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Theses
Southern Cross University Year
Polydora and Dipolydora (Polychaeta:
Spionidae) of estuaries and bays of
subtropical eastern Australia: A review
and morphometric investigation of their
taxonomy and distribution
Lexie Margaret WalkerSouthern Cross University
This thesis is posted at ePublications@SCU.
http://epubs.scu.edu.au/theses/92
Walker, LM 2009, ‘Polydora and Dipolydora (Polychaeta: Spionidae) of estuaries and baysof subtropical eastern Australia: A review and morphometric investigation of their taxonomyand distribution’, MSc thesis, Southern Cross University, Lismore, NSW.
Copyright LM Walker 2009
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Polydora and Dipolydora (Polychaeta: Spionidae) of
estuaries and bays of subtropical eastern Australia: A
review and morphometric investigation of their
taxonomy and distribution.
Submitted by
Lexie Margaret Walker, B.Sc. (UNSW, 1983), Grad. Dip. Ed. (SCU, 1997)
to the
School of Environmental Science and Management
Southern Cross University
to satisfy the requirements for admission to the degree of
M.Sc. by Thesis
on
5th May, 2008
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STATEMENT
I certify that this is an original work wholly completed subsequent to my initial enrolment
in the course for which it is submitted and that the work has never previously been used
in seeking a degree at this university or any other university. To the best of my knowledge
it contains no material previously published or written by another person except where
due acknowledgement is made in the thesis. Any contribution made to the research by
others with whom I have worked at SCU or elsewhere is explicitly acknowledged in the
thesis. The intellectual content of this thesis is the product of my own work, except to the
extent that assistance from others in the project’s design and conception or in style,
presentation and linguistic expression is acknowledged.
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ABSTRACT
The aim of this thesis was to review the current state of knowledge, occurrence and
distribution of two polydorid (Polychaeta: Spionidae) genera, Polydora and Dipolydora, in
estuaries of subtropical eastern Australia. The existing taxonomy of the polydorid group is
confused and descriptions include many relative terms. A numerical taxonomy approach
using Primer 6 was taken to identify species groups and characters. A standardized set of
multivariate morphological characters was developed and resemblance analysis functions
used to create species cluster groups. SIMPER (similarity percentages) analysis on the
same dataset was used to identify the diagnostic characters for each of these species
cluster groups and to identify new characters which could be useful in species diagnosis,
particularly for frequently occurring incomplete specimens.
Prior to this study 3 Polydora species and 4 Dipolydora species were recorded from
subtropical eastern Australia. The present study found 12 Polydora species and 10
Dipolydora species from estuaries and bays of subtropical eastern Australia.
Two Polydora species are new (Polydora sp. P1 and Polydora cf. woodwicki) and 7 are
potentially new species (P. cf. latispinosa; P. cf. websteri; P. sp. P2S; P. sp. P3S; P. sp. P4S;
P. sp. P5S and P. sp. P6S) having been described from single specimens. Polydora cornuta
Bosc, 1902 is recorded from New South Wales for the first time. Polydora cf. calcarea is
reported from mudblisters in oysters. The Australian form of Polydora hoplura Claparède,
1870 and P. haswelli Blake and Kudenov, 1978 are described more fully than in existing
literature.
Three Dipolydora species are new (Dipolydora sp. D1; D. cf. flava and D. sp. D2) and 4
from single specimens are potentially new (D. cf. aciculata/ cf. giardi; D. sp. D3S; D. sp.
D4S and D. sp. D5S). Dipolydora tentaculata (Blake and Kudenov, 1978) and Australian
forms of Dipolydora flava (Claparède, 1870) and Dipolydora socialis (Schmarda, 1861) are
described more fully than in existing literature.
Prior to this study one Dipolydora and no Polydora were recorded from the Tweed-
Moreton bioregion. This study reports 6 species of Dipolydora and 5 species of Polydora
from this bioregion.
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Dipolydora penicillata (Hutchings and Rainer, 1979) and Carazziella victoriensis Blake and
Kudenov, 1978 are recommended for synonymy.
It is recommended that Polydora ciliata (Johnston, 1838) be reinstated and Dipolydora
ciliata (Johnson, 1838) removed from the Australian polychaete checklist following
identification of an error in description translation.
Important gaps in collections were identified for polydorids associated with oysters from
estuaries over the whole subtropical region; and for polydorids from the Tweed-Moreton
IMCRA bioregion.
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Contents CHAPTER 1: INTRODUCTION .............................................................................................................. 7
History of polydorid collection in Australia .................................................................................... 9
Currently recorded Australian polydorid species......................................................................... 15
Polydora and Dipolydora in Australia ........................................................................................... 16
The Dipolydora penicillata problem ......................................................................................... 18
The Dipolydora ciliata problem ................................................................................................ 18
Polydorid taxonomy ..................................................................................................................... 19
The Australian and east coast subtropical climate zones ............................................................ 27
Terrestrial ................................................................................................................................. 27
Marine waters, bioregions and estuaries of the subtropical east coast of Australia .............. 29
The aim of this review .................................................................................................................. 32
CHAPTER 2: MATERIALS AND METHODS ......................................................................................... 33
Materials ...................................................................................................................................... 33
Data collection ............................................................................................................................. 34
Re-identification of material .................................................................................................... 36
Analysis ......................................................................................................................................... 40
Analysis package....................................................................................................................... 40
Dataset preparation for analysis .............................................................................................. 41
Part 1: Resemblance between specimens and position of specimens of questionable identity . 42
Part 2: Final analysis and identifications ...................................................................................... 44
CHAPTER 3: RESULTS ........................................................................................................................ 45
Part 1: Resemblance between specimens and position of specimens of questionable identity . 45
Species clusters ........................................................................................................................ 47
SIMPER analysis ........................................................................................................................ 58
Conclusions on the identity of the re-identified material ........................................................ 60
Part 2: Final analysis and identifications ...................................................................................... 68
Species clusters – Polydora and Dipolydora ............................................................................. 68
Species clusters: Polydora ........................................................................................................ 70
Species clusters: Dipolydora ..................................................................................................... 72
SIMPER analysis of final dataset - Resemblance between Dipolydora and Polydora .............. 74
SIMPER analysis – between Polydora species .......................................................................... 75
SIMPER analysis – between Dipolydora species ...................................................................... 79
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CHAPTER 4: Descriptions of Polydora and Dipolydora species ........................................................ 86
Polydora Bosc, 1802 ................................................................................................................. 86
Dipolydora Verrill, 1879 ......................................................................................................... 119
Distributions of Polydora and Dipolydora in estuaries of subtropical eastern Australia ........... 151
CHAPTER 5: DISCUSSION ................................................................................................................ 154
REFERENCES ................................................................................................................................... 166
Appendices ..................................................................................................................................... 180
Acknowledgements
I wish to thank and acknowledge the many people who have given wise guidance,
support, care and inspirational conversation through the course of this research: Dr Pat
Hutchings, Anna Murray, collection managers and technical staff of the Australian
Museum, Dr Chris Glasby of Museum and Art Gallery of the Northern Territory, Dr Daniel
Bucher, Dr Alison Specht and technical staff at Southern Cross University; Dr Robin Wilson
and technical staff at Museum Victoria; Dr Mal Bryant and technical staff at the
Queensland Museum; Dr Matthew McArthur; Dr Geoff Read; Dr Wayne O’Connor and Dr
Tim Glasby (NSW DPI Fisheries) also thanks to all my family and caring friends... truly
appreciated...and Peter, who turned the lights on..
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The Problem
A recent desktop report to NSW Fisheries (Walker, 2005) on the potential for
translocation of marine pest species between subtropical coastal bioregions in QLD and
NSW found that there were many unidentified specimens of polydorids (Polychaeta:
Spionidae) in museum collections within Australia. Whether these were pest species,
poorly identified species or species new to science was unknown. In addition to this,
unusual and disjoint distribution patterns were noted for some species. Possible
explanations for this include variable collecting effort, targeted research collecting,
misidentification, translocation events, geographical barriers, past geological events or
other natural causes. As these collections are the main source of information on marine
biodiversity in Australia, and the polydorids are known world-wide to include serious pest
species, it was considered important for socio-economic and environmental reasons to
investigate this further. A starting point for this investigation is to clarify exactly:
what do we already know?
what is in museum collections?
what are the apparent distribution patterns?
where are the knowledge and collection gaps?
for the polydorids in subtropical eastern Australia. This thesis aims to begin to answer
these questions by focusing on the two most species-rich polydorid genera, Polydora and
Dipolydora. Ongoing research will consider the remaining polydorid genera known to
occur in Australia: Boccardia, Pseudopolydora, Carazziella, Boccardiella and Polydorella.
CHAPTER 1: INTRODUCTION
Polydorids are a group within the subfamily Spioninae of the family Spionidae, one of the
largest families in the class Polychaeta. Polydorids occur as tube-builders in soft
sediments or associated with, or boring into, calcareous substrates (Blake, 1996d) such as
mollusc shells, calcareous algae, corals or bryozoans (Blake and Evans, 1973). Some
species, such as Dipolydora socialis, are reported as both tube-building in sediments and
as boring into molluscs (Blake, 1971). Several polydorid species are considered pests of
commercially farmed molluscs including oysters, mussels and abalone. In Australia, four
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polydorid species that occur with Saccostrea commercialis, the Sydney Rock Oyster, cause
mud blisters inside the valves of the oyster making them unsuitable for retail (Nell, 2001).
Some muddy substrate dwellers, such as Dipolydora socialis, can reproduce rapidly, the
population developing into a raft of tubes causing physical alteration of the surrounding
environment (Blake, 1971). Polydorids are classified as interface feeders (Dauer et al.
1981), being either surface deposit or suspension feeders depending on water quality.
Reproduction is asexual or, more commonly, sexual. In polydorids, larval development
occurs by brooding in the tube with or without nurse eggs; or by broadcasting larvae at
various stages of development into the plankton where they become planktotrophic
(Blake and Arnovsky, 1999). In some species either or both of these strategies may be
used depending on environmental conditions (Blake, 1996d). Reproductive characters for
Australian Polydora and Dipolydora species are listed in Appendix 1. The flexibility of
polydorids in habit, feeding and reproduction has probably played a part in their
widespread distribution and abundance in estuarine systems around the world (Blake,
1996d).
Woodwick (1963a) described the polydorids as “spionids in which a modified fifth
segment contains enlarged and specialized setae”. Blake (1996d), in his revision of
Polydora, considered the degree of modification of fifth setiger, structure and number of
major spines in setiger 5, first appearance of branchiae and, to a lesser extent, structure
and first appearance of hooded hooks important characters in distinguishing genera of
polydorids. More recently Radashevsky and Hsieh (2000) questioned the use of
“modification of segment 5” as a principal diagnostic feature of the polydorid complex
given the ambiguity of the term and the morphology of some Pseudopolydora species
where neighbouring segments are morphologically similar to segment 5. They
recommended the use of consistent and clearly defined characters such as modification
of post-chaetal lamellae and characteristics of each group of noto- and neurochaetae
(anterior, posterior, and superior or inferior) on chaetiger 5 as described by Mesnil
(1896), Söderström (1920) and Radashevsky and Fauchald (2000). This type of data would
be suitable for cladistic analyses allowing evolutionary relationships between polydorid
genera and other spionids to be investigated and synapomorphies for the group to be
identified. There are no published cladistic analyses of the entire polydorid group but a
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preliminary cladistic analysis of polydorids by Williams (unpublished work from PhD
thesis 2000 in Williams, 2004) concluded that most previous classifications of Boccardia,
Carazziella, Dipolydora, Polydora and Tripolydora were supported.
Worldwide there are currently 9 recognized polydorid genera. In the most recent revision
of Polydora, Blake (1996d) resurrected the genus Dipolydora Verrill, 1879 (formerly
contained within Polydora) and split both into a further 5 species groups based on habitat
and morphological features. Blake (1978; 1996d) also considers that Polydorella should
be placed within Pseudopolydora. Polydorella, originally described from Australia
(Augener, 1914), was synonymized with Pseudopolydora in the most recent revision of
the Australian Spionidae (Blake and Kudenov 1978). However, the genus has since been
resurrected (Tzetlin and Britayev, 1985) and, in addition to the two Australian species,
three new species have been described from areas outside Australia (Tzetlin and Britayev,
1985; Radashevsky, 1996; Williams, 2004). There are currently a total of 147 recognized
species (Appendix 2) within the 9 polydorid genera as follows: Amphipolydora (2 species),
Tripolydora (1 species), Polydorella (5 species), Boccardiella (7 species), Carazziella (13
species), Pseudopolydora (18 species), Boccardia (21 species), Dipolydora (36 species) and
Polydora (44 species).
History of polydorid collection in Australia
The first record of polydorid spionids in Australia was made in 1885 by William Haswell,
at that time lecturer in Zoology and Comparative Anatomy at Sydney University. The
animals were found in mud-filled blisters on the inner surface of oyster shells from
Hunter River oyster beds in New South Wales where oysters where “dying in large
numbers owing to the attacks of some parasite” (Haswell 1885). At this time 24 species
of Polydora had been described from the northern hemisphere, as Polydora, Leucodore,
Leucodorum or Spio, [Bosc, 1802(east coast North America) in Blake, 1996d; Templeton,
1836 (Ireland); Johnston, 1838 (England); Oersted, 1843 (Denmark) in Blake, 1996d;
Grube, 1855 (Germany);1878 (Philippines) in Blake, 1996d; Quatrefages, 1865 (France) in
Blake, 1996d; Claparède, 1869;1870 (Italy); Verrill 1879;1880;1881(east coast North
America); Webster, 1879a; 1879b (east coast North America); Jacobi, 1883(Germany) in
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Blake, 1996d) and one from the southern hemisphere (Schmarda, 1861 (Chile)]. Haswell
cautiously identified the abundant organism in the blisters as Polydora ciliata “identical
with the European Polydora ciliata of Johnston”. He thought this “strange” and noted
that reports of P. ciliata (as Leucodore ciliata) from other parts of Europe generally did
not indicate a “shell-invading habit” and that in those reports of P. ciliata occurring in
various mollusc shells there was no mention of the same amount of destruction as
caused by the animal in the Hunter River. Although the description of the mud-blister
damage in his report was quite detailed, he gave no description of the animal itself.
Several specimens identified as P. ciliata remain in the collection of the Australian
Museum, Sydney, which appear to be from Haswell’s collection, although no definite
information to that effect exists. One specimen of a new species, Polydora (Leucodore)
polybranchia was described from the oyster mud-blisters (Haswell 1885) but no types
were designated. The review of Polydora by Carazzi (1895) referred Polydora (Leucodore)
polybranchia Haswell, 1885 to Polydora (Boccardia) polybranchia. Whitelegge (1890)
included Johnston’s 1838 description of Polydora ciliata (as Leucodore ciliatus) and a
description of the eggs and appearance of the reproductive animal in his report of P.
ciliata occurring in diseased farmed oysters with mudblisters.
Augener (1914), a German polychaete expert, described Polydorella prolifera n. gen., n.
sp. from south-western Australia and also recorded, Polydora armata Langerhans, 1880
from the collection of a German research expedition to south-west Australia. Type
material of Augener is held in the Zoologisches Museum der Universitat, Hamburg,
Germany. The following 60 years saw no new polydorid species described from Australia
and very few collections or reports made. Reports regarding Polydora ciliata in mudworm
continued (Roughley, 1922; 1925; Wilson, 1928). Wilson (1928) described the larvae of
Polydora ciliata and P. hoplura from blisters in oysters from the River Yealm, England and
discussed Whitelegge’s (1890) description of Polydora ciliata eggs from New South Wales
suggesting that, although poorly illustrated, they were more like those described by
Söderström (1920) of Polydora ligni.
As environmental research activity increased in the late 1960s more information became
available on the distribution and diversity of polydorid species. In Victoria, the Port Phillip
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Bay Environmental Survey commenced in 1969. The Marine Pollution Studies Group
within the Fisheries and Wildlife Division of the Ministry of Conservation in Victoria
studied the Port Phillip Bay benthos initially recording one Boccardia sp., eleven Polydora
spp. and Pseudopolydora kempi (Poore et al., 1975). Later, Polydora socialis,
Pseudopolydora paucibranchiata and Boccardia limnicola were identified from near the
outfall of Werribee sewerage treatment farm (Poore and Kudenov, 1978a) and “Polydora
species” from the Port of Melbourne (Poore and Kudenov, 1978b). Blake and Woodwick
(1976) described Boccardia limnicola n. sp. from the freshwater Lake Bong Bong. In New
South Wales, Hutchings and Recher (1974) described the fauna and ecology of Careel
Bay, Pittwater, after a survey of the area through 1972-1973 by the New South Wales
Division of the Australian Littoral Society. Two polydorids, Boccardia sp. and Polydora sp
were recorded. In 1977, Margaret Skeel from the Brackish Water Fish Culture Research
Station at Port Stephens made important collections of “mudworms” from diseased
oysters in New South Wales, Victoria and Tasmania. These were reported as Polydora n.
sp. and Boccardia (Skeel, 1977); Polydora Websteri (sic) (Skeel, 1978); and Polydora
websteri, Polydora haswelli, Polydora hoplura and Boccardia chilensis (Skeel, 1979). Skeel
(1979) described the reproductive strategies of these four polydorids in Australia and
compared these with reports of Polydora websteri from North America, Polydora hoplura
and P. ciliata from Europe (Appendix 1). In Queensland, the Gladstone Environmental
Survey was commissioned by the Queensland Electricity Commission. The survey ran
from 1974-1983. The reference collection checklist (Saenger, 1988) includes 10 polydorid
species: Polydora cf. flava, P. socialis, P. cf. websteri, P. sp. I, P. sp. II, P. sp. III,
Pseudopolydora kempi, Pseudopolydora paucibranchiata, Pseudopolydora sp. I and
Pseudopolydora sp. II. Investigations into the complexity of the benthic biota near Peel
Island in Moreton Bay by researchers from Queensland University and the C.S.I.R.O.,
Division of Tropical Agronomy occurred from March 1970 - March 1971. One Polydora sp.
and three unidentified F. Spionidae were found (Stephenson et al., 1974). In 1972-1974
Moreton Bay macrobenthos at Bramble Bay and Middle Banks were surveyed prior to
land reclamation and sand dredging in the redevelopment of Brisbane airport. Polydora
sp. 1 was recorded at Bramble Bay (Stephenson et al., 1976) and Middle Banks
(Stephenson et al., 1978).
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In 1978, James Blake, from the University of the Pacific, California, and Jerry Kudenov,
from the Marine Studies Group, Ministry for Conservation, Victoria, published the first
account of the Spionidae of south-eastern Australia including a revision of the genera of
the Polydora-complex worldwide. Material was examined from new collections made
through 1977, existing collections and previous surveys in south-eastern Australia. The
south-eastern region was defined as south of Moreton Bay, Queensland, excluding the
material from the Gladstone survey. Two-thirds of the south-eastern Australian Spionidae
species were found to be endemic. Twenty-nine polydorid species in six genera were
recognized: Boccardia (3), Boccardiella gen. nov. (2), Pseudopolydora (5), Polydora (15),
Carazziella gen. nov. (4) and Tripolydora (1). Tripolydora does not occur in Australia but
was included as part of the worldwide generic revision. Polydorella was synonymized with
Pseudopolydora (Blake and Kudenov, 1978). Records of Polydora ciliata were considered
to be the North American pest species, Polydora websteri Hartman in Loosenoff and
Engle, 1943 (Blake and Kudenov, 1978). Fifteen new species were described: Boccardiella
(B. bihamata), Pseudopolydora (P. glandulosa, P. stolonifera), Polydora (P. protuberata, P.
tentaculata, P. aciculata, P. notialis, P. pilocollaris, P. haswelli, P. latispinosa, P.
woodwicki) and Carazziella (C. victoriensis, C. phillipensis, C. hymenobranchia, C.
hirsutiseta) (Blake and Kudenov, 1978). Cosmopolitan species listed were: Polydora flava,
P. socialis, P. giardi, P. armata, P. ligni, P. hoplura, Pseudopolydora kempi,
Pseudopolydora paucibranchiata, Boccardia polybranchia and from the southern
hemisphere, Boccardia chilensis. Pseudopolydora paucibranchiata was considered a
possible introduced species. Boccardia proboscidea was recorded for the first time from
the southern hemisphere, later being reported as a possible introduction by Blake and
Kudenov (Blake and Kudenov, 1981). Petch (1995) found that worldwide populations of
Boccardia proboscidea from temperate waters including Australia and North America
could not be separated based on morphological features.
Reproduction and larval development in the Australian Boccardia chilensis and B.
proboscidea was investigated in Australia by Blake and Kudenov (1981). Blake and
Kudenov (1981), Skeel (1979) and Whitelegge (1890) appear to be the only published
research on reproduction and larval development of Australian polydorids.
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Since the publication of Blake and Kudenov (1978) two new polydorid species have been
described from Australia. Hutchings and Rainer (1979) described Polydora penicillata n.
sp. from the Careel Bay fauna in New South Wales. Hutchings and Turvey (1984) recorded
Boccardia fleckera n. sp., plus an additional two Pseudopolydora spp. described only to
genus, following extensive collecting in South Australia in 1979. Between August 1975
and March 1976, Hartmann-Schröder, from the Zoologisches Museum der Universität,
Hamburg, Germany, collected polychaetes from intertidal sandy substrates along much of
the coast of Australia. The survey commenced in Derby, Western Australia, travelling
down the west coast, across the south coast and up the east coast to Gladstone,
Queensland also including Heron Island on the Great Barrier Reef. Fifty-four stations were
sampled and the material returned to Hamburg, Germany for examination. A series of
publications documented the expedition and its findings (Hartmann-Schröder, 1979;
1980; 1981; 1982; 1983; 1984; 1985; 1986; 1987; 1989; 1990; 1991). No new species of
polydorids were found although several undetermined incomplete Polydora sp. were
recorded. New records for Australia included Polydora (Polydora) ciliata (Johnston, 1838)
from Exmouth, Western Australia (Hartmann-Schröder, 1979) and Polydora
(Pseudopolydora) antennata Claparède, 1870 recorded from the west and south coast
(Hartmann- Schröder, 1981; 1983; 1986). No polydorid material from these collections is
located in Australia. The Hawkesbury River Survey of the Australian Museum Marine
Ecology Department (1977-1981) recorded 10 polydorid species (Hutchings and Murray,
1984), six of these were endemic species described by Blake and Kudenov (1978).
In mid-1990, a growing awareness and concern regarding introduced marine pests
resulted in a collaboration between the Australian Association of Port and Marine
Authorities, CSIRO Centre for Research on Introduced Marine Pests (CRIMP), state and
territory agencies and NSW Fisheries developed with the aim of establishing baseline
data on the distribution and abundance of selected marine pest species. A standardized
protocol to monitor ports for known and potential marine pest species (Hewett and
Martin, 1996; 2001) was designed and the Australian Port Survey program began. By
2004, 41 ports around Australia had been surveyed at an estimated cost of AUD$10
million (Hutchings and Glasby, 2004) including those of Eden in 1996 and 1999-2002
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(Pollard and Rankin, 2003), Newcastle in 1997, Botany Bay in 1998 (Pollard and
Pethebridge, 2002a), Port Kembla in 2000 (Pollard and Pethebridge, 2002) and Sydney
Harbour in 2001(AMBS 2002). The protocol was designed to target particular pest
species, but survey reports indicate that much other useful material has been collected,
including non-pest polydorid species. This material was lodged with museums and other
government agencies, consulting companies and universities. A proposal that all material
should eventually be lodged in museums to be properly curated making it accessible for
taxonomic research into the future (Hutchings and Glasby, 2004) has come into effect
(Hutchings, pers. comm.) although some material could not be located or was in poor
condition and had to be discarded (C. Glasby, pers. comm.). Funding supplied by the
National Heritage Trust and the AMIT was given to each of the state museums for
incorporation and checking of pest species.
Severe disease events in abalone culture facilities in Tasmania occurred throughout 1995-
1997 and 1998- 2000. These are described in Lleonart et al. (2003) who record Boccardia
knoxi, previously only known from New Zealand, and Polydora hoplura from the
mudblisters. Unfortunately, it appears that no specimens were lodged with museums and
the record remains unverified. McDiarmid et al. (2004) investigated the ecology of
polychaetes infesting wild abalone shells in Victoria. Eleven polydorid species were found
and specimens lodged at Museum Victoria. Known taxa included Boccardia chilensis,
Dipolydora armata, Polydora giardi and Polydora woodwicki. Undescribed taxa were
given Museum of Victoria species codes and included Boccardia MoV 3833, Dipolydora
MoV 3834, D. MoV 3835, D. MoV 3836, D. MoV 3838, Polydora MoV 3842 and
Pseudopolydora MoV 3837.
Oyster culture, mudworm disease and culture techniques which can limit mudworm
growth in farmed oysters has been well documented by scientists from NSW Department
of Primary Industries, Port Stephens Fishery Centre (Nell and Smith, 1988; Nell, 2001;
2007a; 2007b). Nell (2001) and Ogburn et al. (2007) suggest that importation of New
Zealand rock oysters to replenish depleted oyster beds in the 1880s brought the disease
causing mudworms into estuaries in New South Wales and southern Queensland. Nell
(2007b) records Polydora websteri, Polydora hoplura and Boccardia chilensis as the main
15
disease causing agents in NSW and southern Queensland oysters and Boccardia knoxi in
Pacific oysters grown sub-tidally in Tasmania.
Currently recorded Australian polydorid species
In the most recent checklist of Australian polychaetes, 33 polydorid species are
recognized (Hutchings and Johnston 2003b) (Table 1; Appendix 3). Hutchings and
Johnston (2003a), in Wilson et al. (2003), list 5 genera, the recently separated (Blake
1996d) Polydora and Dipolydora species remaining amalgamated under Polydora.
Hutchings and Johnston (2003b) list the 33 polydorid species in 6 genera following Blake
(1996d) as follows: Boccardiella (2 species), Carazziella (4 species), Pseudopolydora (6
species), Boccardia (4 species), Dipolydora (11 species) and Polydora (6 species).This
checklist includes Polydorella within Pseudopolydora after Blake (1978; 1996d). There are
no records of Amphipolydora or Tripolydora from Australia. The interactive key of Wilson
et al. (2003) includes 49 Australian polydorid species in 7 genera: Polydorella (1 species),
including Polydorella stolonifera after Radashevsky (1996) but placing Polydorella
prolifera, the type species for the genus, within Pseudopolydora (sensu Blake and
Kudenov, 1978), Boccardiella (4 species), Carazziella (4 species), Pseudopolydora (10
species), Dipolydora (17 species) and Polydora (8 species). Additional to the checklist of
Hutchings and Johnston (2003a) are 12 species with MoV (Museum of Victoria) species
codes (Boccardia (1), Boccardiella (2), Dipolydora (5), Polydora (1) and Pseudopolydora
(3)), two Pseudopolydora identified as sp. 1 and sp. 2, Polydora cf. ligni, Dipolydora cf.
ciliata and D. cf. pilocollaris.
Three polydorid species are listed as introduced marine species to Australian waters
(NIMPIS 2002): Boccardia proboscidea (Hartman, 1940) from California, Polydora ciliata
Johnston, 1838 from Great Britain, and Pseudopolydora paucibranchiata, Okuda, 1937
from Japan. Hayes et al. (2005) list Polydora websteri, Polydora cornuta, Boccardia
proboscidea and Pseudopolydora paucibranchiata as pests of low impact and low
invasion potential relative to other non-native domestic marine pest species. The
Australian Museum port survey of Sydney Harbour final report (Australian Museum
Business Services, 2002) recommends that Boccardia chilensis Blake and Woodwick, 1971
originally described from Chile also be included as a known introduced marine pest.
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Table 1: Number of taxa in polydorid genera worldwide, from the most recent Australian
revisions and checklists and from subtropical east coast estuaries *research currently in
progress indicates that additional species will be found
Number of taxa
Genus
Worldwide
Australia
Subtropical east coast Australian estuaries
Blake and Kudenov (1978)
Hutchings and Johnston (2003a)
Wilson et al. (2003)
ABRS Fauna online 2008 (Hutchings and Johnston 2003b)
this study
Polydora 44 15 17 8 6 12
Dipolydora 36 0 0 17 11 10
Boccardia 21 3 4 5 4 1*
Pseudopolydora 18 5 6 10 6 3*
Carazziella 13 4 4 4 4 2*
Boccardiella 7 2 2 4 2 1*
Polydorella 5 0 0 1 0 0
Amphipolydora 2 0 0 0 0 0
Tripolydora 1 0 0 0 0 0
Total number 147 29 33 49 33 29*
Polydora and Dipolydora in Australia
The most recent revision of Polydora species worldwide (Blake, 1996d) breaks both
Polydora and Dipolydora into 5 groups based on morphology and habitat. Australian
representatives in each group are listed in Tables 2 and 3 for Polydora and Dipolydora
respectively.
17
Table 2: Polydora species currently recognized in Australia grouped according to the
latest revision by Blake (1996d).
Group Australian records
A. Polydora cornuta/nuchalis group
P. cornuta Bosc, 1802
B. Polydora ciliata/websteri group
P. haswelli Blake and Kudenov, 1978
P. hoplura Claparède, 1870
P. latispinosa Blake and Kudenov, 1978
P. websteri Hartman, 1943
P. woodwicki Blake and Kudenov, 1978
C. Polydora alloporis group
No records to date for this group in Australia
D. Polydora bioccipitalis group
No records to date for this group in Australia
E. Polydora colonia/spongicola group No records to date for this group in Australia
Table 3: Dipolydora species currently recognized in Australia grouped according to the
latest revision by Blake (1996d).
Group Australian records
A. Dipolydora giardi group
D. giardi (Mesnil, 1896)
B. Dipolydora concharum/coeca/flava/socialis group
D. flava (Claparède, 1870)
D. protuberata (Blake and Kudenov, 1978)
D. tentaculata (Blake and Kudenov, 1978)
D. socialis (Schmarda, 1861)
C. Dipolydora barbilla/bidentata group
D. pilocollaris (Blake and Kudenov, 1978)
D. Dipolydora armata group
D. aciculata (Blake and Kudenov, 1978)
D. armata (Langerhans, 1880)
D. notialis (Blake and Kudenov, 1978) (D. notialis is poorly known (Blake, 1996d))
E. Dipolydora commensalis group
No records to date from Australia
18
No new species of Polydora or Dipolydora have been described from Australia since the
publication of Blake (1996d), although Wilson et al. (2003) include Polydora cf. ligni,
Dipolydora cf. pilocollaris and an additional Polydora and five Dipolydora named with
MoV codes and associated voucher material.
Two species of Dipolydora, Dipolydora penicillata (Hutchings and Rainer, 1979) and
Dipolydora ciliata (Johnston, 1838) [=Leucodore ciliatus Johnston, 1838], are included in
the current Australian checklist (Hutchings and Johnston, 2003b) but are not mentioned
in Blake’s revision of Polydora (Blake, 1996d).
The Dipolydora penicillata problem
Wilson et al. (2003) in the formatted description notes for Dipolydora penicillata, derived
from the original description in Hutchings and Rainer (1979) as Polydora penicillata,
noted that the description does not comply with the generally accepted definitions for
Polydora or Dipolydora in recent revisions (Blake and Kudenov, 1978; Blake, 1996). The
results of the present study suggest that Dipolydora penicillata (Hutchings and Rainer,
1979) is synonymous with Carazziella victoriensis Blake and Kudenov, 1978.
The Dipolydora ciliata problem
Specimens recently referred to as Dipolydora cf. ciliata in the interactive key of Wilson et
al. (2003) and Dipolydora ciliata Johnston, 1838 in the current ABRS checklist (Hutchings
and Johnston 2003b) were described by Hartmann-Schröder (1979) as Polydora
(Polydora) ciliata, from Broome in Western Australia. This description in German
(Hartmann-Schröder, 1979) is brief and does not include figures. Only the description of
the pygidium gives any indication of placement into either Polydora or Dipolydora after
the revision of Polydora by Blake (1996d). Polydora species as currently defined by Blake
(1996d) have pygidia that are “saucer-shaped or disc-like, border usually entire except for
dorsal gap.” while species of Dipolydora, as defined by Blake (1996d), have pygidia that
are “disc-like, cuff-shaped, with 2, 3, or 4 lobes of various forms, or with 4 or more small
papillae”. The description of P. (P.) ciliata by Hartmann-Schröder (1979) describes the
pygidia of the specimens as “Pygid mit trichterformigem [sic] Lappen, der mediodorsal
eingeschnitten ist”. This can be translated as, “pygidium a funnel-shaped lobe with
mediodorsal incision” (using Babylon (Dallakian and Yanyuk, 2004-2006)). Wilson et al.
19
(2003) describe the pygidium of Dipolydora cf. ciliata, from the same Hartmann-Schröder
(1979) description in German, as “3-lobed with mid- dorsal incision”. This error in
translation mistakenly places the specimens in Dipolydora as defined by Blake (1996d).
The original identification of these specimens as Polydora appears to be correct and
Polydora ciliata should be reinstated in the Australian polychaete checklist.
Blake (1996d) includes Polydora ciliata (Johnston, 1838) in the Polydora ciliata/websteri
Group but notes that Polydora ciliata has been recorded from widespread localities and
that it is possible that these records are actually of several species. This suspicion is
supported and confirmed by genetic and morphological studies (Mustaquim, 1986; 1988;
Manchenko and Radashevsky, 1993; 1998; Radashevsky and Pankova, 2006).
Radashevsky and Pankova (2006) reinstated Polydora calcarea (Templeton, 1836) to
include the boring form recorded by them as P. cf. ciliata (Manchenko and Radashevsky,
1993) from the Sea of Japan and several other early records of boring forms of P. ciliata.
They concluded that, world-wide, current records of P. ciliata described as boring should
be verified. In Australia, where specimens were available, Blake and Kudenov (1978)
referred early records of P. ciliata boring into oysters from the Hunter River, New South
Wales (Haswell, 1885 [not Johnston, 1838]) to Polydora websteri as discussed later in this
thesis. Where only a written record was available (Whitelegge, 1890; Roughley, 1922)
verification of the occurrence of P. ciliata in New South Wales at that time was, and still
is, impossible. The P. ciliata specimens recorded from Broome, Western Australia by
Hartmann-Schröder (1979) were non-boring, being collected from fine sand on tidal
mudflats. As such, the identity of those specimens should remain as Polydora ciliata
(Johnston 1838) and await verification.
Polydorid taxonomy
Since the work of Linnaeus just 250 years ago, the science, and art, of taxonomy has been
influenced by many technological and communications advancements. The problems of
polydorid taxonomy in Australia reflect this situation. Rapid changes in technology in the
fields of communications, biology, microscopy and analysis have placed in stark contrast
the work of early taxonomists with those of the present. Yet, reference back to these
early works and collections is still an essential part of the taxonomic process. As new
20
species are discovered, more information is produced and the types of information
available changes it becomes increasingly difficult to follow this process of reference to
literature and examination of voucher specimens from the past. Descriptions, including
original descriptions, are often brief, eg. Dipolydora flava (Claparède, 1870) and written
in languages other than English, eg. Polydora hoplura Claparède, 1870. Type material can
be difficult to access and is sometimes lodged in collections outside Australia, eg.
Polydorella prolifera Augener, 1914 type is lodged in Zoologisches Museum der
Universität, Hamburg, Germany. As type material ages and is examined by various
researchers the condition of the specimens may deteriorate. Occasionally, type material
may not be deposited at all as is the case with Boccardia polybranchia Haswell, 1885 or
may have been lost so that no reference specimen exists to enable comparison with
potential new species. Without type material and a detailed description it is impossible to
confirm or deny the validity of an identification. In these cases a species name may be
declared as unknown, a “nomen dubium”.
Improved communications and ease of access to literature has made it possible to
complete detailed reviews and comparison of descriptions of species collected from
wide-spread locations in a relatively short period of time. Where morphological variation
is great and observable this should help alleviate the confusion caused by multiple
descriptions of the same species under different names. Radashevsky (2005) redescribed
Polydora cornuta including adult and larval morphology using material from temperate
and subtropical zones worldwide concluding that morphological variation in the species
reflected size dependent characters in one species rather than different sibling species.
Advances in biotechnology, particularly in molecular biology, have been useful in
separating species within groups whose members are morphologically very similar such
as the Polydora websteri/ciliata Group (Blake, 1996d). Polydora ciliata is often reported
as boring into calcareous substrates but is also reported as tube building in muddy
sediments. It has recently been shown, using morphological and molecular methods, that
the almost identical boring and the non-boring forms of Polydora ciliata are in fact two
different species (Radashevsky and Pankova, 2006) indicating that Polydora ciliata may
not be as widespread as has been reported in the past. Confirmation of genetic
21
uniqueness can also lead to more detailed examination of the morphology and habit of
the species in question giving validity to characters which may have been previously
overlooked or considered unimportant.
The polydorids are generally considered difficult to identify. Part of the problem is that
published descriptions and identification keys rely heavily on the presence or absence of
a few particular characters. For Dipolydora and Polydora these basic elements are
posterior spines or needle-packets, pygidial characters and the type of accessory
structures on the chaetiger 5 modified spines. Animals are often collected without
posteriors and chaetiger 5 spines are known to wear and be inconsistent in appearance
(Kendall, 1980). Specimens without a posterior end and with worn accessory structures
on the chaetiger 5 modified spines can be impossible to identify confidently using existing
keys. As microscopical techniques have improved less obvious characters such as palp
morphology (Worsaae, 2001) and various ciliation patterns (Jelsing, 2003; Purshke and
Hausen, 2007) have been suggested as being potentially useful in systematics. Further
problems exist in descriptions of features using relative terms such as “small” and “large”
which lack meaning to a taxonomist unfamiliar with the organism being described.
Characters that could be quantified, such as number of capillary chaetae accompanying
hooded hooks on the neuropodia, are rarely enumerated often being simply described as
“present”. This is a loss of potentially useful information.
Multivariate methods in taxonomy were first developed and used by Fischer (1936).
Increased computing power and novel statistical analysis software has overcome the
problem of applying the required complex and repetitive computations so that the use of
multivariate morphometric techniques to assess similarities and establish groups based
on multivariate characters is becoming more common (Petch, 1995; MacCord and
Amaral, 2005; Ford and Hutchings, 2005; Garraffoni and Comargo, 2006; Glasby and
Glasby, 2006). Use of statistical techniques in taxonomy demands standardized collection
of character data, provides accurate assessment and comparison of variations in
measurable characters of specimens and can indicate which characters are important in
the resemblance between species or groups. Many specimens and many character
variables can be analysed and compared simultaneously. This frees the taxonomist from
22
making potentially inaccurate value-based judgments regarding the importance of
particular characters or of the similarity between specimens. It may also confirm the
judgments made in the work of early taxonomists.
Information on the geographical distribution of a species is often very useful in taxonomy.
For polydorids in Australia, and possibly elsewhere, this is not the case. Many polydorid
species are adaptable in habitat occupied, feeding and reproductive strategy. They occur
as fouling organisms of submerged marine objects or vessels that have accumulated
sediment and as epifauna of molluscs, including commercially farmed oysters, molluscs
and abalone. This adaptability combined with the increased movement of marine vessels
and the intentional transportation of bivalve stock globally and nationally (Ogburn, 2007)
over the last 100-150 years has meant that polydorids have potentially been accidentally
translocated around the globe. Collection of taxonomic information on polydorids and
the increased movement of these vessels have developed simultaneously. As most of
these vessel and stock movements have not been recorded in any systematic fashion, the
country of origin, or region of origin in Australia, of many of the polydorid species is really
an unknown. Several factors exacerbate this problem. Records of widespread polydorid
species in Australia have been made based on descriptions from the literature rather
than by comparison with type material or specimens from the type location. When
reviews and redescriptions of frequently reported, widespread “cosmopolitan” species
occur Australian material is rarely included. Australian polydorid material has been
examined only twice in reviews and redescriptions of polydorid species (Petch, 1995;
Williams, 2004). Williams (2004) reviewed reproduction and morphology in the poorly
known genus Polydorella. Petch (1995) used morphometric methods to examine the pest
species, Boccardia proboscidea, from worldwide locations finding no difference between
populations from temperate climates. Redescriptions of widespread Dipolydora and
Polydora species have never included Australian material (Table 2). Morphological and
molecular investigations of global populations could clarify if widely distributed species
are from the same population but the geographical origin of that population would still
remain unresolved. Until further knowledge is available it may be appropriate to consider
all species of polydorids originally described from Australia as native, or endemic, to
23
Australia and all other polydorid species recorded as occurring in Australia as cryptogenic,
i.e. neither demonstratively native nor introduced (Hilliard, 2004).
Table 4: Polydora species recorded as occurring in Australia, type locations, recent
redescriptions and distributions (excluding Polydora penicillata Hutchings and Rainer,
1979 = Carazziella victoriensis Blake and Kudenov, 1978)
Species Type location
Type material condition
Most recent redescription
Location of most recently described material
Recorded distribution
Australian material included in redescription
Polydora ciliata (Johnston, 1838)
Berwick Bay, England; soft sediments in rock crevices
Unknown.
No record of a type being deposited
Johnston (1838) – non-boring form
North and south Atlantic; Australia (Western Australia)
No
Polydora cornuta Bosc., 1802 (= P. ligni Webster, 1880)
Charleston Harbour, South Carolina, USA
Lost
Neotype (USNM 98587) designated by Blake and Maciolek (1987)
Radashevsky (2005)
Canada Atlantic coast, USA Atlantic Coast, USA Pacific coast, Gulf of Mexico, Brazil, Argentina, Germany (North Sea), Romania (Black Sea), Russia (Sea of Japan), Korea (Yellow Sea),Taiwan (Taiwan Province), off Mainland China (Fuchien Province)
East west, and gulf coasts of North America; Caribbean Sea; Argentina; northern and southern Europe; Australia (Victoria, New South Wales (this study); China; Korea; Japan; India; Brazil; the gulf coast of Mexico; Pacific coast of Russia (Radashevsky (2005))
No
24
Polydora haswelli Blake and Kudenov, 1978
Nth Chinaman’s Beach, Sydney Harbour, Australia
Good
AM W7283
1. Blake and Kudenov (1978)
2.Radashevsky, Lana et al. (2006) describe P. cf. haswelli
1. Australia (New South Wales)
2. Brazil
Australia (New South Wales); New Zealand (Read, unpublished)
Brazil (P. cf. haswelli)
Yes, endemic
2. No, but comments included on basis of description in Blake and Kudenov (1978) and pers. comm. with Blake.
Polydora hoplura Claparède, 1870
Gulf of Naples, Italy
Unknown. None deposited?
1.Read (1975)
2.Blake and Kudenov (1978)
1.New Zealand (Wellington and Marlborough Sounds)
2.Australia (New South Wales; Tasmania)
Eastern Atlantic; South Africa, Europe, New Zealand, Australia (New South Wales, Tasmania)
1.No
2. State that material agrees well with descriptions in 1.
Polydora latispinosa Blake and Kudenov, 1978
Port Phillip Bay, Point Cook, Victoria, Australia
Good
NMV G2874
Blake and Kudenov (1978)
Known only from type locality.
Not reported since original description
Yes, endemic
Polydora websteri Hartman in Loosanoff & Engle, 1943
Long Island Sound, Connecticut, USA
Good
Lectotype LACM-AHF POLY 1628
Radashevsky (1999)
Gulf of Mexico, Lemon Bay, Florida, USA; Long Island Sound, Connecticut, USA
East, west and gulfs coast of North America; west coast of South America; Australia (New South Wales); New Zealand?
Redescription of type material only
Polydora woodwicki Blake and Kudenov, 1978
Port Phillip Bay, Point Cook, Victoria, Australia; in Haliotus shell
NMV G2873
Blake and Kudenov (1978)
Australia (Port Phillip Bay, Victoria)
Yes, endemic
25
Table 5: Dipolydora species recorded as occurring in Australia, type locations,
redescriptions and distributions (Dipolydora ciliata (Johnston, 1838) included in Table 4 as
Polydora ciliata (Johnston, 1838)
Species Type location
Type material condition
Most recent redescription
Location of most recently described material
Recorded distribution
Australian material included in redescription
D. aciculata (Blake and Kudenov, 1978)
Bourne Creek, Kilcunda, Victoria, Australia; in Haliotis shell
Holotype NMV G2872, missing prostomium and peristomium;
Paratypes NMV 2873, good
Blake and Kudenov, 1978
Kilcunda, Victoria
Known only from type locality.
Not reported since original description
Yes, endemic
D. armata (Langerhans, 1880)
Madeira Island, Portugal
Syntypes ZMH V-1547
1.Radashevsky and Noguiera (2003);
2.Bick (2001)
1.Brazil
2. Cala Olivera, Ibiza (northwestern Mediterranean)
Portugal; Spain; Gulf of Naples, Italy; Brazil; Belize; Taiwan; Vietnam; Thailand; Japan; Mexico; New Zealand (Otago, Marlborough Sounds, Wellington); Arctic and Antarctic Seas; South Africa
No
(New Zealand – yes)
Dipolydora giardi (Mesnil, 1896)
St Martin, La Manche, Northern France
No types designated, no depository indicated (Radashevsky and Petersen (2005)
Radashevsky and Petersen (2005)
New material: Gulf of Naples, Italy
Recently discovered old material:
ZMUC POL-653
Mer de la Manche, France
La Manche, France; Gulf of Naples, Italy; Cap de Creus, Mediterranean Spain; Lundy Island, G. Britain; California; N.S.W. and Victoria Australia; Chile; Ecuador; New Zealand (Otago and Wellington)
No. Comments included on basis of description in Blake and Kudenov (1978)
26
D. flava (Claparède, 1870)
Gulf of Naples, Italy
Unknown.
None deposited?
Rainer, 1973 (as Polydora dorsomaculata n.sp., syn. in Blake and Kudenov, (1978))
Channel to north of Quarantine Island, Otago, New Zealand
Lower east coast, Bass Strait, Australia; New Zealand; Indian Ocean; Europe, Ceylon, Sumatra, Japan
no
D. notialis (Blake and Kudenov, 1978)
Tipara, South Australia from Haliotus roei shell
Holotype NMV G2877
Blake and Kudenov, 1978
Tipara, South Australia
South Australia
Not reported since original description
Yes, endemic
D. pilocollaris (Blake and Kudenov, 1978)
Port Phillip Bay, Victoria, Australia
Holotype NMV G2878
Paratypes NMV G2879-2880
Blake and Kudenov, 1978
Port Philip Bay, Victoria, Australia
Known only from type locality
Not reported since original description
Yes, endemic
D. protuberata (Blake and Kudenov, 1978)
Port Philip Bay, Victoria, Australia
Holotype NMV G2870
Paratypes NMV G2871
Blake and Kudenov, 1978
Port Philip Bay, Victoria, Australia
Known only from type locality
Not reported since original description
Yes, endemic
D. socialis (Schmarda, 1861)
Viña del Mar, Chile
Lost? Blake, 1996d
Of type: Mesnil, 1896
California, USA New South Wales, Victoria, Australia; the Sea of Japan; western Pacific; east and west coasts North America; Gulf of Mexico; Chile; Faulkland Islands
No
D. tentaculata (Blake and Kudenov, 1978)
Botany Bay, Towra Beach, New South
Wales, Australia
Holotype NMV G 2885
Paratype NMV G2886 QM G11597
Blake and Kudenov, 1978
Botany Bay, Towra Beach, New South Wales; Moreton Bay, Middle Banks, QLD, Australia
Queensland (Moreton Bay), New South Wales (Botany Bay, Hawkesbury River)
Yes, endemic
27
The Australian and east coast subtropical climate zones
The estuarine environment is an eco-tone between terrestrial/freshwater and marine
environments. Both the terrestrial and marine coastal conditions are considered here.
Terrestrial
Stern et al. (2007) modified Koppen’s scheme and presented it as it applies to Australia
with six climate classes: equatorial, tropical, subtropical, desert, grassland and temperate.
The subtropical class occurs on the east coast of Australia from Gladstone, Queensland
(23:51 151:16) south to Batemans Bay, New South Wales (35:43 150:11) and on the west
coast of Australia from Kalbarri (27:40 114:12) to Pinjarra (32:37 115:52) in Western
Australia. The subtropical climate class has four sub-classes: no dry season, distinctly dry
summer, distinctly dry winter and moderately dry winter. On the west coast a distinctly
dry summer subtropical climate occurs. On the east coast the majority of the subtropical
region is classed as no dry season with some small areas nearing the tropical climate class
classified as distinctly dry winter (Figure 1).
28
Figure 1: Australian climate classes
29
Marine waters, bioregions and estuaries of the subtropical east coast of Australia
The region covered by this study includes four of the IMCRA meso-scale bioregions
(Thackway and Cresswell, 1998): Tweed-Moreton, Manning Shelf, Hawkesbury Shelf and
Batemans Bay Shelf (Table 6). Rainfall over the region shows strong seasonality in the
north, with a predominantly Summer-Autumn pattern, becoming weaker in more
southern areas. No seasonal pattern is obvious in the most southern regions. The rains
from east Australian subtropical regions drain into the Pacific Ocean.
The warm East Australia current flows at more than 1.5km/hour in a southerly direction
along the coast, through the Coral Sea, before slowing and turning at the Manning Shelf
bioregion (Thackway and Cresswell, 1998) into the Tasman Sea becoming the Tasman
Current. Eddies of the East Australia Current continue into the Hawkesbury bioregion
where tropical Coral Sea and temperate Tasman Sea water masses meet (Thackway and
Cresswell, 1998). The southern region is mainly influenced by coastally trapped waves
setting northwards (Thackway and Cresswell, 1998).
One hundred and twenty estuaries occur through the New South Wales bioregions from
Tweed-Moreton to Batemans Shelf (West et al., 1985). Twenty-eight in Tweed-Moreton,
seventeen in Manning Shelf, twenty-two in Hawkesbury Shelf and fifty-three in Batemans
Shelf. These are predominantly sand barrier estuaries in Tweed-Moreton and Manning
Shelf, drowned river valleys in Hawkesbury Shelf and saline coastal lagoons in Batemans
Shelf bioregion (Thackway and Cresswell, 1998). In Queensland, the Tweed-Moreton
bioregion includes three major passage landscapes, with six rivers flowing into Southern
Moreton Bay (Thackway and Cresswell, 1998).
Over half of Australia’s estuaries are considered near pristine (National Land and Water
Resources Atlas (NLWRA), 2002). Fifteen near pristine estuaries occur from the Tweed-
Moreton to Batemans Shelf marine bioregions (Table 6), ten of these occur in Tweed-
Moreton. In New South Wales it is estimated that 64% of estuaries have poor water
quality mainly due to increased sedimentation and nutrient levels resulting from
anthropogenic activities such as land clearing and river flow alteration (Zann, 1995).
30
Table 6: Estuaries of the IMCRA bioregions in Australian east coast subtropical climate
region, rainfall patterns, coastal water influences and location of near pristine estuaries
(compiled from West et al., 1985; Thackway and Cresswell, 1998; NLWRA, 2002). *not
included in West et al., 1985
Bioregion Latitude U (upper) L (lower)
Rainfall Estuaries Near Pristine estuaries
Coastal water influences
Tweed-Moreton
TMN
U: to just north of Port of Battle Creek, QLD (~24°S)
L: North of Nambucca Heads, NSW (30° 39’S)
1400 – 2000mm
Predominantly summer-autumn
QLD: three major passage landscapes Southern Moreton Bay (Nerang, Coomera, Pimpama, Albert, Logan, Brisbane Rivers); Pumicestone Passage; Great Sandy Strait
NSW: 28 estuaries.
Dominated by sand barrier river estuaries
Colloseum Inlet (23.988°S)
Pancake Creek/Jenny Lind Creek (24.012°S)
Rodd’s Harbour (24.02°S)
Round Hill Creek (24.164°S)
Eurimbula Creek (24.17°S)
Jerusalem Creek (29.208°S)
Lake Arragan and River (29.565°S)*
Lake Cakora/Lagoon (29.602°S)*
Sandon River (29.674°S)
Station Creek (29.951 °S)
Dominated by East Australia Current. Longshore southward flow of tropical Coral Sea waters into temperate NSW continental shelf waters. Localised upwelling off Evans Head (29° 17´ S)
Manning Shelf
MAN
U: to north of 30° 39’S (north of Nambucca Heads)
L: Stockton (32° 54’S)
Predominantly summer-autumn rains
17 estuaries.
Large coastal river estuaries are predominantly sand barrier type
Khappinghat Creek (32.01°S)
Dominated by East Australia Current which breaks away in a south-easterly direction. Localised upwelling off Laurieton (31°
31
39´ S)
Hawkesbury Shelf
HAW
U: to 32° 54’S (Stockton)
L: Shellharbour (34° 35’S)
No significant pattern of rainfall
22 estuaries.
Large estuaries predominantly drowned river valleys type.
Eddies of the East Australia Current flowing southward and coastally trapped waves setting northward. Coral Sea and Tasman Sea water masses meet
Batemans Shelf
BAT
U: north to 34° 35’S (Shellharbour)
L: Tathra (36° 48’ S)
No seasonal pattern of rainfall.
53 estuaries.
Relatively small estuaries of the saline coastal lagoon type.
Termeil Lake (35.462°S)
Meroo Lake (35.484°S)
Willinga Lake (35.5°S)
Lake Tarourga (36.115°S)*
Mainly influenced by coastally trapped waves setting northwards.
32
The aim of this review
This review is focused on two commonly encountered polydorid genera, Dipolydora and
Polydora, in estuaries and bays of subtropical east coast Australia. The remaining five
Australian polydorid genera, Pseudopolydora, Boccardia, Boccardiella, Carazziella and
Polydorella are the subject of ongoing research.
This review aims to answer the following questions:
What information is currently available on polydorid spionids of east coast
subtropical estuaries and bays?
What information is held in museum collections about Dipolydora and Polydora in
the region?
How many species or morphospecies of Dipolydora and Polydora occur in
collections from east coast subtropical estuaries and bays?
Which morphological characters are most useful in identifying Polydora and
Dipolydora morphospecies from this region? Are there any “new” characters that
are useful? Is it possible to identify animals which are incomplete?
What are the distributions of the species or morphospecies of Dipolydora and
Polydora through the area in relation to the marine bioregions?
Where are the gaps in our knowledge of polydorid spionids in subtropical eastern
Australia and how can they be addressed?
33
CHAPTER 2: MATERIALS AND METHODS
Materials
Dipolydora and Polydora material was selected from museum database searches for
polydorid genera (Polydora, Dipolydora, Carazziella, Pseudopolydora, Boccardia,
Boccardiella and Polydorella) between Gladstone, Queensland (~24°S) and Tathra, New
South Wales (36° 48’ S). This represents the extent of the subtropical area of the east
coast of Australia based on marine bioregions (Table 1). Holotype and paratype material
of species recorded from Australian subtropical regions were examined when available
(Dipolydora tentaculata; Polydora haswelli; Polydora penicillata (recommended for
synonymy with Carazziella victoriensis in this study)). Type material of Carazziella
victoriensis was examined. Type material of species not recorded from New South Wales
were included when available (Dipolydora aciculata; Dipolydora protuberata; Polydora
latispinosa; Polydora woodwicki; Carazziella phillipensis). In addition, unregistered
material in museum collections identified as Spionidae, from the region of interest was
examined for polydorids. Only material from estuaries and coastal embayments was
included. Collection databases at the Australian Museum (AM), Sydney, Museum Victoria
(MV), Melbourne, the Queensland Museum (QM), Brisbane and the Museum and Art
Gallery of the Northern Territory, Darwin were searched. Collection databases of the
South Australian Museum and the Western Australian Museum and Tasmanian Museum
were not searched although all of these have small polychaete collections from the study
area (Hutchings, pers. comm.). Museum collections outside Australia, such as
Zoologisches Museum der Universitat, Hamburg, Germany, which hold Australian
material from international expeditions, were also not searched. New South Wales DPI
Fisheries provided freshly collected material from Camden Haven Estuary. A total of 125
specimens were examined. One hundred and fourteen of these were considered suitable
for the analysis. Eleven specimens were excluded from the analysis due to poor
condition.
Material for the study also came from collections deposited in museums from major
monitoring projects and collection programs including those of:
34
Queensland Electricity Commission, Calliope River and Auckland Ck, Gladstone, QLD
1974-1983
Marine Pollution Studies Group, Port Phillip Bay, Victoria, 1975
New South Wales Fisheries, Botany Bay Surveys, 1973
Australian Museum Hawkesbury River Surveys, 1977-1983
Maritime Service Board, Hunter River, Newcastle, 1983
CSIRO CRIMP port surveys, Newcastle, 1997
NSW Fisheries, Port Kembla Surveys, 2000
Sydney Ports Survey, 2001
Data collection
A morphological character X specimen data matrix was created as an Excel 2007
spreadsheet. The initial set of morphological characters was based on the character set
for polydorids in the DELTA interactive key of Wilson et.al (2003) and developed as
examination of specimens proceeded. A full list of characters used in the analysis is given
in Appendix 4. Specimens were observed whole on a concave slide in 70% ethanol using
Zeiss light microscopes at the Australian Museum and Museum Victoria, and an Olympus
microscope at Southern Cross University. Specimen data was entered directly into the
Excel 2007 spreadsheet as follows:
1. Label information including registration number, identity and collection details.
Where multiple specimens occurred in a single registered lot each specimen was
given the unique identifier of the registration number followed by .1, .2, .3, etc as
required. In some instances collection station numbers followed the registration
number eg. W31957 2-2-3.
2. Body dimensions: measurements at the Australian Museum and Museum Victoria
were made using a projection of the image through a camera lucida onto a drawn
35
scale. The scale was made for each magnification using a stage micrometer. At
Southern Cross University measurements were made using an eyepiece
micrometer.
3. Prostomium (first segment) and peristomium (second segment) details:
descriptions of the anterior edge of prostomium; lateral lobes; shape of anterior
ventral edge of peristomium and lobes on inner dorsal edge of peristomium
(Diagram 1).
Diagram 1: Dorsal prostomium (left) and ventral peristomium (right) details
4. Branchiae, range of chaetigers with branchiae, chaetiger with longest branchiae.
5. Parapodial lobes/lamellae, anterior to posterior: descriptions of shape (Diagram
2), length in relation to chaetal length.
Diagram 2: Parapodial lobes/lamellae shapes
digitiform triangular/conical globular fan
Lateral lobe
Anterior ventral edge
Inner dorsal edge lobe
Anterior edge of
chaetiger 1
Anterior edge
Palp
(attached to
peristomium)
eyes
caruncle
Prostomium
(first segment)
36
6. Parapodial chaetae, anterior to posterior: Chaetae type, descriptions and counts
for chaetiger 5 modified spines, noto- and neuro-chaetae, extent and number of
neuropodial capillary chaetae and hooded hooks per chaetiger.
7. Hooded hooks: the angle of tooth to main stem of a total of 10 hooks from
posterior, mid and anterior segments of the specimen was measured by drawing
the hook angle using camera lucida and measuring using a protractor. Care was
taken to only measure hooks lying perpendicular to the line of view. Occasionally
no hooks were lying in the correct orientation and so limited or no measurement
was made. Angles were recorded as >90°, >=90°, 90°, <=90° or <90°.
8. Reproductive segments: presence and extent of gametogenic segments was
recorded. Ova were easily seen but sperm packets were identified unreliably.
9. Specimens were re-identified using Blake and Kudenov (1978), Wilson et al. (2003)
and Hutchings and Murray (1984). Changes to original identification were noted.
Unidentified specimens and those of questionable identity were given unique
species codes.
Descriptive notes on specimens were recorded through the spreadsheet.
Images were taken at the same time as observations for specimens examined at Southern
Cross University using an Olympus D12 digital camera. Polydora sp 1 and Polydora cf.
calcarea were imaged at the Museum and Art Gallery of the Northern Territory using a
QImaging 5 MPixel digital camera. Combine ZM (Hadley, 2007) was used for some images
of Polydora sp. 1. Image notes were recorded on the Excel 2007 datasheet. Line drawings
were made using tracings of photographic images.
Re-identification of material
A total of 125 individuals were examined for the analysis. Forty-nine specimens of
Polydora and 57 specimens of Dipolydora were included in the analysis. Although much of
the material had been previously identified some identifications were incorrect and some
samples were polyspecific, having more than one species under the same name. This
37
occurred most often in material from the older large surveys. Anything that was doubtful
was recorded as such.
Four specimens of Polydora, 5 specimens of Dipolydora and a single Carazziella cf.
victoriensis were excluded from the analysis dataset due to the specimens being
damaged, incomplete, contracted or fragile. This made it impossible to examine many
morphological characters and created many missing values for those specimens in the
data matrix. Many specimens had far posterior regions damaged or missing so characters
recorded from this body region were excluded from the Part 1 analysis. A full list of
specimens examined but not included in the analysis is given in Appendix 7.
Combinations of traditionally used diagnostic characters from both Polydora and
Dipolydora in individual specimens also caused problems in making confident
identifications. Presence or absence of a constriction on the hooded hooks is an
important generic character in separating Polydora from Dipolydora (Blake, 1996d).
Specimens that had been identified as Polydora species, but where constriction on the
hooded hooks was not obvious, were re-identified as Dipolydora or Dipolydora?.
Similarly, specimens that had been identified as Dipolydora species but that had hooded
hooks exhibiting a constriction, were re-identified as Polydora species. The problems of
identification of the material examined for this study are explained below.
Polydora haswelli Blake and Kudenov, 1978 material
Six Polydora haswelli paratypes had questionable identity. Four (AM W13042-8, AM
W13042-9, AM W13042-10, AM W13042-11) and an associated juvenile (AM W13042-12)
were re-identified as Dipolydora? based on the lack of a constriction on the hooded
hooks. A fifth P. haswelli paratype (AM W13042-2) with no constriction on the hooded
hooks and notochaetae on chaetiger 1 was re-identified as Dipolydora sp. ????. The sixth
(AM W13042-1), with no constriction on the hooded hooks, was re-identified as D.
socialis. Another Polydora haswelli specimen (AM W23667) bearing resemblance to the
description of Dipolydora aciculata (Blake and Kudenov, 1978) was re-identified as
Dipolydora sp. “different” as it was different to all the other P. haswelli material . The
holotype and paratype of D. aciculata were included in the analysis for comparison.
38
Polydora woodwicki Blake and Kudenov, 1978 and P. cf. woodwicki material
Polydora woodwicki is described as having a rounded prostomium and branchiae which
extend for only 21-23 segments (Blake and Kudenov, 1978). It is the only described
Australian Polydora species with a rounded prostomium. AM W26122 identified as
Polydora woodwicki had a truncate prostomium and branchiae which extended from
segments 7 - 70. This specimen was re-identified and included in the analysis as Polydora
“not woodwicki”. AM W27868, an incomplete specimen identified as Polydora cf.
woodwicki was re-identified as Polydora cf. cornuta on the basis of chaetiger 5 spines.
Two specimens identified as Dipolydora cf. armata (AM W26151, AM W26152) were re-
identified as Polydora cf. woodwicki each having a constriction on the hooded hooks and
a rounded or truncate anterior prostomial margin. Dipolydora armata (AM W29649) had
a constriction on the hooded hooks placing it in Polydora and the prostomial margin of P.
cf. woodwicki but with chaetiger 5 spines and branchial distribution similar to Polydora
hoplura (Blake and Kudenov, 1978). AM W29649 was re-identified as P. cf.
woodwicki/hoplura/armata. Subsequent examination of the P. woodwicki holotype
confirmed that none of these specimens were P. woodwicki and they remained in the
analysis as P. cf. woodwicki and P. cf. woodwicki/hoplura/armata. The holotype of
Polydora woodwicki (MV F42873) was in poor condition but was included in the analysis
for comparison with these and other specimens.
Polydora cf. websteri material
Eleven specimens from Gladstone identified as Polydora cf. websteri were considered a
potential new species, and were re-identified as Polydora sp. 1. This potential new
species showed similarities to Polydora woodwicki in posterior spine form and pygidial
characters. One more specimen from this group (AM W199279-15) was small, more
damaged and its identity was uncertain. This specimen was included in the analysis as
Polydora sp. ??. Five specimens identified as Polydora cf. websteri (AM W199279-3, AM
W199279-10, AM W199279-11, AM W199279-16, AM W199279-24) had no constriction
on the hooded hooks, only simple spines on chaetiger 5 and an obvious gizzard. The
39
obvious gizzard is considered characteristic of Dipolydora socialis (Blake, 1996d). These
specimens were re-identified as Dipolydora cf. socialis.
Four specimens identified as Polydora cf. websteri (AM W16919-1, AM W16919-2, AM
W16919-3, AM W199279-1) were re-identified as Polydora cf. cornuta as they all
possessed an occipital antenna, which is absent in Polydora websteri (Blake, 1971; Blake
and Kudenov, 1978).
Two P. cf. websteri specimens (AM W199279-5, AM W199279-6) had no constriction on
the hooded hooks, and could not be identified from Blake and Kudenov (1978) and were
considered the same as Dipolydora sp. “new” (see below under Dipolydora socialis
material).
Dipolydora giardi (Mesnil, 1986) and D. cf. giardi material
Dipolydora giardi (AM W29944) possessed hooded hooks with a constriction and was
also re-identified as Polydora cf. cornuta (in error as occipital antenna is recorded as
absent). Dipolydora cf. giardi (AM W26115) showed some characters of Dipolydora
aciculata and was re-identified as D cf. aciculata / cf. giardi. The holotype (MV G2872)
and paratype (MV G2873-1) of Dipolydora aciculata were included in the analysis to
assess the resemblance of D. cf. aciculata / cf. giardi to D. aciculata.
Dipolydora flava Claparède, 1870 material
Dipolydora flava specimens (AM W29651-1 and AM W29651-2) were re-identified as D.
cf. flava as specimens had an occipital antenna described as absent in D. flava (Blake and
Kudenov, 1978). Three Dipolydora flava (AM W29950, AM W31920, AM W31921) lacked
needle packets and were re-identified as D. cf. socialis.
Dipolydora socialis (Schmarda, 1861) material
Dipolydora socialis (AM W31957 2-2-3.1) had packets of needle spines, as described for
Dipolydora flava (Blake and Kudenov, 1978), and an occipital antenna so was re-identified
as Dipolydora cf. flava as above. Seven Dipolydora socialis specimens having emergent
packets of needle spines and no occipital antenna were re-identified as Dipolydora flava.
40
Polydora sp. (QM G10642-3) was re-identified as Dipolydora cf. socialis. Eight Dipolydora
socialis specimens identified as D. socialis using Blake and Kudenov (1978) possessed
conspicuous asymmetric companion chaetae and had no gizzard. These were re-
identified as Dipolydora sp. “new”.
Dipolydora tentaculata (Blake and Kudenov, 1978) material
One unregistered specimen identified as Dipolydora tentaculata (QMunreg1) had needle
packets, no occipital antenna (present in D. tentaculata) and simple spines. This specimen
was re-identified as Dipolydora flava/socialis/tentaculata (in error as pygidium is
recorded as being 4-lobed on QMunreg1 not 3-lobed as in both D. flava and D. socialis
(Blake and Kudenov, 1978) and unknown in D. tentaculata Blake and Kudenov, 1978)).
The holotype (MV G2885) and paratype (MV G2886) of D. tentaculata were included in
the analysis.
Dipolydora pilocollaris (Blake and Kudenov 1978) material
Three specimens of Dipolydora pilocollaris are included. The Dipolydora pilocollaris
paratype (AM W17068) was included for comparison.
Dipolydora penicillata (Hutchings and Rainer, 1979) material
Twelve specimens, including type material, identified as Dipolydora penicillata (AM
W8258, holotype; AM W8256, paratype) were re-identified as Carazziella cf. victoriensis.
Paratypes of Carazziella phillipensis (AM W17066) and C. victoriensis (AM W17067) were
included in the analysis dataset for comparison.
Analysis
Analysis package
PRIMER 6 (Plymouth Routines in Multivariate Ecological Research) (Clarke and Gorley,
2005) is a statistical package containing routines for univariate, multivariate and graphical
analyses. The package caters for both biological and physical data matrices. Few
assumptions are made about the form of the data in the methods used making it easy to
use and robust. Originally designed for analysis of species by sample data in community
ecology studies in the marine environment the package is suitable for analysis of any
41
multivariate matrices including for morphometric measurements in taxonomic
discrimination (Clarke and Gorley, 2005). Subsets of a dataset are easily extracted for
independent analysis. PRIMER 6 is used in this study to compare means of a large
number of specimens across a large character set with the intention of identifying like
groups. It can do this quickly: standardizing the multivariate data, producing resemblance
matrices, creating groups using cluster analysis (CLUSTER) and graphically representing
these groups in multidimensional scale graphs (MDS). The groups are clusters of similar
specimens and may be interpreted as clusters of specimens of the same species. It is not
possible to draw conclusions about evolutionary relationships using the type of data in
this study and this is not the purpose, or intention, of this thesis. In addition the
characters responsible for discriminating between, and within, clusters can be identified
using the similarity/distance functions (SIMPER). These characters can be interpreted as
being the key diagnostic characters for the species cluster groups.
Dataset preparation for analysis
A dataset suitable for PRIMER 6 (Clarke and Gorley, 2005) analysis was created from the
initial matrix. Morphological characters were numbered and character descriptions were
converted into presence/absence data. Missing values in the dataset were considered as
absent characters. Where very few missing values were recorded for a character, means
were used to fill in any missing values and retain the character in the dataset (112, 126
and 142). The characters of the most posterior segments were excluded as only 24 of the
114 specimens included in the analysis were complete. The analysis dataset was
multivariate and included presence/absence, ratios, measurements and counts. The list
of morphological characters and associated character number used in the final dataset
for Polydora and Dipolydora is included in Appendix 4.
The dataset was analysed in two parts:
Part1) Resemblance analysis to identify species cluster groups and SIMPER analysis to
clarify and correct the position of specimens of questionable identity.
Part 2) Resemblance analysis of final corrected dataset and SIMPER analysis to identify key
diagnostic features of the species cluster groups.
42
Part 1: Resemblance between specimens and position of specimens of questionable
identity
Two hundred and seven characters were analysed for 114 specimens. Characters
included presence of an anterior dorsal furrow (9), chaetiger 5 dimensions (14-16, 20),
presence and location of dorsal ciliary organs (21-27), shape of anterior prostomial
margin (30-34),presence of anterior prostomial furrows (35,36), shape of anterior
peristomial inner dorsal margin (38-43), shape of peristomial lateral margins (38-
43),shape of anterior peristomial ventral margin (5,57,59-61, 63-74, 76,77,80), presence
of pigmentation (86-90),presence and number of eyes (91-93), caruncle length (94-104),
presence of occipital antenna (105),presence and location of gizzard (106-109),
distribution and length of branchiae (110, 112-114), shape and distribution of parapodial
lobes (115-147) and type and distribution of chaetae (152-165, 168-171, 173-234, 236-
240, 243-253). Character details are included in Appendix 4 .
The dataset was analysed using Primer 6.1.1 (Clarke and Gorley, 2005). The Primer
sample data matrix was setup using the Excel spreadsheet data classified as “Other” data
type. Individual specimens were considered as “samples” and morphological characters
were considered “variables”. Two factors, genus and species, were added based on the
re-identification of the specimens made during examination. Doubtful identifications
were included with their unique species code.
The dataset was normalized and a lower triangular resemblance matrix created analyzing
between samples using DI Euclidean distance as the resemblance measure. This distance
measure was used as it is the Primer 6 default distance measure for environmental data
which this species X morphological character data most resembles. No dummy measure
was used as it is realistic to consider that two samples (specimens) could be identical
(morphologically).
A hierarchical cluster analysis was run using CLUSTER on the samples (specimens) of the
resemblance matrix. Group average was used as the cluster mode as some variability
within a species cluster group was expected. SIMPROF was selected to test for structure
within the clusters (Permutations for mean profile: 1000; Simulation permutations: 999;
Significance level: 5%). Separate analyses were done for, 1) the whole dataset including
43
all specimens 2) Polydora specimens and 3) Dipolydora specimens. Specimens of
questionable genus were included in all analyses.
A scatter gram was produced as needed using the resemblance matrix and non-metric
multidimensional scaling using the MDS analysis (Kruskal stress formula: 1; Minimum
stress: 0.01) to better visualize the distances between the specimens. Two dimensional
and three dimensional scatter plots were produced as required. Separate plots were
created for 1) the whole dataset 2) Polydora specimens and 3) Dipolydora specimens.
Similarity percentage – species (morphological characters) contributions were calculated
using SIMPER one-way (factor: genus) and two-way (factors: genus x species) on the
resemblance matrix (Cut off for low contributions: 90.00%). This analysis shows which
morphological characters are contributing most to resemblance between the genera for
the one-way analysis, and between species within genera for the two-way analysis. The
output generated lists the characters in order of highest to lowest percentage
contribution.
The cluster analysis results were used to identify specimens belonging to the same
species, to advise, or indicate, potential generic placement for those species that were
unidentified or of questionable identity and to inform of any possible misidentifications.
Specimens clustering with material of known identity (including type material),
based on the reference literature used, were considered to be of the same species
(but see discussion).
Specimens which were like known species but which did not cluster with those
species were considered to be potential new species. The SIMPER analysis results
of characters contributing most to the resemblance were used to investigate the
placement of these specimens further.
Specimens occurring in clusters without type material or material of known
identity were considered new species.
Isolated individuals were considered potential new species (but see discussion)
but were investigated further using the results of the SIMPER analysis as below.
44
The results of the SIMPER analysis were used to investigate the characters contributing
most to the resemblance between genera and species for specimens of questionable
identity. These important characters were used in two ways, 1) the analysis dataset was
checked for data entry errors around the character and 2) the original descriptive dataset
was checked to see if the specimen had characters that were aberrant but had been
missed in preparation of the analysis dataset, eg on a damaged or juvenile specimen.
Those specimens were removed from the final dataset for SIMPER analysis.
Part 2: Final analysis and identifications
The resemblance and SIMPER analysis was rerun on the corrected dataset. The SIMPER
analysis results generated information on every character making a contribution of more
than 0.01% to the distance between genera or species within genera. It was necessary to
set a limit to the amount of information used for ease of data management. The most
important diagnostic characters were assumed to be those making the greatest
percentage contribution to the distance up to a total cumulative contribution of 10 - 15%.
This limit level resulted in 1-5 characters being selected as most important for each
resemblance. Characters making an equal contribution over this limit were considered
equally important. Important characters were tabulated for genera, Polydora species and
Dipolydora species to allow comparison of species resemblance characters across genera
and between species within genera. Equally important characters are enclosed within
brackets. A character or characters consistently occurring as important in distinguishing
between a genus or a species and others was considered a diagnostic character for that
genus or species. These characters are reported under “SIMPER diagnosis” in the
taxonomic descriptions in order of decreasing importance. Taxonomic descriptions and
distribution maps for the species and potential new species were made. Posterior
characters were described from complete specimens occurring in the species cluster
groups when present. Descriptions of potential new species are brief and include only
SIMPER diagnostic characters and notes on conspicuous features. Further research and
collecting effort is required to clarify the placement of these single specimen cluster
groups.
45
CHAPTER 3: RESULTS
Part 1: Resemblance between specimens and position of specimens of questionable
identity
The PRIMER resemblance analysis produced distinct clusters of Polydora, Dipolydora and
Carazziella (Figure 1). There appears to be no single point at which a branch splits into
Polydora, Dipolydora, Dipolydora? or Carazziella clusters (Figure 2). Specimens bearing
least resemblance to each other and branching off early were from both Polydora and
Dipolydora. Some branches at lower levels contained mixed genera particularly within the
branches around Polydora haswelli clusters (Figure 2) which also held 3 of the 5
Dipolydora? specimens. All Carazziella specimens were contained within a single cluster
(Figure 4). Branching into identifiable groups is at the species level. In this analysis some
Dipolydora species may bear more overall resemblance to a Polydora species than
another Dipolydora species.
Polydora and Dipolydora Part 1 genus clustersNormalise
Resemblance: D1 Euclidean distance
GenusCarazziella
Dipolydora
Polydora
Dipolydora?
2D Stress: 0.22
Figure 1: MDS scattergram of Polydora including (Carazziella cf. victoriensis syn.
Dipolydora penicillata), Dipolydora and questionable genus Dipolydora? based on Part 1
identifications.
46
Figure 2: Polydora, Dipolydora and Dipolydora? clusters based on Part 1 identifications. (A
is the closed Carazziella cluster)
Poly
dora
and D
ipoly
dora
Part
1
cf. latispinosaarmata
cf. aciculata/ cf. giardi
not woodwicki
cf. cornutacf. cornuta
cf. woodwicki/hoplura/armata
tentaculatatentaculata
giardi? juv
cf. cornuta
flava/socialis/tentaculata(A)
giardi
cf. socialiscf. socialis
pilocollaris
pilocollaris
pilocollariscf. cornuta
cf. cornuta
cf. cornutahoplura
hoplura
aciculata
???sp. new
sp. new
sp. newsp. new
sp. new
sp. new
sp. newsp. new
sp. new
sp. newflava
flava
flava
flavaflava
flava
flavacf. calcarea
cf. calcarea
cf. calcarea
cf. calcareacf. calcarea
cf. calcarea
cf. calcarea
sp. 1??
sp. 1
sp. 1sp. 1
sp. 1
sp. 1
sp. 1sp. 1
sp. 1
sp. 1sp. 1
woodwicki
latispinosa
socialis????
sp. different
socialishaswelli
haswelli
?
haswellihaswelli
haswelli
haswellihaswelli
haswelli
???
giardi?haswelli
haswelli
???cf. woodwicki
cf. woodwicki
cf. flava
cf. flavacf. flava
pilocollaris
protuberatasocialis
socialis
socialis
cf. socialiscf. socialis
cf. socialis
cf. socialiscf. socialis
socialis
socialis
socialissocialis
cf. socialis
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alise
Resem
bla
nce:
D1 E
uclidean d
ista
nce
Ge
nu
sC
ara
zzie
lla
Dip
oly
dora
Poly
dora
Dip
oly
dora
?
47
Species clusters
Distinct clusters were formed for:
Polydora cf. cornuta (AM W16919-1, AM W16919-2, AM W16919-3) (Figure 3:
Group A)
Polydora hoplura Claparède, 1870(Figure 3: Group B)
Polydora cf. calcarea (Figure 3: Group C)
Polydora sp. 1 (Figure 3: Group D), including Polydora ?? (AM W199279-15)
Polydora haswelli Blake and Kudenov, 1978 - All specimens were on an
independent branch but in two subclusters: (AM W7283 holotype, +five
paratypes, W24940-1, W24940-2 + Polydora ? (AM W29948)) (Figure 3: Group E)
and (W23666, W29949 + three of the four P. haswelli paratypes re-identified as
questionable genus Dipolydora? (AM W13042-9, AM W13042-10, AM W13042-
11) (Figure 3: Group F)
Polydora cf. woodwicki (Figure 3: Group G)
Dipolydora cf. flava (AM W29651 – 2, AM W29651 – 1, AM W31957 2-2-3.1)
(Figure 3: Group H)
Dipolydora socialis (Schmarda, 1861) and D. cf. socialis specimens formed one
branch but with mixed clusters (Figure 3: Group I)
Dipolydora flava (Claparède, 1870) (Figure 3: Group J)
Dipolydora sp. “new” (Figure 3: Group K)
Dipolydora pilocollaris (Blake and Kudenov, 1978) (Figure 3: Group L)
D. cf. socialis (AM W29950, AM W31921) (Figure 3: Group M)
Carazziella cf. victoriensis (including the C. victoriensis Blake and Kudenov, 1978
and C. phillipensis Blake and Kudenov, 1978 types) (Figure 3: Group N)
48
Dipolydora tentaculata (Blake and Kudenov, 1978) (MV G2885 holotype, MV
G2886 paratype) (Figure 3: Group O)
Specimens occurring as individuals on separate branches were:
Polydora cf. latispinosa (AM W31464-1)
Polydora not woodwicki (AM W26122)
Polydora cf. cornuta (AM W199279-1)
Polydora cf. cornuta (AM W29944)
Polydora cf. cornuta (AM W27868)
Polydora woodwicki/hoplura/armata (AM W29649)
Polydora woodwicki Blake and Kudenov, 1978 (MV F42873, holotype)
Polydora latispinosa Blake and Kudenov, 1978 (MV G2874 holotype)
Dipolydora socialis (Schmarda, 1861) (AM W31957 2-1-3.2 juv.)
Dipolydora sp. “different” (AM W23667)
Dipolydora socialis (AM W13042-1 Polydora haswelli paratype)
Dipolydora sp ???? (AM W13042-2 Polydora haswelli paratype)
Dipolydora pilocollaris (Blake and Kudenov, 1978) (AM W17068 paratype)
Dipolydora protuberata (Blake and Kudenov, 1978) (AM W17069 paratype)
Dipolydora? sp. ??? (AM W13042-8, Polydora haswelli paratype)
Dipolydora aciculata (Blake and Kudenov, 1978) (MV G2873-1 paratype)
Dipolydora giardi (Mesnil, 1896) (AM W31083)
Dipolydora flava/tentaculata/socialis (QMunreg)
Dipolydora? giardi? juv. (AM W13042-12)
49
Dipolydora cf. aciculata/cf. giardi (AM W26115)
Dipolydora armata (Langerhans, 1880) (AM W31965)
Open branches of Carazziella, Polydora and Dipolydora clusters can be seen in Figures 4,
5 and 6 respectively.
50
Identifications A: Polydora cf. cornuta+cf. cornuta+cf. cornuta B: Polydora hoplura+hoplura
Poly
dora
and D
ipoly
dora
Part
1 s
pecie
s c
luste
rs
AM W31464-1
AM W26122
AM W27868
AM W29944
AM W29649
AM W13042-12
AM W199279-1
(A)
(B)
AM W13042-8
(C)
(D)
MV F42873
VM G2874
AM W31957 2-1-3.2 juv
AM W13042 -2
AM W23667
AM W13042-1
(E)
(F)
(G)
(H)
AM W17068
AM W17069
(I)
(J)
(K)
MV G2873-1
(L)
(M)
AM W31083
(N)
QMunreg 1
(O)
AM W26115
AM W31965
Sa
mp
les (
ind
ivid
ua
l re
gis
tra
tio
ns)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
Ge
nu
sC
ara
zzie
lla
Dip
oly
dora
Poly
dora
Dip
oly
dora
?
51
C: Polydora cf. calcarea+cf. calcarea+cf. calcarea+cf. calcarea+cf. calcarea+cf. calcarea+cf. calcarea D: Polydora sp. 1+??+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1+sp. 1 E: Polydora haswelli+haswelli+?+haswelli (holotype)+haswelli (paratype)+haswelli (paratype)+haswelli (paratype)+haswelli (paratype)+haswelli (paratype) F: Dipolydora? ??? (AM W13042-9, Polydora haswelli paratype)+Dipolydora? giardi? (AM W13042-11, Polydora haswelli paratype)+Polydora haswelli+haswelli+Dipolydora? ??? (AM W13042-10, Polydora haswelli paratype) G: Polydora cf. woodwicki+cf. woodwicki H: Dipolydora cf. flava+cf. flava+cf. flava I: Dipolydora socialis+cf. socialis+cf. socialis+cf. socialis+cf. socialis+socialis+cf. socialis+socialis+socialis+socialis+socialis+cf. socialis+socialis J: Dipolydora flava+flava+flava+flava+flava+flava+flava K: Dipolydora sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new”+sp. “new” L: Dipolydora pilocollaris+pilocollaris+pilocollaris M: Dipolydora cf. socialis+cf. socialis N: Carazziella victoriensis (paratype)+cf. victoriensis+cf. victoriensis+cf. victoriensis+ Carazziella phillipensis (paratype) +cf. victoriensis+cf. victoriensis+cf. victoriensis+cf. victoriensis+cf. victoriensis+cf. victoriensis (Dipolydora penicillata paratype) +cf. victoriensis (Dipolydora penicillata holotype) +cf. victoriensis O: Dipolydora tentaculata (holotype) +tentaculata (paratype) Registration numbers A:AM W16919-1+AM W16919-2+AM W16919-3 B:MV F43060-2+MV F43060 - 3 C:NSWCH3+NSWCH1+NSWCH4+NSWCH2+NSWCH6+NSWCH5+NSWCH7 D:AM W199279-4+AM W199279-15+AM W199279-12+AM W199279-13+AM W199279-14+AM W199279-21+AM W199279-22+AM W199279-2+AM W199279-20+AM W199279-8+AM W199279-7+AM W199279-9 E:AM W24940-1+AM W24940-2+AM W29948+AM W13042-3+AM W13042-4+AM W13042-7+AM W13042-5+AM W13042-6+AM W7283 F:AM W13042-10+AM W13042-11+AM W23666+AM W29949+AM W13042-9 G:AM W26151+AM W26152 H:AM W29651 - 2+AM W29651 - 1+AM W31957 2-2-3.1 I:AM W31957 2-2-3.3+AM W31957 2-1-3.2+AM W199279-3+AM W199279-10+AM W199279-11+AM W199279-16+AM W199279-24+QM G10642-3+AM W31957 2-2-3.2+AM W31957 2-2-2.1+AM W31957 2-1-3.1+AM W31957 2-2-4 .1+AM W31920 J:AM W31945-1+AM W31947-2+AM W31947-4+AM W31947-3+AM W31947-1+AM W31945-2+AM W31933 -1 K:AM W199279-5+AM W199279-6+AM W31957-2.3.3-4+AM W31957-2.3.3-5+AM W31957-2.3.3-6+AM W31957- 2.3.3-1+AM W31957- 2.3.3-2+AM W31957-2.3.3-3+AM W31957 2-3-3.1.77+AM W31957 2-3-3.2.77 L:AM W26119-3+AM W26119-1+AM W26119-2 M:AM W29950+AM W31921 N:AM W17067+AM W19112-9+AM W19112-7+AM W19112-8+AM W17066+AM W19112-6+AM W19112-5+AM W19112-3+AM W19112-4+AM W19112-2+AM W8258+AM W8256+AM W19112-1 O:VM G2885+VM G2886
Figure 3 (previous page): Polydora (including Carazziella cf. victoriensis syn. Polydora
penicillata), Dipolydora and Dipolydora? clusters based on Part 1 identifications. Distinct
clusters have been closed (Groups A – O). Group members and their registration numbers
indicated beneath graph.
52
Resemblance clusters including type material
The holotype (AM W8256) and paratype (AM W8258) of Dipolydora penicillata, identified
as Carazziella cf. victoriensis in this study, grouped closely with the paratype of
Carazziella victoriensis (AM W17067). The cluster also included the paratype of
Carazziella phillipensis (AM W17066) (Figure 4).
The Polydora haswelli holotype (AM W7283) and five paratypes formed a close cluster
(Figure 3: Group E, Figure 5). A second neighbouring cluster contained three P. haswelli
paratypes re-identified in this study as? sp ??? (AM W13042-9, AM W13042-10) and D?
Dipolydora giardi? (AM W13042-11) and two non-type specimens of P. haswelli (Figure 3:
Group F).
The Dipolydora tentaculata holotype (VM G2885) and paratype (VM G2886) clustered
together. No other specimens occurred in this cluster (Figure 3: Group O).
Resemblance clusters containing specimens of doubtful genus Dipolydora?
Polydora haswelli Blake and Kudenov, 1978 material
Resemblance and cluster analyses for Polydora and Dipolydora? (Figure 5) and Dipolydora
and Dipolydora? (Figure 6) indicate that three of the Dipolydora? specimens group
together with the second Polydora haswelli cluster (Figure 3: Group F) which neighbours
the type material cluster (Figure 3: Group E ). Either large variation exists within the P.
haswelli group or this neighbouring group is a potential new species of Polydora. Figure 6
also suggests that Dipolydora sp. ????, D. sp. “different” and Dipolydora socialis (AM
W13042-1), all previously P. haswelli, remain as species within genus Polydora. A second
juvenile Dipolydora socialis (AM W31957 2-1-3.2 juv.) also resembles this group.
Dipolydora? sp ??? (AM W13042-8), Dipolydora ???? (AM W13042-2) and Dipolydora sp.
“different” most closely resembled each other (Table 4) but did not cluster with each
other or any other species.
Dipolydora? giardi? juv. (AM W13042-12) did not cluster with P. haswelli.
53
Figure 4: Carazziella cf. victoriensis (syn. Dipolydora penicillata) cluster Part 1
resemblance analysis. Carazziella victoriensis paratype (AM W17067), Carazziella
phillipensis paratype (AM W17066), Dipolydora penicillata holotype (AM W8256),
Dipolydora penicillata paratype (AM W8258). (Other Polydora and Dipolydora clusters
closed. Refer to Figure 3 for group members)
Cara
zzie
lla c
luste
r P
art
1
AM W31464-1
AM W26122
AM W27868
AM W29944
AM W29649
AM W13042-12
AM W199279-1
(A)
(B)
AM W13042-8
(C)
(D)
MV F42873
VM G2874
AM W31957 2-1-3.2 juv
AM W13042 -2
AM W23667
AM W13042-1
(E)
(F)
(G)
(H)
AM W17068
AM W17069
(I)
(J)
AM W199279-5
AM W199279-6
(K)
MV G2873-1
(L)
(M)
AM W31083
AM W17067
AM W19112-9
AM W19112-7
AM W19112-8
AM W17066
AM W19112-6
AM W19112-5
AM W19112-3
AM W19112-4
AM W19112-2
AM W8258
AM W8256
AM W19112-1
QMunreg 1
(N)
AM W26115
AM W31965
Sa
mp
les (
ind
ivid
ua
l re
gis
tra
tio
ns)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
Ge
nu
sC
ara
zzie
lla
Dip
oly
dora
Poly
dora
Dip
oly
dora
?
54
Figure 5: Part 1 Polydora species clusters and the resemblance to the questionable genus
Dipolydora? (giardi?, giardi? juv, ???) specimens.
Poly
dora
and D
ipoly
dora
? P
art
1
AM W31464-1
AM W26122
AM W29649
AM W29944
AM W13042-12
AM W199279-1
AM W27868
MV F43060-2
MV F43060 - 3
AM W16919-1
AM W16919-2
AM W16919-3
AM W13042-8
NSWCH3
NSWCH1
NSWCH4
NSWCH2
NSWCH6
NSWCH5
NSWCH7
AM W26151
AM W26152
AM W24940-1
AM W24940-2
AM W29948
AM W7283
AM W13042-3
AM W13042-4
AM W13042-7
AM W13042-5
AM W13042-6
AM W13042-10
AM W13042-11
AM W23666
AM W29949
AM W13042-9
AM W199279-4
AM W199279-15
AM W199279-12
AM W199279-13
AM W199279-14
AM W199279-21
AM W199279-22
AM W199279-2
AM W199279-20
AM W199279-8
AM W199279-7
AM W199279-9
MV F42873
MV G2874
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
sp
ecie
sg
iard
i? juv
gia
rdi?
???
cf.
corn
uta
not
woodw
icki
hasw
elli
? hoplu
ra
sp.
1
??
woodw
icki
cf.
woodw
icki/hoplu
ra/a
rmata
cf.
woodw
icki
cf.
latispin
osa
latispin
osa
cf.
calc
are
a
55
Figure 6: Part 1 Dipolydora species clusters and their resemblance to the questionable
genus Dipolydora? (giardi?, giardi? juv, ???) specimens.
Dip
oly
dora
and D
ipoly
dora
? P
art
1
w31965
W26115
AM W13042-12
MV G2885
MV G2886
QMunreg 1
AM W31083
AM W26119-3
AM W26119-1
AM W26119-2
G2873-1
AM W29950
AM W31921
AM W199279-5
AM W199279-6
AM W31957-2.3.3-4
AM W31957-2.3.3-5
AM W31957-2.3.3-6
AM W31957- 2.3.3-1
AM W31957- 2.3.3-2
AM W31957-2.3.3-3
AM W31957 2-3-3.1.77
AM W31957 2-3-3.2.77
W31945-1
AM W31947-2
AM W31947-4
AM W31947-3
AM W31947-1
W31945-2
W31933 -1
w29651 - 2
w29651 - 1
w31957 2-2-3.1
AM W17068
AM W17069
AM W31957 2-2-3.3
AM W31957 2-1-3.2
v199279-3
AM W199279-10
AM W199279-11
AM W199279-16
AM W199279-24
MV G10642-3
AM W31957 2-2-3.2
AM W31957 2-2-2.1
AM W31957 2-1-3.1
AM W31957 2-2-4 .1
AM W31920
AM W13042-8
AM W31957 2-1-3.2 juv
AM W23667
AM W13042 -2
AM W13042-1
AM W13042-10
AM W13042-11
AM W13042-9
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
sp
ecie
sg
iard
i? juv
gia
rdi?
???
arm
ata
acic
ula
ta
cf.
acic
ula
ta/
cf.
gia
rdi
cf.
fla
va
flava
gia
rdi
pilo
colla
ris
pro
tubera
ta
cf.
socia
lis
socia
lis
flava/s
ocia
lis/t
enta
cula
ta
tenta
cula
ta
sp.
new
sp.
diffe
rent
????
56
SIMPER analysis – between genera
Table 1: Resemblance (average squared distance) within and between genera using Part
1 identifications and number of contributing characters of a possible 207 characters
contributing more than 0.1% to the distance (in brackets).
Carazziella Polydora Dipolydora Dipolydora?
Carazziella 121.76 (71)
Polydora 483.85 (124) 179.11(145)
Dipolydora 480.49 (131) 429.56 (162) 189.42 (165)
Dipolydora? 441.98 (69) 397.02(115) 398.84 (122) 146.11 (52)
Table 2: Characters that combined contribute to 15% of the distance between Carazziella,
Polydora, Dipolydora and Dipolydora? using Part 1 identifications. Highest value first (%
contribution). Characters with equal % contribution in brackets.
Carazziella Polydora Dipolydora Dipolydora?
Carazziella
Polydora (212, 213)(2.03%)
188, (180, 181),
55, (209, 214,215)
Dipolydora (212, 213) (2.04%)
188, (180, 181),
55, (209, 214, 215)
244(0.97%)
152, 246, 249, 251,(202, 110),204, 252,
194, 198, 191, 200, (123, 139, 48,
248),(239, 245, 227, 220, 59)
Dipolydora? 196 (5.16%)
247, 153, 60
196 (5.74%)
247, 153, 60
196 (5.72%)
247, 60
57
Important characters contributing to distance between genera
The Carazziella characters were very distinct dominating the characters contributing to
distance between Carazziella and the other genera. Most important characters were
characters 212 (chaetae chaetiger 5: spine form - ventral row - distally swollen) and 213
(chaetae chaetiger 5: spine form - ventral row - bristle-topped upper half of swelling), 188
(chaetae chaetiger 5: number of major spines – ventral), 180 (chaetae chaetiger 5: spine
row shape – straight dorsally), 181 (chaetae chaetiger 5: spine row shape – shallow curve
ventrally).
Dipolydora? characters were also dominant in distancing Dipolydora? from other genera.
Most important characters were 196 (chaetae chaetiger 5: form of dorsal spine - wrinkled
on convex surface), 247 (hooded hooks – dentition – bi), 60 (anterior peristomial ventral
margin -medial v, width slightly less than the base of the prostomium, depth 0.5 width),
153 (chaetae pre-chaetiger 5: chaetiger 1 - neuropodia 1 with capillary chaetae.)
Polydora and Dipolydora:
Polydora and Dipolydora resemblance was not strongly dominated by one character,
the greatest percentage contribution being 0.97%.This highest character contribution
percentage was low compared to those of Carazziella and Dipolydora? (Table 2).
Characters which contributed most to distance were 244 (hooded hooks with constriction
on stem), 152 (chaetae pre-chaetiger 5: chaetiger 1 - notopodia 1 with capillary chaetae),
246 (hooded hooks: maximum number of hooks per chaetiger), 249 (hooded hooks: hh-
angle main fang to stem - >90), 251 (hooded hooks: hh-angle main fang to stem - <90),
202 (chaetae chaetiger 5: type of accessory structure - dorsal row - sub-distal blunt
tooth), 110 (dorsal branchiae commence).
Characters 244, 251 and 202 are considered Polydora characters. Characters 152 and 249
reflect currently accepted Dipolydora characters (Blake, 1996d) although one Dipolydora
species, Dipolydora pilocollaris, has no capillary chaetae on the first notopodia (Blake and
Kudenov, 1978).
58
SIMPER analysis
Resemblance between Polydora species
The resemblance (average squared distance) cross the Polydora group was 82.6. One
hundred and twenty six of the 207 characters contributed more than 0.1% to the
resemblance.
Table 3: Resemblance (average squared distance) between Polydora species using Part 1
identifications. 1. cf. cornuta 2. not woodwicki* 3. haswelli 4. ?* 5. hoplura 6. sp. 1 7. ??*
8. woodwicki* 9. cf. woodwicki/hoplura/armata* 10. cf. woodwicki 11. cf. latispinosa* 12.
latispinosa* 13. cf. calcarea
* distance within the group was not able to be determined for single specimens
Resemblance between Dipolydora and Dipolydora? specimens
Dipolydora? and Dipolydora were analysed together as re-identifications indicated that
the Dipolydora? specimens were more similar to Dipolydora than Polydora. The average
squared distances within species cluster groups for Dipolydora and Dipolydora? were 98.3
and 118.23 respectively.
The Dipolydora? specimens did not resemble any of the other Dipolydora specimens.
1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13.
1. 224.27
2. 673.74 *
3. 402.17 614.24 76.56
4. 386.48 601.94 158.48 *
5. 497.07 650.10 385.48 370.86 10.15
6. 382.93 644.81 299.45 276.34 376.39 65.01
7. 369.81 606.78 268.03 252.88 331.46 133.54 *
8. 369.54 597.52 270.25 248.38 378.75 261.84 230.25 *
9. 578.56 795.73 485.02 493.88 572.87 462.42 480.40 488.35 *
10. 450.35 672.24 341.60 328.14 387.11 327.26 305.78 277.57 478.20 74.29
11. 743.07 1001.95 662.75 599.59 726.01 646.77 648.42 575.14 900.43 676.95 *
12. 411.71 646.41 327.48 292.70 382.75 327.12 272.69 200.51 560.17 326.10 583.59 *
13. 464.14 662.33 333.48 317.94 442.86 349.35 309.84 274.96 573.27 365.71 687.08 335.76 44.15
59
Table 4: Resemblance (average squared distance) between Dipolydora and Dipolydora?
species using Part 1 identifications. A: 1. armata 2. aciculata (type) 3. cf. aciculata/ cf.
giardi 4. cf. flava 5. flava 6. giardi 7. giardi? 8. giardi? juv. B: 9. pilocollaris 10.
protuberata 11. cf. socialis 12. socialis 13. flava/socialis/ tentaculata 14. tentaculata 15.
sp. “new” 16. “different” 17. ??? 18. ???? * single specimens so distance within the
cluster group was not able to be determined.
A.
1 2 3 4 5 6 7 8
1 *
2 689.54 *
3 867.12 654.01 *
4 663.72 444.90 687.75 60.74
5 646.42 401.34 596.36 385.15 101.27
6 706.98 425.99 560.30 445.89 432.70 84.14
7 * * * * * * *
8 * * * * * * 343.67 *
9 678.39 390.71 643.61 433.33 410.19 481.87 * *
10 656.23 417.23 638.44 354.39 340.89 429.97 * *
11 634.90 379.15 638.61 290.93 347.90 423.67 * *
12 620.33 385.24 611.25 308.34 341.07 419.36 * *
13 792.11 516.78 797.60 446.97 475.66 567.54 * *
14 814.69 505.97 729.07 514.47 502 512.37 * *
15 723.74 435.06 655.16 405.90 392.73 487.44 * *
16 648.84 406.71 609.22 372.35 390.30 409.87 * *
17 * * * * * * 213.87 409.19
18 605.61 417.20 606.00 438.84 379.67 394.03 * *
B.
9 10 11 12 13 14 15 16 17 18
9 110.85
10 383.57 122.48
11 366.43 325.38 *
12 371.86 293.91 255.53 41.28
13 519.18 439.02 419.42 387.94 75.69
14 554.43 440.26 479.10 438.79 512.82
15 432.23 357.90 346.36 345.25 468.89 521.76 75.69
16 397.56 272.89 342.47 323.94 427.55 502.63 333.15 *
17 * * * * * * * * *
18 423.58 368.55 366.22 341.99 481.57 519.94 429.95 284.72 284.72 *
60
Conclusions on the identity of the re-identified material
Polydora haswelli material (and the questionable genus Dipolydora?)
Dipolydora? is marginally closer to Polydora (average squared distance 397.02) than
Dipolydora (average squared distances 398.84) and so the Dipolydora? species are
considered to be species of Polydora.
The SIMPER analysis results (Table 2) identified the Part 1 characters most important in
the resemblance between Dipolydora? and both Polydora and Dipolydora as being
characters: 196 (chaetae chaetiger 5: form of dorsal spine – wrinkled on convex surface),
247 (hooded hooks: dentition – bi), 153 (chaetae pre-chaetiger 5: chaetiger 5 -
neuropodia 1 with capillary chaetae) and 60 (anterior peristomial ventral margin: medial
v, width less than the base of the prostomium, depth 0.5 width). Examination of the
descriptive dataset shows that AM W13042-8 is the only individual showing character
196 in the analysis. Characters 247 and 153 are present on all specimens in the analysis
except AM W13042-12, the juvenile attached to AM W13042-11. Character 60 is present
in AM W13042-10 and AM W3947-1 (Dipolydora flava). Character 66 (anterior
peristomial ventral margin: broad shallow v) is the next most important character and is
present on AM W13042-8, -9 and Dipolydora pilocollaris (W26119-1, -2, -3).
The identity of AM W13042-12 is inconclusive. This presumed juvenile had chaetigers 8-
11 expanded and attached to AM W13042-11. The absence of characters 247
(neurochaetae on chaetiger 1) and 153 (bidentate hooded hooks) in only AM W13042-12
suggests that these are juvenile characters. Absence of neurochaetae on chaetiger 1 may
also be a misinterpretation of the juvenile morphology and should be re-examined. AM
W13042-12 was excluded from the final SIMPER analysis.
The identity of AM W13042-8 is inconclusive. The specimen is recorded as being in good
condition, 9mm, 62 chaetigers and having packets of needle spines first emergent from
chaetiger 2. Polydora haswelli is not recorded as having packets of needle spines. It
appears that AM W13042-8 is an individual of a different species or a variant of an
existing species. This animal is included in the final dataset analysis as Polydora sp. P2S
(P=Polydora 2=second species S=single specimen species).
61
AM W13042-10 is recorded as being in good condition and with packets of needle spines
first emergent from chaetiger 21. AM W13042-11 is complete, the posterior is thought to
be compressed or regenerating and a juvenile (AM W13042-12) was pressed firmly to the
posterior end. Needle packets are emergent first at chaetiger 19.
The Dipolydora? specimens AM W13042-9, AM W13042-10 and AM W13042-11 clustered
together and are included in the final dataset with the two non-type P. haswelli (W23666,
W29949) in the cluster as Polydora cf. haswelli (Table 5). Although they cluster with
Polydora haswelli no pairs of groups with samples were present for SIMPER analysis to
occur that could identify the characters contributing to the resemblance.
Dipolydora sp. “different” (AM W23667) and D. ???? (AM W13042-2) occur as isolated
individuals and are included in the final dataset as Dipolydora sp D3S and Dipolydora sp.
D4S respectively (D=Dipolydora 3, 4 =third, fourth species S=single specimen species).
Polydora sp.? (AM W29948) clusters within a P. haswelli cluster group (average squared
distance Part 1 =158.48) (Figure 5) and is included as Polydora haswelli in the final
dataset.
62
Table 5: Specimens of the questionable genus (Dipolydora?), the specimens they most
closely resemble and their identity in the final dataset.
Registration Questionable species code
Length mm
Polydora haswelli, Holotype 11.2
Included in final dataset as
Original identification
Part 1
AM W13042-8
D?. sp.??? Polydora haswelli paratype
9 Dipolydora aciculata/ OR Dipolydora sp. “new”
Polydora sp. P2S
AM W13042-9
D?. sp.??? Polydora haswelli paratype
13 Polydora haswelli, non-type
(Fig. Group F)
Polydora cf. haswelli
AM W13042-10
D?. sp.??? Polydora haswelli paratype
5.6 Polydora haswelli, non-type
(Fig. Group F)
Polydora cf. haswelli
AM W13042-11
Dipolydora? giardi?
Polydora haswelli paratype
13.4 Polydora haswelli, non-type
(Fig. Group F)
Polydora cf. haswelli
AM W13042-12
Dipolydora? giardi? juv.
Small juvenile attached to AM W13042-12
2.1 Dipolydora tentaculata OR Dipolydora flava/socialis/tentaculata
Not included
63
Dipolydora socialis (AM W13042-1) occurs in isolation to other D. socialis specimens. It
resembles most closely the Polydora haswelli clusters (Figure 3) but does not cluster with
either of them. AM W13042-1 is included in the final dataset as Polydora sp. P3S. The
final SIMPER analysis may clarify its position.
Polydora woodwicki Blake and Kudenov, 1978 and P. cf. woodwicki material
The holotype of P. woodwicki occurred on an isolated branch most closely resembling the
holotype of Polydora latispinosa. P. not woodwicki (AM W26122) occurred on an isolated
branch most closely resembling P. cf. cornuta (AM W27868) (originally identified as P. cf.
woodwicki) which also occurred on an isolated branch. P. cf. woodwicki/hoplura/armata
(AM W29649) was not similar to any other specimen. AM W26122 and AM W29649 are
either individuals of new species or variants of existing species. They are included in the
final dataset as Polydora sp. P4S and Polydora sp. P5S. The occipital antenna, present in P.
cornuta, is recorded as absent for AM W27868 so this specimen is included as Polydora
cf. woodwicki in the final dataset.
Polydora cf. websteri material
Polydora sp. 1 specimens clustered together (average squared distance Part 1 = 65.01)
and it appears to be a valid new species. It is included in the final dataset as Polydora sp.
P1. Polydora sp.?? (AM W199279-15) clusters within the Polydora sp. 1 cluster and is
included as P. sp. P1 in the final dataset.
Polydora cf. websteri specimens re-identified as Dipolydora cf. socialis ((AM W199279-3,
AM W199279-10, AM W199279-11, AM W199279-16, AM W199279-24) clustered with
other D. socialis and are included in the final dataset as D. socialis.
The re-identified Polydora cf. cornuta group had an average squared distance of 224.27
(Table 3). The average squared distance within the other Polydora species ranged from
10.15 – 76.56 (Table 3). This indicates that the group had members which were not
similar. Three of the specimens re-identified as Polydora cf. cornuta (AM W16919-1, AM
W16919-2 and AM W16919-3) clustered together. These specimens agree quite well with
the description of Polydora cornuta in Blake and Kudenov 1978 (as Polydora ligni) and are
included in the final dataset as Polydora cornuta. AM W199279-1 also identified as
Polydora cf. cornuta occurred on an isolated branch bearing little resemblance to any
64
other specimen. Apart from having a small occipital antenna, this specimen fits the
description of P. websteri in Blake (1971). It is included in the final dataset as Polydora cf.
websteri.
Dipolydora giardi (Mesnil, 1986) and D. cf. giardi material
AM W29944 re-identified as Polydora cf. cornuta occurred on an isolated branch. It has a
constriction on the hooded hooks so is a Polydora species. It is unlike any other Polydora
in this study and is included in the final dataset as Polydora sp P6S.
Dipolydora cf. aciculata/ cf. giardi (AM W26115) was also an isolated individual in the
cluster analysis showing large distances from other specimens (Table 4). It did not cluster
with the D. aciculata type material although it closely resembles the description of
posterior chaetigers of D. aciculata (Blake and Kudenov, 1978) in having short acicular
spines replacing hooded hooks in the neuropodia and 2-3 acicular spines present in
notopodia. 17.32% of the resemblance (average squared distance=654.01) between AM
W26115 and D. aciculata was in character 207 (chaetae chaetiger 5 - type of accessory
structure-dorsal row - accessory tooth on crest) which was present in AM W26115 and
absent in D. aciculata. 20.53% of the resemblance between D. giardi and AM W26115
(average squared distance = 560.3) was also due to character 207 in addition to 10.26%
from 9 (anterior dorsal furrow first 11 chaetigers) and 23 (dorsal ciliary organ- strip
extending posterior to caruncle), both present in AM W26115 and absent in D. giardi. An
additional small spur on the chaetiger 5 spine crest is described in some specimens of D.
giardi (Blake and Kudenov, 1978) although not present on the D. giardi specimen in this
study. AM W26115 is included in the final dataset as Dipolydora cf. aciculata/ cf. giardi.
Dipolydora flava Claparède, 1870 material
Material re-identified as Dipolydora cf. flava (AM W29651-1, AM W29651-2 and AM
W31957 2-2-3.1) clustered closely together (average squared distance = 60.74). They did
not strongly resemble the D. flava cluster (average squared distance between the cluster
groups = 385.15) or Dipolydora tentaculata (average squared distance between the
cluster groups = 514.4), the only described Australian Dipolydora species with an occipital
tentacle. The D. cf. flava cluster was most similar to Dipolydora cf. socialis (average
65
squared distance between the cluster groups = 290.9) (Table 4), although this should be
considered with caution as the D. cf. socialis cluster group contains dissimilar members as
described below. Characters contributing most to the distance between D. cf. flava and
D. cf. socialis were 71 (anterior peristomial ventral margin - broad “u”, base 0.5x segment
width with 5 lobes on edge, almost as deep as it is wide), 69 (anterior peristomial ventral
margin -shallow open “u”)and 102 (caruncle (prostomial dorsal extension)- mid 4).
Character 71 is present in D. cf. flava and characters 69 and 102 are present in D. cf.
socialis.
AM W29950 and AM W31921 re-identified as D. cf. socialis occurred together in an
isolated cluster indicating that the D. cf. socialis cluster group is multispecific in this Part 1
analysis. These two specimens are included in the final dataset as Dipolydora sp D2.
Characters distinguishing them from D. socialis were identified after SIMPER analysis of
the final corrected dataset. AM W31920 re-identified as D. cf. socialis clusters well within
the Dipolydora socialis cluster and is included as D. socialis in the final dataset.
Dipolydora socialis (Schmarda, 1861) material
The material re-identified as D. flava clustered together (average squared distance =
101.27). QM G10642-3 re-identified as Dipolydora cf. socialis clustered well within the
Dipolydora socialis cluster and is included in the final dataset analysis as D. socialis.
Dipolydora sp. “new” clustered together (average squared distance = 75.69) and the
group is considered a good new species. It is included in the final dataset as Dipolydora
sp. D1. The juvenile D. socialis specimen (AM W31957 2-1-3.2 juv.) clustered away from
the Dipolydora socialis cluster group and most closely resembled Dipolydora ????. AM
W31957 2-1-3.2juv. was removed from the final analysis dataset.
Dipolydora tentaculata (Blake and Kudenov, 1978) material
The specimen re-identified as Dipolydora flava/socialis/tentaculata (QMunreg1) occurred
on an isolated branch next to the cluster of type material of Dipolydora tentaculata.
Characters contributing most to the distance between QMunreg1 and the D. tentaculata
types (average squared distance = 512.82) were 225 (chaetae chaetiger 5: companion
66
chaetae type/s: very thin spines) and 104 (caruncle: mid 5). Character 225 is present in
QMunreg1 and character 104 is present in D. tentaculata. D. tentaculata is distinguished
from D. flava and D. socialis in having hooded hooks in which the angle (of the main fang
to the spine stem) changes in a single fascicle (Blake and Kudenov 1978). This character
does occur in QMunreg1 but also occurs in D. socialis specimens examined in this study.
The distance to D. flava (average squared distance = 475.66) was also dominated by
character 225 (23.97% contribution). QMunreg1 was most similar to D. socialis (average
squared distance = 387.9), character 225 contributing 29.39% to the distance between
the two. QMunreg1 is included in the final dataset as Dipolydora sp. D5S (Dipolydora sp.
5, single specimen).
Dipolydora pilocollaris (Blake and Kudenov 1978) material
The Dipolydora pilocollaris paratype (AM W17068) clustered on a branch with the
Dipolydora protuberata paratype (AM W17069). Three specimens identified prior to this
study as D. pilocollaris (AM W26119) were included in the analysis. These specimens
clustered together but not with the D. pilocollaris type material. The average squared
distance within the D. pilocollaris specimens was 110.85, contributing most to this
distance were characters 49 (anterior peristomial lateral margin: lateral lobes reduced so
that lateral peristomium appears to taper up to prostomium) and 68 (anterior peristomial
ventral margin: medial “u”, equal to width of dorsal lobe, similar depth). Character 49, a
Pseudopolydora character) was found to have been incorrectly included in the dataset in
this single instance and was considered invalid. The distance within the cluster is not
great indicating that D. pilocollaris and D. cf. pilocollaris are very similar but as the three
specimens (AM W26119) do not cluster with the type material they are included in the
final analysis dataset as Dipolydora cf. pilocollaris and described as Dipolydora cf.
pilocollaris2 to avoid confusion with D. cf. pilocollaris in Wilson et al. 2003).
Dipolydora penicillata (Hutchings and Rainer, 1979) material
Specimens of D. penicillata re-identified as Carazziella cf. victoriensis in this study
clustered together with type material of Carazziella victoriensis and C. phillipensis.
Carazziella was clearly distinguished from other material in this study at the genus level
67
(average squared distance = 121.76) (Table 1). Within this Carazziella cluster group, the
average squared distance between C. cf. victoriensis and the C. victoriensis paratype was
226.87 (Table 6). The average squared distance between C. cf. victoriensis and the C.
phillipensis paratype was 246.64 with characters 210 (chaetae chaetiger 5: type of
accessory structure - dorsal row: bristles over crest, tip bristle free) and 144 (parapodial
lamellae/lobes: chaetiger 6+: neuro digitiform) contributing most to this distance.
Characters 210 and 144 are variably present in C. cf. victoriensis and present in the single
C. phillipensis paratype. This leads to the question of whether Carazziella phillipensis is
synonymous with C. victoriensis. The average squared distance between the two
paratypes is 134.32, less than that between Carazziella cf. victoriensis and either of the
paratypes. The results of this analysis suggest that the three are synonymous, but it is not
the purpose of this thesis to review Carazziella so this matter will be addressed in a later
study. Carazziella cf. victoriensis is considered to be synonymous with Carazziella
victoriensis in this study and so it is recommended that Dipolydora penicillata Hutchings
and Rainer, 1979 be synonymised with Carazziella victoriensis Blake and Kudenov, 1978.
Table 6: Resemblance (average squared distance) between Carazziella cf. victoriensis, C.
phillipensis paratype and C. victoriensis paratype
cf. victoriensis phillipensis* victoriensis*
cf. victoriensis 124.11
phillipensis* 246.64 *
victoriensis* 226.87 134.32 *
68
Part 2: Final analysis and identifications
Species clusters – Polydora and Dipolydora
Distinct clusters were formed for Polydora and Dipolydora (Figures 7and 8). P. cf.
woodwicki and Dipolydora sp. D4S and D. sp. D3S were the only specimens that were on
branches with mixed genera.
Polydora and Dipolydora genus clustersNormalise
Resemblance: D1 Euclidean distance
GenusCarazziella
Dipolydora
Polydora
2D Stress: 0.22
Figure 7: MDS scattergram of Polydora including (Carazziella cf. victoriensis syn.
Dipolydora penicillata), Dipolydora.
69
Figure 8: Polydora and Dipolydora specimen clusters based on corrected dataset. (A-
closed cluster of Carazziella specimens)
Poly
dora
and D
ipoly
dora
specie
s
armata
cf. aciculata/ cf. giardi
tentaculata
tentaculata
(A)D5S
giardi
D2
D2
cf. pilocollaris
cf. pilocollaris
cf. pilocollaris
aciculata
D1
D1
D1
D1
D1
D1
D1
D1
D1D1
flava
flava
flava
flava
flava
flava
flava
cf. flava
cf. flava
cf. flava
pilocollaris
protuberata
socialis
socialis
socialissocialis
socialis
socialis
socialis
socialis
socialis
socialis
socialis
socialis
socialis
cf. woodwicki
cf. woodwicki
D3S
D4S
P3S
haswellihaswelli
haswelli
haswelli
haswelli
haswelli
haswelli
haswelli
haswelli
cf. haswelli
cf. haswelli
haswelli
haswelli
cf. haswelli
P1
P1
P1
P1P1
P1
P1
P1
P1
P1
P1
P1
woodwicki
latispinosa
cf. calcarea
cf. calcarea
cf. calcarea
cf. calcarea
cf. calcarea
cf. calcareacf. calcarea
P2S
hoplura
hoplura
cornuta
cornuta
cornuta
cf. websteri
P5S
P6S
P4S
cf. woodwicki
cf. latispinosa
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
Ge
nu
sC
ara
zzie
lla
Dip
oly
dora
Poly
dora
70
Species clusters: Polydora
Distinct clusters were formed for:
Polydora hoplura Claparède, 1870 (Figure 9: Group A)
Polydora cornuta (Figure 9: Group B)
Polydora cf. calcarea (Figure 9: Group C)
Polydora cf. woodwicki (Figure 9: Group D)
Polydora haswelli Blake and Kudenov, 1978 (Figure 9: Group E)
Polydora sp. P1 (Figure 9: Group F)
Specimens occurring on isolated branches were:
Polydora latispinosa Blake and Kudenov, 1978 (MV G2874 holotype)
Polydora woodwicki Blake and Kudenov, 1978 (MV F42873, holotype)
Polydora cf. latispinosa
Polydora cf. woodwicki
Polydora cf. websteri
Polydora sp. P2S (AM W13042-8, Polydora haswelli paratype)
Polydora sp. P3S (AM W13042-1 Polydora haswelli paratype)
Polydora sp. P4S
Polydora sp. P5S
Polydora sp. P6S
Polydora cf. haswelli – the position of this cluster of individuals is unresolved.
71
A: hoplura (MV F43060-2+MV F43060 – 3) B: cornuta (AM W16919-1+AM W16919-2+AM W16919-3) C: cf. calcarea (NSWCH3+NSWCH1+NSWCH4+NSWCH2+NSWCH6+NSWCH5+NSWCH7) D: cf. woodwicki (AM W26151+AM W26152) E: haswelli (AM W24940-1+AM W24940-2+AM W29948+ AM W7283, holotype +AM W13042-3 paratype+AM W13042-4, paratype +AM W13042-7, paratype +AM W13042-5, paratype +AM W13042-6, paratype) F: P1 (AM W199279-4+AM W199279-15+AM W199279-12+AM W199279-13+AM W199279-14+AM W199279-21+AM W199279-22+AM W199279-2+AM W199279-20+AM W199279-8+AM W199279-7+AM W199279-9)
Figure 9: Polydora species clusters using corrected dataset
Poly
dora
specie
s c
luste
rs
AM W31464-1
AM W26122
AM W29649
AM W29944
AM W199279-1
AM W27868
(A)
(B)
AM W 13042-8
(C)
(D)
AM W13042-1
(E)
AM W13042-10
AM W13042-11
AM W23666
AM W29949
AM W 13042-9
(F)
MV F42873
MV G 2874
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alise
Resem
bla
nce:
D1 E
uclidean d
ista
nce
specie
sh
asw
elli
ho
plu
ra
wo
od
wic
ki
cf.
wo
od
wic
ki
cf.
la
tisp
ino
sa
latisp
ino
sa
cf.
ca
lca
rea
cf.
ha
sw
elli
co
rnu
ta
cf.
we
bste
ri
P6
S
P4
S
P5
S
P3
S
P2
S
P1
72
Species clusters: Dipolydora
Distinct clusters were formed for:
Dipolydora tentaculata (Blake and Kudenov, 1978 ) (MV G2885 holotype, MV
G2886 paratype Figure 10: Group A)
Dipolydora cf. pilocollaris 2(Figure 10: Group B)
Dipolydora sp. D1 (Figure 10: Group C)
Dipolydora flava (Claparède, 1870) (Figure 10: Group D)
Dipolydora cf. flava (Figure 10: Group E)
Dipolydora socialis (Schmarda, 1861) (Figure 10: Group F)
Dipolydora sp. D2 (Figure 10: Group G)
Specimens occurring on isolated branches were:
Dipolydora protuberata (Blake and Kudenov, 1978) (AM W17069 paratype)
Dipolydora pilocollaris (Blake and Kudenov, 1978) (AM W17068 paratype)
Dipolydora aciculata (Blake and Kudenov, 1978) (MV G2873-1 paratype)
Dipolydora giardi (Mesnil, 1896)
Dipolydora cf. aciculata/ cf. giardi
Dipolydora armata (Langerhans, 1880)
Dipolydora sp. D3S
Dipolydora sp. D4S
Dipolydora sp. D5S
73
A: tentaculata (MV G2885, holotype +MV G2886, paratype) B: cf. pilocollaris (AM W26119-3+AM W26119-1+AM W26119-2) C: D1 (AM W199279-5+AM W199279-6+AM W31957-2.3.3-4+AM W31957-2.3.3-5+AM W31957-2.3.3-6+AM W31957- 2.3.3-1+AM W31957- 2.3.3-2+AM W31957-2.3.3-3+AM W31957 2-3-3.1.77+AM W31957 2-3-3.2.77) D: flava (AM W31945-1+AM W31947-2+AM W31947-4+AM W31947-3+AM W31947-1+AM W31945-2+AM W31933 -1) E: cf. flava (AM W29651 - 2+AM W29651 - 1+AM W31957 2-2-3.1) F: socialis (AM W31957 2-1-3.2+AM W199279-3+AM W199279-10+AM W199279-11+AM W199279-16+AM W199279-24+QM G10642-3+AM W31957 2-2-3.2+AM W31957 2-2-2.1+AM W31957 2-1-3.1+AM W31957 2-2-4 .1+AM W31920+AM W31957 2-2-3.3) G: D2 (AM W31921+AM W29950)
Figure 10: Dipolydora species clusters using corrected dataset
Dip
oly
dora
specie
s c
luste
rs
AM W31965
AM W26115
(A)
QMunreg 1
AM W31083
(B)
MV G2873-1
AM W23667
AM W13042 -2
(C)
(D)
(E)
(F)
AM W17068
AM W17069
(G)
Sa
mp
les (
sp
ecie
s)
05
10
15
20
25
30
Distance
Norm
alis
e
Resem
bla
nce:
D1 E
uclid
ean d
ista
nce
sp
ecie
sarm
ata
acic
ula
ta
cf.
acic
ula
ta/
cf.
gia
rdi
cf.
fla
va
flava
gia
rdi
pilo
colla
ris
pro
tubera
ta
socia
lis
tenta
cula
ta
D2
D3S
D1
D4S
D5S
cf.
pilo
colla
ris
74
For both Polydora and Dipolydora non-type material occurring as isolated individuals are
potentially new species. They may also be variants or damaged specimens of existing
species. Further material is required before their identity can be confidently established.
SIMPER analysis of final dataset - Resemblance between Dipolydora and Polydora
The average squared distance between Polydora and Dipolydora was 426.37 (Table 7).
The characters that contribute most to the distance between Dipolydora and Polydora in
order of decreasing contribution to distance are:
244 (hooded hooks with constriction on stem) contributes 0.88%
152 (chaetae pre-chaetiger 5: chaetiger 1 - notopodia 1 with capillary chaetae)
249 (hooded hooks: hh-angle main fang to stem - >90)
246 (hooded hooks: maximum number of hooks per chaetiger)
251 (hooded hooks: hh-angle main fang to stem - <90)
252 (hooded hooks: angle variable in the same row)
110 (dorsal branchiae commence)
194 (chaetae chaetiger 5: form of dorsal spine: slight sub-distal swelling)
202 (chaetae chaetiger 5: type of accessory structure - dorsal row - sub-distal blunt tooth)
The above characters in addition to characters 204, 248, 227, 73, (239, 35), (219, 133),
(191,223, 198, 116) and 245 (see Appendix 4) contribute to 15% of the distance between
Polydora and Dipolydora.
75
Table 7: Resemblance (average squared distance) within and between Polydora and
Dipolydora and number of characters (total of 207) contributing more than 0.1% to the
distance (in brackets).
Polydora Dipolydora
Polydora 179.24(162)
Dipolydora 426.37 (160) 189.99(165)
SIMPER analysis – between Polydora species
The resemblances (average squared distance) between Polydora species ranged from
10.15 (P. hoplura) to 170.56 (for P. cf. woodwicki) (Table 8). Suggesting that P. cf.
woodwicki is highly variable or that the cluster group still has more than one species. The
Polydora cf. haswelli cluster group is most similar to Polydora haswelli cluster group.
76
Table 8: Resemblance (average squared distance) between Polydora species. A: 1.
cornuta 2. cf. websteri* 3. P6S* 4. P4S* 5. cf. woodwicki 6. haswelli 7. hoplura 8. P1 B: 9.
woodwicki* 10. P5S* 11. cf. haswelli 12. P3S* 13. P2S* 14. cf. latispinosa* 15. latispinosa*
16. cf. calcarea
* distance within the group was not able to be determined for single specimens.
A.
B.
9. 10. 11. 12. 13. 14. 15. 16.
9. *
10. 488.35 *
11. 272.88 530.72 88.98
12. 297.00 537.20 228.70 *
13. 350.80 596.16 272.38 335.62 *
14. 575.14 900.43 655.23 689.85 753.37 *
15. 200.51 560.17 282.23 357.90 418.83 583.59 *
16. 274.96 573.27 368.22 385.27 460.04 687.08 335.76 44.15
1. 2. 3. 4. 5. 6. 7. 8.
1. 69.92
2. 480.23 *
3. 621.35 610.06 *
4. 728.99 731.92 710.79 *
5. 451.52 460.26 494.88 585.75 170.56
6. 382.42 402.69 478.73 613.12 352.17 77.05
7. 494.16 501.71 554.28 650.10 406.05 384.15 10.15
8. 342.5 375.67 483.49 641.64 351.78 294.79 372.64 65.30
9. 355.01 393.26 429.86 597.52 294.74 268.26 378.75 259.21
10. 577.97 546.97 657.42 795.73 496.48 485.82 572.87 463.92
11. 404.32 403.09 533.45 636.06 370.71 237.45 390.43 315.15
12. 422.27 406.88 524.77 638.6 377.27 247.5 471.31 335.27
13. 491.28 509.58 609.37 722.00 445.72 333.21 493.59 423.28
14. 703.13 761.48 838.12 1001.95 701.10 657.01 726.01 646.90
15. 380.71 434.18 504.49 646.41 347.23 324.32 382.75 322.58
16. 458.03 463.89 503.94 662.33 391.44 332.07 442.86 346.06
77
Table 9: Characters making the greatest percentage contribution to the resemblance
between Polydora species for type material and species which had more than one
representative 1. cornuta 2. cf. websteri* 3. P6S* 4. P4S* 5. cf. woodwicki 6. haswelli 7.
hoplura 8. P1 9. woodwicki*paratype 10. P5S* 11. cf. haswelli 12. P3S* 13. P2S* 14. cf.
latispinosa* 15. latispinosa*paratype 16. cf. calcarea. Greatest contribution first (%
contribution). Characters making equal % contribution in brackets.
Characters for all Polydora species in Appendix 5.
* distance within the group was not able to be determined for single specimens.
1. 5. 6. 7. 8. 9. 11. 14 15. 16.
1.
5. 162 (8.57%) 232, 236
6. 162 (10.12%) (236, 23)
(9, 140) (5.44%) 199
7. 206 (11.64%) 162 74
206 (14.16%) 74 195
206 (14.97%) 74 195
8. 162 (11.29%) 232 236
(9, 140) (5.45%) (59, 199)
51 (3.36%) 59 61
206 (15.43%) 74 195
9. 162 (10.90%) 232 236
(9, 140) (6.5%) 130
198 (3.92%) 51 61
206 (15.18%) 74 195
198 (4.06%) 130 59
11. 162 (9.57%) 232 236
(9,140, 60) (5.17%)
60 (8.07%) 51 61
206 (14.735) 74 195
60 (6.08%) 59 48
60 (7.02%) 198 130
14. 27, 46, 156) (16.21%)
(27,46, 156) (16.26%)
(27,46, 156) (17.35%)
(27,46, 156) (15.70%)
(27,46, 156) (17.62%)
(27,46, 156) (19.82%)
(27,46, 156) (17.40%)
15. (43, 162) (10.16%) 232
43 (11.14%) 9 140
43 (11.935) 198 15
206 (15.03%) 43 74
43 (11.99%) 198 15
43 (19.29%) 230 15
43 (13.71%) 60 230
(27, 46, 156) (19.53%)
16. 162 232 (8.45%) 236
(9, 140) (4.9%) 123
(123, 139) (5.18%) 77
206 (12.99%) 74 195
(123, 139) (4.97%) 77
(123, 139) (6.26%) 77
60 (5.21%) (123, 139)
(27, 46, 156) (16.59%)
43 (11.52%) 123 139
78
Important consistent characters contributing to distance for Polydora species in the
corrected dataset analysis were:
Polydora cornuta Bosc, 1802:
162 (chaetae pre-chaetiger 5: chaetiger 2-4, neuro 2-4 becoming short and spine-
like on chaetiger 4)
232 (chaetae chaetiger 6+: short spine-like chaetae in some notopodia)
Polydora cf. woodwicki:
9 (anterior dorsal furrow first 11 chaetigers)
140 (parapodial lamellae/lobes: chaetiger 6+: notopodial lobes rounded)
Polydora haswelli Blake and Kudenov, 1978:
51 (anterior peristomial lateral margin: lateral lobes rounded, level with
prostomium, forming a blunt anterior end)
61 (anterior peristomial ventral margin: narrow v becoming parallel sided slit to
level with chaetiger 1)
Polydora hoplura Claparède, 1870:
206 (chaetae chaetiger 5: type of accessory structure - dorsal row: subdistal flange
equal to the width of the spine tip, attached to the spine stem laterally, not
attached to the back of the spine stem)
74 (anterior peristomial ventral margin: deep narrow U, appearing shallowly lobed
(7or 9) or segmented on lower edge, width equal to anterior lobe width and 0.5x
depth)
195 (chaetae chaetiger5: from of dorsal spine: sub-distally inflated)
79
Polydora sp. P1
59 (anterior peristomial ventral margin: straight edge with a medial v-shaped
notch)
Polydora cf. haswelli:
60 (anterior peristomial ventral margin: medial v, width slightly less than the base
of the prostomium, depth 0.5 width)
Polydora cf. calcarea:
123 (parapodial lamellae/lobes: chaetiger 2-4 noto- digitiform)
139 (parapodial lamellae/lobes: chaetiger 6+ noto-broad triangular)
77 (anterior peristomial ventral margin: semicircular scallop, base level with
anterior chaetiger 1)
SIMPER analysis – between Dipolydora species
Resemblance (average squared distance) within the Dipolydora cluster group was 88.74.
Resemblance between the Dipolydora species ranged from 6.07 for Dipolydora cf.
pilocollaris2 to 161.06 for Polydora sp. D2 (Table 10).
80
Table 10: Resemblance (average squared distance) between Dipolydora species for type
material and species having more than one representative A: 1. armata* 2. aciculata*
(type) 3. cf. aciculata/ cf. giardi* 4. cf. flava 5. flava 6. giardi* 7. pilocollaris* paratype 8.
protuberata* paratype B: 9. socialis 10. D5S* 11. D2 12. tentaculata 13. D1 14. D3S* 15.
D4S* 16. cf. pilocollaris * single specimens
A.
1 2 3 4 5 6 7 8
1 *
2 689.54 *
3 867.12 654.01 *
4 663.72 444.90 687.75 60.74
5 646.42 401.34 596.36 385.15 101.27
6 706.98 425.99 560.30 445.89 432.70 *
7 591.75 342.02 597.90 355.85 310.48 406.47 *
8 656.23 417.23 638.44 354.39 340.89 429.97 230.63 *
9 617.90 372.09 614.58 280.39 325.71 408.29 408.29 261.29
10 792.11 516.78 797.60 446.97 475.66 567.54 413.25 439.02
11 710.55 466.82 720.90 353.28 454.23 505.53 396.18 417.64
12 814.69 505.97 729.07 514.47 502.00 512.37 435.70 440.26
13 723.74 435.06 655.16 405.90 392.73 487.44 344.33 357.90
14 648.84 406.71 609.22 372.35 390.30 409.87 303.15 272.89
15 605.61 417.20 606.00 438.84 379.67 394.03 316.80 368.55
16 707.27 406.94 658.85 459.16 443.42 507.01 315.15 434.56
B.
9. 10. 11. 12. 13. 14 15 16
81
9. 96.80
10. 373.59 *
11. 315.91 534.08 161.06
12. 442.71 512.85 567.61 41.28
13. 325.59 468.89 443.78 521.76 75.69
14. 320.22 427.55 443.56 512.63 333.15 *
15 349.45 481.57 469.14 519.94 429.95 284.72 *
16 378.10 554.49 504.57 594.00 461.53 429.04 459.17 6.70
Table 11: Characters making the greatest percentage contribution, and the size of the
greatest percentage contribution, to the resemblance between Dipolydora species for
type material and species which had more than one representative A: 1. armata* 2.
aciculata* (type) 3. cf. aciculata/ cf. giardi* 4. cf. flava 5. flava 6. giardi* 7. pilocollaris*
paratype 8. protuberata* paratype 9. socialis 10. D5S* 11. D2 12. tentaculata 13. D1 14.
D3S* 15. D4S* 16. cf. pilocollaris
Characters making equal % contribution in brackets. * species represented by a single
specimen
Characters for all Dipolydora species in Appendix 6.
1. 2. 4. 5. 6. 7. 8. 9. 11. 12. 13. 16.
1.
2. (134, 155, 205) (16.53%)
4. (134, 155, 205) (17.18%)
71 (8.7%) 68 210 232
5. (134, 155, 205) (17.64%)
(68, 210) (5.89%) 232 131
71 (10.04%) 131 161 135
6. (134, 155,
43 (9.08%)
(42, 71)
42 (8.945)
82
205) (16.12%)
199 (68, 195, 210)
(8.68%) 199 195
199 195 125
7. (134, 155, 205) (19.26%)
208 (8.565) 68 210 232
71 (10.87%) 208 146 24
208 (9.43%) 131 161 135
42 (9.52%) 199 208 195
8. (134, 155, 205) (17.37%)
(68, 108, 95, 210) (5.67%)
71 (10.92%) 95 108 146
(108, 95) (6.93%) 131 161
42 (9.0%) 199 (108, 95, 195)
208 (12.69%) 108 95 144
9. (134, 155, 205) (18.45%) 144
(68, 210) (6.35%) 232 (224, 51)
71 (13.80%) 24 146 40
131 (5.28%) 161 135 106
42 (9.475) 199 195 125
208 (11.20%) 38 141 106
(108, 95) (7.54%) (144, 38)
11. (134, 155, 205) (16.04%) 69
(69, 102) (12.21%) 68 210
(69, 102) (16.13%) 71 109
(69, 102) (12.55%) 109 131
(69, 102) (11.28%) 42 199
(69, 102) (14.39%) 208 109
(69, 102) (13.65%) 108 95
(69, 102) (18.04%) 109 40
12. (134, 155, 205) (13.99%)
104 (11.37%) (43, 42) 68
104 (11.18%) (43 42, 71)
104 (11.46%) (43, 42) 109
104 (11.22%) 43 199 195
104 (13.20%) (43, 42) 208
104 (13.06%) (43, 42) 95
104 (12.99%) (43, 42) 109
104 (10.13%) (69, 102) 43
13. (134, 155, 205) (15.75%) 187
(68, 210) (5.43%) 232 224
71 (9.53%) 146 24 40
131 (4.38%) 161 135 223
42 (7.94%) 199 195 125
208 (8.50%) 223 38 160
(108, 95) (6.60%) 223 38
223 (3.8%) 160 65 193
(69, 102) (12.84%) 109 40
104 (11.02%) (43, 42) 109
16. (134, 155, 205) (16.12%) 49
49 (9.51%) 208 210 232
49 (8.43%) 71 208 68
49 (8.72%) 208 68 131
(49, 42) (7.63%) (199, 208)
49 (12.27%) 68 66 122
49 (8.90%) 208 68 108
49 (10.23%) 208 68 66
(69, 102) (11.30%) 49 208
104 (9.68%) (43, 49, 42)
49 (8.38%) 208 68 66
83
Important consistent characters (Table 11) contributing to distance for Dipolydora species
in the corrected dataset analysis were:
Dipolydora tentaculata (Blake and Kudenov, 1978) (MV G2885 holotype, MV G2886
paratype):
104 (caruncle (prostomial dorsal extension): to mid chaetiger 5)
43 (anterior peristomial inner dorsal margin: no obvious lobe, a pair of lobes
between the lateral lobes, half the height of the u-shaped division formed
between the lateral lobes)
42 (anterior peristomial inner dorsal margin: a single medial lobe, ventral to this a
pair of lateral lobes forming a medial narrow ”v”)
Dipolydora cf. pilocollaris:
208 (chaetae chaetiger 5: type of accessory structure - dorsal row: a scale
extending just over convex (outer) may terminate in a rough broad edge)
68 (anterior peristomial ventral margin: medial “u”, equal to width of dorsal
lobe, similar depth)
Dipolydora sp. D1:
223 (chaetae chaetiger 5: companion chaetae type/s: pennoned strongly
geniculate, tapered)
160 (chaetae pre-chaetiger 5: chaetiger 2-4: noto 4-some capillary chaetae with
fibrous edge)
Dipolydora flava (Claparède, 1870)
131 (parapodial lamellae/lobes: chaetiger 2-4: neuro-broad rounded)
84
161 (chaetae pre-chaetiger 5: chaetiger 2-4: noto 2-4-glandular lobes with needle
spines)
135 (parapodial lamellae/lobes: chaetiger 5: neuropodial lobe), this on one
specimen only
Dipolydora cf. flava
71 (anterior peristomial ventral margin: broad “u”, base 0.5x segment width with
5 lobes on edge, almost as deep as it is wide)
146 (parapodial lamellae/lobes: chaetiger 6 +: neuro-conical (triangular))
24 (dorsal ciliary organ: strip across anterior edge of chaetiger 4, just posterior to
caruncle)
Dipolydora socialis (Schmarda, 1861)
125 (parapodial lamellae/lobes: chaetiger 2-4: noto-rounded)
106 (gizzard chaetiger: 13-16)
Dipolydora sp. D2
69 (anterior peristomial ventral margin: shallow open “u”)
102 (caruncle (prostomial dorsal extension) to mid chaetiger 4)
Single specimen cluster groups
Dipolydora protuberata Blake and Kudenov, 1978 (AM W17069 paratype)
108 (gizzard chaetiger: 17-22)
95 (caruncle (prostomial dorsal extension) to anterior chaetiger 2)
85
Dipolydora pilocollaris (Blake and Kudenov, 1978) (AM W17068 paratype)
208 (chaetae chaetiger 5: type of accessory structure - dorsal row: a scale
extending just over convex (outer) may terminate in a rough broad edge)
Dipolydora aciculata (Blake and Kudenov, 1978) (MV G2873-1 paratype)
210 (chaetae chaetiger 5: type of accessory structure - dorsal row: bristles over
crest, tip bristle free)
68 (anterior peristomial ventral margin: medial “u”, equal to width of dorsal
lobe, similar depth)
232 (chaetae chaetiger 6+: short spine-like chaetae in some notopodia)
Dipolydora giardi (Mesnil, 1896)
42 (anterior peristomial inner dorsal margin: a single medial lobe, ventral to this a
pair of lateral lobes forming a medial narrow “v”)
199 (chaetae chaetiger 5: type of accessory structure - dorsal row: lateral cheek-
like swelling with subdistal flange inside, appearing like a tooth from some angles)
195 (chaetae chaetiger 5: form of dorsal spine: sub-distally inflated)
Dipolydora armata (Langerhans, 1880)
134 (parapodial lamellae/lobes: chaetiger 5: notopodial lobe present)
155 (chaetae pre-chaetiger 5: chaetiger 1: neuropodia 1-capillary chaetae- 5
broad)
205 (chaetae chaetiger 5: type of accessory structure - dorsal row: subdistal flange
hooding the spine crest and with lateral wing-like arms extending forward from
spine stem)
86
CHAPTER 4: Descriptions of Polydora and Dipolydora species
Polydora Bosc, 1802
Type species: Polydora cornuta Bosc, 1802. See Blake and Maciolek, 1987.
Diagnosis (Blake, 1996d): Prostomium entire or incised anteriorly, extending posteriorly
as caruncle; eyes present or absent; chaetiger 1 without notochaetae. Chaetiger 5 greatly
modified, with major spines of 1 type (except post-larval or juveniles of a few species in
which first major spine is large, falcate, and differs from second and subsequent spines
that develop and eventually replace it), usually accompanied by slender companion
chaetae; spines arranged in a single curved row. Posterior notopodial spines present or
absent. Neuropodial hooded hooks bidentate with conspicuous angle between teeth,
with constriction and manubrium on shaft, beginning from chaetiger 7-14. Pygidium
saucer-shaped or disc-like, border usually entire except for dorsal gap. Anterior part of
digestive tract without gizzard-like structure.
SIMPER diagnosis (from this study): SIMPER analysis in this study identified the following
characters as contributing most to the distance between Polydora and Dipolydora:
Character 244 (hooded hooks with constriction on stem) – present in Polydora
Character 152 (chaetae pre-chaetiger 5: chaetiger 1 - notopodia 1 with capillary chaetae)
– absent in Polydora
Character 246 (hooded hooks: maximum number of hooks per chaetiger) – in this study
maximum number of hooded hooks is greater in Polydora (13 in Polydora sp. P1) than
Dipolydora (7 in D. tentaculata) but this character may be related to development and
requires further investigation.
Characters 251and 249 (hooded hooks: angle of main fang to stem <90°) – present in
Polydora
Character 202 (chaetae chaetiger 5: type of accessory structure - dorsal row - sub-distal
blunt tooth) – present in Polydora
87
Polydora cf. calcarea
(Figures 1, 2, 3)
Material examined: NEW SOUTH WALES: Camden Haven Estuary, Gogley’s Lagoon,
31.636S, 152.837E, mud blisters in valves of Ostrea angasi, vii.2007, coll: NSW DPI
Fisheries (7 unregistered specimens, included in this study as NSWCH 3, NSWCH1,
NSWCH4, NSWCH2, NSWCH6, NSWCH5, NSWCH7)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 123 (parapodial lamellae/lobes: chaetiger 2-4 noto- digitiform)
Character 139 (parapodial lamellae/lobes: chaetiger 6+ noto-broad triangular)
Character 77 (anterior peristomial ventral margin: semicircular scallop, base level with
anterior chaetiger 1)
Description: Two complete specimens, other specimens broken. Opaque white with
circulatory system conspicuous as red medial areas dorsally from just past chaetiger 5 to
around chaetiger 8, branchiae long with conspicuous cirri. Largest specimen 11.2mm for
65 chaetigers. Palps detached but with specimens, size range from 1.2-2.0mm.
Anterior prostomial margin weakly bifid; two pairs of eyes present or absent, when
present anterior pair farther apart than posterior pair; caruncle (prostomial dorsal
extension) extending to posterior chaetiger 2 to posterior chaetiger 3 (in largest
specimen); occipital antenna absent. Dorsal ciliary organ present as strip on either side of
caruncle (Fig. 2A) and posteriorly as broken strips along segment edge; anterior dorsal
furrow through first 11 chaetigers absent; anterior prostomial transverse furrow absent;
anterior dorsal furrow absent; posterior ventral furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin with a single medial lobe; lateral
margin with conspicuous lateral lobes, lateral lobes rounded, anterior edge level with
88
base of prostomial lobes, lateral lobes are long and fused with chaetiger 1 ventrally,
dorsally lobes appear to overshadow prostomium; ventral margin a “v” becoming a “u”,
anterior edge width of “v” equal to anterior width of prostomium; “u” 0.33x anterior
edge width, twice as deep as wide.
Chaetiger 5, 1.5x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length 0.1x
dorsal width.
Pigmentation: palps – distal three quarters of outer edge of ciliary groove with fine black
line, appearing wavy medially; line is actually made of a single row of fine zigzags;
parapodia – absent; prostomium – absent; peristomium – absent; dorsal thoracic –
absent; ventral thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to chaetiger 53 in larger specimen; longest at chaetiger
22; not overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform. Chaetiger 2-4: notopodia digitiform, similar
size on chaetigers 2-4, about 0.33x length of chaetae; neuropodial lobe digitiform about
0.33x length of chaetae. Chaetiger 5 without noto- or neuropodial lobes. Chaetiger 6+:
notopodial lobe broad triangle on chaetigers 6-9 then decreasing in size, absent after
chaetiger 15; neuropodial lobe not conspicuous and absent posteriorly.
Chaetae: Chaetiger 1: notochaetae absent; neurochaetae capillary chaetae. Chaetiger 2-
4: notochaetae two tiers of capillary chaetae, ventral tier shorter, shortest chaetae 0.5x
length of longest capillary chaetae on dorsal tier; neurochaetae two tiers of long and
short capillary chaetae, 0.5x length of longest notochaetae. Chaetiger 5: dorsally 5
bilimbate capillary chaetae, similar to shortest notochaetae on chaetiger 4, adjoining end
of spine row; ventrally about 8-9 broad aristate chaetae forming two tiers, chaetae are
smaller than hooded hooks on more posterior segments. Chaetiger 5 spines (Fig. 3): one
row of spines in a shallow curve; orientation longitudinal; 5-8 spines; spines can be
variable in same row, falcate or bluntly falcate, with lateral flange present on only one
89
side of spine that may be continuous from upper lateral side scooping to subdistal inner
edge or appear tooth-like in some orientations or be above a subdistal tooth, sub-distal
blunt tooth present on most spines in spine row, spines on ventral side of spine row have
largest blunt tooth, spines on dorsal side often with apparently simple falcate spine with
variably eroded lateral flange; companion chaetae present, close to spines, pennoned
with thin stem. Chaetiger 6+: notochaetae in two tiers, similar to anterior chaetigers to
chaetiger 9, then becoming longer and fewer in number to chaetiger 15, there reducing in
length and number to 2-3 long capillary chaetae and about 5 shorter capillary chaetae,
becoming longer again around chaetiger 30, posteriorly 1-3 long fine capillary chaetae
length approximately 0.5x body width, with several fine developing capillary chaetae in
each fascicle; neuropodia to chaetiger 6 with 5-7 capillary chaetae. Neuropodial hooded
hooks from chaetiger 7 to end, with constriction on stem, 1-11 in a row, 1-4 in posterior
row, bidentate, angle of main fang to stem ≤90°, angle not variable in same row.
Pygidium: collar with a “u”-shaped dorsal gap with triangular lobe/cirrus (?) just inside
gap (Fig. 2B), ventral section flaring out, a pair of dorso-lateral notches.
One specimen with gametogenic segments contained eggs in chaetigers 28-60, two other
incomplete specimens with egg bearing segments commencing at chaetigers 38 and 39.
Ecology: in mud blisters of oysters (Fig. 1)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Manning Shelf. It is very
likely more widely distributed than this but no material suitable for examination was
present in museum collections.
Remarks: A species which was “identical with the European P. ciliata of Johnston” was
cautiously reported by Haswell (1885) as occurring abundantly in diseased and dying
oysters in Hunter River oyster beds near Newcastle. On the inner surface of the shell of
these oysters were mud blisters which contained fine black mud and one or more
specimens of the worm. Further similar reports were made by Whitelegge (1890), a
zoologist of the Australian Museum, who examined material from Newcastle. He was
convinced that the worms were entering oysters that were lying loose on the surface or
partially buried on the low-water mark. Whitelegge (1890) includes Johnston’s 1938
90
description of Leucodore ciliatus, an account of the eggs of the Newcastle oyster worm
and a supplementary note on the occurrence of worms in mudblisters from oysters from
the Clarence River in northern New South Wales. He also refers the reader to McIntosh
(1868) for “a very lengthy description of the tentacles, bristles, hooks and the anal
segment” of Leucodore ciliata. The anal segment illustration (McIntosh 1868, Plate XVIII
Fig. 6.) though, appears similar to the distinctive three-lobed pygidium of Dipolydora
socialis as figured by Blake (1996d, Figure 4.34; 1971, Fig.13). Wilson (1928) thought
Whitelegge’s figures of egg sacs found with the worms identified as Polydora ciliata
(Johnston, 1838) to be more like those of Polydora ligni Webster, 1879 as described by
Söderström (1920). So, it appears that at that time several species may have been
combined under the name P. ciliata. Roughley (1922) reported on technical aspects of
oyster farming and the problems of Polydora ciliata in oysters from the Georges River,
Botany Bay. He believed that the problem mudworms had appeared at the same time as
stock of Sydney Rock Oyster was brought from New Zealand to replenish New South
Wales beds.
Unfortunately, no specimens of known origin have remained from these early records.
One lot (G11390), registered in the Australian Museum old register as “Old collection:
registered in Dec. 1908 as Polydora ciliata (from oyster shell) “, contained the labels
“Newcastle, oyster worm, 1927” with the specimens. These may be specimens examined
by either Haswell (1885) or Whitelegge (1890). Unfortunately, the single anterior
fragment is missing chaetiger 5 spines on both sides making it impossible to confirm an
identity. The Australian Museum, which may have held a permanent slide of the missing
chaetiger 5 spines, has no registered specimens of polydorids in its dry collection. It
appears from other labels included with the specimen that in 1964 the specimen was
sent by E. Pope, Curator of Worms, to K.H. Woodwick, at Fresno State College, Fresno,
California, who identified the specimen as Polydora websteri (Hartman, in Loosanoff and
Engle, 1943) but the material does not seem to have been returned to the Australian
Museum. Hartman had described P. websteri in 1943 with the aim of avoiding confusion
between the American species and the morphologically similar European Polydora ciliata,
based on the description of Polydora ciliata by Fauvel (1927). Subsequently, Blake (1971),
finding no specimens of Polydora ciliata in his studies, referred all records of Polydora
91
ciliata from the east coast of North America to Polydora websteri Hartman, in Loosanoff
and Engle, 1943. Blake and Kudenov (1978), in their revision of Australian Spionidae,
remark that the specimens examined by them, the pre- 1908 AM G11390 (3 specimens)
and NMV G3057 (4 specimens) from Crassostrea commercialis in Tuross Lake in southern
New South Wales, agreed well with the North American species, P. websteri. The Tuross
Lake specimens are currently on extended loan to an institution outside of Australia and
were unavailable for examination in this study. AMG11390 and NMV G3057 were the
only two registered lots of the widely reported pest species Polydora websteri from
subtropical east coast of Australia potentially available for this study in Australian
museums. One of these is no longer identifiable, the other out of the country and
reportedly not in good condition. Enquiries to the NSW DPI Fisheries found that there
were no reference collections of polydorid pest species kept by that organization (Wayne
O’Connor, pers. comm.). Fortunately, being such a common pest, NSW DPI Fisheries
were quickly able to supply oysters with mudworm infestations from the Camden Haven
Estuary in central New South Wales for examination. The abundant organism from the
infested oysters was identical to the description of Polydora ciliata (Johnston 1838) as
reported by Haswell (1885) and Whitelegge (1890). Whitelegge (1890) also reports that
the “ova appear to be matured in the body of the worm and commence on about the
thirtieth segment”, a character also seen in the Camden Haven material. This is almost
certainly the animal referred to by Haswell (1885) and Whitelegge (1890) as Polydora
ciliata (Johnston, 1838).
Recent investigations have indicated that boring and non-boring forms of Polydora ciliata
(Johnston, 1838) are different species. Radashevsky and Pankova (2006) referred boring
forms of Polydora cf. ciliata from the Sea of Japan and Europe to Polydora calcarea
(Templeton, 1836) and recommended that world-wide records of boring P. ciliata should
be verified. As such, material in this study considered identical to Polydora ciliata
(Johnston, 1838) and having a boring habit is referred to as Polydora cf. calcarea.
Polydora cf. calcarea is distinguished from P. cornuta and P. sp. P1 in lacking an occipital
antenna; from complete specimens of P. hoplura, with which it co-occurs in oysters with
mud blisters, and P. latispinosa, by the lack of posterior modified spines. It is similar to
92
the most recent descriptions of Polydora websteri Hartman, in Loosanoff and Engle, 1943
(Radashevsky, 1999) and Polydora haswelli Blake and Kudenov, 1978. In all three species
the chaetiger 5 modified spines have a lateral flange on one side. In P. cf. calcarea and P.
haswelli, the basal edge of the flange adjoins a single subdistal tooth. In P. cf. calcarea, it
seems that this tooth is sometimes only present, or well developed, on one of the spines
in the spine row. The flange often appears damaged or worn. Polydora cf. calcarea also
has a conspicuous dorsal ciliary organ, long flat branchiae which overlap and a finely
zigzagged dark line along the edges of the feeding groove of the palps. P. haswelli has
banded palps, sometimes unpigmented, although this may be due to preservation and
storage. Due to the morphological similarity of some members of the Polydora ciliata/
websteri Group confirmation of the identity of Polydora cf. calcarea as the European P.
calcarea (Templeton, 1838) would require greater and more extensive collection effort to
accurately assess morphological variation at different life stages, comparison with
European specimens and examination using similar methods to those of Radashevsky and
Pankova (2006) including investigation of spermatid aggregates and spermatozoan
morphology of the species and enzyme electrophoresis or molecular studies.
93
Figure 1: Habitat of Polydora cf. calcarea (right). Compacted mud extracted from mud
blister on the inner shell of an oyster from Camden-Haven estuary. Boccardia chilensis
(left) also occurs in this habitat.
94
Figure 2: Polydora cf. calcarea, Camden Haven Estuary (A. anterior dorsal, arrows
indicating paired strips of dorsal ciliary organ B. posterior ventral, arrow indicating “u”-
shaped dorsal gap
A
B.
95
Figure 3: Polydora cf. calcarea, chaetiger 5 spines: light micrograph, left; line drawing of
same, right
Polydora cornuta Bosc, 1802
Type locality: Charleston Harbour, South Carolina, USA; intertidal
Material examined: NEW SOUTH WALES: Towra Beach, Botany Bay, Stn 298, 2.4m at sand
spit, Zostera, sand, 17.iv.1973, coll: NSW Fisheries (AM W16919, 3 specimens)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 162 (chaetae pre-chaetiger 5: chaetiger 2-4, neuro 2-4 becoming short and
spine-like on chaetiger 4)
Character 232 (chaetae chaetiger 6+: short spine-like chaetae in some notopodia)
Description: Three specimens without posterior segments. Longest 3.4mm for 33
chaetigers, 0.4mm wide at chaetiger 5. Specimen with most chaetigers 3mm for 39
chaetigers, 0.3mm wide at chaetiger 5. Palp length 1-1.1mm.
Anterior prostomial margin strongly bifid; two pairs of eyes, anterior pair farthest apart;
caruncle (prostomial dorsal extension) extending to posterior chaetiger 2 or 3; occipital
antenna present. Dorsal ciliary organ may be conspicuous on some specimens as strip
extending posterior to end of caruncle; anterior dorsal furrow through first 11 chaetigers
absent; anterior prostomial transverse furrow absent.
tooth
Flange on one side of
spine tip
96
Anterior peristomial margins: inner dorsal margin with no conspicuous lobes; lateral
margin with conspicuous lateral lobes; ventral margin broad “v”, width of open edge
0.33x segment width.
Chaetiger 5 length 3x length of chaetiger 6; dorsally fused with chaetiger 6 or overlapping
chaetiger 6 but not fused; lateral length 0.5x dorsal width.
Pigmentation: palps - base of palp with tan speckles; parapodia – absent; prostomium -
tan speckles extending posterior from point adjacent to palp attachment point;
peristomium - tan speckling laterally; dorsal thoracic - chaetigers 1-3 lightly speckled with
tan; ventral thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to at least chaetiger 30 (posterior branchiae missing in
these incomplete specimens); longest at chaetiger 12; overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform; 0.5x to 1.0x length of neuropodial capillary
chaetae. Chaetiger 2-4: notopodial lobe conical, 0.2x chaetal length, longer on chaetigers
2 and 3, becoming broad on 4; neuropodial lobe absent. Chaetiger 5 without noto- or
neuropodial lobes. Chaetiger 6+: noto- and neuropodial lobes absent.
Chaetae: Chaetiger 1: notochaetae absent; neuropodia with capillary chaetae. Chaetiger
2-4: notochaetae two tiers, broad capillary chaetae, ventral tier short and spine-like,
geniculate on chaetiger 2, shortest chaetae 0.5x length of longest capillary chaetae on
dorsal tier; neurochaetae two tiers of capillary chaetae, becoming short and spine-like on
chaetiger 4. Chaetiger 5: no dorsal fascicle of capillary chaetae; ventrally three inferior
short spines. Chaetiger 5 spines: one row of spines in a shallow curve, endpoints anterior
dorsal, posterior ventral; orientation longitudinal; 5-6 spines; bluntly falcate; low
subdistal flange; companion chaetae present, brush-tipped and/or truncate. Chaetiger
6+: notochaetae limbate capillary chaetae; 3-4 with fibrous tips and aristate to chaetiger
14, then lengthening and becoming finer, several short spines chaetigers 6-8; from
chaetiger 27 broad short spine-like capillary chaetae; neuropodial capillary chaetae to
97
chaetiger 6, seven broad short geniculate capillary chaetae equal in length to hooded
hooks on more posterior chaetigers. Neuropodial hooded hooks from chaetiger 7 to at
least chaetiger 39 (posterior missing), with constriction on stem, 5-7 in a row, bidentate,
angle of main fang to stem <90°, angle not variable within single row, hooded hooks are
deeply embedded with only a short part of stem and hook emerging.
One specimen, 3mm for 39 chaetigers, gametogenic from chaetigers 27-39.
No noto- or neuropodial spines (although posterior incomplete)
Pygidium missing on all three specimens
Ecology: This material in sand substrate amongst the seagrass, Zostera. Elsewhere, tube-
building in mud and sand flats (Blake, 1996d), on mollusc shells, from decaying wood
amongst rocks with oysters, on polychaete (Merceriella and Diopatra) tubes, in mud
between Zostera, intertidal on fine sand, hull fouling (Radashevsky, 2005) .
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Hawkesbury Shelf,
VICTORIA – IMCRA bioregion - Victorian Embayments; temperate and sub-tropical zones
worldwide (Radashevsky, 2005).
Remarks: These Australian specimens examined are much smaller than the descriptions
of adult Polydora cornuta in Blake, (1971), Blake and Kudenov (1978) and Radashevsky
(2005). The specimens agree well with these descriptions but the spine-like capillary
chaetae, pigmentation and short ventral capillary chaetae have not been previously
described for adults. However, the description of recently settled P. cornuta juveniles
(Radashevsky, 2005) from Brazilian material with pigmentation on the prostomium and
lateral peristomium and occasional presence of one or two short ventral capillary chaetae
on recently settled 18-22 chaetiger juveniles, does agree with the three specimens in this
study. In the Brazilian juveniles the smallest males were 22 chaetigers with gametes in
chaetigers 13-14 and the largest female at 37 chaetigers was gametogenic at chaetigers
14-24. Mature individuals were recorded as having gametes develop from chaetigers 13-
15 to 15-33. The single Australian specimen with gametes differs in that gametes develop
between chaetigers 27-39.
98
Polydora haswelli Blake and Kudenov, 1978
Type locality: Nth Chinaman’s Beach, Sydney Harbour, Australia; sand
Material examined: NEW SOUTH WALES: Sydney Harbour, near N. Chinamens Beach, 8-
10m, 33° 49´S, 151° 14.9´E, sand, 08.v.1971, coll: P. Hutchings (HOLOTYPE, AM W7283);
Sydney Harbour, near N. Chinamens Beach, 8-10m, 33 °49´S, 151° 14.9´E, sand,
08.v.1971, coll: P. Hutchings (PARATYPES AM W13042, 5 specimens); Plantation Point,
Jervis Bay, NSW 1516, intertidal rock platform, 35° 4.8´S, 150° 41.8´E, between Galeolaria
tubes, 24/x/1998, coll: A. Murray (AM W24940, 2 specimens); Brotherson Dock Berth
4,Botany Bay NSW BBD4 P2-0, 0.5m, 33° 58.2´S, 151° 12.7´E, 20.x.1998, coll: NSW
Fisheries (AM W29948, 1 specimen)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 51 (anterior peristomial lateral margin: lateral lobes rounded, level with
prostomium, forming a blunt anterior end)
Character 61 (anterior peristomial ventral margin: narrow “v” becoming parallel sided slit
to level with chaetiger 1)
Description: Only the holotype is complete but it is broken at around chaetiger 41.
11.2mm for 72 chaetigers, 1.0mm wide at chaetiger 5. Palp length not recorded.
Anterior prostomial margin weakly bifid; two pairs of eyes present or absent, when
present anterior pair not further apart than posterior pair; caruncle (prostomial dorsal
extension) extending to posterior chaetiger 2 or posterior chaetiger 3; occipital antenna
absent. Dorsal ciliary organ absent; anterior dorsal furrow through first 11 chaetigers
absent; anterior prostomial transverse furrow absent.
Anterior peristomial margins: inner dorsal margin with no conspicuous lobes; lateral
margin with conspicuous lateral lobes, lateral lobes rounded, level with prostomium,
forming a blunt anterior end; ventral margin a narrow “v” becoming a parallel-sided slit
to level with chaetiger 1.
99
Chaetiger 5, 1.5-2.5 times length of chaetiger 6; dorsally fused with chaetiger 6; lateral
length 0.33x dorsal width.
Pigmentation: palps – holotype with 7 spots from base to distal end; parapodia – absent;
prostomium – when present 2 tan spots , lateral to palp attachment, pair of stripes either
side of prostomium, absent on holotype and one paratype; peristomium - absent; dorsal
thoracic - paired lateral spots, tan, chaetigers 2-4, absent on holotype and two paratypes;
ventral thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7, absent in posterior 0.5x of body, to chaetiger 32 in
holotype; longest at chaetiger 15 – 23; meeting but not overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe globose (rounded); neuropodial
lobe globose to broadly rounded, 0.5x length of neuropodial capillary chaetae. Chaetiger
2-4: notopodial lobe rounded less than 0.5x length of notochaetae, longer on chaetigers 2
and 3; neuropodial lobe absent or less than 0.5x neurochaetae length, globose on
chaetigers 2 and 3. Chaetiger 5 without noto- or neuropodial lobes. Chaetiger 6+: noto-
and neuropodial lobes absent.
Chaetae: Chaetiger 1: notochaetae absent; neuropodia with capillary chaetae. Chaetiger
2-4: notochaetae 2 tiered fan of capillary chaetae, longer dorsally, 0.5x as long ventrally;
neurochaetae short capillary chaetae, 0.33x notopodial chaetae length. Chaetiger 5:
dorsally 3-4 short bilimbate capillary chaetae anterior to spine row; ventrally 4 short
capillary chaetae similar to chaetiger 6. Chaetiger 5 spines: one row of spines in a shallow
curve, endpoints anterior dorsal, posterior ventral; orientation transverse/longitudinal or
longitudinal (holotype); 5-7 spines; spines may vary within single row and may be simple,
simple curved or bluntly falcate; sub-distal tooth and variously eroded weak lateral flange
on one side of spine; companion chaetae present, limbate, occasionally geniculate,
aristate or tapered. Chaetiger 6+: similar to chaetiger 4 until chaetiger 8, number of
capillary chaetae then reducing to 2 to 3 long capillary chaetae and about 4 shorter
capillary chaetae 0.5x length of longer; posterior 20 chaetigers with longer capillary
chaetae and some short newly emergent (?) capillary chaetae, no chaetae with fibrous
100
edge; neuropodia to chaetiger 6 with 4-6 capillary chaetae. Neuropodial hooded hooks
from chaetiger 7 to end, with constriction on stem, 5-11 in a row, 1-4 in row posteriorly,
bidentate, angle of main fang to stem <90°, angle not variable in same row, no posterior
spines.
Pygidium: thin rounded flap of glandular tissue, as wide as 6th segment forward from
posterior, length equal to that of last 6 posterior segments combined.
One specimen with gametogenic segments 28-48. Three paratypes with gametogenic
segments commencing at chaetigers 21, 25 and 30.
Ecology: sandy substrates, amongst Galeolaria tubes, wharf pile scrapings. Also recorded
by Blake and Kudenov (1978) from mud blisters in oysters from the Camden-Haven
Estuary, New South Wales. Unfortunately, that material unavailable for this study.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregions – Hawkesbury Shelf,
Batemans Shelf
Remarks: The material agrees well with descriptions in Blake and Kudenov (1978). There
appears to be great variation in pigmentation. The holotype has lost the distinctive
pigmentation of the prostomium and peristomium but the pigmentation on the palps is
still present but faint. It is not known whether lack of pigmentation is real or an artefact
of preservation. Polydora cf. haswelli recorded from Chile (Radashevsky et al., 2005) and
Polydora neocaeca Williams and Radashevsky, 1999 are similar to P. haswelli but, as
noted by those authors, the accessory structures on the chaetiger 5 spines are different.
Both P. cf. haswelli and P. neocaeca are recorded and illustrated as having a lateral
accessory flange on chaetiger 5 spines. While there is some variability in chaetiger 5
spines within the same row in the P. haswelli specimens examined in this study all have a
subdistal tooth and a variously eroded lateral accessory flange on one side of the spine.
Collection and examination of more fresh material in Australia may expose the same
levels of variation seen in Chilean material.
Three of the P. haswelli paratypes (W13042-9, W13042-10 and W13042-11) did not
cluster with the cluster group containing the Polydora haswelli holotype and were
101
analysed in this study as Polydora cf. haswelli (not Radashevsky, 2005). The P. cf. haswelli
specimens were more similar to Polydora haswelli than any other species. SIMPER
analysis identified ventral and lateral peristomial characters 60 (anterior peristomial
ventral margin: medial “v”, width slightly less than the base of the prostomium, depth 0.5
width), 61 (above) and 51 (above) as important in separating the two species cluster
groups. Polydora cf. haswelli specimens were also recorded as having packets of
emergent needle spines and hooded hooks without a constriction. Parapodial lamellae
and lobes were similar to P. haswelli. It is possible that P. cf. haswelli represent
reproductively mature forms of Polydora haswelli Blake and Kudenov, 1978 as one
specimen had a juvenile firmly pressed against the posterior. Developmental studies of
Polydora haswelli are necessary to aid in resolving the identity of the Polydora cf. haswelli
cluster group.
Polydora hoplura Claparède, 1870
Type locality: Gulf of Naples, Italy; shell-boring
Material examined: NEW SOUTH WALES: Coffs Harbour Jetty, Coffs Harbour, 30 18.4’ S
153 8.5’ E, 9 March 1992, 2.5 m, Salmacina tubes on jetty pilings, P.B. Berents, A.
Murray & R.T. Springthorpe, hand collected, SCUBA and snorkel, 0.5-5 m, AM (W26121);
TASMANIA: Simmonds Bay, Tasmania, (label reads Summons Beach), 43° 08´S, 147° 21´E,
on Crassostrea gigas, 17.vi.1977, coll: M. Skeel (MV F43060, 2 specimens)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 206 (chaetae chaetiger 5: type of accessory structure - dorsal row: subdistal
flange equal to width of spine tip, attached to spine stem laterally, not attached to back
of spine stem)
Character 74 (anterior peristomial ventral margin: deep narrow “u”, appearing shallowly
lobed (7or 9) or segmented on lower edge, width equal to anterior lobe width and 0.5x
depth)
102
Character 195 (chaetae chaetiger5: from of dorsal spine: sub-distally inflated)
Description: Both specimens incomplete. Largest specimen, missing pygidium, 21mm for
101 chaetigers, 1.2mm wide at chaetiger 5. Palps missing.
Anterior prostomial margin weakly bifid; two pairs of eyes present or absent, when
present anterior pair farther apart than posterior pair; caruncle (prostomial dorsal
extension) extending to anterior chaetiger 3; occipital antenna present. Dorsal ciliary
organ present as strip on either side of caruncle and as a transverse strip at segment
edge; anterior dorsal furrow through first 11 chaetigers absent; anterior prostomial
transverse furrow present; anterior dorsal furrow present. Posterior ventral furrow over
last 20 chaetigers present in larger specimen.
Anterior peristomial margins: inner dorsal margin with a single medial lobe, ventral to
this a pair of laterally flattened lobes meeting at anterior 0.33x of length forming a medial
“v”, continuing closely parallel; lateral margin with lateral lobes not notably enlarged;
ventral margin a deep narrow “u”, appearing shallowly lobed (7or 9) or segmented on
lower edge, width equal to anterior lobe width and 0.5x depth.
Chaetiger 5, 1.8 times length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
one quarter dorsal width.
Pigmentation: palps – absent; parapodia – absent; prostomium – light brown
pigmentation line along dorsal furrow in one specimen; peristomium - absent; dorsal
thoracic - absent; ventral thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to chaetiger 87 in larger specimen; longest at chaetiger
10; not overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, equal to length of
chaetiger 1 neuropodial lobe; neuropodial lobe broadly conical lobe, almost equal to
neurochaetae length. Chaetiger 2-4: notopodial lobe broad triangular lobe, 0.33x chaetal
length; neuropodial lobe triangular, 0.5x chaetal length on chaetigers 2 and 3, reduced on
103
chaetiger 4. Chaetiger 5 without noto- or neuropodial lobes. Chaetiger 6+: notopodial
lobe a posterior triangular lobe 0.25-0.33x chaetae length to chaetiger 10, then reduced
but still conspicuous posteriorly; neuropodial lobes absent for most of length, much
reduced posterior triangular lobe present in last 6 chaetigers.
Chaetae: Chaetiger 1: notochaetae absent; neuropodia with a spreading fascicle of
capillary chaetae across face of lobe. Chaetiger 2-4: notochaetae 3-tiered fan of capillary
chaetae, ventral chaetae 0.5x length dorsal chaetae; neurochaetae two tiers of capillary
chaetae, 0.5x length of longest notochaetae. Chaetiger 5: dorsally a cluster of short
capillary chaetae similar to shorter anterior neurochaetae; ventrally 6 broad thick
capillary chaetae. Chaetiger 5 spines: one row of spines in a shallow curve, ventral end
curved posteriorly; orientation longitudinal; 5-7 spines; spines bluntly falcate, sub-distally
inflated; subdistal flange equal to width of spine tip, attached to spine stem laterally, not
attached to back of spine stem; companion chaetae present, limbate, aristate or tapered.
Chaetiger 6+: notochaetae to chaetiger 9 similar to anterior noto-chaetigers, then
thinning to two to four longer capillary chaetae and several shorter capillary chaetae, last
9 chaetigers with falcate golden spines just under 0.5 x length of capillary chaetae;
neuro-chaetiger 6, 7 capillary chaetae. Neuropodial hooded hooks from chaetiger 7 to
end, absent on last 5 chaetigers, with constriction on stem, 5-10 in a row, 5 in posterior
row, bidentate, angle of main fang to stem ≤90°, angle not variable in same row.
Pygidium: pygidium present on posterior fragment in same lot - upright collar, dorsal gap
approx. 0.33x segment width
No specimens with gametogenic segments.
Ecology: inhabiting mud blisters in oysters; mud blisters in Atrina (Bivalvia: Pinnidae)
shells and epifauna on the shells of other molluscs (New Zealand)(Read, 1975).
Distribution: Australian: NEW SOUTH WALES: IMCRA bioregion - Hawkesbury Shelf (Point
Henry Pier), Manning Shelf; TASMANIA: IMCRA bioregion - Bruny; Europe (Claparède,
1870); South Africa (Day, 1967); New Zealand (Read, 1975)
104
Remarks: As noted by Blake and Kudenov (1978) these Tasmanian specimens agree well
with the description of P. hoplura in Read (1975). Polydora hoplura is often reported as
occurring in mud-blisters in oysters from New South Wales. Specimen AM W26121 was
examined but accidentally omitted from the analysis. MV G3059 was not examined or
included in the analysis through oversight. It is surprising that such a commonly reported
species is so poorly represented in museum collections.
Polydora cf. latispinosa
Material examined: NEW SOUTH WALES: Saint Georges Basin, inside Anadara trapezia
shell, 13.vii.2006 coll: Lena Collins (AM W31464-1)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 27 (body with medial glandular ridge)
Character 46 (anterior peristomial lateral margin: lateral lobes with deep medial division)
Character 156 (chaetae chaetiger 1: neurochaetae 2 capillary chaetae)
Remarks: Deeply embedded packets of spines are present beneath notochaetae. P. cf.
latispinosa did not group with the type material of P. latispinosa Blake and Kudenov,
1978. This was the largest specimen examined in this study 52.6mm for 255 chaetigers
and 13mm wide at chaetiger 5.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
105
Polydora cf. websteri
(Figure 4)
Material examined: Calliope River, Queensland, 23° 51´S, 151° 10´E, QEB 1974-1983, coll:
P. Saenger (AM W199279-1)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 67 (anterior peristomial ventral margin: deep “v”)
Character 24 (dorsal ciliary organ: strip across anterior edge of chaetiger 4, posterior to
caruncle)
Character 95 (caruncle (prostomial dorsal extension) to anterior chaetiger 2)
Remarks: This specimen has chaetiger 5 spines with a lateral flange on one side (Fig. 4) as
described for Polydora websteri Hartman, 1943 in Loosenoff and Engle in Blake (1996d)
and Radashevsky (1999). It is differs from P. websteri in possessing a small occipital
antenna.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Tweed-Moreton
Figure 4: Polydora cf. websteri (AM W199279-1) chaetiger 5 spines: light micrograph, left;
line drawing of same, right
Lateral flange
Companion chaetae
106
Polydora cf. woodwicki
(Figure 5)
Material examined: NEW SOUTH WALES: Coffs Harbour Jetty, NSW729, 30° 18.4’ S, 153°
8.5’ E, orange sponge on jetty pilings, 9.iii.1992, coll: R.T.Springthorpe (AM W26151);
Coffs Harbour Jetty, NSW732, 30° 18.4’ S, 153° 8.5’ E, shelly gravel near base of jetty
pilings, 9.iii.1992, coll: R.T.Springthorpe (AM W26152)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 9 (anterior dorsal furrow first 11 chaetigers present)
Character 140 (parapodial lamellae/lobes: chaetiger 6+: notopodial lobes rounded)
Description: 2 incomplete specimens. Largest, 2.3mm for 22 chaetigers, 0.4mm wide at
chaetiger 5. Smaller 1.9mm for 15 chaetigers, 0.6mm wide at chaetiger 5. Palp length
0.7mm.
Anterior prostomial margin rounded; eyes absent; caruncle (prostomial dorsal extension)
extending to posterior chaetiger 2; occipital antenna absent. Dorsal ciliary organ present
as transverse strip at anterior edge of segments; anterior dorsal furrow through first 11
chaetigers present or absent.
Chaetiger 5, 1.5-2.5x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
0.25x dorsal width.
Anterior peristomial margins: inner dorsal margin single medial lobe, ventral to this a pair
of lobes forming a medial “v” where they meet; lateral margin lateral lobes not notably
enlarged; ventral margin a semicircular scallop, base level with anterior chaetiger 1.
Buccal area funnel-shaped from anterior peristomium to chaetiger 4 where digestive
tract visible continuing medially as parallel-sided structure.
107
Pigmentation: palps - absent; parapodia – absent; prostomium - absent; peristomium -
scattered dark lateral pigmentation present or absent; dorsal thoracic - absent; ventral
thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to at least chaetiger 22 (posterior branchiae missing in
these specimens); longest at chaetiger 8; not overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform; neuropodial lobe
conical equal to chaetae length. Chaetiger 2-4: notopodial lobe conical, 0.2x chaetal
length; neuropodial lobe triangular, 0.5x neurochaetae length on chaetigers 2 and 3;
reduced and broad on chaetiger 4. Chaetiger 5 without noto- or neuropodial lobes.
Chaetiger 6+: posterior notopodial lobe, 0.2x length of chaetae, reducing posteriorly;
neuropodial lobes absent.
Chaetae: Chaetiger 1: notochaetae absent; neuropodia with capillary chaetae. Chaetiger
2-4: notochaetae two tiers, capillary chaetae, ventral tier shorter, shortest 0.5x length of
longest capillary chaetae on dorsal tier; neurochaetae two tiers of long and short capillary
chaetae, 0.5x length of longer notochaetae. Chaetiger 5: dorsally 2-4 anterior broad
thick capillary chaetae similar to chaetiger 6 and 3 fine capillary chaetae of equal length
with fibrous tips; ventrally three short broad capillary chaetae. Chaetiger 5 spines (Fig. 5):
one row of spines in shallow curve, ventral end curved posteriorly; orientation
perpendicular; 3-5 simple spines; lateral cheek-like swelling on some spines; continuous
flange from upper lateral sides of spine scooping to subdistal inner edge; companion
chaetae present, limbate, brush-tipped, truncated or tapered or with distal fibrous edge.
Chaetiger 6+: two tiers, dorsal tier long aristate capillary chaetae and ventral tier
geniculate and aristate capillary chaetae 0.5x length of dorsal tier capillary chaetae, some
tips slightly geniculate; neuropodial capillary chaetae to chaetiger 6, 6 capillary chaetae.
Neuropodial hooded hooks from chaetiger 7 to at least chaetiger 22 (posterior hooded
hooks missing in these specimens), with constriction on stem variable, anterior hooded
hooks without constriction, posterior hooded hooks with constriction, 3-5 in single row,
bidentate, angle of main fang to stem 90°, angle not variable in same row.
108
One specimen with eggs in segment 21.
One specimen had posterior notopodia with 2-3 acicular spines.
Pygidium missing on both specimens.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Manning Shelf (Coffs
Harbour).
Remarks: This anomalous species, with the constriction on the hooded hooks present in
posterior but absent in anterior chaetigers, is the only Polydora that resembled
Dipolydora species more than Polydora species. Polydora cf. woodwicki has chaetiger 5
spines that are similar to those of Dipolydora armata Langerhans, 1880 as described in
Blake (1978) but differ from D. armata in having companion chaetae on chaetiger 5 and
branchiae that extend farther than chaetigers 11-12. These specimens resemble the
description of Polydora woodwicki Blake and Kudenov, 1978 in having a rounded
prostomium but are one fifth the size although mature as some contained gametes.
Other Polydora species with a continuous flange around the concave side of the chaetiger
5 spines are included in the Polydora colonia/spongicola Group of Blake (1996d). This
group includes P. colonia Moore, 1907, P. spongicola Berkeley and Berkeley, 1950 and P.
narica Light, 1969. The group members are commensal on sponges or other polychaetes.
Polydora cf. woodwicki almost certainly belongs with this group. The specimens were
collected from an encrusting sponge on jetty pilings, the other specimen from shelly
gravel at the base of the pilings. They differ from P. narica Light, 1969 in which the
caruncle extends to posterior chaetiger 4 and in the spine form as illustrated for P. narica
Light, 1969 in Blake (1996d). The spine-form is most similar to P. spongicola described by
Blake (1996d) as “typically with broad sub-terminal collar on concave side” but P. cf.
woodwicki differs from P. spongicola in lacking eyes. P. spongicola is reported as having
large re-curved notopodial spines in posterior notopodia (Blake, 1996d). P. colonia is
reported as having no eyes but the spine-form “two unequal teeth and a sub-terminal
collar extending half-way around the spine” (Blake, 1971) is different to Polydora cf.
woodwicki. Further collecting of fresh complete material and examination of type
material is necessary to confirm the identity of these two specimens.
109
Figure 5: Polydora cf. woodwicki (AM W26151), chaetiger 5 spines: light micrograph. Left;
line drawing of same, right
Polydora sp. P1
(Figures 6, 7, 8)
Material examined: QUEENSLAND: Calliope River, 23°51´S, 151° 10´E, QEB code 97, 1974-
1983, coll: P. Saenger (AM W199279 in part, 12 specimens)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 59 (anterior peristomial ventral margin: straight edge with medial v-shaped
notch)
Companion chaetae
Spine tip
Continuous flange
110
Description: Six complete specimens. Largest specimen 4.7mm for 44 chaetigers, 0.5mm
wide at chaetiger 5. Palp missing on largest specimen, on other specimens, palp length
range 0.6-1.2mm.
Anterior prostomial margin weakly or strongly bifid; two pairs of eyes present, anterior
pair farther apart than posterior pair (Fig. 6A); caruncle (prostomial dorsal extension)
extending to anterior chaetiger 2 to mid chaetiger 3; occipital antenna present. Dorsal
ciliary organ present as transverse strip at segment edge; anterior dorsal furrow through
first 11 chaetigers absent; anterior prostomial transverse furrow absent; anterior dorsal
furrow absent; posterior ventral furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin with a single medial lobe; lateral
margin with conspicuous lateral lobes, lateral lobes corner angular (Fig. 6B); ventral
margin straight edge with a medial v-shaped notch.
Chaetiger 5, two to three times length of chaetiger 6 (Fig. 6B); dorsally fused with
chaetiger 6; lateral length 0.2-0.4 times dorsal width.
Pigmentation: palps – absent or base of palp dark grey; parapodia – absent or parapodia
1 and 2 dark grey; prostomium – absent or dark grey; peristomium – absent or grey
pigment; dorsal thoracic – absent or orange/tan pigment patch across chaetiger 6 and 7
on one specimen; ventral thoracic – absent.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to chaetiger 43 in larger specimen; longest at chaetiger
18; overlapping dorsally in anterior segments.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe globular; neuropodial lobe
globular lobe, 0.2x chaetal length. Chaetiger 2-4: conical lobe, 0.5x chaetal length;
neuropodial lobe digitiform, 0.5x chaetae length on chaetigers 2 and 3, reduced on 4.
Chaetiger 5 without noto- or neuropodial lobes. Chaetiger 6+: notopodial lobe posterior
triangular lobe, 0.5x chaetae length, absent after chaetiger 6; neuropodial lobe appears
to be digitiform, 0.5x chaetae length on 6 then reducing posteriorly; posteriorly
111
parapodia eventually becomes an expanded semicircular disc with noto- and neuro-
chaetae running along ridge.
Chaetae: Chaetiger 1: notochaetae absent; neurochaetae capillary chaetae. Chaetiger 2-
4: notochaetae two tiers of broad capillary chaetae, ventral tier shorter, shortest are 0.5x
length of longest capillary chaetae on dorsal tier, long and short strongly geniculate,
chaetae in chaetigers 2-4 have posterior orientation; neurochaetae two tiers, broad
capillary chaetae, ventral tier shorter, shortest 0.5x length of longest capillary chaetae on
dorsal tier, long and short geniculate. Chaetiger 5: dorsally no chaetae; ventrally inferior
ventral tuft of four tiny broad capillary chaetae. Chaetiger 5 spines (Fig. 7): one row of
spines in a shallow curve, endpoints anterior dorsal, posterior ventral; orientation
longitudinal; 5-6 spines; spines simple or bluntly falcate, sub-distal blunt tooth;
companion chaetae present, limbate, brush-tipped, truncated or tapered; distal edge
fibrous; companion chaetae were difficult to see, they appeared to be close and low
between spines. Chaetiger 6+: notochaetae in two tiers, dorsally 2-3 long capillary
chaetae, ventrally a cluster of short broad limbate aristate capillary chaetae to chaetiger
10, then ventral cluster becomes fine again, after chaetiger 10 notochaetae have an
anterior orientation; neuropodia with two tiers of capillary chaetae, similar to chaetiger
4. Neuropodial hooded hooks from chaetiger 7 to end, with constriction on stem
although constriction on hooded hooks is weak in more anterior segments, 2-13 in a row,
2-4 in posterior row, bidentate, angle of main fang to stem <90°, angle not variable in
same row. Notochaetae on posterior 18 segments are splayed out tufts of 2-3 long spine-
like capillary chaetae and approximately 10 straight, spine-like capillary chaetae 0.6 x
length of longer capillary chaetae giving posterior a “sparkly” appearance (Fig. 6C, Fig. 8).
Pygidium: a cuff, raised ring of thin tissue, broadly bilobed ventrally, two? rows of short
cilia on inner edge of lobes, cilia extending just past edge of lobes.
Six specimens with gametogenic segments commencing at chaetiger 15-20. One
specimen with eggs in chaetigers 18-40.
Ecology: tube-dwelling in muddy substrates
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Distribution: Australian: QUEENSLAND: IMCRA bioregion – Tweed-Moreton (Calliope
River, Gladstone)
Remarks: This new species belongs in the Polydora cornuta/nuchalis Group of Blake
(1996), having hooded hooks with a constriction, possessing an occipital antenna, no
notochaetae on chaetiger 1, notopodial capillary chaetae absent on chaetiger 5, major
spines of chaetiger 5 falcate or straight, with an accessory tooth on the concave side and
no lateral flange. It differs from P. cornuta Bosc, 1802 in having a cuff-like pygidium, from
P. cirrosa Rioja, 1943 and P. nuchalis Woodwick, 1953 and Polydora cavitensis Williams,
2007 in having an accessory tooth on chaetiger 5 spines. Polydora sp. P1 differs from all
four in the spinous appearance of the posterior 18 notochaetigers. This spiny posterior is
similar to Polydora woodwicki Blake and Kudenov, 1978 described as having “posterior
notochaetae including both long and short stiff, erect capillary setae, imparting spinous
appearance to the posterior end”. SIMPER analysis results (Appendix 4) indicate that
characters 198 (lateral flange appearing tooth-like), 130(chaetiger 2-4 neuropodial lobes
globular) and 59(as above) contribute most to distinguishing between P. sp P1 and P.
woodwicki. Character 198 is present in P. woodwicki; characters 130 and 59 are present
in P. sp. P1. The two also differ in P. sp. P1 having a bilobed prostomium, lacking a flange
on chaetiger 5 spines and possessing an occipital antenna rather than having a rounded
prostomium, flange on chaetiger 5 spines and no occipital antenna as in P. woodwicki .
There are also similarities to Polydora robi Williams, 2000 which has coiled bundles of
posterior spines rather than the open splayed fascicles of Polydora sp P1.
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Figure 6: Polydora sp. P1 (AM W199279-2) A. whole animal, dorsal view B. anterior,
ventral view C. posterior, dorsal view showing spinous posterior segments
A
B
C
114
Figure 7: Polydora sp. P1, (AM W199279-4) chaetiger 5 spines
Figure 8: Polydora sp. P1, ventral posterior (AM W199279-2)
tooth
Companion chaetae
115
Polydora sp. P2S
Material examined: NEW SOUTH WALES: Sydney Harbour, near N. Chinamens Beach, 8-
10m, 33° 49´S 151° 14.9´E, sand, 08.v.1971, coll: P. Hutchings (AM W13042-8, Polydora
haswelli, PARATYPE)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 196 (chaetae chaetiger 5: form of dorsal spine: wrinkled on convex surface)
Remarks: This is another of the Polydora haswelli Blake and Kudenov, 1978 types that
clustered away from the cluster group containing the P. haswelli holotype. Morphological
variation at different developmental stages and in reproductive condition needs to be
investigated to clarify why the type material is not clustering together.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Polydora sp. P3S
No images available.
Material examined: NEW SOUTH WALES: Sydney Harbour, near N. Chinamens Beach, 8-
10m, 33° 49´S 151° 14.9´E, sand, 08.v.1971, coll: P. Hutchings (AM W13042-1, Polydora
haswelli, PARATYPE)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 157 (chaetae pre-chaetiger 5: chaetiger 2-4: notopodia, number of tiers of
capillary chaetae)
Character 108 (gizzard: chaetiger 17-22)
Remarks: This is another of the Polydora haswelli Blake and Kudenov, 1978 types that
clustered away from the cluster group containing the P. haswelli holotype. Morphological
variation at different developmental stages and in reproductive condition needs to be
116
investigated to clarify why the type material is not clustering together. This specimen was
re-identified as Dipolydora socialis prior to the Part 1 analysis but overall the specimen
clustered more closely with Polydora than Dipolydora and so was included with Polydora
in the final corrected dataset analysis.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Polydora sp. P4S
(Figure 9)
Material examined: NEW SOUTH WALES: Richmond River, Ballina, NSW661, old wharf
between Cherry & Martin Sts., 3m, 28° 52.5’ S, 153° 33.6’ E, loose rocks with silt and
oysters, 05.iii.1992, coll: S. J. Keable, hand collected, SCUBA (AM W26122)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 54 (anterior lateral peristomial margin: lateral edge “3”-shaped (Fig. 9B)
Character 203 (chaetae chaetiger 5: dorsal spine accessory structures: lateral tooth
at curve of stem (Fig. 9A)
Character 140 (parapodial lamellae/lobes: chaetiger 6+: notopodial lobe rounded)
Remarks: This specimen is similar to Polydora woodwicki Blake and Kudenov, 1978 but
differs in having branchiae which extend from chaetiger 7 to 70. P. woodwicki is
described (Blake and Kudenov, 1978) as having branchiae which continue for only 21-23
segments.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Tweed-Moreton
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Figure 9: Polydora sp. P4S (AM W26122), A. chaetiger 5 spines showing lateral tooth at
curve of stem B. anterior ventral showing “3”-shaped peristomial lateral margin
Polydora sp. P5S
(Figure 10)
Material examined: NEW SOUTH WALES: Botany Bay, Channel Marker 4, BB CH4 P1-7,
33 ° 59' 18"S, 151° 12' 36"E, pylon/piling scraping, 21.x.1998, coll: NSW Fisheries (AM
W29649)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 201 (continuous flange joined to spine at neck of fang and along convex spine
edge almost to tip. The flange projects perpendicular to the spine axis a distance almost
equal to the width of the spine. A conspicuous inner tooth meets the flange at the outer
edge (Fig.10).
Character 24 (dorsal ciliary organ: strip across anterior edge of chaetiger 4, posterior to
caruncle)
A
B
Lateral tooth Lateral peristomial
margin
118
Remarks: This specimen resembles Polydora woodwicki Blake and Kudenov, 1978 in
having a truncate prostomium. The chaetiger 5 spines have some resemblance to those
of Polydora hoplura Claparède, 1870 but the specimen lacks an occipital antenna and
falcate posterior spines. The chaetiger 5 spine form and lack of occipital antenna indicate
that it may belong in the Polydora colonia/ spongicola Group (Blake, 1996d) in which the
chaetiger 5 spines have a continuous flange around the concave side of the spine tip.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Figure 10: Polydora sp P5S (AM W29649) chaetiger 5 spines and companion chaetae: light
micrograph left; line drawing of same, right
Polydora sp. P6S
Material examined: NEW SOUTH WALES: Brotherson Dock, Berth 1/1A, Botany Bay, NSW
BB BD1, 33° 58.3´S, 151° 12.4´E, 19.x.1998 (AM W29944)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 137 (chaetiger 6+: notopodia digitiform)
Remarks: This specimen is similar to Polydora cornuta (Bosc, 1802) but lacks on occipital
antenna. The spines resemble those of Dipolydora giardi (Mesnil, 1896) in being falcate
tooth
Continuous flange
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with a subdistal swelling and a lateral flange which may appear tooth-like. The hooded
hooks have a constriction, there are no notochaetae on chaetiger 1 and the angle of the
main fang to the stem of the hooded hooks is <90° so the specimen is within the genus
Polydora. The resemblance analysis indicates that it is closest to the Polydora cf. haswelli
species cluster.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Dipolydora Verrill, 1879
Type species: Polydora concharum Verrill, 1879, designated by Verrill, (1881)
Diagnosis (Blake, 1996d): Prostomium entire or incised anteriorly, extending posteriorly
as caruncle; eyes present or absent. Chaetiger 1 with notochaetae. Chaetiger 5 with
major spines of 1 type, with or without companion chaetae; spines arranged in a single
curved row. Posterior notopodial spines present or absent. Neuropodial hooded hooks
bidentate, usually with a re-curved shaft with constriction and manubrium; main fang
forming wide angle with shaft and narrow, acute angle with apical tooth; hooks present
from chaetiger 7-17. Pygidium disc-like, cuff-shaped, with 2, 3, or 4 lobes of various
forms, or with 4 or more papillae. Anterior part of digestive tract sometimes with
enlarged, thick gizzard-like structure.
SIMPER diagnosis (from this study): SIMPER analysis in this study identified the following
characters as contributing most to the distance between Polydora and Dipolydora:
Character 244 (hooded hooks with constriction on stem) – absent in Dipolydora
Character 152 (chaetae pre-chaetiger 5: chaetiger 1 - notopodia 1 with capillary chaetae)
– present in Dipolydora (except Dipolydora pilocollaris (Blake and Kudenov, 1978))
Character 246 (hooded hooks: maximum number of hooks per chaetiger) – in this study it
appears that maximum number of hooded hooks per chaetiger is greater in Polydora (13
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in Polydora sp. P1) than in Dipolydora (7 in D. tentaculata Blake and Kudenov, 1978) but
this character may be related to development and requires further investigation.
Characters 251and 249 (hooded hooks: angle of main fang to stem <90°) – absent in
Dipolydora.
Character 202 (chaetae chaetiger 5: type of accessory structure - dorsal row - sub-distal
blunt tooth) – absent in Dipolydora specimens in this study, present in the Dipolydora
giardi Group (Blake, 1996d).
Dipolydora cf. aciculata/ cf. giardi
(Figures 11, 12)
Material examined: NEW SOUTH WALES: Cook Island, Temp: 20.5°C., on SCUBA by hand,
Northeast of Mary's Rock, 17m, 28°11.42'S, 153°34.79'E, red alga: Desilia pulchra, coll:
R.T. Springthorpe (AM W26115)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 207 (chaetae chaetiger 5 spines: form of accessory structure on dorsal spine:
accessory tooth on crest) (Fig. 11)
Character 9 (anterior dorsal furrow on first 11 chaetigers present)
Character 23 (dorsal ciliary organ: strip extending posterior to caruncle)
Remarks: The specimen is similar to D. aciculata Blake and Kudenov (1978) in having
branchiae present on anterior 1/3 of body, 4-lobed pygidium, caruncle to posterior
chaetiger 3, posterior unidentate neuropodial spines replacing hooded hooks from
around chaetiger 51 (Fig. 12) and 2 - 3 notopodial acicular spines from around chaetiger
30. The specimen is similar to Dipolydora giardi in the nature of the chaetiger 5 spines,
121
incised prostomium and 1-2 capillary chaetae accompanying hooded hooks in
neuropodia.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Tweed Moreton
Figure 11: Dipolydora cf. aciculata / cf. giardi (AM W26115) chaetiger 5 spines and
companion chaetae as in D. giardi.
Figure 12: Dipolydora cf. aciculata / cf. giardi (AM W26115) posterior neuropodial
acicular spines as in D. aciculata.
Dipolydora flava (Claparède, 1870)
(Figures 13, 14)
Acicular spine
Capillary chaeta
Accessory tooth on spine crest
122
Type locality: Gulf of Naples, Italy; in sandy tube in association with Hyalinoecia rigida
(Polychaeta: Onuphidae)
Material examined: NEW SOUTH WALES: No. 2 Products Wharf, Port Kembla Inner
Harbour, PK PW2 P1-7, 7m, -34.45417, S150.8875E, pylon/piling scraping, 14.v.2000, coll:
NSW Fisheries, CRIMP Ports Integrated Project (AM W31945, 2 specimens); Grain Berth,
Port Kembla Inner Harbour, 3m, -34.45417S, 150.89333E, pylon/piling scraping,
08.v.2000, coll: NSW Fisheries, CRIMP Ports Integrated Project (AM W31947, 4
specimens); No. 1 Products Wharf, Port Kembla Inner Harbour, PK PW2 P1-7, 7m, -
34.46333S, 150.89417E, pylon/piling scraping, 10.v.2000, coll: NSW Fisheries, CRIMP
Ports Integrated Project (AM W31933, in part, 1 specimen)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 131 (parapodial lamellae/lobes: chaetiger 2-4: neuro-broad rounded)
Character 161 (chaetae pre-chaetiger 5: chaetiger 2-4: noto 2-4-glandular lobes with
needle spines)
Character 135 (parapodial lamellae/lobes: chaetiger 5: neuropodial lobe), this seen on
one specimen only
Description: Two specimens with posterior segments but without palps. Specimen in best
condition (AM W31945-1) 10.6mm for 88 chaetigers, 0.5mm wide at chaetiger 5. Palp
missing AM W31945-1), range on other specimens 1-1.5mm. Body opaque, white,
rounded in cross section, chaetae not conspicuous.
Anterior prostomial margin weakly to strongly bifid; two pairs of eyes present, anterior
pair farthest apart; caruncle (prostomial dorsal extension) extending to posterior
chaetiger 3; occipital antenna absent. Dorsal ciliary organs present or absent, when
present a strip on either side of caruncle and as transverse strip at segment edge;
anterior dorsal furrow through first 11 chaetigers absent; anterior prostomial transverse
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furrow absent; anterior dorsal furrow absent; posterior ventral furrow over last 20
chaetigers absent.
Anterior peristomial margins (Fig. 13): inner dorsal margin a single medial lobe, ventral to
this a pair of lobes forming a medial “v” where they meet; lateral margin with lateral
lobes rounded, anterior edge level with base of prostomial lobes; ventral margin a deep
rounded “u”.
Chaetiger 5, 1.7x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length 0.5x
dorsal width.
Pigmentation: variable, either absent or, on one specimen, with heavy black speckling on
prostomium, peristomium, dorsal thoracic, ventral thoracic and posterior segments.
Gizzard absent.
Dorsal branchiae from chaetiger 8 to chaetiger 34 in complete animal; longest at
chaetiger 9-10; overlapping dorsally in anterior chaetigers.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe triangular. Chaetiger 2-4: notopodial lobe conical just less
than 0.5x chaetae length, additional low flattened superior and dorsal glandular lobes
with dark pigmentation; neuropodial lobe chaetigers 2 and 3 with broad rounded
lobe/lamellae 0.2x chaetae length. Chaetiger 5 without noto- or neuropodial lobes, one
specimen with rounded lobe similar to chaetigers 6 and 7. Chaetiger 6+: notopodial lobe
absent; neuropodial lobe chaetigers 6 to 7 with rounded lobe, 0.25x chaetiger length. A
raised ridge runs from notopodia to neuropodia, chaetae sit on anterior edge of segment;
segmentation is deep and segment edges angular.
Chaetigers 4 to 14 ventral surface with a glandular band extending across anterior edge
of segment between parapodia; dorsal surface with paired oval glandular patches from
chaetiger 11.
124
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers of capillary chaetae, ventral tier shorter,
shortest chaetae 0.5x length of longest capillary chaetae on dorsal tier; some glandular
lobes with fine needle spines; neurochaetae two tiers of long and short capillary chaetae,
0.5x length of longer notochaetae. Chaetiger 5: dorsally 2 broad geniculate capillary
chaetae; ventrally 3-5 short capillary chaetae, 0.5 x length of chaetiger 6 neurochaetae.
Chaetiger 5 spines (Fig. 14A): one row, shallow curve, ventral end curved posteriorly;
orientation transverse/longitudinal; 2-4 spines; spines bluntly falcate, sub-distal swelling;
companion chaetae present, limbate, aristate, tapered. Chaetiger 6+: notochaetae two
tiers, long and short broad limbate aristate capillary chaetae, shortest 0.25 length of
longest, some geniculate, strongly directed posteriorly to chaetiger 17, then number of
chaetae reduces to 2-3 long and 2-3 short capillary chaetae per notopodia, directed
anteriorly, only long capillary chaetae conspicuous in most posterior segments, packets of
fine needle spines, occasionally emergent, from chaetiger 19-21 to end, needle spines
emerge from a glandular lobe posterior and ventral to notopodia (Fig. 14B); neuropodia
with capillary chaetae to chaetiger 9-11, number per chaetiger : 5-8,1,1,1,1,1 similar for
other specimens, hooded hooks accompanied by 1 - 3 fine capillary chaetae in last
chaetigers. Neuropodial hooded hooks from chaetiger 7 to end, without constriction on
stem, 1-5 in a row, bidentate, bidentate and unidentate posteriorly, angle of main fang to
stem >90°, angle variable within single row. To about chaetiger 22 chaetae are positioned
equidistant from anterior and posterior edge of segment, after about chaetiger 22 both
neuro- and notochaetae are positioned on angular anterior edge of segment, at around
chaetiger 49 angular edge and chaetae become located in medial position in segment so
that segment lateral edge is strongly angled with chaetae sitting along lateral ridge
formed by the angle.
Pygidium: 3 lobes, single ventral, 2 smaller dorsal; lateral notches closer to dorsal side;
not as wide as final segments.
Gametogenic segments from chaetiger 30 in two specimens.
125
Ecology: pylon scrapings from wharves; in Ostrea lutaria shell (Rainer, 1973); in sandy
tube associated with the onuphid polychaete, Hyalinoecia rigida (Claparède, 1870)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Hawkesbury Shelf;
VICTORIA: IMCRA bioregion – Victorian Embayments; Europe; New Zealand; Indian
Ocean; Japan
Remarks: These Dipolydora flava specimens agree in part with descriptions of Dipolydora
flava (Claparède, 1870) from Australia (Blake and Kudenov, 1978), New Zealand (Rainer,
1973 as Polydora dorsomaculata) and the original description of Claparède (1870). They
differ from the description of Blake and Kudenov (1978) and Claparède (1870) in
possessing eyes. They differ from Blake and Kudenov (1978) and Rainer (1973) and agree
with Claparède (1870) in having needle spines occurring from around chaetiger 19-21
rather than only in posterior regions. They differ from the description of Rainer (1973)
and agree with Blake and Kudenov (1978) in having a three-lobed pygidium rather than a
slightly flared collar with a dorsal notch. Claparède (1870) makes no comment on the
pygidium. Fine needle spines were observed in some specimens from glandular areas on
chaetigers 2-4. This character has not been previously described by Blake and Kudenov
(1978), Rainer (1973) or Claparède (1870). Further molecular research is required to
investigate the variation in material described as D. flava (Claparède, 1870) worldwide.
126
Figure 13: Dipolydora flava (AM W29651-2) anterior ventral view showing peristomium
margins (inner dorsal margin single lobe obscured by ventral lobes).
A
Lateral margin
Ventral margin
Ventral lobe
Simple spine
Companion chaeta
127
Figure 14: Dipolydora flava (AM W29651-2) A. chaetiger 5 spines and companion chaetae
B. emergent packet of needle spines beside base of notopodial capillary chaetae
Dipolydora cf. flava
Material examined: NEW SOUTH WALES: Newcastle, Queens Wharf, AUNTLQWP3-7, 7m,
32.56°S, 151.47°E, scraping, 28.viii.1997 coll: CSIRO-CRIMP (AM W29651, 2 specimens);
Hawkesbury River, New South Wales, -33.5°S, 151.2°E, 2.ii.1982, coll: Hawkesbury River
Survey, 2-2-3 (AM W31957 2-2-3 in part, 1 specimen included in the analysis as AM
W31957 2-2-3.1)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 71 (anterior peristomial ventral margin: broad “u”, base 0.5x segment width
with 5 lobes on edge, width and depth equal)
Character 146 (parapodial lamellae/lobes: chaetiger 6 +: neuropodia-conical (triangular))
Emergent
needle spines
B
Capillary
chaetae
128
Character 24 (dorsal ciliary organ: strip across anterior edge of chaetiger 4, just posterior
to caruncle)
Description: Three specimens one (AM W29651-2) with posterior segments but without
palps. 16.5mm for 105 chaetigers, 0.9mm wide at chaetiger 5. No palps on any specimen.
Body tan, inter-segmental bands conspicuous ventrally.
Anterior prostomial margin strongly bifid; eyes absent; caruncle (prostomial dorsal
extension) extending to posterior chaetiger 3; occipital antenna present. Dorsal ciliary
organs present as a strip on either side of caruncle or as strip across anterior edge
chaetiger 4; anterior dorsal furrow through first 11 chaetigers absent; anterior prostomial
transverse furrow absent; anterior dorsal furrow present; posterior ventral furrow over
last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin a single medial lobe, ventral to this a
pair of lobes forming a medial “v” where they meet; lateral margin with lateral lobes not
enlarged, rounded, lateral edge below more anterior edge; ventral margin broad “u”,
base 0.5x segment width, 5 lobes on edge, depth equal to width.
Chaetiger 5, 2.5x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
0.33x dorsal width.
Pigmentation: palps – unknown, prostomium – absent, peristomium – absent, sparsely
scattered black pigment on dorsal thoracic and ventral thoracic segments.
Gizzard chaetigers 17-22 in complete specimen, 15-19 in incomplete specimens.
Dorsal branchiae from chaetiger 8 to chaetiger 53; longest at chaetiger 20-25; not
overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform, >0.5x chaetae length. Chaetiger 2-4:
notopodial lobe posterior triangular, 0.2x chaetae length; neuropodial lobe triangular
129
0.5x chaetae length on chaetigers 2 and 3, broad on chaetiger 4. Chaetiger 5 without
noto- or neuropodial lobes. Chaetiger 6+: notopodial lobe posterior triangular, 0.2x
chaetae length, reduced posteriorly; neuropodial lobe triangular, 0.2x chaetae length,
absent after chaetiger 8.
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers of capillary chaetae, ventral tier shorter,
shortest chaetae 0.5x length of longest capillary chaetae on dorsal tier; neurochaetae two
tiers of long and short capillary chaetae, 0.25x length of longer notochaetae. Chaetiger 5:
dorsally 4 broad capillary chaetae; ventrally seven thick anterior geniculate chaetae.
Chaetiger 5 spines: one row, straight; orientation transverse; 3-5 spines; spines simple
curved, one specimen with low sub-distal flange; companion chaetae present, limbate,
truncate, aristate or tapered. Chaetiger 6+: notochaetae to chaetiger 17, two tiers, short
broad limbate aristate capillary chaetae, some geniculate, posterior to chaetiger 17, 2-3
long and 2-3 short capillary chaetae per notopodia, long capillary chaetae only in most
posterior segments, notopodial packets of needle spines from chaetiger 25 emerging
from lobe posterior and ventral to notopodia; neuropodia with capillary chaetae to
chaetiger 12, number per chaetiger: 7-8,1,1,1,1,1. Neuropodial hooded hooks from
chaetiger 7, accompanied by 1 - 2 fine capillary chaetae in posterior chaetigers, hooded
hooks absent last few chaetigers, without constriction on stem, 1-5 in single row,
bidentate, bidentate and unidentate posteriorly, angle of main fang to stem ≥90°, angle
variable within single row.
Pygidium: present but damaged.
Gametogenic segments from chaetiger 15-80 in one specimen.
Ecology: Subtidal wharf scrapings, sandy mud, salinity 33-35‰.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Hawkesbury Shelf
130
Remarks: Specimens are similar to the description of Dipolydora flava (Claparède, 1870)
in Blake and Kudenov (1978) but from differ from this description and that of D. flava
from New Zealand (Rainer, 1973 as Polydora dorsomaculata) in having packets of needle
spines from around chaetiger 20 rather than only in posterior segments, capillary chaetae
accompanying hooded hooks in neuropodia posterior to chaetiger 7, and chaetiger 5
spine row straight rather than curved. They also differ from the description by Rainer
(1973) in length of chaetiger 5 compared to surrounding segments being greater (2.5x
rather than 1.5x), and lack of eyespots. They differ from the descriptions of Blake and
Kudenov (1978), Rainer (1973) and Claparède in possessing an occipital antenna. D. cf.
flava differs from Dipolydora flava (Claparède, 1870) as described in this study in having 5
lobes on the peristomial ventral margin, shape of the neuropodial lobes and in lacking
pre-chaetiger 4 needle spines. Dipolydora cf. flava differs from Dipolydora tentaculata
(Blake and Kudenov, 1978) described as having an occipital antenna in possessing packets
of needle spines. D. cf. flava is similar to Dipolydora socialis (Schmarda, 1861) as
described in Blake (1971) and Blake and Kudenov (1978) in possessing a conspicuous
gizzard but differs in having an occipital antenna and packets of needle spines.
Resemblance analysis identifies Polydora socialis as the most similar species cluster group
to D. cf. flava. Characters 71, 24,146 (above) and 40 (anterior peristomial inner dorsal
margin: a single medial lobe, ventral to this a pair of lobes forming a medial “v”)
contribute most to the distance between the two. Dipolydora cf. flava belongs within the
Dipolydora concharum/coeaca/flava/socialis Group of Blake (1996).
Dipolydora cf. pilocollaris 2
Material examined: NEW SOUTH WALES: Richmond River, Ballina,NSW661,old wharf
between Cherry & Martin Sts., 3m, 28° 52.5’ S, 153° 33.6’ E, loose rocks with silt and
oysters, 05.iii.1992, coll: S.J.Keable, hand collected, SCUBA (AM W26119, 3 specimens)
131
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 208 (chaetae chaetiger 5: type of accessory structure - dorsal row: a scale
extending just over convex (outer) terminating in a rough broad edge)
Character 68 (anterior peristomial ventral margin: medial “u”, equal to width of dorsal
lobe, similar depth)
Description: Three specimens, two (AM W29651-2) with posterior segments, one of
these with palps (AM W 26119-2). The complete specimen 2.7mm for 45 chaetigers,
0.3mm wide at chaetiger 5. Palps 1.3mm.
Anterior prostomial margin weakly bifid; eyes absent; caruncle (prostomial dorsal
extension) extending to anterior chaetiger 3; occipital antenna absent. Dorsal ciliary
organs absent; anterior dorsal furrow through first 11 chaetigers absent; anterior
prostomial transverse furrow present; anterior dorsal furrow absent; posterior ventral
furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin with no conspicuous lobes; lateral
margin with lateral lobes rounded, anterior edge level with base of prostomium; ventral
margin medial “u”, equal to width of dorsal lobe, similar depth.
Chaetiger 5, 1.5x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
0.25x dorsal width.
Pigmentation: palps – dark at base of palps, prostomium – absent, scattered black
pigment spots on peristomium, dorsal thoracic and posterior segments.
Gizzard absent.
Dorsal branchiae from chaetiger 7 to chaetiger 32; longest at chaetiger 10; not
overlapping dorsally.
132
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform posterior to broad lamellae, >0.5x chaetae
length. Chaetiger 2-4: notopodial lobe posterior triangular, 0.2x chaetae length;
neuropodial lobe digitiform, 0.3- 0.5x chaetae length, longer on chaetiger 3. Chaetiger 5
without noto- or neuropodial lobes. Chaetiger 6+: notopodial lobe posterior triangular,
0.5x chaetae length, to chaetiger 14, absent posteriorly; neuropodial lobe absent.
Chaetae: Chaetiger 1: notochaetae absent; neurochaetae capillary chaetae. Chaetiger 2-
4: notochaetae two tiers of capillary chaetae, aristate, geniculate; ventral tier shorter,
shortest chaetae 0.5x length of longest capillary chaetae on dorsal tier; neurochaetae
spreading fascicle of capillary chaetae. Chaetiger 5: dorsally 4 broad capillary chaetae, in
close proximity to spine row; ventrally seven geniculate chaetae. Chaetiger 5 spines: one
row, straight; orientation longitudinal; 3 spines; spines falcate, sub-distal swelling, a scale
extending just over convex (outer) terminating in a rough broad edge; companion
chaetae present, geniculate, sigmoid, tapered, aristate. Chaetiger 6+: similar to anterior
notopodia to chaetiger 8, then chaetae fewer and finer to chaetiger 14, then 2-3 long and
2-3 short capillary chaetae per fascicle; neuropodia with capillary chaetae to chaetiger 8,
number per chaetiger: 5-6,1,1; hooded hooks accompanied by 1 - 3 fine capillary chaetae
in last chaetigers. Neuropodial hooded hooks from chaetiger 7 to end, without
constriction on stem, 1-5 in single row, bidentate, bidentate and unidentate posteriorly,
angle of main fang to stem >90°, angle not variable within single row.
Pygidium: four lobes; 2 ventral, 2 dorsal, ventral lobes smaller.
Gametogenic segments absent.
Ecology: loose rocks with silt and oysters
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion – Tweed-Moreton
133
Remarks: Specimens are similar to Dipolydora pilocollaris (Blake and Kudenov, 1978) in
parapodial lobe characters and size (AM W17068, paratype: 3.4mm for 21 chaetigers,
0.7mm wide, posterior missing). D. cf. pilocollaris2 differs from the paratype in having
branchiae which commence at chaetiger 7 not chaetiger 8. It differs from the description
in Blake and Kudenov (1978) in having branchiae absent in posterior quarter rather than
posterior half and in the ventral pair of pygidial lobes being smaller than the dorsal pair
rather than the reverse. D. cf. pilocollaris2 differs from D. cf. pilocollaris in Wilson et al.
(2003) in having branchiae on more than the first 20 chaetigers and in not occurring on
abalone shells. D. cf. pilocollaris2 shares the SIMPER diagnostic character 208 (above)
with the paratype (AM W17068) but this character differs from the description of D.
pilocollaris in Blake and Kudenov (1978) and of D. cf. pilocollaris in Wilson et al. (2003) in
lacking a prominent tuft of bristles from the scale on the concave surface of the chaetiger
5 spine. D. cf. pilocollaris is similar to Dipolydora convexa (Blake and Woodwick, 1971) in
lacking the tuft of bristles on chaetiger 5 spines and in having unidentate hooks
posteriorly, but differs from D. convexa (Blake and Woodwick, 1971) in lacking posterior
armature. Examination of fresh material to assess 1) if the lack of bristles on the chaetiger
5 spines is an artefact of storage and 2) the variation in branchial distribution will help to
clarify the position of these specimens.
Dipolydora socialis (Schmarda, 1861)
(Figures 15, 16)
Type locality: Viña del Mar, Chile
Material examined: NEW SOUTH WALES: Hawkesbury River, -33.5S, 151.2E, Hawkesbury
River Survey, 2-1-3.1, 20.viii.1980 (AM W31957 2-1-3, 3 specimens, one juvenile);
Hawkesbury River, -33.5S, 151.2E, Hawkesbury River Survey, 2-2-2, 12.i.1977 (AM
W31957 2-2-2, one specimen); Hawkesbury River, -33.5S, 151.2E, Hawkesbury River
Survey, 2-2-3, 2.ii.1982 (AM W31957 2-2-3, 2 specimens); Hawkesbury River, -33.5S,
151.2E, Hawkesbury River Survey, 2-2-4, 17.v.1982 (AM W31957 2-2-4 .1); Newcastle,
134
Queens Wharf, New South Wales, AUNTLQWP3-6, 6m, -32.9S, 151.8E, 28.viii.1997, coll:
CSIRO-CRIMP (AM W31920); QUEENSLAND: Calliope River, Queensland, 23° 51´S, 151°
10´E, QEB 1974-1983, coll: P. Saenger (AM W199279 in part, 5 specimens included in
analysis as AM W199279-3, AM W199279-10, AM W199279-11, AM W199279-16, AM
W199279-24); Bramble Bay, SEQ #13, 26° 06´S, 152° 17´E, vi 1973, coll: S. Cook (QM
G10642-3)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 125 (parapodial lamellae/lobes: chaetiger 2-4: noto-rounded)
Character 106 (gizzard chaetiger: 13-16)
Description: Two adult specimens with posterior, both without palps. Largest 8.9mm for
51 chaetigers. 0.6mm wide at chaetiger 5. Palp length range 0.2mm on 2mm long juvenile
specimen, 2.7-2.9mm on larger specimen.
Anterior (Fig. 15) prostomial margin weakly to strongly bifid; eyes absent; caruncle
(prostomial dorsal extension) extending to anterior margin of chaetiger 4 on average,
range from posterior chaetiger 2 to posterior chaetiger 4; occipital antenna absent.
Dorsal ciliary organs present as strip on either side of caruncle; anterior dorsal furrow
through first 11 chaetigers absent; anterior prostomial transverse furrow absent; anterior
dorsal furrow present; posterior ventral furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin a single medial lobe; lateral margin
with lateral lobe not notably enlarged; ventral margin broad “u”, base 0.5x segment
width, almost as deep as it is wide.
Chaetiger 5, 1.25x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
0.4x dorsal width.
Pigmentation: variable, either absent or with sparsely scattered black pigment spots on
dorsal thoracic and ventral thoracic segments or over whole length of body.
135
Gizzard visible over 3-4 segments usually from chaetiger 13 but occasionally from
chaetiger 15, and in one specimen from chaetiger 25.
Dorsal branchiae from chaetiger 8 to end in complete animal; several specimens with
branchiae from chaetiger 9; longest at chaetiger 16; not overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform, more than 0.5x chaetae length. Chaetiger 2-4:
notopodial lobe conical 0.2x chaetae length; neuropodial lobe triangular 0.33x chaetae
length on chaetigers 2 and 3, reduced and broad on chaetiger 4. Chaetiger 5 without
noto- or neuropodial lobes. Chaetiger 6+: notopodial lobe posterior triangular, 0.2x
chaetae length, reducing posteriorly; neuropodial lobe absent.
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers, dorsal long fine aristate capillary chaetae
and ventral geniculate and aristate capillary chaetae 0.5x length of dorsal capillary
chaetae, some tips strongly geniculate; neurochaetae two tiers of long and short capillary
chaetae, 0.5 length of longer notochaetae. Chaetiger 5: dorsally 4 anterior broad thick
capillary chaetae; ventrally 4 broad thick capillary chaetae similar to short capillary
chaetae of neurochaetae on chaetiger 4. Chaetiger 5 spines (Fig. 16): one row, shallow
curve, ventral end curved posteriorly; orientation longitudinal; 6 spines; spines simple,
curved, slight sub-distal swelling; companion chaetae present, limbate, sigmoid,
truncated, pennoned, aristate, tapered. Chaetiger 6+: notochaetae two tiers, dorsal tier
3-4 long capillary chaetae, ventral tier a cluster of short broad limbate aristate capillary
chaetae 0.3-0.5 length of longer chaetae in dorsal tier, some geniculate, this pattern
continues to posterior but number of chaetae reduces to 2-3 long and 2-3 short capillary
chaetae per notopodia; neuropodial capillary chaetae to chaetiger 8-10, number per
chaetiger: 5-7,2,2,2,2 similar for other specimens, hooded hooks accompanied by 1 - 3
fine capillary chaetae in last chaetigers. Neuropodial hooded hooks from chaetiger 7 to
end, without constriction on stem, 1-5 in a row, bidentate, bidentate and unidentate
posteriorly, angle of main fang to stem ≥90°, angle variable within single row.
136
Pygidium: On larger specimen, brush like disc flared ventrally, golden with fibrous edge;
two lobes dorsally; larger than last posterior chaetiger. All other specimens with damaged
or missing posteriors.
Gametogenic segments from chaetiger 15-20 in three specimens.
Ecology: coarse sand, soft or sandy mud; Posidonia seagrass beds; intertidal to about
400m (Blake, 1996d)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregions: Hawkesbury;
QUEENSLAND: IMCRA bioregions: Tweed-Moreton; east and west coasts of North
America; Gulf of Mexico; Chile; Falkland Islands; western Pacific, the Sea of Japan
Remarks: These Australian specimens agree with the description of Dipolydora socialis
(Schmarda, 1861) in Blake (1996d). However, the specimens and description differ from
the illustration of Leucodore socialis from the type locality (Schmarda, 1861). The
illustration indicates a weakly bi-lobed prostomium, no caruncle and branchiae on all
chaetigers except the modified chaetiger which is chaetiger 6, not chaetiger 5 as in other
polydorids. Manchenko and Radashevsky (2002) report that in the original description by
Schmarda (1861) and the most recent redescription of the type material by Mesnil (1896)
pygidium shape and caruncle length is not mentioned. Re-examination of material from
the type location is required to confirm that Schmarda (1861) and Blake (1971; 1996d)
have described the same species and to be certain of the identity of Australian material
known as Dipolydora socialis (Schmarda, 1861). Until further research confirms
otherwise the description of Dipolydora socialis (Schmarda, 1861) in Blake (1996d) is
accepted.
This study examined only a small proportion of the large amount of material identified as
Dipolydora socialis in Australian museum collections. One new species, Dipolydora sp. D1,
was discovered. It is expected that additional new species are likely to be found with
further examination of this material.
137
Figure 15: Dipolydora socialis (AM W13037) anterior dorsal
Figure 16: Dipolydora socialis (AM W31957 2-1-3.2) chaetiger 5 spines and companion
chaetae
caruncle
branchia
Gizzard
(internal)
Simple spine
Companion chaetae
138
Dipolydora tentaculata (Blake and Kudenov, 1978)
Type locality: Botany Bay, Towra Beach, New South Wales Australia; in Halophila zone
Material examined: NEW SOUTH WALES: Towra Beach, Botany Bay, New South Wales,
BB321, 34° 03´S, 151° 10´E, Halophila zone, iv 1973, coll: R. Rands, NSW State Fisheries
(HOLOTYPE MV G2885, PARATYPE MV G2886).
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 104 (caruncle (prostomial dorsal extension): mid 5)
Character 43 (anterior peristomial inner dorsal margin: no conspicuous lobe, a pair of
lobes between lateral lobes, 0.5x height of u-shaped division formed between lateral
lobes)
Character 42 (anterior peristomial inner dorsal margin: a single medial lobe, ventral to
this a pair of lateral lobes forming a medial narrow “v”)
Description: Two specimens, holotype and paratype, posterior and palps missing.
Holotype 19.9mm for 80 chaetigers, 1.2mm wide at chaetiger 5. Palp length unknown.
Body appearing papillated over much of surface.
Anterior prostomial margin strongly bifid; eyes absent; caruncle (prostomial dorsal
extension) extending to mid chaetiger 5; occipital antenna present. Dorsal ciliary organs
absent; anterior dorsal furrow through first 11 chaetigers absent; anterior prostomial
transverse furrow absent; anterior dorsal furrow absent; posterior ventral furrow over
last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin a single medial lobe, not always
conspicuous, ventral to this a pair of lateral lobes forming a medial narrow “v” 0.5x height
of U-shaped division formed between lateral lobes; lateral margin with conspicuous
lateral lobes; ventral margin semicircular scallop (“u”-shaped), base level with anterior
chaetiger 1.
139
Chaetiger 5, 1.5x length of chaetiger 6; not dorsally fused with chaetiger 6, overlapping
chaetiger 6 but not fused; lateral length 0.33x dorsal width.
Pigmentation: palps – absent; parapodia – absent; prostomium – absent; peristomium –
absent; dorsal thoracic – absent; ventral thoracic – absent.
Gizzard from chaetiger 17-22.
Dorsal branchiae from chaetiger 8 to at least chaetiger 80; longest at chaetiger 22; not
overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe conical, similar in size to
neuropodial lobe; neuropodial lobe broadly triangular lobe greater than 0.5x chaetae
length. Chaetiger 2-4: notopodia broad triangular lamellae less than 0.5x length of
chaetae and a long digitiform like lobe 0.5x length of chaetae; neuropodial lobe broadly
triangular lobe less than 0.5x length of chaetae, similar to notopodia, appears to increase
in size from chaetiger 2 to chaetiger 4. Chaetiger 5 without noto- or neuropodial lobes.
Chaetiger 6+: notopodial lobe triangular lobe 0.2x length of chaetae; neuropodial lobe
absent.
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers of capillary chaetae, ventral tier shorter,
shortest chaetae 0.5x length of longest capillary chaetae on dorsal tier; neurochaetae two
tiers of short capillary chaetae 0.5x length of longest notochaetae. Chaetiger 5: dorsally 4
anterior short broad capillary chaetae; ventrally 5 capillary chaetae. Chaetiger 5 spines:
one row, almost straight; orientation longitudinal; 6-7 spines; spines simple, slight
subdistal swelling; companion chaetae present, limbate, truncated, with distal fibrous
edge. Chaetiger 6+: notochaetae similar to anterior notochaetae to about chaetiger 13,
then fewer notochaetae per chaetiger and reduced in size, more posterior notopodia
with 4 long capillary chaetae, 2 medium length capillary chaetae and several shorter
capillary chaetae; neuropodial capillary chaetae to chaetiger 14, number per chaetiger to
chaetiger 14: 8,0,0,2,0,0,2, 2,1. Neuropodial hooded hooks from chaetiger 7 to at least
chaetiger 80, without constriction on stem, 4-7 in a row, bidentate, angle of main fang to
stem <90°, angle variable within single row.
140
Pygidium: missing
No gametogenic segments.
Ecology: Halophila zone
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion - Hawkesbury Shelf;
QUEENSLAND: IMCRA bioregion - Tweed-Moreton (Blake and Kudenov, 1978)
Remarks: Agrees with original description in Blake and Kudenov (1978). Specimens from
Moreton Bay (QM G11597) not examined.
Dipolydora sp. D1
(Figures 17, 18, 19)
Material examined: NEW SOUTH WALES: Hawkesbury River, - 33.5S, 151.2E, 27.v.1983,
Hawkesbury River Survey 2-2-3 (AM W31957-2.3.3, 6 specimens); Hawkesbury River, -
33.5S, 151.2E, 12.i.1977, Hawkesbury River Survey 2-2-4 (AM W31957, in part, 2
specimens analysed in this study as AM W31957 2-3-3.1.77+AM W31957 2-3-3.2.77);
QUEENSLAND: Calliope River, Queensland, 23° 51´S, 151° 10´E, coll: P. Saenger, QEB
1974-1983 (AM W199279, in part, 2 specimens analysed in this study as AM W199279-5
and AM W199279-6)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 223 (chaetae chaetiger 5: companion chaetae type/s: pennoned strongly
geniculate, tapered)
Character 160 (chaetae pre-chaetiger 5: chaetiger 2-4: noto 4-some capillary chaetae
with fibrous edge)
141
Description: Two complete specimens. Largest complete specimen (AM W31957- 2-3-3.1)
8.3mm for 38 chaetigers, 0.8mm wide at chaetiger 5. Palp length 4.1mm. Body is
translucent, palps broad translucent ribbons (Figure 17) with tapered tips, ciliary strip a
fine line.
Anterior prostomial margin strongly bifid; two pairs of eyes present, anterior pair farthest
apart (Figure 18); caruncle (prostomial dorsal extension) extending to posterior chaetiger
4; occipital antenna absent. Dorsal ciliary organs present as strip on either side of
caruncle and as transverse strip at segment edge; anterior dorsal furrow through first 11
chaetigers absent; anterior prostomial transverse furrow absent; anterior dorsal furrow
present; posterior ventral furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin a single medial lobe, ventral to this a
pair of laterally flattened lobes meeting at anterior 0.33x of length forming a medial “v”,
continuing closely parallel, lateral lobes rounded, anterior edge level with base of
prostomial lobes; lateral margin with conspicuous lateral lobes; ventral margin broad “v”
becoming a narrow “v”-shaped slit medially. Anterior ventral edge of chaetiger 1 a deep
rounded “u” .
Chaetiger 5, 1.8x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length 0.4x
dorsal width.
Pigmentation: variable, absent on largest complete specimen, palps – absent; parapodia
– brown spot at base of neuropodia to chaetiger 15; prostomium – fine brown line at
edge of furrow between lobes; peristomium – sparse minute brown spots anterior
lateral; dorsal thoracic – paired fine brown lines medially to chaetiger 6 -15; additional
lateral spot closer to neuropodia; chaetigers 1-5 with spots rather than lines; ventral
thoracic – medial spot chaetigers 3-4; paired lines on 5; paired lateral spots chaetiger 6.
Gizzard absent.
Dorsal branchiae from chaetiger 8 to chaetiger 36 in complete animal, other specimens
branchiae commence on 7 or 9; longest at chaetiger 13; not overlapping dorsally.
142
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe triangular. Chaetiger 2-4: notopodial lobe conical 0.2x
length of chaetae; neuropodial lobe a triangular lobe 0.2x length of chaetae on chaetigers
2 and 3, reduced and broad on chaetiger 4. Chaetiger 5 without noto- or neuropodial
lobes. Chaetiger 6+: notopodial lobe posterior triangular , 0.2x length of chaetae,
reducing posteriorly; neuropodial lobe absent.
Chaetigers 4 to 14 ventral surface with a glandular band extending across anterior edge
of segment between parapodia; dorsal surface with paired oval glandular patches from
chaetiger 11.
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers of capillary chaetae, ventral tier with
shorter capillary chaetae, shortest 0.5x length of longest capillary chaetae on dorsal tier,
some notochaetae on chaetiger 4 with fibrous edge, some geniculate; neurochaetae two
tiers of long and short capillary chaetae, 0.5x length of longer notochaetae. Chaetiger 5:
dorsal fascicle absent; ventrally 5-6 anterior capillary chaetae similar to more anterior
chaetigers. Chaetiger 5 spines (Figure 19B): one row, but companion chaetae conspicuous
and may initially appear to be two spine rows; row almost straight, ventral end curved
posteriorly; orientation transverse; 12 spines; spines simple, pipette-shaped tip
(extended, tapering but tip blunt), slight sub-distal swelling; companion chaetae present,
distally swollen, strongly asymmetrical, tapering to one side at almost right angles.
Chaetiger 6+: notochaetae short bilimbate capillary chaetae to chaetiger 10, several in
each bundle having a fibrous edge, then number of capillary chaetae reducing from
chaetiger 8 or 9 to fascicles composed of 1-2 longer capillary chaetae and 5-7 capillary
chaetae varying from 0.5 - 0.75x length of longer capillary chaetae, some with finely
fibrous edge; posteriorly up to 5 long fine capillary chaetae, sometimes with fibrous edge;
neuropodia capillary chaetae to chaetiger 13, number per chaetiger for largest specimen:
11,6,6,6,6,4,2,1, similar for other specimens. Neuropodial hooded hooks (Figure 19A)
from chaetiger 7 to end, without constriction on stem, 1-5 in a row, bidentate, bidentate
and unidentate posteriorly, angle of main fang to stem ≥90°, angle variable within single
row.
143
Pygidium: 2 lateral lobes, shallow, not wider than final chaetigerous segment, two
notches, ventral deeper.
Gametogenic segments 13-37.
Ecology: sandy mud substrates
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury Shelf;
QUEENSLAND: IMCRA bioregion: Tweed-Moreton
Remarks: This species belongs to the Dipolydora concharum/coeaca/flava/socialis Group
of Blake (1996d). It is similar to Dipolydora socialis (Schmarda, 1861) in Blake (1996d) but
differs from it in having no visible gizzard, in the form of the pygidium which has 2 lobes
not three and in the form of the companion chaetae which are conspicuous, pennoned,
strongly geniculate and tapering. SIMPER analysis (Appendix 6) identifies characters 223
and 160 (both described above), 65 (anterior peristomial ventral margin: broad “v”
becoming a narrow slit medially) and 193 (chaetae chaetiger5: form of dorsal spine:
pipette-tip (flattened)) as contributing most to the difference between D. socialis and D
sp. D1.
Figure 17: Dipolydora sp. D1 (AM W31957 2-3-3.1) whole animal, dorsal view
prostomium
palps
144
Figure 18: Dipolydora sp. D1 (AM W31957 2-3-3.1) anterior ventral, left; anterior dorsal,
right. Line drawings of same below.
Grooved
palp
Grooved palps
Peristomial ventral margin Chaetiger 1 ventral margin
Eyes
145
Figure 19: Dipolydora sp. D1 (AM W31957 2-3-3.1) A. hooded hook row, mid-body B.
chaetiger 5 spines and companion chaetae
Dipolydora sp. D2
Material examined: NEW SOUTH WALES: Newcastle, Merewether Berth dock,
AUNTLMBP1-3, 3m, -32.93°S, 151.76°E, 27.vii.1997, coll: CSIRO-CRIMP (AM W31921, 1
specimen); Newcastle, Dyke Berth 4 AUNTLD4P2-7, 7m, 32.55°S, 151.48°E, 27.viii.1997,
coll: CSIRO-CRIMP (AM W29950, 1 specimen)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
A
B
Simple spines
Companion chaetae
146
Character 69 (anterior peristomial ventral margin: shallow open “u”)
Character 102 (caruncle (prostomial dorsal extension): mid 4)
Description: Two adult specimens, one with posterior, both without palps. Largest 9.6mm
for 68 chaetigers. 0.5mm wide at chaetiger 5. Palps missing. Body thickened and opaque,
many segments with barrel-like structure (reproductive?).
Anterior prostomial margin strongly bifid; eyes absent; caruncle (prostomial dorsal
extension) extending to mid-chaetiger 4; occipital antenna absent. Dorsal ciliary organs
present as strip on either side of caruncle; anterior dorsal furrow through first 11
chaetigers absent; anterior prostomial transverse furrow absent; anterior dorsal furrow
absent; posterior ventral furrow over last 20 chaetigers absent.
Anterior peristomial margins: inner dorsal margin a single medial lobe, ventral pair of
lobes forming a medial “v”; lateral margin lateral lobe not notably enlarged; ventral
margin broad “u”, base 0.5x segment width, almost as deep as it is wide.
Chaetiger 5, 1.75x length of chaetiger 6; dorsally fused with chaetiger 6; lateral length
0.35x dorsal width.
Pigmentation: parapodia – orange tips, sparsely scattered black pigment spots on
peristomium, ventral thoracic from chaetiger 9 and around posterior.
Gizzard visible from chaetiger 25 - 29. 15-19 on smaller specimen.
Dorsal branchiae from chaetiger 9 to end in complete animal; longest at chaetiger 15; not
overlapping dorsally.
Parapodial lamellae/lobes: Chaetiger 1: notopodial lobe digitiform, similar to lobe on
neuropodia 1; neuropodial lobe digitiform, more than 0.4x chaetae length. Chaetiger 2-4:
notopodial lobe conical 0.2x chaetae length, longer on chaetigers 1 and 3, broad on 4;
neuropodial lobe triangular 0.33x chaetae length on chaetigers 2 and 3, reduced and
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broad on chaetiger 4. Chaetiger 5 without noto- or neuropodial lobes. Chaetiger 6+:
notopodial lobe absent; neuropodial lobe absent.
Chaetae: Chaetiger 1: notochaetae present, capillary chaetae; neurochaetae capillary
chaetae. Chaetiger 2-4: notochaetae two tiers, capillary chaetae, ventral tier 0.5x longest
capillary chaetae on dorsal tier; neurochaetae two tiers of long and short capillary
chaetae, 0.25x length of longer notochaetae. Chaetiger 5: dorsally 4 anterior broad thick
capillary chaetae; ventrally three superior broad geniculate capillary chaetae. Chaetiger 5
spines: one row, straight; orientation transverse/perpendicular; 5 spines; spines simple,
curved, slight sub-distal swelling; companion chaetae present, limbate, aristate, tapered.
Chaetiger 6+: notochaetae two tiers, dorsal tier 3-4 long capillary chaetae, ventral tier a
cluster of short broad limbate aristate capillary chaetae 0.3-0.5 length of longer chaetae
in dorsal tier, some geniculate, this pattern continues to posterior but number of chaetae
reduces to 2-3 long and 2-3 short capillary chaetae per notopodia; neuropodial capillary
chaetae to chaetiger 9, number per chaetiger: 7,1,2,1 similar for other specimen, hooded
hooks accompanied by 1 - 3 fine capillary chaetae in last chaetigers. Neuropodial hooded
hooks from chaetiger 7 to end, without constriction on stem, 2-4 in a row, bidentate,
bidentate and unidentate posteriorly, angle of main fang to stem ≥90°, angle variable
within single row.
Pygidium: not noted
Gametogenic segments, body thickened and opaque, many segments with barrel-like
structure (reproductive?).
Ecology: docking berth, subtidal, scrapings (?)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregions: Hawkesbury
Remarks: These two specimens resemble the Dipolydora socialis species cluster more
than any other species cluster in the analysis. SIMPER analysis identified characters 69
and 102 (above) as making an equal percentage contribution to the distance between D.
socialis species cluster and D. sp D2. Characters 109 (gizzard chaetigers 25-29) and 40
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(single medial lobe, ventral pair of lobes forming a medial “v”) are the next most
important characters. D. protuberata (Blake and Kudenov, 1978) and D. tentaculata
(Blake and Kudenov, 1978) also are described as having a gizzard-like structure at
chaetigers 17-18 and 19-20 respectively. No posterior notopodial spines were present as
in D. protuberata (Blake and Kudenov, 1978) and no occipital antenna was present as in
D. tentaculata (Blake and Kudenov, 1978). It is possible that these specimens are showing
morphological changes due to reproductive state. There is also a possibility of
environmental pollution in this location. Changes may be due to toxicological effects.
Once again studies into both these areas would be useful in understanding morphological
variation in Australian Dipolydora socialis and species with very similar form.
SIMPER diagnostic characters of additional subtropical species of Dipolydora represented
by only a single individual.
Dipolydora sp. D3S
Material examined: NEW SOUTH WALES: Botany Bay (AM W23667)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 95 (caruncle (prostomial dorsal extension): to anterior chaetiger 2)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Remarks: This specimen is another resembling Polydora haswelli but with hooded hooks
without a constriction.
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Dipolydora sp. D4S
Material examined: NEW SOUTH WALES: Sydney Harbour, near N. Chinamens Beach, 8-
10m, 33° 49´S 151°S 14.9´E, sand, 08.v.1971, coll: P. Hutchings (AM W13042-2, Polydora
haswelli PARATYPE)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 195 (chaetae chaetiger 5: accessory structure on dorsal spine: sub-distally
inflated)
Remarks: This specimen is contracted, covered with formalin crystals and branchiae are
stuck onto body- only line of vascular tissue obvious. This poor condition may have led to
its distance from the other P. haswelli types.
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Hawkesbury
Dipolydora D5S
Material examined: QUEENSLAND: Middle Banks, northern Moreton Bay, 27° 11´S, 153°
19´ E, Nov 83-Nov 84, coll: P. Saenger and S. Park (QMunreg1)
SIMPER diagnosis: reveals that the following characters are most important in defining
the species.
Character 225 (chaetae chaetiger 5: companion chaetae type/s: fine spines)
Character 228 (chaetae chaetiger 5: companion chaetae type/s: feathery and bifurcate)
Distribution: Australia: NEW SOUTH WALES: IMCRA bioregion: Tweed-Moreton
Remarks: This large specimen (12.8mm for 93 chaetigers) is another belonging in the
Dipolydora concharum/coeca/flava/socialis Group (Blake, 1996d).Previous to this study
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the specimen had been identified as D. tentaculata but there is no evidence of an
occipital tentacle. It is similar to D. flava in having packets of needle spines but the
resemblance analysis indicates that it is more similar to D. pilocollaris type material. It
differs from D. pilocollaris in characters 225 and 228 (above) and in not having the scale-
like tooth over the convex surface of the chaetiger 5 spines. Very little material is
available from Moreton Bay; further collecting would almost certainly reveal new species.
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Distributions of Polydora and Dipolydora in estuaries of subtropical eastern Australia
Twelve Polydora species and 10 Dipolydora species were found in material examined
from subtropical eastern Australia. Distribution maps for the species found in this study
based around IMCRA bioregions are provided in Figures 20 and 21 respectively.
The greatest number of Dipolydora species occurred in the more northern bioregion
while the greatest number of Polydora species occurred in the more southern bioregion
(excluding Batemans Shelf for which no specimens were located in museum collections).
The present study has identified 6 Dipolydora species and 5 Polydora species from the
Tweed-Moreton IMCRA bioregion.
Four species of Dipolydora and a single Polydora species, Polydora cf. calcarea, were
found in collections from the Manning Shelf IMCRA bioregion.
Four Dipolydora species and 6 Polydora species were found in collections from the
Hawkesbury Shelf IMCRA bioregion.
No Polydora or Dipolydora material was included in the analysis from the Batemans Shelf
IMCRA bioregion. Material was examined but it was not suitable for the analysis.
Dipolydora socialis (Schmarda, 1861), Dipolydora sp. D1 and Dipolydora tentaculata
(Blake and Kudenov, 1978) were the only Dipolydora species found in more than one
bioregion. Dipolydora socialis (Schmarda, 1861) and Dipolydora sp. D1 occurred in the
Tweed-Moreton and neighbouring Manning Shelf bioregions. Dipolydora tentaculata
(Blake and Kudenov, 1978) occurred in the Hawkesbury Shelf and Tweed-Moreton
bioregions. No Polydora species was found in more than one bioregion.
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TMN Tweed- Moreton north to 24°S
MAN Manning Shelf north to 30° 39’S
HAW Hawkesbury Shelf north to 32° 54’S
BAT Batemans Shelf north to 34° 35’S to 36° 48’S
IMCRA Bioregions subtropical climate zones
Tweed-Moreton
Polydora hoplura Claparède, 1870
Polydora cf. websteri
Polydora cf. woodwicki
Polydora sp. P1
Polydora sp. P4S
Manning Shelf
Polydora cf. calcarea
Hawkesbury Shelf
Polydora cornuta Bosc, 1802
Polydora haswelli Blake and Kudenov, 1978
Polydora cf. latispinosa
Polydora sp. P2S
Polydora sp. P5S
Polydora sp. P6S
Figure 20: Distribution of Polydora species in estuaries of subtropical eastern Australia.
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TMN Tweed- Moreton north to 24°S
MAN Manning Shelf north to 30° 39’S
HAW Hawkesbury Shelf north to 32° 54’S
BAT Batemans Shelf north to 34° 35’S to 36° 48’S
IMCRA Bioregions in subtropical climate zones
Tweed-Moreton
Dipolydora socialis (Schmarda, 1861)
Dipolydora tentaculata (Blake and Kudenov, 1978)
Dipolydora cf. aciculata / cf. giardi
Dipolydora cf. pilocollaris 2
Dipolydora sp. D1
Dipolydora sp. D5S
Manning Shelf
Dipolydora socialis (Schmarda, 1861)
Dipolydora cf. flava
Dipolydora sp. D1
Dipolydora sp. D2
Hawkesbury Shelf
Dipolydora flava (Claparède, 1870)
Dipolydora socialis (Schmarda, 1861)
Dipolydora tentaculata (Blake and Kudenov, 1978)
Dipolydora sp. D3S
Dipolydora sp. D4S
Figure 21: Distribution of Dipolydora species in estuaries of subtropical eastern Australia.
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CHAPTER 5: DISCUSSION
The aim of this thesis was to review the current state of knowledge, occurrence and
distribution of two polydorid genera, Polydora and Dipolydora, in estuaries of subtropical
eastern Australia. A novel approach using resemblance analysis of a standardized set of
multivariate morphological characters was used to create species cluster groups. The
same dataset was used to identify the diagnostic characters for each of these species
cluster groups and to indicate previously overlooked characters which could be useful in
species diagnosis, particularly for incomplete specimens. This review process was
commenced with the intention of identifying base knowledge and knowledge gaps in
order to plan a strategy to address these gaps in the future.
What information is currently available on polydorid spionids of east coast
subtropical estuaries and bays?
The first comprehensive work on the polydorid spionids of Australia was that of Blake and
Kudenov (1978) who described the Spionidae of south-eastern Australia. They found a
high level of endemicity within the south-eastern Australian spionid fauna and concluded
their work anticipating further discoveries of new species, noting that the shell-boring
habitat, sponge and coral microhabitats had not been investigated at all in Australia.
These concluding comments are largely still valid. Surprisingly, only one additional species
(Dipolydora penicillata (Hutchings and Rainer, 1979) suggested to be synonymised with
Carazziella victoriensis Blake and Kudenov, 1978 in this study) has been described in the
30 years since that time even though species lists and specimens deposited at museums
from ecological studies, long- and short- term monitoring of coastal marine environments
and adventitious collecting events indicate that specimens defying identification using
existing literature have been found, eg. McDiarmid et al. (2004). A comparison of
worldwide numbers of Polydora species to Dipolydora species (Table 1) indicates that the
number of described Australian Polydora species is low compared to the number of
described Dipolydora species.
The most recent research into reproductive behaviour in Polydora and Dipolydora species
in Australia concerned polydorid mudworms was that of Skeel (1977; 1978; 1979). The
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mudworms, particularly Polydora species, are considered a serious pest of commercially
important bivalve fisheries to the extent that culture methods for oysters are specifically
designed to address the mudworm problem. The studies of the 1970s reported briefly on
Polydora haswelli, Polydora hoplura and Polydora websteri from Australian oysters. This
research in combination with the earlier research of Whitelegge (1890) and Roughley
(1922; 1925) form the basis of various advice sheets written since that time (Nell, 2001;
2007a; 2007b). Dipolydora and Polydora species in Australia for which information on
reproductive behaviour is unavailable are: Polydora latispinosa, Polydora woodwicki,
Dipolydora aciculata, Dipolydora armata, Dipolydora giardi, Dipolydora flava, Dipolydora
notialis, Dipolydora pilocollaris, Dipolydora protuberata, Dipolydora socialis and
Dipolydora tentaculata. Reproductive strategies in Australian species of Polydora and
Dipolydora and the same species from other locations are given in Appendix 1. Although
not all polydorids are of commercial importance, information on reproduction, larval and
juvenile development for all polydorid species would provide invaluable information for
taxonomists and fisheries managers alike. Knowledge of the morphology of the larvae
would make identification of potential pest species in plankton samples possible and may
provide a useful starting point for investigations into distribution and dispersal distances
of the larvae.
What information is held in museum collections about Dipolydora and Polydora in
the region?
The specimens examined in this study were from museum collections accumulated over a
period of just over 100 years. It is possible that the distribution range of some of the
species recorded in the present study no longer reflect the current situation. Loss, or
gain, of species in any area may be a result of anthropogenic influences such as habitat
alteration or climate change attributable to human induced global warming. They may
also reflect natural cycles or disease events. Museum collections also strongly reflect
“human” factors such as proximity to population centres, pleasant and easily accessible
collecting locations, interests of museum curators, funding body concerns or areas of
infrastructure development such as ports and marinas. This “human” factor must be
taken into consideration if using these collections to draw conclusions on species
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distributions or the biodiversity of particular regions. To be confident of species
distributions new or continued sampling programs in previously collected locations
followed by a program of long term standardized repeated monitoring in each of the
bioregions would be necessary. This type of sampling program would be especially useful
in accurately identifying the limits of the distribution ranges and their dynamic over time
for species in this study.
The subtropical region covered by the present study from Tweed-Moreton IMCRA
bioregion to Batemans Shelf IMCRA bioregion has 120 estuaries in New South Wales
(NLWRA, 2002) and 3 passageways with 6 rivers flowing into Moreton Bay in Queensland
(Thackway & Cresswell, 1998). Fifteen estuaries in this region are considered pristine
(Table 6). This study examined specimens from 19 non-pristine estuaries through this
subtropical region. No material from pristine estuaries was found in any museum
collection. Given the statement of Blake and Kudenov (1978) regarding high endemicity in
the Australian fauna, studies such as the present one based predominantly on museum
collections probably indicate only a small proportion of the existing biodiversity. An
extended sampling program of previously uncollected estuaries along the coast would
provide a more complete picture of the diversity of Polydora and Dipolydora species on
the subtropical eastern coast.
Museum collections are invaluable in providing a focal point for developing biodiversity
knowledge. Examination of these collections can provide indications of taxonomic
problems and collection gaps. One such collection gap identified in the course of this
research was the lack of polydorid specimens from oyster farms. The collections of Skeel
from the 1970s remain the only material from oyster farms from the east coast of
Australia. From this collection there is only one lot (NMV G3057) containing the widely
reported Polydora websteri, considered a major pest species of oysters. The only other
material identified as Polydora websteri in museum collections is the very important,
aged and incomplete specimen, AM W11390, thought to be collected by Haswell before
1900. Registered lots identified as Polydora cf. websteri in this study invariably were
multispecific and contained no specimens of Polydora websteri. Clearly, it is essential to
have multiple specimens in good condition of important pest species in museum
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collections. A focussed collecting effort is required to survey bivalve culture facilities in
estuaries of the east coast of Australia.
How many species or morphospecies of Dipolydora and Polydora occur in collections
from east coast subtropical estuaries and bays?
The resemblance analysis and similarity-percentage functions in Primer 6 proved to be a
useful tool in examining the polydorid fauna. Good species cluster groups were formed
containing known species. Anything unusual was immediately obvious as an isolated
individual in the dendrogram plot. Juvenile specimens, damaged individuals, mis-
identifications and data entry errors were all found in this way.
Prior to this study 3 Polydora species and 4 Dipolydora species were recorded from
subtropical eastern Australia. The present study found 12 Polydora species and 10
Dipolydora species for subtropical eastern Australia.
Two Polydora species are new (Polydora sp. P1 and Polydora cf. woodwicki) and 7 are
potentially new species (P. cf. latispinosa; P. cf. websteri; P. sp. P2S; P. sp. P3S; P. sp. P4S;
P. sp. P5S and P. sp. P6S) having been described from single specimens. Polydora cornuta
Bosc, 1902 is recorded from New South Wales for the first time. The identity of Polydora
cf. calcarea requires confirmation through allozyme analysis from the morphologically
similar P. ciliata (Johnston, 1838) previously recorded from Australia but synonymized
with Polydora websteri Hartman, 1943 in Loosenoff and Engle by Blake and Kudenov in
1978. Specimens fitting the description of Polydora websteri Hartman, 1943 in Loosenoff
and Engle were not seen in the present study. Polydora hoplura Claparède, 1870 and P.
haswelli Blake and Kudenov, 1978 are redescribed.
Three Dipolydora species are new (Dipolydora sp. D1; D. cf. flava and D. sp. D2) and 4 are
potentially new (D. cf. aciculata/ cf. giardi; D. sp. D3S; D. sp. D4S and D. sp. D5S).
Expanded descriptions are provided for Dipolydora tentaculata (Blake and Kudenov,
1978), Dipolydora flava (Claparède, 1870) and Dipolydora socialis (Schmarda, 1861).
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Which morphological characters are most useful in identifying Polydora and
Dipolydora morphospecies from this region? Are there any “new” characters that
are useful? Is it possible to identify animals which are incomplete?
The SIMPER function of Primer 6 was used to identify diagnostic characters for Polydora
and Dipolydora species in this study. This proved to be a very useful method allowing
many specimens to be examined together using a standardised and detailed set of
characters, thus avoiding the biases toward particular characters and knowledge
assumptions which cause confusion in existing keys.
Although the intention of the study was to examine only species level characters, generic
characters separating Polydora and Dipolydora were also assessed after some Polydora
haswelli Blake and Kudenov, 1978 material was noted to lack a constriction on the
hooded hooks, placing the material within the genus Dipolydora using existing diagnoses
(Blake, 1996d). These anomalous specimens were analysed as Dipolydora?. Resemblance
analysis resolved that the specimens when all characters were considered, clustered with
Polydora. For the material in this study diagnostic characters most important in
separating the two genera were (in order of importance): presence/absence of a
constriction on the hooded hooks; presence/absence of capillary notochaetae on
chaetiger 1; maximum number of hooded hooks in a single hook row; angle of the main
fang to the stem of the hooded hooks <90° and presence or absence of a subdistal blunt
tooth. These characters are in agreement with characters used currently to separate
Polydora and Dipolydora. Maximum number of hooded hooks in a hook row is an
additional character to the current diagnoses and in this study it was found that the
maximum number of hooded hooks is greater in Polydora (13 in Polydora sp. P1) than
Dipolydora (7 in D. tentaculata). This is possibly a character worthy of further
investigation including consideration of variation in number of hooded hooks due to
developmental stage. It is clear though that multiple characters should be considered
when separating Polydora and Dipolydora as they are currently defined. The constriction
on the hooded hooks, although the most important character, is not always obvious and
absence of capillary notochaetae on chaetiger 1, currently considered a Polydora
characteristic is seen in Dipolydora pilocollaris. Further investigation of polydorid genera
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using resemblance and SIMPER may help to clarify these anomalies. Evidence that this
may also be a useful technique for separation of genera is seen in the clear separation of
the Carazziella victoriensis / phillipensis species cluster group from other Polydora and
Dipolydora specimens in the analysis.
Characters important in separating species within Dipolydora and Polydora were (in no
particular order): chaetiger 5 spine and companion chaetae (Dipolydora only) characters;
shape of the peristomial margins; caruncle length; chaetae and lobes of the pre-chaetiger
5 chaetigers; presence of gizzard (Dipolydora only); dorsal ciliary organ/s location;
presence of an anterior furrow through the first 11 chaetigers; and for Polydora only,
chaetiger 6 + shape of the notopodial lobes.
For Polydora the characters occurring most often as important in separating species were
chaetiger 5 spine characters, peristomial margin characters and chaetiger 6+ notopodial
lobe shape. The peristomial ventral margins are rarely included in character descriptions
although occasionally illustrations are provided with descriptions. It appears from this
study that these anterior characters may be very useful for identification of Polydora. In
this study posterior spine characters were not included in the analysis, even for complete
specimens, as so few specimens were complete. The species cluster groups for specimens
which did have posterior spines, eg. Polydora hoplura still remained good and distinct
from other species. I think this indicates that it may not be essential to have posterior
characters to identify Polydora species. Further investigation and more concise definition
of the character states for the ventral peristomial margins should provide a very useful
character for identification. This investigation would best be done using live material to
exclude artefacts of storage and preservation from the character state definitions as may
have occurred in this study.
Characters of the parapodial lobes, particularly those posteriorly from chaetiger 6, were
shown to be important for separating Polydora species in this study. This is another
character which is generally overlooked and may be worthy of further investigation.
Characters important in separating Dipolydora species are chaetiger 5 spine characters,
caruncle length and peristomial margin shapes. Peristomial margin shapes are a character
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which has been overlooked for Dipolydora and is should be investigated further to better
define character states using fresh material.
The character states defined in this study, although based initially on the character states
used in the DELTA interactive key for polydorids (Wilson et al. 2003) were developed as
examination of the specimens in the study proceeded. It would now be useful to re-
examine each of the characters in isolation across the full set of species to further refine
the descriptions and states. Examination of fresh or live material would be beneficial for
soft tissue characters. It would also be very useful to establish clearly the variations in
chaetiger 5 spine characters in a single spine row both with regard to spine development
and emergence within the row and in the erosion patterns of older spines.
Developmental studies associated with SEM imaging could be undertaken to achieve this.
It should be recognized that the results of the SIMPER analysis are based on the factors
set preceding the analysis. In this type of analysis used for taxonomic purposes those
factors are “genus” and “species” so correct identification is important. Using the
resemblance analysis prior to the SIMPER analysis to cluster, and so name, specimens
accurately according to morphological characteristics is essential. The distance measure
within a factor group will give some indication of whether multiple species are mistakenly
included in that factor group. Including valid species, preferably type material, in the
analysis will provide a standard reference point for assessing acceptable distance within a
factor group. Further investigation of this method of analysis could attempt to discover if
there is a distance limit defining “species” for the polydorid group, or any other group
under investigation.
What are the distributions of the species or morphospecies of Dipolydora and
Polydora through the area in relation to the marine bioregions?
The results of the present study indicate that prior to this study a single Dipolydora
species, Dipolydora tentaculata (Blake and Kudenov, 1978) and no Polydora species had
been described from the Tweed-Moreton IMCRA bioregion. There have been no
descriptions or records of Polydora or Dipolydora from the northern New South Wales
region of the Tweed-Moreton IMCRA bioregion, although their presence has been noted
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in reports of mudworm disease in oysters in the Clarence River (Whitelegge, 1890). The
present study has identified 6 Dipolydora species including D. socialis (Schmarda, 1861)
and Dipolydora tentaculata (Blake and Kudenov, 1978) and 5 Polydora species from the
Tweed-Moreton region. There is still no material deposited in museum collections from
live mollusc shells or hermit crabs for the Tweed-Moreton region.
Prior to the present study the only Polydora species recorded from the Manning Shelf
IMCRA bioregion was Polydora haswelli Blake and Kudenov, 1978. These specimens were
collected from mudblisters in live oysters and were not available for examination for the
present study. Their identity must be questioned though as the type material for P.
haswelli Blake and Kudenov, 1978 was collected from sand. It is increasingly seen through
use of molecular techniques that species reported to occur in multiple habitats are
actually morphologically similar but genetically different species (Manchenko and
Radashevsky, 1993; 1998; 2002; Radashevsky and Pankova, 2002). In the present study
Polydora cf. calcarea was found abundantly in fresh material from the Camden-Haven
Estuary. This is discussed in detail in the taxonomic descriptions section. The lack of
Polydora species in the Manning Shelf region is almost certainly a result of lack of
collecting effort as suitable estuarine habitat including soft sandy mud substrates,
seagrass beds and oyster leases occur throughout estuaries in the bioregion. Four species
of Dipolydora were found including D. socialis (Schmarda, 1861) in collections from the
Manning shelf IMCRA bioregion.
Six Polydora species were found in collections from the Hawkesbury Shelf IMCRA
bioregion. These included P. haswelli Blake and Kudenov, 1978 and P. hoplura Claparède,
1870 plus four potential new species represented by a single specimen. This may reflect
greater collecting effort around a highly populated region. Five Dipolydora species
occurred in the Hawkesbury Shelf bioregion including D. socialis (Schmarda, 1861), D.
flava (Claparède, 1870) and D. tentaculata (Blake and Kudenov, 1978).
It appears that Dipolydora species are spread more widely along the east coast. D.
socialis, D. tentaculata and D. sp. D1 occurring across two or more bioregions. Dipolydora
flava is restricted to the Hawkesbury Shelf bioregion.
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Polydora species appear to be restricted to particular bioregions, but Polydora hoplura is
recorded from Tasmania (Blake and Kudenov, 1978). Specimens identified as Polydora
haswelli from Batemans Shelf bioregion were examined in this study but were considered
unsuitable for the analysis so have not been included here.
The distributions given on these maps are a reliable indication of species presence in a
bioregion, but absence from a bioregion is not so reliable. Absence may indicate a lack of
collecting effort in certain habitats (Table 1). This is particularly noted for Polydora cf.
woodwicki collected from an encrusting sponge in the Tweed-Moreton bioregion. Based
on existing records this habitat type appears to have not been collected from the
Manning, Hawkesbury or Batemans Shelf bioregions and so comparison between these
three regions and Tweed-Moreton may not be a true reflection of species distribution. It
should be considered though, that museum records are based on presence only. Habitats
and regions may have been sampled but no organisms found. Unfortunately, there seems
to be no easily accessible record kept of areas and habitats which have been sampled
which would allow a reliable species absence to be noted. Absence may also be due to
fauna in newly collected material being overlooked by inexperienced sorters.
Additionally, many areas are collected by private consultancy companies and university
students but this material, although of great value for public and scientific knowledge, is
not required to be deposited in museums. Until all habitats in all regions are collected, or
the problems noted above addressed, statements on species absence from a bioregion
should be treated cautiously.
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Table 1: Polydorid habitats sampled (+) and found to contain Polydora or Dipolydora
species and those not yet sampled (-) within IMCRA bioregions in estuaries of subtropical
eastern Australian based on museum records.
Habitat sampled
IMCRA bioregion soft substrates Sponges live mollusc/hermit
crab shells
Tweed-Moreton + + -
Manning Shelf + - +
Hawkesbury Shelf + - +
Batemans Shelf - - -
Where are the gaps in our knowledge of polydorid spionids in subtropical eastern
Australia and how can they be addressed?
The most important knowledge and collection gap found in this study and that which
should be addressed as quickly as possible is of the polydorid pest species occupying
mudblisters in oysters. It is incredible that there is no easily accessible reference
collection specifically for these pest species which have been reported over the last 120
years. Estuaries where they are noted to have occurred have not been sampled in any
methodical or standardised way to confirm that it is the same suite of species that is
occurring in all estuaries. There have been no collections made of oyster material from
northern New South Wales or Queensland. Studies of reproduction and larval
development in mudworms have not been thoroughly investigated and reported. Also
indicative of the neglect of this area is that CRIMP surveys specifically designed to collect
marine pest species did not collect from estuarine aquaculture facilities even though the
164
polydorid causing mudblisters in oysters appears to have been one of the first introduced
marine pest species reported from Australia (Haswell, 1885; Ogburn, 2007).
It is anecdotally discussed that movement of oyster stock along the coast has confounded
the natural distribution patterns of polydorids inhabiting bivalves in estuaries. There has
never been a study to investigate this statement and currently there is no evidence to
support it. A systematic collection from oyster leases at least in each of the bioregions in
each season for two years and associated studies into reproduction and larval
development would provide a good start to addressing this knowledge and collection
gap. Molecular work may be necessary to identify the species involved. A molecular study
comparing populations of mudworms from Australia, Europe and North America may be
able to ascertain the origin of these pests. This would provide accurate base knowledge
for oyster growers, taxonomists, aquaculture facility managers and marine pest
researchers and managers.
Museum collections of polydorids have been made chiefly from surveys of benthos in
estuaries along the subtropical coast (Table 1). Other habitat types have not been
consistently collected for each of the subtropical bioregions. For this reason distribution
data may not accurately reflect absence of a species from a region - it may simply reflect
lack of collection in a particular habitat. It would be useful to collect from each bioregion
for a particular habitat type. This may find species from the Polydora alloporis Group
(borers in mollusc shells or commensal with corals), Polydora bioccipitalis Group (borers
in gastropod shells especially hermit crabs), Polydora colonia/spongicola Group
(commensal with sponges and other polychaetes) and Dipolydora commensalis Group
(borers in mollusc shells) of Blake (1996d) that have not been recorded from Australia
(Chapter 1: Tables 2 and 3) providing information of use to researchers investigating
trends in global biogeography and biodiversity.
There is a lack of information and collection material from the Tweed-Moreton bioregion
estuaries. Based on the results of this study it appears that new species will occur in these
areas. Unfortunately, apart from the Gladstone QEB surveys, lots from this region contain
only one or two individuals. More specimens are required to provide an accurate
description and to confirm if these are indeed new species.
165
The gaps in our knowledge of Polydora and Dipolydora in Australia are largely a result of
lack of research in the area since the late 1970s. Fortunately, this lull has not occurred
globally and international research into polydorids has progressed with development of
molecular techniques, investigation of new characters of use in polydorid taxonomy and
attempts to refine knowledge on widely reported “cosmopolitan” species. In Australia, a
standardised collecting program focussed on particular habitats would be a first step
toward gaining a better understanding of this important polychaete group.
.....................................................
166
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complex (Polychaeta : Spionidae). Zoologischer Anzeiger 240(1):47-59.
Zann LP. 1995. Our sea, our future: Major findings of the state of the marine environment
report for Australia, for the Department of the Environment, Sport and Territories,
Ocean Rescue 2000 Program, Great Barrier Reef Marine Park Authority,
Townsville
180
Appendices
1Reproductive strategies of Polydora and Dipolydora species
2 Current worldwide species list of described Polydora and Dipolydora
3 Checklist of Australian polydorid Spionidae
4 PRIMER analysis character list and character numbers
5 Characters contributing most to distance between Polydora species
6 Characters contributing most to distance between Dipolydora species
7 Re-identification and reasoning for all specimens examined
APPENDIX 1: Published information on reproductive characters of Polydora and Dipolydora species from Australia and related species from
other locations (there is no information on reproduction in Australian Carazziella, Boccardiella).
Species Egg size (µm)
Type of development
Number of egg sacs; Number of eggs/sac; Total number of eggs
% egg development; Total larvae
Shape of nectochaete
Larval size; segments /µm
Development time: larvae in
sac (days)
Metamorphosis size (no. days planktonic)
Locality References
Polydora ciliata
135 Broadcaster 15-20 15-20
225-400
100 - 3 segments (256µm)
5 England 3.
Polydora ciliata
Ca. 120 Brooded without nurse eggs;
planktotrophic, released at the 3-
chaetiger stage
Strings of 15-20 capsules, each with
about 20 eggs
Narrow, tapering
posteriorly
18-19 chaetigers, 1340µm (6 weeks)
England; Denmark;
Sweden; North Sea
3., 10., 6., 7. In 5.
Polydora ciliata
? Brooded without nurse eggs;
planktotrophic
Narrow, tapering
15 chaetigers North Sea and off Belgium
11. in 5.
Polydora cf. ciliata
84-99 Brooded normally without nurse
eggs; planktotrophic, released at 3-
chaetiger stage
Strings of 14-25 capsules, each with about 10-170 eggs
Narrow, slightly
expanded in middel
17-18 chaetigers Ca. 1225µm
Sea of Japan 12. in 5.
Polydora cornuta
100-110 Broadcaster; planktotrophic
Capsules joined in a string or loosely held
together Up to 30 eggs/capsule
3 chaetiger (260µm)
17-18 chaetigers 1200-1300µm long
Brazil 13.
Polydora cornuta
? Brooded without nurse eggs,
planktotrophic
Individual capsules free from one another attached by 2 threads
Elongate, narrow, tapering
18 chaetigers (2600µm)
Sweden;North Sea
15. , 6. 7. In 5.
Polydora cornuta
120 Brooded, without nurse eggs;
planktotrophic; released at 3-4
chaetigers, polytelic
Capsules in loose bead-like string; some
singly attached with>132 eggs per
capsule
Thick, weakly expanded in
middle, tapering
17 chaetigers (1300µm) ca. 2
weeks
Maine, Damariscotta
Estuary
2.
Polydora cornuta
86-93.8 Brooded, without nurse eggs;
planktotrophic; released at 3-4
chaetigers, polytelic
Capsules in loose bead-like string; some
singly attached with30-60 eggs per capsule
? ? Florida 16. 17. In 5.
Polydora haswelli
111 ± 8 broadcaster 1 -
1200-2400
100 - 3 250± 20
9.3 ± 2.2 Australia: New South Wales
1.
Polydora cf. haswelli
~ 100 Planktotrophic; released at 3
chaetigers
Capsules lined in a string attached to
inner wall of burrow by two stalks
- - 3 chaetiger ~280 (based on P.ecuadoriana)
- 16-18 chaetigers Brazil 18.
Polydora hoplura
140 brooder 50 60
3000
“few” - 15 1670
- Europe 1.
Polydora hoplura
140 brooder 1 -
500-600
5 - 3-15 -
12.0 ± 2.7 Australia: Tasmania
3.
Polydora latispinosa
Australia -
Polydora websteri
101 ± 25
brooder 19.3 ± 7.2 83.0 ± 37.5 1500-2000
4 - 3-15 230-880
13.8 ± 4.0 Australia: New South Wales,
Tasmania
1.
Polydora websteri
140 broadcaster 10+ 50-55
500-550+
100 - 3 350-400
5 42 days at 15°C Eastern North America
2.
Polydora woodwicki
Australia -
Dipolydora aciculata
Australia -
Dipolydora armata
Australia -
Dipolydora armata
102.35 ± 0.88
Brooded with nurse eggs:
planktic larvae released at
various sizes
5-28 -
50-100
3- 14 3-7
Elongate, narrow
3-18 chaetigers (82% of sacs with
larvae ≤10 chaetigers )
200-300
20 chaetigers 3500+µm
Barbados, West Indies
4.
Dipolydora armata
ca. 90 Brooded with nurse eggs;
polytelic
Non compartmentalized
cylinder,
“few” 7 16-21chaetigers (1500-1800µm)
Brazil 14.
Dipolydora giardi
Australia -
Dipolydora giardi
80 Brooded without nurse eggs;
planktotrophic released at 3
chaetiger stage
1 elongated, hollow, transparent cylinder
Elongate hollow
cylinder with separate unpaired filaments
Approx. 8 17 chaetigers, 980µm, ca. 24
days, may delay metamorphosis
for 5 months
Central California 8.
1. Skeel, 1979 2. Blake, 1969 3. Wilson, 1928
4. Lewis 1998 5. Blake and Arnovsky, 1999 6. Hannerz, 1956
7. Plate and Husemann, 1994 8. Day and Blake, 1979 9. Carrasco, 1979
10. Thorson, 1946 11. Delcour and Maurice, 1982 12. Radashevsky, 1983
13. Radashevsky, 2005 14.Radashevsky and Noguiera, 2003 15. Söderström, 1920
16. Rice and Simon,1980 17. Rice, 1981 18. Radashevsky et al., 2006
Dipolydora flava
Australia -
Dipolydora flava
- Brooded; planktotrophic
- - Elongate, thickened
18 chaetigers 1500µm;
21 chaetigers 2100
Sweden; North Sea
6. and 7. in 5.
Dipolydora notialis
Australia -
Dipolydora pilocollaris
Australia -
Dipolydora protuberata
Australia -
Dipolydora socialis
Australia -
Dipolydora socialis
120 Brooded without nurse eggs;
planktotrophic released at 3
chaetiger stage; polytelic
Strings of capsules - -
Thick not fusiform
3 280µm (4 segments)
15 chaetigers, 1200-1400µm
Maine, Damariscotta
estuary
2.
Dipolydora socialis
? Brooded; planktotrophic
? Thick, weakly fusiform
26 chaetigers, 2250µm
Chile, Bahia de Concepcion
9. in 5.
Dipolydora tentaculata
Australia -
APPENDIX 2: Current valid species of polydorids in Australia and worldwide
Amphipolydora Blake, 1983
Valid species
Amphipolydora abranchiata (Hartman, 1953)
Amphipolydora vestalis Paterson and Gibson, 2003
Australia
No records to date
*Polydorella Augener, 1914
Valid species
Polydorella dawydoffi Radashevsky, 1996
Polydorella kamakamai Williams, 2004
*Polydorella prolifera Augener, 1914
Polydorella smurovi Tzetlin and Britayev, 1985
*Polydorella stolonifera (Blake and Kudenov, 1978)
Australia
*Polydorella prolifera Augener, 1914
*Polydorella stolonifera (Blake and Kudenov, 1978)
*Species listed under Pseudopolydora in Blake and Kudenov (1978)
Tripolydora Woodwick, 1964
Valid species
Tripolydora spinosa Woodwick, 1964
Australia
No records to date
Pseudopolydora Czerniavsky, 1881
Valid species (Blake, 1996; Blake and Kudenov, 1978; Read, 1975; Radashevsky and Hsieh, 2000)
P. achaeta Radashevsky and Hsieh, 2000
P. antennata Claparède, 1870
P. bassarginensis Zachs, 1933
P. corniculata Radashevsky and Hsieh, 2000
P. derjugini Zachs, 1933
P. diopatra Hsieh 1992
P. gibbsi Light, 1974
P. gigeriosa Radashevsky and Hsieh, 2000
P. glandulosa Blake and Kudenov, 1978
P. kempi (Southern, 1921)
P. cf. kempi japonica Imajima and Hartman, 1964
P. novaegeorgiae (Gibbs, 1971)
P. orientalis Annenkova, 1937
P. paucibranchiata (Okuda, 1937)
P. primigenia Blake, 1983
*P. prolifera (Augener, 1914)
P. pulchra (Carazzi, 1895)
P. reticulata Radashevsky and Hsieh, 2000
*P. stolonifera Blake and Kudenov, 1978
P. vexillosa Radashevsky and Hsieh, 2000
Australia
P. antennata Claparède, 1870
P. kempi (Southern, 1921)
P. glandulosa Blake and Kudenov, 1978
P. paucibranchiata (Okuda, 1937)
*P. stolonifera Blake and Kudenov, 1978
*P. prolifera (Augener, 1914)
*Also listed under Polydorella
Polydora Bosc, 1802
Valid species after Blake (1996)
P. aura Sato-Okoshi 1998
P. calcarea (Templeton, 1836) after Radashevsky and Pankova, 2006
P. carinhosa Radashevsky, Lana and Nalesso, 2006
P. fusca Radashevsky and Hsieh, 2000
P. manchenkoi Radashevsky and Pankova, 2006
P. mabinii Williams 2001 (P. ciliata/websteri group)
P. neocaeca Williams and Radashevsky, 1999 (P. ciliata/websteri group)
P. robi Williams, 2000
P. triglanda Radashevsky and Hsieh, 2000
P. umangivora Williams 2001 (P. ciliata/websteri group)
P. villosa Radashevsky and Hsieh, 2000
Blake (1996) after his revision of the genus Polydora defined 5 species groups (A-E) for the genus:
A. Polydora cornuta/nuchalis group
Valid species
P. cornuta Bosc, 1802
P. cirrosa Rioja, 1943
P. nuchalis Woodwick, 1953
Australia
P. cornuta Bosc, 1802
B. Polydora ciliata/websteri group
Valid species
P. aggregata Blake, 1969
P. brevipalpa Zachs, 1933 (Blake (1996) considers this sim. to P. websteri) _
P. ciliata (Johnston, 1838)
P. curiosa Radashevsky, 1994
P. ecuadoriana Blake, 1983
P. glycymerica Radashevsky, 1993
P. haswelli Blake and Kudenov, 1978
P. hermaphroditica Hannerz, 1956
P. hoplura Claparède, 1870
P. latispinosa Blake and Kudenov, 1978
P. limicola Annenkova, 1934
P. pacifica Takahashi, 1937
P. pygidialis Blake and Woodwick, 1972
P. rickettsi Woodwick, 1961
P. spondylana Mohammad, 1973
P. vulgaris Mohammad, 1972+
P. websteri Hartman, 1943
P. woodwicki Blake and Kudenov, 1978
Australia
P. ciliata (Johnston, 1838)
P. haswelli Blake and Kudenov, 1978
P. hoplura Claparède, 1870
P. latispinosa Blake and Kudenov, 1978
P. websteri Hartman, 1943
P. woodwicki Blake and Kudenov, 1978
C. Polydora alloporis group
Valid species
P. alloporis Light, 1970a
P. cavitensis Pillai, 1965
P. hornelli (Willey, 1905)
P. kaneohe Ward, 1981
P. wobberi Light, 1970b
Australia
No records to date
D. Polydora bioccipitalis group
Valid species
P. bioccipitalis Blake and Woodwick, 1972
P. maculata Day, 1963
P. gaikwadi Day 1973b
P. heterochaeta Rioja, 1939*
Australia
No records to date
*P. heterochaeta was described from a planktonic postlarvae and the adult has never been
described. The larvae is said to be similar to the larvae of P. ciliata, P. websteri and very similar to
larvae of P. hoplura (Blake, 1996)
E. Polydora colonia/spongicola group
Valid species
P. colonia Moore, 1907
P. narica Light, 1969*
P. spongicola Berkeley and Berkeley, 1950
Australia
No records to date
*There are no other confirmed records of P. narica.
Dipolydora Verrill, 1879
Valid species occurring after Blake (1996).
D. anoculata (Moore, 1907) after Radashevsky and Petersen, 2005
D. carunculata Radashevsky, 1993
D. huelma Sato-Okoshi and Takatsuka, 2001
D. melanopalpa Manchenko and Radashevsky, 2002
Blake (1996) groups Dipolydora into five groups (A-E).
A. Dipolydora giardi group
Valid species
D. giardi (Mesnil, 1896)
D. bifurcata (Blake, 1981)
D. magellanica (Blake, 1983)
D. tetrabranchia (Hartman, 1945)
D. tridenticulata (Woodwick, 1964)
D. trilobata (Radashevsky, 1993)
Australia
D. giardi (Mesnil, 1896)
B. Dipolydora concharum/coeca/flava/socialis group
Valid species
D. concharum (Verrill, 1879)
D. alborectalis (Radashevsky, 1993)
D. cardalia (Berkeley, 1927)
D. capensis (Day, 1955)
D. coeca (Oersted, 1843)
D. flava (Claparède, 1870)
D. goreensis (Augener, 1918)
D. normalis (Day, 1957)
D. peristomialis (Hartman, 1975)
D. pilikia (Ward, 1981)
D. protuberata (Blake and Kudenov, 1978)
D. saintjosephi (Eliason, 1920)
D. socialis (Schmarda, 1861)
D. tentaculata (Blake and Kudenov, 1978)
D. vulcanica (Radashevsky, 1994b)
Australia
D. flava (Claparède, 1870)
D. protuberata (Blake and Kudenov, 1978)
D. tentaculata (Blake and Kudenov, 1978)
D. socialis (Schmarda, 1861)
C. Dipolydora barbilla/bidentata group
Valid species
D. barbilla (Blake, 1981)
D. bidentata (Zachs, 1933)
D. langerhansi (Mesnil, 1896)
D. pilocollaris (Blake and Kudenov, 1978)
Australia
D. pilocollaris (Blake and Kudenov, 1978)
D. Dipolydora armata group
Valid species
D. aciculata (Blake and Kudenov, 1978)
D. akaina Blake, 1996
D. armata (Langerhans, 1880)
D. blakei (Maciolek, 1984)
D. caulleryi (Mesnil, 1886)
D. notialis (Blake and Kudenov, 1978)
D. quadrilobata (Jacobi, 1883)
Australia
D. aciculata (Blake and Kudenov, 1978)
D. armata (Langerhans, 1880)
D. notialis (Blake and Kudenov, 1978)*
*D. notialis is poorly known (Blake, 1996)
E. Dipolydora commensalis group
Valid species
D. commensalis (Andrews, 1891)
D. elegantissima (Blake and Woodwick, 1972)
D. hartmanae (Blake, 1971)
D. antonbruunae (Blake, 1893)
Australia
No records to date
Boccardiella Blake and Kudenov, 1978
Valid species (Blake and Kudenov, 1978)
B. bihamata Blake and Kudenov, 1978
B. hamata (Webster, 1879a)
B. ligerica (Ferronière, 1898)
B. limnicola (Blake and Woodwick, 1976)
B. magniovata (Read, 1975)
B. occipitalis Blake, 1983
B. truncata (Hartman, 1936)
Australia
B. bihamata Blake and Kudenov, 1978
B. limnicola (Blake and Woodwick, 1976)
Boccardia Carazzi, 1893
20 described species (Blake, 1996)
Valid species (Blake and Woodwick, 1971; Read 1975; Blake and Kudenov, 1981; Hutchings and
Turvey, 1984; Blake, 1983; Blake, 1986; Blake, 1996)
B. acus (Rainer, 1973)
B. androgyna Read, 1975
B. anophthalma (Rioja 1962)
B. basilaria Hartman, 1961
B. berkeleyorum Blake and Woodwick, 1971
B. chilensis Blake and Woodwick, 1971
B. columbiana Berkeley, 1927
B. fleckera Hutchings and Turvey, 1984
B. galapagense Blake, 1986
B. knoxi (Rainer, 1973)
B. lamellata Rainer, 1973
B. natrix (Söderström), 1920
B. otakouica Rainer, 1973
B. perata (Khlebovitsch, 1959)
B. polybranchia (Haswell, 1885)
B. proboscidea Hartman, 1940
B. pseudonatrix Day, 1961
B. pugettensis Blake 1979
B. semibranchiata Guerin, 1990
B. syrtis (Rainer, 1973)
B. tricuspa (Hartman, 1939)
Australia (Blake and Kudenov, 1978)
B. fleckera Hutchings and Turvey, 1984
B. polybranchia (Haswell, 1885)
B. proboscidea Hartman, 1940
B. chilensis Blake and Woodwick, 1971
Carazziella Blake and Kudenov, 1978
12 species known (Blake, 1996)
Valid species (Blake and Kudenov, 1978; Blake, 1979; Blake 1996)
C. calafia Blake, 1979
C. carrascoi Blake, 1979
C. citrona (Hartman, 1941)
C. hirsutiseta Blake and Kudenov, 1978
C. hobsonae Blake 1979
C. hymenobranchiata Blake and Kudenov, 1978
C. quadricirrata (Rainer, 1973)
C. patagonica Blake, 1979
C. phillipensis Blake and Kudenov, 1978
C. proberti Blake, 1984
C. reishi (Woodwick, 1964)
C. spongilla Sato-Okoshi, 1998
C. victoriensis Blake and Kudenov, 1978
Australia (Blake and Kudenov, 1978)
C. hirsutiseta Blake and Kudenov, 1978
C. hymenobranchiata Blake and Kudenov, 1978
C. phillipensis Blake and Kudenov, 1978
C. victoriensis Blake and Kudenov, 1978
APPENDIX 3: Current Australian checklist of the Polydora-complex (polydorids) within the family Spionidae, extract from Hutchings, PA and Johnson RT, (2003b) Australian Faunal Directory: Checklist for Spionidae, Generation Date: March 19, 2008
SPIONIDAE (in part – polydorid species only)
Boccardia Carazzi, 1893 Perialla Kinberg, 1866 Boccardia Carazzi, 1893 Paraboccardia Rainer, 1973 Boccardia chilensis Blake & Woodwick, 1971
Boccardia (Boccardia) chilensis Blake & Woodwick, 1971
Boccardia jubata Rainer, 1973 Boccardia fleckera Hutchings & Turvey, 1984
Boccardia fleckera Hutchings & Turvey, 1984
Boccardia polybranchia (Haswell, 1885) Polydora (Leucodore) polybranchia
Haswell, 1885 Polydora euryhalina Hartmann-Schröder,
1960 Boccardia proboscidea Hartman, 1940
Boccardia proboscidea Hartman, 1940 Boccardiella Blake & Kudenov, 1978
Boccardiella Blake & Kudenov, 1978 Boccardiella bihamata Blake & Kudenov, 1978
Boccardiella bihamata Blake & Kudenov, 1978
Boccardiella limnicola Blake & Woodwick, 1976 Boccardia limnicola Blake & Woodwick,
1976 Carazziella Blake & Kudenov, 1978
Carazziella Blake & Kudenov, 1978 Carazziella hirsutiseta Blake & Kudenov, 1978
Carzziella hirsutiseta Blake & Kudenov, 1978
Carazziella hymenobranchiata Blake & Kudenov, 1978
Carazziella hymenobranchiata Blake & Kudenov, 1978
Carazziella phillipensis Blake & Kudenov, 1978 Carazziella phillipensis Blake & Kudenov,
1978 Carazziella victoriensis Blake & Kudenov, 1978
Carazziella victoriensis Blake & Kudenov, 1978
Dipolydora Verrill, 1879 Dipolydora Verrill, 1881 Dipolydora aciculata (Blake & Kudenov, 1978)
Polydora aciculata Blake & Kudenov, 1978 Dipolydora armata (Langerhans, 1880)
Polydora armata Langerhans, 1880 Polydora monilaris Ehlers, 1904
Dipolydora ciliata (Johnston, 1838) Leucodore ciliatus Johnston, 1838
Dipolydora flava (Claparède, 1870) Polydora flava Claparède, 1870 Polydora pusilla Saint-Joseph, 1894 Polydora dorsomaculata Rainer, 1973
Dipolydora giardi (Mesnil, 1896) Polydora giardi Mesnil, 1896
Dipolydora notialis (Blake & Kudenov, 1978) Polydora notialis Blake & Kudenov, 1978
Dipolydora penicillata (Hutchings & Rainer, 1979) Polydora penicillata Hutchings & Rainer,
1979 Dipolydora pilocollaris (Blake & Kudenov, 1978)
Polydora pilocollaris Blake & Kudenov, 1978
Dipolydora protuberata (Blake & Kudenov, 1978) Polydora protuberata Blake & Kudenov,
1978 Dipolydora socialis (Schmarda, 1861)
Leucodore socialis Schmarda, 1861 Polydora plena Foster, 1971
Dipolydora tentaculata (Blake & Kudenov, 1978) Polydora tentaculata Blake & Kudenov,
1978 Polydora Bosc, 1802
Polydora Bosc, 1802 Diplotis Montagu, 1815 Leucodora Johnston, 1838 Leipoceras Möbius, 1874 Protopolydora Czerniavsky, 1881 Pseudoleucodore Czerniavsky, 1881 Polydora haswelli Blake & Kudenov, 1978
Polydora haswelli Blake & Kudenov, 1978 Polydora hoplura Claparède, 1870
Polydora hoplura Claparède, 1870 Polydora latispinosa Blake & Kudenov, 1978
Polydora latispinosa Blake & Kudenov, 1978
Polydora ligni Webster, 1879
Polydora ligni Webster, 1879 Polydora websteri Hartman, 1943
Polydora websteri Hartman, 1943 Polydora woodwicki Blake & Kudenov, 1978
Polydora woodwicki Blake & Kudenov, 1978
Pseudopolydora Czerniavsky, 1881 Pseudopolydora Czerniavsky, 1881 Carazzia Mesnil, 1896 Polydorella Augener, 1914 Neopygospio Berkeley & Berkeley, 1954 Pseudopolydora antennata (Claparède, 1870)
Polydora antennata Claparède, 1870 Pseudopolydora glandulosa Blake & Kudenov,
1978 Pseudopolydora glandulosa Blake &
Kudenov, 1978 Pseudopolydora kempi (Southern, 1921)
Polydora (Carazzia) kempi Southern, 1921 Neopygospio laminifera Berkeley &
Berkeley, 1954 Pseudopolydora paucibranchiata (Okuda, 1937)
Polydora (Carazzia) paucibranchiata Okuda, 1937
Pseudopolydora prolifera (Augener, 1914) Polydorella prolifera Augener, 1914
Pseudopolydora stolonifera Blake & Kudenov, 1978
Pseudopolydora stolonifera Blake & Kudenov, 1978
APPENDIX 4: Polydora and Dipolydora characters for PRIMER analysis
character number
data type character sub-character
9 0/1 anterior dorsal furrow first 11 chaetigers
14 ratio lateral length chaetiger 5: lateral length chaetiger6
15 0/1 chaetiger 5 dorsally fused with 6
16 0/1 chaetiger 5 overlapping 6 but not fused
20 ratio chaetiger 5 lateral length:dorsal width
21 0/1 dorsal ciliary organ 22 0/1
strip on either side of caruncle
23 0/1
strip extending posterior to caruncle
24 0/1
strip across ant edge of chaetiger 4, just posterior to caruncle
25 0/1
transverse strip at segment edge
27 0/1
medial glandular ridge
30 0/1 anterior prostomial margin Rounded
32 0/1
weakly bifid
33 0/1
truncate, rounded slightly raised corners
34 0/1
strongly bifid
35 0/1 anterior prostomial dorsal furrow
36 0/1
anterior prostomial transverse furrow
38 0/1 anterior peristomial inner dorsal margin dorsal edge bilobed
39 0/1
a single medial lobe
40 0/1
a single medial lobe, ventral to this a pair of lobes forming a medial small v where they meet
41 0/1
a single medial lobe, ventral to this a pair of laterally flattened lobes meeting at anterior 0.33x of length forming a medial v, continuing closely parallel
42 0/1
a single medial lobe, ventral to this a pair of lateral lobes forming a medial narrow v
43 0/1
no obvious lobe , a pair of very small lobes between the lateral lobes, half the height of the u-shaped division formed between the lateral lobes
44 0/1 anterior peristomial lateral margin large lateral lobes
45 0/1
lateral lobes not notably enlarged
46 0/1
large lateral lobes deeply divided medially
47 0/1
lateral lobes distally with conical
tip/supports
48 0/1
lateral lobes corner angular
49 0/1
lateral lobes reduced so that lateral peristomium appears to taper up to prostomium
51 0/1
lateral lobes rounded, level with prostomium, forming a blunt anterior end
52 0/1
lateral lobes rounded, anterior edge level with base of prostomial lobes
53 0/1
lateral lobes rounded, anterior edge clearly below more anterior lobe
54 0/1
lateral lobes edge 3 shaped
55 0/1 anterior peristomial ventral margin undivided
57 0/1
parallel sided slit
59 0/1
straight edge with a very tiny v medially
60 0/1
medial v, width slightly less than the base of the prostomium, depth 0.5 width
61 0/1
narrow v becoming parallel sided slit to level with chaetiger 1
63 0/1
broad v becoming a parallel sided slit
64 0/1
broad v, opening 1/3rd anterior edge width
65 0/1
broad v becoming a narrow v shaped slit medially
66 0/1
broad shallow v 67 0/1
deep v
68 0/1
small medial u, equal to width of dorsal lobe, similar depth
69 0/1
shallow open U
70 0/1
broad u, base 0.5x segment width, almost as deep as it is wide
71 0/1
broad u, base 0.5x segment width with 5 small lobes on edge, almost as deep as it is wide
73 0/1
deep rounded U
74 0/1
deep 0rrow U, appearing shallowly lobed (7or 9) or segmented on lower edge, width equal to anterior lobe width and 0.5x depth
76 0/1/
v becoming a u; anterior edge width equal to anterior width of prostomium; width of u 0.33xanterior edge width; depth of u 2x width of u
77 0/1
semicircular scallop, base level with anterior chaetiger 1
80 0/1
medial division extends posteriorly to chaetiger 1
86 0/1 pigmentation parapodia 87 0/1
prostomium
88 0/1
peristomium
89 0/1
dorsal thoracic
90 0/1
ventral thoracic
91 0/1 eyes present/absent
92 count
number
93 0/1
anterior pair widest apart
95 0/1 caruncle (prostomial dorsal extension) ant 2
97 0/1
post 2 98 0/1
ant 3
99 0/1
mid 3 100 0/1
post 3
101 0/1
ant 4
102 0/1
mid 4
103 0/1
post 4
104 0/1
mid 5
105 0/1 occipital antennae present/absent
106 0/1 gizzard chaetiger 13-16
107 0/1
15-19
108 0/1
17-22
109 0/1
25-29
110 count dorsal branchiae commence 112 count
longest at chaetiger
113 0/1
overlapping dorsally 114 0/1
overlapping dorsally anteriorly
115 0/1 parapodial lamellae/lobes chaetiger 1 notopodia present
116 0/1
noto1-digitiform
117 0/1
noto1-conical
118 0/1
noto1-globular
119 0/1
chaetiger 1 neuro1-digitiform
120 0/1
neuro1-conical
121 0/1
neuro1-globular
122 0/1
neuro1-broad rounded lamellae
123 0/1
chaetiger 2-4 noto- digitiform
124 0/1
noto-conical
125 0/1
noto-rounded
126 ratio
noto- length lobe:longest chaetae
127 0/1
noto-longer on 2,3 becoming broad on 4
128 0/1
chaetiger 2-4 neuro-digitiform
129 0/1
neuro- conical
130 0/1
neuro- globular
131 0/1
neuro-broad rounded
132 ratio
neuro-length lobe:longest chaetae
133 0/1
neuro longer on 2,3 becoming broad on 4
134 0/1
chaetiger 5 noto
135 0/1
neuro
136 0/1
chaetiger 6 +
noto-small digitiform inferior lobe between the setae and the branchiae
137 0/1
noto-digitiform
138 0/1
noto- conical(triangular) posterior lobe
139 0/1
noto-broad triangular
140 0/1
noto-rounded
141 0/1
noto-spreading fan
142 0/1
noto length lobe:longest chaetae
143 0/1
noto -lobes change shape and become reduced at chaetiger
144 0/1
chaetiger 6 + neuro-digitiform
145 0/1
neuro-globule
146 0/1
neuro-conical (triangular)
147 0/1
neuro present after chaetiger 18
152 0/1 chaetae pre-chaetiger 5 chaetiger 1 notochaetae1 with capillaries
153 0/1
chaetiger 1 neurochaetae 1with capillaries
154 0/1
neuro 1-capills spreading fascicle
155 0/1
neuro 1-capills 5 broad
156 0/1
neuro 1-capills 2
157 0/1
chaetiger 2-4 noto 2-4 number of tiers of capills
158 0/1
noto 2-4-fanned
159 0/1
noto2-4- some geniculate capills
160 0/1
noto 4-some capills with fibrous edge
161 0/1
noto 2-4-glandular lobes with needle spines
162 0/1
chaetiger 2-4
neuro 2-4 becoming short and spine-like on chaetiger 4
163 0/1
neuro 2-4 two tiers of long and short capills, 0.5 length of longer notochaetae
164 0/1
neuro 2-4 with geniculate capills
165 0/1
short caps, about 1/3 notopodial chaetae length
168 count chaetae chaetiger 5
5 dorsal-number of chaetae
169 0/1
5 dorsal-broad capills
170 0/1
5 dorsal-geniculate
171 0/1
5 dorsal-adjoining spine row
173 0/1
5 ventral-number
174 0/1
5 ventral- tiers
175 0/1
5 ventral- broad capills
176 0/1
5 ventral- fine capills
177 0/1
spine row shape
shallow curve, endpoints anterior dorsal, posterior ventral
178 0/1
very shallow curve
179 0/1
straight
180 0/1
straight dorsally
181 0/1
shallow curve ventral
182 0/1
orientation longitudinal
183 0/1
transverse
184 0/1
perpendicular
185 count
number of rows of spines
186 count
number of spine types
187 count
number of major spines dorsal
188 count
ventral
189 0/1
form of dorsal spine simple
190 0/1
simple curved spines
191 0/1
bluntly falcate
192 0/1
falcate
193 0/1
pipette-tip (flattened)
194 0/1
slight subdistal swelling
195 0/1
subdistally inflated
196 0/1
wrinkled on convex surface
197 0/1
type of accessory structure - dorsal row
low subdistal flange on inner edge
198 0/1
lateral flange appearing toothlike in some orientations
199 0/1
lateral cheeklike swelling with subdistal flange inside, appearing like a tooth in some angles
200 0/1
continuous flange, upper lateral sides of spine scooping to subdistal inner edge
201 0/1
continuous flange joined to spine at neck of fang and along convex spine edge almost to tip. The flange projecting perpendicular to the spine axis a distance almost equal to the width of the spine. It appears as if
a large inner tooth meets the flange at the outer edge.
202 0/1
sub-distal blunt tooth
203 0/1
small lateral tooth at curve of stem
204 0/1
subdistal tooth adjoining on one side only a variously eroded lateral flange which extends toward spine tip
205 0/1
subdistal flange hooding the spine crest and with lateral winglike arms extending forward from spine stem
206 0/1
large subdistal flange equal to the width of the spine tip, attached to the spine stem laterally, doesn't appear to be attached to the back of the spine stem
207 0/1
accessory tooth on crest
208 0/1
a scale extending just over convex (outer) may terminate in a rough broad edge
209 0/1
band of bristles around collar (subterminal)
210 0/1
bristles over crest, tip bristle free
211 0/1
bristle covering from throat to tip
212 0/1
spine form - ventral row distally swollen
213 0/1
bristle-topped upper half of swelling
214 0/1
lip around dorsal side of swelling
215 0/1
concavity on anterior side
216 0/1
companion chaetae present/absent
217 0/1
companion chaetae type/s limbate
218 0/1
geniculate
219 0/1
sigmoid
220 0/1
brush-tipped
221 0/1
truncated
222 0/1
pennoned
223 0/1
pennoned strongly geniculate, tapered
224 0/1
very thin
225 0/1
very thin fine spines
226 0/1
tapered
227 0/1
aristate
228 0/1
feathery and bifurcate
229 0/1
distal fibrous edge
230 count chaetae chaetiger 6+ chaetae change length or form at
231 0/1
ventral fascicle with short broad limbate or aristate up to change point
232 0/1
short spinelike chaetae in some notopodia
233 0/1
some embedded spine bundles after chaetiger 18
234 0/1
fibrous edge
236 0/1
neuropodial capillaries 237 count
number neuropodial capillaries chaet 6
238 count
chaet 7
239 count
chaet 10
240 count
chaet 16
243 count hooded hooks from neuro
244 0/1
hooded hooks with constriction on stem
245 count
minimum number of hh 246 count
max number of hooks
247 0/1
dentition bi
248 0/1
dentition uni
249 0/1
hh-angle main fang to stem >90
250 0/1
90
251 0/1
hh-angle main fang to stem <90
252 0/1
angle variable in same row
253 0/1 notopodial spines notopodial packets of fine needle spines
APPENDIX 5: Characters making the greatest percentage contribution, and the size of the greatest percentage contribution, to the resemblance between Polydora species. A: 1. cornuta 2. cf. websteri* 3. P6S* 4. P4S* 5. cf. woodwicki 6. haswelli 7. hoplura 8. P1 B: 9. woodwicki* 10. P5S* 11. cf. haswelli 12. P3S* 13. P2S* 14. cf. latispinosa* 15. latispinosa* 16. cf. calcarea. Characters making equal % contribution in brackets. *single specimen species
A:
1. 2. 3. 4. 5. 6. 7. 8.
1.
2. 67 (23.74%) 24, 172
3. 137 (18.35%) 162, 199
(67, 137) (18.69%) 24
4. (54, 203) (15.64%) 140
(54,67,203) (15.58%)
(54, 137, 203) (15.58%)
5. 162 (8.57%) 232, 236
67 (24.77%) 24, 95
137 (23.04%) 228, 195
(54,203) (19.46%) 140
6. 162 (10.12%) (236, 23)
67 (28.31%) 24, 95
137 (23.81%) 228, 199
(54, 203) (18.59%) 140
(9, 140) (5.44%) 199
7. 206
(11.64%) 162 74
67
(22.72%) 206 24
137
(20.57%) 206 74
(54, 203)
(17.54%) 140
206
(14.16%) 74 195
206
(14.97%) 74 195
8. 162 (11.29%)
232 236
67 (30.35%)
24 95
137 (23.58%)
199 228
(54,203) (17.77%)
140
(9, 140) (5.45%)
(59, 199)
51 (3.36%)
59 61
206 (15.43%)
74 195
9. 162 (10.90%) 232
236
67 (28.99%) 24
95
137 (26.52%) 199
228
(54,203) (19.08%) 140
(9, 140) (6.5%) 130
198 (3.92%) 51
61
206 (15.18%) 74
195
198 (4.06%) 130
59
10. 201 (19.72%) 24
162
(201,67) (20.84%) 95
(201,137) (17.34%) 24
(201, 54, 203) (14.33%)
201 (22.96%) 24
9
201 (23.475) 24
63
201 (19.09%) 206
24
201 (24.57%) 24
33
11. 162 (9.57%) 232
236
67 (28.28%) 24
95
137 (21.37%) 199
228
(54,203) (17.92%) 140
(9,140, 60) (5.17%)
60 (8.07%) 51
61
206 (14.735) 74
195
60 (6.08%) 59
48
12. 157 (15.19%) 162 108
67 (28.02%) 24 157
137 (21.72%) 157 199
(54, 203) (17.85%) 140
157 (9.275) 108 9
157 (10.28%) 108 51
157 (16.70%) 206 74
157 (10.43%) 108 117
13. 196 (23.20%) 162 232
(196,67) (22.37%) 24
(196,137) (18.715) (199, 228)
(196,54, 203) (15.79%)
196 (25.58%) (9, 140)
196 (34.21%) 66 233
196 (23.10%) 206 74
196 (26.93%0 66 233
14. 27, 46, 156) (16.21%)
(27,46, 156, 67) (14.97%)
(27,46, 156, 137) (13.065)
(27,46, 156, 54) (11.38%)
(27,46, 156) (16.26%)
(27,46, 156) (17.35%)
(27,46, 156) (15.70%)
(27,46, 156) (17.62%)
B.
15 (43,
162) (10.16%) 232
67
(26.26%) 43 24
137
(22.6%) 43 199
(54,
203) (17.64$) 140
43
(11.14%) 9 140
43
(11.935) 198 15
206
(15.03%) 43 74
43
(11.99%) 198 15
16. 162
232 (8.45%) 236
67
(24.57%) 24 95
137
(22.62%) 199 228
(54,
203) (17.21%) 140
(9,
140) (4.9%) 123
(123,
139) (5.18%) 77
206
(12.99%) 74 195
(123,
139) (4.97%) 77
9. 10. 11. 12. 13 14 15. 16.
10. 201 (23.34%) 24 63
11. 60 (7.02%) 198 130
201 (21.48%) 24 63
12. 157 (11.78%) 108 198
201 (21.22%) 24 157
157 (15.30%) 108 60
13. 196
(32.50%) 66 198
(196,201)
(19.12%) 24
196
(41.85%) 60 112
196
(33.97%) 157 108
14. (27,46, 156)
(19.82%)
(27,46, 156, 201)
(12.66%)
(27,46, 156)
(17.40%)
(27,46, 156)
(16.53%)
(27,46, 156, 196)
(15.13%)
15. 43 (19.29 %) 230
15
201 (20.35%) 43 24
43 (13.71%) 60 230
43 (10.81%) 157 108
196 (27.22%) 43 66
(27, 46, 156) (19.53%)
16. (123, 139) (6.26%)
77
201 (19.89%) 24
63
60 (5.21%) (123,
139)
157 (9.08%) 108
123
196 (24.78%) 123
139
(27, 46, 156)
(16.59%)
43 (11.52%) 123
139
43(11.52) 123, 139,
77
APPENDIX 6: Characters making the greatest percentage contribution, and the size of the greatest percentage contribution, to the resemblance between Dipolydora species. A: 1. armata* 2. aciculata* (type) 3. cf. aciculata/ cf. giardi* 4. cf. flava 5. flava 6. giardi* 7. pilocollaris* paratype 8. protuberata* paratype B: 9. socialis 10. D5S* 11. D2 12. tentaculata 13. D1 14. D3S* 15. D4S* 16. cf. pilocollaris
Characters making equal % contribution in brackets. * species represented by a single specimen
A:
1. 2. 3. 4. 5. 6. 7. 8.
1.
2. (134, 155, 205) (16.53%)
3. (134,
155, 205, 207) (13.15%)
207
(17.43%) 9, 23 68
4. (134,
155, 205) (17.18%)
71
(8.7%) 68 210 232
207
(16.58%) 9 23 71
5. (134,
155, 205) (17.64%)
(68,
210) (5.89%) 232 131
207
(19.12%) (9, 23) 125
71
(10.04%) 131 161 135
6. (134,
155, 205) (16.12%)
43
(9.08%) 199 (68, 195, 210)
207
20.35%) 9 23 42
(42,
71) (8.68%) 199 195
42
(8.94%) 199 195 125
7. (134, 155, 205) (19.26%)
208 (8.56%) 68 210 232
207 (19.07%) 9 23 208
71 (10.87%) 208 146 24
208 (9.43%) 131 161 135
42 (9.52%) 199 208 195
8. (134, 155, 205) (17.37%)
(68, 108, 95, 210) (5.67%)
207 (9.86%) 9 (23, 108)
71 (10.92%) 95 108 146
(108, 95) (6.93%) 131 161
42 (9.0%) 199 (108, 95, 195)
208 (12.69%) 108 95 144
9. (134, 155, 205) (18.45%) 144
(68, 210) (6.35%) 232 (224, 51)
207 (18.55%) 9 23 125
71 (13.80%) 24 146 40
131 (5.28%) 161 135 106
42 (9.475) 199 195 125
208 (11.20%) 38 141 106
(108, 95) (7.54%) (144, 38)
10. (225 134, 155, 205) (14.39%)
225 (22.06%) 228 68 210
(225, 207) (14.29%) (9, 23)
225 (25.5%) 71 228 101
225 (23.97%) 228 (131, 101)
225 (20.09%) 42 199 228
225 (27.59%) 228 208 101
225 (25.97%) 228 (108 95)
11. (134, 155,
(69, 102)
207 (15.81%)
(69, 102)
(69, 102)
(69, 102)
(69, 102)
(69, 102)
B.
205)
(16.04%) 69
(12.21%)
68 210
9
23 69
(16.13%)
71 109
(12.55%)
109 131
(11.28%)
42 199
(14.39%)
208 109
(13.65%)
108 95
12. (134, 155, 205)
(13.99%)
104 (11.37%) (43,
42) 68
207 (15.64%) 9
23 104
104 (11.18%) (43
42, 71)
104 (11.46%) (43,
42) 109
104 (11.22%) 43
199 195
104 (13.20%) (43,
42) 208
104 (13.06%) (43,
42) 95
13. (134, 155, 205)
(15.75%) 187
(68, 210) (5.43%)
232 224
207 (17.40%) 9
23 125
71 (9.53%) 146
24 40
131 (4.38%) 161
135 223
42 (7.94%) 199
195 125
208 (8.50%) 223
38 160
(108, 95) (6.60%)
223 38
14. (134, 155, 205)
(17.57%)
(68, 95, 210)
(5.81%) 232
207 (18.71%) 9
23 95
71 (10.39%) 95
24 40
95 (6.06%) 131
161 135
42 (9.44%) 199
95 195
208 (9.66%) 95
146 38
108 (8.66%) 146
(144, 38)
15 (134, 155, 205)
(18.82%) 136
236 (13.78%) 68
195 210
207 (18.81%) (236,
9, 23)
236 (13.10%) 71
195 132
236 (15.15%) 195
131 161
236 (14.60%) 42
199 125
236 (18.15%) 208
195 132
236 (15.60%) 108
95 195
16. (134,
155, 205) (16.12%) 49
49
(9.51%) 208 210 232
207
(17.30%) 9 23 49
49
(8.43%) 71 208 68
49
(8.72%) 208 68 131
(49,
42) (7.63%) (199, 208)
49
(12.27%) 68 66 122
49
(8.90%) 208 68 108
9. 10. 11. 12. 13 14 15. 16.
10. 225 (30.51%) 228
76 101
11. (69, 102)
(18.04%) 109 40
225 (20.99%)
69 102 228
12. 104
(12.99%) (43, 42) 109
225
(22.23%) 104 43 42
104
(10.13%) (69, 102) 43
13. 223 (3.8%) 160 65 193
225 (24.31%) 228 101 146
(69, 102) (12.84%) 109 40
104 (11.02%) (43, 42) 109
14. 95 (6.81%) 146 193 233
225 (26.66%) 228 95 101
(69, 102) (12.85%) 95 109
104 (11.44%) (43, 42) 95
95 (7.09%) 146 223 160
15. 236 (16.46%) 195 233 57
225 (23.67%) 236 228 195
236 (12.265) (69, 102) 195
(236, 104) (11.06%) (43, 42)
236 (13.38%) 195 223 160
236 20.20%) 95 195 146
16. 49
(10.23%) 208 68 66
225
(20.56%) 49 228 208
(69,
102) (11.30%) 49 208
104
(9.68%) (43, 49, 42)
49
(8.38%) 208 68 66
49
(9.02%) 208 (68, 95)
236
(12.52%) 49 208 68
Appendix 7: Re-identification and reasoning for all specimens examined *these identifications based on existing descriptions as PRIMER analysis was restricted to Polydora and Dipolydora species only ** not included in PRIMER analysis
Registration number Original identification
Proposed identification prior to analysis/reasoning
Identification after final analysis
AM W 13042-8 Polydora haswelli
Dipolydora? Polydora P2S
AM W 13042-9 Polydora haswelli
Dipolydora? Polydora haswelli
AM W11714-1 Polydora flava Accidental omission Dipolydora flava (label)
**Consistent with Dipolydora flava
AM W11714-2 Polydora flava Accidental omission Dipolydora flava (label)
**Consistent with Dipolydora flava
AM W13042 -2 Polydora haswelli
Dipolydora sp. Dipolydora D4S
AM W13042-1 Polydora haswelli Paratype
Dipolydora socialis Polydora sp. P3S
AM W13042-10 Polydora haswelli
Dipolydora? Polydora haswelli
AM W13042-11 Polydora haswelli Paratype
Dipolydora? (giardi) Polydora haswelli
AM W13042-12 Polydora haswelli (attached juvenile with AM W13042-11)
? ** juvenile specimen removed from analysis
**Not determined
AM W13042-3 Polydora haswelli Paratype
Polydora haswelli
AM W13042-4 Polydora haswelli Paratype
Polydora haswelli
AM W13042-5 Polydora haswelli Paratype
Polydora haswelli
AM W13042-6 Polydora haswelli Paratype
Polydora haswelli
AM W13042-7 Polydora haswelli Paratype
Polydora haswelli
AM W13044-1 Boccardiella bihamata
* *Boccardiella bihamata
AM W13044-2 Boccardiella bihamata
* *Boccardiella bihamata
AM W13044-3 Boccardiella bihamata
* *Boccardiella bihamata
AM W13044-4 Boccardiella bihamata
* *Boccardiella bihamata
AM W13044-5 Boccardiella bihamata
* *Boccardiella bihamata
AM W13044-6 Boccardiella bihamata
* *Boccardiella bihamata
AM W15717 Polydora protuberata
Dipolydora protuberata ** some important character information missing
**Dipolydora protuberata
AM W16367 Carazziella hirsutochaeta
Carazziella hirsutochaeta
AM W16368 Carazziella hirsutochaeta
Carazziella hirsutochaeta
AM W16919-1 Polydora cf. websteri
Polydora cf. cornuta but check re short ventral spines on chaetiger 5(ventral fascicle of unilimbate capillaries in B&K 1978) 27/07Rad2005 ventral capills occasionally present in 18-22 chaetiger recently settled juveniles)
Polydora cornuta
AM W16919-2 Polydora cf. websteri
Polydora cf. cornuta but check re short ventral spines on chaetiger 5(ventral fascicle of unilimbate capillaries in B&K 1978) 27/07Rad2005 ventral capills occasionally present in 18-22 chaetiger recently settled juveniles)
Polydora cornuta
AM W16919-3 Polydora cf. websteri
Polydora cf. cornuta but check re short ventral spines on chaetiger 5(ventral fascicle of unilimbate capillaries in B&K 1978) 27/07Rad2005 ventral capills occasionally present in 18-22 chaetiger recently settled juveniles)
Polydora cornuta
AM W17066 Carazziella phillipensis
Carazziella phillipensis
AM W17067 Carazziella victoriensis
Carazziella victoriensis
AM W17068 Polydora Data entry error: originally Dipolydora
pilocollaris Paratype
entered Dipolydora protuberata 7 April 08 corrected ID to pilocollaris
pilocollaris
AM W17069 Polydora protuberata Paratype
Dipolydora protuberata Dipolydora protuberata
AM W19112 extra This was probably AM W170067.
Carazziella victoriensis
AM W19112-1 Polydora penicillata
As above Carazziella victoriensis
AM W19112-2 Polydora penicillata
As above Carazziella victoriensis
AM W19112-3 Polydora penicillata
As above Carazziella victoriensis
AM W19112-4 Polydora penicillata
As above Carazziella victoriensis
AM W19112-5 Polydora penicillata
As above Carazziella victoriensis
AM W19112-6 Polydora penicillata
As above Carazziella victoriensis
AM W19112-7 Polydora penicillata
As above Carazziella victoriensis
AM W19112-8 Polydora penicillata
As above Carazziella victoriensis
AM W19112-9 Polydora penicillata
As above Carazziella victoriensis
AM W194020 Boccardia fleckera
*Boccardia fleckera
AM W199279-1 Polydora cf. websteri (Queensland Electricity Board code number 97)
photos 54-57 Polydora not websteri cf. cornuta (occipital antennae is tiny and on top of hump of caruncle above eyes ; number of hh based on Blake96, agrees with Rad 99; cf. cornuta but spine doesn't match B&K78)
Polydora cf. websteri
AM W199279-10 Polydora cf. websteri (Queensland Electricity Board code number 97)
Dipolydora cf. socialis Dipolydora socialis
AM W199279-11 Polydora cf.websteri (Queensland Electricity Board code number 97)
Dipolydora cf. socialis Dipolydora socialis
AM W199279-12 Polydora cf.websteri
Polydora “sparklebum” cf. (spines and posterior cf.
Polydora sp. P1
(Queensland Electricity Board code number 97)
woodwickii; occ. ant)
AM W199279-13 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-14 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-15 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora Polydora sp. P1
AM W199279-16 Polydora cf.websteri (Queensland Electricity Board code number 97)
Dipolydora cf. socialis Dipolydora socialis
AM W199279-17 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf. paucibranchiata
AM W199279-18 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf. paucibranchiata
AM W199279-19 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora *Pseudopolydora sp.
AM W199279-2 Polydora cf.websteri (Queensland Electricity Board code
photo52-51 (Polydora “sparklebum”)
Polydora sp. P1
number 97)
AM W199279-20 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-21 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ.ant)
Polydora sp. P1
AM W199279-22 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-23 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf. paucibranchiata
AM W199279-24 Polydora cf.websteri (Queensland Electricity Board code number 97)
Dipolydora cf. socialis Dipolydora socialis
AM W199279-25 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata (these are probably the real paucibranchiata based on Okuda's description AM)
*Pseudopolydora cf. paucibranchiata
AM W199279-26 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf. paucibranchiata
AM W199279-27 Polydora cf.websteri (Queensland Electricity Board code number 97)
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf. paucibranchiata
AM W199279-28 Polydora cf.websteri
*Pseudopolydora cf. paucibranchiata
*Pseudopolydora cf.
(Queensland Electricity Board code number 97)
paucibranchiata
AM W199279-3 Polydora cf. websteri (Queensland Electricity Board code number 97)
Dipolydora socialis Dipolydora socialis
AM W199279-4 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora cornuta (NB8th July "sparklebum" not cornuta no occ ant)
Polydora sp. P1
AM W199279-5 Polydora cf.websteri (Queensland Electricity Board code number 97)
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W199279-6 Polydora cf.websteri (Queensland Electricity Board code number 97)
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W199279-7 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-8 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W199279-9 Polydora cf.websteri (Queensland Electricity Board code number 97)
Polydora “sparklebum” cf. (spines and posterior cf. woodwickii; occ. ant)
Polydora sp. P1
AM W23666 Polydora haswelli
AM W23667 Polydora haswelli
Dipolydora sp. , looks different to everything else,
Dipolydora D3S
not able to ID from B& K, similar to D. aciculata (holotype at Vic Mus, NMV G2872, Paratypes NMV G2873)
AM W24817 Polydora cf. websteri
*Pseudopolydora *Pseudopolydora sp.
AM W24940-1 Polydora haswelli
Polydora haswelli
AM W24940-2 Polydora haswelli
Polydora haswelli
AM W26115 Polydora cf. giardi
cf. aciculata (branchiae present on anterior 1/3, 4-lobed pygidium, caruncle to post 3, posterior neuropodial spines, notopodial acicular spines), cf. giardi (spines, incised prostomium, capillary neurosetae 1-2 accompanying hh;
Dipolydora cf.aciculata / cf. giardi
AM W26119-1 Polydora pilocollaris
Dipolydora pillocollaris Dipolydora cf. pillocollaris
AM W26119-2 Polydora pilocollaris
Dipolydora pillocollaris Dipolydora cf. pillocollaris
AM W26119-3 Polydora pilocollaris
Dipolydora pillocollaris Dipolydora cf. pillocollaris
AM W26121 Polydora hoplura
Accidental omission Polydora hoplura
AM W26122 Polydora woodwicki
not woodwicki (too many branchiae?)
Polydora sp. P4S
AM W26124 Polydora flava Jar contains W26123 . Jar registered as Polydora flava but W26123 reidentified as Prionospio cf. wambiri
**See cell to left
AM W26151 Dipolydora cf. armata
I suspect that this is Polydora cf. woodwicki, spec is small and incomplete so can't be sure, also hh appear in anterior to be without constriction and posteriorly with constriction? NB 6.7.07 have looked at woodwicki types and this is not it.
Polydora cf. woodwicki
AM W26152 Dipolydora cf. armata
I suspect that this is Polydora cf. woodwicki, spec is small and incomplete so can't be sure, also hh appear in anterior to be without constriction and posteriorly
Polydora cf. woodwicki
with constriction?
AM W27868 Polydora cf. woodwicki
cf. P. cornuta (Rad redescription, same spines, maybe occ. Ant. detached?)nb.10/9/07 agrees with P. woodwicki description but no posterior & not on Abalone
Polydora cf. woodwicki
AM W29649 Dipolydora armata
Polydora cf. woodwicki (diffs: branchiae to end, spine flange location) maybe hoplura (spines same, but no occ ant or post spines…RECHECK)
Polydora sp. P5S
AM W29651 - 1 Polydora flava ? Incomplete, appears very similar to anterior end of AM W29651-2
Dipolydora cf. flava
AM W29651 - 2 Polydora flava needle spines are emergent, occipital antennae present (absent in P. flava?)
Dipolydora cf. flava
AM W29944 Polydora giardi Polydora cornuta? Polydora sp. P6S
AM W29948 ? Polydora haswelli
AM W29949 Polydora haswelli Polydora haswelli
AM W29950 Polydora flava Dipolydora cf. socialis Dipolydora sp. D2
AM W31083 Dipolydora giardi
Dipolydora giardi
AM W31461 Spionidae Not included in analysis Polydora hoplura
**
AM W31463 Spionidae Not included in analysis Dipolydora armata
**
AM W31464-1 Polydora latispinosa; should be cf. latispinosa(18th april08)
Polydora cf. latispinosa
AM W31920 Polydora flava Dipolydora cf. socialis Dipolydora socialis
AM W31921 Polydora flava Dipolydora cf. socialis Dipolydora sp. D2
AM W31922 Polydora flava *Boccardiella sp. (cf.. magniovata CHECK limnicola)
*Boccardiella sp.
AM W31933 -1 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31933-2 Dipolydora socialis
*Boccardia chilensis *Boccardia chilensis
AM W31933-3 Dipolydora socialis
*Boccardiella sp? Spines cf. chilensis, one spine row, occ. ant, no posterior spines, no constriction, branchiae
*Boccardiella sp.
origin very close to notopodia base
AM W31945-1 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31945-2 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31947-1 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31947-2 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31947-3 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31947-4 Dipolydora socialis
Dipolydora flava Dipolydora flava
AM W31957- 2.3.3-1 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957- 2.3.3-2 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957 2-1-3.1 Polydora socialis
Dipolydora socialis (outside label)
Dipolydora socialis
AM W31957 2-1-3.2 Polydora socialis
Dipolydora socialis (outside label)
Dipolydora socialis
AM W31957 2-1-3.2 juv Polydora socialis
** Dipolydora socialis (outside label); juvenile specimen removed from analysis
** Dipolydora socialis
AM W31957 2-2-2.1 Polydora socialis
Dipolydora socialis (outside label)
Dipolydora socialis
AM W31957 2-2-3.1 Polydora socialis
Dipolydora cf. flava Dipolydora cf. flava
AM W31957 2-2-3.2 Polydora socialis
Dipolydora socialis Dipolydora socialis
AM W31957 2-2-3.3 Polydora socialis
Dipolydora socialis Dipolydora socialis
AM W31957 2-2-4 .1 Polydora socialis
Dipolydora socialis (outside label)
Dipolydora socialis
AM W31957 2-3-3.1.77 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of
Dipolydora sp. D1
main fang from stem, keys to socialis in B&K78 AM)
AM W31957 2-3-3.2.77 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957-2.3.3-3 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957-2.3.3-4 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957-2.3.3-5 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31957-2.3.3-6 Polydora socialis
Dipolydora Pseudopolydora??Carazziella smooth and bristled asymm (Dipolydora sp; lack of constriction on hh, angle of main fang from stem, keys to socialis in B&K78 AM)
Dipolydora sp. D1
AM W31965 Polydora armata
Dipolydora armata (label) Dipolydora armata
AM W7033 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-1 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-2 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-3 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-4 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-4 Boccardiella limnicola
* *Boccardiella limnicola
AM W7034-5 Boccardiella limnicola
* *Boccardiella limnicola
AM W7283 Polydora haswelli Holotype
Polydora haswelli
AM W8256 Polydora penicillata
As above Carazziella victoriensis
AM W8258 Polydora penicillata
Morphology indicates Carazziella sp. cf. victoriensis or phillipensis
Carazziella victoriensis
F123107 Carazziella MOV 3294
* *Carazziella MOV 3294
F124867 Pseudopolydora glandulosa MOV no 1160
* *Pseudopolydora glandulosa MOV no 1160
F42873 Polydora woodwicki Holotype
Polydora woodwicki
F43060 - 3 Polydora hoplura
Polydora hoplura
F43060-1 Polydora hoplura
Accidental omission **Polydora hoplura
F43060-2 Polydora hoplura
Polydora hoplura
G 2874 Polydora latispinosa Holotype
Polydora latispinosa
G10439 Polydora sp. Dipolydora tentaculata (but with posterior spines cf. aciculata). Specimen in three pieces. Not included in analysis
** see box at left
G10618 Polydora *Pseudopolydora paucibranchiata (based on original description this is probably not paucibranchiata (original 10 hh)
*Pseudopolydora sp.
G10642-1 Polydora sp 1 *Pseudopolydora paucibranchiata
*Pseudopolydora sp.
G10642-2 Polydora sp 1 *Pseudopolydora paucibranchiata
*Pseudopolydora sp.
G10642-3 Polydora sp 1 Dipolydora cf. socialis (no posterior)
Dipolydora socialis
G11390 Polydora websteri
??P. hoplura?? specimen lacking both spine packets on chaetiger 5
Not able to confirm
G2872 Polydora aciculata Holotype
Dipolydora aciculata Dipolydora aciculata
G2873-1 Polydora Dipolydora aciculata Dipolydora
aciculata Paratype
aciculata
G2885 Polydora tentaculata Holotype
Dipolydora tentaculata Dipolydora tentaculata
G2886 Polydora tentaculata Paratype
Dipolydora tentaculata Dipolydora tentaculata
NSW Fisheries Camden Haven specs -1 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -2 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -3 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -4 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -5 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -6 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
NSW Fisheries Camden Haven specs -7 Polydora calcarea (described as P. ciliata in Fauvel 1927)
Polydora cf. calcarea
QMunreg 1 Polydora tentaculata
Dipolydora flava/socialis needle packets/no occ./simple spines/
Dipolydora sp. D5S
QMunreg2 Polydora (2) *Boccardia chilensis *Boccardia chilensis
QMunreg3 Polydora (2) *Boccardia chilensis *Boccardia chilensis
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