); NMV F101921, stn GAB-098 (2 colonies);

NMV F101922, stn GAB-101 (1 colony).

Etymology, cognatus (L.) - related, referring to the similarities

of the species with some descriptions of “C. angulopora , \

Diagnosis. Conescharellina with large, often flattened

colonies; antapical surface flat or hollow, with large cancelli.

Zooid orifices with a narrow sinus and large condyles.

Avicularia large, lateral, interzooidal, with subtriangular ros-

trum orientated adapically and laterally. Root pores frequent,

lunate.

Description. Colonies very large, usually flattened, occasion-

ally conical, particularly early in astogeny. Calcification

smooth. Zooids in irregular quincuncial series, tending to

appear radial in very large colonies. Primary orifice an elon-

gated oval with a narrow sinus and prominent condyles,

peristome raised laterally and antapically but only a little adapi-

cally; adapical pore just outside, or on edge of the peristome.

Avicularia interzooidal, lateral and adapical, large, rostrum sub-

triangular, directed adapically and laterally (usually in the same

direction); bar with 1-3 ligulae. Lunate root pores frequent

among orifices, apparently replacing avicularia, each with a

pair of small lateral avicularia with a ligulate bar. Antapical sur-

face with large, central cancelli, or hollow, covered by series of

cancelli and minute avicularia. Proliferal region growing edge

with prominent frontal shields visible at all stages of growth.

Colony diameter up to 10 mm, height 3 mm, number of

whorls (radial) 18, number of zooids per whorl, at least 10.

Remarks. C. cognata differs from the accepted character of

Maplestone’s concept of C. angulopora (see above) that it

resembles in several features: its colony shape and the variable

but generally slightly wider primary orifice. There is a variation

in orifice shape between colonies from Bass Strait, where they

are very narrow, and those from Southern Australia. The large

avicularia can be seen at the growing edge to be interzooidal

(Fig. 3D) and are usually consistently orientated in one direc-

tion within a single colony, although they vary within samples.

The pair associated with each root pore is orientated laterally.

Root pores are abundant, apparently replacing avicularia. In the

larger colonies, the hollow antapical surface is completely

covered by series of cancelli, interspersed with minute,

rounded avicularia, totally unlike the solid surface of the

conical colonies of C. angulopora (sensu Maplestone).

C. cognata is common among the samples from GAB and

BSS stations and in the Maplestone collection from South

Australia. Gordon (1985, 1989) described and figured speci-

mens from the Kermadec region and from New Zealand, that he

assigned to C. angulopora. The colonies were not conical but

flat. The antapical surface had no avicularia and a small central

cancellate area. Gordon used an identical description for both

sets of specimens and figured developing ovicells in one of the

colonies from the Kermadec region (1985: fig. 23). Although

closely similar to the colonies from South Australia, these spec-

imens differ in the more elongated shape of the primary orifices

and the characters of the antapical surface.

Conescharellina ecstasis sp. nov.

Figures 4, 5A-D.

Holotype. NMV F98987, stn SLOPE-6.

Paratypes. NMV F101923, stn SLOPE-6 (45 colonies, many with

roots and opercula and mandibles).

Other specimens. NMV F101924, stn SLOPE-2 (7 colonies, 4 with

roots); NMV F98988, stn SLOPE-7 (10 colonies, 2 with roots); NMV

Figure 4. Conescharellina ecstasis sp. nov. NMV 98987, holotype.

Radial series of zooid orifices with adapical pores and large avicular-

ia, scale = 200pm.

152

P. E. Bock and P. L. Cook

Figure 5. A-C, Conescharellina ecstasis sp. nov. A, NMV 98988. Adapical view, scale = 500 pm. B-C, NMV 98987, holotype. B, lateral view of

colony, growth direction arrowed, scale = 1 mm. C, detail of adapical surface showing secondary calcification, avicularia and lunate root pore

with three small avicularia, scale = 200 pm. D-F, Conescharellina diffusa sp. nov. D, NMV F98990. Adapical view of colony showing lunate

root pores and broken ovicell (arrow), scale = 500 pm. E, NMV F98989, holotype. Lateral view of large colony, growth direction arrowed,

scale = 1 mm. F. Conescharellina aff. diffusa sp. nov. NMV P311803, Bairnsdale, Victoria, Miocene. Lateral view of colony, scale = 500 pm.

F101925, stn SLOPE-39 (1 colony with root); NMV F101926, stn

SLOPE-40 (2 colonies); NMV F101927, stn SLOPE-45 (1 colony);

NMV F101928, stn SLOPE-48 (1 colony); NMV F101929, stn

SLOPE-53 (3 colonies).

Etymology, ecstasis (L.) - joy, with reference to the appearance

of the orifice and paired lateral avicularia.

Diagnosis. Conescharellina with large, conical colonies, solid

antapically. Zooid orifices in radial series, primary orifice with

a rounded sinus. Avicularia very large, paired, elongated, lateral

to orifice, orientated laterally and adapically. Root pores lunate.

Description. Colonies large, conical, wider than high, domed

and stellate in early astogeny. Calcification finely mamillate.

Zooids in apparently radial series, peristomes prominent mar-

ginally, particularly in young, stellate colonies. Primary orifice

with a distinct, rounded sinus and paired condyles, adapical

pore on the edge of the peristome. Avicularia paired, lat-

eral, very large, with acutely triangular rostra, nearly always

directed laterally and adapically; bar without a ligula. Adapical

region with large avicularia and lunate root pores, each with a

pair of small lateral avicularia. Antapical region solid, with

radiating series of small avicularia and a few cancelli.

Colony diameter up to 4.7 mm, height 2.3 mm, number of

whorls 6, number of zooids per whorl 8.

Remarks. The colonies of C. ecstasis sp. nov. are recognisable

immediately, even to the naked eye, by the pairs of large avic-

ularia, with mandibles of dark brown cuticle. The orientation of

the rostra varies a little; those of the one specimen from stn

SLOPE-45 being almost horizontal, like the rostra of

C. biarmata. In contrast, one of the two colonies from stn

SLOPE-7 has rostra directed almost adapically. Young colonies

are stellate, with prominent peristomes, especially marginally,

giving the colony a “ Trochosodon-Mke ” appearance (Fig. 5A).

C. ecstasis differs completely from C. biarmata in colony

shape and size, the characters of the primary orifice, and lack

of avicularian ligulae. Except for stn SLOPE-6, only a few

specimens of C. ecstasis were present at each of the eight

SLOPE stations. The two colonies of C. biarmata sensu stricto

from stn SLOPE- 19 were easily distinguished by their much

Australian Conescharellinidae (Bryozoa)

153

smaller dimensions and orientation of avicularia. All records of

C. ecstasis are from deep water. The SLOPE stations range

from south-eastern New South Wales, to eastern Victoria and

Tasmania. The bathymetric range of records is from 400 m to

1096 m.

Conescharellina diffusa sp. nov.

Figures 5D-F, 6A-B

Holotype. NMV F98989, South Australia (no other details), from box

labelled “ Bipora philippinemE^ in Maplestone’s hand.

Paratypes. NMV F101930, South Australia, as above (3 colonies).

Other specimens. NMV F101931, South Australia, Maplestone

Collection (55 colonies); NMV F101932, Kangaroo Island, South

Australia, Maplestone Collection (1 colony); NMV, F101933, probably

NSW, Maplestone Collection (3 colonies); NMV F101934, stn BSS-

065 (8 colonies, 5 with roots); NMV F101935, stn BSS-171 (1 colony);

NMV F101936, stn SLOPE-49, (2 colonies); NMV F101937, stn

GAB-020 (1 colony); NMV F101938, stn GAB-067 (1 colony); NMV

F101939, stn GAB-069 (1 colony); NMV F101940, stn GAB-118

(3 colonies); NMV F101941, stn GAB-129 (2 colonies); NMV

F98990, Dampier DA-2-37-01, North-western Australia (2 colonies,

one with 8 ovicells, both with roots); NMV FI 01942, Dmitri

Mendeleev collection, Tasmania (1 colony).

C. aff. diffusa : Specimens from the Tertiary of Victoria with

very similar but not identical characters: Bairnsdale, NMV

P311803 (Fig. 5F), P311804, plus 20 additional colonies;

Muddy Creek (three colonies).

Etymology, diffusus (L.) - extended, dispersed, with reference

to the wide distribution of this species.

Diagnosis. Conescharellina with large, often elongated, conical

colonies, zooid orifices radial, surrounded by a rim of peris-

tome. Root pores frequent, lunate, without avicularia.

Avicularia in series alternating with orifices, small, rounded to

subtriangular, bar without a ligula; non-palatal area with

spinous processes. Ovicells fragile, with a wide ectooecial rim.

Description. Colony often large, conical, very narrow or

domed, higher than wide. Zooid orifices apparently radial,

alternating with radial series of rounded avicularia.

Calcification smooth. Primary orifice with a fairly deep, round-

ed sinus and paired condyles, surrounded by a peristome rim;

adapical pore outside peristome. Ovicells fragile, with a wide

ectooecial rim and a semitransparent entooecial frontal area.

Avicularia paired, lateral and slightly adapical, or in series

alternating with orifices. Orientation lateral and adapical; ros-

trum rounded to subtriangular, bar without ligula, but with 3 or

more very fine, spinous processes on the non-palatal side.

Antapical surface solid, with a small, central cancellate area.

Lunate pores frequent, occurring in series with the avicularia

but without any accompanying small avicularia.

Recent colonies with up to 28 zooid whorls and more than

14 zooids per whorl, height up to 5.0 mm, diameter 3.5 mm.

Fossil colonies rounded, height 2.5 mm, diameter 3 mm.

Remarks. The Recent colonies are among the largest and most

widely distributed of the Australian species examined. The

avicularia appear to be unique in possessing small calcareous

spine-like structures on the non-palatal side of the bar. Some

Figure 6. Conescharellina diffusa sp. nov. NMV F98989, holotype. A,

detail of zooid orifices with adapical pores, lunate root pores and avic-

ularia with non-palatal spinules (arrow), scale = 200 pm. B, detail of

orifices with adapical pores; orifice at left shows broken laminae of

ectooecium and overlying entooecium of developing ovicell; orifice at

right shows adapical pore surrounded by ectooecial lamina of ovicell

(arrow). Note non-palatal spinules of avicularia, scale = 200 pm.

154

P. E. Bock and P. L. Cook

zooids show evidence of the development of an ectooecial

lamina surrounding the adapical pore; others appear to have

also developed an entooecial layer above the pore (Fig. 6B).

Usually, colonies are distinctly higher than wide but five of

eight from Bass Strait (stn 64) are shorter and more rounded in

outline. All five have long roots, in one case, anchored ter-

minally to a fragment of a “scrupocellariid” bryozoan. The two

colonies from north-western Australia are also conical but

somewhat rounded; their avicularia show non-palatal spinous

processes. One colony has eight fragile and only partially com-

plete ovicells (Fig. 5D), showing that these have a very thinly

calcified entooecium and a wider ectooecium than those of

Trochosodon fecundus sp. nov. and C. stellata sp. nov.

Records of C. diffusa are widely separated. It ranges from

north-west Australia to New South Wales, the west and central

Australian Bight, South Australia, Bass Strait, and Tasmania,

from 15 m. (north-west Australia) to 200 m. (Tasmania).

The fossil colonies are much smaller and domed; their avic-

ularia do not possess any non-palatal projections and it appears

probable that, although closely related, they are not referable to

C. diffusa sensu stricto (Fig. 5F). The specimen of “Bipora

cancellata ” from Baimsdale described by MacGillivray (1895:

89, pi. 12 fig. 1; NMV P22727) appears to be conspecific with

those listed here from Baimsdale and Muddy Creek.

Conescharellina obscura sp. nov.

Figures 7, 8A-C

Bipora philippinensis. — Maplestone, 1910: 6, pi. 1 figs 2, 2a (not

Busk, 1854).

Holotype NMV F98991, stn BSS-155.

Paratype NMV F98992, stn BSS-155.

Other specimens. BMNH as C. philippinensis, 1909.11.12.12 and

13, Green Point, Port Jackson, NSW (Maplestone Collection, possibly

from Whitelegge’s material, 2 colonies with ovicells); BMNH

1899.5.1.1148, Port Jackson (Hincks Collection, 5 colonies); BMNH,

Figure 7. Conescharellina obscura sp. nov. A, NMV F98991, holo-

type. Lateral and adapical view of large colony, scale= 500 pm.

as L. cancellatus, 1934.10.20.88, Port Stephens, NSW (Vine

Collection, 2 colonies); NMV F101943, stn GAB-048; (1 colony with

ovicells); NMV F101944, stn GAB-074; (3 colonies); NMV F101945,

stn GAB-108 (1 colony); NMV F101946, stn GAB-113 (2 colonies);

NMV F101947, stn GAB-118 (2 colonies); NMV F101948, stn GAB-

131 (1 colony); NMV F101949, Dampier DA-2-09-02 (1 colony with

root); NMV FI 01950, Dampier DA-2-73 -01 (2 colonies with roots).

Etymology, obscura (L.) - hidden, referring to the confusion of

records with those of C. cognata, C. stellata and C. philip-

pinensis.

Diagnosis. Conescharellina with flat, often large colonies.

Zooid orifices oval with a short sinus, peristomes not promi-

nent. Avicularia rounded, lateral and antapical, near the orifice,

bar with 1-3 ligulae. Root pores lunate. Ovicells globular, very

fragile, with an extensive area of entooecium frontally.

Description. Colony fairly flat, even lenticular, distinctly wider

than high. Zooid orifices quincuncial, sunken in surrounding

peristome, that is not prominent. Orifice oval, with a short,

Figure 8. Conescharellina obscura sp. nov. A-B, NMV P98992. A, adapical view of small colony, showing lunate root pores, scale = 500 _m. B,

detail of zooid orifices, avicularia, and one lunate root pore, scale = 200 pm. C, BMNH 1899.5.1.1148, Hincks Collection, Port Jackson, New

South Wales. Detail of zooid orifices with adapical pores, avicularia and one lunate root pore. Note the wide orifice sinuses, scale = 200 pm.

Australian Conescharellinidae (Bryozoa)

155

rounded sinus and small condyles. Adapical pore within peris-

tome calcification. Avicularia rounded, lateral and antapical

near the orifice, bar with 1-3 ligulae. Lunate root pores fre-

quent adapically, with 1, occasionally 2, lateral avicularia.

Antapical surface covered with small cancelli.

Colonies up to 14 mm in diameter but usually much

smaller, maximum number of zooid whorls estimated as 15,

and zooids per whorl 20.

Remarks. The close but superficial similarity in colony form

means that records of this species were originally confused

with those of C. cognata, that also has large, flat colonies with

numerous antapical cancelli. The colonies from the GAB sta-

tions are very large, ranging from 9 to 12 mm in diameter and

are hollow antapically; those from stns GAB-074 and

GAB- 118 are more domed and nearly solid antapically. In the

smaller colonies from stn BSS-155, the lunate root pores are

present but they are very rare or absent from the large colonies.

Again, this is in contrast to C. cognata, where they are common

throughout colony growth. Although the large, flat colonies

resemble those described in “C. crassa” by Tenison Woods

(1880), they differ in orifice shape and the types of avic-

ularia and root pores present. Antapical oral avicularia are

also found only in C. pustulosa, from which C. obscura differs

in colony shape and size, orifice shape and form of the

antapical cancelli. Whitelegge (1887) described speci-

mens from Port Jackson as Bipora philippinensis, with a

depressed conical shape and orifices with a wide sinus.

Avicularia with subcircular mandibles occurred in pairs and

sometimes on the antapical side of each orifice. The anta-

pical surface had cancelli and avicularia. Ovicells were present

and were “globose and smooth, with a faint fimbriated stigma

in front.”

Colonies referred to this species were observed alive by

Whitelegge (1887: 347) for three days. He noted pairs of

“tubular filaments” attached to annelid tubes and to fragments

of shell. He thought that these roots originated from avicularia

and did not recognise the function of the lunate root pore that

he also reported. Maplestone (1910: 6, pi. 1, fig. 2) illustrated

ovicells in specimens of Whitelegge’ s material that he also

referred to C. philippinensis but noted that he could see no

frontal stigma. Maplestone ’s specimens from Port Jackson, in

the BMNH collection, are probably part of Whitelegge’ s mater-

ial and are here referred to C. obscura sp. nov., whereas those

in the NMV collection from South Australia, also labelled

Bipora philippinensis, are here assigned to another new

species, C. diffusa. The two slides from Green Point

(1909.11.12.12 and 13) each contain a single, fairly flat colony,

less than 2.5 mm in diameter, with most of the opercula and

mandibles intact. They each include two fragile ovicells and up

to eight partially developed ovicells. These are globular, like

those figured by Maplestone (1910). They appear to have an

extensive area of frontal entooecial calcification and a series of

minute pores close to the ovicell base, that may mark the limit

of the ectooecium. There is no sign of any striations or a

“stigma”. The other, smaller specimens from the BMNH col-

lections are obviously conspecific but have no ovicells. They

too, have most of the opercula and mandibles present; only one

preparation shows the primary orifice clearly. One of the five

specimens from Port Jackson (Fig. 8C, Hincks collection,

BMNH 1899.5.1.1148) is similar in characters to specimens

from Bass Strait (Fig. 8B, stn BSS-155) except for the greater

width of the orifice sinus. C. obscura occurs from north-west

Australia and across the Great Australian Bight to New South

Wales, from a depth range of 12 to 125 metres.

Conescharellina stellata sp. nov.

Figures 9A-I

Holotype. NMV F98993, stn GAB-019.

Paratypes. NMV F98994, stn GAB-019 (8 colonies).

Other specimens. NMV F98995, stn GAB-128 (1 colony).

Etymology, stellata (L.) - starry, referring to the appearance of

the colonies from the adapical side.

Diagnosis. Conescharellina with small, domed colonies.

Orifices with rounded sinus and distinct condyles, surrounded

by a raised peristome laterally and sometimes antapically.

Avicularia lateral, rounded; bar without a ligula, non-palatal

area sometimes filled by a lamina. Ovicells fragile, with a

depressed, marginally striated entooecium.

Description. Colony small, domed, wider than high; zooid

orifices quincuncial. Calcification smooth and slightly tuber-

culate, adapical region sometimes with small, secondarily

thickened mamillae. Peristomes raised laterally, forming a

prominent, stellate pattern, especially at the colony margin;

sometimes extended adapically and very prominent. Primary

orifices rounded with a fairly wide sinus and small to distinct

condyles. Adapical pore large, on the edge or outer face of the

peristome, surrounded by a rim of calcification, sometimes

slightly asymmetrically placed. Avicularia rounded, lateral and

paired, widely separated from the peristomes; bar without a

ligula, non-palatal side sometimes with a thin lamina, occa-

sionally pierced by a pore. Lunate root pores tending to occur

adapically, each with a pair of closely apposed, rounded lateral

avicularia. Ovicells present on subperipheral zooids, very

fragile, with a raised, smooth, transparent ectooecium and a

depressed entooecium, striated marginally, forming pores at

the ectooecial junction. Antapical surface with a small central

cancellate area.

Colony diameter 1-1.5 mm, height 1 mm, number of alter-

nating whorls 6, number of zooids per whorl 6-8.

Remarks. The colonies have a very regular, stellate appearance

from the adapical side. The peristomes are usually well devel-

oped laterally but, in one colony (from stn GAB -128), they are

also extensive antapically, forming a funnel. The ovicells are

extremely fragile and were detached soon after initial scanning

electron microscopy. The lower face of the ectooecium shows

that it was almost certainly covered by cuticle and apposed but

not attached to the surface of the zooid adapically (Figs 9H, I).

Although closely similar to the smaller colonies of C. obscura

in several features, C. stellata differs in the form and distribu-

tion of the avicularia, that do not include a solitary one on the

antapical side of the peristome. The avicularia also differ in the

lack of ligulae and the presence of a lamina filling the non-

156

P. E. Bock and P. L. Cook

Figure 9. Conescharellina stellata sp. nov. A-C NMV F98993, holotype. A, adapical view of colony showing prominent bilabiate marginal peri-

stomes and lunate root pores, scale = 500 pm. B, detail of marginal zooid orifice with adapical pore. Avicularium with porous non-palatal lami-

na (arrow), compare Figure 9E, scale = 200 pm. C, lateral view of colony showing lunate root pore, avicularia, scale = 200 pm. D-E, NMV

F98995. D, adapical view of colony with prominent, rounded, funnel-shaped marginal peristomes, scale = 500 pm. E, marginal view showing

extended peristomes, orifices and avicularia with non-palatal laminae, one with pore (arrow), compare Figure 9B, scale = 200 pm. F-I, NMV

P98994. F, Colony with cuticle in situ and two ovicells near margin, scale = 100 pm, G, detail of ovicells, scale = 100pm. H, I, frontal and basal

surfaces of detached ovicell. H, showing ridges in margin of entooecium, forming pores at junction with ectooecium, with basal smooth ectooe-

cium. I, with fracture at original margin of adapical pore, scales = 50 pm.

Australian Conescharellinidae (Bryozoa)

157

palatal area. This is not always developed in the specimens

from stn GAB -01 9 but is constantly present in the distinctly

larger, more prominent avicularia of the colony from stn GAB-

128. The lamina may be pierced by a pore in both populations.

C. stellata has been found from the western end of the Great

Australian Bight at a depth of 59 m and from the central region

at a depth of 304 m. Although these two populations show

differences in detail, the number of colonies does not allow any

estimate of its significance.

Conescharellina plana sp. nov.

Figures 10A-D

Holotype. NMV F98996, stn SLOPE-2.

Paratypes. NMV F98997, stn SLOPE-2 (26 colonies).

Other specimens. NMV F101951, stn BSS-167 (4 colonies, 1

with root and ovicells); NMV F101952, stn BSS-169; 3 colonies,

1 with root); NMV F101953, stn SLOPE-6 (5 colonies); NMV

F101954, stn SLOPE-7 (1 colony); NMV F101955, stn SLOPE-40 (57

colonies, 2 with roots); NMV F101956, stn SLOPE-56 (13 colonies);

NMV F101957, stn GAB-020 (6 colonies); NMV F101958, stn GAB-

030 (2 colonies, 1 with root); NMV F101959, stn GAB-044 (1 colony

with root); NMV F101960, stn GAB-049 (1 colony).

Etymology, planus (L.) - smooth, with reference to the lack of

raised peristomes above the colony surface.

Diagnosis. Conescharellina with large, slightly flattened

colonies, solid antapically. Zooid orifices in radial rows, deeply

sunken within a circular peristome, that is not raised above the

colony surface. Avicularia paired, small, with a minute ligula.

Root pores numerous, circular, surrounded by avicularia.

Ovicells fragile with a fairly wide ectooecial rim.

Figure 10. Conescharellina plana sp. nov. A-C, NMV F98996, holotype. A, adapical view of colony, scale = 1 mm. B, detail of root pore with

surrounding avicularia, scale = 200 pm. C, detail of colony margin showing orifices, peristomes, and avicularia, scale = 200 pm. D, NMV F98997,

paratype, antapical surface of colony, scale = 500 pm.

158

P. E. Bock and P. L. Cook

Description. Colonies large, slightly flattened, wider than high.

Zooid orifices in marked, apparently radial series, peristomes

tubular, deep but not prominent at the surface. Calcification

smooth to finely tuberculate. Primary orifice with a small,

rounded sinus, deeply hidden at the base of the circular peris-

tome, adapical pore just outside the edge of the peristome.

Ovicells fragile, with a fairly wide ectooecial rim and a semi-

transparent entooecial frontal area. Avicularia paired, close to

the edge of the peristome, adapical and antapical, very small,

rounded, with a minute ligula. Root pores numerous, large, cir-

cular, surrounded by a circlet of 3-4 small avicularia. Antapical

surface solid and flat, with small, scattered avicularia.

Colony diameter 4.5 mm, height 2 mm. Number of whorls

5, number of zooids per whorl 8.

Remarks. The large, circular root pores of C. plana are similar

in appearance to those of Conescharellina ebumea, C. perculta ,

C. humerus and Crucescharellina australis that are also sur-

rounded by a circlet of small, rounded avicularia. The peris-

tomes of C. plana are unusual in being elongated but not promi-

nent and the colony surface is smooth. Two colonies exhibit a

single, marginal zooid each, with a prominent peristomial avic-

ularium (Fig. IOC). Only one of the two ovicells present in the

colony from stn BSS-167 is complete; a deeply pigmented

embryo is visible through the thin frontal calcification.

Colonies of C. plana are widely distributed off the southern and

eastern coasts of Australia occurring from the western

Australian Bight to the eastern border of Victoria, through Bass

Strait, from depths ranging from 80 to 1096 m.

Conescharellina perculta sp. nov.

Figures 11A-D

Holotype. NMV F98998, slide labelled E3195 (Locality unknown,

probably off New South Wales).

Paratypes. NMV F98999, locality as above.

Etymology, percultus (L.) - highly adorned, with reference

to the patterning of the numerous avicularia and colony

calcification.

Diagnosis and description. Colonies small, discoid, distinctly

wider than high, with a mamillate centre and prominent mar-

ginal peristomes, calcification delicate and finely tuberculate.

Orifices quincuncial, becoming radial. Primary orifice with a

rounded sinus. Peristomes elongated and tubular, raised anta-

pically and prolonged into a spout, prominent at the colony

margin. Adapical pore present on outer face of the peristomes.

Root pores circular, surrounded by up to 5 small avicularia. All

avicularia small, rostrum rounded, bar with a minute ligula.

Each orifice with 1 adapical, 1 lateral and 1 antapical pair of

avicularia. Further pairs of lateral and antapically placed

avicularia, that are visible from the antapical surface, are

accompanied by pairs of pores.

Largest colony about 2.3 mm wide and 0.5 mm high, with 6

astogenetic generations and probably up to 11-12 zooids per

whorl at margin.

Remarks. The locality from that the three small colonies of

were collected also provided two well preserved colonies of

C. biarmata and therefore is inferred to have been collected

from New South Wales. C. perculta is distinguished by its

delicate, semitransparent, finely tuberculate calcification, with

numerous avicularia surrounding the spout- shaped peristomial

orifices. As in C. ebumea, C. ocellata and T. diommatus, the

marginal peristomes can be recognised from the antapical

surface by the pattern or outline of the associated paired

avicularia. In many other respects, such as the depth of the

peristome, the distribution of circum-oral avicularia, and

type of root pore, C. perculta greatly resembles C. plana, from

which it differs principally in colony size, the patterning and

shape of the orifices, and nature of the antapical surface,

including the peristomes. The circular root pores, with

their surrounding avicularia, resemble those of C. ebumea,

C. plana and C. humerus, as well as those of Crucescharellina

australis.

Conescharellina pustulosa sp. nov.

Figures 12A-D

Holotype. NMV F99000, stn SLOPE-2.

Paratypes. NMV F99001, stn SLOPE-2 (9 colonies).

Other specimens. NMV F99002, unlabelled Maplestone specimens,

probably from NSW (5 colonies); NMV F101961, stn BSS-158 (1

colony); NMV F101962, stn BSS-169 (1 colony); NMV F101963, stn

SLOPE-40 (14 colonies); NMV F101964, stn SLOPE-45 (1 colony);

NMV F101965, stn GAB-049 (1 colony).

Etymology, pustula (L) - a bubble or blister, with reference to

the calcification of the zooid surfaces.

Diagnosis. Conescharellina with small, domed colonies.

Zooids orifices with a subtriangular sinus; surrounded by

raised, pustular secondary calcification, that also covers the

antapical surface. Avicularia antapical, peristomial, raised, bar

without a ligula.

Description. Colonies small, domed. Surface irregular, formed

by raised, pustular secondary calcification. Zooids in quin-

cuncial series, peristomes raised laterally, prominent only at

colony margins. Primary orifice with a wide subtriangular sinus

and minute condyles. Adapical pore outside the peristome rim.

One small peristomial avicularium lateral and antapical, nearly

vertical to the rim of the peristome, rostrum rounded, bar with-

out a ligula; other occasional avicularia scattered among the

zooids. Adapical region with a few, lunate root pores; antapical

region with more pustular secondary calcification, cancelli and

avicularia.

Colony diameter 2.2 mm, height 1.2 mm, number of whorls

6, number of zooids per whorl 8.

Remarks. The pustular calcification occurs among the zooid

orifices and is a prominent feature of the antapical surface of

the small colonies. The only other species in the samples

examined that possesses an antapical peristomial avicularium is

C. obscura, but this is not placed on the edge of the peristome

as in C. pustulosa. C. pustulosa bears a close but superficial

similarity to C. papulifera Harmer (1957: 734, pi. 47 figs 7-9,

text-fig. 70C). Harmer did not describe or figure the primary

orifice, that he was not certain was visible. He mentioned

Australian Conescharellinidae (Bryozoa)

159

Figure 11. Conescharellina perculta sp. nov. A, NMV F98998, holotype. A, adapical view of colony, scale = 200 (am. B, NMV P98999, antapi-

cal view of marginal peristomes, scale = 200 pm. C-D, NMV F98998, holotype. C, detail of orifices, peristomes, and avicularia, scale = 200 pm.

D, detail of root pore, scale = 200 pm.

paired avicularia and radially costulate zooid orifices but did

not figure them. Specimens of C. papulifera (BMNH,

1964.3.2.3, paratype?, Java Sea and 1964.3.2.2, Siboga stn 77,

Borneo Bank, 59 m) have been examined. They are minute;

their dimensions being less than half of those of C. pustulosa at

the same astogenetic stage. The peristomes are raised and

tubular, arranged in radial series; the primary orifices are not

visible. Minute avicularia alternate with the zooid orifices and

none are antapical and peristomial.

C. pustulosa is distributed from the coasts of New South

Wales to Bass Strait and the Great Australian Bight, from a

wide depth range of 36 to 800 m. There is, however, little

variation in the characters of the colonies from the different

populations.

Conescharellina ocellata sp. nov.

Figures 13A-D

Holotype. NMV P311805; Miocene, Balcombe Bay, Victoria (see

appendix).

Paratype. NMV P311806; Balcombe Bay, Victoria.

Other specimens. Victoria, Balcombe Bay (55 colonies); Batesford

Quarry (see appendix) (45 colonies).

Etymology, ocellata (L.) - having little eyes, with reference to

the appearance of the antapical peristomial avicularia.

Diagnosis. Conescharellina with minute, slightly domed

colonies. Zooid orifices radial. Avicularia small, paired, lateral,

with a minute ligula. Paired avicularia visible on the antapical

side of the marginal peristomes.

160

P. E. Bock and P. L. Cook

Figure 12. Conescharellina pustulosa sp. nov. A-B, NMV F99000, holotype. A, adapical view of colony, scale = 500 pm. B, detail of two mar-

ginal orifices, showing thick secondary calcification and peristomial avicularia, scale = 200 pm. C, NMV F99001, paratype, group of orifices,

adapical pores with developing ectooecial lamina arrowed, scale = 200 pm. D, NMV F99002, colony lateral view, growth direction (antapical)

arrowed, scale = 500 pm.

Description. Colonies very small, domed, orifices appearing to

be in radial series. Primary orifice with a short, narrow sinus

and distinct paired condyles, with a laterally and antapically

raised peristome. Adapical pore on the edge of the peristome.

Avicularia small, rounded, paired and antapical, lateral to the

sinus, bar with a minute ligula, orientated laterally and ada-

pically. Adapical region with a few rounded pores, inferred to

be root pores, with 2-3 small adjacent avicularia. Antapical

region with a few avicularia and central cancelli; and with

prominent, paired avicularia present on the antapical side of the

peristome of marginal zooids.

Diameter of colony 2.4 mm, height 1.6 mm, up to 6 radial

whorls, 5-7 zooids per whorl.

Remarks. C. ocellata is easily recognisable in unworn colonies

by the paired antapical avicularia on the marginal peristomes.

C. ocellata resembles C. eburnea and Trochosodon diammotos

in this character but differs in all its dimensions and in the

shape of the primary orifice. It differs from C. macgillivrayi in

its domed colonies, radially arranged orifices, shape of the

primary orifice, and rarity of antapical cancelli. It differes from

C. humerus in the position and shape of the primary orifice and

position of the lateral avicularia, especially in antapical view.

Conescharellina macgillivrayi sp. nov.

Figs 13E-F, 14A

Bipora philippinensis. — MacGillivray, 1895: 89, pi. 12 fig. 2.

Holotype. NMV P311810, Miocene, Balcombe Bay, Victoria.

Paratypes. NMV P3 11811, Balcombe Bay.

Other specimens, Miocene, Balcombe Bay (approximately 209

colonies); Miocene, Bairnsdale (12 colonies); Miocene, Batesford

Quarry (22 colonies); Miocene, Heywood Bore (approximately 63

worn colonies); Miocene, Muddy Creek (13 colonies), Miocene,

Australian Conescharellinidae (Bryozoa)

161

Figure 13. A-D, Conescharellina ocellata sp. nov. A-B, NMV P311805. A, adapical view of colony, scale = 500 pm. B, detail of orifices, avic-

ularia and septular pores, scale = 200 pm. C-D, NMV P311806. C, antapical view, scale = 500 pm. D, marginal view showing peristomes and

paired avicularia (arrow), scale = 200 pm. E-F, Conescharellina macgillivrayi sp. nov. NMV P311810. E, adapical view of colony, scale = 500

pm. F, detail of orifices with adapical pores, peristomes and avicularia, scale = 200 pm.

Puebla Clay, Torquay (16 colonies); Miocene, Mount Schanck, South

Australia (approximately 100 colonies).

Etymology. Named for P.H. MacGillivray.

Diagnosis. Conescharellina with fairly flat, small colonies.

Zooid orifices arranged quincuncially, marginal peristomes

prominent. Avicularia rare. Root pores circular. Antapical

surface with cancelli.

Description. Colonies fairly flat, distinctly wider than high,

peristomes prominent marginally. Calcification finely tuber-

culate. Zooid orifices obviously quincuncial, sinus short and

rounded, with paired condyles. Peristomes raised laterally and

antapically; adapical pore outside the peristome. Root pores

round, small, adapical. Avicularia rare, lateral or antapical,

rounded, bar with a minute ligula. Antapical surface with large

cancelli and minute avicularia.

Colony diameter 1.9 mm, height 1.25 mm, number of

whorls 6, zooids per whorl 8.

Remarks. Colonies of C. macgillivrayi are the most numerous

of the fossils found in the Victorian and South Australian

Tertiary samples, although many colonies are worn and their

identity has had to be inferred from their proportions and

orifice pattern. Comparison of the colonies with MacGillivray’s

(1895) specimen of “ Bipora philippinensis ” (NMV P 27728)

indicates that they are conspecific and it is also possible that

this species is the “ Lunulites cancellatus ” of Waters (1882b:

512, pi. 22 figs 10, 11) from Baimsdale, although this was fig-

ured with more numerous avicularia. None of his specimens

has been examined.

162

P. E. Bock and P. L. Cook

Figure 14. Conescharellina macgillivrayi sp. nov. and C. humerus sp. nov. A, C. macgillivrayi NMV P3 11811, antapical view of colony, showing

cancelli, scale = 500 _m. B-F. C. humerus, B, NMV P311814, adapical view of colony, showing root pore, scale = 500 pm. C-D. NMV P311812.

C, Adapical view of colony, scale = 500 mm. D, detail of orifices and avicularia, scale = 200 pm. E, NMV P3 1 1 8 14, detail of root pore (see Figure

14B), scale = 100 pm. F, NMV P3 11813, detail of marginal zooids, showing profiles of peristomes and avicularia, scale = 100 pm.

Conescharellina humerus sp. nov.

Figures 14B-F

Holotype. NMV P311812, Miocene, Balcombe Bay, Victoria .

Paratypes. NMV P3 11813, P3 11814, Miocene, Balcombe Bay.

Other specimens. Miocene, Balcombe Bay (43 colonies); Miocene,

Batesford Quarry (170 colonies); Miocene, Muddy Creek (7 colonies);

Miocene, Paraatte Bore (8 colonies); Miocene, Puebla Clay, Torquay

(16 colonies); Miocene, Mount Schanck, South Australia (approxi-

mately 125 colonies).

Etymology, humerus (L.) - a shoulder, with reference to the

outline of the lateral avicularia and peristome from antapical

view.

Diagnosis. Conescharellina with slightly domed colonies.

Zooid orifices radial. Avicularia small, lateral, forming a

“shoulder” visible on marginal peristomes. Round root pore

near the centre adapically, surrounded by small avicularia.

Description. Colonies small, slightly domed, distinctly wider

than high. Orifices radially arranged towards the margin of the

colony. Primary orifices with a distinct, deep, rounded sinus

and paired condyles, peristome raised laterally, adapical pore

outside peristome. Avicularia small, paired, rounded, lateral

and antapical, directed inwardly, bar with ligula, subrostral

chamber prominent, visible as a lateral “shoulder” in marginal

zooids. A fairly large, rounded root pore near the centre of the

adapical region, surrounded by a circlet of six avicularia.

Antapical surface cancellate centrally, otherwise smooth, with

small avicularia.

Colony diameter 3.3 mm, height 1.5 mm, number of whorls

4, number of zooids per whorl 7.

Australian Conescharellinidae (Bryozoa)

163

Remarks. The colonies of C. humerus are widely distributed in

the Victorian Tertiary but are not as numerous as those of

C. macgillivrayi. C. humerus is immediately recognisable by

the profile of the marginal peristomes formed by the prominent

lateral avicularian rostra. The rounded root pore with circlet

of avicularia is reminiscent of those found in C. eburnea,

C. plana, C. perculta and in Crucescharellina australis.

Bipora Whitelegge, 1887

Bipora Whitelegge, 1887: 340 (part). — Levinsen, 1909: 312. —

Harmer, 1957: 754.

Type species. Flabellipora [sic] flabellaris Levinsen, 1909

(subsequent designation by Levinsen, 1909).

Description. Colony fan-shaped, laterally flattened, zooids

arranged in 2 apposing, frontally budded expanses, separated

by a series of cancelli, visible antapically. Orifices sinuate, with

paired condyles, surrounded by a peristome that is not promi-

nent. Avicularia small and rounded, with a bar but no ligula.

Root pores lunate, paired, adapical. Ovicells not known, but

adapical pore present.

Remarks. Whitelegge (1887) described seven species that he

assigned to Bipora but did not indicate a type species. He

assigned specimens from Port Jackson to “Bipora (?) elegans ”

of d’Orbigny (1852) somewhat doubtfully, remarking “if this

species proves to be different (as I think it will) from the fossil

form described by d’Orbigny as Flabellopora elegans, it can

remain as B. elegans Waters”. D’Orbigny’s species was not a

fossil: Whitelegge’s reference was to a remark by Waters

(1887a: 71) who mentioned receiving a specimen of

“ Flabellopora elegans ” from New South Wales that grew in an

“irregular subcrescentic form with two layers of zooecia

separated by a cellular structure formed of avicularian cells”.

This specimen was apparently from Brazier, as Waters (1887:

200) listed specimens from Port Stephens (from approximately

13-15 m depth), collected by him, some of which had “between

the layers a cancellous structure”. Waters’ figures (pi. 5 figs

13-17) leave no doubt that they represent “Bipora flabellaris”,

even though Waters (1889) remarked that Whitelegge had

“favoured me with further specimens of Flabellopora elegans,

d’Orb., and I feel no doubt as to the correctness of my identifi-

cation”. However, Waters later (1905, 1921) amended this view

and stated that he had adopted Levinsen’ s name. Levinsen

(1909) had somewhat informally and irregularly designated

Flabellipora [sic] elegans Waters (1887) not d’Orbigny (1852),

that he then renamed Flabellipora flabellaris, as the type

species of Bipora. Harmer (1957: 755) remarked that “ Bipora

is a genus of uncertain validity” but that B. flabellaris was the

only species mentioned by Whitelegge (1887: 346), as Bipora

(?) elegans, that would be available as type species, as all the

other species had subsequently been referred either to

Conescharellina or Flabellopora. Presumably, the type speci-

mens of B. flabellaris are among those figured by Waters

(1887). Harmer (1957: 755) incorrectly listed the registration

numbers of some specimens in the collections of the Natural

History Museum. The numbers should read “99.5.1.1147” indi-

cating Hincks’ material and “97.5.1.807” indicating

Bracebridge Wilson material. Harmer concluded that there

seemed to be “sufficient reason for regarding Bipora, with this

genotype” (i.e. B. flabellaris ) “as a distinct genus of

Conescharellinidae”. Lu (1991) described three species of

Zeuglopora from the South China Sea as Bipora.

Maplestone (1904: 209) listed specimens of “ Bipora ele-

gans ” among his own collection of fossils from Jimmy’s Point,

Victoria. No specimens of Maplestone ’s material are extant and

it cannot be established whether or not this is the only fossil

record of Bipora.

Bipora flabellaris Levinsen, 1909

Figures 15A-E

Bipora (?) elegans . — Whitelegge, 1887 (not d’Orbigny, 1852).

Flabellopora elegans. — Waters, 1887: 200.

Flabellipora flabellaris Levinsen, 1909. — Livingstone, 1924: 211.

Specimens examined. NMV F99003, stn GAB -020 (2 colonies); NMV

F101966, stn GAB-030 (2 colonies); NMV F99004, stn GAB-116 (1

colony); NMV F101967, stn GAB-118 (1 colony).

Description. Colony fan-shaped, composed of 2 apposed

zooidal faces, separated by an intervening cancellated and avic-

ularian layer. Adapical region often extrazooidally thickened,

with rhizoids arising from small lunate pores. Zooid primary

orifice with a subtriangular sinus and paired condyles. Adapical

pore present outside peristome. Peristome raised laterally.

Avicularia paired, lateral and antapical, rostrum rounded,

directed adapically, bar without ligula.

Colonies up to 10 mm wide, 8 mm deep.

Remarks. Many of the specimens examined here are worn.

Only one, from stn GAB -020, has three long roots (width 0.25

mm, length 2.0 mm), that arise from the adapical region of a

large colony from 155 m depth.

B. flabellaris is obviously very closely related to species

of Conescharellina. The early growth stages are hardly

distinguishable, except for the slight flattening of the colony.

Later stages, however, emphasise the cancellated region, that

curves round the antapical edge and protrudes beyond the

orifices of the zooidal series of each face, producing the

typical fan-shaped colony. All the specimens examined here

appear to belong to one species but it is possible that other

forms of Bipora may eventually be found from the Australian

region.

Trochosodon Canu and Bassler, 1927

Trochosodon Canu and Bassler, 1927: 11. — Canu and Bassler, 1929:

493.— Harmer, 1957: 744.

Type species. Trochosodon linearis Canu and Bassler, 1927

(original designation).

Description. Colonies forming a low cone, orifices both radial-

ly and quincuncially arranged, antapical marginal series of

zooids tubular, projecting, often prominent; frequently without

avicularia. Adapical pores and kenozooids present, among large

rounded root pores; lunate pores also reported to be present.

Avicularia and ovicells present. Antapical cancelli usually rare

or absent.

164

P. E. Bock and P. L. Cook

Figure 15. Bipora flabellaris (Levinsen, 1909). A-D, NMV F99003. A, young colony, direction of growth arrowed, scale =100 (am. B, antapical

view of colony, showing narrow kenozooidal region (arrowed), scale = 1 mm. C, detail of orifices with adapical pores, and avicularia, scale =

200 pm. D, adapical region, showing lunate root pores, scale = 200 pm. E, NMV F99004, large worn colony at late growth stage, showing

flabelliform shape and everted antapical region, direction of growth arrowed, scale = 2 mm.

Remarks. Canu and Bassler (1927, 1929) considered that

Trochosodon was characterised by the absence of avicularia.

The type species, T. linearis (Canu and Bassler, 1927: 11, 42,

pi. 1 fig. 12; 1929: 493, pi. 1 figs 11-13), was from Sibuko Bay,

Borneo ( Albatross stn 5586), from a depth of 247 fathoms (625

m). The unique figured colony was 2.5 mm in diameter, with

approximately 6-8 zooids per whorl and a strong tendency for

the orifices to be arranged radially. The marginal peristomes

were prominent and the antapical surface was convex, with

little structure except some scattered pores, that may have been

minute avicularia. No avicularia appear to have occurred on the

adapical surface near the peristomes. Canu and Bassler (1929:

494, pi. 70 figs 7-10) also described T. quincuncialis from the

same station. It was distinguished by its quincuncially arranged

orifices but pi. 70, fig. 10 clearly shows some series to be

radially arranged; it is probable that the two species are syn-

onymous. Canu and Bassler (1927, 1929) gave no details of the

primary orifices except that they were sinuate. Their figures

were all retouched but pi. 70 fig. 12 perhaps shows a few ada-

pical pores. Harmer (1957: 744) noted difficulties in defining

Trochosodon , remarking “it is not easy to establish a clear dis-

tinction between this genus and Ga^scda ^^” , maintaining

that all the abyssal species, including T. linearis that he

assigned to the genus, possessed avicularia. However, he could

not have examined the unique type specimen of T. linearis

sensu stricto and it seems possible that his Siboga specimens

belonged to a distinct species (see below). Generally, the dis-

tinguishing features of Trochosodon include prominent, tubular

marginal peristomes and virtual or complete absence of anta-

pical cancelli. Gordon (1989) introduced several abyssal

Australian Conescharellinidae (Bryozoa)

165

species from the New Zealand region but the only authentic

past records from Australia appear to be those of T. ampulla

(Maplestone), described below and three hitherto unnamed

species from Cape York, Queensland, figured by Cook and

Lagaaij (1976) and Cook (1981), from Challenger stn 185,

from 279 m, a locality that was not mentioned by Busk (1884).

These colonies are here referred to T. aster sp. nov.,

T. anomalus sp. nov. and T. praecox sp. nov., bringing the total

of species described from Australia to seven. Ovicells were

described by Harmer (1957) in T. optatus (see below). These

appear to be asymmetrical and have ridged frontals, resembling

those of specimens of T. fecundus sp. nov. from north-western

Australia, that are, however, symmetrically developed.

Similarly ridged ovicells were reported in C. striata by Silen

(1947) but these were also asymmetrically developed, like

those of C. catella, as described by Harmer (1957), and almost

certainly of T. asymmetricus sp. nov. A suite of independent

character states distinguishing “ Trochosodon ” from

“ Conescharellina ” is thus far from complete or consistent.

However, the wide diversity of species assigned to

Conescharellina itself, suggests that this genus will certainly

require eventual revision, including a definition of its type

species and a review of all other taxa referred to it (see Silen,

1947: 34). Until this is accomplished, it is probably wisest to

maintain Trochosodon for a group of species that are fairly

consistent and differ slightly from most other forms assigned

to Conescharellina. Australian species are introduced here

from New South Wales, Victoria, north-west Australia

and Queensland. Although they exhibit a mosaic of character-

istics some of which can be regarded as “typical” of

Conescharellina, they are considered here to be distinct enough

to be assigned provisionally to a generic group and referred to

Trochosodon. Harmer (1957) considered this to be mainly an

abyssal genus but material included here also derives from

shallow depth.

Trochosodon ampulla (Maplestone, 1909)

Figures 16A-C

Bipora ampulla Maplestone, 1909: 269, pi. 76 figs 4a, b, 5a, b.

Conescharellina ampulla. — Livingstone, 1924: 212.

Trochosodon ampulla. — Canu and Bassler, 1929: 493. — Harmer,

1957: 744.

Specimens examined. BMNH 2000.2.23.1 (part of material sent by

Maplestone to the Natural History Museum, 1 colony). NMV F99005,

F99006, labelled by Maplestone, almost certainly part of the

type material from NSW; and NMV F101968, same collection

(6 additional colonies).

Description. Colony forming a very low dome, distinctly

wider than high. Orifices quincuncially arranged, rapidly

obscured by extrazooidal and kenozooidal calcification.

Prominent, flask shaped, marginal zooids with elongated,

tubular peristomes. Primary orifice slightly elongated, with

a small pointed sinus. Adapical pore present outside peristome.

A few scattered pores (root pores?) present adapically.

Avicularia small, often paired, placed laterally and adapically

beside each peristome, rostrum almost semicircular, bar with a

minute ligula. Antapical surface with a small central region of

cancelli.

Colonies up to 4.7 mm in diameter and 1.6 mm in height;

with approximately 8 whorls, each with 10 zooids.

Remarks. The specimens examined are somewhat worn and

show few primary orifices, deeply hidden by the elongated

peristome. The peristomes of the marginal zooids are tubular

and prominent, the calcification is thickened and there are only

one or two apparent adapical pores. Antapical cancelli are

usually confined to a small, central area, although a much

larger area was figured by Maplestone (1909). The species is

distinguished by large size and stellate colony form with very

prominent tubular marginal peristomes.

Figure 16. Trochosodon ampulla (Maplestone, 1909). A, NMV F99005, adapical view of colony, scale = 1 mm. B-C, NMV F99006. B, lateral

view of larger colony, scale = 1 mm. C, detail of orifice and avicularia, scale =100 pm.

166

P. E. Bock and P. L. Cook

Specimens additional to the types suites of T. ampulla

have not been reported since its first description. This, taken

together with Maplestone’s labelling of the NMV specimens

and the occurrence in this collection of other, apparently unique

records from New South Wales, of Zeuglopora lanceolata etc.,

strongly suggests that these specimens are part of the original

type suite.

Trochosodon fecundus sp. nov.

Figures 17A-F

Holotype. NMV F99007, Dampier Archipelago, stn DA-2-75-02.

Paratypes. NMV F99008, F99009, Dampier Archipelago, stn

DA-2-75-02.

Etymology, fecundus (L.) - fertile, prolific, with reference to

the numerous ovicells present in the specimens.

Diagnosis. Trochosodon with peristomes raised laterally and

arranged quincuncially. Zooid orifices concealed, with a very

wide, shallow sinus. Avicularia rounded. Ovicells prominent,

symmetrical, with a thin marginal ectooecium. Root pores

lunate.

Description. Colony forming a low cone, wider than high, with

prominent peristomes, particularly at the margin. Calcification

smooth to finely mamillate. Orifices in irregular, quincuncial

series, oval, with a pair of minute condyles that delineate a

broad, very shallow sinus. Peristomes raised laterally and

antapically, forming a partial, shallow tube. One avicularium

near and lateral to each orifice, rostrum semicircular, often

orientated adapically, with a bar but no ligula. Adapical pore

symmetrically placed. Ovicells fragile, symmetrical, promi-

nent, with an ectooecial layer visible marginally, that extends

laterally to form paired leaflike lobes above the orifice and the

lateral part of the peristomes. Entooecium flat and smooth

frontally, with raised marginal striations forming a series of

pores where it meets the edge of the ectooecium. Small lunate

root pores present. Antapical surface with large cancelli.

Colonies with maximum diameter 2.25 mm and height 0.75

mm, number of whorls 6-7, number of zooids per whorl 10-12.

Remarks. The extreme fragility of the ovicell calcification

makes it impossible to treat specimens with bleach before

electron microscopy. The striated ovicells are similar to those

Figure 17. T. fecundus sp. nov. A, NMV F99009, paratype. Detail of orifice and adapical pore, scale = 50 pm. B-D, NMV F99007, holotype. B,

Adapical view of colony with ovicells and lunate root pores, scale = 500 pm. C. detail of orifices and avicularia, note one adapical pore with

developing ectooecial lamina (arrowed), scale = 200 pm. D, ovicells, orifices and lunate root pore, scale = 200 pm. E, NMV F99009, paratype.

Adapical view of small colony with associated “anascan” ancestrula (arrowed), scale = 500 pm. F, NMV 99008, paratype, detail of ovicell,

showing marginal ectooecium with lateral lappets and ridged entooecium, scale =100 pm.

Australian Conescharellinidae (Bryozoa)

167

Figure 19. Trochosodon asymmetricus sp.nov. NMV F99011, paratype. A, detail of orifices and asymmetrically placed, tubular adapical pores,

scale =100 pm. B, adapical region showing rounded root pores and avicularia, scale = 200 pm. C, detail of marginal peristomes, orifices, adapi-

cal pores, and avicularia, scale = 200 pm.

Figure 18. Trochosodon asymmetricus sp.nov. NMV F99011, paratype.

Adapical view of colony, scale = 1 mm.

figured in Trochosodon optatus by Harmer (1957: 747, pi. 48

figs 16-18, text-figs 77, 78). As the locality from that these

colonies were collected is off the north-west coast of Australia,

it is therefore not very remote from the type locality of

T. optatus, from the coast of Java ( Siboga stn 318, Kangeang

Island, 88 m). However, examination of two of the colonies

from this station (BMNH 1964.3.2.12 part), shows that they

differ in having raised, radial series of zooid orifices, minute

avicularia, and only rare, adapical, lunate root pores. The

principal differences occur in the relationships and position of

the ovicells, that are not exactly as described by Harmer (1957).

They are, in fact, asymmetrically developed, like the ovicells of

C. striata Silen (1947) but, unlike that and other similar

species, have no obvious orifice. Instead, the ovicell opens into

the base of the peristome through a laterally placed foramen.

The peristome is long and tubular and completely obscures the

ovicell opening. The frontal entooecium is striated, as figured

by Harmer, and resembles that of C. striata and T. fecundus.

The ectooecial wall of the ovicell of T. optatus is closely

apposed to the walls of both the neighbouring peristomes; the

ovicells are wedged in between them and difficult to observe.

The only other species observed in that the ovicell orifice

opens into the peristome is T. praecox (see below), and that has

symmetrical ovicells.

The occurrence of an ancestrula with seven marginal spines

(Fig. 17E) on the adapical centre of two colonies is unique.

They are not referable to Trochosodon ; this suggests that they

are extraneous and belong to another, possibly “anascan” species.

T. fecundus is known only from north-western Australia

from 20 m.

Trochosodon asymmetricus sp. nov.

Figures 18, 19A-C

Holotype. NMV F99010, stn SLOPE-6 (colony with root).

Paratypes. NMV F99011 (figured), stn SLOPE-6 (1 worn colony).

Other specimens. NMV F101969, stn SLOPE-7 (1 colony).

Etymology, asymmetros (Gr.) - without symmetry, referring to

the position of the adapical pore.

168

P. E. Bock and P. L. Cook

Diagnosis. Trochosodon with radial series of peristomes, alter-

nating with minute, rounded avicularia. Zooid orifices deeply

concealed. Adapical pore asymmetrically placed. Root pores

circular.

Description. Colony domed, very small, wider than high;

orifices apparently arranged radially; calcification granular.

Peristomes raised, tubular, with intervening radial series of

minute, rounded avicularia; occasional, asymmetrically placed

adapical avicularia; bar without ligula. Primary orifice oval,

deeply concealed, with a short, rounded sinus. Adapical pore

tubular, present in peripheral and subperipheral zooids,

asymmetrically placed inside the margin of the peristome.

Ovicells inferred to be asymmetrical. Root pores adapical,

circular, with a rim and 1 adjacent avicularium. Antapical sur-

face with occasional short radial series of isolated cancelli,

derived from the frontal septular pores of the antapical surface

of the zooids.

Colony diameter 2.5 mm, height 1.5 mm, 5 whorls of 8-9

zooids per whorl.

Remarks. T. asymmetricus is the only species among those

examined (except T. optatus, see above, and a few zooids of

C. stellata ), that exhibits an asymmetrically placed adapical

pore. No ovicells have been found but it may be inferred that

these, too, would be in an asymmetrical position between the

rows of zooid orifices, as are the ovicells of T. optatus Harmer

(1957), together with those of C. striata, C. brevirostris and

C. longirostris of Silen (1947), as well as the specimens

described by Harmer (1957) assigned to C. catella Canu and

Bassler (1929). The tubular appearance of the adapical pore

resembles that figured by Livingstone (1925) in “C. crassa ”.

There are more zooids per whorl than in Trochosodon

anomalus but there are several closely similar characters shared

by these two species. Both have finely tuberculate calcification

and similar radial series of avicularia alternating with the

orifices. The primary orifice is also almost identical in appear-

ance (compare Figs 19 A, 24F). However, the adapical pores are

completely different in position, so it is inferred that the types

of ovicells would be an important distinction between the two

taxa. T. asymmetricus occurs from two adjacent stations from

the New South Wales slope, from 770 to 1096 m.

Trochosodon diommatus sp. nov.

Figures 20, 21A-C

Holotype. NMV F99012, figured specimen, stn SLOPE-7.

Paratype. NMV F99013, F99014, figured specimens, stn SLOPE-7.

Other specimens. NMV F101970, stn SLOPE-6 (4 colonies, 3 very

young); NMV F101971, stn SLOPE-7 (22 colonies, 10 with roots);

NMV F101972, stn SLOPE-45 (1 colony with root).

Etymology, di - two and ommatos - an eye (Gr.), referring to

the paired antapical peristomial avicularia.

Diagnosis. Trochosodon with stellate, radial peristomes, calci-

fication smooth to finely tuberculate. Zooid orifices deeply

concealed, with a narrow sinus. Frontal avicularia minute;

a prominent pair on the antapical surface of the marginal

peristomes.

Figure 20. Trochosodon diommatus sp. nov. NMV F99012, holotype.

adapical view of colony showing lunate root pores, scale = 500 pm.

Description. Colony stellate, fairly flat, distinctly wider than

high, with prominent marginal peristomes. Orifices quincuncial

at first, becoming radial. Primary orifice at the base of the long,

tubular but not prominent peristome, with an elongate, fairly

narrow sinus and large, paired condyles. An adapical pore

present on the edge of the peristome of some peripheral zooids.

Avicularia single, lateral and antapical between the peristomes,

rostrum semicircular, with a bar but no ligula; other small avic-

ularia scattered. Lunate root pores frequent in the adapical

region, each with a pair of avicularia laterally. Antapical sur-

face with marginal pores and avicularia; a pair of avicularia on

the antapical surface of each peristome (cf. C. ocellata and

C. eburnea).

Colony with up to 4 whorls and 4-5 zooids per whorl.

Diameter up to 4.7 mm, height up to 1.5 mm.

Remarks. Ovicells have not been seen in T. diommatus but the

central position of the adapical pore suggests that they would

be symmetrical, like those of T. fecundus, rather than asymmet-

rical, as in T. asymmetricus. Several colonies from stn SLOPE-

7 have roots present; these are 0.5-1 .0 mm long. T. diommatus

is easily distinguished by the presence of the pair of minute

avicularia on the antapical side of the marginal zooid peris-

tomes. It resembles two other species in the presence of antapi-

cal peristomial avicularia. It differs completely from fossil

C. ocellata in dimensions and arrangement of the zooid ori-

fices, that have a longer, more acutely subtriangular sinus. It

differs from C. eburnea in its long peristomes and narrow ori-

ficial sinus, as well as the form of its root pores. T. diommatus

Australian Conescharellinidae (Bryozoa)

169

Figure 21. Trochosodon diommatus sp. nov. A. NMV F99012, holo-

type. Detail of orifices and adapical pores, scale = 200 pm. B, NMV

F99013, paratype, antapical view of marginal peristomes, showing

paired avicularia, scale =200 pm. C. NMV F99014, paratype, detail of

peristome, orifice, and lunate root pores, scale = 200 pm.

has a similar distribution to T. asymmetricus, with the addition

of a record from 800 m depth off Tasmania.

Trochosodon aster sp. nov.

Figures 22A-C, 23

Trochosodon sp. 1. — Cook and Lagaaij, 1976, pi. 1 figs 3, 4.

Holotype. BMNH 1976.1.6.2 part, Challenger stn 185, Cape York,

Queensland, 279 m.

Paratypes. BMNH 1976.1.6.2. part (20 colonies) and BMNH

1969.1.2.2 (7 colonies). NMV F99015, F99016, F99017, and F101973,

same locality (7 colonies).

Etymology, aster (L.) - a star, referring to the budding pattern.

Diagnosis and description. Colonies very small, stellate, bud-

ded in alternating zooid triads early in astogeny, orifices

becoming radial later, fairly flat, but mamillate and raised cen-

trally. Primary orifice almost circular, with a wide sinus, usual-

ly obscured by the elongated peristome, that has a pair of small,

rounded, lateral avicularia. Adapical pores present, root pores

lunate, rare. Calcification mamillate, on adapical and antapical

surfaces.

Colony diameter up to 2 mm, height 0.3 mm, number of

whorls up to 4 and 3-4 zooids per whorl.

Remarks. The colonies from Cape York are heavily calcified

and often somewhat worn. They range in size from 0.25 mm to

nearly 2 mm in diameter and have long marginal peristomes

that bear small avicularia laterally. One colony, figured by

Cook and Lagaaij (1976), shows lunate root pores among the

adapical mamillae. T. aster resembles T. pacificum Lu (1991:

74, pi. 21 fig. 4) from the South China Sea but differs in the

presence of lunate root pores and minute lateral peristomial

avicularia. T. aster also has some characteristics similar to

those described for T. linearis from the East Indies by Harmer

(1957). Two of his specimens have been examined (BMNH

1964.3.2.10, Strait of Makassar, Siboga stn 88, 1301 m, and

1964.3.2.11 , the Banda Sea, stn 227, 2081 m). These are pro-

portionally larger than T. aster, with bilabiate peristomes. One

colony was figured by Harmer (1964.3.2.11, pi. 48 fig. 14,

text-fig.75), who gave a very detailed description of the early

astogeny. The colony has a central, rounded root pore. Canu

and Bassler’s (1929) unique type specimen of T. linearis was

dredged from 635 m depth, from Borneo. The description is not

adequate to decide its synonymy with Harmer’s specimens, that

he could not have compared with the type.

Trochosodon anomalus sp. nov.

Figures 24A-F

Holotype. NMV F99018 stn SLOPE-7.

Paratypes. NMV F99019, stn SLOPE-7 (2 colonies).

Other specimens. BMNH 1976.1.6.2, Challenger stn 185, Cape

York, Australia, 279 m (26 colonies); NMV F101974, Challenger stn

185 (10 colonies).

Etymology. From anomalos (Gr.) - irregular, inconsistent, deviat-

ing, with reference to the combination of character states found

in several genera, that are uniquely possessed by this species.

170

P. E. Bock and P. L. Cook

Figure 22. A-D Trochosodon aster sp. nov. A, NMV F99015, paratype, young colony showing alternating triad structure; root pores and avic-

ularia arrowed, scale = 200 pm. B, NMV 99016, paratype, older colony, scale = 500pm. C, NMV P99017, paratype, colony showing adapical

calcification, scale = 500 pm.

Figure 23. Trochosodon aster sp. nov. NMV F99015, paratype, Detail

of orifice and avicularia, scale =100 pm.

Diagnosis and description. Colonies very small, less than 2 mm

in height and diameter but appearing to be higher than wide.

Calcification mamillate. Zooids arranged in alternating whorls,

each of 3 zooids, appearing to be in radial series; peristomes

elongated and prominent. Primary orifice with a shallow,

rounded sinus and paired condyles, adapical pore symmetric-

ally placed on the edge of the peristome. Avicularia paired,

lateral, widely separated from the orifices, alternating in radial

series; rostra rounded, bar without a ligula. Adapical region

with avicularia and small rounded pores; antapical region with

a few avicularia only.

Colony diameter 0.5-1. 5 mm, height 0.5-1. 5 mm, number

of whorls 2-4, number of zooids per whorl 3.

Remarks. Specimens of T. anomalus are of great interest as they

include characteristics “typical” of both Trochosodon and

Conescharellina\ in some features they even resemble species

of Batopora, from which they are readily distinguished by the

presence of an adapical pore. The arrangement of the radial

series of avicularia suggests assignment to Conescharellina but

the lack of basal cancelli and the presence of prominent,

tubular peristomes allow its inclusion here with other species of

Trochosodon.

The numerous colonies from Cape York indicate that the

distribution of this tiny species extends from Queensland to

New South Wales, from lower shelf to slope depths. The simi-

larities between T. anomalus and T. asymmetricus are described

above.

Trochosodon praecox sp. nov.

Figures 25A-F

Trochosodon sp. 2. — Cook and Lagaaij, 1976, pi. 1 figs 5, 6.

Trochosodon sp. — Cook, 1981, pi. C fig. 4.

Holotype. BMNH 2003.11.27.1 (specimen figured by Cook and

Lagaaij, 1976 and Cook, 1981), Challenger stn 185, Cape York,

Queensland, Australia, 279 m.

Australian Conescharellinidae (Bryozoa)

171

Figure 24. Trochosodon anomalus sp.nov. A-B, NMV F99018, holotype. A, lateral view of colony, direction of growth arrowed, scale = 500 _m.

B, antapical view showing developing zooid, scale = 500 pm. C-D, BMNH 2003.11.27.2. C, lateral view of colony, growth direction arrowed,

scale = 500 pm. D, antapical view, scale = 500 pm. E-F, NMV F99018, holotype. E, adapical region showing root pores and avicularia, scale =

200 pm. F, detail of orifices and avicularia, scale = 200 pm.

Paratypes. BMNH 2003.11.27.4 (as above), 1969.1.2.1, 1976.1.6.2

(part); NMV F99020-F99022. (67 colonies in total).

Etymology, praecox (L.) - precocious, immature, referring to

the production of ovicells at the earliest astogenetic stages in

these minute colonies.

Diagnosis and description. Colonies minute, domed, with

mamillate calcification, that forms a raised mound adapically

and covers the antapical surface. Primary orifice obscure, with

a rounded sinus. Peristomes elongated; tubular and marginally

prominent. Root pores adapical, rare, rounded. Lateral peris-

tomial avicularia paired, very small, rostrum rounded. Ovicells

developed on zooids of the second and third whorls, symme-

trical, globular, very well calcified, opening into the peristome

through a foramen. Ectooecium narrow, marginal; entooecium

with frontal and marginal pores. Antapical surface granular and

mamillate.

Colony diameter 0.50-0.80 mm, height 0.25-0.50 mm, number

of whorls 2, number of zooids per whorl 3.

Remarks. T. praecox is known from more than 60 specimens,

retrieved from one of the unstudied sediment samples from the

Challenger collection, stored in the the Natural History

Museum Mineralogy Department. Busk (1884) reported no

bryozoan specimens from stn 185 from Cape York. Like T.

optatus, T. praecox has ovicells that open into a tubular peris-

tome but are symmetrical in development. They resemble those

of C. africana Cook, 1966 (also Cook, 1981), that also has ovi-

cells that differ from the ovicells observed in most other species

in their relatively robust calcification. Of the 68 specimens

examined, 17 have at least one completely developed ovicell.

Some of the smallest colonies have two or three ovicells, devel-

oped on second or third astogenetic zooid generations.

Although the specimens are all worn, scanning electron

172

P. E. Bock and P. L. Cook

Figure 25. T. praecox sp. nov., Cape York. A, NMV F99020, paratype, detail of orifices and avicularia. with two complete ovicells and one

peristomial foramen (arrowed), scale = 200 pm. B-C, NMV F99022, paratype, B, marginal zooids of colony with complete ovicell, avic-

ularia arrowed, scale = 200 pm. C, detail of ovicells, showing ectooecial margin and central foramina of entooecium (arrowed); note avicularia,

scale =100 pm. D-E, NMV F99021, paratype. D, young colony with one peristomial foramen, root pores arrowed, scale = 200 pm. E, detail of

foramen and adapical pore (arrowed), scale = 100 pm. F, NMV P99022, paratype. Colony with one complete and two developing ovicells

(arrowed), scale = 200 pm.

microscopy has revealed details of ovicell development. Zooids

apparently develop an elongated peristome, with a foramen on

its adapical surface. This is in contact with an adapical pore

(Fig. 25E) at its edge. An ectooecial and an entooecial lamina

then grow together, one “below” the other, from the adapical

pore in the adapical direction (Fig. 25C).The two laminae then

curve in an antapical direction, forming a capsule. The fusion

of the laminae with the lateral and antapical edges of the fora-

men finally closes it, forming the complete ovicell (Fig. 25B).

It should be noted that this does not produce a peristomial ovi-

cell, the walls of which are expansions of the frontal shield. The

ovicells of T. praecox comprise a separate development of

ectooecium and entooecium. Other characteristics that are

distinct enough for colonies to be recognised from additional,

better preserved material, should it ever become available, are

also revealed by scanning electron microscopy. The minute size

of the colonies of T. praecox is comparable to those of accom-

panying foraminiferans; Cook (1981) noted the close similarity

in appearance among them.

The occurrence of reproductive precocity in interstitial

bryozoans with very small colonies has been described and

discussed by Winston and Hakansson (1986: 43).

Australian Conescharellinidae (Bryozoa)

173

Trochosodon gordoni sp. nov.

Trochosodon multiarmatus . — Gordon, 1989: 83, pi. 49 figs D-F

(not Bipora multiarmata Maplestone, 1909: 268).

Etymology. Named for Dr Dennis P. Gordon.

Description. Colony domed, wider than high, calcification fine-

ly mamillate. Zooid orifices in quincunx, sinus rounded, peris-

tomes raised slightly marginally. Rounded avicularia and large

frontal septular pores scattered among orifices. Circular root

pores adapical. Antapical surface with a few avicularia.

Remarks. Gordon (1989) identified specimens from New

Zealand as Maplestone’s species and assigned them all to

Trochosodon. He appears to have confused the dimensions of

his colonies with those of Conescharellina multiarmata , that

are always “higher than wide”, not “typically wider than high”,

as he described. The broad orifice sinus of the New Zealand

species is also quite unlike that of C. multiarmata. Gordon’s

species lacks any basal cancelli and has one or two large, cir-

cular, central rhizoid pores adapically. It appears to be referable

to Trochosodon but certainly not to Conescharellina multiar-

mata. Gordon figured a specimen from Station P927

(40°50.1'S, 168°14.8'E, 1005-1009 m, western South Island,

New Zealand) and reported it from numerous other localities

from southern New Zealand, from a range of 540 to 1676 m

depth.

Crucescharellina Silen, 1947

Crucescharellina Silen, 1947: 44.

Agalmatozoum Hanner, 1957: 757.

Type species. Crucescharellina japonica Silen, 1947 (original

designation).

Description. Colonies are cruciform or star-shaped and may

have branches that bifurcate terminally. The adapical zooid

orifices are sinuate and interspersed with lunate or rounded root

pores. The antapical growing edges are positioned at the limits

of the branches but an antapical surface, that is the equivalent

of the exposed frontal wall of conescharelliniform colonies, is

also continuous and present on the “lower, non-orificial side”

of colonies. It is inferred that the colonies live, in fact, with this

antapical surface upward with the orifices directed downward,

because the rhizoids that occur among them are inferred to

anchor the colonies above or into the surface of the bottom

sediments. Rounded or acute avicularia occur, that are occa-

sionally large and spathulate. The orifices possess an adapical

pore but ovicells have not been seen. Roots were figured in

C. japonica by Silen (1947: pi. 1 fig. 11) and the position of the

root pores suggests that the mode of life is similar to that

inferred for the genus Euginoma (Hayward 1978), that also

occurs from abyssal depths (d’Hondt and Schopf ,1984).

Remarks. Crucescharellina was introduced by Silen (1947) for

C. japonica from near the Goto Islands, Japan, from a depth of

175 m. Only one colony was found; it was stellate but each

branch originated from a narrow neck, one or two zooids in

width. The branches rapidly expanded and then bifurcated,

each subbranch starting with one or two zooids. The

subbranches also expanded rapidly, so that within two as

togenetic generations, the segments were 4 zooids wide. Lunate

root pores were present but these were not associated with

branch bifurcations and no large, spathulate avicularia were

described. Gordon and d’Hondt (1997: 73, figs 221-223)

described “C. japonica ” from the Philippines from 640-668 m.

They too, had only one colony. It differed in having much less

expanded branches, regular lunate root pores, and rare large

axillary avicularia. The primary orifice had a shallow sinus and

paired condyles. Silen (1947: 44) stated that he referred

Trochosodon decussis Canu and Bassler (1929: 495, pi. 71 figs

7-10, from 456 m, east of Mindanao in the Philippines) to his

genus Crucescharellina. Harmer (1957) was unaware of Silen’s

work, that was not available to him during the war of

1939-1945, and introduced Agalmatozoum for Trochosodon

decussis Canu and Bassler (1929). Colonies of this species

were cruciform, with triserial branches, and were described

with lunate root pores and an elliptical secondary orifice.

Avicularia or small pores were present antapically but no large

avicularia were mentioned in the original description. Harmer

(1957) listed more than ten colonies of A. decussis from seven

localities in the Sulu, Banda, and Celebes Seas. The depths

were nearly all abyssal, ranging from 535 to 3112 m. The

branches of the colonies were mostly biserial and the root pores

were circular, placed regularly at bifurcations, and surrounded

by a ring of small avicularia. In addition, large, axillary spathu-

late avicularia sometimes occurred on the lateral sides of

branches. The species from the Siboga area described by

Harmer (1957) as A. decussis strongly resembles

Crucescharellina australis from Australia described below, not

the original form from the Philippines described by Canu and

Bassler (1929). Gordon and d’Hondt (1997: 74, figs 224-227)

introduced another very similar stellate species, C. aster, with

biserial branches, from several New Caledonian and New

Zealand localities at a depth range of 760 to 1573 m. The root

pores were central and rounded but no large avicularia were

present. Their material included numerous colonies, that they

noted resembled “clusters of snowflakes”. A single preparation

of a colony in the Natural History Museum collection (BMNH

1963.8.18.18) closely resembles the description of C. aster but

has slightly more extended, spiny peristomes. The specimen is

from Challenger stn 169, off New Zealand (37°34'S, 179°22'E,

1295 m), a station that was not mentioned by Busk (1884).

Gordon (1989: 84, pi. IE figs 50B-E) described another biser-

ial species, C. jugalis, from northern New Zealand, from a

depth range of 1217-1357 m. The colonies were irregularly

branched but had circular root pores very similar to those of the

Australian C. australis and A. decussis sensu Harmer (1957).

Although there is no doubt of the synonymy of the two

genera Crucescharellina and Agalmatozoum, there are uncer-

tainties as to the identity of the various taxa referred to them in

these previous descriptions. Among other records. Cook (1981)

figured one of two very young, cruciform colonies from Cape

York, from 279 m (BMNH 1976.1.6.2, part), as Agalmatozoum

species. These, with the specimens of C. australis described

here from Point Hicks, Victoria and from eastern Tasmania,

remain the only records of Crucescharellina from Australian

waters to date.

174

P. E. Bock and P. L. Cook

Labracherie and Sigal (1975) mentioned a form similar to

Crucescharellina obtained from Lower Eocene samples col-

lected from a deep-sea drilling south of Madagascar

(33°37.21'S, 45°09.60'E, 1030 m). This was not described fur-

ther but is not too remote from the Recent south-west Indo-

Pacific records and, unlike the European Eocene species men-

tioned above, may represent an early form of

Conescharellinidae.

Crucescharellina australis sp. nov.

Figures 26A-E

Holotype. NMV F99024, stn SLOPE-27.

Paratypes. NMV F99025, stn SLOPE-27 (8 colonies).

Other specimens. NMV F101975, stn SOELA-S03/84/74, E.

Tasmania, 320 m.

Etymology, australis (L.) - southern, referring to the distribu-

tion of the species.

Diagnosis. Crucescharellina with biserial branches; zooid ori-

fice with a shallow sinus. Avicularia small and rounded; occa-

sionally large, axillary spathulate. Root pores circular, placed at

branch bifurcations.

Description. Colonies probably cruciform, present material

with four branches. Branches biserial, bifurcating at each fourth

to fifth astogenetic generation. Primary orifice with a wide,

shallow sinus, and minute condyles; obscured at the base of a

long peristome. These are sometimes raised antapically (i.e.

towards the end of a branch) and may have 4-5 small, spinous

processes on their margins. Avicularia small, rounded, near

each peristome, bar without a ligula. Rare enlarged, spathulate

avicularia placed in the axils between branches. Root pores reg-

Figure 26. Crucescharellina australis sp.nov. A-D, NMV F99024, holotype. A, colony from adapical surface, scale = 2 mm. B, detail of branch

showing orifices, avicularia and root pore (arrowed), scale = 500 pm. C, detail of root pore, scale = 200 pm. D, detail of orifice with adapical

pore, direction of growth arrowed, scale =100 pm. E, NMV F99025, paratype, large spatulate axillary avicularium, scale =100 pm.

Australian Conescharellinidae (Bryozoa)

175

ularly placed at bifurcation of branches; circular, with a

distinct rim, surrounded by 3-4 small avicularia. Adapical pore

within the calcification of the edge of the peristome, present in

many zooids, but no ovicells seen. Antapical surface finely

granular, with approximately one small rounded avicularium

per zooid.

Branches 0.6 mm width, length 6 mm.

Remarks. The specimens from stn SLOPE-27 comprise nine

colony fragments. The largest show evidence of having once

been cruciform but only four complete branches are now

present. Two colonies have enlarged, spathulate avicularia in

the axil between two branches, very similar to that shown in his

“A. decussis ” by Harmer (1957, pi. 49 fig. 13). The colony

structure, type and distribution of root pores, and the large avic-

ularia of C. australis make it virtually certain that it is con-

specific with some specimens of A. decussis sensu Harmer but

it is distinct from T. decussis Canu and Bassler (1929) and the

other species mentioned above. Some fragmentary preparations

of Harmers’ Siboga material, labelled A. decussis , from the

BMNH collection, have been examined, The specimens are all

slightly worn, and none possesses large avicularia. A cruciform

colony and fragments from Siboga stn 211 (BMNH

1964.3.2.21, south of Celebes, 1158 m.), most closely resem-

bles C australis. Other fragments from Siboga stn 102 (BMNH

1964.3.2.23, Sulu Archipelago, 535 m.) differ in having triser-

ial branches and distinctly elongated avicularia near the zooid

orifices. Some partially decalcified fragments from Siboga stn

221 (BMNH 1964.3.2.25, Banda Sea, 2798-3112 m.) also

resemble C. australis but have raised peristomes on the anta-

pical side of the orifices. They possess t hr ee long roots (over 20

mm), that emanate from the root pores. It is possible that

Harmer’ s material, identified as A. decussis , may belong to

more than one species. Circular root pores surrounded by small

avicularia also occur in Conescharellina eburnea, C. plana,

C. perculta and C. humerus.

Zeuglopora Maplestone, 1909

Zeuglopora Maplestone, 1909: 272. — Canu and Bassler, 1929:

510.— Harmer, 1957: 755.

Type species. Zeuglopora lanceolata Maplestone, 1909

(original designation).

Description. Colony similar to Flabellopora, ligulate, appar-

ently composed of 2 laminae but in fact consisting of a pair of

alternating and interdigitating expanses of frontally budded

zooid series. Single or small groups of marginal zooids

enlarged and prominent, forming a serrated edge, occasionally

with enlarged avicularia. Orifices oval, with paired condyles

forming a subtriangular antapical sinus; peristome tubular. A

rounded adapical pore is present but ovicells are unknown.

Avicularia usually small, rounded, with a bar but no ligula.

Colony anchored by 1 or 2 roots, arising from lunate pores in

the adapical region.

Remarks. Both Canu and Bassler (1929) and Harmer (1957)

analysed the colony structure of Zeuglopora and maintained its

distinction from Flabellopora.

Zeuglopora lanceolata Maplestone, 1909

Figures 27, 28A-C

Zeuglopora lanceolata Maplestone, 1909: 272, pi. 78, fig. 11. —

Harmer, 1957: 757.

Bipora lanceolata. — Livingstone, 1924: 211.

Specimens examined. NMV F99026, 1 colony labelled by

Maplestone, probably part of the type material from NSW.

Diagnosis and description. As for the genus, serrated edges of

colony formed by prominent zooids, that occur in alternating

unequal pairs. Primary orifice with a subtriangular sinus and

well developed paired condyles; obscured at the base of a

tubular peristome, that is most prominent adapically. A

rounded adapical pore, at a little distance from the edge of the

peristome, is present in some central and antapical zooid

orifices. Surface of zooids mammilate, interspersed with

minute, rounded avicularia, with a bar but no ligula. Root pores

lunate, adapical, paired, large and surrounded by extrazooidal

calcification.

Colony length 7 mm, breadth 2.25 mm. Number of astoge-

netic generations 12-13, number of zooids per generation 10.

Remarks. The single colony from New South Wales resembles

Maplestone’s (1909) description. The orifices of the enlarged

marginal zooids are surrounded by up to five small avicularia

and resemble the root pores of C. eburnea and C. plana.

However, the adapical end of the colony shows that the actual

root pores resemble those of Flabellopora and consist of large

lunate pores surrounded by massive, secondary extrazooidal

calcification. The adapical pore is large and a little offset in

position towards the colony margin. It seems probable that any

ovicell would be slightly asymmetrically placed. Harmer

(1957: 737) examined a colony that was part of Maplestone’s

(1909) type material in the BMNH collection (BMNH

1909.11.12.3). It has a large adapical foramen that was filled

with detritus. It seems almost certain that this colony actually

possessed paired, lunate adapical root pores like other speci-

mens (Fig. 28C). There are no additional records of Z. lanceo-

lata from Australia but Canu and Bassler (1929: 511, pi. 75, fig.

6) described a very similar colony from deep water (630 m)

north-east of Borneo as Z. lanceolata, an identification

accepted by Harmer (1957). The locality was remote from

Australia; the figured colony is slightly narrower than those

from Australia, with more prominent marginal zooids than the

type specimens. The character of the primary orifices, root

pores and avicularia are uncertain. Harmer (1957) also

described a new species, Z. arctata, from two minute colonies

from 82 m off Java, where the bottom sediment consisted of

fine grey mud. Each colony had large marginal avicularia, and

a long single root emanating from the adapical region. Cadee

(1987: 52) noted the occurrence of several hundred colonies of

Z. arctata together with an undescribed species from soft-

bottom sediments in the Banda Sea but gave no detailed

descriptions. Lu (1991) described several species of Zeuglo-

pora (as Bipora) from the South China Sea. His B. pagoda

(p.70, pi. 18 fig. 2) and B. trinodata (p. 71, pi. 18 fig. 3) both

resemble the unnamed species mentioned by Cadee (1987).

176

P. E. Bock and P. L. Cook

Figure 27. Zeuglopora lanceolata Maplestone, 1909. NMV F99026.

Lateral view of colony, direction of growth arrowed, scale = 1 mm.

Flabellopora d’Orbigny, 1851

Flabellopora d’Orbigny, 1851: 52.

Flabillopora, d’Orbigny, 1852: 186 (bis) (lapsus). — Canu and

Bassler, 1929: 495.— Harmer, 1957: 749.— Silen, 1947: 47.— Lu,

1991: 72.

Type species. Flabellopora elegans d’Orbigny, 1851 (mono-

typy).

Description. Colony leaf-like or trilobed, superficially appear-

ing to be bilaminar, anchored by root systems originating from

lunate pores on the adapical edge. Zooids in alternating and

interdigitating frontally budded series, orifices sinuate, the

sinuses orientated antapically towards the growing edge. A

small adapical pore sometimes present. Avicularia in patterns

among orifices, usually small and rounded, with a bar but no

ligula. Ovicells unknown but presumably originating from

adapical pores.

Remarks. F. elegans was recorded by d’Orbigny (1851: 53)

from about 20 m. “pres de Ouantang et d’ Hainan” in the China

Sea. Later, d’Orbigny (1852: 186 bis) mentioned additional

specimens from “dans le detroit de Malaca et a Manille” [sic].

Harmer (1957: 751) noted that Waters’ (1905: 9, pi. 1 fig. 5)

figured specimen from the d’Orbigny collection was from

Malacca. It was therefore certainly not of the type specimen

and may not even have been of the same species. Waters’ fig-

ure, like those of Conescharellina from the d’Orbigny collec-

tion (see above), was semidiagrammatical and included only

three zooid orifices. A photograph of the type specimen (Taylor

and Gordon, 2002, fig.3D) closely resembles d’Orbigny’s 1852

illustration but provides no details of the primary orifices or

distribution of avicularia. As in the case of Conescharellina, the

generic characters of d’Orbigny’s descriptions and illustrations

are unmistakeable but the details of specific characters are

obscure and require examination and redescription of the type

specimen.

Delicate roots up to 25 mm in length were described by

Harmer (1957), who noted their origin from lunate pores. In

one of the trilobed colonies he illustrated, as F. irregularis (pi.

49 fig. 6), thirteen roots occur along the adapical edge of the

colony. Ovicells are unknown in Flabellopora although

d’Orbigny (1852: 186 bis) mentioned the presence of a “pore

ovarien”. It is not known if this is the equivalent of the “proxi-

mal pore” of Harmer (1957) or the adapical pore, that is now

known to be the origin of ovicells in Conescharellina and

Trochosodon. Harmer (1957: 749, text-fig. 79) illustrated the

central region of a colony expanse, that showed hemispherical

areas of calcification adapically to zooid orifices. The sur-

rounding calcification was raised into “lozenge-shaped” areas,

a term used by Canu and Bassler (1929). Harmer suggested that

each hemispherical calcification might represent the basal part

(i.e. the ectooecium) of an ovicell. He figured and mentioned

the presence of “proximal pores” but did not appear to associ-

ate them with ovicells. Adapical pores have been found fre-

quently in the specimens examined here but they are usually

associated with zooids that are marginal in position; they are

not distributed in the centre of colony expanses, or surrounded

by raised “lozenges”.

Australian Conescharellinidae (Bryozoa)

177

Figure 28. Zeuglopora lanceolata Maplestone, 1909. NMV F99026. A, detail of orifice with adapical pore, scale = 100 pm. B, detail of large,

marginal zooid orifice with surrounding avicularia, direction of growth arrowed, scale = 200 pm. C. adapical region showing lunate root pores,

scale =200 pm.

Flabellopora umbonata (Haswell, 1881)

Figures 29A-C

Eschara umbonata Haswell, 1881: 41, pi. 2 figs 5, 6.

Bipora umbonata . — Whitelegge, 1887: 345. — Livingstone, 1924:

209. — Livingstone, 1926: 98, pi. 5 figs 4, 5.

Bipora mamillata Maplestone, 1909: 270, pi. 77 fig. 7.

Conescharellina mamillata . — Bretnall, 1922: 191.

Specimens examined. NMV F99395, stn SLOPE-40 (1 colony);

NMV FI 01 976, stn GAB -020 (2 young and 1 trilobed colony with

roots, plus fragments); NMV F101977, stn GAB-030 (2 young and

2 trilobed colonies with roots, plus fragments); NMV F101978, stn

GAB-045 (1 trilobed colony with roots, plus fragments); NMV

F101979, stn GAB-056 (1 colony with fragments); NMV F101980, stn

GAB-067 (1 trilobed colony); NMV F101981, stn GAB-074

(5 fragments); NMV F101982, stn GAB-084 (2 trilobed colonies plus

fragments); NMV F101983, stn GAB-088 (1 colony plus

fragments); NMV FI 01 984, stn GAB -093 (1 large colony); NMV

F101985, stn GAB-112 (1 colony); NMV F101986, stn GAB-117

(1 trilobed colony plus fragments); NMV F101987, stn GAB-119

(2 young and 3 trilobed colonies); NMV F101988, stn GAB- 128

(1 trilobed colony); NMV F101989, stn SOELA-S03/84/74, E.

Tasmania, 320 m (1 colony).

Description. Colony leaf-shaped, sometimes trilobed. Zooid

frontal shield continuous, without zooid borders; calcification

smooth with umbonate mamillae occurring among the orifices

and the avicularia. Orifices almost circular, patent, with a

rounded sinus, peristomial rim raised, narrow; adapical

pores present. Avicularia small, rostra subtriangular or rounded,

bar without a ligula. Septular pores rare, scattered. Root

pores lunate, on the adapical edge, surrounded by thickened

calcification.

Remarks. Livingstone (1924, 1926) and Harmer (1957) exam-

ined specimens from Queensland that were reported to be from

Haswell’s original material, although only fragments of this

were preserved. Maplestone’s (1909) type specimen of Bipora

mamillata was unique but Livingstone (1924) mentioned

“types” that may have had a different provenance. Both

Livingstone and Harmer were convinced that F. umbonata was

identical with B. mamillata , Maplestone (1909) however had

noted some differences, both within Haswell’s suite of speci-

mens and between them and his colonies from New South Wales.

The numerous specimens examined here are often slightly

worn and very few possess roots. There are small differences

among specimens but these seem to be the result of astogenet-

ic position, ontogenetic thickening and wear. Some specimens

have larger umbonate mamillae among orifices than others and

some have larger avicularia but none of these differences is cor-

related with locality. The specimens from stations GAB -020,

GAB-030 and GAB-119 include several very young colonies.

These are lanceolate and consist principally of parallel series of

antapically directed zooids with few laterally inclined series.

Later growth illustrated the development of paired lateral lam-

inae, giving the typically trilobed shape. One regenerated

colony from station GAB -093 has a diameter of 28 mm and has

developed nine thickened rays from an irregular central area. It

is possible that other species of Flabellopora occur in

Australian waters but have not yet been recognised.

Ptoboroa Gordon and d’Hondt, 1997

Ptoboroa Gordon and d’Hondt, 1997: 70, pi. 47F, G, 48A.

Type species. Trochosodon pulchrior Gordon, 1989: 81

(original designation).

178

P. E. Bock and P. L. Cook

Figure 29. A-C, Flabellopora umbonata (Haswell, 1881). NMV

F99395, stn GAB-056 . A. Colony, growth direction arrowed, scale =

1 mm. B. Antapical region showing orifices and avicularia, adapical

pore arrowed, scale = 200 pm. C. Adapical region, root pore arrowed,

scale = 1 mm.

Remarks. No species of Ptoboroa has been found in the present

collections. P. pulchrior from New Zealand is strikingly similar

to a species of Batopora from stations SLOPE-6 and SLOPE-7

but differs in the possession of an adapical pore, indicating its

closer association with the Conescharellinidae, particularly

Trochosodon , in the development and form of its ovicells (Bock

and Cook, in press).

Summary and discussion

Six of the seven genera of Conescharellinidae ( Cones -

charellina, Bipora, Trochosodon, Zeuglopora, Cruces-

charellina and Flabellopora ) are represented in Australian

waters (see Appendix). The seventh genus, Ptoboroa, is at pres-

ent known only from New Zealand and New Caledonia (Bock

and Cook, in press). Records occur from north-west Australia,

Cape York, Queensland, the coasts of New South Wales and

Victoria, and Western Australia to Tasmania. Numerous Tertiary

samples from Victoria and South Australia have been examined.

Although many species, particularly of Conescharellina,

have been described from the western Pacific region by Canu

and Bassler (1929), Silen (1947) and Harmer (1957), the pres-

ent collections include an unexpectedly high proportion

(approximately 66%) of new taxa. There are apparently

several explanations for this diversity. First, the number of

colonies examined is greatly in excess of any other named col-

lection.The total number is more than 1940, of which 52% are

Recent specimens. A few species are represented by only one to

three colonies, occurring from a single locality (for example

Conescharellina perculta, Trochosodon ampulla, T. fecundus

and Zeuglopora lanceolata), and 30 localities provided

specimens of only one species. In contrast, seven taxa are rep-

resented by more than 60 colonies (for example C. cognata

(177), C. plana (120), C. biarmata (98), C. multiarmata (85),

C. diffusa (82), C. ecstasis (71) and T. praecox (67), while stn

SLOPE-7 and stn GAB -020 include specimens of six species

each). The Tertiary accumulations of colonies include 438 of

C. macgillivrayi from seven localities, 360 of C. humerus from

five localities and 102 of C. ocellata from two localities. The

large number of specimens allows comparisons among and

within populations, and subsequent definition of taxa with con-

fidence, within a range of variation. Second, the reports of

Australian species made in the nineteenth and early twentieth

centuries were from fairly restricted geographical and bathy-

metrical localities, mostly from New South Wales. The exten-

sive geographic range of species examined here has revealed

new taxa and also allowed investigation of their variation. For

example, there are differences between populations of C. dif-

fusa, that has a range fom north-west Australia to Tasmania but

its essential characteristics are consistent. Third, the interstitial

or semi-interstitial mode of life, particularly at very deep local-

ities, may reduce the possibility of wide dispersal of some

larvae but the distribution of C. ecstasis, from New South

Wales to Tasmania from a depth range of 400 to 1096 m, sug-

gests that some other factors are involved. Fourth, examination

of colonies using the scanning electron microscope, often for

the first time, has refined definitions of previously described

species and has revealed characters and character states

essential for future investigations.

Australian Conescharellinidae (Bryozoa)

179

The Australian fauna of Conescharellinidae appears to be

quite distinct from the Indo- West-Pacific fauna to the north and

the New Zealand fauna to the east. Further investigations may

show similarities among the deeper water faunas of New

Caledonia and the eastern Australian coast, for example, among

species of Crucescharellina. There are also some tenuous links

with New Zealand illustrated by populations of C. cognata.

Little is known of fossil populations of Conescharellinidae;

only Conescharellina has an established fossil record. The

three most abundant fossil species are not only very similar to

one another in characters, they appear to have little in common

with any of the Recent forms. All have very small colonies that

are apparently astogenetically mature; all have rounded root

pores with a circlet of avicularia. C. ocellata resembles C.

eburnea in possessing a pair of antapical avicularia on margin-

al peristomes but there is no evidence of any descendant

sequence among the specimens examined.

Acknowledgements

Dr K. J. Tilbrook provided the initial impetus for this study by

finding specimens of Maplestone species in the collections of

the Natural History Museum, that were kindly lent by Mary

Spencer Jones. Chris Rowley, Museum Victoria, has been help-

ful in facilitating access to that collection. Samples from

Dampier were from collections made by Dr Gary Poore.

Samples from the GAB series are provided through the

activities of Dr Yvonne Bone (University of Adelaide) and the

Master and crew of RV Franklin. The contribution of CSIRO is

gratefully acknowledged.

References

Accordi, B. 1947. Nuove forme di Briozoi eocenici. Studi Trentini di

Scienze Naturali Acta Geologica 25: 103-110.

Arnold, P., and Cook, P.L. 1997. Some Recent species of the genus

Anaskopora Wass, 1975 (Bryozoa: Cribriomorpha) from

Queensland. Memoirs of the Queensland Museum 42: 1-11.

Banta, W.C. 1972. The body wall of cheilostome Bryozoa, V. Frontal

budding in Schizoporella unicornis floridana. Marine Biology 14:

63-71.

Bock, P.E., and Cook, P.L. 1996. The genus Selenariopsis Maplestone,

1913 (Bryozoa, Ascophorina). Records of the South Australian

Museum 29: 23-31.

Bock, P.E., and Cook, P.L. 2000. Lekythoporidae (Bryozoa,

Cheilo stomata) from the Tertiary and Recent of southeastern

Australia. Memorie di Scienze Geologiche 52: 167-174.

Bock, P.E., and Cook, P.L. 2001. Revision of the multiphased genus

Corbulipora MacGillivray (Bryozoa: Cribriomorpha). Memoirs of

Museum Victoria 58: 191-213.

Bock, P.E., and Cook, P.L, in press. New species of the bryozoan

genera Batopora and Lacrimula (Batoporidae) from Australia.

Proceedings of the Royal Society of Victoria.

Bretnall, R.W. 1922. Studies on Bryozoa 2. 1. On a collection of

Bryozoa from 26-38 fathoms off Norah Head. Records of the

Australian Museum 13: 189-192.

Brown, D.A. 1958. Fossil cheilostomatous Polyzoa from south-west

Victoria. Memoirs of the Geological Survey of Victoria 10: 1-90.

Brown, K.M., Schmidt, R., and Bone, Y. 2002. Observations on eco-

logical adaptations of Lanceopora smeatoni (MacGillivray), from

West Island, South Australia. Pp. 61-65 in: Wyse Jackson, P.N.,

Buttler, C.J., and Spencer-Jones, M. (eds), Bryozoan Studies 2001.

A.A. Balkema Publishers: Lisse, Abingdon, Exton, Tokyo.

Busk, G. 1854. Catalogue of marine Polyzoa in the collection of the

British Museum, II. Cheilostomata (part). Trustees of the British

Museum (Natural History): London, pp. viii, 55-120.

Busk, G. 1884. Report on the Polyzoa collected by H.M.S. Challenger

during the years 1873-1876. Part 1. The Cheilostomata. Report on

the Scientific Results of the Voyage of the H.M.S. “Challenger”,

Zoology 10: 1-216.

Cadee, G.C. 1987. The shallow soft-bottom faunas of the Java Sea and

Banda Sea. Pp. 49-56 in: Ross, J.R.P. (ed.), Bryozoa: Present and

Past. Western Washington University: Bellingham.

Canu, F., and Bassler, R.S. 1917. A synopsis of American Early

Tertiary Cheilostome Bryozoa. United States National Museum

Bulletin 96: 1-87.

Canu, F., and Bassler, R.S. 1927. Classification of the cheilostomatous

Bryozoa. Proceedings of the United States National Museum 69:

1-42.

Canu, F., and Bassler, R.S. 1929. Bryozoa of the Philippine region.

United States National Museum Bulletin 100: 1-685.

Cheetham, A.H. 1966. Cheilostomatous Polyzoa from the Upper

Bracklesham Beds (Eocene) of Sussex. Bulletin of the British

Museum (Natural History) (Geology) 13: 1-115.

Cook, P.L. 1966. Some “sand fauna” Polyzoa (Bryozoa) from Eastern

Africa and the northern Indian Ocean. Cahiers de Biologie Marine

1: 207-223.

Cook, P.L. 1979. Mode of life of small, rooted “sand fauna”

colonies of Bryozoa. Pp. 269-281 in: Larwood, G.P., and

Abbott, M.B. (eds), Advances in Bryozoology . Academic Press:

London.

Cook, P.L. 1981. The potential of minute bryozoan colonies in the

analysis of deep sea sediments. Cahiers de Biologie Marine 22:

89-106.

Cook, P.L. 1985. Bryozoa from Ghana. A preliminary survey. Annales

Musee Royale de VAfrique Centrale, Sciences Zoologiques,

Tervuren 238: 1-315.

Cook, PL., and Bock, PE. 1994. The astogeny and morphology of

Rhabdozoum wilsoni Hincks (Anasca, Buguloidea). Pp. 47-50 in:

Hayward, P. J., Ryland, J.S., and Taylor, P.D., (eds), Biology and

Palaeobiology of Bryozoans. Olsen and Olsen: Fredensborg.

Cook, P.L., and Chimonides, P.J. 1981. Morphology and systematics of

some rooted cheilostome Bryozoa. Journal of Natural History 15:

97-134.

Cook, P.L., and Chimonides, P.J. 1985. Larval settlement and early

astogeny of Parmularia (Cheilostomata). Pp. 71-78 in: Nielsen, C.,

and Larwood, G.P. (eds), Bryozoa: Ordovician to Recent. Olsen and

Olsen: Fredensborg.

Cook, P.L., and Chimonides, P.J. 1987. Recent and fossil Lunu-

litidae (Bryozoa, Cheilostomata), 7. Selenaria maculata (Busk) and

allied species from Australasia. Journal of Natural History 21:

933-966.

Cook, P.L., and Lagaaij, R. 1976. Some Tertiary and Recent

conescharelliniform Bryozoa. Bulletin of the British Museum

(Natural History), Zoology 29: 317-376.

Gordon, D.P 1984. The marine fauna of New Zealand: Bryozoa:

Gymnolaemata from the Kermadec Ridge. New Zealand

Oceanographic Institute Memoir 91: 1-198.

Gordon, D.P. 1985. Additional species and records of Gymnolaemate

Bryozoa from the Kermadec region. Records of the New Zealand

Oceanographic Institute 4: 160-183.

Gordon, D.P. 1989. The marine fauna of New Zealand: Bryozoa:

Gymnolaemata (Cheilostomida Ascophorina) from the western

south Island continental shelf and slope. New Zealand

Oceanographic Institute Memoir 97: 1-158.

180

P. E. Bock and P. L. Cook

Gordon, D.P., and d’Hondt, J.-L. 1997. Bryozoa: Lepraliomorpha and

other Ascophorina from New Caledonian waters. Memoires du

Museum National d’Histoire Naturelle, Paris 176: 9-124.

Gregory, J.W. 1893. On the British Palaeogene Bryozoa. Transactions

of the Zoological Society of London 13: 219-279.

Grischenko, A.V., Gordon, D.P., and Taylor, P.D. 1998 (1999). A

unique new genus of cheilostomate bryozoan with reversed-polari-

ty zooidal budding. Asian Marine Biology 15: 105-117.

Hageman, S.J., Bone, Y., McGowran, B., and James, N.P. 1996.

Bryozoan species distributions on the cool-water Lacepede Shelf,

southern Australia. Pp. 109-116 in: Gordon, D.P., Smith, A.M., and

Grant-Mackie, J.A. (eds), Bryozoans in Space and Time. NIWA:

Wellington.

Harmer, S.F. 1957. The Polyzoa of the Siboga Expedition, Part 4.

Cheilostomata Ascophora II. Siboga Expedition Reports 28d:

641-1147.

Has well, W.A. 1881. On some Polyzoa from the Queensland Coast.

Proceedings of the Linnean Society of New South Wales 5: 33-44.

Hayward, P.J. 1978. The morphology of Euginoma vermiformis Jullien

(Bryozoa, Cheilostomata). Journal of Natural History 12: 97-106.

Hayward, P.J., and Cook, P.L, 1979. The South African Museum’s

Meiring Naude Cruises. Part 9, Bryozoa. Annals of the South

African Museum 79: 43-130.

Hincks, T. 1880. A history of the British Marine Polyzoa. Van Voorst:

London, cxli + 601 pp.

Hincks, T. 1881. Contributions towards a general history of the marine

Polyzoa. (Part VI. Polyzoa from Bass’s Straits continued, no title).

Annals and Magazine of Natural History (5) 8: 122-129 (separate

pp. 63-70).

Hincks, T. 1892. Contributions towards a general history of the marine

Polyzoa. Appendix. Annals and Magazine of Natural History (6) 9:

327-334 (separate pp. 190-197).

d’Hondt, J.-L. 1985. Contribution a la systematique des Bryozoaires

Eurystomes. Apports recents et nouvelles propositions. Annales des

Sciences Naturelles, Zoologie and Biologie Animale (13) 7: 1-12.

d’Hondt, J.-L., and Schopf, T.J.M. 1984. Bryozoaires des grandes pro-

fondeurs recueillis lors des campagnes oceanographiques de la

Woods Hole Oceanographic Institution de 1961 a 1968. Bulletin du

Museum National d’Histoire Naturelle, Paris (4) 6A (4): 907-973.

Jelly, E.C. 1889. A synonymic catalogue of the Recent marine Bryozoa.

Dulau and Company: London. 322 pp.

Kirkpatrick, R. 1890. Reports on the zoological collections made in

Torres Straits by Professor A.C.Haddon, 1888-1889. Hydroida and

Polyzoa. Scientific Proceedings of the Royal Dublin Society, new

series 6: 603-626.

Labracherie, M. 1975. Sur quelques bryozoaires de l'Eocene inferieur

nord-aquitain. Revista Espahola de Micropaleontologia 1:

127-164.

Labracherie, M., and Sigal, J. 1975. Les Bryozoaires cheilostomes des

formations Eocene Inferieur du Site 246 (crosiere 25, Deep Dea

Drilling Project). Pp. 449—4-66 in: Pouyet, S. (ed.), Bryozoa 1974

( Documents de Laboratoires de Geologie Faculte de sciences de

Lyon, HS 3) Universite Claude Bernard: Lyon.

Levinsen, G.M.R. 1909. Morphological and systematic studies on the

cheilostomatous Bryozoa. Nationale Forfatterers Forlag:

Copenhagen. 431 pp.

Livingstone, A. A. 1924. Studies on Australian Bryozoa. No. 1.

Records of the Australian Museum 14: 189-212.

Livingstone, A.A. 1925. Studies on Australian Bryozoa. No. 2.

Records of the Australian Museum 14: 301-305.

Livingstone, A.A. 1926. Studies on Australian Bryozoa. No. 3.

Records of the Australian Museum 15: 79-99.

Livingstone, A.A. 1928. Bryozoa lfom South Australia. Records of the

South Australian Museum 4:11 1-124.

Lu Linhuang 1991. Holocene bryozoans from the Nansha sea area.

Pp. 11-81, 473-486 in: Multidisciplinary Oceanographic

Expedition Team of Academia Sinica to the Nansha Islands (eds),

Quaternary Biological Groups of the Nansha Islands and the

Neighbouring Waters. Zhongshan University Publishing House:

Guangzhou.

MacGillivray, PH. 1895. A monograph of the Tertiary Polyzoa of

Victoria. Transactions of the Royal Society of Victoria 4: 1-166.

Maplestone, C.M. 1904. Tabulated list of the fossil cheilostomatous

Polyzoa in the Victorian Tertiary deposits. Proceedings of the Royal

Society of Victoria (new series) 17: 182-219.

Maplestone, C.M. 1909. The results of deep-sea investigations in the

Tasman Sea. The expedition of the H.M.C.S “ Miner ”, 5. The

Polyzoa. Records of the Australian Museum 7: 267-273.

Maplestone, C.M. 1910. On the growth and habits of the Biporae.

Proceedings of the Royal Society of Victoria (new series) 23: 1-7.

Neviani, A. 1895. Briozoi Eocenici del calcare nummulitico di

Mosciano presso Firenze. Bolletino della Societa Geologica

Italiana 14: 119-129.

Neviani, A. 1901. Briozoi neogenici delle Calabrie. Palaeontographia

Italica 6(1900), 115-266.

d’Orbigny, A. 1851-1854. Paleontologie frangaise. Terrains Cretaces,

V, Bryozoaires. Victor Masson: Paris (1851: 1-188; 1852, 185

bis-472; 1853, 473-984; 1854, 985-1192).

Pizzaferri, C., and Braga, G. 2000. Nuove osservazioni sullo sviluppo

astogenetico di Batopora rosula (Reuss), Bryozoa Cheilostomatida

del Miocene del Pedeappennino Parmense. Annali dei Museo

Civico -Rovereto 14 (1998): 55-88.

Reuss, A.E. 1867. Uber einige Bryozoen aus dem deutschen

Unteroligozan. Sitzungsberichte der Akademie der Wissenschaften

in Wien (Abt. 1 ) 55: 216-234.

Ryland, J.S. 1982. Bryozoa. Pp. 743-769 in: Parker, S.P. (ed.),

Synopsis and classification of living organisms. McGraw-Hill: New

York.

Silen, L. 1947. Conescharellinidae (Bryozoa Gymnolaemata) collected

by Prof. Dr Sixten Bock’s expedition to Japan and the Bonin Islands

1914 . Arkiv for Zoologi 39A: 1—59.

Stoliczka, F. 1862. Oligocane Bryozoen van Latdorf in Bernburg.

Sitzungberichte der Kaiserlichen Akademie Wissenschaften Wien

45, Abt. 1: 71-94.

Taylor, P.D., and Gordon, D.P. 2002. Alcide d’Orbigny’s work on

Recent and fossil bryozoans. Comptes Rendus Palevol 1 (7):

533-547.

Tenison Woods, J.E. 1880. On some Recent and fossil species of

Australian Selenariidae (Polyzoa). Transactions of the Royal

Society of South Australia 3: 1-12.

Waters, A.W. 1881. On fossil chilostomatous Bryozoa from south-west

Victoria, Australia. Quarterly journal of the geological society

(London) 37: 309-347.

Waters, A.W. 1882a. On fossil Chilostomatous Bryozoa from Mount

Gambier, South Australia. Quarterly Journal of the Geological

Society ( London ) 38: 257-276.

Waters, A.W. 1882b. On fossil chilostomatous Bryozoa from

Baimsdale, (Gippsland). Quarterly Journal of the Geological

Society (London) 38: 502-513.

Waters, A.W. 1887. Bryozoa from New South Wales, North Australia,

etc. Part II. Annals and Magazine of Natural History (5) 20:

181-203.

Waters, A.W. 1889. Bryozoa from New South Wales, North Australia,

etc. Part IV. Annals and Magazine of Natural History (6) 4: 1-24.

Waters, A.W. 1904. Bryozoa. Resultats du Voyage du S.V. “ Belgica ”,

Zoologie. Expedition Antarctique Beige 4: 1-114.

Waters, A.W. 1905. Notes on some Recent Bryozoa in d’Orbigny’s

collection. Annals and Magazine of Natural History (7) 15: 1-16.

Australian Conescharellinidae (Bryozoa)

181

Waters, A.W. 1919. On Batopora and its allies. Annals and Magazine

of Natural History (9) 3: 79-94.

Waters, A.W. 1921. Observations on the relationships of the (Bryozoa)

Selenariidae, Conescharellinidae, etc., fossil and Recent. Journal of

the Linnean Society (Zoology) London 34: 399—427.

Whitelegge, T. 1887. Notes on some Australian Polyzoa. Proceedings

of the Linnean Society of New South Wales 2: 337-347.

Whitelegge, T. 1888. Notes on some Australian Polyzoa. Annals and

Magazine of Natural History (6) 1: 13-22.

Winston, J.E., and Hakansson, E. 1986. The interstitial fauna of the

Capron Shoals, Florida. American Museum Novitates 2865: 1-98.

Zagorsek, K. 2001. Upper Eocene Bryozoa from the Alpine Foreland

Basin in Salzburg, Austria (Borehole Helmberg-1). Osterreichische

Akademie der Wissenschaften, Schriftenreihe der Erdwissenschaft-

lichen Kommissionen Band 14: 509-609.

Zagorsek, K., and Kazmer, M. 2001. Eocene Bryozoa from Hungary.

Courier For schungsinstitut Senckenb erg 231: 1-159.

Appendix. Station numbers, details of localities, latitude, longitude, depth, with distribution of species

Maplestone specimens

South Australia, C. biarmata, C. magniarmata, C. cognata, C. diffusa

Kangaroo I., South Australia, C. cognata.

Locality assumed to be NSW and includes part of type material described by Maplestone (1909). “C. angulopora ” sensu

Maplestone, C. biarmata, C. magniarmata, C. cognata, C. diffusa, C. pustulosa, Trochosodon ampulla, Zeuglopora lanceolata

BMNH 1976.1.6.2, Cape York, Queensland, Challenger stn 185, 279 m, Trochosodon fecundus, T. anomalus, T. praecox

Museum Victoria Bass Strait Survey

BSS-055, 39°9'S, 143°26', 85 m, C. cognata.

BSS-065, 39°5'S, 142°33' 207 m, C. cognata.

BSS-117, 40°38'S, 145°23’E 36 m, C. pustulosa.

BSS-130, 39°38'S, 145°5.01'E 66 m, C. cognata.

BSS-155, 38°24'S, 144°54.03'E, 70 m, C. cognata.

BSS-158, 38°34'S, 144°54.03'E, 82 m, C. cognata, C. pustulosa.

BSS-159, 39°46'S, 146°18’E, 80 m, C. cognata.

BSS-161, 39°47'S, 147°19.3'E, 60 m, C. cognata.

BSS-162, 40°9.4'S, 147°32'E, 51 m, C. cognata.

BSS-167, 39°44.8'S, 148°40.6'E, 124 m, C. magniarmata; C. plana.

BSS-169, 39°2.4'S, 148°30.6'E, 120 m, C. multiarmata; C. plana, C. pustulosa.

BSS-170, 38°52.6'S, 148°25.2'E, 140 m, C. biarmata, C. multiarmata, C. magniarmata, C. cognata.

BSS-171, 38°53.7'S, 147°55.2'E, 71 m, C. magniarmata, C. cognata, C. diffusa.

BSS-176, 38°54.3'S, 147°13.4'E, 58 m, C. cognata.

Museum Victoria eastern Australian continental slope, RV Franklin , 1986

SLOPE-2, off Nowra, NSW, 34°57.90’S, 151°8’E, 503 m, C. eburnea, C. multiarmata, C. ecstasis, C. plana, C. pustulosa

SLOPE-6, off Nowra ,NSW, 34°51.90'S, 151°12.60E, 770 m, C. ecstasis, C. plana, T. asymmetricus, T, diommatus

SLOPE-7, off Nowra, NSW, 34°52.29'S, 151°15.02'E, 1096 m, C. multiarmata, C. ecstasis, C. plana, T. asymmetricus, T. diom-

matus, T. anomalus

SLOPE-19, off Eden, NSW, 37°07.3'S, 15°20.2'E, 520 m, C. biarmata

SLOPE-27, S of Point Hicks, Vic., 38°25'S, 149°E, 1500 m, Crucescharellina australis

SLOPE-39, S of Point Hicks, Vic., 38°19.1'S, 149°14.3'E, 600 m, C. multiarmata, C. ecstasis, C. pustulosa

SLOPE-40, S of Point Hicks, Vic., 38°17.7'S, 149°11.3'E, 400 m, C. multiarmata, C. ecstasis, C. plana, C. pustulosa, Flabellopora

umbonata

SLOPE-45, off Freycinet Peninsula, Tas., 42°02.2'S, 148°38.7E, 800 m C. ecstasis, C. pustulosa, T. diommatus

SLOPE-48, off Freycinet Peninsula, Tas., 41°57 5'S, 148°37.9'E, 400 m, C. multiarmata, C. ecstasis

SLOPE-49, off Freycinet Peninsula, Tas., 41°56.5'S, 148°37.9'E, 200 m, C. diffusa

SLOPE-53, 54 km ESE, of Nowra, NSW, from 34°52.77S, 151°15.04'E, 996 m to 34°54.03' 151°19.05E, 990 m, C. ecstasis.

SLOPE-56, 44 km E, of Nowra, NSW, from 34°55.79'S, 151°08.06'E, 429 m to 34°56.06'S, 151°07.86'E, 466 m, C. plana.

Great Australian Bight, Y. Bone collection, RV Franklin , 1995

GAB-015, 33°20'S, 130°00E, 203 m, C. magniarmata

GAB-019, 33°22'S, 129°19E, 301 m, C. cognata, C. stellata

GAB-020, 33°20'S, 129°18'E, 157 m, C. magniarmata, C. cognata, C. diffusa, C. plana, Bipora flabellaris, Flabellopora

umbonata

GAB-030, 33°13'S, 128°29'E, 137 m, C. multiarmata, C. cognata, C. plana, Bipora flabellaris, Flabellopora umbonata

GAB-044, 33°25'S, 125°58E, 163 m, C. plana

182

P. E. Bock and P. L. Cook

Appendix. Continued

GAB-045, 33°25'S, 125°58'E, 143.5 m, C. cognata, Flabellopora umbonata

GAB-048, 33°53'S, 125°22'E, 182 m, C. obscura

GAB-049, 33°53'S, 125°22'E, 156 m, C. cognata, C. plana, C. pustulosa

GAB-056, 33°19'S, 125°43'E, 72.5 m, C. magniarmata, Flabellopora umbonata

GAB-067, 33°22'S, 124°23'E, 50 m, C. cognata, C. diffusa, Flabellopora umbonata

GAB-069, 33°43'S, 124°23'E, 65.5 m, C. diffusa

GAB-074, 34°15'S, 124°24'E, 117-125 m, C. obscura, Flabellopora umbonata

GAB-084, 34°20'S, 124°08'E, 96 m, Flabellopora umbonata

GAB-088, 34°35’S, 123°38’E, 98 m, Flabellopora umbonata

GAB-093, 34°32’S, 122°58'E, 95 m, Flabellopora umbonata

GAB-098, 34°39'S, 122°26'E, 156 m, C. cognata

GAB-101, 34°33S, 121°33E, 236 m, C. cognata

GAB-108, 34°29’S, 121°32'E, 101 m, C. obscura

GAB-112, 34°20’S, 119°55'E, 65 m, Flabellopora umbonata

GAB-113, 34°36'S, 119°55'E, 106 m, C. obscura

GAB-116, 34°37'S, 119°21'E, 66 m, Bipora flabellaris

GAB-117, 34°35'S, 119°00'E, 65.5 m, Flabellopora umbonata

GAB-118, 34°59'S, 119°00'E, 87 m, C. diffusa, C. obscura, Bipora flabellaris

GAB-119, 35°00'S, 119°00’E, 149 m, Flabellopora umbonata

GAB- 128, 35°07'S, 116°52'E, 59 m, C. stellata, Flabellopora umbonata

GAB- 129, 35°07'S, 116°20'E, 70 m, C. diffusa

GAB-131, 35°07'S, 115°51'E, 160 m, C. obscura

Dampier Archipelago, north-western WA, G.C. B. Poore collection, 1999

DA-2-09-02, 20°20.5'S, 117°05.4'E, 33 m, off Delambre I., C. obscura

DA-2-75-02, 20°32.17'S, 116°33.63'E, 20.5 m, off Goodwyn I., T.fecundus

DA-2-73-01, 20°40.0'S, 116°27.7'E, 12.5 m, off Eaglehawk I., C. obscura

DA-2-37-01, 20°36.5'S, 116°35.0'E, 15 m, off Enderby I., C. diffusa

Other Museum Victoria collections

Off Tasmania?, RV Dmitri Mendeelev, C. diffusa

S03/84/74, off eastern Tas., RV Soela, 42°41'S, 148°25.0'E, 320 m, Flabellopora umbonata, Crucescharellina australis

Fossil localities from Victoria and South Australia

Bairnsdale (Skinner’s): Mitchell River bank, about 12 km W of Bairnsdale, Vic., 37°47.9'S, 147°29.5'E. C. macgillivrayi, C. off

diffusa

Balcombe Bay: also known as Fossil Beach, Mornington, Mount Martha and possibly “Schnapper Point” (MacGillivray); on coast

of Port Phillip Bay, about 3 km S of Mornington, Vic., 38°14.5'S, 145°01.7'E. Fyansford Clay. Age: Balcombian; Middle

Miocene, (Langhian). C. ocellata, C. macgillivrayi, C. humerus

Batesford Quarry: upper levels of Batesford Limestone Quarry, 7 km W of Geelong, Vic., 38°06.5'S, 144°17.3'E. Fyansford Clay.

Age: Balcombian; Middle Miocene, (Langhian). C. ocellata, C. macgillivrayi, C. humerus

Heywood No. 10 Bore, Mines Department of Victoria, 38°07.9'S, 141°37.6'E. Age: Miocene. C. macgillivrayi.

Mount Schanck: limestone quarry about 1 km W of Mount Schanck, about 15 km S of Mount Gambier, SA, 37°57'S, 140°43.2'E.

Gambier Limestone. Age: Early Miocene, (Longfordian). C. macgillivrayi, C. humerus

Muddy Creek: Clifton Bank, Muddy Creek, 8 km W of Hamilton, Vic., 37°44.6'S, 141°56.4'E. Muddy Creek Marl (= Gellibrand

Marl). Age: Balcombian. C. macgillivrayi, C. humerus, C. off. diffusa

Paaratte No.l Bore. Mines Department bore in the Parish of Paaratte, located in the village of Port Campbell, Vic., 38°36.8'S,

143°00.0'E. Age: Middle Miocene. C. humerus

Puebla: coastal section, about 3 km W of Torquay, Vic., 38°21.4'S, 144°17.8'E. Jan Juc Formation. Age: Longfordian; Early

Miocene, (Aquitanian). C. macgillivrayi, C. humerus

Memoirs of Museum Victoria 61(2): 183-208 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia

Erich M. G. Fitzgerald

School of Geosciences, P.O. Box 28E, Monash University, Clayton, Victoria 3800, Australia

(erichmgf@mail.earth.monash.edu.au) and Geosciences, Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001,

Australia (efitzger @ museum, vie .go v. au)

Abstract Fitzgerald, E.M.G. 2004. A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia. Memoirs of Museum

Victoria 61(2): 183-208.

The stratigraphy, age, lithology, and vertebrate fauna of all 56 pre-Pleistocene fossil Cetacea-bearing localities in

Australia are reviewed. The majority of these localities occur in the state of Victoria, and are Miocene in age. The most

complete cetacean fossils have been recovered from coastal exposures of the Upper Oligocene Jan Juc Formation, south-

west of Torquay in the Torquay Basin (Victoria). The inadequately known fossil record of cetaceans in Australia is due

to a lack of research, and not a lack of potentially fossiliferous rock outcrop.

Keywords Cetacea, Archaeoceti, Mysticeti, Odontoceti, Australia, localities, fossil record, Tertiary, Oligocene, Miocene, Pliocene

Introduction

Australia has an incompletely known fossil record of cetaceans

(whales, dolphins, and porpoises; Order Cetacea). The oldest

fossil cetaceans from Australia are Early Oligocene, with the

fossil record being best known from the Late Miocene-Early

Pliocene (Fig.l for correlation of cetacean-bearing strati-

graphic units). The majority of Australian fossil cetaceans

described (but not necessarily published) have been derived

from only a few locations. These fossil sites occur within the

Paleogene-Neogene marine sedimentary basins along the

southern margin of Victoria. Other described fossil cetaceans

have been collected from South Australia and Tasmania,

although fossil cetaceans from these states are less well

represented in museums.

Mahoney and Ride (1975) provided a synopsis of fossil

cetacean genera and species described from Australia with

information on the history of collection and study. They did not

include all cetaceans in institutional palaeontology collections.

Fordyce (1982a, 1982b) published important taxonomic

reviews of Australian fossil cetaceans based on museum col-

lections. Bearlin (1987, 1988) continued studies on Australian

fossil Cetacea, with analyses of Neogene mysticete specimens

and their provenance. In all of these studies, research was con-

ducted primarily on specimens in Museum Victoria, Melbourne

(NMV), and to a lesser extent on fossils in the South Australian

Museum and the School of Earth Sciences, University of

Melbourne. The fossil cetaceans in the palaeontology collec-

tions of the Tasmanian Museum, Hobart and Queen Victoria

Museum, Launceston remain largely u nk nown.

The Australian fossil record of cetaceans is so poorly known

because little systematic prospecting has been carried out. All

significant fossil cetaceans have been discovered by accident,

often by amateur palaeontologists or members of the public.

Initial steps to advance research are: 1, to identify where fossil

cetaceans have previously been discovered in Australia; 2,

determine the faunal compositions of these localities based on

fossils in museum and other collections; 3, conduct a census of

the sedimentary geology of these localities; and 4, determine

the geological age of the fossil-bearing localities.

This review aims to document localities in Australia from

which pre-Pleistocene fossil cetaceans have been recovered.

Because this review is principally based on museum and

university collections, it is unlikely that all localities have been

recognised. Several large private collections exist that include

significant material. If accurate locality and stratigraphic data

occur with these specimens, it is likely that the real number

of fossil cetacean localities is much larger. This review is

intended to serve as a companion to Rich and co-workers ’

(1991) review of Tertiary terrestrial mammal localities.

Overview of localities and the fossil record

The fossiliferous localities in Australia that have yielded

cetaceans occur within the offshore and onshore marine, and in

one case freshwater, sedimentary basins in south-eastern and

southern central Australia. These areas of Paleogene-Neogene

sedimentation correspond to the: Otway, Torquay, Port Phillip,

and Gippsland basins in Victoria; St Vincent, Murray, Gambier

and Lake Eyre basins in South Australia; and the Bass Basin in

184

E. M. G. Fitzgerald

Figure 1. Correlation chart of Oligocene-Pliocene Australian fossil cetacean-bearing stratigraphic units. Abbreviations: Fmn=Formation;

Lst=Limestone; Mbr=Member; Sst=Sandstone. Broken lines indicate uncertain maximum and mi nimum age ranges of stratigraphic units. Ages,

chronostratigraphy, and planktonic foram zones modified from Alley et al. (1995) and Holdgate and Gallagher (2003).

Tasmania (Figs 2, 3). The most abundant and diverse fossil

records of cetaceans occur in the Otway, Torquay, and Port

Phillip basins of Victoria. However, the most complete and

best-preserved fossil cetacean material is derived from the

Torquay Basin in central coastal Victoria.

The most northern localities are those in the Lake Frome

area in north-eastern South Australia (31°S), whereas the most

southern localities occur offshore southern Tasmania (45 °S).

The stratigraphic record of cetaceans is most complete for the

Miocene, with several localities in South Australia, Victoria,

and Tasmania representing the Early, Middle, and Late

Miocene (see below). However, most Miocene sites are either

Middle or Late Miocene in age. The Late Oligocene record is

becoming better known, with a higher diversity of cetaceans

than previously recognised (Fitzgerald, 2003, 2004). It is

important to note that most of the Late Oligocene fossil

cetaceans can probably be referred to Mammalodontidae, and a

new undescribed group of archaic toothed mysticetes. No fos-

sil cetaceans have been recorded from the Eocene of Australia

(Fordyce, 1982a, 1991). Other poorly documented stratigraph-

ic intervals are the Early Oligocene, early Early Miocene and

Late Pliocene.

In Australia, the fossil record of mysticetes is much better

known than that for odontocetes (Bearlin, 1987; Fitzgerald,

2004). The Australian Late Oligocene-Late Pliocene record of

Mysticeti is particularly informative because it documents

stem-group toothed mysticetes (Berta et al., 2003), the archaic

baleen-bearing “cetotheres” in the Miocene, early records of

Balaenopteridae in the Mid-Late Miocene (Bearlin, 1988), and

the transition to a relatively modern mysticete fauna composed

of balaenids and balaenopterids in the Late Miocene-early

Pliocene (Bearlin, 1987, 1988; Fordyce, 1991).

The Australian fossil record of Odontoceti is known only in

broad outlines. This reflects the paucity of complete skulls, pre-

serving diagnostic features, in museum and university collec-

tions. There are only two records of fairly complete and diag-

nostic fossil odontocete skulls in Australia: an undescribed

squalodontid from Victoria (Bearlin, 1982), and Prosqualodon

davidis (Flynn, 1923, 1948) from Tasmania. The skull of the

latter specimen is now lost (Mahoney and Ride, 1975) and the

former skull lacks the rostrum and most of the dorsal surface of

the cranium. However, the remainder of the holotype specimen

of P. davidis, consisting of much of the postcranial skeleton and

mandibles, is still extant in the collection of the Department of

Geology, University of Tasmania. Most odontocete fossils are

represented by isolated mandibles, teeth, periotics and tympan-

ic bullae. An exception is an incompletely prepared, partially

articulated, postcranial skeleton from the Late Oligocene of

Victoria (NMV P48861). Although lacking cranial elements

(apart from several teeth), the morphology of the scapula indi-

cates that this specimen may represent a primitive eurhinodel-

phinid odontocete.

Tertiary fossil Cetacea localities in Australia

185

Figure 2. Tertiary sedimentary basins with fossil Cetacea localities in

Victoria and Tasmania. Modified from Abele et al. (1988) and

Holdgate and Gallagher (2003).

Taxonomy

Systematic biologists have yet to reach a consensus on cetacean

systematics and taxonomy. This is largely due to the publica-

tion of numerous divergent phylogenetic hypotheses, utilising

both molecular and morphological data sets, over the last two

decades (e.g., Amason and Gullberg, 1994; Geisler and Luo,

1996; Heyning, 1989, 1997; Messenger and McGuire, 1998;

Milinkovitch et al., 1993; Zhou, 1982). Higher classification

schemes were reassessed by Geisler and Sanders (2003) but the

more conventional classification of Fordyce and Muizon

(2001) is followed herein. The classification of non-cetacean

mammals follows McKenna and Bell (1997). For non-

mammalian tetrapods, osteichthyans, and most chondrich-

thyans, the classification of Carroll (1988) is used. Some mod-

ifications to chondrichthyan taxonomy presented in Carroll

(1988) and Kemp (1991) have been made following a recent

review by Purdy and colleagues (2001). Fossil species of great-

toothed sharks are referred to the genus Carcharodon, and not

Carcharocles, following Gottfried and Fordyce (2001).

Faunal lists

Few Australian workers have studied Cainozoic marine verte-

brate faunas in detail, with notable exceptions being Bearlin

(1987), Chapman and Cudmore (1924), Chapman and

Pritchard (1904, 1907), Fordyce (1984, 1991), Kemp (1970,

1982, 1991), and Pledge (1985). Kemp (1991) outlined the var-

ious biases affecting the taxonomic compositions of chondrich-

thyan faunas in Australia, and his conclusions may be applied

to cetaceans. Most of the substantial Tertiary marine vertebrate

holdings of Museum Victoria were obtained prior to 1980.

Indeed, most of the collection of Cainozoic marine vertebrate

fossils dates from the first half of the twentieth century. It is,

therefore, inevitable that identifications and alpha taxonomic

Figure 3. Tertiary sedimentary basins with fossil Cetacea localities in

South Australia. Modified from Alley et al. (1995).

studies were influenced by the taxonomic references of the time

and contemporary philosophical approaches to systematics.

The faunal lists presented herein reflect these biases. However,

given the paucity of new material (apart from cetaceans) added

to museum collections over the last 30 years, it is unlikely that

the taxonomy requires major reassessment. Of more singular

importance is the incomplete nature of the faunal lists, due to

the lack of identification of many specimens. It is highly

probable that the faunal lists represent artificially impoverished

faunas, particularly of marine tetrapods and osteichthyan fish.

For cetaceans and other marine tetrapods, faunal lists have

been compiled from my personal observations and consultation

of published (Bearlin, 1988; Fordyce, 1982a, 1984, 1991;

Pledge, 1985) and unpublished (Bearlin, 1987) studies.

Terrestrial and freshwater vertebrate faunas were adopted from

Rich and others (1991), and unpublished data provided by

K. Piper. Chondrichthyan and osteichthyan faunas were adapt-

ed from information in Kemp (1978, 1991), Pledge (1985) and

Stinton (1958, 1963) as well as my own observations. Although

Pledge (1967) reviewed South Australian Tertiary chon-

drichthyans, his published localities for chondrichthyans can-

not be correlated with South Australian cetacean localities.

Many of the identifications of marine tetrapods, and their taxo-

nomic affinities, are preliminary.

Information on the Australian Cainozoic fossil record of

marine birds may be found in: Finlayson (1938), Glaessner

(1955), Jenkins (1974, 1990), Simpson (1957, 1959, 1965,

1970), Rich (1975), Vickers-Rich (1991), Vickers-Rich and

Rich (1993) and Wilkinson (1969). For more detailed informa-

186

E. M. G. Fitzgerald

tion on Cainozoic fossil marine fish see Chapman (1913,

1917a, 1930), Chapman and Cudmore (1924), Chapman and

Pritchard (1904, 1907) and Kemp (1991).

Localities

For the purposes of this review, a locality is defined as a limit-

ed area where fossils have been derived from one particular

stratigraphic unit. However, there is some variation in the use

of the term “locality” when applied to sites that have yielded

fossil cetaceans. For example, cetacean fossils have been

recovered from an area around Bird Rock, Torquay, which

encompasses outcrop in the shore platform, and in the cliff sec-

tions, at varying heights above the base of the cliff.

Furthermore, fossils may have been derived from an area in the

immediate vicinity of the Bird Rock stack, or up to 150 m

south-west of Bird Rock. This contrasts with localities such as

Clifton Bank, along Muddy Creek, where there is only a small

area of outcrop within a limited area. In most cases, field col-

lection data for specimens is not detailed enough to enable pre-

cise limits to be placed on the area of a locality. Hence, these

localities have a relatively broad definition.

Stratigraphy

Unless otherwise indicated, stratigraphic nomenclature follows

that of Abele and colleagues (1988) and Holdgate and

Gallagher (2003) for Victoria, and Alley and co-workers (1995)

for South Australia. Determination of the stratigraphic position

of most localities was based on work in Abele and colleagues

(1988, and references therein), data presented by Alley and oth-

ers (1995), and Lukasik and James (1998).

Australian fossil Cetacea localities

VICTORIA (Fig. 4)

1. The Otway Basin (Fig.5)

1.1 Dutton Way, Portland

Geographic location. This site consists of several disjunct

points along south-south-east facing sea cliffs and beach on the

north-west side of Portland Bay, north of Portland (38°19'S,

141°38'E). Fragmentary material has also been collected from

reefs and the seafloor in Portland Bay, in water depths of up to

100 m.

Stratigraphic position. The fossiliferous horizon that has

yielded fossil cetaceans has been identified as a phosphatic

nodule bed at the base of the Whaler’s Bluff Formation.

Age. Early Pliocene. The onshore Whaler’s Bluff Formation

has been dated to zones N18-N19 (Mallett, 1977). Dating of the

offshore deposits of the Whaler’s Bluff Formation indicate that

sedimentation continued into the mid-Pliocene (zones N19-

N21) (Holdgate and Gallagher, 2003). It is likely that most of

the fossil cetacean material was derived from the nodule

horizon and therefore has a maximum age of earliest Pliocene.

Lithology. The main lithologies present include a basal phos-

phatic nodule bed, fossiliferous clays, oyster beds, and sandy

limestones (Holdgate and Gallagher, 2003).

Figure 4. Areas with fossil cetacean-bearing localities in Victoria and

Tasmania. Solid circles represent fossil Cetacea localities. Modified

from Abele et al. (1988) and Holdgate and Gallagher (2003).

Figure 5. Fossil Cetacea localities in the Otway Basin, western

Victoria. Solid circles represent fossil Cetacea localities.

Material. All worn, isolated elements: cranial fragments, peri-

otics, tympanies, teeth, vertebrae, rib fragments.

Fauna. Carcharodon megalodon, Carcharodon carcharias,

Isurus hastalis, Isurus sp., Palorchestidae, Vombatidae,

Macropodidae, Ektopodontidae, ?Phocidae, Balaenidae,

Balaenoptera sp., Megaptera sp., Physeteridae, Ziphiidae,

Delphinidae: 2 species.

1.2 Arch Site, Grange Burn (Fig.6 for localities 1.2-1. 4)

Geographic location. Resting on a quartz porphyry bar at the

base of a low cliff on the southern bank of Grange Burn, oppo-

site a natural arch, immediately north of “The Caves” property,

8 km west of Hamilton, western Victoria (near 37°43'30"S,

141°56'0"E) (Bearlin, 1987).

Stratigraphic position. Bochara Limestone Member, Port

Campbell Limestone, unconformably overlying Devonian

Rocklands Rhyolite.

Tertiary fossil Cetacea localities in Australia

187

Figure 6. Fossil Cetacea localities in the Grange Bum area, west of

Hamilton in western Victoria. Solid circles represent fossil Cetacea

localities.

Age. Early Middle Miocene. The foraminifera Lepidocyclina

howchini in the matrix suggests a zone N8-N9 (Batesfordian)

age for the cetacean fossils from this locality (Bearlin, 1987).

Lithology. Yellow-brown bryozoal calcarenite.

Material. One partial skeleton.

Fauna. Carcharodon megalodon, Isurus hastalis, Pelocetus sp.

1.3 Clifton Bank

Geographic location. Low cliffs above the riverbank on the

western and eastern sides of Muddy Creek (south of the junc-

tion of Grange Burn and Muddy Creek), and due west of the

“Clifton” property, located 8 km west of Hamilton (near

37°43'30"S, 141°56'0"E).

Stratigraphic position. Muddy Creek Marl Member, Port

Campbell Limestone. In more complete sections along Grange

Burn, e.g. at Pat’s Gully (see Gill, 1957), the Muddy Creek

Marl is conformably underlain by the Bochara Limestone (Gill,

1957), and disconformably overlain by the Grange Burn

Formation.

Age. Middle Middle Miocene to early Late Miocene. Sr/Sr

dating of shells from 1 m below the base of the Grange Burn

Formation, within the Muddy Creek Marl, yielded an age of

10.8 Ma (Dickinson et al., 2002). The presence of the

foraminiferan Orbulina suturalis indicates a zone N10 age

(Middle Miocene, Serravallian, 12-14 Ma) (Abele et al., 1988).

However, the Muddy Creek Marl can range as high as zone

N16 (up to ~8 Ma) (Singleton et al., 1976). Acacia pollen have

been recorded from the base of the Muddy Creek Marl, at

Clifton Bank, which indicate that this member lies above the

top of the Cyatheacidites annulatus Zone (Harris, 1971); this

suggests that the Muddy Creek Marl Member is younger than

15 Ma. Gill (1957) suggested a Middle or Lower Miocene age

(Balcombian) for the Muddy Creek Marl.

Lithology. Richly fossiliferous grey silty marl.

Material. Fossil cetacean elements are generally very well pre-

served, although no articulated skulls and skeletons have yet

been recovered from this unit. Elements preserved include

vertebrae, ribs, and periotics.

Fauna. Carcharias sp., Carcharias taurus, Odontaspis acutis-

sima, Isurus hastalis, Isurus retroflexus, Isurus desori, Isurus

planus, Isurus cf. paucus, Isurus sp., Carcharodon megalodon,

Carcharodon sp., Galeocerdo aduncus, Notorynchus primi-

genius, Dasyatidae, Heterenchelys regularis, Muraenesox

obrutus, Hypomesus glaber, Merluccius fimbriatus,

Trachichthodes salebrosus, Sebastodes fissico status, Sillago

pliocaenica, Diodon formosus, Balaenidae, Scaldicetus lodgei,

cf. Physeter sp., Odontoceti indet.

1.4 Forsyth’s Bank to Fossil Rock Stack

Geographic location. River bank and river bed exposures along

and in Grange Burn, south-east of “The Caves” homestead, 8

km west of Hamilton (37°43'42±03"S, 14r56'40±04"E.

Stratigraphic position. Grange Burn Formation, uncon-

formably overlying the Muddy Creek Marl. The base of the

Grange Burn Formation is marked by a phosphatic nodule bed

horizon (Gill, 1957; Dickinson et al., 2002; Holdgate and

Gallagher, 2003). In most sections where the top of the Grange

Burn Formation is exposed, a basalt layer disconformably over-

lies the marine sediments (Gill, 1957). Further to the south-east

along the Grange Bum, a terrestrial palaeosol facies is present

between the Grange Burn Formation and the basalt.

Age. The Grange Bum Formation has generally been con-

sidered as Kalimnan (Early Pliocene) in age, due to the com-

position of the rich invertebrate macrofauna (Gill, 1957).

Foraminifera dates indicate a zone N17 age (Mallett, 1977).

The basalt above the Grange Burn Formation has been dated to

4.35 Ma using K-Ar (Turnbull et al., 1965). More recently,

Sr/Sr dates from the base of the Grange Burn Formation indi-

cated a 4.0-5 .0 Ma maximum age for the formation (Dickinson

et al., 2002). These data indicate an earliest Early Pliocene age

for the Grange Burn Formation.

Lithology. Shelly marl and sandy to pebbly limestone

(Holdgate and Gallagher, 2003).

Material. As is typical of most Mio-Pliocene nodule bed fossil

vertebrate material in Victoria, fossils are often rolled, polished,

and broken. Almost all specimens represent isolated elements,

with associated material being very rare. Typically preserved

elements include partial rostra, cranial fragments, isolated

periotics and tympanic bullae, teeth, incomplete mandibles,

vertebrae, and ribs.

188

E. M. G. Fitzgerald

Fauna. Heterodontus cainozoicus, Carcharodon megalodon,

Carcharodon carcharias, Isurus escheri , Isurus hastalis,

Myliobatis sp., Edaphodon sweeti, Ischyodus cf. dolloi,

Kurrabi sp., Phocidae, “Cetotheriidae”, Balaenoptera sp., cf.

Scaldicetus sp., cf. Physeter sp., cf. Mesoplodon sp.,

?Delphinidae.

1.5 Spring Creek

Geographic location. Bed of Spring Creek, near Minhamite, 40

km south-east of Hamilton (near 37°59'S, 142°20'E).

Stratigraphic position. Unnamed unit (Abele et al., 1988). This

unit is probably laterally equivalent to the Grange Burn

Formation. Gill (1957) mentioned a Kalimnan-aged location

“from Goodwood station near Minhamite Railway Station 25

miles (40.2 km) SE of Ha mi lton” (p. 152).

Age. Presumed Late Miocene-Early Pliocene. T.A. Darragh

(pers. comm, to G.G. Simpson, cited in Simpson (1970))

has suggested a Cheltenhamian (Upper Miocene) or

older age for the Spring Creek beds near Minhamite.

Simpson (1970) suggested that the Spring Creek locality

was equivalent in age to the Black Rock Sandstone at

Beaumaris.

Lithology. Fossiliferous green-grey marly fine sand

approximately 1 m thick (Abele et al., 1988).

Material. Worn and polished isolated elements: periotics and

indeterminate bone fragments.

Fauna. Pseudaptenodytes macraei, Balaenoptera sp.

1.6 Kawarren

Geographic location. Old “Alkemade’s Quarry”, slightly north

of Kawarren railway station, on the steep north bank of Loves

Creek, Kawarren, about 19 km south of Colac (near 38°29'S,

143°35'E).

Stratigraphic position. Clifton Formation (Abele et al., 1988;

McHaffie and Inan, 1988).

Age. In the north-east margins of the Port Campbell

Embayment, the Clifton Formation is Late Oligocene (Abele et

al., 1988).

Lithology. The Clifton Formation is generally a medium-

coarse-grained calcarenite, with about 10% quartz and limonite

sand (Tickell et al., 1992). At the Kawarren Quarry, this unit

consists of friable, pale yellow limestone interbedded with

harder crystalline bands (McHaffie and Inan, 1988). All of the

vertebrate fossils have been recovered from the more friable

calcarenite layers.

Material. Generally well-preserved isolated elements, exhibit-

ing a low degree of rolling. Elements preserved include teeth,

ribs and bone fragments.

Fauna. Carcharias sp., Isurus sp., Isurus desori, Carcharoides

sp., Carcharoides totuserratus, Carcharodon angustidens,

?Mysticeti new family.

1.7 Leigh River

Geographic location. Outcrop on the eastern bank of the Leigh

River, about 5 km north of Shelford, 46 km north-east of

Geelong (near 38°00'S, 143°58'E).

Stratigraphic position. Middle section of the Gellibrand Marl

(Abele et al., 1988; Dickinson et al., 2002).

Age. Probably Early Miocene. In the north-east part of the Port

Campbell Embayment, the Gellibrand Marl ranges in age from

Late Oligocene to Middle Miocene (Abele et al., 1988).

Outcrop along the Leigh River is Early Miocene in age, as the

Late Oligocene-aged base of the formation is not exposed,

and the youngest Middle Miocene (Bairnsdalian) section

has probably been eroded, or not deposited, due to the Leigh

River locality being at the embayment margin (Abele et al.,

1988).

Lithology. Marl, calcareous silt, clay, and sand with minor cal-

carenite layers.

Material. One well-preserved isolated tooth.

Fauna. Isurus sp., Cetacea indet.

1.8 Hopkins River

Geographic location. North end of outcrop in a quarry on the

west bank of the Hopkins River, 150 m south of the Princes

Highway bridge over the Hopkins River, near Allansford, about

10 km east of Warrnambool (near 38°23'S, 142°35'E).

Stratigraphic position. Port Campbell Limestone sensu stricto

(Abele et al., 1988; Tickell et al., 1992).

Age. Middle-Late Miocene, zones N11-N17 (Holdgate and

Gallagher, 2003).

Lithology. Yellow fine-grained calcarenite.

Material. A fairly well-preserved posterior part of mandible

with two large conical teeth in place.

Fauna, cf. Scaldicetus sp.

1.9 Gibson’s Steps

Geographic location. Cliff at Gibson’s Steps, approximately 12

km east of Port Campbell, western Victoria (near 38°40'S,

143°07'E).

Stratigraphic position. Port Campbell Limestone (Bearlin,

1987; Abele et al., 1988; Tickell et al., 1992).

Age. Bairnsdalian, Middle Miocene, zone N10 (Bearlin, 1987).

Lithology. Yellow-grey fine-grained calcarenite.

Material. One partial skeleton

Fauna. “Cetotheriidae”.

1.10 Curdie

Geographic location. “Kurdeez” Quarry (Victorian

Agricultural Lime Ltd), 5 km north-north-west of Curdie, near

Timboon (approximately 38°27'S, 142°56'E).

Tertiary fossil Cetacea localities in Australia

189

Stratigraphic position. Port Campbell Limestone.

Age. Balcombian, correlated with zone N10, Serravallian,

Middle Miocene (Bearlin, 1987).

Lithology. Calcarenite.

Material. One partial skeleton.

Fauna. ?Balaenopteridae.

1.11 Princetown Beach

Geographic location. Near Point Ronald, Princetown, 18 km

east of Port Campbell, western Victoria (38°42'S, 143°09'E).

Stratigraphic position. Nodule bed within the Clifton

Formation.

Age. Late Oligocene. Below the unnamed nodule horizon,

shells have given Sr/Sr dates of 27.4 Ma (Holdgate

and Gallagher, 2003). Immediately above the nodule bed

Sr/Sr dates average -24.0 Ma. Foraminifera from the

Clifton Formation indicate a zone P21b-P22 age (Late

Oligocene).

Lithology. Thin horizon of phosphate and limonite nodules

within limestone- sandy limestone matrix.

Material. One worn and polished incomplete rib.

Fauna. Odontaspis sp., Cetacea indet.

1.12 Castle Cove

Geographic location. One of two possible locations: (1) type

locality for the Calder River Limestone along the south-eastern

bank of the Calder River, north-west of Hordern Vale, or (2) on

the coast 1.7 km south-east of Castle Cove. Both sites are on

the western side of Cape Otway, and east of Point Reginald, in

the Aire district (near 38°47'S, 143°25'E). Etheridge (1878)

noted that the holotype tooth of Parasqualodon wilkinsoni was

found west of the Aire River.

Stratigraphic position. Calder River Limestone. Fordyce

(1982a) suggested that the holotype specimen of

Parasqualodon wilkinsoni was derived from the Calder

River Limestone, despite the fact that there is no direct

evidence that would indicate such a derivation. However,

there are two lines of evidence that indirectly suggest the

provenance of the holotype of Parasqualodon wilkinsoni'. (1)

the holotype tooth preserves distinctive features suggesting

affinities with Prosqualodon davidis (indeed, this isolated

tooth may be congeneric with P. davidis, or even con-

specific); the Prosqualodontidae is only known from Late

Oligocene-earliest Miocene deposits; (2) McCoy (1867a)

stated that the holotype tooth was collected from sandy beds at

Castle Cove on the Cape Otway coast; the only fossiliferous

marine sediments in this area, which are of suitable age to yield

a prosqualodontid tooth, are those of the Calder River

Limestone.

Age. Late Oligocene.

Lithology. Sandy bryozoal calcarenite with a thin discontinuous

basal layer of phosphatic nodules and quartz pebbles (Abele et

al., 1988).

Material. One tooth.

Fauna. Prosqualodon sp. (= Parasqualodon wilkinsoni ).

2. The Torquay Basin (Fig.7)

2.1 Split Point (Fig.8 for localities 2. 1-2.7)

Geographic location. Towards lighthouse, at Split Point, near

Aireys Inlet, 49 km south-west of Geelong (near 38°28'S,

144°06'E).

Stratigraphic position. Point Addis Limestone Member (Webb,

1995).

Age. Late Oligocene.

Lithology. Yellow sandy bryozoal calcarenite.

Material. One incomplete anterior caudal vertebra, and one

worn tooth.

Fauna. Carcharias macrotus, Carcharoides totuserratus,

Cetacea indet.

2.2 Point Addis

Geographic location. Cliffs on southern side of Point Addis,

south-west of Torquay, central coastal Victoria (38°23'S,

144°15'E).

Stratigraphic position. Point Addis Limestone. Most vertebrate

fossils have been collected from the base of the upper member

of the Point Addis Limestone (Nicolaides and Wallace, 1997;

Webb, 1995).

Age. Late Oligocene (Abele et al., 1988; Holdgate and

Gallagher, 2003).

Lithology. Ferruginous intraclastic conglomerate with abraded

shelly and vertebrate skeletal components (Nicolaides and

Wallace, 1997; Webb, 1995). This horizon directly overlies a

regionally extensive hardground (Nicolaides and Wallace,

1997; Webb, 1995).

Material. Isolated elements: teeth, tympanic bullae, rare post-

cranial remains. The material is usually broken and worn.

Fauna. Mammalodontidae.

2.3 Bells Headland

Geographic location. 300 m south-west of Bells Beach, south-

west of Torquay (near 38°22'S, 144°16'E).

Stratigraphic position. Lower beds of the Point Addis

Limestone (Abele et al., 1988).

Age. Late Oligocene.

Lithology. Sandy bryozoal calcarenite (Abele et al., 1988).

190

E. M. G. Fitzgerald

Figure 7. Fossil Cetacea localities in the Torquay and Port Phillip

basins, central coastal Victoria. Solid circles represent fossil Cetacea

localities. Modified from Holdgate and Gallagher (2003).

Material. Fossil preservation is generally very good. Cetacean

material consists of: one skull and associated ear bones;

associated vertebrae, and bone fragments.

Fauna. Mysticeti new family; genus and species 2.

2.4 Bells Beach

Geographic location. On shore platform, low tide mark, base of

low cliffs at north-east end of Bells Beach, south-west

of Torquay, central coastal Victoria (38°22'S, 144°17'E).

Stratigraphic position. Point Addis Limestone

Age. Late Oligocene

Lithology. Yellow bryozoal calcarenite (Abele et al., 1988).

Material. One partially articulated incomplete skeleton,

isolated periotics, and vertebrae.

Fauna. Carcharodon angustidens, Mammalodontidae genus

and species indet 1 .

2.5 Rocky Point

Geographic location. Low cliffs at Rocky Point, small head-

land at northern-most end of Bells Beach, about 250 m NE of

Bells Beach, south-west of Torquay (near 38°22'S, 144°17'E).

Stratigraphic position. ?Lower Jan Juc Marl.

Age. ?Late Oligocene.

Lithology. Yellow-orange silty and sandy marls.

Material. Indeterminate bone fragments.

Fauna. Cetacea indet.

2.6 Deadman’s Gully

Geographic location. Cliff exposures 600 m south-west

Fishermans Steps, near Deadman’s Gully, south-west

Torquay, central coastal Victoria (near 38°20'S, 144°18'E).

Figure 8. Late Oligocene fossil Cetacea localities on the coast south-

west of Torquay, in central coastal Victoria. Shaded areas represent

coastal outcrop of the Oligocene-Miocene Torquay Group. Solid

circles represent fossil Cetacea localities.

Stratigraphic position. Jan Juc Marl, based on the presence of

the molluscs Liratomina intertexta and Chione halli (which are

restricted to the Jan Juc Marl, T.A. Darragh, pers. comm.).

Age. Late Oligocene.

Lithology. Grey-yellow glauconitic and pyritic carbonate-

cemented calcarenite.

Material. One partially articulated incomplete skeleton: skull,

mandibles, teeth, tympanies, thyrohyal, atlas, axis, cervical ver-

tebra, ribs, scapulae, radius.

Faunal. Mysticeti new family, genus and species 1.

2.7 Bird Rock

Geographic location. Bluff adjacent to Bird Rock stack and

cliffs to the south-west, and shore platform exposures from

Bird Rock in the north-east, extending 250 m to the south-west;

south-west of Torquay, central coastal Victoria (38°20'54"S,

144°18'35"E).

Stratigraphic position. Upper Jan Juc Marl (Holdgate and

Gallagher, 2003; Webb, 1995).

Age. Late Oligocene. Siesser (1979) suggested that the

Oligocene-Miocene boundary occurs 2.5 m below the Jan Juc

Marl-Puebla Clay contact. Li and colleagues (1999) noted that

the local nannoplankton datum used by Siesser (1979) to esti-

mate the position of the Oligocen-Miocene boundary, is no

longer valid because the age of this datum is -24.5 Ma, while

the recognised date of the Oligocene-Miocene boundary is

of 23.8-23.9 Ma (Berggren et al., 1995). Li and others (1999) pro-

of vided foraminiferal biofacies evidence indicating that the top of

the Jan Juc Marl corresponded to the Late Oligocene-Early

Tertiary fossil Cetacea localities in Australia

191

Miocene boundary. Furthermore, the Jan Juc Marl-Puebla

Clay contact has yielded Sr/Sr dates of 23 ± 1 Ma (Kelly

et al., 2001), consistent with the Oligo-Miocene boundary

occurring at the Jan Juc Marl-Puebla Clay contact, and not

within the Jan Juc Marl. Foraminifera from the Jan Juc Marl

correlate this formation with international foram zones

P21-P22, and therefore indicate a Late Oligocene age

(Holdgate and Gallagher, 2003). Sr/Sr dates have yielded an

age of 23.9-27.4 Ma for the Jan Juc Marl exposed at Bird Rock

(Dickinson, 2002; Holdgate and Gallagher, 2003). These data

support a Late Oligocene (Chattian) age for the Jan Juc Marl,

and indicate that the beginning of the Early Miocene occurs

immediately above the top of the Jan Juc Marl, in the

Puebla Clay.

Lithology. Silty glauconitic marl and clayey sandy glauconitic

calcarenite (Abele, 1979; Glover, 1955; Holdgate and

Gallagher, 2003; Raggatt and Crespin, 1955; Webb, 1995).

Material. Fossil preservation is generally fairly good, however

fossils from the upper beds exposed in the bluff are often cor-

roded. Typical elements preserved include ribs, vertebrae,

teeth, tympanies, and periotics. Less common are skulls and

partially articulated skeletons.

Fauna. Heterodontus cainozoicus, Carcharias elegans,

Carcharias taurus, Carcharias sp., Odontaspis incurva,

Carcharoides totuserratus, Carcharodon angustidens, Isurus

desori, Isurus planus, Isurus sp., Dasyatis sp., Myliobatis sp.,

Megalops lissa (Stinton, 1958), Pterothrissus pervetustus

(Stinton, 1958), Heterenchelys regularis (Stinton, 1963),

Astroconger rostratus (Stinton, 1958), Urconger rectus

(Stinton, 1963), Merluccius fimbriatus, Gadus refertus

(Stinton, 1958), Ophidion granosum (Stinton, 1963),

Trachichthodes salebrosus (Stinton, 1958), Cleidopus carver-

nosus (Stinton, 1963), Sillago pliocaenica (Stinton, 1958),

Coelorhynchus elevatus, Xiphias sp., Aves indet., Mysticeti

family indet., Mammalodon colliveri (Pritchard, 1939),

Mammalodon sp., Mammalodon new species 1, Mysticeti new

family, Prosqualodon sp., ?Eurhinodelphinidae.

2.8 Waurn Ponds Quarry

Geographic location. Waurn Ponds Quarry, operated by Blue

Circle Southern Cement Ltd, south of Waurn Ponds (near

38°12'S, 144°16'E).

Stratigraphic position. Waurn Ponds Limestone Member of the

Jan Juc Formation; a lateral equivalent of the lower part of the

Point Addis Limestone (Holdgate and Gallagher, 2003;

Nicolaides and Wallace, 1997).

Age. Late Oligocene (-24-27 Ma) (Abele et al., 1988; Holdgate

and Gallagher, 2003; Nicolaides and Wallace, 1997).

Lithology. Bryozoal calcarenite with some interbedded marls.

These sediments are capped by a subaerial exposure surface

that is laterally equivalent to a similar horizon at Point Addis.

At Waurn Ponds Quarry, this horizon is heavily cemented

(Abele et al., 1988; Holdgate and Gallagher, 2003; Nicolaides

and Wallace, 1997; Webb, 1995).

Material. The completeness and quality of preservation of

fossil cetacean material varies. However, the vast majority

of cetacean fossils consist of disarticulated and isolated ele-

ments. Some fossils preserve fine surface detail on bones and

teeth, whereas others are highly polished and worn with no

surface detail present.

Fauna. Carcharias taurus, Carcharias macrotus, Carcharoides

totuserratus, Carcharodon angustidens, Isurus desori, Isurus

sp., ILamna cattica, Lamna sp., Carcharhinus sp., Galeocerdo

sp., Notorynchus cepedianus, Myliobatis sp., Diprotodontidae

(D. Pickering and T. Rich pers. comm.), cf. Mammalodon

colliveri, Mammalodon sp., Mammalodontidae sp. indet.,

“Cetotheriidae”, Mysticeti new genus, Mysticeti family indet.,

cf. Squalodontidae indet., Odontoceti family indet.

2.9 Ocean Grove

Geographic location. Near Ocean Grove, on the Bass Strait

side of the Bellarine Peninsula (near 38°15'S, 144°3FE).

Stratigraphic position. Puebla Clay (Abele et al., 1988).

Age. Early Longfordian (Early Miocene).

Lithology. Calcareous sand (Abele et al., 1988).

Material. Bone fragments.

Fauna. Cetacea indet.

3. The Port Phillip Basin (Fig.7)

3.1 Batesford Quarry

Geographic location. Australian Cement Company quarry

south of Batesford, on the western bank of the Moorabool

River, west of Geelong (38°06'S, 144°17'E).

Stratigraphic position. Batesford Limestone; unconformably

overlies Palaeozoic granite, and grades conformably up into the

overlying Fyansford Formation (Abele et al., 1988; Holdgate

and Gallagher, 2003; Webb, 1995).

Age. The lower 21 m of the Batesford Limestone is

Longfordian (Lower Miocene) in age, whereas the upper 12 m

of the formation is the Lepidocyclina-beanng type section of

the uppermost Lower to lower Middle Miocene Batesfordian

local marine stage (Holdgate and Gallagher, 2003).

Lithology. Fossil cetaceans have been recovered from the basal

beds, which are composed of calcareous sand and gravel.

The upper Batesfordian part of the formation consists of

biocalcarenite (Abele et al., 1988; Bowler, 1963; Webb, 1995).

Material. In both upper and lower parts of the Batesford

Limestone, cetacean fossils are generally well preserved with

only slight mi neralisation of original bone. The majority of

cetacean fossils have been found in the Batesfordian-age upper

beds. Rib fragments and vertebrae are the most commonly rep-

resented elements, with tympanic bullae being poorly repre-

sented. Elements of the appendicular skeleton and cranial

remains are very rare. Only two skeletons represented by assoc-

iated elements have been recovered from Batesford Quarry.

192

E. M. G. Fitzgerald

Fauna. Orectolobus sp., Carcharias taurus, Carcharodon

megalodon, Isurus desori, Isurus hastalis, lsurus oxyrinchus,

Isurus cf. oxyrinchus, Isurus planus, Isurus retroflexus, Isurus

cf .paucus, Isurus sp., Lamnidae incerta sedis, Carcharhinus cf.

brachyurus, Carcharhinus sp., Galeocerdo aduncus, Sphyrna

sp., Notorynchus primigenius, Pristiophorus lanceolatus,

Labrodon sp., Spheniscidae, Diprotodontidae indet., “Ceto-

theriidae” indet., Physeteridae, ?Squalodontidae new genus and

new species.

3.2 Moorabool River

Geographic location. North-west of Geelong (near 37°56'S,

144°09'E).

Stratigraphic position. Lower Maude Formation (Abele et al.,

1988).

Age. Latest Late Oligocene-earliest Miocene (zones P22-N4)

(Abele et al., 1988; Holdgate and Gallagher, 2003).

Lithology. Shelly bryozoal calcarenite.

Material. Cetacean material consists of one tympanic bulla and

bone fragments.

Fauna. Heterodontus cainozoicus, Carcharias sp., Isurus sp.,

cf. Mammalodontidae.

3.3 North Shore, Corio Bay

Geographic location. Outcrop along north shore of Corio Bay,

near Geelong (near 38°06'S, 144°24'E).

Stratigraphic position. Fyansford Formation (Abele et al.,

1988).

Age. Youngest part of the Fyansford Formation, representing

zone N12 (Bairnsdalian, Middle Miocene) (Abele et al., 1988).

Lithology. Basal sandy calcarenite passing upwards into cal-

careous silt and clay with sandy calcarenite interbeds (Abele et

al., 1988).

Material. Bone fragments.

Fauna. Lamna sp., Cetacea indet.

3.4 Curlewis

Geographic location. Cliff sections along the northern side of

the Bellarine Peninsula, near Curlewis (near 38° ll’S,

144°30E).

Stratigraphic position. Fyansford Formation (Abele et al.,

1988).

Age. Early Middle Miocene (Abele et al., 1988).

Lithology. Calcareous clay and marl with thin bryozoal

calcarenite interbeds (Abele et al., 1988).

Material. Bone fragments.

Fauna. Heterodontus cainozoicus, Isurus hastalis,

Trachichthodes salebrosus (Stinton, 1958), Diodon sp., Cetacea

indet.

3.5 Beaumaris

Geographic location. Coastal exposures located east of

Rickett’s Point on the west shore of Beaumaris Bay, north-east

shore of Port Phillip Bay (37°59'S, 145°03E).

Stratigraphic position. Basal Black Rock Sandstone (Abele et

al., 1988; Holdgate and Gallagher, 2003).

Age. Cheltenhamian to Kalimnan (latest Late Miocene-earliest

Early Pliocene) (Abele et al., 1988; Dickinson et al., 2002; Gill,

1957; Holdgate and Gallagher, 2003; Mallett, 1977).

Lithology. Most fossil vertebrates have been recovered from a

layer of ferruginous and phosphatic nodules in a matrix of

quartz sand and gravel. This nodule bed occurs at the base of

the Black Rock Sandstone. Less abundant fossil vertebrates

occur in the sediments immediately above the nodule horizon,

this layer consisting of calcareous sandstone and sandy marl

(Abele et al., 1988; Gill, 1957; Singleton, 1941).

Material. Cetacean fossils derived from the nodule bed consist

of fragmentary, isolated elements that are usually highly worn

and polished, having undergone significant post-mortem trans-

port. These fossils include indeterminate bone fragments, ribs,

vertebrae, forelimb elements, teeth, rostral and cranial frag-

ments, periotics, tympanic bullae, and incomplete mandibles.

Vertebrate fossils from the sandy beds overlying the nodule

horizon are better preserved than the nodule material, with a

lesser degree of replacement of bone by secondary mineralis-

ation. The sandy bed material includes ribs, vertebrae, teeth,

mandibles, periotics, tympanic bullae, middle-ear ossicles, and

rare partially articulated skeletons.

Fauna. Heterodontus cainozoicus, Carcharias taurus,

Carcharodon megalodon, Parotodus benedenii, Isurus desori,

Isurus hastalis, Isurus retroflexus, I Lamna, Megascyliorhinus

sp., Carcharhinus cf. brachyurus, Carcharhinus sp.,

Galeocerdo aduncus, Pristiophorus lanceolatus, Myliobatis

sp., Edaphodon mirabilis, Edaphodon cf. mirabilis, Edaphodon

sweeti, Ischyodus cf. dolloi, Sillago pliocaenica (Stinton,

1958), Lactarius tumulatus (Stinton, 1963), ?Trionychidae,

Pseudaptenodytes macraei, IPseudaptenodytes minor ,

Spheniscidae, Diomedea thyridata (Wilkinson, 1969),

Zygomaturus gilli (Woodburne, 1969; Stirton, 1967), Kolopsis

sp. cf. K. torus (Rich et al., 2003), Phocidae (Fordyce and

Flannery, 1983), “Cetotheriidae” indet., cf. Balaena,

Balaenidae indet., Balaenoptera sp., Megaptera sp.,

Physetodon baileyi (probably Physeteridae indet.), Scaldicetus

macgeei, Physeteridae indet., Ziphiidae, Delphinidae indet.

4. Gippsland Basin (Fig.9)

4.1 Merrimans Creek

Geographic location. Approximately 25 m from the surface, in

the Gippsland Cement Quarry on Merrimans Creek, about 19

km south-east of Rosedale, Gippsland, Victoria (near 38°15'S,

146°51E).

Stratigraphic position. Gippsland Limestone Formation

(Holdgate and Gallagher, 2003).

Tertiary fossil Cetacea localities in Australia

193

Figure 9. Fossil Cetacea localities in the Gippsland Basin, eastern Victoria. Solid circles represent fossil Cetacea localities.

Age. Longfordian; Early Miocene (planktonic foraminiferal

zones N5-N7).

Lithology. Light grey fossiliferous marly limestone and

interbedded limestone and marl.

Material. Fragmentary mandible and probable vertebrae.

Fauna. Isurus hastalis, Mysticeti.

4.2 Rose Hill

Geographic location. On Mitchell River, near Bairnsdale, East

Gippsland, Victoria (near 37°49'S, 147°36'E).

Stratigraphic position. Tambo River Formation.

Age. Mitchellian (Late Miocene), representing planktonic

foraminiferal zones N16-N17. Dickinson (2002) recorded Sr/Sr

dates averaging 6.0 Ma for the top of the type section of the

Tambo River Formation at Swan Reach.

Lithology. Uniform marl and marly limestone, and glauconitic

sandy coquinas (Abele et al., 1988).

Material. Bone fragments.

Fauna. Cetacea indet.

4.3 Jemmys Point

Geographic location. Low cliffs at Jemmys Point, between the

south-east end of “The Narrows”, and the confluence of North

Arm and Cunningham Arm, west of Lakes Entrance, East

Gippsland (37°53’S, 147°58E).

Stratigraphic position. Jemmys Point Formation (Abele et al.,

1988; Singleton, 1941; Wilkins, 1963).

Age. Kalimnan (Early Pliocene); planktonic foraminiferal

zones N18-N19 (Holdgate and Gallagher, 2003).

Lithology. Sandy clay with shell beds (Abele et al., 1988;

Carter, 1985; Holdgate and Gallagher, 2003; Wilkins, 1963).

Material. Isolated bone fragments, vertebrae, and one

incomplete skull.

Fauna. Carcharodon carcharias, Notorynchus cepedianus,

Mesoplodon longirostris.

4.4 North Arm

Geographic location. “Golden Point” property, North Arm,

Lake King, near Lakes Entrance, East Gippsland (close to

37 0 51'S, 148°58E) (Bearlin, 1987).

Stratigraphic position. Jemmys Point Formation.

Age. Kalimnan (Early Pliocene); planktonic foraminiferal

zones N18-N19 (Holdgate and Gallagher, 2003).

Lithology. Calcareous shelly sand, with phosphorite con-

cretions (Abele et al., 1988; Carter, 1985).

194

E. M. G. Fitzgerald

Material. One incomplete skeleton comprising: almost

complete skull, mandibles, periotics, tympanic bullae, com-

plete series of cervical vertebrae, isolated thoracic and lumbar

vertebrae, and ribs.

Fauna. Megaptera new species 1 .

4.5 Trident Arm

Geographic location. Trident Arm of Lake Tyers, east of Lakes

Entrance, East Gippsland, Victoria (near 37°49'S, 148°08'E).

Stratigraphic position. Phosphatic horizon at the base of the

Jemmys Point Formation (M. Wallace, pers. comm.).

Age. Kalimnan (Early Pliocene); planktonic foraminiferal

zones N18-N19 (Carter, 1985; Holdgate and Gallagher, 2003).

Lithology. Cemented phosphatic and glauconitic calcarenite

with Ostrea shells.

Material. One partial cranium.

Fauna. ?Balaenidae.

4.6 Newmerella

Geographic location. Railway cutting 3 km west of

Newmerella, Orbost district, East Gippsland, Victoria (near

37°45'S, 148°24'E).

Stratigraphic position. Probably the Lindenow Sandstone

Member, Gippsland Limestone Formation.

Age. Longfordian (Early Miocene), planktonic foraminiferal

zones N5-N7 (Holdgate and Gallagher, 2003).

Figure 10. Areas with fossil cetacean-bearing localities in South

Australia. Modified from Alley et al. (1995).

Lithology. Ferruginised fine quartz sandstone (Abele et al.,

1988)

Material. Cetacean fossils are relatively well preserved. The

most significant specimens occur in private collections and

these include a mysticete skull and mandibles, and postcranial

elements (Bearlin, 1987). Material in Museum Victoria

includes the posterior part of a large mandible, and one large

vertebra.

Fauna. Carcharodon megalodon, ?”Cetotheriidae”.

South Australia (Fig. 10)

5. St Vincent Basin (Fig. 11)

5.1 Port Willunga

Geographic location. 300-400 m south of the old jetty, Port

Willunga Beach, approximately 45 km south of Adelaide (this

is the locality data for the cranium; the exact location for the

radius is unknown, but it was collected in the vicinity of Port

Willunga) (Pledge, 1994; N. Pledge, pers. comm.) (near

35°19'S, 138°27’E).

Stratigraphic position. Ruwarung Member, Port Willunga

Formation (N. Pledge, pers. comm.).

Figure 11. Fossil Cetacea localities in the St Vincent and Murray

basins in south-east South Australia. Solid circles represent fossil

Cetacea localities. Modified from Alley et al. (1995).

Tertiary fossil Cetacea localities in Australia

195

Age. Willungan (Early Oligocene; Rupelian), 30-32 Ma (Alley

et al., 1995; Holdgate and Gallagher, 2003).

Lithology. Bryozoal marl limestone containing chert nodules

(Alley et al., 1995; N. Pledge pers. comm.).

Material. One right radius (Pledge, 1994) and one incomplete

cranium.

Fauna. Cetacea indet., cf. Aetiocetidae (N. Pledge, pers.

comm.).

5.2 Abattoirs Bore

Geographic location. Abattoirs Bore, Dry Creek, about 10 km

north of Adelaide (34°50'00"S, 138°36'35"E) (Howchin, 1919;

Bearlin, 1987).

Stratigraphic position. Dry Creek Sand.

Age. Late Pliocene (Alley et al., 1995).

Lithology. Shelly sand.

Material. One isolated periotic.

Fauna. Carcharodon carcharias, Balaenidae (Howchin, 1919).

6. Murray Basin (Figs 11, 12)

6.1 Fred’s Landing

Geographic location. Boat launching area about 3 km down-

stream from Tailem Bend, on the east bank of the Murray River

(35°17’S, 139°27’E) (Pledge, 1994).

Stratigraphic position. Upper Buccleuch Formation (Pledge,

1994) .

Age. Early Oligocene.

Lithology. Green and slightly glauconitic fine-grained lime-

stone (Pledge, 1994; Alley et al., 1995).

Material. One heterodont tooth.

Fauna. Cetacea indet.

6.2 Wellington

Geographic location. Near Wellington, on the Murray River,

south-east of Adelaide (Pledge and Rothausen, 1977) (near

35°19’S, 139°23’E).

Stratigraphic position. Ettrick Formation (Pledge and

Rothausen, 1977).

Age. Late Oligocene (Alley et al., 1995).

Lithology. Glauconitic marl, calcareous clay, and mudstone,

with silt and sand (Pledge and Rothausen, 1977; Alley et al.,

1995) .

Material. Several heterodont teeth (possibly associated).

Fauna. Hexanchus agassizi. Cetacea indet. ( -Metasqualodon

harwoodi).

Figure 12. Fossil Cetacea localities along the Murray River north-east

of Adelaide. Solid circles represent fossil Cetacea localities. Modified

from Lukasik and James (1998).

6.3 Wongulla

Geographic location. Cliffs on east bank of Murray River

opposite Wongulla, approximately 5 km south of Devon Downs

(near 34°41’S, 139°35’E).

Stratigraphic position. Upper Member, Mannum Formation.

Age. Early Miocene (Lukasik and James, 1998).

Lithology. Yellow-orange calcarenite.

Material. Anterior portions of two large mandibles and

associated incomplete vertebrae.

Fauna. Mysticeti.

6.4 Blanchetown

Geographic location. Close to the north end of the channel

between Notts Island and Murray River cliffs, about 4 km south

of Blanchetown Bridge, north-east of Adelaide (N. Pledge,

pers. comm.) (near 34°28’S, 139°36’E).

Stratigraphic position. Upper Member, Mannum Formation

(Lukasik and James, 1998).

Age. Cetacean fossils were recovered about 3-4 m below the

Lepidocyclina Zone (N. Pledge, pers. comm.), and therefore,

pre-Batesfordian. The minimum age of the upper Mannum

Formation is earliest Longfordian, based on the presence of

Operculina victoriensis at the base of this unit. The Upper

Member of the Mannum Formation is overall no younger than

Early Miocene (Lukasik and James, 1998).

Lithology. Fine-grained calcarenite.

196

E. M. G. Fitzgerald

Material. One mandible with two posterior teeth in situ and

four associated vertebrae. These fossils represent one individ-

ual, and other bones remain unprepared at the South Australian

Museum.

Fauna. ?Squalodontidae.

6.5 MacBean’s Pound

Geographic location. About 6.5 km from Blanchetown, Murray

River (near 34°25'20"S, 139°37’05"E).

Stratigraphic position. Upper Member, Mannum Formation.

Age. Early Miocene.

Lithology. Bryozoal calcarenite.

Material. Mandibles.

Fauna. ?”Cetotheriidae”.

6.6 Murbko

Geographic location. Murray River cliffs near Murbko

Homestead, 22.5 km north-east of Blanchetown (34 o 07'55"S,

139°39’05"E).

Stratigraphic position. Glenforslan Formation (sensu Lukasik

and James, 1998).

Age. The presence of Lepidocyclina howchini indicates a

Batesfordian (early Middle Miocene) age (Lukasik and James,

1998).

Lithology. Bryozoal calcarenite.

Material. One almost complete cranium.

Fauna, cf. Parietobalaena sp.

6.7 Waikerie

Geographic location. 100 m downstream from the Sunlands

Pumping Station, 8 km west of Waikerie, Murray River (near

34°09'S, 139°55’E).

Stratigraphic position. Pata Formation.

Age. Balcombian (Middle Miocene) (Lukasik and James,

1998).

Lithology. Yellow-orange fine calcarenite with muddy bands.

Material. One incomplete scapula.

Fauna. Mysticeti indet.

6.8 Winkie

Geographic location. Riverbed, Murray River, Gerard Mission,

near Winkie (near 34°22’45"S, 140°28'30"E).

Stratigraphic position. ?Bookpurnong Formation (Bearlin,

1987).

Age. ?Mio-Pliocene.

Lithology. ?Marl, silty clay and minor fine sand (Alley et al.,

1995).

Material. Incomplete cranium and mandible (Bearlin, 1987).

Fauna. Balaenopteridae.

6.9 Sunlands Pumping Station

Geographic location. Beds immediately above a hardground of

cemented sand and oyster shells, 3-4 m above the base of the

cliffs, Sunlands Pumping Station, 8 km west-north-west of

Waikerie, Murray River (near 34°09’S, 139°55'E) (Pledge,

1985).

Stratigraphic position. Loxton Sand (Alley et al., 1995; Pledge,

1985).

Age. Early Pliocene.

Lithology. Poorly sorted coarse sand containing pebbles.

Material. Isolated elements comprising vertebrae and teeth

(Pledge, 1985).

Fauna. Heterodontus cf. cainozoicus, Orectolobus gippsland-

icus, Carcharias sp., Carcharodon cf. megalodon, Isurus

hastalis, Lamna cf. cattica, Mustelus sp., Carcharhinus cf.

b rachyurus, Galeocerdo aduncus, Galeorhinus cf. australis,

Sphyrna sp., Pristiophorus lanceolatus, cf. Myliobatis sp.,

Labroidei indet., Monacanthidae indet., Diodon formosus, cf.

Zygomaturus, Phascolarctos maris, Dorcopsis sp., cf. Dugong

sp. (Pledge, 1992; Domning, 1996), Mysticeti indet.,

?Delphinidae indet.

6.10 Overland Corner

Geographic location. Near Overland Corner, Murray River,

about 10 km north-west of Lake Bonney (near 34°09'20"S,

140°20'E).

Stratigraphic position. ?Glenforslan Formation.

Age. ?Batesfordian (early Middle Miocene).

Lithology. Bryozoal calcarenite.

Material. One incomplete cranium.

Fauna. Mysticeti.

7. Gambier Basin

7.1 Mount Gambier (Fig. 13)

Geographic location. The old “Knights and Pritchard’s”

dimension stone quarry, near Marte, 10 km west of Mount

Gambier (near 37°48'S, 140°09'E).

Stratigraphic position. Camelback Member, Gambier

Limestone (Alley et al., 1995).

Age. Late Oligocene (planktonic foraminiferal zones P21-P22)

(Li et al., 2000).

Lithology. Pale cream-white porous bryozoal calcarenite.

Tertiary fossil Cetacea localities in Australia

197

Figure 13. Fossil Cetacea locality in the Gambier Basin, extreme

south-east of South Australia. Solid circles represent fossil Cetacea

localities. Modified from Alley et al. (1995).

Material. Fossil preservation is generally very good. Cetaceans

are represented by isolated teeth, vertebrae, and cranial

fragments.

Fauna. Carcharodon angustidens, Spheniscidae, “ Squalodon ”

gambierensis, Mysticeti indet., Prosqualodontidae.

8. Lake Eyre Basin (Fig. 14)

8.1 Lake Namba

Geographic location. 4 km south of Ericmas Quarry, which

occurs at the base of low bluffs just south of track to Billeroo

Waterhole on western side of Lake Namba, Lake Frame area,

South Australia (31°14'S, 140°14'E) (Fordyce, 1983; T.Rich,

pers. comm.).

Stratigraphic position. Ericmas Fauna, base of upper member,

Namba Formation, equivalent to unit 4 of Namba Formation at

Lake Pinpa.

Age. Late Oligocene-Middle Miocene (Callen and Tedford,

1976). Palynofloras suggest a Miocene-Pliocene age range for

the Namba Formation (Alley et al., 1995). Fordyce (1983) sug-

gested that the Namba Formation was Middle Miocene in age,

and no older than Batesfordian-Balcombian. Recently,

Woodbume and colleagues (1993) have indicated that the

Etadunna Formation (a correlate of the Namba Formation) is

perhaps 24-28 Ma. If the latter dates are accurate, they would

imply that the Namba Formation is Late Oligocene in age.

However, due to the inconsistent results of various dating

Figure 14. Fossil Cetacea localities east of Lake Frome, in the Lake

Eyre Basin, north-east South Australia. Solid circles represent fossil

Cetacea localities. Modified from Tedford et al. (1977).

methods, the Namba Formation is considered herein as Late

Oligocene-Middle Miocene in age.

Lithology. Illite and kaolinite clays (Alley et al., 1995).

Material. One right periotic.

Fauna. Neoceratodus djelleh, Neoceratodus eyrensis,

Neoceratodus gregoryi, Neoceratodus sp. 1, Neoceratodus sp.

4, Osteichthyes indet., Chelidae, Crocodylia, Columbidae,

Anseriformes, Obdurodon insignis, Dasyuridae, Raemeother-

ium yatkolai, Madakoala devisi, Madakoala wellsi, Pildra

antiquus, Pildra secundus, Petauridae, Eurhinodelphinidae.

8.2 Lake Pinpa

Geographic location. Western margin and floor of Lake Pinpa

(31°10'S, 140°14'E) (Fordyce, 1983).

Stratigraphic position. At top of unnamed member 1, Namba

Formation (Fordyce, 1983).

Age. Late Oligocene-Middle Miocene (Alley et al., 1995;

Callen and Tedford, 1976; Woodburne et al., 1993).

Lithology. Illite and kaolinite clays.

Material. One incomplete associated skeleton, and isolated left

periotics and radius.

Fauna. Neoceratodus djelleh, Neoceratodus forsteri,

Neoceratodus eyrensis, Neoceratodus gregoryi, Neoceratodus

nargun, Neoceratodus sp. 1, Neoceratodus sp. 2, Neoceratodus

sp. 3, Neoceratodus sp. 4, Osteichthyes indet., Emydura sp.,

Meiolaniidae, Crocodylia, Passeriformes, Rallidae, Podiciped-

idae, Burhinidae, Phoenicopteridae, Anatidae, Pelecanus

tirarensis, Phalacrocoracidae, Obdurodon insignis, Muramura

sp., Ilaria illumidens, Ilariidae indet., Madakoala devisi,

?Potoroidae, ?Macropodidae, Petauridae, Pildra antiquus,

Chunia cf. illuminata, Pilkipildra handae, Miralina cf. minor,

Eurhinodelphinidae.

198

E. M. G. Fitzgerald

8.3 Lake Yanda

Geographic location. Western side of Lake Yanda, Lake Frome

area, South Australia (31°0rS, 140°19'E).

Stratigraphic position. Yanda Fauna, near the contact between

the two unnamed members of the Namba Formation (Rich et

al., 1991).

Age. Late Oligocene-Middle Miocene (Alley et al., 1995;

Callen and Tedford, 1976; Woodburne et al., 1993).

Lithology. Illite and kaolinite clays (Alley et al., 1995).

Material. One incomplete tympanic bulla.

Fauna. Neoceratodus djelleh, Neoceratodus eyrensis,

Neoceratodus gregoryi, Neoceratodus sp. 1, Neoceratodus sp.

4, Osteichthyes indet., Crocodylia, Phoenicopteridae, Anatidae,

Dasylurinja kokuminola, Ilaria sp., Djilgaringa thompsonae,

Miralina cf. minor, Eurhinodelphinidae.

Tasmania

9. Bass Basin

9.1 Fossil Bluff (Fig.4)

Geographic location. Coastal cliff section at Fossil Bluff, Table

Cape, near Wynyard, north-east of Launceston, Bass Strait

coast of Tasmania (40°58.8'S, 145°43.9'E). Strictly speaking,

the Fossil Bluff locality is situated approximately 40-50 km

south of the southern margin of the Bass Basin (N. Kemp, pers.

comm.).

Stratigraphic position. Fossil Bluff Sandstone.

Age. Longfordian (Early Miocene). The Fossil Bluff Sand-

stone has a planktonic foraminiferal fauna which corresponds

to zone N4 (earliest Early Miocene; Aquitanian = 21-23.9 Ma)

(Quilty, 1980). Fordyce (2003) indicated that the Aquitanian in

south-east Australia is comparable to the Chattian (Late

Oligocene) in New Zealand. The odontocete genus Pros-

qualodon occurs in the Waitakian (latest Oligocene-earliest

Miocene) of New Zealand (Fordyce, 1984, 1991), the Late

Oligocene of Victoria and perhaps South Australia (Fitzgerald,

2004, and herein), the Early Miocene of Tasmania (Flynn,

1923; Fitzgerald, 2004), and the Early Miocene of Patagonia,

Argentina (Lydekker, 1899; Cabrera, 1926; Fordyce, 2002b;

Muizon, 2002).

Lithology. Fine siltstones and shales, and glauconitic calcare-

ous sandstone (Kemp, 1991).

Material. Isolated teeth, vertebrae, ribs, partially articulated

postcranial skeletons, and one associated skeleton comprising:

skull, mandibles, teeth, almost complete right forelimb, ribs, 2+

thoracic vertebrae, 4+ lumbar vertebrae, and other elements.

The latter specimens represent the holotype of Prosqualodon

davidis. Unfortunately the skull is now lost (see Mahoney and

Ride (1975) for details). Isolated teeth from Fossil Bluff, that

represent prosqualodontids congeneric, and perhaps conspecif-

ic, with Prosqualodon davidis, are present in Museum Victoria

and the South Australian Museum.

Fauna. Heterodontus cainozoicus, Carcharias taurus,

Ischyodus mortoni, Wynyardia bassiana, Prosqualodon

davidis, Prosqualodon cf. davidis. The holotype of Scaptodon

lodderi was discovered near this locality.

9.2 Cameron Inlet (Fig.4)

Geographic location. Excavations of drains around Memana,

near Cameron Inlet, east coast of Flinders Island, Bass Strait,

Tasmania (near 39°59'S, 148°05'E).

Stratigraphic position. Cameron Inlet Formation (Sutherland

and Kershaw, 1971).

Age. Late Early-Late Pliocene (Sutherland and Kershaw,

1971).

Lithology. Fine silty coquina limestones and sands (Sutherland

and Kershaw, 1971).

Material. Vertebrae, ribs, incomplete skulls, partial mandibles,

teeth, periotics, and tympanic bullae.

Fauna. Carcharodon megalodon, Carcharodon carcharias,

Isurus hastalis, Isurus oxyrinchus, Carcharhinus sp., cf.

Balaenoptera, cf. Megaptera, Physeter sp., Ziphiidae indet.,

?Delphinidae.

10. South Tasman Rise (Fig. 15)

10.1 Tasman Fracture Zone

Geographic location. Pipe dredge 147DR013, 2700-3900 m

depth, northern end of eastern Tasman Fracture Zone, south-

west of Tasmania (45°07'S, 144°31'E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995). The one cetacean from

this site (a probably indeterminate ziphiid) may indicate a max-

imum age for these sediments of Late Miocene. Exon and oth-

ers (1995) suggested that fossil ziphiids from the South Tasman

Rise represented a similar fauna to that of Cameron Inlet, which

has a mid-Late Pliocene age (Sutherland and Kershaw, 1971).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One incomplete rostrum.

Fauna. Ziphiidae indet.

10.2 Seamount east of South Tasman Rise

Geographic location. Dredge 147DR038, 2050-2300 m depth,

on a seamount east of South Tasman Rise (near 45°43.0’S,

149°00’E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One incomplete rostrum.

Fauna. Ziphiidae indet.

Tertiary fossil Cetacea localities in Australia

199

Figure 15. Fossil Cetacea localities offshore southern Tasmania. Solid

circles represent fossil Cetacea localities. Modified from Exon et al.

(1997).

10.3 Eastern South Tasman Rise

Geographic location. Dredge site 147DR50, 2050-2300 m

depth, east- west trending high on the eastern side of the South

Tasman Rise (near 45°30'S, 147°20E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One incomplete rostrum, and one skull.

Fauna. ?Mysticeti, Ziphiidae.

10.4 North-east South Tasman Rise

Geographic location. Dredge site 147DR052, 2200-2370 m

depth, north-east South Tasman Rise, south-west of Tasmania

(near 45°38.1'S, 146°24.5E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One small periotic.

Fauna. Odontoceti.

10.5 East Tasman Rise

Geographic location. Dredge site 147DR043, 3030-3600 m

depth, eastern scarp of East Tasman Rise, south of Tasmania

(near 43°54’S, 15U18E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One incomplete tympanic bulla (Exon et al., 1995).

Fauna. ?Mysticeti indet.

11. East Tasman Plateau (Fig. 15)

11.1 Eastern Scarp

Geographic location. Dredge 147DR043, 3030-3600 m depth,

eastern scarp of East Tasman Plateau, south-east of Tasmania

(near 43°54’S, 151°18E) (Exon et al., 1995).

Stratigraphic position. Unnamed lithological unit.

Age. Late Neogene (Exon et al., 1995).

Lithology. ?Silicified limestones (Exon et al., 1995).

Material. One incomplete skull.

Fauna, cf. Mesoplodon sp.

Discussion

Fordyce (1984) identified 21 and Fitzgerald (2004) 26 fossil

Cetacea localities in Australia. Further discoveries, and more

study of museum collections has resulted in the recognition of

56 Tertiary fossil Cetacea localities in Australia. Thirty-two of

these localities occur in Victoria, 16 in South Australia, and

eight in and around Tasmania. All are geographically distinct

and generally include only one stratigraphic unit that has

yielded fossil cetaceans. The number of recognised Australian

fossil Cetacea localities compares with approximately 58-60

recognised Australian Tertiary terrestrial mammal localities

(Rich et al., 1991; pers. obs.). That almost all the fossil

cetacean-bearing localities occur in the south-east of the conti-

nent, while extensive areas of Cainozoic marine sedimentary

rock outcrop occur outside this region, points to a potentially

rich record of fossil cetaceans in unexplored areas. Moreover,

it is highly probable that many localities in south-east Australia

remain unrecognised. This is suggested by the broad distribu-

tion of cetacean-bearing localities within marine sedimentary

basins. However, a perhaps less fortuitous aspect of this local-

ity distribution and the faunal lists and material derived from

these localities is the lack of cetacean fossil density at these

localities.

Few localities have yielded diagnostic, reasonably complete

skulls and/or skeletons. These localities are: Victoria; Arch Site

(Grange Burn), Gibson’s Steps, Curdie, Bells Headland, Bells

Beach, Deadman’s Gully, Bird Rock, Batesford Quarry, North

Arm, Newmerella; South Australia; Murbko, Lake Pinpa; and

Tasmania; Fossil Bluff. The source stratigraphic units (and

lithologies) at the latter localities occur elsewhere, and it may

be expected that more reasonably complete specimens will be

discovered at these localities in future. The most important g

eneral area for fossil cetaceans in Australia is the coastline

south-west of Torquay, in the Torquay Basin, Victoria. Eight

skulls/skeletons have been collected from this area, with all

derived from the Upper Oligocene Jan Juc Marl and its lateral

200

E. M. G. Fitzgerald

equivalents. Limitations to conducting fieldwork to prospect

for, and recover, fossil cetaceans in this area include: extreme

rarity of shore platforms being scoured by storm tides to

remove sand and algae, thus uncovering larger areas of acces-

sible outcrop; and dangerously unstable, steep, and high cliff

sections. Despite these limitations, systematic prospecting of

the coastline in this area will likely yield more important fossil

cetaceans.

The Australian pre-Pleistocene fossil record of cetaceans, as

currently understood, can be summarised thus: Eocene - not

present; Early Oligocene - poorly known, but great potential;

Late Oligocene - poorly known at present but will probably be

the second-best known period of cetacean evolution in

Australian waters; Early Miocene - poorly known, but great

potential; Middle Miocene — poorly known, but great poten-

tial; Late Miocene — reasonably well-known; Early Pliocene

— best known period of cetacean evolution in Australian

waters; and Late Pliocene — poorly known.

Although the fossil record of Cetacea in Australia is known

only in the broadest of terms, the identification of 56 recog-

nised pre-Pleistocene fossil cetacean-bearing localities

emphasises the wide distribution of cetacean fossils, and the

potential for the elucidation of the details of this fossil

record. These localities indicate geographic areas that may be

fruitful for future discoveries, as well as stratigraphic horizons

that are known to produce fossil cetaceans. Fossil cetaceans

have arguably received the least attention of all groups of

Australian fossil vertebrates, and yet they have, potentially, the

richest pre-Pleistocene fossil record of all mammalian taxa in

Australia.

Acknowledgements

T.H. Rich and an anonymous reviewer are thanked for their

critical review of the manuscript. The author thanks T.H. Rich,

D. Henry, and N.W. Longmore for access to specimens in their

care at Museum Victoria. R. Bearlin, T. Darragh, R. Schmidt,

and M. Wallace, are thanked for providing valuable informa-

tion. S. Wright is thanked for allowing the author to study his

important fossil collection. N. Kemp provided unpublished

photographs of the Flinders Island fossil cetaceans, and

assisted with information on Tasmanian fossil Cetacea.

N. Pledge graciously provided the author with locality infor-

mation on South Australian fossil Cetacea. D. Bohaska (United

States National Museum of Natural History) is thanked for his

assistance with literature. K. Piper and L. Schwartz are thanked

for discussions on Australian Tertiary ma mm als. All of the most

important Australian fossil cetacean specimens were discov-

ered, and in many cases collected, by amateur palaeontologists,

non-specialists, and other members of the public. Without their

efforts over the last 150 years, it would be impossible to carry

out this research. These fossil collectors are thanked for their

diligence, and foresight shown in donating their discoveries to

relevant institutions. This work forms part of a Ph.D. thesis

undertaken in the School of Geosciences, Monash University,

and Museum Victoria, and was financially supported by an

Australian Postgraduate Award. Scientific Editor G.C.B. Poore

provided helpful editorial advice on the manuscript.

References

Abele, C. 1979. Geology of the Anglesea area, central coastal Victoria.

Geological Survey of Victoria Memoir 31: 1-73.

Abele, C., Gloe, C.S., Hocking, J.B., Holdgate, G., Kenley, P.R.,

Lawrence, C.R., Ripper, D., Threlfall, W.F., and Bolger, P.F. 1988.

Tertiary. Pp. 251-350 in: Douglas, J.G. and Ferguson, J.A. (eds),

Geology of Victoria. Victorian Division, Geological Society of

Australia Incorporated: Melbourne.

Alley, N.F., Lindsay, J.M., Barnett, S.R., Benbow, M.C., Callen, R.A.,

Cowley, W.M., Greenwood, D., Kwitko, G., Lablack, K.L.,

Lindsay, J.M., Rogers, P.A., Smith, PC., and White, M.R. 1995.

Tertiary. Pp. 151-218 in: Drexel, J.F., and Preiss, W.V. (eds), The

geology of South Australia. Volume 2. The Phanerozoic. Bulletin of

the Geological Survey of South Australia 54.

Anonymous. 1939. A fossil whale from Australia. Nature 143: 525.

Arnason, U., and Gullberg, A. 1994. Relationships of baleen whales

established by cytochrome b gene sequence comparison. Nature

367:726-728.

Bames, L.G., and Mitchell, E.D. 1978. Cetacea. Pp. 582-602 in:

Maglio, V.J., and Cooke, H.B.S. (eds), Evolution of African

Mammals. Harvard University Press: Cambridge.

Bearlin, R.K. 1982. A new “shark-toothed dolphin” (Order Cetacea,

Family Squalodontidae) from Batesford, Victoria, Australia. B.Sc.

(Honours) Thesis, Monash University: Clayton. 133 pp.

Bearlin, R.K. 1987. The morphology and systematics of Neogene

Mysticeti from Australia and New Zealand. Ph.D. Thesis,

University of Otago: Dunedin. 212 pp.

Bearlin, R.K. 1988. The morphology and systematics of Neogene

Mysticeti from Australia and New Zealand (abstract of Ph.D.

Thesis, University of Otago, Dunedin). New Zealand Journal of

Geology and Geophysics 31: 257.

Berggren, W.A., Kent, D.V., Swisher, C.C. Ill, and Aubry, M.-P.

1995. A revised Cenozoic geochronology and chronostrati-

graphy. Pp. 129-212 in: Berggren, W.A., Kent, D.V., Aubry,

M.-R, and Hardenbol, J. (eds), Geochronology, Time Scales

and Global Stratigraphic Correlation. SEPM Special Publication

54.

Berta, A., Demere, T., and Gatesy, J. 2003. Systematics and evolution

of the Mysticeti. Journal of Vertebrate Paleontology 23

(Supplement to Number 3): 33A-34A.

Bowler, J.M. 1963. Tertiary stratigraphy and sedimentation in the

Geelong-Maude area, Victoria. Proceedings of the Royal Society of

Victoria 76: 69-137.

Cabrera, A. 1926. Cetaceos fosiles del Museo de La Plata. Revista del

Museo de La Plata 29: 363-411.

Callen, R.A., and Tedford, R.H. 1976. New late Cainozoic rock units

and depositional environments, Lake Frome area, South Australia.

Transactions of the Royal Society of South Australia 100: 125-167.

Camp, C.L., Taylor, D.N., and Welles, S.P. 1942. Bibliography of fos-

sil vertebrates 1934-1938. Geological Society of America Special

Paper 42: 1-663.

Carroll, R.L. 1988. Vertebrate Paleontology and Evolution. W.H.

Freeman and Company: New York. 698 pp.

Carter, A.N. 1985. A model for depositional sequences in the Late

Tertiary of southeastern Australia. Special Publication, South

Australian Department of Mines and Energy 5: 13-27.

Chapman, F. 1912. On the occurrence of Scaldicetus in Victoria.

Records of the Geological Survey of Victoria 3: 236-238.

Chapman, F. 1913. Note on the occurrence of the Cainozoic shark,

Carcharoides, in Victoria. Victorian Naturalist 30: 142-143.

Chapman, F. 1917a. New or little-known Victorian fossils in the

National Museum. Part XX. — Some Tertiary fish-teeth.

Proceedings of the Royal Society of Victoria 29: 134—141.

Tertiary fossil Cetacea localities in Australia

201

Chapman, F. 1917b. New or little-known Victorian fossils in the

National Museum. Part XXI. — Some Tertiary cetacean remains.

Proceedings of the Royal Society of Victoria 30: 32-43.

Chapman, F. 1918. On an apparently new type of cetacean tooth from

the Tertiary of Tasmania. Proceedings of the Royal Society of

Victoria 30: 149-152.

Chapman, F. 1929. Illustrated Guide to the Collection of Fossils

Exhibited in the National Museum of Victoria. H.J. Green,

Government Printer: Melbourne. 55 pp.

Chapman, F. 1930. Note on chimaeroid fish remains from the

Australian Tertiaries. Annals and Magazine of Natural History

Series 10, 8: 139.

Chapman, F., and Cudmore, F.A. 1924. New or little-known fossils in

the National Museum. Part XXVII. — Some Cainozoic fish remains,

with a revision of the group. Proceedings of the Royal Society of

Victoria 36: 107-162.

Chapman, F., and Pritchard, G.B. 1904. Fossil fish remains from the

Tertiaries of Australia. Part I. Proceedings of the Royal Society of

Victoria 17: 267-297.

Chapman, F., and Pritchard, G.B. 1907. Fossil fish remains from the

Tertiaries of Australia. Part II. Proceedings of the Royal Society of

Victoria 20: 59-75.

Cozzuol, M.A. 1996. The record of the aquatic mammals in southern

South America. Pp. 321-342 in: Arratia, G. (ed.), Contributions of

Southern South America to Vertebrate Paleontology. Miinchner

Geowissenschaftliche Abhandlungen, Reihe A, Geologie und

Palaontologie, 30.

Dickinson, J.A. 2002. Neogene tectonism and phosphogenesis across

the SE Australian margin. Ph.D. Thesis, University of Melbourne:

Melbourne. 229 pp.

Dickinson, J.A., Wallace, M.W., Holdgate, G.R., Gallagher, S.J., and

Thomas, L. 2002. Origin and timing of the Miocene-Pliocene

unconformity in south-east Australia. Journal of Sedimentary

Research 72: 288-303.

Domning, D.P. 1996. Bibliography and index of the Sirenia and

Desmostylia. Smithsonian Contributions to Paleobiology 80:

1-611.

Etheridge, R., Jr. 1878. A catalogue of Australian Fossils (Including

Tasmania and the Island of Timor ) Strati graphic ally and

Zoologically Arranged. Cambridge University Press: Cambridge.

232 pp.

Exon, N.F., Marshall, J.F., McCorkle, D.C., Alcock, M., Chaproniere,

G.C.H., Connell, R., Dutton, S.J., Elmes, M., Findlay,

C., Robertson, L., Rollet, N., Samson, C., Shafik, S., and Whitmore,

G.P. 1995. AGSO Cruise 147 report: Tasman Rises geological

sampling cruise of Rig Seismic: stratigraphy, tectonic history and

palaeoclimate of the offshore Tasmanian region. Australian

Geological Survey Organisation Record 1995/56.

Exon, N.F., Moore, A.M.G., and Hill, P.J. 1997. Geological framework

of the South Tasman Rise, south of Tasmania, and its sedimentary

basins. Australian Journal of Earth Sciences 44: 561-577.

Finlayson, H.H. 1938. On the occurrence of a fossil penguin in

Miocene beds in South Australia. Transactions of the Royal Society

of South Australia 62: 14—17.

Fitzgerald, E.M.G. 2003. Oligocene Cetacea (Mammalia) from the

southern margin of Australia. Geological Society of Australia

Abstracts 70: 17.

Fitzgerald, E.M.G. 2004. The fossil record of cetaceans (Mammalia)

on the Australian continent. Fossil Collector Bulletin 72: 5-32.

Flynn, T.T. 1920. Squalodont remains from the Tertiary strata of

Tasmania. Nature 106: 406-407.

Flynn, T.T. 1923. A whale of bygone days. Australian Museum

Magazine 1: 266-272.

Flynn, T.T. 1932. A new species of fossil cetacean from Tasmania.

Geological Magazine 69: 327-329.

Flynn, T.T. 1948. Description of Prosqualodon davidi Flynn, a fossil

cetacean from Tasmania. Transactions of the Zoological Society of

London 26: 153-197.

Fordyce, R.E. 1978 (1979). A review of austral archaeocete whales

[Abstract]. New Zealand Geological Society Miscellaneous

Publication 23A: [unpaginated].

Fordyce, R.E. 1982a. A review of Australian fossil Cetacea. Memoirs

of the National Museum of Victoria 43: 43-58.

Fordyce, R.E. 1982b. The Australasian marine vertebrate record

and its climatic and geographic implications. Pp. 595-627 in:

Rich, P.V., and Thompson, E.M. (eds), The Fossil Vertebrate

Record of Australasia. Monash University Offset Printing Unit:

Clayton.

Fordyce, R.E. 1983. Rhabdosteid dolphins (Mammalia: Cetacea) from

the Middle Miocene, Lake Frome area, South Australia. Alcheringa

7: 27-40.

Fordyce, E. 1984. Evolution and zoogeography of cetaceans in

Australia. Pp. 929-948 in: Archer, M., and Clayton, G. (eds),

Vertebrate Zoogeography and Evolution and Zoogeography in

Australasia. Hesperian Press: Perth.

Fordyce, R.E. 1988. Taxonomic status of Victorian fossil whales

assigned to the genus Cetotolites McCoy, 1879. Memoirs of the

Museum of Victoria 49: 59-65.

Fordyce, R.E. 1991. The Australasian marine vertebrate record and its

climatic and geographic implications. Pp. 1165-1190 in: Vickers-

Rich, P., Monaghan, J.M., Baird, R.F., and Rich, T.H. (eds),

Vertebrate Palaeontology of Australasia. Pioneer Design Studio in

cooperation with the Monash University Publications Committee:

Melbourne.

Fordyce, R.E. 2002a. Oligocene archaeocetes and toothed mysticetes:

Cetacea from times of transition. Pp. 16-17 in: Fordyce, R.E., and

Walker, M. (eds), Abstracts, Third Conference on Secondary

Adaptation to Life in Water. Geological Society of New Zealand

Miscellaneous Publication 114A.

Fordyce, R.E. 2002b. Fossil sites. Pp. 471-482 in: Perrin, W.F.,

Wiirsig, B., and Thewissen, J.G.M. (eds), Encyclopedia of Marine

Mammals. Academic Press: San Diego.

Fordyce, R.E. 2003. Cetacean evolution and Eocene-Oligocene oceans

revisited. Pp. 154—170 in: Prothero, D.R., Ivany, L.C., and Nesbitt,

E.A. (eds), From Greenhouse to Icehouse: the Marine

Eocene-Oligocene Transition. Columbia University Press: New

York.

Fordyce, R.E., and Flannery, T.F. 1983. Fossil phocid seals from the

late Tertiary of Victoria. Proceedings of the Royal Society of

Victoria 95: 99-100.

Fordyce, R.E., and Muizon, C. de. 2001. Evolutionary history of

cetaceans: a review. Pp. 169-233 in: Mazin, J.M., and Buffrenil, V.

de. (eds), Secondary Adaptation of Tetrapods to Life in Water.

Verlag Dr. Friedrich Pfeil: Miinchen.

Geisler, J.H., and Luo, Z. 1996. The petrosal and in ner ear of

Herpetocetus sp. (Mammalia: Cetacea) and their implications for

the phylogeny and hearing of archaic mysticetes. Journal of

Paleontology 70: 1045-1066.

Geisler, J.H., and Sanders, A.E. 2003. Morphological evidence for the

phylogeny of Cetacea. Journal of Mammalian Evolution 10:

23-129.

Gill, E.D. 1957. The stratigraphical occurrence and palaeoecology of

some Australian Tertiary marsupials. Memoirs of the National

Museum of Victoria 21: 135-203.

Glaessner, M.F. 1947. A fossil beaked whale from Lakes Entrance,

Victoria. Proceedings of the Royal Society of Victoria 58: 25-34.

202

E. M. G. Fitzgerald

Glaessner, M.F. 1955. Pelagic fossils ( Aturia , penguins, whales) from

the Tertiary of South Australia. Records of the South Australian

Museum IT. 353-372.

Glover, J.E. 1955. Petrographical study of rock samples from the

coastal section between Torquay and Airey’s Inlet, Victoria.

Proceedings of the Royal Society of Victoria 67: 149-164.

Gottfried, M.D., and Fordyce, R.E. 2001. An associated specimen

of Carcharodon angustidens (Chondrichthyes, Lamnidae) from

the Late Oligocene of New Zealand, with comments on

Carcharodon interrelationships. Journal of Vertebrate Paleontology

21: 730-739.

Hall, T.S. 1911. On the systematic position of the species of Squalodon

and Zeuglodon described from Australia and New Zealand.

Proceedings of the Royal Society of Victoria 23: 257-265.

Harris, W.K. 1971. Tertiary stratigraphic palynology, Otway Basin. Pp.

67-87 in: Wopfner, H., and Douglas, J.G. (eds), The Otway Basin

of Southeastern Australia. Special Bulletin, Geological Surveys of

South Australia and Victoria.

Heyning, J.E. 1989. Comparative facial anatomy of beaked whales

(Ziphiidae) and a systematic revision among the families of extant

Odontoceti. Natural History Museum of Los Angeles County,

Contributions in Science 405: 1-64.

Heyning, J.E. 1997. Sperm whale phylogeny revisited: analysis of the

morphological evidence. Marine Mammal Science 13: 596-613.

Holdgate, G.R., and Gallagher, S.J. 2003. Tertiary: a period of transi-

tion to marine basin environments. Pp. 289-335 in: Birch, W.D.

(ed.), Geology of Victoria. Geological Society of Australia Special

Publication 23, Geological Society of Australia (Victoria Division):

Melbourne.

Howchin, W. 1919. [Tympanic bone of Balaena from the Pliocene of

South Australia.] Transactions and Proceedings of the Royal

Society of South Australia 43: 430.

Jenkins, R.F. 1974. A new giant penguin from the Eocene of Australia.

Palaeontology 17: 291-310.

Jenkins, R.F. 1990. Anthropomis nordenskjoeldi Nordenskjoeld’s giant

penguin. Pp. 183-187 in: Rich, P.V. and van Tets, G.F. (eds),

Kadimakara: Extinct Vertebrates of Australia. Princeton University

Press: Princeton.

Kelly, J.C., Webb, J.A., and Maas, R. 2001. Isotopic constraints on the

genesis and age of autochthonous glaucony in the Oligo-Miocene

Torquay Group, south-eastern Australia. Sedimentology 48:

325-338.

Kemp, N.R. 1970. Studies on the dentition of Australian Tertiary and

Recent sharks. M.Sc. Thesis, University of Melbourne: Melbourne.

354 pp.

Kemp, N.R. 1978. Detailed comparisons of the dentitions of extant

hexanchid sharks and Tertiary hexanchid teeth from South Australia

and Victoria, Australia (Selachii: Hexanchidae). Memoirs of the

National Museum of Victoria 39: 61-83.

Kemp, N.R. 1982. Chondrichthyans in the Tertiary of Australia. Pp.

88-117 in: Rich, P.V., and Thompson, E.M. (eds), The Fossil

Vertebrate Record of Australasia. Monash University Offset

Printing Unit: Clayton.

Kemp, N.R. 1991. Chondrichthyans in the Cretaceous and Tertiary of

Australia. Pp. 497-568 in: Vickers-Rich, R, Monaghan, J.M.,

Baird, R.F., and Rich, T.H. (eds), Vertebrate Palaeontology of

Australasia. Pioneer Design Studio in cooperation with the Monash

University Publications Committee: Melbourne.

Kohler, R., and Fordyce, R.E. 1997. An archaeocete whale (Cetacea:

Archaeoceti) from the Eocene Waihao Greensand, New Zealand.

Journal of Vertebrate Paleontology 17: 574—583.

Li, Q., Davies, P.J., and McGowran, B. 1999. Foraminiferal sequence

biostratigraphy of the Oligo-Miocene Janjukian strata from

Torquay, southeastern Australia. Australian Journal of Earth

Sciences 46: 261-273.

Li, Q., McGowran, B., and White, M.R. 2000. Sequence and biofacies

packages in the mid-Cenozoic Gambier Limestone, South

Australia: reappraisal of foraminiferal evidence. Australian Journal

of Earth Sciences 47: 955-970.

Lukasik, J.J., and James, N.P. 1998. Lithostratigraphic revision and

correlation of the Oligo-Miocene Murray Supergroup, western

Murray Basin, South Australia. Australian Journal of Earth

Sciences 45: 889-902.

Lydekker, R. 1899. On the skull of a shark-toothed dolphin from

Patagonia. Proceedings of the Zoological Society of London 1899:

919-922.

Mahoney, J.A., and Ride, W.D.L. 1975. Index to the genera and species

of fossil Mammalia described from Australia and New Guinea

between 1838 and 1968. Western Australian Museum Special

Publication 6: 1-250.

Mallett, C.W. 1977. Studies in Victorian Tertiary Foraminifera:

Neogene planktonic faunas. Ph.D. Thesis, University of

Melbourne: Melbourne. 381 pp.

McCoy, F. 1866. Notes sur la zoologie et la palaeontologie de Victoria,

par Frederick M’Coy. Traduit de 1’ anglais par E. Lissignol.

Masterman: Melbourne. 35 pp.

McCoy, F. 1867a. On the occurrence of the genus Squalodon in the

Tertiary strata of Victoria. Geological Magazine 4: 145.

McCoy, F. 1867b. On the recent zoology and palaeontology of

Victoria. Annals and Magazine of Natural History 20: 175-202.

McCoy, F. 1875. Squalodon wilkinsoni (McCoy). Prodromus of the

palaeontology of Victoria; or, figures and descriptions of the

Victorian organic remains. Decade 2: 7, 8. Geological Survey of

Victoria: Melbourne.

McCoy, F. 1879a. Cetotolites. Prodromus of the palaeontology of

Victoria; or, figures and descriptions of the Victorian organic

remains. Decade 6: 13-17. Geological Survey of Victoria:

Melbourne.

McCoy, F. 1879b. Physetodon baileyi (McCoy). Prodromus of the

palaeontology of Victoria; or, figures and descriptions of the

Victorian organic remains. Decade 6: 19-21. Geological Survey of

Victoria: Melbourne.

McHaffie, I.W., and Inan, K. 1988. Limestone. Pp. 572-579 in:

Douglas, J.G., and Ferguson, J.A. (eds), Geology of Victoria.

Victorian Division, Geological Society of Australia Incorporated:

Melbourne.

McKenna, M.C., and Bell, S.K. 1997. Classification of mammals

above the species level. Columbia University Press: New York. 631

pp.

Messenger, S.L., and McGuire, J.A. 1998. Morphology, mole-

cules, and the phylogenetics of cetaceans. Systematic Biology 47:

90-124.

Milinkovitch, M.C., Ortf, G., and Meyer, A. 1993. Revised phylogeny

of whales suggested by mitochondrial ribosomal DNA sequences.

Nature 361: 346-348.

Mitchell, E.D. 1989. A new cetacean from the Late Eocene La Meseta

Formation, Seymour Island, Antarctic Peninsula. Canadian Journal

of Fisheries and Aquatic Sciences 46: 2219-2235.

Muizon, C. de. 2002. River dolphins, evolutionary history. Pp.

1043-1049 in: Perrin, W.F., Wiirsig, B., and Thewissen, J.G.M.

(eds), Encyclopedia of Marine Mammals. Academic Press: San

Diego.

Nicolaides, S., and Wallace, M.W. 1997. Submarine cementation and

subaerial exposure in Oligo-Miocene temperate carbonates,

Torquay Basin, Australia. Journal of Sedimentary Research 67:

397-410.

Tertiary fossil Cetacea localities in Australia

203

Pledge, N.S. 1967. Fossil elasmobranch teeth of South Australia and

their stratigraphic distribution. Transactions of the Royal Society of

South Australia 91: 135-160.

Pledge, N.S. 1985. An Early Pliocene shark tooth assemblage in South

Australia. Special Publication, South Australian Department of

Mines and Energy 5: 287-299.

Pledge, N.S. 1992. First record of fossil sirenians in southern Australia.

Fossil Collector Bulletin 37: 6.

Pledge, N.S. 1994. Cetacean fossils from the Lower Oligocene of

South Australia. Records of the South Australian Museum 27:

117-123.

Pledge, N.S., and Rothausen, K. 1977. Metasqualodon harwoodi

(Sanger, 1881)-a redescription. Records of the South Australian

Museum 17: 285-297.

Pritchard, G.B. 1939. On the discovery of a fossil whale in the older

Tertiaries of Torquay, Victoria. Victorian Naturalist 55: 151-159.

Purdy, R.W., Schneider, V.P., Applegate, S.P., McLellan, J.H., Meyer,

R.L., and Slaughter, B.H. 2001. The Neogene sharks, rays, and

bony fishes from Lee Creek Mine, Aurora, North Carolina. Pp.

71-202 in: Ray, C.E., and Bohaska, DJ. (eds), Geology and

Paleontology of the Lee Creek Mine, North Carolina, III.

Smithsonian Contributions to Paleobiology 90.

Quilty, P. 1980. New rotaliid foraminiferids from the Oligo-Miocene

of Tasmania. Alcheringa 4: 299-3 1 1 .

Raggatt, H.G., and Crespin, I. 1955. Stratigraphy of Tertiary rocks

between Torquay and Eastern View, Victoria. Proceedings of the

Royal Society of Victoria 67: 75-142.

Rich, P.V. 1975. Antarctic dispersal routes, wandering continents, and

the origin of Australia’s non-passeriform avifauna. Memoirs of the

National Museum of Victoria 36: 63-125.

Rich, T.H., Archer, M., Hand, S.J., Godthelp, H., Muirhead, J., Pledge,

N.S., Flannery, T.F., Woodburne, M.O., Case, J.A., Tedford, R.H.,

Turnbull, W.D., Lundelius, E.L., Jr., Rich, L.S.V., Whitelaw, M.J.,

Kemp, A., and Rich, P.V. 1991. Appendix 1. Australian Mesozoic

and Tertiary terrestrial mammal localities. Pp. 1005-1058 in:

Vickers-Rich, P„ Monaghan, J.M., Baird, R.F., and Rich, T.H. (eds),

Vertebrate Palaeontology of Australasia. Pioneer Design Studio in

cooperation with the Monash University Publications Committee:

Melbourne.

Rich, T.H., Darragh, T.A., and Vickers-Rich, P. 2003. The strange case

of the wandering fossil. Bulletin of the American Museum of

Natural History 13: 556-567.

Romer, A.S. 1966. Vertebrate Paleontology. Third Edition. University

of Chicago Press: Chicago. 468 pp.

Sanger, E.B. 1881. On a molar tooth of Zeuglodon from the Tertiary

beds on the Murray River near Wellington, S.A. Proceedings of the

Linnean Society of New South Wales 5: 298-300.

Siesser, W.G. 1979. Oligocene-Miocene calcareous nannofossils from

the Torquay Basin, Victoria, Australia. Alcheringa 3: 159-170.

Simpson, G.G. 1957. Australian fossil penguins, with remarks on pen-

guin evolution and distribution. Records of the South Australian

Museum 13: 51-70.

Simpson, G.G. 1959. A new fossil penguin from Australia.

Proceedings of the Royal Society of Victoria 71: 113-119.

Simpson, G.G. 1965. New record of a fossil penguin in Australia.

Proceedings of the Royal Society of Victoria 79: 91-93.

Simpson, G.G. 1970. Miocene penguins from Victoria, Australia, and

Chubut, Argentina. Memoirs of the National Museum of Victoria

31: 17-23.

Singleton, F.A. 1941. The Tertiary geology of Australia. Proceedings

of the Royal Society of Victoria 53: 1-125.

Singleton, O.P., McDougall, I., and Mallett, C.W. 1976. The

Pliocene-Pleistocene boundary in southeastern Australia. Journal

of the Geological Society of Australia 23: 299-311.

Stinton, F.C. 1958. Fish otoliths from the Tertiary strata of Victoria,

Australia. Proceedings of the Royal Society of Victoria 70: 81-93.

Stinton, F.C. 1963. Further studies of the Tertiary otoliths of Victoria,

Australia. Proceedings of the Royal Society of Victoria 76: 13-22.

Stirton, R.A. 1967. New species of Zygomaturus and additional obser-

vations on Meniscolophus, Pliocene Palankarinna fauna, South

Australia. Australian Bureau of Mineral Resources, Geology and

Geophysics Bulletin 85: 129-147.

Sutherland, F.L., and Kershaw, R.C. 1971. The Cainozoic geology of

Flinders Island, Bass Strait. Papers and Proceedings of the Royal

Society of Tasmania 105: 151-175.

Tate, R. 1892. [Zeuglodon tooth from Tasmania.] Transactions of the

Royal Society of South Australia 15: 265.

Tedford, R.H., Archer, M., Bartholomai, A., Plane, M., Pledge, N.S.,

Rich, T„ Rich, R, and Wells, R.T. 1977. The discovery of Miocene

vertebrates, Lake Frome area, South Australia. BMR Journal of

Geology and Geophysics 2: 53-57.

Tickell, S.J., Edwards, J., and Abele, C. 1992. Port Campbell

Embayment 1:100 000 map geological report. Geological Survey of

Victoria Report 95.

Turnbull, W.D., Lundelius, E.L., and McDougall, I. 1965. A potassi-

um-argon dated Pliocene marsupial fauna from Victoria, Australia.

Nature 206: 816.

Uhen, M.D. 1998. Middle to Late Eocene basilosaurines, and dorudon-

tines. Pp. 29-61 in: Thewissen, J.G.M. (ed). The Emergence of

Whales. Plenum Press: New York.

Vickers-Rich, P. 1991. The Mesozoic and Tertiary history of birds on

the Australian plate. Pp. 721-808 in: Vickers-Rich, P, Monaghan,

J.M., Baird, R.F., and Rich, T.H. (eds), Vertebrate Palaeontology of

Australasia. Pioneer Design Studio in cooperation with the Monash

University Publications Committee: Melbourne.

Vickers-Rich, R, and Rich, T.H. 1993. Wildlife of Gondwana. Reed:

Chats wood. 276 pp.

Webb, J.A. (ed). 1995. Cool-water carbonates of the north-eastern

Otway Basin, southeastern Australia. Australasian

Sedimentologists Group Field Guide Series No. 6. Geological

Society of Australia: Sydney. 56 pp.

Wilkins, R.W.T. 1963. Relationships between the Mitchellian,

Cheltenhamian and Kalimnan Stages in the Australian Tertiary.

Proceedings of the Royal Society of Victoria 76: 39-59.

Wilkinson, H.E. 1969. Description of an Upper Miocene albatross

from Beaumaris, Victoria, Australia, and a review of fossil

Diomedeidae. Memoirs of the National Museum of Victoria 29:

41-51.

Woodburne, M.O. 1969. A lower mandible of Zygomaturus gilli from

the Sandringham Sands, Beaumaris, Victoria, Australia. Memoirs of

the National Museum of Victoria 29: 29-39.

Woodburne, M.O., MacFadden, B.J., Case, J.A., Springer, M.S.,

Pledge, N.S., Power, J.D., Woodburne, J.M., and Springer, K.B.

1993. Land mammal biostratigraphy and magnetostratigraphy of

the Etadunna Formation (Late Oligocene) of South Australia.

Journal of Vertebrate Paleontology 13: 483-515.

Zhou, K. 1982. Classification and phylogeny of the Superfamily

Platanistoidea, with notes on evidence of the monophyly of the

Cetacea. Scientific Reports of the Whales Research Institute 34:

93-108.

204

E. M. G. Fitzgerald

Table 1 . Taxonomy of vertebrates from Tertiary fossil cetacean-bearing localities. * indicates provisional classification.

Class

Order

Suborder

Family

Genus and species

Chondrichthyes

Galeomorpha

Heterodontoidea

Heterodontidae

Heterodontus cainozoicus Chapman and Pritchard, 1904

Orectoloboidea

Orectolobidae

Orectolobus gippslandicus Chapman and Cudmore, 1924

Lamnoidea

Odontaspididae

Carcharias elegans Agassiz, 1843

Carcharias taurus Rafinesque, 1810

Carcharias macrotus Agassiz, 1843

Odontaspis acutissima Agassiz, 1843

Odontaspis incurva Davis, 1888

Lamnidae

Carcharodon angustidens Agassiz, 1 843

Carcharodon megalodon Agassiz, 1835

Carcharodon carcharias Linnaeus, 1758

Isurus hastalis Agassiz, 1838

Isurus retroflexus Agassiz, 1843

Isurus desori Agassiz, 1 843

Isurus escheri Agassiz, 1843

Isurus planus Agassiz, 1843

Isurus oxyrinchus Rafinesque, 1810

Isurus paucus Guitart-Monday, 1966

Parotodus benedenii Le Hon, 1871

Carcharoides totuserratus Ameghino, 1901

Lamna cattica Philippi, 1846

Carcharhinoidea

Parascylidae

Megascyliorhinus sp.

Triakidae

Mustelus sp.

Galeorhinus australis Macleay, 1881

Carcharhinidae

Carcharhinus brachyurus Gunther, 1870

Galeocerdo aduncus Agassiz, 1843

Sphyrnidae

Sphyma sp.

Hexanchoidea

Hexanchidae

Hexanchus agassizi Cappetta, 1976

Notorynchus primigenius Agassiz, 1843

Notorynchus cepedianus Peron, 1807

Squalomorpha

Pristiophoroidea

Pristiophoridae

Pristiophorus lanceolatus Davis, 1888

Batoidea

Myliobatoidea

Myliobatidae

Myliobatis sp.

Dasyatidae

Dasyatis sp.

Chimaeriformes

Chimaeroidei

Chimaeridae

Edaphodon sweeti Chapman and Pritchard, 1907

Edaphodon mirabilis Chapman and Cudmore, 1924

Ischyodus dolloi Leriche, 1902

Ischyodus mortoni Chapman and Pritchard, 1907

Osteichthyes

Elopiformes

Elopoidei

Megalopidae

Megalops lissa Stinton, 1958

Albuloidei

Pterothrissidae

Pterothrissus pervetustus Stinton, 1958

Anguilliformes

Heterenchelyidae

Heterenchelys regularis Stinton, 1958

Muraenesocidae

Muraenesox obrutus Stinton, 1958

Congridae

Astroconger rostratus Stinton, 1958

Urconger rectus Frost, 1928

S al m on i formes

Argentinoidei

Argentinidae

Hypomesus glaber Stinton, 1963

Gadiformes

Gadoidei

Merluccidae

Merluccius fimbriatus Stinton, 1958

Gadidae

Gadus refertus Stinton, 1958

Ophidiiformes

Ophidiidae

Ophidion granosum Stinton, 1958

Beryciformes

Berycoidei

Trachichthyidae

Trachichthodes salebrosus Stinton, 1958

Monocentridae

Cleidopus carvernosus Stinton, 1958

Scorpaeniformes

Scorpaenoidei

Scorpaenidae

Sebastodes fissicostatus Stinton, 1963

Perciformes

Percoidei

Sillaginidae

Sillago pliocaenica Stinton, 1952

Lactariidae

Lactarius tumulatus Stinton, 1958

Labroidei

Labridae

Labrodon sp.

Scombroidei

Xiphiidae

Coelorhynchus elevatus Stinton, 1956

Xiphias sp.

Tetraodontiformes

Balistoidei

Monacanthidae

Tetraodontoidei

Diodontidae

Diodon formosus Chapman and Pritchard, 1907

Dipnoi

N eoceratodontidae

Neoceratodus djelleh Kemp, 1982

Neoceratodus eyrensis White, 1925

Neoceratodus gregoryi White, 1925

Neoceratodus forsteri Krefft, 1870

Neoceratodus nargun Kemp, 1983

Tertiary fossil Cetacea localities in Australia

205

Table 1. Continued.

Class

Order

Suborder

Family

Genus and species

Reptilia

Chelonia

Pleurodira

Chelidae

Emydura sp.

Cryptodira

Meiolaniidae

Trionychidae*

Crocodylia

Aves

Columbiformes

Passeriformes

Columbidae

Gruiformes

Ralli

Rallidae

Podicipediformes

Podicipedidae

Charadriiformes

Burhinidae

Phoenicopteridae

Anseriformes

Anatidae

Pelecaniformes

Pelecani

Pelecanidae

Pelecanus tirarensis Miller, 1966

Sulae

Phalacrocoracidae

Procellariiforme s

Diomedeidae

Diomedea thyridata Wilkinson, 1969

Sphenisciformes

Spheniscidae

Pseudaptenodytes macraei Simpson, 1970

Pseudaptenodytes minor Simpson, 1970

Mammalia

Monotremata

Ornithorhynchidae

Obdurodon insignis Woodbume and Tedford, 1975

Dasyuromorphia

Dasyuridae

Dasylurinja kokuminola Archer, 1982

Diprotodontia

Wynyardiidae

Wynyardia bassiana Spencer, 1901

Muramura sp.

Palorchestidae

Diprotodontidae

Raemeotherium yatkolai Rich et al., 1978

Zygomaturus gilli Stirton, 1967

Kolopsis torus Woodbume, 1967

Ilariidae

Vombatidae

Ilaria illumidens Tedford and Woodbume, 1987

Phascolarctidae

Madakoala devisi Woodbume et al., 1987

Madakoala wellsi Woodbume et al., 1987

Phascolarctos marts Pledge, 1987

Potoroidae

Macropodidae

Kurrabi sp.

Dorcopsis sp.

Petauridae

Pseudocheiridae

Pildra antiquus Woodbume et al., 1987

Pildra secundus Woodbume et al., 1987

Ektopodontidae

Chunia illuminata Woodbume and Clemens, 1986

Pilkipildridae

Pilkipildra handae Archer et al., 1987

Djilgaringa thompsonae Archer et al., 1987

Miralinidae

Miralina minor Woodbume et al., 1987

Carnivora

Phocidae

Sirenia

Dugongidae

Dugong* sp.

Cetacea

Archaeoceti

Basilosauridae*

Metasqualodon harwoodi* Sanger, 1881

“ Squalodon ” gambierensis Glaessner, 1955

Mysticeti

Mammalodontidae

Aetiocetidae*

Mammalodon colliveri Pritchard, 1939

“Cetotheriidae”

Pelocetus sp.

Parietobalaena sp.

Balaenidae

Balaena sp.

Balaenopteridae

Balaenoptera sp.

Megaptera sp.

Odontoceti

Prosqualodontidae

Prosqualodon sp. McCoy, 1866

Prosqualodon davidis Flynn, 1923

Squalodontidae

Eurhinodelphinidae

Physeteridae

Physetodon bailey i McCoy, 1879b

S captodon lodderi Chapman, 1918

Scaldicetus lodgei Chapman, 1917b

Scaldicetus macgeei Chapman, 1912

Scaldicetus sp.

Physeter sp.

Ziphiidae

Mesoplodon longirostris Cuvier, 1 823

Ziphius sp.

Delphinidae

206

E. M. G. Fitzgerald

Table 2. Classification of formally described Australian fossil Cetacea.

Taxon

Previous classification

Suborder Archaeoceti

Family Basilosauridae

“ Squalodon ” gambierensis Glaessner, 1955

Suborder incerta sedis

Metasqualodon harwoodi Sanger, 1881

Cetotolites leggei McCoy, 1879a (nomen dubium; Fordyce, 1988)

Cetotolites nelsoni McCoy, 1879a (nomen dubium; Fordyce, 1988)

Cetotolites pricei McCoy, 1879a (nomen dubium; Fordyce, 1988)

Cetotolites rugosa McCoy, 1879a (nomen dubium; Fordyce, 1988)

Suborder Mysticeti

Family Mammalodontidae

Mammalodon colliveri Pritchard, 1939

Suborder Odontoceti

Family Physeteridae

Physetodon baileyi McCoy, 1879b

Scaptodon lodderi Chapman, 1918

Scaldicetus lodgei Chapman, 1917b

Scaldicetus macgeei Chapman, 1912

Family Prosqualodontidae

Prosqualodon davidis Flynn, 1923

Prosqualodon sp. McCoy, 1866

Family Delphinidae

Gen. et sp. indet. Chapman, 1917b

Odontoceti: Squalodontidae, Glaessner, 1955; Odontoceti:

Squalodontidae, Pledge and Rothausen, 1977; Archaeoceti:

Basilosauridae, Fordyce 2002a; Archaeoceti: Basilosauridae, Fitzgerald

2004

Archaeoceti: Basilosauridae, Sanger, 1881; Odontoceti: Squalodontidae

Hall, 1911; Odontoceti: Physeteridae, Chapman, 1929; Odontoceti:

Squalodontidae, Pledge and Rothausen, 1977; Odontoceti:

?Squalodontidae, Fordyce, 1991; Autoceta: incertae sedis, Fitzgerald

2004

Odontoceti: Ziphiidae, McCoy, 1879a

Archaeoceti: Basilosauridae, Anonymous, 1939; Cetacea: incertae

sedis,

Camp et al., 1942; Archaeoceti: Basilosauridae, Romer, 1966;

Odontoceti: Squalodontidae, Pledge and Rothausen, 1977; Archaeoceti:

Dorudontidae (=Basilosauridae: Domdontinae of Barnes and Mitchell,

1978; Uhen, 1998), Fordyce, 1979; Mysticeti: incertae sedis, Fordyce,

1982a; Mysticeti: Mammalodontidae, Mitchell, 1989; Mysticeti: incerta

sedis, Fordyce, 1991

Odontoceti: Physeteridae, McCoy, 1879b; Odontoceti: Physeteridae,

Fordyce, 1982a

Odontoceti: Physeteridae, Chapman, 1918; Odontoceti: Physeteridae,

Fordyce, 1982a

Odontoceti: Physeteridae, Chapman, 1917b; Odontoceti: Physeteridae,

Fordyce, 1982a

Odontoceti: Physeteridae, Chapman, 1912; Odontoceti: Physeteridae,

Fordyce, 1982a

Odontoceti: Squalodontidae, Flynn, 1920,, 1923,, 1932,, 1948;

Odontoceti: Prosqualodontidae: Prosqualodon australis, Cozzuol,

1996; Odontoceti: Prosqualodontidae, Muizon 2002

Odontoceti: Squalodontidae: Phocodon wilkinsoni, McCoy, 1866;

Odontoceti: Squalodontidae: Squalodon wilkinsoni, McCoy, 1867a„

1875; Odontoceti: Squalodontidae: Parasqualodon wilkinsoni, Hall,

1911; Odontoceti: Squalodontidae: Prosqualodon sp., Pledge and

Rothausen, 1977; Odontoceti: Squalodontidae: IP ro squalodon davidis,

Fordyce, 1982a; Odontoceti: Squalodontidae: Prosqualodon davidis,

Fitzgerald 2004

Odontoceti: Delphinidae: Steno cudmorei. Chapman, 1917b;

Odontoceti: Delphinidae: Gen. et sp. indet., Fordyce, 1982a

Tertiary fossil Cetacea localities in Australia

207

Table 3. Summary of Australian Tertiary aquatic mammal taxa, localities, stratigraphy, ages, and references. Abbreviations: SA, South

Australia; Tas., Tasmania; Vic., Victoria; Fmn, Formation; Lst, Limestone; Sst, Sandstone; M, Miocene; O, Oligocene; P, Pliocene. * indicates

provisional classification.

Taxa

Locality

Stratigraphy

Age

References

CETACEA

ARCHAEOCETI

Basilosauridae*

“Squalodon” gambierensis

Mount Gambier; SA

Gambier Lst

Late 0

Glaessner, 1955; Pledge and

Rothausen, 1977; Fordyce, 2002a

MYSTICETI

Mammalodontidae

Mamrnalodon colliveri

Bird Rock; Vic.

Jan Juc Marl

Late 0

Pritchard, 1939; Fordyce, 1982a, 1984;

Mitchell, 1989; Fitzgerald, 2004

Mammalodon sp. indet.

Bird Rock, Waurn Ponds

Jan Juc Marl; Waum Ponds

Late 0

Fordyce, 1982a, 1988

Quarry; Vic.

Lst

Mamrnalodon sp. nov. 1

Bird Rock; Vic.

Jan Juc Marl

Late 0

Gen. et sp. indet. 1

Bells Beach; Vic.

Point Addis Lst

Late 0

Pledge, 1994

Gen. et sp. indet.

Point Addis, Waurn Ponds

Point Addis Lst;

Late 0

Fordyce, 1982a, 1988

Quarry; Vic.

Waurn Ponds Lst

cf. Mammalodontidae

?Aetiocetidae*

Moorabool River; Vic.

Maude Fmn

Late O-Early M

Gen. et sp. nov.

Cetotheriidae

Port Willunga; S A

Port Willunga Fmn

Early 0

Gen. et sp. indet.

Forsyth’s Bank to Fossil

Grange Burn Fmn; Port

Early P; Middle

Bearlin, 1987, 1988; Fitzgerald, 2004

Rock Stack, Gibson’s Steps,

Campbell Lst;

M; Late 0;

Waurn Ponds Quarry

Waurn Ponds Lst,

Middle M; Late

Batesford Quarry,

Batesford Lst; Black Rock

M-Early P;

Beaumaris, Newmerella;

Sst; Gippsland Lst;

Early M; Early

Vic.; MacBean’s Pound; SA

Fmn; Mannum Fmn

Pelocetus sp.

Arch Site, Grange Bum; Vic Bochara Lst

Middle M

Bearlin, 1987, 1988

cf. Parietobalaena sp.

Balaenidae

Murbko; SA

Glenforslan Fmn

Middle M

Bearlin, 1987, 1988

Gen. et sp. indet.

Dutton Way (Portland),

Whalers Bluff Fmn;

P; Middle-Late

Howchin, 1919; Fordyce, 1982a, 1984;

Clifton Bank, Forsyth’s

Muddy Creek Marl;

M; Early P;

Bearlin, 1987

Bank to Fossil Rock Stack

Grange Burn Fmn;

Late M-Early

Beaumaris, Trident Arm;

Black Rock Sst;

P; Early P;

Vic.; Abattoirs Bore; SA

Jemmy s Point Fmn;

Dry Creek Sand

Late P

cf. Balaena sp.

Balaenopteridae

Beaumaris; Vic.

Black Rock Sst

Late M-Early P

Gill, 1957; Fordyce, 1982a

Gen. et sp. indet.

Curdie; Vic.; Winkie; SA

Port Campbell Lst;

Middle M; ?

Bearlin, 1987, 1988

?Bookpurnong Fmn

M-P;

Balaenoptera sp.

Dutton Way (Portland),

Whalers Bluff Fmn;

P; Early P;

Fordyce, 1982a; Bearlin, 1987

Forsyth’s Bank to Fossil

Grange Burn Fmn;

Late M-Early

Rock Stack, Spring Creek

Unnamed unit;

P; Late

Beaumaris; Vic.

Black Rock Sst

M-Early P

cf. Balaenoptera sp.

Cameron Inlet; Tas.

Cameron Inlet Fmn

Fordyce, 1982a

Megaptera sp.

Dutton Way (Portland), .

Whalers Bluff Fmn;

P; Late M-

Bearlin, 1987

Beaumaris; Vic

Black Rock Sst;

Early P

Megaptera sp. nov. 1

North Arm; Vic.

Jemmy s Point Fmn

Early P

Bearlin, 1987, 1988; Fitzgerald, 2004

cf. Megaptera sp.

Family nov.

Cameron Inlet; Tas.

Cameron Inlet Fmn

Gen. et sp. nov. 1

Deadman’s Gully; Vic.

Jan Juc Marl

Late 0

Fitzgerald, 2004

Gen. et sp. nov. 2

ODONTOCETI

Bells Headland; Vic.

Point Addis Lst

Late 0

Kohler and Fordyce, 1997

Physeteridae

Gen. et sp. indet.

Dutton Way (Portland),

Whalers Bluff Fmn;

P; Early P;

Fordyce, 1982a, 1984

Forsyth’s Bank to Fossil

Grange Bum Fmn;

Middle M;

Rock Stack, Batesford

Batesford Lst;

Late M-Early

Quarry, Beaumaris; Vic.

Black Rock Sst

Physeter sp.

Cameron Inlet; Tas.

Cameron Inlet Fmn

Fitzgerald, 2004

208

E. M. G. Fitzgerald

Table 3. continued.

Taxa

Locality

Stratigraphy

Age

References

cf. Physeter sp.

Clifton Bank, Forsyth’s

Bank to Fossil Rock Stack;

Vic.

Muddy Creek Marl;

Grange Burn Fmn

Middle-Late

M; Early P

Fordyce, 1982a; Fitzgerald, 2004

cf. Scaldicetus sp.

Forsyth’s Bank to Fossil

Rock Stack, Hopkins River

Beaumaris; Vic.

Grange Burn Fmn;

Port Campbell Lst;

Black Rock Sst

Early P;

Middle-Late

M; Late M-

Early P

Fitzgerald, 2004

Ziphiidae

Gen. et sp. indet.

Dutton Way (Portland),

Whalers Bluff Fmn;

P; Late M-

Sutherland and Kershaw, 1971; Exon

et al.

Beaumaris; Vic.; Cameron

Inlet, Tasman Fracture

Zone, Seamount East of

South Tasman Rise,

Eastern South Tasman

Rise; Tas.

Black Rock Sst;

Cameron Inlet Fmn;

Unnamed unit

Early P; P;

Late Neogene

1995; Fitzgerald, 2004

cf. Mesoplodon sp.

Forsyth’s Bank to Fossil

Grange Burn Fmn;

Early P;

Chapman, 1917b; Fordyce, 1982a;

Rock Stack; Vic.;

Eastern Scarp; Tas.

Unnamed unit

Late Neogene

Exon et al., 1995

Mesoplodon longirostris

Jemmys Point; Vic.

Jemmys Point Fmn

Early P

Glaessner, 1947; Fordyce, 1982a

Squalodontidae*

Gen. et sp. indet.

Waurn Ponds Quarry; Vic.;

Blanchetown; SA

Waum Ponds Lst;

Mannum Fmn

Late 0;

Early M

?Gen. et sp. nov.

Batesford Quarry; Vic.

Batesford Lst

Middle M

Bearlin, 1982; Fordyce, 1982a, 1984;

Fitzgerald, 2004

Prosqualodontidae

Prosqualodon davidis

Fossil Bluff; Tas.

Fossil Bluff Sst;

Early M;

Flynn, 1920, 1923, 1932, 1948;

Mahoney and Ride, 1975

Prosqualodon sp.

Castle Cove, Bird Rock;

Calder River Lst; Jan Juc

Late 0; Late

McCoy, 1866, 1867a, 1867b, 1875;

Vic.; Fossil Bluff; Tas.

Marl; Fossil Bluff Sst

0; Early M

Tate, 1892; Hall, 1911; Flynn, 1948;

Fordyce, 1982a, 1982b, 1984;

Fitzgerald, 2003, 2004

Gen. et sp. indet.

Mount Gambier; SA

Gambier Lst

Late 0

Hall, 1911; Fordyce, 1984

Eurhinodelphinidae

Gen. et sp. nov.

Lake Namba, Lake Pinpa,

Lake Yanda; SA

Namba Fmn

Late 0-

Middle M

Tedford et al., 1977; Fordyce, 1982a,

1982b, 1983, 1984

?Gen. et sp. indet.

Bird Rock; Vic.

Jan Juc Marl

Late 0

Delphinidae

Gen. et sp. indet.

Dutton Way (Portland),

Whalers Bluff Fmn;

P; Late M

Chapman, 1917; Fordyce, 1982a;

Beaumaris; Vic.

Black Rock Sst

-Early P

Fitzgerald, 2004

?Gen. et sp. indet.

Forsyth’s Bank to Fossil

Rock Stack; Vic.; Sunlands

Pumping Station; SA;

Cameron Inlet; Tas.

Grange Bum Fmn;

Loxton Sand;

Cameron Inlet Fmn

Early P;

Early P; P

Fordyce, 1982a; Pledge, 1985

CARNIVORA

Phocidae

?Gen. et sp. indet.

Dutton Way (Portland),

Forsyth’s Bank to Fossil

Rock Stack, Beaumaris;

Vic.

Whalers Bluff Fmn;

Grange Bum Fmn;

Black Rock Sst

P; Early P;

Late M-Early

Fordyce and Flannery, 1983

SIRENIA

Dugongidae

cf. Dugong sp.

Sunlands Pumping Station;

Loxton Sand

Early P

Pledge, 1992; Domning, 1996

Memoirs of Museum Victoria 61(2): 209-216 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A new Late Eocene cassiduloid (Echinoidea) from Yorke Peninsula, South

Australia

Francis C. Holmes

15 Kenbry Road, Heathmont, Victoria 3135, Australia and Department of Invertebrate Palaeontology, Museum Victoria,

PO Box 666E, Melbourne, Victoria 3001, Australia (fholmes@bigpond.net.au)

Abstract Holmes, F.C. 2004. A new Late Eocene cassiduloid (Echinoidea) from Yorke Peninsula, South Australia. Memoirs of

Museum Victoria 61(2): 209-216

A new species of cassiduloid, from the Muloowurtie Formation on the east coast of Yorke Peninsula, is described and

tentatively assigned to the genus Rhynchopygus. R7 janchrisorum sp. nov. is the first record of the genus, as redefined

by Smith and Jeffery (2000), to occur outside Europe, and the first confirmed from the Cainozoic. The history of species

previously assigned to Rhynchopygus is briefly discussed and details of further occurrences of the type species, R.

marmini, listed.

Keywords Echinoidea, Cassiduloida, Rhynchopygus, Late Eocene, South Australia, new species

Introduction

Australian Tertiary echinoids, donated to Museum Victoria

by RJ. Foster in 1996, included a single specimen of a cas-

siduloid quite distinct from any other species of this order so far

recorded from Australia. The specimen was collected in the

1970s from “Sliding Rocks” (Fig. 1), the type section of

the Muloowurtie Formation (redefined by Stuart, 1970), 9.9 km

SSW of Ardrossan and 1.1 km NNE of Muloowurtie Point,

on the east coast of Yorke Peninsula, South Australia [MV

locality PL3497].

Materials and methods. Specimen numbers prefixed P, on

which this study is based, are housed in the Invertebrate

Palaeontology Collection, Museum Victoria (NMV).

Measurements were made with a dial calliper to an accuracy of

0.1 mm. Parameters are expressed as a percentage of test length

(%TL) or test width (%TW).

Age and stratigraphy

The early to middle Late Eocene Muloowurtie Formation at

“Sliding Rocks” consists of a 12 m thick sequence of biogenic

calcarinites, quartz sands, calcareous and glauconitic quartz

sands and sandstones with minor thin conglomerates, silts and

clays, disconformably overlying the Lower Cambrian Kalpara

Formation. While it is unclear from which specific horizon

Foster collected his specimen (the holotype), three additional

specimens were found by C. Ah Yee and J. Krause during 2002,

in the uppermost bed of the formation about 800 m north of

“Sliding Rocks”. This latter horizon, consisting of variegated

argillaceous quartz sands interbedded with arenaceous clays,

lies immediately below the disconformity separating the

Muloowurtie Formation from the overlying Throoka Silts.

The fossiliferous beds of the Muloowurtie Formation are

generally considered to be Aldingan (Priabonian) in age lying

within planktonic foraminiferal zones P15 and PI 6, although

the base of the formation is probably late Middle Eocene (P14).

In general, the Muloowurtie Formation is contemporaneous

with the Tortachilla Limestone at Maslin Bay on the south-east

side of the Basin, both having been deposited during the

Tortachilla and Tuketja transgressions. However, the upper-

most beds of the formation are currently considered to be

equivalent to the upper middle part of the Blanche Point

Formation, which overlies the Tortachilla Limestone, a time of

relatively high sea level preceding the onset of the Chinamans

Gully regression.

Associated fauna

Stuart (1970) recorded the echinoids Fibularia gregarta Tate,

1885, Salenidia tertiaria (Tate, 1877) and Eupatagus sp. (?),

together with crinoid plates, brachiopods, bivalves, the

bryozoans Retopora and Cellopora, ostracods and foramini-

ferans from richly fossiliferous sands in the lower part of the

Muloowurtie Formation at “Sliding Rocks”. In addition to the

new cassiduloid, a typical Australian Late Eocene echinoid

fauna consisting of Australanthus longianus (Gregory, 1890),

210

Francis C. Holmes

Figure 1. A and B, general location maps; C, map of St Vincent Basin in relation to Yorke Peninsula, Kangaroo Island and the Fleurieu Peninsula

(Maslin Bay), South Australia; D, location of the northernmost exposure (1), and southernmost exposure (2), of the Muloowurtie Formation, west

side of Gulf St Vincent.

Eurhodia australiae (Duncan, 1877), Gillechinus cudmorei

Fell, 1963, and Schizaster ( Paraster ) tatei McNamara and

Philip, 1980, have been found in the same general area. Two of

these, Gillechinus cudmorei and Salenidia tertiaria occur with

Hemiaster ( Bolbaster ) cf. subidus McNamara, 1987 and the

new cassiduloid in the uppermost bed of the Formation.

Apatopygus vincentinus (Tate, 1891) was recorded from

“Muloowurtie”, near Ardrossan, although no specific localities

were given with Tate’s syntypes of this species. The neolam-

padoid Pisolampas concinna Philip, 1963, is also found in the

Formation in the vicinity of Harts Mine, approximately 3.8 km

south of “Sliding Rocks”.

A list of echinoid species recorded from the three major Late

Eocene formations in the St Vincent Basin, the Tortachilla

Limestone, Kingscote Limestone (lowest unit) and

Muloowurtie Formation, is given in the Appendix.

The difference in the number of species recorded from each

of these formations is no doubt primarily due to collecting bias;

the Tortachilla Limestone in the coastal cliffs along Maslin Bay

being well exposed and easily accessible from Adelaide.

Although the Kingscote (Kangaroo Island) and Muloowurtie

(Yorke Peninsula) deposits have been known for over 120 years

(Tepper, 1879; Tate, 1883), virtually no systematic study of

their echinoid fauna has been carried out. It is only in com-

paratively recent times that detailed stratigraphic information

relating to these deposits has been published (Kingscote

Limestone, Milnes et al., 1985; Muloowurtie Formation, Stuart,

1970). The variation in sedimentary lithology within the Basin,

a consequence of small basin size, narrow basin width, and the

paralic nature of deposition (Cooper, 1985), rather than any

minor age difference, must also be considered in relation to

species distribution and difference in size, preservation, and

number of specimens of individual species found in the three

formations. As well as the apparent unique occurrence of the

new cassiduloid in the Muloowurtie Formation, the lack of

Echinolampas posterocrassa posterocrassa Gregory, 1890, one

of the most common species in the Kingscote and Tortachilla

Limestones, may, in this context, be quite significant.

Systematic Palaeontology

Order Cassiduloida Claus, 1880

Family Faujasiidae Lambert, 1905

Rhynchopygus d’ Orbigny, 1856

Type species. Cassidulus marmini Agassiz, in Agassiz and

Desor, 1847, by monotypy.

Diagnosis. See Smith and Jeffery (2000: 191)

Remarks. Because of its monobasal apical system (Fig. 3) and

lack of known phyllode detail, the new Late Eocene species

from South Australia can only tentatively be assigned to the

genus; although Smith and Jeffery (2000: 192) suggested that

the apical system of Rhyncopygus donetzensis Faas, 1918, may

indeed be monobasal or at least have very reduced genital

plates. Rhyncopygus, has been used in the past 150 years as a

genus or subgenus to accommodate nearly 40 species, ranging

A new Late Eocene Australian echinoid

211

in age from early Late Cretaceous (Turanian) to Recent.

Virtually all of these species have subsequently been reassigned

to other genera, namely Cassidulus Lamark, 1801, Eurhodia

Haime, 1853, Procassidulus Lambert, 1918, Rhyncholampas

Agassiz, 1869, and even the holasteroid Corystus Pomel, 1883.

The type species of all these genera, except Eurhodia , have at

some time been assigned to Rhynchopygus. The most recent

review of Rhynchopygus (Smith and Jeffery, 2000) noted that

the differently shaped and positioned periprocts of three species

included in the genus by Kier (1962), the type species

R. marmini, R. lapiscancri (Leske, 1778) and R. macari

(Smiser, 1935), preclude uniting them into a single genus-level

taxon; referring only to R. marmini and R. donetzensis as

belonging to the genus. Although R. donetzensis was listed by

Lambert and Thiery (1925: 588), no reference to this important

species was made by Kier (1962). R. lapiscancri has since been

assigned by van der Ham et al. (1987) to Procassidulus and

R. macari by Smith and Jeffery (2000) to Rhyncholampas.

Without comment the latter authors also assigned both

Rhynchopygus and Procassidulus to the Faujasiidae, rather than

the Cassidulidae.

Both Mortensen (1948: 201) and Kier (1962:161) blame the

inaccurate illustrations of d’Orbigny (1856: pi. 927) for the

early taxonomic problems in defining Rhynchopygus. While

this is unquestionably true, d’Orbigny(1856) and Desor

(1855-1858) did illustrate and refer to the prominent lip-like

projection of the test that occurs in interambulacrum 5 adjacent

to the periproct; the main feature now considered to distinguish

Rhynchopygus from genera which have many other character-

istics in common. However, Mortensen (1948) considered this

projection to be of no generic value and Kier (1962) did not

even mention it in his generic diagnosis, but added to the con-

fusion by describing the periproct opening as either transverse

or longitudinal, presumably to accommodate other species then

assigned to the genus.

Mortensen (1948) considered Rhynchopygus a synonym of

Cassidulus while Kier (1962) regarded the tetrabasal apical

system as a major feature separating the two genera. In addition

he deemed Procassidulus, a genus retained by Mortensen, to be

a synonym of Rhynchopygus.

Rhyncopygus? janchrisorum sp. nov.

Figures 2A-F, 3A, B, 4A-C, 5, 6A

Type material. Holotype, NMV P145616 from Late Eocene (Aldingan,

Priabonian) Muloowurtie Formation, “Sliding Rocks”, Yorke

Peninsula, South Australia. Paratypes. NMV P312113 to P312115

from the uppermost bed of the same formation, NNE of “Sliding

Rocks” at approximately 34°34.17'S, 137°53.40'E (Fig. 1).

Description. Test moderately small, oval in outline at the

ambitus, anterior and posterior evenly rounded, widest point

central. Aboral surface moderately inflated, except for a

depression posterior to the periproct, with the apex just anteri-

or of the apical disk at the proximal end of slightly swollen

ambulacrum III. Adoral surface slightly depressed around

peristome and along the posterior paired ambulacra I and V.

Aboral tubercules very small, about 0.15mm diameter, and

closely spaced with a density of about 12-15 tubercles per

mm2. Adoral tubercles, where visible near the margin, are also

closely spaced but larger, about 0.6 mm diameter. Naked

granular zone in ambulacrum III and interambulacrum 5.

Apical system monobasal, centre of disk 42^-5 %TL from

anterior margin, with 4 gonopores in contact with the apical

disk but extending into the first pair of interambulacral plates,

anterior pair closer together than posterior pair. Ocular plates

relatively small and about equal in size. Approximately 80

hydropores (Fig. 3).

Petals moderately short, broad and unequal in length.

Longest in ambulacrum III (approx. 62% radius with about 30

pores per tract) and shortest in anterior pair II and IV (approx.

50% radius with about 20 pores per tract). Inner pores of pore

pairs oval, outer pores slot like, with alignment noticeably

oblique in ambulacra II and IV. At widest point interporiferous

zone in ambulacra II, III and IV about equal in width to zones

of pore pairs, parallel sided and open ended in ambulcrum III

and narrowing distally in II and IV. Posterior pair of petals have

narrower poriferous zones curving outwards distally with porif-

erous zones lb and Va noticeably wider than la and Vb.

Poriferous zones in individual petals equal in length. Anterior

pair of petals diverge between 134° and 140°, posterior pair

between 306° and 310°. The aboral swelling in ambulacrum III

forms a low ridge along line of perradial suture for full length

of petal.

Periproct supramarginal, transverse, situated at the anterior

end of a pronounced posterior anal depression in interam-

bulacrum 5, beneath a wide semi-circular projecting lip (about

19%TW) extending to nearly 20%TL from the posterior

margin. The anal depression di mi nishes posteriorly and barely

reaches the margin.

Peristome small, pentagonal, centre of opening situated

37-40%TL from the anterior margin, Floscelle well developed

with very pronounced pointed and inflated bourrelets, anterior

pair wide and wedge-shaped, posterior single and pair elon-

gated and near parallel sided. Phyllodes deeply sunken proxi-

mally but with pronounced ridge between bourrelets at edge of

peristomal opening. Detail of phyllode pores unknown.

Etymology. For Janice Krause and Christopher Ah Yee of

Hamilton, Victoria.

Remarks. The description is based on four specimens, all of

which have been subject to diagenetic compression resulting in

radial cracks along adradial and interradial sutures. In the case

of the holotype these cracks extend between half and two-thirds

distance between ambitus and distal end of petals. Compared

with a similarly compressed specimen of Australanthus lon-

gianus from the same locality, it is estimated that the holotype

of Rhyncopygusl janchrisorum sp. nov. would have been about

24.0 mm long, 20.5 mm (85.5%TL) wide and a minimum of

10.5 mm (44%TL) high compared with the 26.8 mm long, 22.9

mm wide and 7.8 mm high dimensions of the compressed

fossil test (Fig. 4). Rhyncopygusl janchrisorum is easily dis-

tinguished from the type species of the genus, R. marmini, in

having a larger, less elongated and less inflated test, a

monobasal apical disk, far more pronounced and broader

petals, and prominent bourrelets. It differs from R. donetzensis,

based on the illustrations in Smith and Jeffery (2000),

212

Francis C. Holmes

Figure 2. Rhynchopygusl janchrisorum sp. nov. A-C, adapical, right lateral and posterior views of holotype NMV P145616; D, adoral view of

paratype NMV P312115; E, adoral view of paratype NMV P312114; F, adapical view of paratype NMV P312113. All specimens form the Late

Eocene Muloowurtie Formation, Yorke Peninsula, South Australia. Scale bar 10 mm.

Figure 3. Apical plate structure. A and B, Rhynchopygusl janchriso-

rum sp. nov., Late Eocene holotype NMV P145616; C, R. marmini,

Late Cretaceous USNM 19559 from Port Brechay, La Manche, France

(drawing adapted from Kier, 1962). Scale bars 1mm.

Figure 4. Rhynchopygusl janchrisorum sp. nov. A-C, reconstruction

of adapical, right lateral and posterior characteristics of holotype NMV

P145616 (actual specimen outline shown with broken line). Scale bar

10 mm.

A new Late Eocene Australian echinoid

213

Figure 5. Rhynchopygus 1 } janchrisorum sp. nov. Drawing of adapical

surface of NMV P312113 showing plate structure distal to petals.

Scale bar 10 mm.

primarily in being far less tumid and having a longer and

broader petal in ambulacrum III, shorter posterior petals and

considerably wider ambulacral plates at the ambitus (Fig. 5).

The test of R.l janchrisorum is longer and proportionately

wider than the other two species and has a more anterior

periproct and projecting lip.

The occurrence of a similar but much less pronounced

canopy above the periproct is also present in Hardouinia

( Fauraster ) priscus Lambert, in Lambert and Thiery, 1924.

Kier (1962: 143), in his description of the poorly preserved and

compressed test, noted that the periproct is very wide and low;

however, Smith and Jeffery (2000: 205) referred to this as an

external feature that funnels into an opening only a little wider

than tall and list Hardouinia waageni Holland and Feldman,

1967, and Hardouinia nuratensis Moskvin, 1984, as synonyms

of this species. Apart from this feature, H. (F.) priscus differs

from R.l janchrisorum in its near circular outline with slightly

truncated posterior, its tetrabasal apical system and smaller and

equal sized petals with narrower poriferous tracts and much

wider interporiferous zone

Distribution and age of Rhynchopygus

The type species R. marmini recorded from France, Belgium,

and The Netherlands, has until now been considered to occur

only in the Late Maastrichtian, becoming extinct at the

Cretaceous/Tertiary boundary (Smith et al., 1999: 136). Recent

investigations (Philippe Mercier, pers. comm., 2004) showed

that R. marmini occurs in the Late? Santonian Calcarenite de la

Bouchardiere, Craie de Villedieu, at La Richardiere, Dissay

(Sarthe), France; and that three specimens (NMV P3 11748)

from cliffs on the right bank of the Gironde Estuary, between

the south side of Pointe Suzac northwards to Pointe de

Vallieres, Royan and St Palais-sur-Mer, France, are Late

Campanian. In the southeast Netherlands and northeast

Belgium, R. marmini has so far been recorded only from the

Nekum and Meerssen members of the Maastricht Formation

(Jagt, 2000: 263).

A partial specimen, R. sp., is recorded from the

Maastrichtian section, west of Cabo Major, near Santander

(Cantabria), Spain (Smith et al., 1999: 105), and the Museum

national d’Histoire naturelle, Paris, holds a single specimen

preserved in flint, labelled as originating from the Danian of St

Cristophe (sic), Loire et Cher, France, which Smith and Jeffery

(2000) suspected to be reworked Maastrichtian. Philippe

Mercier (pers. comm.) suggested this latter locality is probably

St Christophe in Eure et Loire (near Loire et Cher) but is

uncertain of the local stratigraphy.

The second species, R. donetzensis, is recorded from the

Maastrichtian (Cretace Superieur) of the Donetz Basin, in the

vicinity of Krymskoie and Serebrianca, Ukraine (Faas, 1918).

Savchinskaya (1974) referred to the species occurring in the

Maastrichtian of the Don Basin, Russia. As with R. marmini.

Smith and Jeffery (2000: 192) did not show this species

crossing the K/T boundary.

The additional information on French localities and the dis-

covery of R1 janchrisorum, extends the range of Rhynchopygus

from the Late? Santonian to the Late Eocene, an interval of

approximately 85 million years (Table 1).

Figure 6. Comparative drawings of A, Late Eocene Rhynchopygus ? janchrisorum sp. nov. from Yorke Peninsula, South Australia; B, Late

Cretaceous (Late Maastrichtian) R. donetzensis from the Severny Donetz Basin, Ukraine (adapted from Smith and Jeffery, 2000); Cl,

Late Cretaceous (Late? Santonian) R. marmini from La Richardiere, Dissay, France; C2, posterior profile of Late Cretaceous (Late Campanian)

R. marmini from near Royan and St Palais-sur-Mer, France. Based on published illustrations, latter profile typical of Maastrichtian R. marmini

specimens in general. Scale bars 5 mm.

214

Francis C. Holmes

Table 1. Currently known distribution and age of Rhynchopygus

marmini (1); R. sp. (2); R. donetzensis (3); and R1 janchrisorum sp.

nov. (4). Triangle ( ^ ) indicates occurrence in Late (Upper) section of

Stage; inverted triangle, Early (Lower) section ( ▼ ); and square (■),

non specific. Spain (Sp), France (Fr), Belgium (Bel), The Netherlands

(Net), Ukraine (Ukr). Australia (Aus).

Epoch

Stage

Est. Age Sp

Bel Net Ukr

Aus

33.0ma

Priabonian

37.0

■ 4

Bartonian

Eocene

Lutetian

41.3

49.0

Ypresian

54.8

Thanetian

57.9

Paleocene

Selandian

60.9

Danian

65.0

Maastrichtian 2

^ 1

^ 1 ^1 m3

73.0

Campanian

83.0

^ I

Late

Santonian

^ 1

Cretaceous

Coniacian

87.0

89.0

Turonian

91.0

Cenomanian

97.5

Found with R1 janchrisorum (see Appendix), Australanthus

Bittner, 1892, from the Middle/Late Eocene of southern

Australia is the only member of the Faujasiidae previously

recorded from this continent. Kier (1962: 18) considered

Australanthus a possible descendant of Hardounia Haime, in

d’Archiac and Haime, 1853.

Acknowledgements

I am indebted to Christopher Ah Yee and Janice Krause

(Hamilton, Victoria) for collecting and donating the paratypes;

Andrew Smith (Natural History Museum, London) for

sketches and details of Rhynchopygus donetzensis ; and Zang

Wenlong (Primary Industries and Resources, South Australia)

for stratigraphic information on the Muloowurtie Formation. I

thank Museum Victoria staff, David Holloway and Thomas

Darragh (Invertebrate Palaeontology) for continuous help and

encouragement; and Frank Job and Sandra Winchester

(Library) for access to references. I also thank Philippe Mercier

(St Rimay, France) for photographs, measurements, locality

information, and age of R. marmini specimens used for com-

parative purposes; and John Jagt (Nationaal Natuurhistorich

Museum Maastricht, The Netherlands) for current information

on Maastricht Formation specimens from The Netherlands and

Belgium.

References

Agassiz, A. 1869. Preliminary report on the Echini and starfishes

dredged in deep water between Cuba and the Florida Reef by L. F.

De Pourtales. Bulletin of the Museum of Comparative Anatomy,

Harvard University 1(9): 253-308.

Agassiz, L., in Agassiz, L. and Desor, E. 1847. Catologue raisonne des

families, des genres et des especes de la classe des echinodermes.

Annales des Sciences Naturelles 7: 129-168.

Bittner, A. 1892. Uber echiniden des Tertiars von Australien.

Sitzungsberichte der kaiserlichen Akademie der Wissenschaften zu

Wein (math. Naturw. Classe) 101(1): 331-371, pis 1-4.

Claus, C.F.W. 1880. Grundziige der Zoologie (4 edn) vol. 2. N.G.

Elwert: Margburg and Leipzig. 552 pp.

Cooper, B.J. 1985. The Cainozoic St Vincent Basin - tectonics, struc-

ture, stratigraphy. Pp. 35-49 in: Lindsay, J.M. (ed.), Stratigraphy,

palaeontology, malacology: papers in honour of Dr Nell Ludbrook.

Department of Mines and Energy, South Australia, Special

Publication 5.

Desor, E. 1855-1858. Synopsis des echinides fossiles. Ch. Reinwald:

Paris. Kriedel and Niedner: Weisbaden. lxiii + 490 pp., 44 pis.

Duncan, PM. 1877. On the Echinodermata of the Australian Cenozoic

(Tertiary) deposits. Quarterly Journal of the Geological Society of

London 33(1): 43-73, pis 3, 4.

Faas, A. 1918. Rhynchopygus donetzensis n. sp. des depots cretaces

superieurs du basin de la riviere Severny Donetz. Annuarie de la

Societe Paleontologique de Russie 2: 65-84, pi. 3 [In Russian with

French summary].

Fell, H.B. 1963. New genera of Tertiary echinoids from Victoria,

Australia. Memoirs of the National Museum of Victoria 26:

211-217.

Gregory, J.W. 1890. Some additions to the Australian Tertiary

Echinoidea. The Geological Magazine 27: 481-492. pis 13, 14.

Haime, J., in d’Archiac, V. D. and Haime, J. 1853. Descriptions des

animaux fossiles du groupe nummulitique de I’lnde. Gide and

Baudry: Paris. 373 pp, 36 pis.

Ham, R.WJ.M. van der, Wit, W. de, Zuidema, G., and Birgelen, M.

van, 1987. Zeeegels uit het Krijt en Tertiair van Maastricht, Luik en

Aken. Publicaties van het Natuurhistorisch Genootschap in

Limberg 36: 1-91, 24 pis.

Holland, F.D., and Feldman, R.M. 1967. A new species of cassiduloid

echinoid from the Fox Hills Formation (Upper Cretaceous) of

North Dakota. Journal of Paleontology 41: 252-255.

Holmes, F.C. 1993. Australian fossil echinoids: annotated bibliography

and list of genera and species. Occasional Papers of the Museum of

Victoria 6: 27-53.

Jagt, J. W. M. 2000. Late Cretaceous-Early Palaeogene echinoderms

and the K/T boundary in the southeast Netherlands and northeast

Belgium - Part 4: Echinoids. Scripta Geologica 121: 181-375.

Kier, PM. 1962. Revision of the cassiduloid echinoids. Smithsonian

Miscellaneous Collections 144(3): 1-262, pis 1-44.

Lamarck, J.B.1801. Systeme des animaux sans vertebres, ou, Tableau

general des classes, des orders, et des genres de ces animaux. Chez

Deterville: Paris, viii + 432 pp.

Lambert, J., in Doncieux, L. 1905. Catologue descriptif des fossiles

nummulitiques de l’Aube et de 1’ Herault. Annales de T Universite

deLyon 17: 129-164, pi. 5.

Lambert, J. 1918. Considerations sur la classification des Echinides

Atelostomes. 1. Brachygnata (sic) et Procassiduloida. Memoire

A new Late Eocene Australian echinoid

215

Societe Academique, d’ agriculture, des sciences, arts et belle lettres

du departement de l’ Aube 82: 1-48.

Lambert, J., and Thiery, P. 1910-1925. Essai de nomenclature raison-

nee des Echinides. Vols. 1-9. Libraire L. Ferriere: Charmont. 607

pp., 15 pis.

Leske, N.G. 1778. Jacobi Theodori Klein Naturalis disposito

Echinodermatum. Edita et descriptionibus novisque inventis et syn-

onymis auctorum aucta. Officiana Gleditschiana: Lipis. xxii + 278

pp., 54 pis.

McNamara, K.J. 1987. Taxonomy, evolution and functional morpholo-

gy of southern Australian Tertiary hemiasterid echinoids.

Palaeontology 30(2): 319-352.

McNamara, K.J., and Philip, G. M. 1980. Australian Tertiary schizas-

terid echinoids. Alcheringa 4(1): 47-65.

Milnes, A.R., Ludbrook, N.H., Lindsay, J.M., and Cooper, B.J. 1985.

The succession of Cainozoic marine sediments on Kangaroo Island,

South Australia. Transactions of the Royal Society of South

Australia 107(1): 1-35.

Moskvin, M.M. 1984. [Echinoids Domechinus and Hardouinia

(Echinoidea) from the Upper Cretaceous of Middle Asia].

Paleontologicheskii Sbornik 2 1 , 63-68, 1 pi. [In Russian],

Mortensen, T. 1948. A monograph of the Echinoidea 4(1).

Holectypoida, Cassiduloida. C. A. Reitzel: Copenhagen. 371 pp.,

14 pis.

d’Orbigny, A. 1856. Echinoides irreguliers. Paleontologie Frangaise:

descriptions zoologique et geologique de tous les Animaux

Mollusques at Rayonnes fossiles de France. Terrain Cretace 6.

Maisson: Paris. 596 pp., pis 801-1005.

Philip, G.M. 1963. Two Australian Tertiary neolampadids, and the

classification of cassiduloid echinoids. Palaeontology 6(4):

718-726, pis 106-107.

Pomel, N.A. 1883. Classification methodique et genera des echinides

vivants et fossiles. Adolphe Jourdan: Alger. 131 pp., 36 pis.

Savchinskaya, G.Y. 1974. [Atlas of the Upper Cretaceous fauna of the

Don Basin], Nidra Press: Moscow. 639 pp., 128 pis [In Russian],

Smiser, J.S. 1935. A monograph of the Belgian Cretaceous echinoids.

Memoires du Musee Royal d’Histoire Naturelle de Belgique 68:

1-98, pis 1-9.

Smith, A.B., Gallerm, J„ Jeffery, C.H., Ernst, G., and Ward, P. D. 1999.

Late Cretaceous-early Tertiary echinoids from northern Spain:

implications for the Cretaeous-Tertiary extinction event. Bulletin of

the Natural History museum, London (Geology), 55(2): 81-137.

Smith, A.B., and Jeffery, C. H., 2000. Maastrichtian and Palaeocene

echinoids: a key to world faunas. Special papers in palaeontology

63. The Palaeontological Association: London. 406 pp.

Stuart, W.J. Jnr 1970. The Cainozoic stratigraphy of the eastern coastal

area of Yorke Peninsula, South Australia. Transactions of the Royal

Society of South Australia 94: 151-178, figs 2-4.

Tate, R. 1877. On a new species of Belemnites and Salenia from the

Middle Tertiaries of South Australia. Quarterly Journal of the

Geological Society of London 33(2): 256-259.

Tate, R. 1883. The botany of Kangaroo Island. Transactions and

Proceedings of the Royal Society of South Australia 6, 116-171.

Tate, R. 1885. Miscellaneous contributions to the palaeontology of the

older rocks of Australia. Southern Science Record (n.s.) 1: 1-5.

Tate, R. 1891. A bibliography and revised list of the described echi-

noids of the Australian Eocene, with descriptions of some new

species. Transactions and Proceedings and Report of the Royal

Society of South Australia 14: 270-282.

Tepper, O. 1879. Introduction to the cliffs and rocks at Ardrossan,

Yorke’s Peninsula. Transactions and Proceedings and Report of the

Philosophical Society of Adelaide, South Australia 2: 71-79.

216

Francis C. Holmes

Appendix

South Australian Middle?-Late Eocene echinoids recorded from the Tortachilla Limestone, Maslin Bay (TL), lowest unit of the Kingscote

Limestone, Kangaroo Island (KL), and the Muloowurtie Formation, Yorke Peninsula (MF). Named species confirmed from all three formations

are marked ► . Information based on published literature and specimens housed in Museum Victoria Invertebrate Palaeontology collection or

privately owned. References to authors cited but not listed in the main text references can be found in Holmes (1993)

Cidaroida

Cidaridae sp.

Stereocidaris cudmorei Philip, 1964

S. fasten Philip, 1964

S. inermis Philip, 1964

S. sp. ‘C’ Philip, 1964

S. sp. [unidentified]

S. (?) hispida Philip, 1964

S. (?) intricata Philip, 1964

•l

•

•i

• 1 2 3 4

Salenoida

► Salenidia tertiaria ( Tate, 1877)

•

Temnopleuroida

Tatechinus nudus Philip, 1969

Temnopleuridae sp.

Ortholophus bittneri Philip, 1969

•

•l

•

•i

• 1

Clypeasteroida

Fibularia. sp. ‘A’ [Irwin pers. com.]

F. sp. ‘B’ [Irwin pers. com.]

F. sp. ‘C’ [Irwin pers. com.]

•

F. sp. [unidentified, non. F. gregata of Stuart, 1970]

Monostychia sp. ‘A’ [small undescribed species]

• 4

• 1

M. sp.’B’ [medium sized undescribed (?) species]

• 4

Cassiduloida

► Apatopygus vincentinus ( Tate, 1891)

•

► Australanthus longianus (Gregory, 1891)

•

• 4

Echinolampas posterocrassa Gregory, 1890

•

• 4

► Eurhodia australiae (Duncan, 1 877)

•

Rhyncopygusl janchrisorum sp. nov.

•

Neolampadoida

Aphanoporal bassoris Holmes, 1995

•4

•

► Pisolampas concinna Philip, 1963

•

Holasteroida

Corystus dysasteroides (Duncan, 1877)

Giraliaster bellissae Foster and Philip, 1978

G. sulcatus (Hutton, 1873)

G. tertiaria (Gregory, 1890)

•

• 2

Spatangoida

Eupatagus sp.

•3

► Gillechinus cudmorei Fell, 1963

•

• 4

Hemiaster (Bolbaster) subidus McNamara, 1987

Linthia pulchra McNamara, 1985

Prenaster aldingensis Hall, 1907

Protenaster preaustralis McNamara, 1985

•

• 4

Psephoaster lissos McNamara, 1987

Schizaster (Paraster) tatei McNamara and Philip, 1980

•

• 4

Total

27+3?

11+4?

10+3?

1 Not specifically identified, may belong to one of the listed species

2 Although almost certainly Late Eocene, occurrence of species in the formation requires confirmation

3 No reference to occurrence of species in taxonomic literature: may have been incorrectly identified or come from an overlying stratigraphic

unit

4 Previously unpublished identification

Memoirs of Museum Victoria 61(2): 217-227 (2004)

ISSN 1447-2546 (Print) 1447-2554 (On-line)

http://www.museum.vic.gov.au/memoirs/index.asp

A new species of Quinquelaophonte (Crustacea: Copepoda: Harpacticoida:

Laophontidae) from Port Phillip Bay, Victoria, Australia

Genefor K. Walker-Smith

Marine Invertebrates Section, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia and Marine

Biology Laboratory, Museum Victoria, GPO Box 666E, Melbourne, Vic. 3001, Australia

Present address: Invertebrate Zoology, Tasmanian Museum and Art Gallery, GPO Box 1164, Hobart, Tasmania 7001,

Australia and School of Zoology, University of Tasmania, Private Bag 5, Hobart, Tasmania 7001, Australia

(genefor. walker- smith @ utas . au)

Abstract Walker-Smith, G.K. 2004. A new species of Quinquelaophonte (Crustacea: Copepoda: Harpacticoida: Laophontidae)

from Port Phillip Bay, Victoria, Australia. Memoirs of Museum Victoria 61(2): 217-227.

A new species of Quinquelaophonte Wells, Hicks and Coull, 1982 from Port Phillip Bay, is separ-

ated from its cogeners by the presence of five elongate setae on PI exopod-2. The PI exopod-2 of all other species of

Quinquelaophonte has two long setae and three shorter spines. In addition, the PI endopod-2 of the new species has an

accessory seta that is longer than the terminal claw. This condition is also present in Q. wellsi, the only other species of

Quinquelaophonte described from Australia. The new species brings the total number of species of Quinquelaophonte

to ten.

Keywords Quinquelaophonte, Laophontidae, Harpacticoida, marine Copepoda, Port Phillip Bay, Victoria, Australia

Introduction

Australia’s harpacticoid fauna is diverse. In a recent survey of

harpacticoids from Port Phillip Bay more than 50 species were

identified from the shallow subtidal seagrass and surrounding

sandy areas (Walker-Smith, 2003). It is estimated that only a

small percentage of Australian harpacticoid species has been

described; 94 species have been described from marine, estuar-

ine and brackish water ecosystems, although worldwide there

are in excess of 3000 species (Giere, 1993; Huys et al., 1996).

Sixty-four of the species found in Australia are considered

endemic.

Nine species of Quinquelaophonte Wells, Hicks and Coull,

1982 have been described from around the world (Lee, 2003),

including Q. wellsi (Hamond, 1973), which was described from

a saline lake in South Australia. During a survey of

Harpacticoida in Port Phillip Bay, a new species of

Quinquelaophonte common on the surface of the unvegetated

sediment adjacent to the subtidal seagrass, Heterozostera

tasmanica (Martens ex Ascherson) den Hartog, was discovered.

This new species was rarely found among the seagrass itself

(Walker- Smith, 2003). Several other species of Quin-

quelaophonte are considered to be sediment-surface dwellers;

Q. wellsi, Q. candelabrum Wells, Hicks and Coull, 1982 and

Q. longifurcata (Lang, 1965) (Hamond, 1973; Wells et al.,

1982; Lang, 1965).

Seagrass and sediment samples were collected by hand (Walker-

Smith, 2003) and fixed in 4% buffered formalin in sea-water. After (at

least) 48 hours samples were washed over a 63-pm mesh sieve and

retained material was transferred to 70% ethanol. Samples were exam-

ined under a Zeiss Stemi S V 1 1 or a Wild M8 stereomicroscope and

harpacticoids were extracted using fine forceps. Harpacticoids were

dissected in a drop of glycerol on a microslide, using electrolytically-

sharpened tungsten needles. Appendages were mounted in glycerol.

Microslides were examined using three microscopes (Olympus BX50

and Leica DMR compound microscopes with Nomarski interference

contrast, Leitz Dialux 22 compound microscope). Illustrations were

made with the aid of a camera lucida. Once appendages were illu-

strated, they were permanently mounted in Gurr’s Aquamount and

coverslips were sealed with clear nail varnish. Scanning electron

micrographs were taken using a Philips XL20 scanning electron

microscope (KV=10, spot size 3).

Terminology used follows that of Huys and Boxshall (1991).

Abbreviations used are: Al, antennules or first antennae; A2, antennae

or second antennae; Mxl, maxillules; Mx, maxillae; P1-P4, swimming

legs 1-4. Individual segments of P1-P4 rami are written (for example)

as PI exopod-3, which refers to the third or terminal segment of the PI

exopod. P5 and P6 refer to the fifth and sixth legs. Total length meas-

urements are from the base of the rostrum to the posterior margin of the

caudal rami (caudal setae are excluded). Armature formulae (also

known as the setal formulae) for swimming legs are constructed fol-

lowing the methods of Lang (1934) (also see Huys and Boxshall, 1991 :

29). The term “armature” is used to refer collectively to setae and

spines. Type material is held in Museum Victoria (NMV) and the South

Australian Museum (SAM).

218

Genefor K. Walker-Smith

Table 1 . Distribution of Quinquelaophonte species

Species

Distribution

Reference

Q. brevicomis (T. Scott, 1894)

Ghana

T. Scott, 1894

Q. quinquespinosa (Sewell, 1924)

India: Chilka Lake, Orissa

Sewell, 1924

Egypt: Lake Menzaleh

Gurney, 1927

Bermuda

Willey, 1930; Lang, 1948

Tunisia: Goulette

Monard, 1935

Reunion

Chappuis et al., 1956

Angola: Benguela

Candeias, 1959

USA: Puget Sound, Seattle, Washington State

Wieser, 1959

France: Marseilles

Bodin, 1964; Hamond, 1973

Mozambique: Inhaca Island

Wells, 1967

Seychelles: Aldabra

Wells and McKenzie, 1973

Andaman Islands

Wells and Rao, 1987

Q. capillata (Wilson, 1932)

USA: Katama Bay, Marthas Vineyard, Massachusetts;

Wilson, 1932; Coull, 1976, 1986

North Inlet, Georgetown, South Carolina

Coull, 1986

Bahamas: Eleuthera; Crooked Island

Fiers, 1986

Q. longifurcata (Lang, 1965)

USA: California

Lang, 1965

Q. parasigmoides (Bozic, 1969)

Reunion: St Phillippe

Bozic, 1969

Q. wellsi (Hamond, 1973)

Australia: Robe and Beachport, South Australia

Hamond, 1973

Q. candelabrum Wells, Hicks and Coull, 1982

New Zealand: Raion Point, Pauatahanui Inlet, Porima

Harbour, Wellington; Papanui Inlet, Otago Peninsula;

Whangateau Harbour, Northland; Avon-Heathcote Estuary,

Christchurch; Hobson’s Bay, Waitemata Harbour, Auckland

Wells, Hicks and Coull, 1982

Q. bunakenensis Mielke, 1997

Indonesia: Sulawesi

Mielke, 1997

Q. koreana Lee, 2003

Korea: Taean

Lee, 2003

Q. prolixasetae sp. nov.

Australia: Port Phillip Bay, Victoria

present study

Quinquelaophonte Wells, Hicks and Coull, 1982

Quinquelaophonte Wells, Hicks and Coull, 1982: 178-179.

Type species. Laophonte quinquespinosa Sewell, 1924.

Diagnosis. A1 of female with 5 or 6 segments; A2 exopod

reduced, 1- segmented; P2-P4 exopods of male strongly modi-

fied; P2 endopod of male not modified (same as for female); P5

of male reduced to 4-5 setae arising from the somite margin;

caudal rami with 3 terminal setae, only one of which is well

developed (seta V).

Species. Quinquelaophonte brevicomis (T. Scott, 1894);

Q. quinquespinosa (Sewell, 1924); Q. capillata (Wilson, 1932);

Q. longifurcata (Lang, 1965); Q. parasigmoides (Bozic, 1969);

Q. wellsi (Hamond, 1973); Q. candelabrum Wells, Hicks and

Coull, 1982; Q. bunakenensis Mielke, 1997; Q. koreana Lee,

2003; Q. prolixasetae sp. nov.

Distribution. See Table 1 .

Habitat. Marine, intertidal and shallow subtidal; in saline lakes;

sand and mud. Frequently recorded in detritus-rich habitats.

Remarks. Fiers (1986) discovered specimens of Q. quinque-

spinosa (from the West Indies) and noted the specimens had an

“interesting feature”: the inner distal edge of P3 endopod- 1

with a few “long and fragile hairs” (Fiers, 1986: 142). Because

Fiers (1986) believed these “hairs” resembled the inner seta of

Q. parasigmoides , he suggested Q. parasigmoides was within

the range of variability of Q. quinquespinosa and therefore

should be considered synonymous with it. However, Lee

(2003) rejected this, instead believing confirmation of the syn-

onymy required examination of more specimens, from more

localities and I agree.

In his catalogue of marine harpacticoids, Bodin (1997)

listed Paronychocamptus wilsoni Coull, 1976 as a junior syn-

onym of Q. capillata but Lee and Huys (1999) recognized

P. wilsoni as a valid species and I support this. Inspection of

Coull’s (1976) illustrations of P. wilsoni revealed distinct dif-

ferences between this species and Q. capillata. Firstly, the P3

endopod-2 of male P. wilsoni is modified and has a spine-like

distal outgrowth and this modification does not occur in

Q. capillata. Secondly, the P5 exopod of the male is well devel-

oped and has four setae in P wilsoni but is reduced and repre-

sented by five setae in Q. capillata. For all other species of

Quinquelaophonte the P5 exopod of male is reduced. The P5 of

female P. wilsoni has only four setae on the baseoendopod and

five setae on the exopod, while in Q. capillata there are five

setae on the baseoendopod and six setae on the exopod. The

caudal setae of the two species also differ; P. wilsoni has two

well developed terminal setae but Q. capillata has only one

well developed terminal seta (seta V) — the possession of only

one well developed caudal seta is a character state defining

Quinquelaophonte. The setal formula for the swimming legs

also varies between these species (Table 2). Lastly, when Coull

(1986) re-examined Wilson’s type material he discovered the

A2 exopod of Q. capillata had three setae, and not two as

originally reported. The A2 exopod of P. wilsoni has only

two setae.

New species of harpacticoid copepod

219

Table 2. Comparison of the setal formulae of Quinquelaophonte

capillata and Parony choc amp tus wilsoni. Endp = endopod;

exp = exopod.

Segments of swimming legs

P2 endp-2 P3 exp-3 P3 endp-3 P4 exp-3 P4 endp-2

Q. capillata 1.2.0 1.2.3 2.2.1 1.2.3 1.1.1

P. wilsoni 2.2.0 2.2.3 3.2.1 1.2.2 1.2.0

Quinquelaophonte prolixasetae sp. nov.

Figures 1-8

Material examined. Holotype. NMV J52388 (ovigerous female, on 8

slides). Australia, Victoria, Port Phillip Bay: at the end of Grand Scenic

Drive, in front of the Sands Caravan Park, Moolap (38°09.92'S

144°28.42'E). Collected from the surface of unvegetated sediments,

adjacent to sub tidal seagrass beds of Heterozostera tasmanica (water

depth approximately 1 m), G. K. Walker-Smith, 17 Nov 1997.

Paratypes. NMV J52389 (1 male, allotype, on 6 slides), NMV

J53020 (1 female on 5 slides), NMV J53110 (1 female on 5 slides),

NMV J53111 (1 female on 1 slide), NMV J53112 (1 male on 1 slide),

NMV J53021 (20 adult females and 2 juveniles), NMV J53022

(9 males), SAM C6096 (20 females, including 2 ovigerous and 4 juve-

niles), SAM C6097 (9 males), SAM C6098 (8 females on a SEM stub),

SAM C6099 (4 males on an SEM stub). All paratypes collected with

the holotype.

Other material. NMV J48528 (13 specimens), NMV J48529 (3

specimens), NMV 48530 (5 specimens), NMV J48531 (12 specimens),

NMV J48532 (29 specimens), NMV J48533 (3 specimens), NMV

J48535 (103 specimens). Collection data as for holotype.

Diagnosis. A1 of female 6-segmented; A2 exopod with 3 setae;

PI exopod-2 with 5 thin, elongate setae; PI endopod-2 acces-

sory setae longer than terminal claw; P3 exopod-3 of female

with 1 inner seta; P3 endopod-2 of female with 5 setae; P4

exopod-3 of female without inner seta; P4 endopod-2 with

3 setae; P5 exopod of female with 6 setae; P5 of male with

5 setae.

Adult dimensions. Females: mean length 0.85 mm ± 0.05 mm

(n = 22). Males: mean length 0.82 mm ± 0.03 mm (n = 21).

Description of female. Body tapering posteriorly (Fig. 1A-B).

Rostrum fused to cephalothorax, with 2 sensillae. Clear delin-

eation of body somites. Somite margins with setules. Hyaline

frill present on somite preceding anal somite. Anal somite with

anal operculum (Figs 3D, 8D). Caudal rami length 3 times

width (Fig. 3D); 3 setae on lateral margin (Figs 3D, 8C), seta I

minute (difficult to see under compound microscope), seta IV

reduced, seta V well developed and covered with minute

spinules (only visible via SEM) (Figs 3D, 8C), terminal acces-

sory seta (VI) on inner subdistal corner, dorsal seta (VII)

triarticulate at the base.

Antennule 6-segmented (Fig. 2A), aesthetasc fused basally

to seta on segment 4, terminal segment with smaller aesthetasc

fused basally to 2 setae (i.e. tritheck). Antenna with allo-

basis (Fig. 2B), abexopodal seta reduced to a small spine no

different from the surrounding spines, exopod reduced to

single segment with 3 setae. Endopod with 2 pinnate spines

Figure 1. Quinquelaophonte prolixasetae sp. nov., female, paratype

(NMV J53020): A, habitus, dorsal view; B, habitus, lateral view.

laterally and 3 geniculate setae and 2 pinnate spines terminally.

Endopod also with spinules laterally and a subapical hyaline

frill.

Mouthparts. Labrum with setules along anterior margin

(Figs 2G, 7A). Paragnaths as in figures 2H and 7A. Mandible

with well developed gnathobase (Fig. 2C), palp 1- segmented

and with 4 setae (endopod and exopod fused to basis), endopod

represented by 3 setae, exopod represent by 1 seta, basal arma-

ture represent by a larger pinnate seta. Maxillule (Figs 2D, 7A)

arthrite with 6 spines and a row of setae on the posterior

surface, also with 1 seta on lateral margin; coxa with 1 smooth

seta, 1 long spine and a row of spinules on upper sur-

face; endopod and basis fused, endopod represented by

New species of harpacticoid copepod

221

Figure 3. Quinquelaophonte prolixasetae sp. nov., female, holotype (NMV J52388): A, PI; C, P2. B, P2 Q. wellsi (redrawn from Hamond, 1973).

Female, paratype (NMV J53020): D, caudal rami and anal somite.

2 setae, basis endite with 2 smooth setae and 1 long spine;

exopod 1-segmented and with 2 smooth setae. Maxilla (Figs

2E, 7B) syncoxa with 3 endites, first endite with 1 spinose seta,

middle endite with 2 setae and 1 pinnate spine, distal endite

with 1 pinnate spine and 2 setae; allobasis with a pinnate claw

and 2 smooth setae inserted at the base of the claw, also with 3

lateral setae that are remnants of the endopod. Maxilliped (Fig.

2F), prehensile, syncoxa with 2 setae, basis without orna-

mentation, endopod represented by terminal claw with 1 seta

and some distal spinules.

PI (Figs 3 A, 8A-B) coxa with spinules on outer margin.

Basis with 2 rows of spinules and 2 spinulose spines. Exopod

2-segmented, exopod- 1 with 3 rows of spinules and 1 unipin-

nate spine, exopod-2 with 5 elongate, smooth setae. Endopod

2-segmented, endopod- 1 with fine setules along inner margin,

endopod-2 with 1 spinulose claw, 3 setae on lateral margin and

2 short setae at the base of the claw, terminal accessory seta is

more than twice the length of the claw.

P2-P4 exopod 3-segmented, endopod 2-segmented (Figs

3C, 4A-B). P2 endopod reaching just beyond distal margin of

P2 exopod-2 (Fig. 3C). P3 endopod not reaching beyond distal

margin of P3 exopod-2 (Fig. 4A). P4 endopod not reaching

222

Genefor K. Walker-Smith

Figure 4. Quinquelaophonte prolixasetae sp. nov., female, holotype

(NMV J52388): A, P3; B, P4.

beyond distal margin of P4 exopod-2 (Fig. 4B). P3 and P4

endopod-2 without tube pore.

Armature formulae for swimming legs:

Exopod

Endopod

0.1.123

0.120

0.1.123

0.221

0.1.023

0.120

P5 baseoendopod (Fig. 5C), outer setophore with 1 seta,

endopodal lobe with 2 serrate spines and 3 smooth setae,

endopodal lobe not reaching to distal margin on exopod.

Exopod longer than wide, with 3 pinnate setae and 3 smooth

setae, as well as some spinules.

Description of male. Male same as for female except for the

following: A1 subchirocer and without seta on segment- 1 (Figs

21, 7C-F); P2-P4 larger and more chitinised (Figs 5B, 6A-B),

lateral spines longer, exopod-3 almost at right angles to exo-

pod-2. P4 endopod-2 more than 2 times length of endopod-1.

P5 reduced to 5 setae (Fig. 5D).

Etymology. Prolixus (Latin): stretched out, long; plus setae

(Latin): bristles; referring to the five elongate setae on PI

exopod-2.

Figure 5. Quinquelaophonte prolixasetae sp. nov. male, paratype

(NMV J52389): A, PI and intercoxal sclerite; B, P2 and intercoxal

sclerite; D, section of urosome, ventral view showing P5 and P6.

Female, holotype (NMV J52388): C, P5.

Distribution. Australia, Victoria, Port Phillip Bay, specifically:

Blairgowrie, St Leonards, Grassy Point, Point Richards, Clifton

Springs and Moolap.

Remarks. Although all appendages of the holotype have been

mounted on microscope slides, the orientation of the mouth-

parts did not allow for clear illustration, thus mouthparts have

been illustrated using paratypes. Careful comparison of the

holotype and paratypes were made.

Discussion

Quinquelaophonte prolixasetae is the second species of the

genus described from Australia and is distinguished from its

cogeners by the presence of five elongate setae on the PI exo-

pod-2. All other species of Quinquelaophonte have two long

setae and three spines on PI exopod-2 (Pig. 3B).

New species of harpacticoid copepod

223

Quinquelaophonte prolixasetae is most closely related to Q.

wellsi (the other Australian species) sharing the unusual char-

acter of the PI endopod-2 accessory seta longer than the ter-

minal claw. Character states separating Q. prolixasetae from

Q. wellsi are: P3 exopod-3 with one inner seta (two in

Q. wellsi ), P4 exopod-3 without an inner seta (with one in

Q. wellsi ) and the setation of PI exopod-2 as mentioned above.

Many illustrations of species of Quinquelaophonte lack fine

detail, however, in a recent paper (Lee, 2003) several smaller

features, possibly omitted by previous authors, were illustrated.

Lee (2003) noted the abexopodal seta on the A2 of Q. koreana

was a “tiny . . . pinnate seta”. In most illustrations of the A2 (of

other Quinquelaophonte species) no distinction has been made

between the abexopodal seta and the other spinules on the

allobasis. This lack of distinction may be because the difference

went unnoticed or it may simply be that the abexopodal seta

appears exactly like the other allobasis spinules. I was unable

to distinguished the abexopodal seta of Q. prolixasetae from

the other spinules on the ventral margin of the allobasis. The

abexopodal seta of Q. bunakenensis, Q. candelabrum and

Q. wellsi is longer than the neighbouring allobasis spinules

(Mielke, 1997; Wells et al., 1982; Hamond, 1973). The maxil-

lule of Q. prolixasetae also differed from that of Q. koreana

having a short lateral seta on the lateral margin of the arthrite

instead of a long one, as found in Q. koreana. The illustration

of the maxillule arthrite of Q. wellsi did not include a lateral

seta (Hamond, 1973). Quinquelaophonte bunakenensis and

Q. parasigmoides both have a long seta on the distal end of the

maxillule arthrite (Mielke, 1997; Bozic, 1969). Lee (2003)

noted P3 and P4 endopod-2 of Q. koreana possessed a tube

pore. This character state was not observed in Q. prolixasetae

and has not been illustrated for any other species of

224

Genefor K. Walker-Smith

Figure 7. Quinquelaophonte prolixasetae sp. nov. Female (SAM C6098): A, mouthparts: a, labmm; b, paragnath; c, Mxl; Mx; B, Mx. Male (SAM

C6099): C, A1 (open) dorsal view; D, A1 dorsal view; E, A1 ventral view; F, A1 ventral view close up.

Quinquelaophonte. The possession a single well developed

terminal seta (V) on the caudal rami is a diagnostic feature of

Quinquelaophonte. Seta V and seta IV are fused basally in Q.

koreana but are not fused in Q. prolixasetae. The distal half of

seta V of Q. koreana has tiny spinules covering the surface but

the entire length of seta V of Q. prolixasetae, is covered with

minute spinules.

Some of the character states distinguishing Q. prolixasetae

from all other species of Quinquelaophonte are listed in table 3.

Acknowledgements

This work was supported (in part) by a grant from the

Australian Biological Resources Study. I would like to thank

New species of harpacticoid copepod

225

226

Genefor K. Walker-Smith

Figure 8. Quinquelaophonte prolixasetae sp. nov. female (SAM C6098): A, PI dorsal view; B, PI endopod-2 and claw; C, caudal rami, ventro-

lateral view; D, anal operculum and caudal rami, dorsal view.

two anonymous reviewers whose valuable comments lead to

the improvement of this manuscript. Thanks to the staff and

students from the Marine Biology Laboratory at Museum

Victoria for allowing me to use their microscopes. Thanks also

to Lyn Waterhouse from Adelaide Microscopy for her

assistance with the SEM.

References

Bodin, P. 1964. Recherches sur la systematique et la distribution des

Copepodes Harpacticoides des substrates meubles des environs de

Marseille. Recueil des travaux de la Station marine d’Endoume.

Faculte des Sciences de Marseille 51(35): 107-183.

Bodin, P. 1997. Catalogue of the new marine harpacticoid copepods.

L’lnstitut Royal des Sciences Naturelles de Belgique: Bmssels. 304

pp.

Bozic, B. 1969. Copepodes Harpacticoides de la Reunion. Bulletin du

Museum National d'Histoire Naturelle 41(4): 867-882.

Candeias, A. 1959. Contribution to the knowledge of the harpacticoids

(Crustacea, Copepoda) from the littoral of Angola. Publicagoes

Culturais da Companhia de Diamantes de Angola 45: 77-104.

Chappuis, P.A., Delamare Deboutteville, C., and Paulian, R. 1956.

Cmstaces des eaux souterraines littorals d’une resurgence d’eau

douce a la Reunion. Memoires de I’Institut Scientifique de

Madagascar 11 A: 51-78.

Coull, B.C. 1976. On the two laophontid harpacticoid copepods

described by Wilson as Laophonte capillata, with keys to the genus

Paronychocamptus. Transactions of the American Microscopical

Society 95(1): 35-45.

Coull, B.C. 1986. A new species of Pseudobradya and the rediscovery

and correction of Quinquelaophonte capillata (Copepoda:

Harpacticoida). Transactions of the American Microscopical

Society 105(2): 121-129.

Fiers, F. 1986. Harpacticoid Copepoda from the West Indian islands:

Laophontidae (Copepoda, Harpacticoida). Bijdragen tot de

Dierkunde 55(2): 331-426.

Giere, O. 1993. Meiobenthology: the microscopic fauna in aquatic

sediments. Springer- Verlag: Berlin. 328 pp.

Gumey, R. 1927. Zoological results of the Cambridge expedition to the

Suez Canal, 1924. 23. Report on the Crustacea-Copepoda (littoral

and semi-parasitic). Transactions of the Zoological Society of

London 22: 451-577.

Hamond, R. 1973. The harpacticoid copepods (Crustacea) of the

saline lakes in southeast Australia, with special reference to

the Laophontidae. Records of the Australian Museum 28(17):

393-420.

New species of harpacticoid copepod

227

Huys, R., and Boxshall, G.A. 1991. Copepod Evolution. The Ray

Society: London. 468 pp.

Huys, R., Gee, J.M., Moore, C.G., and Hamond, R. 1996. Marine and

brackish water harpacticoid copepods, part 1. Field Studies

Council: Shrewsbury. 352 pp.

Lang, K. 1934. Marine Harpacticiden von der Campbell-Insel und eini-

gen anderen sudlichen Inseln. Acta Universitatis Lundensis, New

Series, Avd. 2, 30:1-56.

Lang, K. 1948. Monographie der Harpacticiden. Hakan Ohlsson’s

Bpktrycheri: Lund and Nordiska Bpkhandeln: Stockholm. 2 vols,

1682 pp.

Lang, K. 1965. Copepoda Harpacticoidea from the Californian Pacific

coast. Kungliga Svenska Vetenskapsakademiens Handlingar (series

4) 10(2): 1-560.

Lee, W. 2003. A marine harpacticoid, Quinquelaophonte koreana sp.

nov. from a sandy beach in Korea (Crustacea: Copepoda).

Zoological Science 20: 657-668.

Lee, W. and Huys, R. 1999. Bathylaophonte gen. nov. from deep-sea

hydrothermal vents and the polyphyly of Paronychocamptus

(Copepoda: Harpacticoida). Cahiers de Biologie Marine 40:

293-328.

Mielke, W. 1997. On a small collection of Laophontidae (Copepoda)

from Sulawesi, Indonesia. Microfauna Marina 1 1 : 223-250.

Monard, A. 1935. Les Harpacticoides marins de la region de

Salammbo. Bulletin de la Station Oceanographique de Salammbo

34: 1-94.

Scott, T. 1894. Report on Entomostraca from the Gulf of Guinea, col-

lected by John Rattray, B.Sc. Transactions of the Linnean Society of

London (Series 2), Zoology 6(1): 1-161.

Sewell, R.B.S. 1924. Fauna of the Chilka Lake. Crustacea Copepoda.

Memoirs of the Indian Museum 5: 771-851.

Walker-Smith, G.K. 2003. The harpacticoid copepod fauna of Port

Phillip Bay (Victoria, Australia) and their contribution to the diet of

juvenile King George whiting ( Sillaginoides punctata :

Sillaginidae). PhD thesis, Department of Zoology, The University

of Melbourne: Victoria, Australia. 278 pp.

Wells, J.B.J. 1967. The littoral Copepoda (Cmstacea) of Inhaca Island,

Mozambique. Transactions of the Royal Society of Edinburgh 67:

189-358.

Wells, J.B.J., Hicks, G.R.F., and Coull, B.C. 1982. Common harpacti-

coid copepods from New Zealand harbours and estuaries. New

Zealand Journal of Zoology 9: 151-184.

Wells, J.B J., and McKenzie, K.G. 1973. Report on a small collection

of benthic copepods from marine and brackish waters of Aldabra,

Indian Ocean. Crustaceana 25(2): 133-146.

Wells, J.B.J., and Rao, G.C. 1987. Littoral Harpacticoida (Crustacea,

Copepoda) from Andaman and Nicobar Islands. Memoirs of the

Zoological Survey of India 16(4): 1-385.

Wieser, W. 1959. Free-living nematodes and other small invertebrates

of Puget Sound beaches. University of Washington Publication in

Biology 19: 1-79.

Willey, A. 1930. Copepod phenology — Observations based on new

material from Canada and Bermuda. Archivio Zoologico Italiano

16: 601-617.

Wilson, C. B. 1932. The copepods of the Woods Hole region

Massachusetts. Bulletin of the United States National Museum 158:

1-365.

129 > Dimorphic brooding zooids in the genus Adeona Lamouroux from Australia

(Bryozoa: Cheilostomata)

Philip E. Bock and Patricia L. Cook

135 > A review of Australian Conescharellinidae (Bryozoa: Cheilostomata)

Philip E. Bock and Patricia L Cook

183 > A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia

Erich M. G. Fitzgerald

209 > A new Late Eocene cassiduloid (Echinoidea) from Yorke Peninsula, South Australia

Francis C. Holmes

217 > A new species of Quinquelaophonte (Crustacea: Copepoda: Harpacticoida: Laophontidae) from

Port Phillip Bay, Victoria, Australia

Genefor K. Walker-Smith

Memoirs of Museum Victoria

Volume 61 Number 1 2004

1 > A molecular and morphological revision of genera of Asterinidae (Echinodermata: Asteroidea)

P. Mark O’Loughlin and Jonathan M. Waters

41 > A new genus of millipedes (Diplopoda: Polydesmida: Dalodesmidae) from wet forests in southern

Victoria, with brief remarks on the Victorian Polydesmida

Robert Mesibov

47 > Biosystematics of Australian mygalomorph spiders: descriptions of three new species of Teyl

from Victoria (Araneae: Nemesiidae)

Barbara York Main

57 > Chirostylidae from north-western Australia (Crustacea: Decapoda: Anomura)

Shane T. Ahyong and Keiji Baba

65 > Sicafodiidae, fam. nov. for Sicafodia stylos, gen. nov., from the marine bathyal of south-eastern

Australia (Crustacea: Amphipoda: Gammaridae)

Jean Just

75 > Pseudidotheidae (Crustacea: Isopoda: Valvifera) reviewed with description of a new species,

first from Australia

Gary C. B. Poore and Tania M. Bardsley

85 > The long-horned caddisfly genus Oecetis (Trichoptera: Leptoceridae) in Australia: two new species

groups and 1 7 new species

Alice Wells

111 > Descriptions of new species and a new genus of leptophlebiid mayflies (Insecta: Ephemeroptera)

from the Northern Territory, Australia

J. C. Dean and P. J. Suter

Volume 61 Number 2 2004

121 > Mitochondrial 1 2S rRNA sequences support the existence of a third species of freshwater blackfish

(Percicthyidae: Gadopsis) from South-eastern Australia

Adam D. Miller, Gretchen Waggy, Stephen G. Ryan and Christopher M. Austin

Continued inside back cover >

Contents