kimberley marine biota. historical data: polychaetes...
TRANSCRIPT
Kimberley marine biota. Historical data: polychaetes (Annelida)
Pat Hutchings1*, Chris Glasby2, Maria Capa1,4 and Alison Sampey3
1 Australian Museum, 6 College Street, Sydney NSW 2010.
2 Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin NT 0801
3 Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC WA 6986
4 Museum of Natural History and Archaeology, Norwegian University of Science and Technology, NO-7491 Trondheim, Norway
* Email: [email protected]
ABSTRACT – We examined all the previously published records of polychaetes from the Kimberley Project Area as well as unpublished records from Australian museum records and updated any recent nomenclatural changes. Over 260 species from 43 families have been reported and we suggest that this number will increase considerably as more survey work is carried out in the area and additional habitats such as soft sediments are properly surveyed. At this stage many of these species appear to have wide distributions with few endemic species recorded, but we anticipate this will increase with taxonomic revisions incorporating both molecular and morphological techniques. We hope this paper will advertise the availability of this material to specialists for incorporation into their Australian revisions.
KEYWORDS: natural history collections, species inventory, Kimberley region, biodiversity, north west Australia, baseline
133–159 (2014) DOI: 10.18195/issn.0313-122x.84.2014.133-15984RECORDS OF THE WESTERN AUSTRALIAN MUSEUM
SUPPLEMENT
INTRODUCTION
The importance of utilising natural history collection datasets to provide baseline biodiversity information for conservation and environmental management decisions is now recognised (Pyke and Ehrlich 2010). The Kimberley region of Australia is currently of great interest for its conservation value, with a number of proposed marine protected areas, and also for its oil and gas reserves, fishing and aquaculture activities, nature based tourism, and proposed development (Department of Environment and Conservation 2009). Consequently, baseline data to characterise the values and assets in the region are needed (Wood and Mills 2008).
The Western Australian Museum (WAM) and other Australian natural history institutions have undertaken marine biodiversity surveys of the species present in the Kimberley, but many of these data and their interpretations are either unpublished or published in specialist taxonomic literature, so not readily accessible to researchers and managers. To address this, WAM instigated an extensive data compilation of major taxa known from the Kimberley region. Wilson (2014) has
reviewed the habitat and historical background of the Kimberley Project Area. Here, we document what is currently known about shallow water polychaete diversity within the Kimberley Project Area.
POLYCHAETES (ANNELIDA)
Polychaetes are a major component of the marine benthos in terms of number of both individuals and species. They range in length from less than 1 mm to over 1 m. Polychaetes are currently classified into more than 80 families with over 15,000 species known worldwide and many remaining to be described. They vary considerably in morphology and life style, including many different types of feeding strategies (carnivores, deposit and filter feeders, herbivores, parasitic or commensal). They exhibit a considerable range of reproductive strategies including both sexual and asexual. Some species live only a few weeks, while others live for several years. They may breed almost continuously, or have a restricted breeding season. Some survive spawning and others do not. They live in a wide variety of marine habitats from the supra-littoral to the deep sea: some live in sediments, others bore
134 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
into hard substrates or are firmly attached to them, while others move freely over the substrate, swim in the water column or are commensal on a range of animals. Some are tubiculous, living in muddy/sandy or calcareous tubes. For all these reasons polychaetes have been widely used as surrogates for the entire benthic community. Because they occupy all levels within the food chain a diverse polychaete assemblage provides a good indication of a healthy environment. In addition, some taxa (e.g. some species of Capitellidae, such as Capitella spp.) are good indicators of polluted environments. As well as marine species, others occur in estuarine areas and freshwater habitats, and some terrestrial species live in damp environments (Beesley et al. 2000; Rouse and Pleijel 2001).
HISTORY
Historical collections of polychaetes from the Kimberley coast are scarce. In the late 1800s and early 1900s two expeditions collected in Western Australia: the German S.M.S. Gazelle between 1874 and 1876 and the south west Australian Hamburg expedition led by Michaelsen and Hartmeyer in 1905. The former did not extend north of the Dampier Archipelago and the latter only reached north to Shark Bay. Apparently, the only European expedition that sampled the Kimberley coast was the Swedish Mjöberg Scientific Expedition (1910–1911), for which the polychaetes were reported by Johansson (1918) and Augener (1922). All the polychaetes described by Augener came from a small area off Cape Jaubert in the southern Kimberley. No further notable collecting of polychaetes occurred until 1975 when Hartmann-Schröder and Hartmann collected at several sites around Broome and Derby as part of their Australia-wide investigation of intertidal polychaetes and ostracods. All the specimens studied by Augener (1922) and Hartmann-Schröder (1979) – over 260 specimen lots – were deposited in the Zoologisches Institut und Zoologisches Museum der Universität
Hamburg, Germany (HZM), except for some paratypes, which are housed at WAM.
The first extensive survey of polychaetes along the Kimberley coast was made in 1988 (Hutchings 1989). Polychaetes were collected from 29 locations with 121 species identified (Table 1). Many of the species collected during this survey were new and have been described subsequently (Hutchings and Glasby 1990; Hutchings and Reid 1990, 1991; Pillai 2009; Murray et al. 2010). Polychaetes were also collected during a 1991 survey of 21 locations along the coast between Broome and Wyndham (referred to as the KIRE 91), by WAM where 54 taxa were identified, to either species or family level by Hanley (1992). Hanley recorded the greatest diversity within the Polynoidae with 20 species identified, including three new species. Other common families were the Nereididae and Terebellidae. Hanley was particularly interested in polynoids commensal with holothurians, crinoids, alcyonarians and gorgonians and with one species commensal in tubes of terebellids. In a second survey to the southern Kimberley in 1994, polychaetes associated with intertidal hard substrates and mangroves were collected. Specimens from both surveys are lodged at the Museum and Art Gallery of the Northern Territory (MAGNT). Hanley (1995) provides a list of the polychaete families collected with the polynoids and the eunicids identified to species. Both of these reports suggest a high diversity of polychaetes with new species as well as widely distributed ones commonly found in similar habitats throughout northern Australia, although many of these widely distributed species need to be carefully re-examined to confirm their identifications.
Additional surveys were carried out by MAGNT and also by WAM in the 1980s and in 2008, but polychaetes were not specifically targeted and no species lists were compiled. These included an initial 1984 survey of Ashmore, Cartier and Hibernia Reefs by MAGNT (Russell 2005), followed by those in 1987 and 1992, yielding many polychaete specimens now housed at MAGNT.
Survey year No. locations No. species No. families Reference
1988 29 121 13 Hutchings 1989*
1991 21 54+ 21 Hanley 1992
1994 10 50+ 22 Hanley 1995
TABLE 1 Number of locations, species and families of the polychaetes sampled during main survey expeditions in the region and presented as reports. *Number of species not presented in the report so the number was extracted from the database.
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 135
All the records discussed above are incorporated into the present study.
Previous research in the region on other taxonomic groups such as fishes (Hutchins 1999), echinoderms (Marsh and Marshall 1983) and corals (Veron and Marsh 1988) revealed some general patterns of the fauna in the region. Firstly, species richness and composition were different inshore compared to offshore. Secondly, more endemic species occur inshore compared to more widespread species offshore: we have tested this using data collected by Hanley for the distribution of commensal species of polynoids.
AIMS
1. Collate records of shallow water (<30 m) polychaete species in the Kimberley Project Area that are verified by specimens lodged in Australian museum collections (1880s – 2009) to provide a baseline diversity dataset;
2. identify collection and taxonomic bias and gaps;
3. test whether there are proportionally more wide ranging species (e.g. Indo-West Pacific or Indo-Pacific) occurring offshore compared to inshore – i.e. species that inhabit clear oligotrophic waters versus endemic or Indo-Australian species that would have higher tolerances to the silty turbid waters occurring along the coast; and
4. explore cross shelf differences in species richness and composition for commensal Polynoid species.
METHODS
The Kimberley Project Area encompasses an area west and north of the Kimberley coast (from south of Broome northward and eastward to the Western Australia–Northern Territory border) extending beyond the 1000 m bathymetric contour, with the coastline forming a natural inshore boundary, as shown in Figure 1 (see Sampey et al. 2014, for a full explanation of the study area).
For the purpose of this synthesis we are using the classical concept and family membership of the polychaetes (e.g. Fauchald 1977, Rouse and Pleijel 2001). Largely based on molecular data, the clitellates (leeches and oligochaetes), sipunculans, and siboglinids have been moved recently into the polychaetes, but these groups are even less well known from the region. So, while clitellates and sipunculans are widespread and probably diverse in the Kimberley, they are poorly known and will not be considered any further in this review.
The methodology follows that outlined by Sampey et al. (2014). Briefly, polychaete species data were sourced from the collection databases of WAM (July 2010), MAGNT (August 2009) and AM (August 2009), as well as the species reported on WAM survey expeditions of 1988, 1991 and 1994 along the Kimberley coast (Hutchings 1989; Hanley 1992, 1995).
Species names represent a hypothesis that is subject to change as new informat ion (morphology, genetic, behaviour, distribution ranges) is discovered (Gaston and Mound 1993); at this stage al l species l isted herein are recognised only on the basis of morphological characters (i.e. morphospecies). The species names and taxonomic placement of the records in the dataset were checked in an attempt to present the currently accepted name and to resolve synonymies and old combinations, but the specimens were not re-examined for this study (Sampey et al. 2014). Species names were checked using online databases (Appeltans et al. 2010; ABRS 2011) to identify the currently accepted taxonomic name, and based either on recently published data or research currently being undertaken. However, this does mean that the species names are more up-to-date for some families, and this reflects the current knowledge of Australian polychaetes.
SPATIAL INFORMATION, COLLECTION DETAILS AND MAPPING
As described in Sampey et al. (2014), data from all sources were collated into a single database. Location and collecting details were checked and verified. The locations of the specimen records were visualised using ARCGIS v9, ArcMap v9.3. Maps of species richness and sampling effort were generated for each main location. Since species richness patterns are highly dependent on sampling effort, we calculated the number of collecting events at a location to provide an indication of relative sampling effort. A collecting event was defined by the season and year of collecting, and the full list of locations, latitude and longitude and other relevant collection information is provided for all taxonomic groups in Table 4 in Sampey et al. (2014).
Throughout this paper ‘inshore‘ refers to locations along the coast, and the numerous islands and reefs found shoreward of the 50 m depth contour (Figure 1). ‘Offshore‘ refers to the shelf edge atolls, which arise from deeper waters (200–400 m) along the continental margin.
136 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
BIOGEOGRAPHIC AND HABITAT CODING
Species were coded for their known habitat and biogeographic range (Sampey et al. 2014). If a species did not conform to a single code then appropriate combinations were used.
The following biogeographic codes were used:
• Western Australian endemic (WA). Currently known only from Western Australian waters, often from the type locality only; may eventually prove to be a Northern Australian endemic with more collecting effort.
• Northern Australian endemic (NA). Found throughout tropical Australian waters.
• Southern Australian endemic (SA). Found throughout temperate Australian waters and its presence in the Kimberley is the northern extent of its known range; although this may represent a lack of data and should be treated with caution.
• Australian endemic (A). Found throughout tropical and temperate Australian waters.
• Indo-Australian (IA). Found throughout Australian and Indonesian waters, may extend to the Philippines.
• Indo-West Pacific (IWP). Found throughout the Red Sea, Indian and Western Pacific Oceans.
• Indo-Pacific (IP). Found throughout the Red Sea, Indian Ocean and throughout the Pacific Ocean.
• West Pacific (WP). Found throughout the Western Pacific, although presence in Kimberley indicates its occurrence in at least the eastern Indian Ocean.
• Central Pacific (CP)*. Found throughout the Central Pacific.
• Tropicopolitan (T)*. Found throughout all tropical oceans.
• Circumglobal (C)*. Found throughout all oceans in both tropical and temperate waters.
Codes indicated by an asterisk (*) are the most problematic and need to be treated with caution; this is especially true for families for which no recent Australian revision has been undertaken. Our knowledge of the distribution and habitat of the polychaete fauna of the Pacific, South East Asia and the Indian Ocean is poor and dominated by widespread species. However, we expect with additional studies, many of these species will be found to consist of several morphologically similar species each with discrete non-overlapping ranges.
Within its distribution range, a species may be restricted to certain habitat types. We provided an indication of the habitats sampled in the Kimberly
region to date by coding species for their preferred habitat, if known, as follows:
• Intertidal (i). Found in the intertidal zone, which is extremely large as a result of the large tidal range.
• Subtidal (s). Found in the subtidal or sublittoral zone.
• Hard substrate (H). Found associated with hard substrates (e.g. rock, coral, rubble).
• Soft substrate (S). Found associated with soft substrates (e.g. sand, mud).
• Estuarine (E). Found in estuarine or brackish waters.
• Epizoic (EZ). Always found in an external association with a particular species of animal.
• Unknown (U).
DATA ANALYSES
To explore composition differences across the shelf, the total number of commensal polynoid species was calculated for each location (reef, island, or atoll) and then visualised using non-metric multi dimensional scaling (nMDS) using the Bray-Curtis distance measure.
RESULTS
NUMBER OF SPECIMENS IN COLLECTIONS
A total of 1,046 registered specimen lots of polychaete species were included in this dataset (excluding lots in overseas museums) (Table 2a). A 'lot' represents between one and many individuals from the same site. The number of lots included was variable across families, ranging from one for 11 families, which usually coincided with that family not being identified to species (e.g. Maldanidae species), to 210 for the Polynoidae, the most consistently collected and identified family in the Project Area (Tables 2 and 3). The number of specimen lots housed in the various institutions was also variable (413 species, 40%, AM; 588 species, 56%, MAGNT; and 45 species, 4%, WAM; Table 2).
Many lots were excluded from the present dataset (814 species, 44%; Table 2b). This was primarily due to incomplete identification or where the records are not consistent with any recognisable nomenclature. This material needs to be re-examined. Three lots were collected in waters deeper than 30 m. A further 125 lots of polychaetes are available in WAM, but these were not included in the dataset due to late cataloguing of samples.
The oldest Kimberley polychaete record housed in
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 137
collections of Australian museums was Myzostoma stochoeides Atkins, 1927, taken off crinoids collected by H.L. Clark at Cape Villaret in 1929 and housed in the AM collections. A number of species, including Perkinsyllis augeneri (Hartmann-Schröder, 1979) collected by Hartmann-Schröder at Broome in 1975, are located in the collections of WAM. The oldest MAGNT Kimberley specimen was Platynereis uniseris Hutchings and Reid, 1991 collected from Ashmore Reef in 1983. This reflects the dearth of collecting in the Kimberley region from the early 1900s to the late 1970s.
Up to 2009, 30 new polychaete species have been described from the Kimberley, belonging to eight families (Capitellidae, Glyceridae, Nereididae, Phyllodocidae, Pilargidae, Serpulidae, Syllidae, and Terebellidae). In addition, several new species and genera of Polynoidae in the MAGNT collection have been flagged as new by R. Hanley, but have yet to be formerly described (Table 3). Work is underway by M. Capa and C. Watson (MAGNT) describing new species of Sabellidae and Chrysopetalidae, respectively. It is anticipated that future collecting events in the Project Area will yield many more new species and records of polychaetes for the area.
SPECIES RICHNESS AND COLLECTING EFFORT
A total of 261 taxa belonging to 43 families of polychaetes were recorded in our dataset (Table 3). Sixteen families have not yet been sorted to morphospecies: Euphrosinidae, Lumbrineridae, Nephtyidae, Sphaerodoridae, Maldanidae, Opheliidae, Orbiniidae, Paraonidae, Questidae, Scalibregmatidae, Chaetopteridae, Magelonidae, Poecilochaetidae, Spionidae, Ampharetidae, and Flabelligeridae (Table 3). In part this reflects the lack of expertise in these families worldwide, although some are currently being investigated, and this paper may help to alert the worldwide polychaete community to the availability of this material for study.
Polychaete data are available for 69 locations in the Project Area (Table 4, Figures 1–3). Species richness ranged from 80 (at Broome) to one (at 12 inshore and one offshore location; Table 4, Figure 2). Collecting effort was highly variable, with nine collecting events occurring at Broome, but only one at 48 other locations (Table 4, Figure 3). The number of polychaete families collected at any one location was also highly variable with 29 families collected at Broome compared to only one family at 16 other locations (Table 4). The presence or absence of polychaete families reported at a location very much depends on the habitats sampled and the collector. More polychaete species have been reported from
inshore (232 species) than offshore (82 species; Tables 3, 6 and 7). However, collecting effort is substantially different for these areas of the shelf, with more collecting inshore compared to offshore or midshelf (25 versus 13 locations, respectively). At the time of writing, no collecting had occurred at Browse Island (midshelf) for polychaetes. The eastern Kimberley has not been sampled for polychaetes, nor have Seringapatam or Imperieuse Reef (Rowley Shoals) (Figure 1). The Polynoidae is likely to have had a more even collecting effort than others (see Discussion) and of the 33 species total, 27 are reported from inshore sites, 21 from offshore sites and 15 occurred in both. There is a weak inshore versus offshore difference in species composition (MDS plot, Figure 4), which may be explained partially because many species recorded here are commensal on echinoderms, which show a similar inshore versus offshore difference in both richness and composition (Sampey and Marsh 2015).
BIOGEOGRAPHY AND HABITATS
The majority of polychaete species were widespread (in all categories C, CP, IA, IP, IWP, T, WP) both inshore (111 species, 49%) and offshore (45 species, 54%) (Table 5).
The proportion of endemics (those in categories WA, A, NA and SA) inshore (87 species, 38%) was more than twice the proportion of endemics offshore (12 species, 14%) (Table 5). The proportion of Indo-Australian species was also higher inshore (20 species, 9%) versus offshore (5 species, 6%) (Table 5). The dominance of widely distributed species is also reflected in the Polynoidae. In this family one, and possibly up to four, species are narrow range endemics (Australaugeneria michaelseni Pettibone, 1969). Three new undescribed taxa identified by Hanley (Table 3, Figure 1, 2), and another three polynoid species occur in the Kimberley as well as in other parts of Australia. Twenty five species of polynoids found in the Kimberley also occur throughout the Indo-West Pacific region.
A high proportion of the species were associated with hard substrates: 70 species (31%) inshore versus 19 species (23%) offshore. However, some species were associated with soft substrates (41 species, 18% inshore versus 4 species, 5% offshore, Table 6). Three species were estuarine (Table 6). Most species had an unknown substrate preference (72 species, 34% inshore versus 27 species, 32% offshore; Table 6). However, it should be noted that many families not represented are those that live exclusively buried in soft sediments, which were not sampled.
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kilometres
Polychaetes
FIGURE 1 Location of records of polychaete species in the Kimberley Project Area of Western Australia. The Project Area boundary is marked in grey. Map projection: GDA94, Scale: 1:6, 250,000.
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 139
DISCUSSION
These results demonstrate that the polychaete fauna of the Project Area remains largely under sampled and undescribed. Collection gaps are methodological since the sampling effort has been concentrated mostly in shallow water hard substrates. Sampling across the shelf, including soft sediments between reefs, certainly would reveal a highly diverse polychaete assemblage. The data are also biased towards larger species, and fine sorting of soft sediments and washings from broken up hard substrates would yield additional species. Further, existing collections reflect surveys that were inventory focussed and somewhat biased toward certain families, which were the interest of the particular workers undertaking the collecting (e.g. Polynoidae).
Identification efforts have been greatest on certain families (e.g. Nereididae, Terebellidae, Polynoidae), which have been the subject of recent taxonomic research. Within the WAM, AM and MAGNT collections there are unidentified polychaetes, some sorted to family and included in this study, others not. This highlights the lack
of funding for taxonomic research and the lack of taxonomic expertise in particular polychaete families. Nevertheless, some of this material (e.g. Sabellidae, Sabellariidae, Oweniidae) is currently being examined with new species to be described. We also anticipate that some of these specimens will be found to represent species endemic to the areas.
It is premature to make any generalisations about the polychaete fauna of the Project Area as so much is still unknown. Our discussion focuses on what is known from the limited families studied to date and highlights some collection and taxonomic gaps, which will provide a basis to focus future research.
SPECIES RICHNESS PATTERNS
The 261 taxa listed in our dataset represent a low species richness estimate of polychaetes for the Project Area if compared to other tropical areas (Hutchings personal observation). Causes for this low diversity are the limited collecting effort mentioned above together with the superficial identification of specimens of particular families.
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50 m
100 m
100 m
100 m
1000
m
50 m
30 m
30 m
DERBYBROOME
WYNDHAM
128°0'E
128°0'E
125°0'E
125°0'E
122°0'E
122°0'E
119°0'E
119°0'E
12°0
'S
12°0
'S
15°0
'S
15°0
'S
18°0
'S
18°0
'S
±
KIMBERLEY
0 50 100 150 200
kilometres
No. of Collecting Events
!( 1!( 2!( 3 - 5
!( 6 - 10!( 11 - 20!( 21+
Polychaetes
FIGURE 3 Number of collecting events for polychaetes at each main collecting location. This was based on a count of the season code and provides an indication of sampling effort. Map projection: GDA94, Scale: 1:6, 250,000.
140 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Included families AM MAGNT WAM Total Excluded families AM MAGNT WAM Total
Acoetidae 2 2 Acoetidae 1 1
Ampharetidae 1 1 Amphinomidae 25 4 29
Amphinomidae 1 5 6 Aphroditidae 1 1 2
Aphroditidae 4 2 1 7 Arenicolidae 2 2
Capitellidae 1 1 Capitellidae 33 33
Chaetopteridae 19 19 Chrysopetalidae 3 53 56
Chrysopetalidae 29 5 34 Cirratulidae 27 27
Cirratulidae 1 1 Dorvilleidae 5 5
Dorvilleidae 1 1 Eunicidae 1 96 2 99
Eunicidae 2 28 30 Euphrosinidae 3 3
Euphrosinidae 1 1 Glyceridae 16 1 17
Flabelligeridae 5 5 Hesionidae 23 1 24
Glyceridae 11 1 12 Lumbrineridae 26 26
Goniadidae 4 1 5 Myzostomidae 1 1
Hesionidae 11 11 Nereididae 2 80 13 95
Lumbrineridae 2 2 Oenonidae 16 16
Magelonidae 1 1 Onuphidae 2 2 3 7
Maldanidae 9 9 Oweniidae 4 1 5
Myzostomidae 1 12 13 Pectinariidae 1 1
Nephtyidae 7 7 Phyllodocidae 26 26
Nereididae 70 54 16 140 Pilargidae 3 3
Oenonidae 1 1 Polynoidae 30 71 2 103
Onuphidae 3 4 7 Sabellariidae 4 1 5
Opheliidae 26 26 Sabellidae 6 32 1 39
Orbiniidae 14 14 Serpulidae 24 14 2 40
Oweniidae 9 9 Sigalionidae 7 7
Paraonidae 2 2 Syllidae 12 84 96
Pectinariidae 1 1 2 Terebellidae 1 45 46
Phyllodocidae 10 1 11 Total 82 697 35 814
Pilargiidae 1 1
Poecilochaetidae 2 2
Polynoidae 4 206 210
Questidae 1 1
Sabellariidae 1 1 2
Sabellidae 31 8 39
Scalibregmatidae 3 3
Serpulidae 96 45 1 142
Sigalionidae 1 2 3
Sphaerodoridae 1 1
Spionidae 14 14
Syllidae 75 11 6 92
Terebellidae 88 64 3 155
Trichobranchidae 1 1
Total 413 588 45 1046
TABLE 2 Number of registered specimen lots of Kimberley polychaetes housed in Australian natural history collections. Included are those lots identified to species or able to be distinguished as a separate species. Excluded lots are those that were incompletely identified or from deepwater (>30 m) locations.
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 141
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Fam
ily:
Am
ph
inom
idae
Chl
oeia
flav
a (P
alla
s, 1
766)
IPU
s●
Eur
ytho
e co
mpl
anat
a (P
alla
s, 1
766)
**T
Us
●●
Har
tman
n-Sc
hröd
er (1
979)
Pse
udeu
ryth
oe o
culif
era
(Aug
ener
, 191
3)*
Ish●
Har
tman
n-Sc
hröd
er (1
979)
Fam
ily:
Dor
vill
eid
aeD
orvi
llea
sp.
●
Fam
ily:
Eu
nic
idae
Eun
ice
afra
Pet
ers,
185
4IP
Uis
●E
unic
e af
ra p
unct
ata
Pete
rs, 1
854*
TU
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Eun
ice
ante
nnat
a Sa
vign
y in
Lam
arck
, 181
8*
●A
ugen
er (1
922)
Eun
ice
aphr
odit
ois
(Pal
las,
178
8)E
unic
e cf
. joh
nson
i Har
tman
, 195
4●
Eun
ice
long
icir
ris
Gru
be, 1
869*
IWP
Us
●A
ugen
er (1
922)
Eun
ice
paup
era
Gru
be, 1
878
Eun
ice
sici
liens
is (G
rube
, 184
0)*
NA
Us
●●
Aug
ener
(192
2)E
unic
e to
rres
iens
is M
cInt
osh,
188
5E
unic
e tu
bife
x C
ross
land
, 190
4**
IWP
Uis
●H
artm
ann-
Schr
öder
(197
9) B
room
e, A
ugen
er (1
922)
Lysi
dice
nin
etta
Aud
ouin
and
Miln
e E
dw
ard
s, 1
833*
*C
Us
●●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Lysi
dice
rob
usta
Sti
mps
on, 1
856*
IWP
Us
●A
ugen
er (1
922)
Mar
phys
a m
ossa
mbi
ca (P
eter
s, 1
854)
Nem
aton
erei
s sp
. nov
. ●
●N
emat
oner
eis
unic
orni
s (G
rube
, 184
0)*
CU
is●
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
alol
a si
cilie
nsis
(Gru
be, 1
840)
●
Fam
ily:
Eu
ph
rosi
nid
aeE
uphr
osin
idae
sp.
●
Fam
ily:
Lu
mb
rin
erid
aeLu
mbr
iner
is s
p.*
●H
artm
ann-
Schr
öder
(197
9) B
room
eLu
mbr
iner
idae
spp
. ●
Fam
ily:
Oen
onid
aeA
rabe
lla ir
icol
or (M
onta
gu, 1
804)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Ara
bello
neri
s br
oom
ensi
s H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
TABL
E 3
Spe
cies
of
poly
chae
tes
reco
rded
fro
m t
he P
roje
ct A
rea.
Cod
es a
re d
efine
d in
Bio
geog
raph
ic a
nd H
abita
t C
odin
g, a
bove
. The
sup
ersc
ript
asso
ciat
ed w
ith t
he E
Z ha
bita
t co
de in
dica
tes
the
anim
al w
ith w
hich
the
pol
ycha
ete
spec
ies
is a
ssoc
iate
d; s
oft
cora
ls1 ,
crin
oids
2 , o
phiu
roid
s3 , h
olot
huria
n Stic
hopu
s4 .
142 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Dri
lone
reis
qua
drio
cula
ta H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Oen
one
fulg
ida
Savi
gny
in L
amar
ck, 1
818*
●A
ugen
er (1
922)
Oen
one
sp.
Fam
ily:
Dor
vill
eid
aeSc
hist
omer
ingo
s ne
glec
ta (F
auve
l, 19
23)*
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Fam
ily:
On
up
hid
ae●
Dio
patr
a cu
prea
cup
rea
(Bos
c, 1
802)
*W
AU
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Dio
patr
a de
ntat
a K
inbe
rg, 1
865*
●A
ugen
er (1
922)
Dio
patr
a lil
liput
iana
Pax
ton,
199
3D
iopa
tra
mac
ulat
a Pa
xton
, 199
3N
AU
s●
Fam
ily:
Aco
etid
aeP
olyo
dont
es a
trom
argi
natu
s H
orst
, 191
7IA
Us
●P
olyo
dont
es s
ibog
ae H
orst
, 191
7IA
Us
●
Fam
ily:
Ap
hro
dit
idae
Aph
rodi
ta a
ustr
alis
Bai
rd, 1
865
IWP
Us
●A
phro
dita
kul
mar
is H
utch
ings
and
McR
ae, 1
993
AU
s●
Laet
mon
ice
mol
ucca
na (H
orst
, 191
6)IA
Us
●P
alm
yra
auri
fera
Sav
igny
in L
amar
ck, 1
818
IPH
s●
Pon
toge
nia
areo
cera
s (H
asw
ell,
1883
)A
Hs
●
Fam
ily:
Ch
ryso
pet
alid
aeA
rich
lidon
han
nelo
reae
Wat
son
Rus
sell,
199
8IP
Hs
●B
haw
ania
am
boin
ensi
s H
orst
, 191
7IW
PH
is●
Bha
wan
ia r
ivet
i (G
ravi
er, 1
908)
IPH
is●
Bha
wan
ia s
p. A
*IP
Hs
●H
artm
ann-
Schr
öder
(197
9) B
room
eC
hrys
opet
alum
rem
anei
Per
kins
, 198
5●
Chr
ysop
etal
um s
p. 1
1 W
atso
n, m
sIP
Hs
●C
hrys
opet
alum
sp.
5 W
atso
n, m
sIP
Hs
●C
hrys
opet
alum
sp.
7 W
atso
n, m
sIP
Hs
●P
alea
notu
s sp
. 8 W
atso
n, m
sIP
Hs
●Tr
epto
pale
par
omol
os W
atso
n, 2
010
IPH
s●
●Tr
epto
pale
sp.
3 W
atso
n, m
sIP
Hs
●
Fam
ily:
Gly
ceri
dae
Gly
cera
der
beye
nsis
Har
tman
n-Sc
hröd
er, 1
979*
Har
tman
n-Sc
hröd
er (1
979)
Der
byG
lyce
ra la
ncad
ivae
Sch
mar
da,
186
1*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Gly
cera
mac
into
shi G
rube
, 187
7IW
PU
is●
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 143
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Gly
cera
trid
acty
la S
chm
ard
a, 1
861
CU
is●
Hem
ipod
ia s
impl
ex (G
rube
, 185
7)C
Uis
●
Fam
ily:
Gon
iad
idae
Gly
cind
e bo
nhou
rei G
ravi
er, 1
904
IWP
Us
●G
lyci
nde
nipp
onic
a Im
ajim
a, 1
967*
IWP
Us
●H
artm
ann-
Schr
öder
(197
9) B
room
eG
onia
da e
mer
ita
Aud
ouin
and
Miln
e E
dw
ard
s, 1
834*
●G
onia
da p
auci
dens
Gru
be, 1
878
●
Fam
ily:
Hes
ion
idae
Hes
ione
spl
endi
da S
avig
ny in
Lam
arck
, 181
8**
IWP
Us
●●
Aug
ener
(192
2)Le
ocra
tes
chin
ensi
s K
inbe
rg, 1
866*
*IW
PU
s●
●A
ugen
er (1
922)
Leoc
rati
des
sp.
●Si
gam
bra
pett
ibon
ea H
artm
ann-
Schr
öder
, 197
9*H
artm
ann-
Schr
öder
(197
9) B
room
e
Fam
ily:
Myz
osto
mid
aeW
A●
Myz
osto
ma
ambi
guum
von
Gra
ff, 1
887
IWP
EZ
2●
Myz
osto
ma
cirr
icos
tatu
m Jä
gers
ten,
193
7IW
PE
Z2
●M
yzos
tom
a na
nsen
i von
Gra
ff, 1
887
IWP
EZ
2●
●M
yzos
tom
a po
lycy
clus
Atk
ins,
192
7T
EZ
2●
●M
yzos
tom
a st
ocho
eide
s Atk
ins,
192
7IW
PE
Z2
●
Fam
ily:
Nep
hty
idae
Nep
htyi
dae
sp.
●
Fam
ily:
Ner
eid
idae
Cer
aton
erei
s au
stra
lis H
artm
ann-
Schr
öder
, 198
5A
Us
●●
Cer
aton
erei
s m
irab
ilis
Kin
berg
, 186
6C
Us
●●
Cer
aton
erei
s pe
rkin
si H
artm
ann-
Schr
öder
, 198
5A
Us
●●
Com
pose
tia
sp.
●Le
onna
tes
crin
itus
Hut
chin
gs a
nd R
eid
, 199
1N
AU
is●
Leon
nate
s in
dicu
s K
inbe
rg, 1
866
IWP
Hs
●M
icro
nere
is b
anse
i (H
artm
ann-
Schr
öder
, 197
9)A
Us
●N
amal
ycas
tis
abiu
ma
(Gru
be, 1
872)
TU
is●
Nea
nthe
s bo
ngco
i Pill
ai, 1
965
IAU
s /E
●N
eant
hes
daw
ydov
i (Fa
uvel
, 193
7)IA
Us
●N
eant
hes
cf. n
anha
iens
is W
u, S
un a
nd Y
ang,
198
1●
Nea
nthe
s pa
chyc
haet
a (F
auve
l, 19
18)
IPH
s●
●N
eant
hes
unifa
scia
ta W
illey
, 190
5IW
PH
s●
Ner
eis
cock
burn
ensi
s Aug
ener
, 191
3SA
Hs
●
144 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Ner
eis
denh
amen
sis A
ugen
er, 1
913
IAH
s●
Ner
eis
jack
soni
Kin
berg
, 186
6*●
Aug
ener
(192
2)N
erei
s se
mpe
rian
a (G
rube
, 187
8)*
Aug
ener
(192
2)N
erei
s (N
eant
hes)
sp.
*H
artm
ann-
Schr
öder
(197
9) B
room
eN
icon
sp.
●
Par
aleo
nnat
es b
olus
(Hut
chin
gs a
nd R
eid
, 199
1)IA
Si●
Per
iner
eis
broo
men
sis
Har
tman
n-Sc
hröd
er, 1
979*
IASi
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
erin
erei
s ai
buhi
tens
is (G
rube
, 187
8)●
Per
iner
eis
helle
ri (G
rube
, 187
8)**
IWP
Uis
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
erin
erei
s ni
grop
unct
ata
(Hor
st, 1
889)
**IW
PU
is●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Per
iner
eis
nunt
ia (S
avig
ny in
Lam
arck
, 181
8)IP
Si●
Per
iner
eis
obfu
scat
a (G
rube
, 187
8)IA
H/
Si●
Per
iner
eis
sing
apor
iens
is (G
rube
, 187
8)IP
Uis
●P
erin
erei
s su
luan
a (H
orst
, 192
4)IW
PH
s●
Per
iner
eis
vanc
auri
ca (E
hler
s, 1
868)
**IW
PH
/Si
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
laty
nere
is p
olys
calm
a C
ham
berl
in, 1
919
IWP
Us
●●
Pla
tyne
reis
uni
seri
s H
utch
ings
and
Rei
d, 1
991
AU
is●
●P
seud
oner
eis
anom
ala
Gra
vier
, 190
1IW
PH
s●
●P
seud
oner
eis
rott
nest
iana
Aug
ener
, 191
3*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pse
udon
erei
s sp
p.
●Q
uadr
icir
ra b
anse
i Har
tman
n-Sc
hröd
er, 1
979*
●H
artm
ann-
Schr
öder
(197
9) B
room
eR
ullie
rine
reis
sp.
●
Sim
plis
etia
aeq
uise
tis
(Aug
ener
, 191
3)SA
E●
Sim
plis
etia
ery
thra
eens
is (F
auve
l, 19
18)
IWP
Ss●
Solo
mon
oner
eis
mar
auen
sis
Gib
bs, 1
971
WP
Si●
Web
ster
iner
eis
sp.
●
Fam
ily:
Ph
yllo
doc
idae
Ana
itid
es m
adei
rens
is (L
ange
rhan
s, 1
879)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Ana
itid
es s
p.
●H
artm
ann-
Schr
öder
(197
9) B
room
eE
teon
e sp
. ●
Eul
alia
mic
roph
yllo
ides
Har
tman
n-Sc
hröd
er, 1
979
WA
UH
artm
ann-
Schr
öder
(197
9) B
room
eE
umid
a sp
. ●
Gen
etyl
lis c
asta
nea
Mar
anze
ller,
1879
*●
●H
artm
ann-
Schr
öder
(197
9) B
room
eN
otop
hyllu
m s
plen
dens
(Sch
mar
da,
186
1)*
●H
artm
ann-
Schr
öder
(197
9) B
room
eN
otop
hyllu
m c
f. sp
lend
ens
(Sch
mar
da,
186
1)●
Phy
llodo
ce (A
nait
ides
) dup
lex
McI
ntos
h, 1
885*
IWP
Uis
●A
ugen
er (1
922)
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 145
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Phy
llodo
ce m
alm
gren
i? G
ravi
er, 1
900
Pro
tom
ysit
ides
usc
hako
vi H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pro
tom
ysti
des?
sp.
●
Fam
ily:
Pil
argi
dae
Siga
mbr
a pe
ttib
onea
e H
artm
ann-
Schr
öder
, 197
9N
AU
is
Fam
ily:
Pol
ynoi
dae
●A
ustr
alau
gene
ria
mic
hael
seni
Pet
tibo
ne, 1
969
WA
Us
●A
ustr
alau
gene
ria
pott
si P
etti
bone
, 196
9IW
PU
s●
●A
ustr
alau
gene
ria
ruti
lans
(Gru
be, 1
878)
IAE
Z1
Gas
trol
epid
ia c
lavi
gera
Sch
mar
da,
186
1IW
PE
Z4
●●
Gau
dich
audi
us c
imex
(Qua
tref
ages
, 186
5)IW
PU
s●
●ge
n. n
ov. s
p. n
ov. 1
Han
ley,
unp
ublis
hed
gen.
nov
. sp.
nov
. 2 H
anle
y, u
npub
lishe
d●
gen.
nov
. sp.
nov
. 4 H
anle
y, u
npub
lishe
d●
Hal
osyd
nops
is p
ilosa
(Hor
st, 1
917)
IWP
Us
●H
arm
otho
e ch
arlo
ttae
Hut
chin
gs a
nd M
urra
y, 1
984
SASs
●H
arm
otho
e di
ctyo
phor
a (G
rube
, 187
8)**
IAH
s●
Aug
ener
(192
2)H
arm
otho
e pr
aecl
ara
(Has
wel
l, 18
83)
SASs
●H
arm
otho
e ve
sicu
dent
a H
anle
y an
d B
urke
, 199
1W
PU
s●
●H
erm
enia
aca
ntho
lepi
s (G
rube
, 187
6)IW
PU
s●
Hol
olep
idel
la c
omm
ensa
lis W
illey
, 190
5*IP
EZ
3●
Aug
ener
(192
2)H
olol
epid
ella
nig
ropu
ncta
ta (H
orst
, 191
5)Ip
hion
e m
uric
ata
(Sav
igny
in L
amar
ck, 1
818)
**IW
PH
s●
●A
ugen
er (1
922)
Iphi
one
ovat
a K
inbe
rg, 1
856
WP
Us
●●
Lagi
sca
hirs
uta
Rul
lier,
1972
IPH
s●
●Le
pida
sthe
nia
spp.
●
●Le
pido
notu
s ad
sper
sus
(Gru
be, 1
878)
IAU
s●
Lepi
dono
tus
cari
nula
tus
(Gru
be, 1
870)
IWP
Us
●●
Lepi
dono
tus
cris
tatu
s (G
rube
, 187
5)IW
PU
s●
●Le
pido
notu
s gl
aucu
s (P
eter
s, 1
854)
IWP
Us
●Le
pido
notu
s (T
horm
ora)
juke
si B
aird
, 186
5*A
Sis●
Aug
ener
(192
2)Le
pido
notu
s oc
ulat
us B
aird
, 186
5*H
artm
ann-
Schr
öder
(197
9) B
room
eLe
pido
notu
s st
ella
tus
Bai
rd, 1
865*
●A
ugen
er (1
922)
Lepi
dono
tus
york
ianu
s Aug
ener
, 192
2**
Par
adyt
e cr
inoi
dico
la (P
otts
, 191
0)IP
EZ
2●
146 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Par
adyt
e te
ntac
ulat
a (H
orst
, 191
5)IA
Us
●●
Par
alep
idon
otus
am
pulli
feru
s (G
rube
, 187
8)IP
Us
●●
Par
alep
idon
otus
ery
thro
mar
is?
Han
ley,
199
1IW
PU
s●
●P
aral
epid
onot
us in
dicu
s (K
inbe
rg, 1
856)
IPU
s●
Scal
iset
osus
mjo
berg
i Aug
ener
, 192
2*IA
Hs
●A
ugen
er (1
922)
Suba
dyte
pap
illife
ra (H
orst
, 191
5)T
horm
ora
juke
sii B
aird
, 186
5IP
Uis
●●
Ver
ruca
pelm
a ni
gric
ans
(Hor
st, 1
915)
IWP
Us
●●
Ver
ruca
pelm
a re
tusa
Han
ley
and
Bur
ke, 1
991
WP
Us
●●
Fam
ily:
Sig
alio
nid
ae●
Eup
holo
e ph
ilipp
ensi
s M
cInt
osh,
188
5IA
Us
Eut
hale
ness
a di
gita
ta (M
cInt
osh,
188
5)IP
His
●
Fam
ily:
Sp
hae
rod
orid
ae●
Spha
erod
orid
ae s
p.
Fam
ily:
Syl
lid
ae●
Alc
yono
sylli
s ph
ili G
lasb
y an
d W
atso
n, 2
001
IAE
Z1
●A
utol
ytus
sp.
A*
NA
Us
●H
artm
ann-
Schr
öder
(197
9) B
room
eA
utol
ytus
sp.
B*
●H
artm
ann-
Schr
öder
(197
9) D
erby
Bra
nchi
osyl
lis b
arin
gabo
oree
n Sa
n M
arti
n et
al.,
200
8●
Bra
nchi
osyl
lis e
xilis
(Gra
vier
, 190
0)IP
Us
●B
ranc
hios
yllis
ocu
lata
Ehl
ers,
188
7T
Us
●B
ranc
hios
yllis
orb
iniif
orm
is S
an M
arti
n et
al.,
200
8N
AU
s●
Bra
nia
arti
cula
ta H
artm
ann-
Schr
öder
, 198
2SA
Uis
●B
rani
a cl
avat
a (C
lapa
rèd
e, 1
863)
*A
Hs
●H
artm
ann-
Schr
öder
(197
9) B
room
eB
revi
cirr
osyl
lis a
ncor
i (Sa
n M
arti
n an
d H
utch
ings
, 200
6)●
Ehl
ersi
a sp
. ●
●E
urys
yllis
tube
rcul
ata
Ehl
ers,
186
4C
Us
Exo
gone
afr
ican
a H
artm
ann-
Schr
öder
, 197
4IP
His
●E
xogo
ne b
revi
ante
nnat
a H
artm
ann-
Schr
öder
, 195
9C
Us
●E
xogo
ne d
ispa
r (W
ebst
er, 1
879)
CU
s●
Exo
gone
exm
outh
ensi
s H
artm
ann-
Schr
öder
, 198
0C
Uis
●E
xogo
ne g
ambi
ae L
aner
a et
al.,
199
4C
U●
Exo
gone
nai
dina
Oer
sted
, 184
5*A
Sis●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Exo
gone
het
eros
etoi
des
Har
tman
n-Sc
hröd
er, 1
979*
*●
Exo
gone
lour
ei B
erke
ley
and
Ber
kele
y, 1
938
IPU
●
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 147
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Exo
gona
sim
plex
Har
tman
n-Sc
hröd
er, 1
960*
NA
His
●H
artm
ann-
Schr
öder
(197
9) B
room
eE
xogo
na v
erug
era
afri
cana
Har
tman
n-Sc
hröd
er, 1
976*
●H
artm
ann-
Schr
öder
(197
9) B
room
eH
aplo
sylli
s sp
ongi
cola
(Gru
be, 1
855)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Lang
erha
nsia
bro
omen
sis
Har
tman
n-Sc
hröd
er, 1
979
Odo
ntos
yllis
aus
tral
iens
is H
artm
ann-
Schr
öder
, 197
9**
AU
s●
Odo
ntos
yllis
lang
erha
nsia
eset
osa
Har
tman
n-Sc
hröd
er, 1
979*
*A
Si●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Opi
stho
dont
a m
oren
a L
ange
rhan
s, 1
879
CU
●O
pist
hosy
llis
brun
nea
Lan
gerh
ans,
187
9**
CU
s●
Opi
stho
sylli
s ja
poni
ca Im
ajim
a, 1
966
IWP
Us
●O
pist
hosy
llis
long
icir
rata
Mon
ro, 1
939
IPU
s●
Opi
stho
sylli
s vi
ridi
s L
ange
rhan
s, 1
879
CU
s●
Par
aehl
ersi
a eh
lers
iaef
orm
is (A
ugen
er, 1
913)
AU
s●
Par
asph
aero
sylli
s in
dica
Mon
ro, 1
937
IWP
Us
●P
erki
nsyl
lis a
ugen
eri (
Har
tman
n-Sc
hröd
er, 1
979)
ASis
●Pe
rkin
sylli
s ha
rtm
anns
chro
eder
ae (S
an M
artin
and
Hut
chin
gs, 2
006)
AU
is●
Pio
nosy
llis
auge
neri
Har
tman
n-Sc
hröd
er, 1
979*
●●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pla
kosy
llis
sp.
WA
●P
rosp
haer
osyl
lis n
atha
ni S
an M
arti
n an
d L
opez
, 199
8W
PU
Pro
spha
eros
yllis
pap
illos
issi
ma
(Har
tman
n-Sc
hröd
er, 1
979)
ASi
●P
rosp
haer
osyl
lis s
expa
pilla
ta (H
artm
ann-
Schr
öder
, 197
9)A
Si●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pse
udoe
xogo
ne?
sp.
●●
Rho
palo
sylli
s ha
mul
ifera
Aug
ener
, 191
3A
Us
Salv
ator
ia k
oori
necl
avat
a Sa
n M
arti
n, 2
005
AU
●Sp
haer
osyl
lis c
apen
sis
dens
opap
illat
a H
artm
ann-
Schr
öder
, 197
9*A
Si●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
cen
troa
mer
ican
a H
artm
ann-
Schr
öder
, 195
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
den
sopa
pilla
ta H
artm
ann-
Schr
öder
, 197
9●
Spha
eros
yllis
eri
nace
us C
lapa
rèd
e, 1
863*
IPU
is●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
hir
suta
Ehl
ers,
189
7●
Spha
eros
yllis
hys
trix
Cla
parè
de,
186
4*SA
H/
Si●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
late
ropa
pilla
ta H
artm
ann-
Schr
öder
, 198
6●
Spha
eros
yllis
long
ipap
illat
a H
artm
ann-
Schr
öder
, 197
9*●
●H
artm
ann-
Schr
öder
(197
9) D
erby
Spha
eros
yllis
mul
tipa
pilla
ta H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
opi
stho
cula
ta H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spha
eros
yllis
pap
illos
issi
ma
Har
tman
n-Sc
hröd
er, 1
979*
●H
artm
ann-
Schr
öder
(197
9) B
room
eSp
haer
osyl
lis s
expa
pilla
ta H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Sylli
s gr
acili
s au
stra
liens
is H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
148 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Sylli
s (T
ypos
yllis
) var
iega
ta G
rube
, 186
0*●
Aug
ener
(192
2)Sy
llis
sp.
Typo
sylli
s (L
ange
rhan
sia)
bro
omen
sis
Har
tman
n-Sc
hröd
er, 1
979*
CU
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Typo
sylli
s (L
ange
rhan
sia)
cor
nuta
(Rat
hke,
184
3)*
●H
artm
ann-
Schr
öder
(197
9) B
room
eTr
ypan
osyl
lis z
ebra
(Gru
be, 1
860)
●Ty
posy
llis
(Lan
gerh
ansi
a) s
p.*
AH
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Typo
sylli
s (T
ypos
yllis
) hya
lina
(Gru
be, 1
863)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Typo
sylli
s (T
ypos
yllis
) lut
ea H
artm
ann-
Schr
öder
, 196
0*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Typo
sylli
s (T
ypos
yllis
) cra
ssic
irra
ta T
read
wel
l, 19
25*
●H
artm
ann-
Schr
öder
(197
9) B
room
eTy
posy
llis
(Typ
osyl
lis) r
ayge
orge
i Har
tman
n-Sc
hröd
er, 1
979*
●H
artm
ann-
Schr
öder
(197
9) B
room
eTy
posy
llis (
Typo
sylli
s) fi
liden
tata
gan
thea
umen
sis H
artm
ann-
Schr
öder
, 197
9*W
A●
Typo
sylli
s (T
ypos
yllis
) pha
robr
oom
ensi
s H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Typo
sylli
s (T
ypos
yllis
) gla
ndul
osa
(Aug
ener
, 191
3)*
WA
●H
artm
ann-
Schr
öder
(197
9) B
room
eTy
posy
llis
(Typ
osyl
lis) p
hary
nxci
rcum
fusa
ta H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Xen
osyl
lis s
cabr
oide
s Sa
n M
arti
n et
al.,
200
8W
A●
Fam
ily:
Ow
enii
dae
●O
wen
ia fu
sifo
rmis
Del
le C
hiaj
e, 1
844*
NA
SsH
artm
ann-
Schr
öder
(197
9) B
room
e, A
ugen
er (1
922)
Ow
enia
mir
raw
a Fo
rd a
nd H
utch
ings
, 200
5●
Fam
ily:
Sab
ella
riid
ae●
Idan
thyr
sus
aust
ralie
nsis
(Has
wel
l, 18
83)
AH
is
Idan
thyr
sus
arm
atus
Kin
berg
, 186
7*A
Hs
●H
artm
ann-
Schr
öder
(197
9) B
room
eLy
gdam
is s
p.
●
Fam
ily:
Sab
elli
dae
●A
mph
igle
na le
nae
Cap
a an
d R
ouse
, 200
7W
AH
i
Bis
pira
man
icat
a (G
rube
, 187
8)W
PH
/Ss
●●
Bis
pira
por
ifera
(Gru
be, 1
878)
WP
H/
Ss●
Bis
pira
ser
rata
Cap
a, 2
008
NA
Si●
Bra
nchi
omm
a ni
grom
acul
ata
(Bai
rd, 1
865)
*A
His
●H
artm
ann-
Schr
öder
(197
9) B
room
eB
ranc
hiom
ma
spp.
●
Cho
ne a
ustr
alie
nsis
Har
tman
n-Sc
hröd
er, 1
979*
*W
ASs
●H
artm
ann-
Schr
öder
(197
9) B
room
eM
egal
omm
a in
terr
upta
Cap
a an
d M
urra
y, 2
009
NA
Hs
●N
otau
lax
spp.
A
Hi
●●
Per
kins
iana
ano
dina
Cap
a, 2
007
WA
Hs
Pot
amill
a ne
glec
ta (S
ars,
185
1)*
NA
Hs
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
seud
opot
amill
a sp
. 2 (s
ensu
Cap
a, 2
007)
●
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 149
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Sabe
llast
arte
indi
ca (S
avig
ny, 1
820)
*N
AH
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Sabe
llast
arte
sp.
nov
. 2 (s
ensu
Cap
a et
. al.
2010
)●
Styl
omm
a ju
ani C
apa,
200
8W
AU
●St
ylom
ma
palm
atum
(Qua
tref
ages
, 186
6)C
PH
s●
Fam
ily:
Ser
pu
lid
ae●
?Ser
pula
sp.
Pill
ai, 2
009
Hyd
roid
es a
dam
afor
mis
Pill
ai, 2
009
WA
Hs
●H
ydro
ides
alb
icep
s (E
hren
berg
and
Gru
be, 1
869)
IWP
His
●H
ydro
ides
exa
ltat
us (M
aren
zelle
r, 18
84)*
*IW
PH
is●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Hyd
roid
es e
xter
nisp
ina
Stra
ugha
n, 1
967
NA
His
●●
Hyd
roid
es k
imbe
rley
ensi
s Pi
llai,
2009
WA
Hs
Hyd
roid
es m
alle
olas
pina
Str
augh
an, 1
967
NA
His
●H
ydro
ides
min
ax (G
rube
, 187
8)IW
PH
is●
Hyd
roid
es p
seud
exal
tatu
s Pi
llai,
2009
WA
Hi
●H
ydro
ides
rec
ta S
trau
ghan
, 196
7N
AH
i●
Hyd
roid
es s
impl
iden
tatu
s Pi
llai,
2009
WA
Hs
●H
ydro
ides
spi
culit
ubus
Pill
ai, 2
009
WA
Hs
●H
ydro
ides
spi
ratu
bus
Pilla
i, 20
09W
AH
i●
Hyd
roid
es tu
berc
ulat
us Im
ajim
a, 1
976
WA
Hs
●K
imbe
rley
a hu
tchi
ngsa
e Pi
llai,
2009
WA
Hs
●P
omat
olei
os k
raus
sii (
Bai
rd, 1
865)
**IP
His/
E●
Pom
atos
tegu
s ac
tino
cero
s (M
örch
, 186
3)T
His
●●
Pro
tula
bis
pira
lis (S
avig
ny, 1
820)
IWP
Hs
●P
rotu
la p
allia
ta (W
illey
, 190
5)IW
PH
s●
?Pro
tula
pal
liata
(Will
ey, 1
905)
*W
AH
s●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pse
udop
rotu
la k
imbe
rley
ensi
s Pi
llai,
2009
●Sa
lmac
ina
sp.
●●
Serp
ula
ampl
iloba
ta P
illai
, 200
9W
AH
s●
Serp
ula
hart
man
ae R
eish
, 196
8IW
PH
is●
●Se
rpul
a ju
kesi
i Bai
rd, 1
865
IAH
is
Serp
ula
nudi
radi
ata
Pilla
i, 20
09W
AH
is●
●Se
rpul
a va
sife
ra H
asw
ell,
1884
AH
is●
Serp
ula
verm
icul
aris
Lin
ne, 1
767*
IWP
His
●●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Serp
ula
wat
soni
Will
ey, 1
905
●Sp
iras
erpu
la s
nelli
i Pill
ai a
nd te
n H
ove,
199
4IA
His
●Sp
irob
ranc
hus
baile
ybro
ckae
Pill
ai, 2
009
WA
Hi
●
150 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Spir
obra
nchu
s co
rnic
ulat
us (G
rube
, 186
2)IW
PH
is●
●Sp
irob
ranc
hus
poly
trem
a (P
hilip
pi, 1
844)
CH
is●
Spir
obra
nchu
s ri
char
dsm
ithi
Pill
ai, 2
009
IAH
s●
Spir
obra
nchu
s te
trac
eros
(Sch
mar
da,
186
1)T
Hs
●V
erm
iliop
sis
cylin
dric
a Pi
llai,
2009
WA
His
●V
erm
iliop
sis
glan
dige
ra G
ravi
er, 1
908
IPH
s●
Fam
ily:
Cap
itel
lid
ae●
Das
ybra
nchu
s ca
ducu
s (G
rube
, 184
6)*
WA
SiH
artm
ann-
Schr
öder
(197
9) B
room
eH
eter
omas
tus
cf. fi
lifor
mis
Cla
parè
de,
186
4*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Not
omas
tus
(Clis
tom
astu
s) b
room
ensi
s H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Not
omas
tus
broo
men
sis
Har
tman
n-Sc
hröd
er, 1
979
●Sc
ypho
proc
tus
orna
tus
Har
tman
n-Sc
hröd
er, 1
979*
●H
artm
ann-
Schr
öder
(197
9) B
room
e
Fam
ily:
Mal
dan
idae
●M
ald
anid
ae s
pp.
●
Fam
ily:
Op
hel
iid
ae●
Arm
andi
a in
term
edia
Fau
vel,
1901
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Arm
andi
a la
nceo
lata
Will
ey, 1
905*
●A
ugen
er (1
922)
Arm
andi
a br
oom
ensi
s H
artm
ann-
Schr
öder
, 197
9*H
artm
ann-
Schr
öder
(197
9) B
room
eO
phel
ina
acum
inat
a H
artm
ann-
Schr
öder
, 197
9●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pol
yoph
thal
mus
pic
tus
(Duj
ard
in, 1
839)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Oph
eliid
ae s
pp.
●
Fam
ily:
Orb
inii
dae
●N
aine
ris
laev
igat
a (G
rube
, 184
0)*
●H
artm
ann-
Schr
öder
(197
9) B
room
eSc
olop
los
(Sco
lopl
os) d
iffici
lis D
ay, 1
977*
●H
artm
ann-
Schr
öder
(197
9) B
room
eO
rbin
iidae
spp
. ●
Fam
ily:
Par
aon
idae
●Pa
raon
idae
sp.
Fam
ily:
Qu
esti
dae
●Q
uest
idae
sp.
●
Fam
ily:
Sca
lib
regm
atid
aeH
ybos
cole
x ve
rruc
osa
Har
tman
n-Sc
hröd
er, 1
979*
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Scal
ibre
gmat
idae
sp.
●
Fam
ily:
Ch
aeto
pte
rid
ae●
Cha
etop
teri
dae
spp
. ●
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 151
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Phy
loch
aeto
pter
us c
f. vi
trar
ius
(Ehl
ers,
190
8)*
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
hylo
chae
topt
erus
sp.
*H
artm
ann-
Schr
öder
(197
9) B
room
e
Fam
ily:
Mag
elon
idae
Mag
elon
idae
sp.
Fam
ily:
Poe
cilo
chae
tid
ae●
Poec
iloch
aeti
dae
sp.
●
Fam
ily:
Sp
ion
idae
Dis
pio
unci
nata
Har
tman
, 195
1*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pol
ydor
a (P
olyd
ora)
arm
ata
Lan
gerh
ans,
188
1*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pol
ydor
a (P
olyd
ora)
cili
ata
(Joh
nsto
n, 1
838)
*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Pri
onos
pio
(Pri
onos
pio)
falla
x Sö
der
strö
m, 1
920*
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
rion
ospi
o (P
rion
ospi
o) la
nceo
lata
Har
tman
n-Sc
hröd
er, 1
979*
●H
artm
ann-
Schr
öder
(197
9) B
room
eSc
olel
epis
(Sco
lele
pis)
bal
ihai
ensi
s H
artm
ann-
Schr
öder
, 197
9*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Spio
nid
ae s
pp.
●
Fam
ily:
Am
ph
aret
idae
●A
mph
aret
idae
sp.
A
mph
icte
is p
hilip
pina
rum
Gru
be, 1
878*
●A
ugen
er (1
922)
Isol
da c
f. pu
lche
lla M
ülle
r, 18
58*
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Fam
ily:
Cir
ratu
lid
ae●
?Cir
rifo
rmia
sp.
*H
artm
ann-
Schr
öder
(197
9) B
room
eD
odec
acer
ia s
p.
●
Fam
ily:
Fla
bel
lige
rid
ae●
Flab
ellig
erid
ae s
pp.
●
Fam
ily:
Pec
tin
arii
dae
●A
mph
icte
ne fa
vona
Hut
chin
gs a
nd P
eart
, 200
2A
Si /H
s●
Pec
tina
ria
(Pec
tina
ria)
cf.
anti
poda
Sch
mar
da,
186
1*●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Fam
ily:
Ter
ebel
lid
ae●
Am
phit
riti
des
ithy
a H
utch
ings
and
Gla
sby,
198
8A
Ss
Eup
olym
nia
koor
angi
a H
utch
ings
and
Gla
sby,
198
8A
Ss●
●E
upol
ymni
a cf
. neb
ulos
a (M
onta
gu, 1
818)
*N
ASs
●H
artm
ann-
Schr
öder
(197
9) B
room
eE
uthe
lepu
s m
arch
inba
r H
utch
ings
, 199
7●
Eut
hele
pus
serr
atus
Hut
chin
gs a
nd G
lasb
y, 1
986
ASs
●H
auch
iella
sp.
●
Lana
ssa
sp.
●
152 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Taxa
Biog
eogr
aphi
c
code
Habi
tat
code
Insh
ore
Offs
hore
Publ
icat
ions
and
loca
tion
Lani
ce b
idew
a H
utch
ings
and
Gla
sby,
198
8A
Ss●
Lani
ce c
onch
ilega
(Pal
las,
176
6)*
NA
Ss●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Lani
ce s
inat
a H
utch
ings
and
Gla
sby,
199
0●
Lani
cola
loba
ta H
artm
ann-
Schr
öder
, 198
6A
Sis●
Loim
ia in
gens
(Gru
be, 1
878)
IASs
●●
Loim
ia m
edus
a (S
avig
ny, 1
817)
*N
ASs
●H
artm
ann-
Schr
öder
(197
9) B
room
e, A
ugen
er (1
922)
Loim
ia o
chra
cea
(Gru
be, 1
878)
●Ly
silla
sp.
●
Nic
olea
am
nis
Hut
chin
gs a
nd M
urra
y, 1
984
ASs
●N
icol
ea v
enus
tula
(Mon
tagu
, 181
8)*
ASs
●●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome
Phi
sidi
a ec
huca
Hut
chin
gs a
nd G
lasb
y, 1
988
●P
ista
ant
hela
Hut
chin
gs a
nd G
lasb
y, 1
990
NA
Ss
Pis
ta a
ustr
alis
Hut
chin
gs a
nd G
lasb
y, 1
988
ASs
●P
ista
cur
tiun
cata
Har
tman
n-Sc
hröd
er, 1
981
ASs
●P
ista
kim
berl
iens
is H
utch
ings
and
Gla
sby,
199
0N
ASs
●●
Pis
ta v
iola
cea
Har
tman
n-Sc
hröd
er, 1
984
ASs
●P
olyc
irru
s br
oom
ensi
s H
artm
ann-
Schr
öder
, 197
9*●
●H
artm
ann-
Schr
öder
(197
9) B
room
eP
olyc
irru
s sp
. ●
Pse
udop
rocl
ea a
ustr
alis
Hut
chin
gs a
nd G
lasb
y, 1
990
ASs
●St
rebl
osom
a ac
ymat
um H
utch
ings
and
Rai
ner,
1979
ASs
●St
rebl
osom
a dy
tiko
s H
utch
ings
and
Gla
sby,
199
0N
ASs
●Te
rebe
lla m
acul
ata
Hut
chin
gs a
nd G
lasb
y, 1
988
NA
Ss●
Tere
bella
pap
pus
Hut
chin
gs a
nd M
urra
y, 1
984
ASs
●Te
rebe
lla p
tero
chae
ta S
chm
ard
a, 1
861*
ASis
●H
artm
ann-
Schr
öder
(197
9) B
room
eTe
rebe
lla ta
ntab
iddy
cree
kens
is H
artm
ann-
Schr
öder
, 198
0●
The
lepu
s ex
tens
us H
utch
ings
and
Gla
sby,
198
7A
Ss●
The
lepu
s pl
agio
stom
a (S
chm
ard
a, 1
861)
*IA
Ss●
Har
tman
n-Sc
hröd
er (1
979)
Bro
ome,
Aug
ener
(192
2)T
hele
pus
robu
stus
(Gru
be, 1
878)
●T
hele
pus
thor
acic
us (G
rube
, 187
0)*
●A
ugen
er (1
922)
Fam
ily:
Tri
chob
ran
chid
aeTe
rebe
llide
s w
oola
wa
Hut
chin
gs a
nd P
eart
, 200
0A
Ss●
Fam
ily:
Ste
rnas
pid
aeSt
erna
spis
scu
ata
(Ran
zani
, 181
7)*
●H
artm
ann-
Schr
öder
(197
9) B
room
e
Fam
ily:
Sac
coci
rrid
aeSa
ccoc
irru
s sp
.*
●H
artm
ann-
Schr
öder
(197
9) B
room
e
*
His
toric
al r
ecor
ds n
ot d
epos
ited
in A
ustr
alia
n m
useu
m c
olle
ctio
ns.
**
His
toric
al r
ecor
ds b
ut a
lso
pres
ent
in A
ustr
alia
n m
useu
m c
olle
ctio
ns.
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 153
TABL
E 4
Spe
cies
ric
hnes
s an
d nu
mbe
r of
col
lect
ing
even
ts a
t ea
ch lo
catio
n.
Loca
tion
Colle
ctin
g
year
rang
e
Species richness
No. coll. events
No. familiesAcoetidaeAmpharetidaeAmphinomidaeAphroditidaeCapitellidaeChaetopteridaeChrysopetalidaeCirratulidaeDorvilleidaeEunicidaeEuphrosinidaeFlabelligeridaeGlyceridaeGoniadidaeHesionidaeLumbrineridaeMagelonidaeMaldanidaeMyzostomidaeNephtyidaeNereididaeOenonidaeOnuphidaeOpheliidaeOrbiniidaeOweniidaeParaonidaePectinariidaePhyllodocidaePilargidaePoecilochaetidaePolynoidaeQuestidaeSabellariidaeSabellidaeScalibregmatidaeSerpulidaeSigalionidaeSphaerodoridaeSpionidaeSyllidaeTerebellidaeTrichobranchidae
Ad
mir
al Is
land
1994
11
1●
Ad
mir
alty
Gul
f19
782
12
●●
Alb
ert I
slan
ds
1988
101
4●
●●
●
Ash
mor
e R
eefO
A19
83–2
002
478
17●
●●
●●
●●
●●●
●●
●●
●●●
Bea
gle
Ree
f19
919
15
●●
●●
●
Bed
ford
Isla
nd19
941
11
●
Ber
noui
lli Is
land
1988
201
5●
●●
●●
Bro
ome
1932
–198
780
929
●●
●●
●●●
●●●
●●
●●●●●●●●
●●●●●
●●●●
Buf
fon
Isla
nd19
8819
15
●●
●●
●
Caf
fare
lli Is
land
1994
21
1●
Cap
e L
eveq
ue19
72–1
987
103
7●
●●
●●
●●
Cap
e V
illar
et19
291
11
●
Car
eeni
ng B
ay19
8810
15
●●
●●
●
Car
tier
Isla
ndO
A19
9234
112
●●
●●
●●
●●●
●●
●
Cas
sini
Isla
nd19
88–1
991
223
8●
●●
●●
●●●
Chu
rchi
ll R
eef
1991
31
2●
●
Cle
rke
Ree
fOA
1982
–198
34
23
●●
●
Coc
kato
o Is
land
1991
71
5●
●●
●●
Con
dill
ac Is
land
1988
–199
116
25
●●
●●
●
Cor
neill
e Is
land
1988
91
5●
●●
●●
Cor
onat
ion
Isla
nd19
887
13
●●
●
Der
by19
754
23
●●
●
154 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
Loca
tion
Colle
ctin
g
year
rang
e
Species richness
No. coll. events
No. familiesAcoetidaeAmpharetidaeAmphinomidaeAphroditidaeCapitellidaeChaetopteridaeChrysopetalidaeCirratulidaeDorvilleidaeEunicidaeEuphrosinidaeFlabelligeridaeGlyceridaeGoniadidaeHesionidaeLumbrineridaeMagelonidaeMaldanidaeMyzostomidaeNephtyidaeNereididaeOenonidaeOnuphidaeOpheliidaeOrbiniidaeOweniidaeParaonidaePectinariidaePhyllodocidaePilargidaePoecilochaetidaePolynoidaeQuestidaeSabellariidaeSabellidaeScalibregmatidaeSerpulidaeSigalionidaeSphaerodoridaeSpionidaeSyllidaeTerebellidaeTrichobranchidae
Des
cart
es Is
land
1988
71
4●
●●●
Ent
ranc
e Is
land
1988
31
2●
●
Fene
lon
Isla
nd19
8824
27
●●
●●
●●●
Geo
rge
Wat
er19
901
11
●
Gre
gory
Isla
nd19
946
12
●●
Hal
e Is
land
1991
21
2●
●
Her
itag
e R
eef
1991
21
2●
●
Hey
woo
d Is
land
1988
11
1●
Hib
erni
a R
eefO
A19
9211
16
●●
●●●
●
Hun
ter
Riv
er19
881
11
●
Irvi
ne Is
land
1988
–199
412
23
●●
●
Jack
son
Isla
nd19
886
12
●●
Jone
s Is
land
1991
121
7●
●●
●●
●●
Jung
ulu
Isla
nd19
886
14
●●
●●
Kin
g G
eorg
e R
iver
1991
41
1●
Kur
i Bay
1991
51
4●
●●
●
Lac
eped
e Is
land
s19
82–1
991
72
5●
●●
●●
Laf
onta
ine
Isla
nd19
8823
16
●●
●●
●●
Leo
nie
Isla
nd19
944
12
●●
Les
ueur
Isla
nd19
916
13
●●
●
Lon
g R
eef
1980
–199
126
38
●●
●●
●●
●●
Lor
d Is
land
1991
–199
415
27
●●
●●
●●
●
Luc
as Is
land
1988
511
9●
●●
●●
●●
●●
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 155
Loca
tion
Colle
ctin
g
year
rang
e
Species richness
No. coll. events
No. familiesAcoetidaeAmpharetidaeAmphinomidaeAphroditidaeCapitellidaeChaetopteridaeChrysopetalidaeCirratulidaeDorvilleidaeEunicidaeEuphrosinidaeFlabelligeridaeGlyceridaeGoniadidaeHesionidaeLumbrineridaeMagelonidaeMaldanidaeMyzostomidaeNephtyidaeNereididaeOenonidaeOnuphidaeOpheliidaeOrbiniidaeOweniidaeParaonidaePectinariidaePhyllodocidaePilargidaePoecilochaetidaePolynoidaeQuestidaeSabellariidaeSabellidaeScalibregmatidaeSerpulidaeSigalionidaeSphaerodoridaeSpionidaeSyllidaeTerebellidaeTrichobranchidae
Mar
et Is
land
s19
49–1
988
52
3●
●●
Mer
mai
d R
eefO
A19
821
11
●
Mis
sion
Bay
1991
71
4●
●●
●
Mon
taliv
et Is
land
s19
88–1
991
362
7●
●●
●●
●●
Mon
tgom
ery
Ree
f19
89–1
994
52
3●
●●
Nap
ier
Bro
ome
Bay
1986
–199
111
24
●●
●●
Parr
y H
arbo
ur19
911
11●
Port
Geo
rge
IV19
8811
14
●●
●●
Port
War
rend
er19
762
11
●
Prud
hoe
Isla
nds
1988
61
2●
●
Quo
ndon
g Po
int
1975
11
1●
Rob
roy
Ree
fs19
88–1
991
72
4●
●●
●
Scor
pion
Isla
nd19
9111
14
●●
●●
Scot
t Ree
fOA
1984
–190
86
23
●●
●
Shir
ley
Isla
nd19
8812
14
●●
●●
Slat
e Is
land
s19
881
11
●
Sund
ay Is
land
1991
–199
418
28
●●
●●
●●
●●
Tallo
n Is
land
1994
11
1●
Tran
quil
Bay
1991
11
1●
Van
sitt
art B
ay19
91–
2006
32
2●
●
Wai
lgw
in Is
land
1988
121
5●
●●
●●
Wild
cat R
eefs
1988
21
2●
●
Will
ie C
reek
1975
–198
45
23
●●
●
Yank
awin
garr
i Isl
and
1991
11
1●
156 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
TABLE 5 Species richness and number of collecting events at each location.
Biogeographic Code Inshore Offshore- 30 27
A 40 7
WA 20 2
NA 21 3
SA 6
C 15 3
CP 1
IA 20 5
IP 25 10
IWP 37 20
T 7 3
WP 6 4
Total 228 84
TABLE 6 Number of species with each habitat code.
Habitat Code Inshore Offshore- 30 27
E 1
EZ 8 7
Hi 5 1
Hi/Si 3
His 20 6
His/E 1
Hs 38 10
Hs/Si 1 1
Hs/Ss 2 1
Si 10
Sis 5
Ss 26 4
U 6
Uis 18 4
Us 53 23
Us/E 1
Total 228 84
Inshore sites (intertidal and shallow subtidal and hard substrate habitats) typically have between 1 and 20 species present, with a few sites having much higher numbers. The offshore sites appear to have higher numbers, but we stress that only three offshore sites have been sampled for polychaetes. At all sites only limited collecting of particular habitats was undertaken and it is premature to discuss patterns of species richness within all the polychaetes. Only ten of the sampled sites have more than 20 species recorded, which is undoubtedly a gross underestimate of the diversity of polychaetes.
For a description of richness and distribution patterns, we suggest it is more informative to look at individual polychaete families, for example the terebellids, but again sampling was limited to the inshore regions (Hutchings and Glasby 1990). Of all the polychaete families for which we have morphospecies data, the Polynoidae currently yield the most reliable information on species richness and distribution patterns. This is because the collecting effort for this family has been greater with more sites sampled than for any other family, as Hanley collected at both inshore (1991, 1994) and offshore (1987, 1992) sites, concentrating almost exclusively on polynoids, and all material was identified. The limited conclusions, which that can be drawn from the data highlight the need for additional sampling, increasing the number of locations and habitats as individual polychaete families or species tend to have specific habitat requirements.
Species richness patterns across locations for the most part reflect the variable collection effort and cannot be interpreted as an accurate measure of species richness at any one location, especially when it is the result of a single collecting event. However, the presentation of the data in this way allows an assessment of which locations in the Kimberley have been surveyed for polychaetes, and provide an indication of the available data. Current integrative studies, incorporating genetic data with the morphological information, are demonstrating that some of the species previously considered widespread are in fact a complex of sibling species, increasing the diversity of known polychaete fauna in Australia and the Kimberley (see for example Capa et al. 2010).
The dominance of widespread species and conversely the lack of narrow range endemics in the Kimberley region reflects, in part, the collection and identification biases discussed above. Additional sampling will be required to confirm whether taxa recorded from regions contiguous with the Kimberley (i.e. southern Western Australia and Northern Territory) are also present in the Kimberley. Examples to date include the sabellariid
KIMBERLEY MARINE BIOTA. HISTORICAL DATA: POLYCHAETES 157
FIGURE 4 A MDS plot of the distribution of polynoids within the region.
Idanthyrsus willora (Hutchings et al. 2012), and sabellids belonging to the genera Amphicorina, Euchone and Pseudobranchiomma amongst others (Capa personal observation). Considering just the 24 terebellids identified to species, 15 are widespread Australian species, seven are restricted to Northern Australia and two are Indo-Australian. Of the 26 Nereididae species with distributional data, five are widespread Australian species, none appear to be restricted to northern Australia and 21 are widespread in the Indo-Pacific. A few nereidid species, Neanthes pachychaeta (Fauvel), Perinereis suluana (Horst) and Perinereis nuntia Savigny in Lamarck, appear to be common in the Kimberley and Indo-Australian region, but are rarely, if ever, encountered in northern Australia (Glasby personal observation).
The discussion above partly ref lects the specialist interests of Russell Hanley, former curator of polychaetes at MAGNT who was particularly interested in commensal species, rather than the abundance of this family across the Kimberley region [in coral reef areas polynoids are infrequently collected (Hutchings personal observation)]. In contrast, Hutchings (1989) made extensive collections of all polychaetes, especially those associated with dead coral substrates, and targeted intertidal collecting was also done when feasible given tidal and safety issues, and subtidal soft sediments were not collected. Also, facilities
for sorting in the field meant that small individuals were often overlooked. However, it should be noted that no comprehensive sampling of all habitats present in the Project Area has been undertaken to date, and this attempt to synthesise the existing data is to facilitate more comprehensive surveys of the area, which are expected, given the recognised limitations in previous sampling, to reveal a very diverse and abundant polychaete fauna.
REGIONAL BIOGEOGRAPHIC COMPARISON
At this stage it is premature to discuss the regional biogeography for the polychaetes as sampling across locations has been too variable.
Some widespread polychaetes are being shown to represent suites of cryptic species (Carlton 2009; Capa et al. 2013). For example, some current work on nereidids has shown that using both morphological and molecular data to redefine species boundaries considerably reduces the distributional ranges (Glasby et al. 2013). We suspect many of the species listed in Table 3 will be found to represent suites of species, in which case diversity estimates for the region will increase. In addition, some of the names of so-called widespread species in Table 3 will almost certainly change as revisions of these species occurs and it is likely some will be found to represent undescribed species.
158 P. HUTCHINGS, C. GLASBY, M. CAPA AND A. SAMPEY
FUTURE DIRECTIONS
As revisions of individual famil ies are undertaken the specimens in historical collections need to be re-examined and their identifications confirmed. Additional habitats need to be sampled, especially soft sediments and those from deeper water, together with environmental information such as sediment characteristics.
While polychaete collections were historically preserved in formalin, contemporary collection practice now routinely preserves material in ethanol to enable subsequent genetic studies, which are changing our taxonomic concepts – see for example Capa et al. (2010, 2013) and Wei et al. (2013). Such studies are expected to continue and the contemporary surveys underway (2009–2014) in the Project Area are undertaking such practices.
A current focus of some conservation research has been utilisation of museum data in large scale biogeographic analyses. To date the Kimberley species data have been available for use in only a very general manner, with most information on assemblages in different areas, or bioregional comparisons being derived from some well known taxa such as fishes. Polychaetes could be a useful taxon to include, but as already identified from our data, the sampling and taxonomic biases limit their value. However, it would be possible to focus on certain families, which have been both elucidated and are also well surveyed in other areas such as terebellids and nereidids. Currently C. Glasby and P. Hutchings are collaborating on such a project to utilise these families in identifying biogeographic regions and comparing them with those already identified using sediment and demersal fish data.
Finally, this historical review of the current status of polychaete knowledge in the Kimberley region can be used to identify where additional survey work is necessary to obtain a comprehensive picture of polychaete distributions in the region.
ACKNOWLEDGEMENTS
We thank our taxonomic colleagues whose substantial collections and identifications of the Kimberley polychaete fauna formed the basis of this dataset. The unpublished data of Russell Hanley on the polynoids that are housed in MAGNT have been particularly helpful. We also thank the Collection Managers in each of the museums for supplying the databases. We express our thanks to Stacey Osborne and Albert Miles for databasing the unregistered Kimberley material in WAM collections and for checking taxonomic and spatial information in this dataset and to Stacy Osborne for assistance in formatting tables for this paper. Also thanks to the reviewers who provided helpful comments.
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MANUSCRIPT RECEIVED 23 DECEMBER 2013; ACCEPTED 12 SEPTEMBER 2014.