Walker

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

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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

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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

117

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

119

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

123

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