RECORDS OF THE WESTERN 85 117–183 (2020) DOI: 10.18195/issn.0313-122x.85.2020.117-183 SUPPLEMENT

Smaller molluscs from a multi-taxon survey (2012–2014) of the shallow marine environments of the tropical Kimberley region,

Peter Middelfart*, Lisa Kirkendale and Clay Bryce

Aquatic Zoology Department, Western Australia Museum, Locked Bag 49, Welshpool DC, Western Australia, Australia * Corresponding author: [email protected]

ABSTRACT – The marine environments of Western Australia’s Kimberley region (coast to continental edge) are rich in spatial diversity and vast in scale affording a broad range of habitats for marine fauna and fora. The macromolluscan (here defned as adult molluscs >10 mm) diversity of this large marine realm has been the subject of numerous surveys over the last 50 years. However, the smaller molluscs (here defned as adult molluscs <10 mm ) have, for the most part, been ignored due to challenges in the collection and study of this assemblage. To begin to redress this imbalance a pilot project specifcally designed to target smaller molluscs was undertaken over the fnal three years (2012–2014) of the survey component of the Woodside Collection Project (Kimberley) 2009–2014. Seventy-seven stations (of 179 surveyed) were sampled yielding 1,883 identifed specimen lots representing 505 micromollusc species from ≤5 mm adult size class (following Geiger et al. 2007), 43 species from >5–10 mm adult size class, 68 species from >10 mm adult size class, which are likely to be juveniles of macromolluscs, as well as 8 species whose adult size could not be determined. This diversity spans 102 family or superfamily level taxa. Twenty-one of these familial level taxa were new to the survey area (see Bryce et al. 2018 for a full description of the Kimberley survey area) and the number of taxa recorded from a further 18 families was doubled. A considerable degree of species rarity characterised this fauna with almost 52% of all species being singletons. Less than 22% of taxa were recorded from three or more stations. A total of at least 26 new species, not yet formally described, as well as an additional 23 new records, for the survey area and/or Australia were discovered, yielding 49 new species-level records. Two formal taxonomic revisions are undertaken in this paper due to the acquisition of new material and a review of type material. Firstly, Haplocochlias minutissumus Pilsbry, 1921 is synonymised under parvissima Hedley, 1899 [as Lophocochlias parvissimus (Hedley, 1899)]. Secondly, the synonymy of Condylocuna io (Bernard, 1915) and Condylocuna tricosa Middelfart, 2002, is revoked. Many putative species require further study to determine their taxonomic status; several are probably new to science.

KEYWORDS: , Indian Ocean, Kimberley survey area, smaller molluscs, micromolluscs, north- west Australia, Woodside Collection Project

INTRODUCTION Wells and Allen 2005; Wilson 2014; Kirkendale et al. 2019), baseline marine research is still necessary The Kimberley marine region is a remote part of to generate core information suitable for the northern Australia with a high degree of habitat complexity. It is vast in area and it extends from the management of the region’s conservation, tourist coast to the edge of the continental shelf. The coast and industrial potential (Kirkendale et al. 2019). is characterised by large river systems, expansive One of the largest projects to document marine estuaries, and coral, rock, sand and mud habitats. biodiversity of the region, indeed any region The wide continental shelf extends offshore to a within Australia, was The Woodside Collection Project series of mid-shelf reefs and shelf-edge atolls with (Kimberley) 2008–2015 (see Sampey et al. (2014) for well-developed coral reefs adjacent to deep-sea a full description of the Project Area and historical basins (Sampey et al. 2014). While some studies collections and Bryce et al. (2018) for survey area have been undertaken of the region’s biota (see and methodology information). 118 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

With 179 stations surveyed (Bryce et al. 2018), Collecting smaller molluscs is logistically the documentation of the molluscs fauna from the complicated due to their size (Middelfart et al. Kimberley survey area has been separated into three 2016), but scrubbing rocks and rock walls, the parts. The published historical dataset documenting use of suction pumps, elutriation of sea water the museum-archived molluscan diversity for the following shaking and scrubbing of seaweed region was completed in 2015 (Willan et al. 2015), and rocks, and sieving sediments are well- which reported on 1,784 species. A publication established methods (Geiger et al. 2007). Visual documenting macromollusc (>10 mm when adult) detection is an additional challenge. While many results is in preparation (Kirkendale et al. in prep.) smaller molluscs range upward from 3 mm in and this present paper focuses on the outcomes length and are just observable with the naked of a pilot project specifcally designed to capture eye, many are only a few hundred micrometres and assess smaller molluscan (<10 mm when adult) long and therefore require the use of high- diversity in the region. powered microscopes to observe and illustrate Smaller molluscs (i.e., molluscs <10 mm the most delicate shell features on these tiny in greatest shell dimension) constitute a key specimens, Consequently, Scanning Electron element in marine ecosystems and often occur Microscopy (SEM) is the only option. Image in abundance and great diversity (Albano et al. stacking methodologies and high-resolution 2011; Middelfart et al. 2016). Their lifestyles are digital cameras are also important tools that can varied and include grazing assemblages (e.g. produce specimen images in <1 mm category Rissooidea), a predatory component (e.g. ), (Geiger et al. 2007). parasites (e.g. ) and commensals (e.g. Galeommatoidea) (Beesley et al. 1998; Morton 1988). Early taxonomic works on smaller molluscs from In soft boreal sediments, microbivalves (e.g. Mysella) the tropical coast of northern Australia includes and microgastropods (e.g. ) may exhibit those by Hedley (1918); Laseron (1956a,b, 1958a,b, densities numbering thousands per square metre 1959) and Iredale and Laseron (1957). Many of (Muus 1967; Ockelmann and Muus 1978; Middelfart these studies included micromolluscs collected pers. obs.). By comparison, tropical reefs systems from throughout Australia (Ponder and Yoo 1977, tend to host high diversity and low abundance 1978; Ponder 1983, 1985, 1990, 1999; Ponder and across most microgastropod taxa (Bouchet et de Keyzer 1992; Middelfart 2002a,b). Interest in al. 2002; Barnes 2019). Smaller molluscs are an the Australian temperate coast was fuelled by important food source for benthic predators and are the numerous endemic taxa discovered there important grazers and bioturbators of sediments (Marshall 1983). The publications listed by Wells (Mann 1982). and Allen (2005) and by Willan (2005, 2015), which A lack of elementary identifcation resources has, include specimens collected in the Kimberley in part, resulted in the exclusion of smaller molluscs Project Area (Sampey et al. 2014 for project area from marine faunal surveys undertaken in Western boundaries), include some smaller molluscs. Australia (WA), and to some extent nationally and Important international publications are the globally. Moreover, identifcation of smaller molluscs revision of the (Geiger 2012) and requires a large time commitment to access the Marine Mollusks in Japan by Okutani (2017). scattered literature contained in library databases. Recent studies have revealed that smaller While numerous published works are available to molluscs represent a substantial component of identify Australian macromolluscs, the literature on worldwide diversity and, once integrated into smaller molluscs is patchy and, in many instances, inventories, increase biodiversity estimates by outdated. Often the original publication, dating back orders of magnitude (Bouchet et al. 2002; Albano et several decades, is the only information available. Access to older literature has been improved al. 2011; Middelfart et al. 2016). This component of with the creation and public availability of the biodiversity has been inadequately characterised, Biodiversity Heritage Library. However, illustrations but it is likely to be very species rich in regions and detailed descriptions of microscopic species of complex spatial and temporal heterogeneity, are often poor, missing or incomplete leading to such as the marine bioregions of the Kimberley identifcation uncertainty. A number of revisions (Wilson 2014). have been published, which clarify the limitations of earlier works and enable identifcation of some AIMS groups (Ponder and Yoo 1981; Marshall 1983, 1988; Ponder 1985, 1990; Dijkstra 1991; Middelfart 2002a, b; The aim of this study was to test semi- Renard and Bouchet 2003; Cossignani 2006; Dijkstra quantitative field collection methods to assess and Maestrati 2008; Geiger 2012; Pizzini et al. 2013; diversity of smaller molluscs in the Kimberley Warén 2013). Project Area. The approach was threefold: SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 119

1. To sample habitats harbouring potential smaller tape would identify a rock or rubble site. A visually molluscs from 77 stations in the survey area. estimated 10 litre volume of rocks or rubble was then 2. To produce a smaller-mollusc inventory for the scrubbed with a laundry brush into the collecting region. bags. On deck, the brushed sediment from each bag 3. To report on morphospecies diversity. was combined and run through a series of sieves to 250 μm mesh size. Any macromolluscs (Ma) over 10 mm were removed by hand. The post-sieved METHODS sediment was preserved in 100% ethanol for further The sampling program to collect smaller molluscs processing at the Western Australian Museum was undertaken from 2012–2014 as an adjunct to (WAM). a macromollusc survey for the Woodside Collection In the laboratory at the WAM, samples from Project (Kimberley) 2008–2015, during which on-site 77 stations were processed using two or more surveys were carried out from 2009–2014 (Bryce techniques (Table 1) for smaller molluscs (live et al. 2018). During the earlier 2009–2011 survey and dead collected specimens). Each subsample period, sediments were retained as a by-product was spread thinly in a 120 mm petri dish and all of sampling other taxa, e.g. during rock breaking molluscs (i.e. M, Ma and Me size categories) were for and small . These sorted using forceps and fne brushes. Because of the sediments were sorted as part of the smaller- potential scale of the project, the volume subsampled mollusc study to extend the sampling to other from each station was calibrated until a workable habitats, as well as to stations not covered in 2013– smaller volume was achieved (Table 1). 2014. The surveyed stations chosen in 2013–2014 had suitable habitat for rock scrubbing in an allowable The fauna encountered in this study was timeframe They were geographically categorised characterised by a morphospecies concept or following Richards et al. (2018) (see Table 2). Operational Taxonomic Unit (OTU) where a ‘normal’ adult specimen or specimen lot, representing a The smallest size category of molluscs retrieved species, was chosen as the morphotype for that (i.e. ≤5 mm adult size class) was classified as micromolluscs (M), following Geiger et al. (2007). taxon. That specimen/lot became the reference Recognition of this fraction is important as it permits for all future identifications of that particular comparison with recent published literature. A taxon. Every subsequent observed shell was slightly larger category of >5–10 mm adult size (Me) compared to the pool of OTUs and matched or was also sorted and tallied. While not formally provided with a new OTU and photographed. The treated by previous authors, it is recognised here morphospecies concept was utilised as a practical because it represents a neglected component of approach to characterise high diversity, with species molluscan diversity that has not previously been identifcation undertaken where possible. Middelfart studied in Western Australia. Additionally, many (2002a,b) used this method on the microbivalve superfamilies include species that span adult groups, Condylocardiinae and Cuninae. The sizes e.g. 1–10 mm in the and technique relies on the ability of the taxonomist to Pyramidelloidea and the micromollusc size class differentiate species, with less focus on taxonomic splits these superfamilies arbitrarily. Consequently identifcation, which can be time consuming and we include this larger size category to begin to reliant on the taxonomic expertise of the researcher. address this issue. Juvenile molluscs with an adult This rationale is similar to barcoding, which involves size >10 mm (Ma) were tallied as the third size sequencing a quickly evolving gene region (such category. as COI or COX1) for species comparison, but with These were included because, similar to the different strengths and weaknesses. For example, >5–10 mm adult size class referred to earlier, they cryptic species are likely to be lumped into one represent diversity often overlooked. However, species, but the involvement of a taxonomist they can be crucial for insights into little known, may offer the additional beneft of a full species yet taxonomically informative life-history stages, identifcation. which may highlight cryptic species (pers. comm. K. Taxa were identifed to species or morphospecies Ockelmann). as defned above. Systematic placement and current The smaller-mollusc sampling program was name assignments were checked against an addition to the already existing macromollusc (World Register of Marine Species), the AFD survey and methods were developed to minimise (Australian Faunal Directory) and the ALA (Atlas additional survey time and effort. Four collecting of Living Australia) to reach a consensus as at 1 bags were designed and constructed to retain March 2018, with WoRMS the ultimate arbiter. sediment while allowing water to drain. The Taxonomic experts were contacted for assistance collectors (C. Bryce and L. Kirkendale), separated with problematic groups (Dan Geiger, Scissurellidae by at least 50 m as defned by the transect survey and , Eatoniellidae, Rissoellidae, 120 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 17 14 3 15 11 4 13 13 4 25 14 11 4 2.5ml sediment sorted 47 18 44 5ml sedimemt sorted 38 13 15 ml sediment sorted 48 30ml sediment sorted Full sample sorted 132 0 3 43 49 Max. depth 0 17 17 20 15 20 16 16.8 16 16 15 18 0 0 14 0 20 0 0 18 13 22 21 18 E125.55891° E125.55947° E125.30443° E123.89345° E125.36195° E125.15963° E123.84163° E124.10310° E124.33061° E124.53304° E124.53580° E124.66444° E124.66100° E124.73986° E123.53458° E123.55461° E123.55514° E123.53898° E123.55955° E123.89476° E124.01858° E125.36179° Lattitude Longitude S14.09939° S14.11046° S14.25298° S13.90069° S14.05731° S14.25455° S15.27486° S14.99347° S14.11754° S14.11675° S14.05740° S15.58797° S15.06938° S14.98867° S14.93606° S14.93457° S14.95373° E124.67258° S14.83227° E124.72337° S14.84764° S14.11258° S14.11369° S14.10808° S13.89635° S13.43058° ations with habitat notes, latitude and longitude, maximum depth (after Bryce et al. 2018) and volume of sorted subsample with identifed number of number of of sorted subsample with identifed and volume Bryce et al. 2018) maximum depth (after and longitude, ations with habitat notes, latitude Sampled st species captured. species Intertidal: reef platform Sloping sand plain Subtidal fore-reef slope Submerged shoal reef patch Dive: Subtidal fore-reef slope Patch reef Subtidal fore-reef slope Subtidal fore-reef slope reefPatch reef patch Dive: Sand and shell rubble plain Patch reef Sub-littoral fore-reef slope Subtidal fore-reef slope Intertidal: reef platform Intertidal: reef platform outcrop Isolated Intertidal: reef platform Subtidal fore-reef slope Intertidal: reef platform Intertidal: reef platform Submerged shoal Submerged shoal

112 113 114 108 111 115 92 95 105 106 110 Station Habitat 91 93 94 96 97 98 99 100 102 103 104 107 109 TABLE 1 TABLE SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 121 48 26 4 42 45 70 27 40 10 15 24 53 25 33 21 38 37 45 18 45 17 36 14 20 20 2.5ml sediment sorted 5ml sedimemt sorted 15 ml sediment sorted 30ml sediment sorted Full sample sorted 30 Max. depth 12 0 15 30 30 15 13 1 20 1 16 15 14 14 6 1.5 6.5 15 21 20 15 15 15 21 22 22 E125.20915° E122.97277° E123.02014° E123.0284° E123.10063° E123.16004° E123.12373° E123.13609° E123.12887° E123.12888° E123.06116° E122.98128° E122.92641° E122.98484° E123.01711° E122.99586° E123.14563° E123.33585° E123.37858° E123.32208° E123.38967° E124.26672° E124.58352° E124.03244° Lattitude Longitude S12.23284° S14.30367° S12.21418° S12.19378° S12.18392° S12.23728° S12.29350° S12.27733° S12.18848° S12.1929° S12.17297° S12.27478° S12.29602° E123.02718° S12.24369° E122.24369° S12.19448° S12.26233° S12.27858° S12.24147° S12.20775° S11.98815° S11.96167° S11.97404° S11.97605° S12.79930° S13.07686° S12.44741° Intertidal Subtidal fore-reef slope Lagoon back reef Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope reef platform Intertidal snorkel: Subtidal fore-reef slope Intertidal: reef platform Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Lagoon back reef Intertidal reef snorkel: platform Tide pool Lagoon Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Submerged shoal Submerged shoal Submerged shoal 123 Station Habitat 116 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 142 143 144 145 146 147 148 122 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 27 13 25 30 23 25 11 38 13 13 12 6 14 33 6 11 36 10 21 18 7 26 16 22 17 24 2.5ml sediment sorted 5ml sedimemt sorted 15 ml sediment sorted 30ml sediment sorted Full sample sorted Max. depth 14 14 20 20 13 20 15 20 15 20 14 0 18 14 0 8 8 0 20 0 18 8 0 17 0 E119.64845° E119.62708° E119.35901° E119.38384° E119.36871° E119.37663° E119.35914° E119.37819° E118.96577° E118.97325° E118.96240° E118.96316° E118.97383° E118.96511° E118.97094° E118.94100 E118.97723° E118.96272° E118.93266°E118.97271° 20 E119.37061° E119.38378° E119.37496° E119.38068° E119.36826° E119.64928° Lattitude Longitude S17.06287° S17.07240° S17.25188° S17.36311° S17.28 432° S17.27991° S17.29773° S17.29298° S17.50697° S17.53597° S17.5374 4° S17.50915° S17.54793° S17.55509° S17.53619° S17.56070 S17.593 47° S17.50573° S17.51110° S17.56666° S17.38771° S17.31697° S17.31431° S17.30587° S17.1546° S17.09053° Subtidal fore-reef slope Lagoon Sub-littoral fore-reef slope Sub-littoral fore-reef slope Lagoon Sub-littoral fore-reef slope Lagoon Subtidal fore-reef slope Subtidal fore-reef slope Subtidal fore-reef slope Lagoon Intertidal Subtidal fore-reef slope Lagoon Intertidal Lagoon Subtidal fore-reef slope Intertidal Subtidal fore-reef slope Subtidal fore-reef slope Intertidal Subtidal fore-reef slope Lagoon Intertidal Lagoon Lagoon Station Habitat 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 174 175 176 SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 123

Skeneidae and Tornidae). The distributions for 700 named species were checked in ALA as well as general literature to assess potential range extensions. 600 All taxa identifed to species level or allocated an OTU were allocated a unique registration 500 number (SXXXXX) and accessioned into the mollusc collection of the WAM. The shells corresponding to OTUs were 400 photographed with a Tucsen 10MP microscope camera or a Canon Eos 6D camera mounted on a variety of microscopes, including Wild Heerbrugg 300

M7A, M3Z, Leica MZ16, M125 and M205. Images Sspecies accumulatio n from the Canon 6D were focus stacked by hand 200 setting the low area and high point focus and imaging fve to ten points in between. These were then combined in Helicon Focus (http://www. 100 heliconsoft.com/heliconsoft-products/helicon- focus/) to achieve the best depth of feld. Samples from this project were only subsampled 0 90 100 110 120 130 140 150 160 170 180 once. Additional sorting effort would certainly Station number generate more specimens and probably additional species. Live taken specimens from samples where FIGURE 1 Species accumulation as a function of only dead shells were found initially is also likely. station number (= sample area/volume). Unsorted sediment samples are housed at the WAM. superfamilies reported here, a number of species RESULTS were represented by juveniles or subadults of This project involved approximately 1,000 hours of macromolluscs. These diversity counts are low laboratory-based sorting, photography, identifcation, compared to data presented in the historic dataset databasing and manuscript preparation. These (solid bars at the top of chart, Figure 2). Conversely, hours do not include the time taken to accession the 21 families were not documented in Willan et material into the WAM mollusc database. al. (2015) or ALA (1 March 2018) as occurring in north-western Australia, (open columns at the Assessment of 77 stations and 1,883 identifed bottom of Figure 2). These were: Anabathridae, specimen lots revealed 505 smaller mollusc species Condylocardiidae, Dialidae, Eatoniellidae, in the ≤5 mm adult size class (M), 43 species from Horaiclavidae, , Leptochitonidae, Litiopidae, >5–10 mm adult size class (Me), 68 juvenile or , Neoleptonidae, , subadult species from the >10 mm adult size class Orbitestellidae, Pickworthiidae, Ringiculidae, (Ma), and 8 species whose adult size could not be Rissoellidae, Scissurellidae, Seguenzioidea determined (Table 2, Column 2). (unplaced), Skeneidae, Sportellidae, Thraciidae A species accumulation graph (Figure 1) was and Tornidae (Table 2, Column 2). Groups with generated to examine the effect each added station numerous microscopic taxa (e.g. rissoids/rissoinids, had on total diversity. The data illustrate the linear cerithiopsids and caecids) had a several-fold increase increase in diversity with each additional station. No in the number of taxa known from the Kimberley. asymptote was reached, indicating more taxa were This finding represents more than 20% of being added at a constant rate for every additional the entire diversity of families globally or an station assessed. approximate 9% increase in total molluscan family For each family, the number of taxa reported in diversity for the Kimberley region (209 families from this study is presented alongside the numbers for Willan et al. 2015 and 21 additional from this paper). that family from the Kimberley historic mollusc In previously documented families, many paper (Willan et al. 2015), to aid comparison (Table 2, additional records came to light, resulting in a Columns 4–5). doubling or more of taxa in the Atlantidae, Caecidae, One hundred mollusc families were found in Capulidae, Cerithiopsidae, Cylichnidae, , this study, compared with 209 families reported Epitonidae, , Galeommatoidea, Iravadidae, in the Kimberley historic mollusc paper (Willan et Liotidae, Philinidae, , , al. 2015), where it was noted that smaller molluscs Rissoidae, , Triphoridae and were underrepresented. Of the 102 families or (Table 2, Column 2). 124 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

Conidae Muricidae Veneridae Tellinidae Pectinidae Nassariidae Mytilidae Arcidae Columbellidae Pyramidellidae Ranellidae Fissurellidae Lucinidae Discodorididae Semelidae Spondylidae Pteriidae Triphoridae Dentaliidae Galeommatoidea Limidae Carditidae Raphitomidae Aplysiidae Haliotidae Haminoeidae Pholadidae Trapezidae Eulimidae Colubrariidae Turritellidae Corbulidae Chamidae Epitoniidae Triviidae Mesodesmatidae Gastrochaenidae Nuculidae Malleidae Vermetidae Hipponicidae

axon Pleurobranchidae T Velutinidae Myidae Cavoliniidae Calliostomatidae Clathurellidae Rissoidae/Rissoinidae Neomeniidae Chaetodermatidae Aplustridae Ungulinidae Crassatellidae Propeamussiidae Cylichnidae Vanikoridae Cystiscidae Hiatellidae Philinidae Atlantidae Iravadiidae Plesiotrochidae Capulidae Scaliolidae Cerithiopsidae Caecidae Ringiculidae Seguenzioidea (unplaced) Sportellidae Juliidae Orbitestellidae Horaiclavidae Litiopidae Leptochitonidae Thraciidae Omalogyridae Eatoniellidae Pickworthiidae Dialidae Scissurellidae Anabathridae Neoleptonidae Philobryidae Rissoellidae Murchisonellidae Skeneidae Condylocardiidae Tornidae 010203040 50 60 70 80 90

Number of taxa

FIGURE 2 The number of species level taxa (grey = historical data, Willan et al. (2015); white bar = this survey data) in each family as presented in Willan et al. (2015) and this study. Families without any representation in this study have been omitted. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 125

The rarity of the species identifed is shown in perparvulum (Watson, 1886) Figure 3, with just over half of the species found only (KimMoll74) (Figure 4C), was the third most once during the entire project, while almost 70% common species and found at 37 localities. It was were found only once or twice. encountered live from the intertidal to 13 m depth, with dead shells found at deeper depths. The only THE TEN MOST COMMON SPECIES readily available information about the distribution of this species is its presence in the Indo-Pacifc Of the most commonly encountered species in this Ocean with ingression into the Mediterranean Sea. project, nine were microgastropods and one was a In Australia it has been recorded from tropical bivalve macromollusc (Figure 4). northern Queensland (Qld), Gulf of Carpentaria The most frequently occurring species overall was and Northern Territory (NT) (Online Zoological Parashiela cf. invisibilis (Hedley, 1899) (KimMoll326) Collections of Australian Museums (OZCAM) and (Figure 4A), which was found live in 43 samples AFD). No records are available from WA and it is across all depths. Records for species of Parashiela here considered a new record for the state. Apataxia are sparse in north-western Australia with only two cerithiiformis Tryon, 1887 (KimMoll314) (Figure records known from the Project Area (ALA, 2018). 4D) is the fourth most common species found The second most common species was Lophocochlias in 30 localities. Scissurella cf. quadrata Geiger and parvissimus (Hedley, 1899) (KimMoll18) (Figure 4B) Jansen, 2004 (KimMoll4) (Figure 4E) was common, identifed from 39 samples and found at all sampled found at 25 stations followed by the caecid, Caecum depths. This species was originally placed in sepimentum de Folin, 1868 (recorded as Caecum by Hedley (1899: 554–555, Figure 67) based on lilianum Hedley, 1903 in most databases in Australia, material collected at Funafuti Atoll in the western see Pizzini et al. (2013) (KimMoll14) (Figure 4F). The Pacifc (see discussion below). type locality for the latter species is Maurice, La Réunion (lectotype, Muséum national d’Histoire

350 naturelle (MNHN) type collection) and it has been recorded from 11 localities in eastern Australia, most commonly at Masthead Reef, Qld (ALA) yet, the only comment about distribution made by Pizzini et al. (2013) was that C. sepimentum exists in Australia. 300 The most common bivalve was Septifer cumingii Récluz, 1849 (KimMoll52) (Figure 4G) and it was the seventh most common mollusc, found at 22 stations.

250 Live specimens were recorded from the intertidal to 13 m depth. Previously known as Septifer australis Laseron, 1956, this name has now been synonymised with S. cumingii (AFD, Huber 2010). Two additional

200 species were found in samples from 21 stations; Pandalosia ephamilla (Watson, 1886) (KimMoll11) (Figure 4H) (Warén, 1983, Figures 71, 72), and the, as yet, unidentifed eatoniellid (KimMoll360) (Figure

No. of instances 4I) (Winston Ponder personal communication). 150

NOTEWORTHY STATIONS With a total of 132 species, station 91/K12 was 100 the most diverse of the 77 stations surveyed. The sample was part of the initial sorting trial for smaller molluscs and proved too labour intensive to complete sorting. Based on the unsorted 50 material remaining, this sample is estimated to contain in excess of 200 species. The site is a high energy deposition plain north-west of Montgomery Reef, between the Bonaparte and 0 Buccaneer Archipelagos (Bryce et al. 2018). The 0510 15 20 25 30 35 40 45 sediment at the site has been sorted by subsea Frequency of occurence in dataset tidal currents leaving a coarse accumulation of comminuted shells and broken coralline rubble FIGURE 3 Number of instances (frequency of occurrence) species were found in the rich in small molluscs. No living specimens were dataset. encountered in the sample. 126 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

FIGURE 4 Most common species found. A) Parashiela cf. invisibilis (Hedley, 1899) (KimMoll326, WAM S94001); B) Lophocochlias parvissimus (Hedley, 1899) (KimMoll18, WAM S94346); C) Cerithidium perparvulum (Watson, 1886) (KimMoll74, WAM S94246); D) Apataxia cerithiiformis Tryon, 1887 (KimMoll314, WAM S94319); E) Scissurella cf. quadrata Geiger and Jansen, 2004 (KimMoll4, WAM S84509); F) Caecum sepimentum de Folin, 1868 (KimMoll14, WAM S84515); G) Septifer cumingii Récluz, 1849 (KimMoll52, WAM S84953); H) Pandalosia ephamilla (Watson, 1886) (KimMoll11, WAM S84851); I) Eatoniellidae (KimMoll360, WAM S68921). Scale bars 500µm. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 127

129, 138 138 129,

129

Offshore

South North 154, 165

** * Midshelf

105

109 109

Stations North 111

111

116* Inshore Central 95

98,

* South 91* 112 91* 91

91* 91* 91* Habitat

S/H S/H S/H No. historical taxa 28 U H No. survey taxa U 12

H 4 1

2 0

2 0

U U U KimMoll96 KimMoll355 OTU KimMoll82 KimMoll49 KimMoll317 KimMoll2 KimMoll1 KimMoll109 KimMoll107 KimMoll110 ?Me M Ma

M

M

Shell size M M ?Me axa from the survey area; species (registration number) = placed or unplaced species with reference specimen for OTU being the registration being the registration OTU specimen for axa from the survey area; species (registration number) = placed or unplaced with reference ) ) )M )M S84585 ) ) ) S84616 S84503 S84502 ) sp. ( sp. ( sp. ( number (bold = image available, regular = no image available); Shell size (M = micromollusc <0.5 mm; Me = medium mollusc 0.5 to 10 mm, Ma = macromollusc mm, Ma = macromollusc (M = micromollusc <0.5 mm; Me medium mollusc 0.5 to 10 Shell size regular = no image available); number (bold = image available, taxa alongside the same indicated Number of taxalevel = species recorded Unit (OTU); Taxonomic mm); KimMollXXX = allocated Operational adult >10 juvenile station with a positive Each et al. (2018). Richards Stations follow et al. (2014). Sampey Habitat codes follow data from the historical data set (Willan et al. 2015). into South, Central and North. are subclassifed in turn which is indicated under the regional divisions of Inshore, Midshelf or Offshore, the OTU identifcation for identifcation. species collected. StationStationmorphotypespecimen * = number with an = live bold numbers in indicated after number (with registration Micromollusc t S84559 S84615 S84613 ) ) S84600 sp. ( sp. ( sp. (

sp. ( S90034 S84908 Leionucula Acar Acar congenita ( Smith,(E.A. 1885) cf. sp. ( sp. ( Chaetoderma Chaetoderma Nucula Nucula Nucula Family: Arcidae Family: Subclass: Pteriomorphia Subclass: Family: Nuculidae Subclass: Protobranchia BIVALV IA Family: Neomeniidae SOLENOGASTRES Family: Chaetodermatidae Taxa and speciesTaxa registration number CAUDOFOVEATA TABLE 2 TABLE 128 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 126, 127, 127, , 126,

126* 143* 123, 128*, 128*, 123, 143, 133, 129, 145 128* 106, 123, 127*, 127*, 123, 106, 142, 130, 128, 143 103*, 133

110*

151

130 105 135, 144 Offshore 166 152*,

170

152*

149 , 156, , 156, 149 165

South North

165 Midshelf

123, 124, 124, 123, 128*, 134

Stations

111*

North 110, 111*

Inshore

Central 99

100, 114*, 116 South 91* 111 114 114* 114* 91 Habitat No. historical taxa H U U U U U H 7 0 U U U U U U U U No. survey taxa U H KimMoll236 KimMoll546 KimMoll442 KimMoll423 KimMoll422 KimMoll332 KimMoll236 KimMoll278 KimMoll301 KimMoll564 KimMoll95 KimMoll277 KimMoll276 KimMoll224 KimMoll223 OTU KimMoll222 KimMoll50 M M M M M M

?Ma ?Ma ?M Ma ?Ma ?Ma M M Shell size M Ma )M ) ) ) ) ) ) ) S84758 ) ) ) ) ) ) ) S84871 S94263 sp. ( S84829 S90285 S84740 S84739 S84738 , fragment) ) sp. ( S84737 S94138 S90346 S90288 S90287 S90138 S84598 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( S94264 S84830 Acar Acar Acar Acar Acar Micromytilus sp. ( Cosa Cosa Cosa Cosa Cosa ? sp. ( Miratacar wendtiMiratacar ( 1907) (Lamy, Bentharca Arca cf. cf. cf. cf. cf. Acar Family: Philobryidae Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 129 , , 127 , 130, 135, 135, , 130, , 140, 145 136 146

134 109 109*

110*

103*, 123 103*, 128* 107*

123*

105, 106, 109 123, 128, 129* 136 106* Offshore

, 151 , 161, 162, 162, , 161, 154, 156, 157 165, 170, 176 149,

152, 165

152

152*

South North

Midshelf

148

Stations

110 111*

110, 111, 113

North

116* , 100, 114 , 100, Inshore

114, 116*

116* 115, 93 114* 114*

Central

91*

South S/H S/H Habitat

No. historical taxa 30

U U U

H H U U U 11 U U H H U No. survey taxa 5 9 U KimMoll52 KimMoll61 KimMoll279 KimMoll103 KimMoll63 KimMoll57 KimMoll51 KimMoll336 KimMoll211 KimMoll297 KimMoll235 KimMoll475 KimMoll286 KimMoll562 KimMoll509 OTU KimMoll333 Me ?M ?M M ?M Ma Ma ?Ma Me M

?M ? Ma M M

Shell size Me ) ) ) ) ) ) ) ) ) )

) ) )

S84757 S84725 S84866 S84560 S84968 S84538 S84564 S84831 S84609 S68867 S84567 S94260 S92528 , fragment) sp. ( sp. ( sp. ( sp. ( S84847 sp. ( sp. ( sp. ( sp.( cf. minimus sp. ( sp. ( ) ) sp. ( Limatula S68900 S84952 Septifer cumingii ( Récluz, 1849 Musculus Musculus Musculus Modiolus Lithophaga Lithophaga malaccanaLithophaga ( 1858) (Reeve, Gregariella coralliophaga Gregariella ( 1791) (Gmelin, Dacrydium cf. Limatula Lima Ctenoides Ctenoides Hayami & Kase, 1993 ( Septifer rudis Dall, Bartsch 1938 & Rehder, ( Family: Mytilidae Family: Taxa and speciesTaxa registration number Limidae Family: 130 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 134 , 123* 126, 128* 126*, 133

109*

, Offshore 156 162 , ,

168* 150 149*

152* 160 South North

153

168* Midshelf

Stations

111*

111 North

110*

* Inshore

116*

114*

116 Central

91* South

91* S/H

Habitat S/H S/H

36 No. historical taxa 13 U U U 3 2 H H H U

U 5 U H H No. survey taxa

2 4

U 6 H KimMoll513 KimMoll437 KimMoll506 KimMoll606 KimMoll73 KimMoll550 KimMoll335 KimMoll283 KimMoll101 KimMoll281 KimMoll72 KimMoll566 OTU KimMoll53 KimMoll304 KimMoll102 KimMoll608 M M

?M Ma ?M ?Ma ?Ma Ma

?M Ma Ma Ma

Shell size Ma Ma

?Ma

) ) ) ) )M ) ) ) S84576 S84575 S94267 S84540) S84884 S94109 S84606 madreporarum S84607 S85534 sp. ( cf. ) ) ) ) ) ) ) sp. ( sp. ( sp. (spat) ( (spat) sp. sp. (spat) ( (spat) sp. sp. (spat) ( (spat) sp. sp. (spat) ( (spat) sp. sp. (spat) ( (spat) sp. S68918 S90321 S94154 S84966 S84839 S84833 S85520 Malleus Malleus Cyclopecten Pinctada Pinctada Pinctada Pinctada Pinctada sp. ( sp. ( cf. sp.( sp. ( sp. ( sp. ( Coralichlamys (Sowerby, 1842) ( 1842) (Sowerby, sp. ( ? ? ? cf. ? cf. ? Family: Propeamussiidae Family: Pectinidae Family: Taxa and speciesTaxa registration number Family: Malleidae Family: Pteriidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 131 , 128, 142 , 128,

105* 138* 128*

134* 129*

126*

127

, Offshore , 167, 169 , 167,

159*

151*, 162, 162, 151*, 166 170, 171

South North 149, 154, 154, 149, 168167,

Midshelf

146, 148

Stations

North

111*

Inshore

114*

100 Central

100, 114, 116*

91*

91* 91* South 91 S/H Habitat S/H

0 No. historical taxa 13 U U U U

15 U U U U

U U 6 9

No. survey taxa

U 2 KimMoll99 KimMoll310 KimMoll439 KimMoll424 KimMoll582 KimMoll496 KimMoll469 KimMoll229 KimMoll542 KimMoll560 KimMoll111 KimMoll94 OTU KimMoll65 KimMoll280 M M M M

?M M M ?M Me

Ma Ma

Shell size )M ) )M

) ) )M ) ) ) )

S92503 S90328 , S84603 ) S90290 S94232 S84618 S84597 S94101 S84569 S84832 ( sp. ( sp. ( S94370 ( sp. ( , fragment) sp. ( sp. ( sp. ( S84746 sp. ( sp. ( S68855 Cunanax Cyclocardia Crassacuna Crassacuna crassisculpta Crassacuna Middelfart, ( 2002 Condylocardia Condylocardia cf. ? Condylocuna tricosa Condylocuna (S84900) 2002 Middelfart, fragment) sp. ( Glans sp. ( Cyclocardia cf. Cyclocardia Cyclocardia Spondylus Spondylus Family: Condylocardiidae Family: Carditidae Family: Subclass: Taxa and speciesTaxa registration number Family: Spondylidae 132 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 139 ,

106, 123 106,

103*

134*

134*

139* 133*

128

128, 136, 138

Offshore 165*

153*, 175

151 South North Midshelf

Stations

110*

North

98 Inshore

97, 100*

Central 114*

91* 91* South 91* 91* 91* 91* S/H Habitat No. historical taxa U H U

4 0 U U U 3 2 U U U U U U U U U No. survey taxa KimMoll596 KimMoll303 KimMoll249 KimMoll234 KimMoll499 KimMoll529 KimMoll498 KimMoll457 KimMoll501 KimMoll100 KimMoll77 KimMoll79 KimMoll76 KimMoll78 KimMoll230 OTU KimMoll80 M M M M M M

M M M M M M M M M Shell size )M )M ) ) ) ) ) )

) ) ) S94016 S68862 S84604 S84580 S84582 S84579 S84583 S84747 sp. ( sp. ( S84883 S84776 S84756 ) ) sp. ( cf. atkinsoni ) sp. ( sp. ( sp. ( S68863 S90425 ) S68871 Salaputium Salaputium S84581 sp. (S94452) sp. Thracia Thracia Thracia sp. ( cf. cf. sp. ( sp. ( Warrana triangulata Middelfart, ( 2002 Warrana pellucida Middelfart, ( 2002 Warrana lunata Middelfart, ( 2002 Warrana brucemarshalli Middelfart, ( 2002 (Tenison-Woods, 1877) 1877) (Tenison-Woods, ( Ovacuna Crassacuna Crassacuna praecalva Crassacuna (Hedley, 1909) ( 1909) (Hedley, Family: Thraciidae Family: Crassatellidae Family: Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 133 , 138* 131 , 126 129*

123*

127*

123,

, Offshore 166 , , 159, 162, 162, , 159, , 154, 157 154, , South North 151

162*

149* 158 165, 171, 167, 176 174*,

164

158 162* 159 171* 171

166* 164*

Midshelf

Stations North

112*

Inshore Central

114*

114*

95* South

91* 91*

Habitat S/H S/H

S/H S/H S/H S/H

No. historical taxa 21 49 13 No. survey taxa

U 1 1 U

2 6 U U U U U 7

U

5 U U 2 U OTU KimMoll536 KimMoll417 KimMoll587 KimMoll210 KimMoll549 KimMoll105 KimMoll98 KimMoll511 KimMoll363 KimMoll588 KimMoll378 KimMoll227 KimMoll610 KimMoll447 KimMoll601 KimMoll592 KimMoll346 M ?

Shell size ?M ?M

M M Ma

?M ?M M ?M Ma

?M ?M

Ma )? ) )M ) ) ) ) S68914 S94415 ) S84724 S84602 S90088 sp. ( sp. ( sp. ( S84998 S90367 ) ) , broken ) ) sp. ( ) ) ) ) ) sp. ( sp. (S85548 sp. ( sp. ( S94060 S94153 S90265 S84611 S94416 S90139 S84744 S94495 S94441 Pillucina Epicodakia Jactellina sp. ( Hiatella Coralliophaga Coralliophaga sp. ( sp. ( sp. ( cf. cf. Epicodakia sp. ( sp. ( sp. ( ? Cadella sp. ( sp. ( Taxa and speciesTaxa registration number Family: Hiatellidae Family: Trapezidae Family: Lucinidae Family: Tellinidae Family: Family: Semelidae 134 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 140 ,

107*

134*

127*, 139

130* 123, 124, 126, 126, 124, 123, 137*

Offshore 168 162* 162,

165 170* South North 149, 174

149, 176

175 Midshelf

Stations , 114*

110

111 North

111*, 113111*, 110 , 111

, 114, 115, 115, , 114, Inshore

97 116*

114, 116*

114* 100*

92, 94, 114, 114, 94, 92, 116* 92, 114, 116* Central

114*

91* 91 South

S/H

Habitat S/H S/H 10 No. historical taxa U

U U U 20 U U U 1 0 U U U No. survey taxa U 4 5 U

H

U 5 66 KimMoll591 KimMoll70 KimMoll288 KimMoll55 KimMoll97 KimMoll482 KimMoll228 KimMoll252 KimMoll68 KimMoll67 KimMoll614 OTU KimMoll414 KimMoll282 KimMoll225 KimMoll479 KimMoll398 KimMoll212 M M M

M M

Ma Ma Ma Ma

Ma Ma Ma Ma Ma

Shell size )M )M )M ) ) ) ) ) ) ) ) , fragment) ) S84745 ) ) ) S94433 S84601 S84742 S84837 S84726 S84573 sp. ( S84571 S84570 S85581 sp. ( S84783 sp. ( S84850 S84558 sp. ( sp. ( S92542 S90201 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( Kurtiella Kellia Borniola Melliteryx Irus Irus Lasaea cf. cf. Kellia Kellia cf. cf. sp. (S92571) sp. Chama sp. ( Chama sp. ( Chama sp. ( Chama sp. (S90256 Venerupis ? ? Antigona Venerupis Superfamily: Galeommatoidea Family: Sportellidae Family: Family: Chamidae Family: Taxa and speciesTaxa registration number Family: Veneridae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 135 138 , 128, 134 , 128, 124

136* 123* 109* 124*

123* 129*

138*

123, 127

105*,

Offshore , 176 , 170, , 170,

South North

155

167, 169167, 149 175

175, 176* Midshelf

Stations

North

110

116* Inshore Central

114*

92, 95, 96, 96, 95, 92, 115,

South 91*

91*

91*

91* S/H Habitat S/H S/H S/H S/H

S/H No. historical taxa

1 4

U

U U U

No. survey taxa U 1 5 U 1 4

U H U U KimMoll81 OTU KimMoll483 KimMoll517 KimMoll54 KimMoll484 KimMoll209 KimMoll104 KimMoll510 KimMoll470 KimMoll108

KimMoll440 KimMoll56 KimMoll604 KimMoll327 KimMoll112 KimMoll531 M

M M M M M M M M Shell size Ma

M

Ma M M M M ) ) ) ) ) ) ) ) ) ) sp.

S84584 S84555 S84614 cf. philippiensis S84541 S68800 S84619 S84610 ) S68906 S85512 S84929 S94062 ) ) ) ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( cf. ovata sp. ( S92587 S68925 S90336 Pseudogaleomma S84722 Gastrochaena sp. ( sp. ( Scintilla Scintilla Mysella Mysella Mysella Mysella ovata ( 1906) (Hedley, Corbula macgillivrayi macgillivrayi Corbula ( Smith, 1885 sp. ( Thiele, ( 1830 cf. ( Mysella Mysella Mysella (S92504) 1906) (Hedley, Mylitta Family: Gastrochaenidae Taxa and speciesTaxa registration number Family: Corbulidae Family: Family: Mesodesmatidae Family: 136 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , , , 140 136 , 130 127* 137 136*

109*

128*, 130 123*, 124, 126 124, 123*, 128,

130*

123* 129*

107*

Offshore

153

South North

Midshelf

Stations

110*

North

113*

Inshore

100* 114

116

Central

91

91* South S/H S/H S/H S/H S/H Habitat

No. historical taxa

4 0

U U U

U

6 0 U U No. survey taxa 2 2 U 1 6 U 1 3 KimMoll394 KimMoll393 KimMoll302 KimMoll3 KimMoll250 KimMoll83 KimMoll426 KimMoll69 KimMoll106 KimMoll522 KimMoll516 KimMoll472 OTU KimMoll359 KimMoll289 ?Me ?M ?M Me

M M M M M

M M

?

Shell size )M )? ) ) ) ) ) ) S84612 S90059 S68923 S92511 S68990 S84535 S90293 S90108 sp. ( ) ) ) ) ) sp. ( ) sp. ( sp. ( sp. ( sp. ( S90193 S90191 S84872 S84777 S84586 S84852 Micropolia Martesia sp. ( sp. ( sp. ( Parachiton sp. ( sp. ( Neolepton Neolepton cf. Micropolia Diplodonta sp. ( Diplodonta sp. ( cf. sp. ( Family: Leptochitonidae Family: POLYPLACOPHORA Family: Neoleptonidae Family: Ungulinidae Family: Pholadidae Taxa and speciesTaxa registration number Family: Myidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 137

, , 143 , 131, 109* 139* , 140 130 , 137, 138, 138, , 137, ,

139*

128

143 139

129*

134 139 103 134, 128, 126, 123, 129,

Offshore 176 , 165, , 165, , 169 ,

150

153, 154, 156, 158 156*

154*

162 159* 167 South North 155, 158, 162, 162, 158, 155, 175 169

Midshelf

148*

148

Stations 113 ,

110*

111*

North

110 , 113

Inshore

Central

91* 91*

91* 91

South

S/H S/H Habitat S/H S/H S/H 21 No. historical taxa 2 3 U 4 U H U U 1 8 U U U

No. survey taxa

5 0

1 ?

KimMoll460 KimMoll404 KimMoll577 KimMoll183 KimMoll182 KimMoll299 KimMoll551 KimMoll167 KimMoll171 KimMoll500 KimMoll583 OTU KimMoll4 KimMoll443

Ma

M M ?M ?M

?Me ?Me Ma M M M

?Me

Shell size )

) ) )? ) ) S90441 ) )

) )

S94281 S84678 S84693 S84694 S84868 S94324 S68864 cf. atkinsoni 94371 S84825 ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( cf. quadrata ) ) sp. ( sp. (Juvenile) ( (Juvenile) sp. S68970 S90213 S84509 Calliostoma Diodora (Tenison-Woods, 1877) 1877) (Tenison-Woods, ( Sukashitrochus Calliostoma Haliotis Puncturella Emarginula Emarginula Scissurella Scissurella Bandel, 1998 ( Bandel, 1998 Scissurella evaensis Geiger & Jansen, 2004 ( Scissurella sp. ( Family: Calliostomatidae Family: Family: Fissurellidae Family: Family: Haliotidae Family: Family: Scissurellidae Family: Subclass: Subclass: Taxa and speciesTaxa registration number 138 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , , , 128 , 140* 134* 127 , 143 , , 130, 133 , 130, , 127*, 128, 127*, 134, 143

128*,129 , 128*,129 132, 134, 135 128 134* 135, 136,

133 128

140*

123, 123, 134 143 145*

142* 131, 136 Offshore , 156 , 170, , 170, South North

154

152*

151, 169* 174 Midshelf

148 146*

Stations , 112 North

112*

109

Inshore Central

98, 100

South

91*

91*

Habitat S/H S/H S/H S/H S/H

S/H S/H S/H No. historical taxa No. survey taxa U 1 0 H

U

U H 6 2

7 3 U

OTU KimMoll415 KimMoll427 KimMoll362 KimMoll418 KimMoll488 KimMoll189 KimMoll494 KimMoll7 KimMoll184 KimMoll374 KimMoll382 KimMoll466 KimMoll563 KimMoll374 M M M M M M M

Shell size M

M ) )M )M )M )M )M ) )

) ) ) ) )

S68934 S85538 S84695 S68954 S94262 S90128 S84700 S68836 S90485 ) S90258 sp. ( sp. ( sp. ( sp. ( sp. ( S90295 S90086 S68837 S68824 sp. ( sp. ( sp. ( sp. ( sp. ( cf. fordiana Austroliotia Austroliotia Austroliotia Austroliotia Austroliotia Brookula Tricolia Tricolia Tricolia Tricolia Tricolia ( 1888) (Pilsbry, ? ? ? Gabrielona pisinna Robertson, ( 1973 Gabrielona sp. ( Bathyliotina schepmani Bathyliotina ( 1953) (Habe, ? ? Superfamily: Seguenzioidea (unplaced) Superfamily: Seguenzioidea Taxa and speciesTaxa registration number Liotiidae Family: Family: Phasianellidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 139 , 105 , 134, , 134, , , 107, 108, 108, , 107, 145* 107, 111*, 128, 128, 111*, 107, 129, 134 116*

126, 129, 135, 135, 129, 126, 143* 103* 128, 128, 106, 134, 131, 129, 138, 143 139, 134*

103*, 104, 108, 138, 129, 128, 143, 142, 139, 145

107, 109* 145 105 109* 144, 136, 135, 145

102*, , Offshore , 165 161 , 152

154* 156* 165 South North 149,

165*

161, 165 154

Midshelf

146*

147 Stations North

110, 113 110

112

111*

Inshore Central

98

93

South

91

Habitat S/H S/H S/H S/H S/H

S/H No. historical taxa 0 41 U U U U

No. survey taxa

U U U U 13 U

15

KimMoll553 KimMoll275 KimMoll62 KimMoll579 KimMoll567 OTU KimMoll444 KimMoll237 KimMoll495 KimMoll491 KimMoll243 KimMoll599 KimMoll17 KimMoll197 KimMoll5 KimMoll345 M M M M M M M M M M M

Shell size Me Me ) )

)M )M ) ) ) ) ) ) ) ) ) S84806 S84510 S84511 S84995 S90353 S84760 S68854 S68831 S94284 sp. ( sp. ( sp. ( sp. ( S94486 S94196 S84826 S84566 sp. ( sp. ( sp. ( sp. ( ) ) sp. ( sp. ( sp. ( sp. ( Eurytrochus Eurytrochus S94333 S84767 Liotella Liotella Liotella Liotella Leucorhynchia tricarinata Melvill & Standen, 1896 ( Munditiella Lodderena Lodderena Lodderena Lodderena Lodderena ornata & McGinty(Olsson 1958) ( Leucorhynchia caledonica ( Crosse, 1867 Munditiella ? ? Taxa and speciesTaxa registration number Family: Skeneidae Family: Trochidae (and Chilodontaidae) (and Trochidae Family: 140 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 106 , 107 143 105* , 140 , , 140, 143 , 139, 148* , 139, , 106*, , 123, 124, 124, , 123,

137* 135* 102, 105 109 130, 132, 133 134 142*,

105*

100*, 143

126, 133* 127* 131 156 Offshore , , 176 155 , , 170 , 151, 152, 151,

South North 176* 149 149 165 152, 165, 167 152

164*

153 149

Midshelf

Stations North

110

112

100 , Inshore Central

114

96

97

South

91*

91* 91* 91*

Habitat S/H

S/H S/H S/H S/H S/H S/H S/H No. historical taxa 44 14

33

U 1 No. survey taxa H H

U U U

U U KimMoll401 KimMoll185 OTU KimMoll258 KimMoll449 KimMoll6 KimMoll8 KimMoll370 KimMoll593 KimMoll292 KimMoll196 KimMoll195 KimMoll263 KimMoll192 KimMoll406 KimMoll505 KimMoll395 ?Ma

Me

Shell size ?Ma Ma ?Ma M M ? ?Me ?Me Me Ma Me ? Me )M ) ) ) ) ) ) ) ) ) ) ) )

S94072 S84542 S90206 S90226 S90393 S84958 S94446 S90197 S84803 sp. ( S84706 S84696 S84920 S84707 ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( ) ) S68886 Talopena Cerithium Clypeomorus S90121 S84703 ? sp. juvenile ( sp. ? Stomatella Stomatella Alcyna ocellata Adams, 1860 ( ? Stomatella impertusa (Burrow, 1815) ( 1815) (Burrow, sp. juvenile ( sp. sp. juvenile ( sp. Jujubinus Jujubinus Ethminolia sp. Juvenile ( Juvenile sp. sp. ( cf.Granata maculata & Gaimard,1834) (Quoy ( Family: Cerithiidae Family:

Family: Turbinidae Family: Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 141 , , 124 139 , , 133, , 133, , , 127, , 127, , 148 140 126 123 136 125 , 136* , 133, , 133, , , 108, 127, 127, , 108,

124, 136, 1 38 102, 140, 143, 145 134, 143 105, 137* 132* 135, 102 133,132*, 134, 140 139, 129, 137* 140, 136* 112* 105* 108*, 139 105*

102 136*

Offshore , 176 , , 175 155 167 151 , 158, , 158, , , 170, , 170,

149, 153,152*, 154, 156 168 171 154,

161

161, 162

South North

150* Midshelf

146*

147 148

Stations

110 , 113

110

111* North

116 Inshore 100* 93, 98, 100 98, , 93, 114,

Central

97, 98, 100* 98, 97,

South

S/H S/H S/H S/H S/H S/H S/H S/H Habitat S/H S/H No. historical taxa U

H

U

U

U U U No. survey taxa

KimMoll259 KimMoll74 KimMoll419 KimMoll323 KimMoll315 KimMoll342 KimMoll308 KimMoll400 KimMoll400 of =?var. KimMoll316 KimMoll617 KimMoll356 KimMoll353 KimMoll316 KimMoll315 KimMoll268 OTU KimMoll256 KimMoll392 ?KimMoll258 Ma M M M M M M M M M M M M M M Shell size Ma M )

) ) )

S90190 ) ) ) ) ) ) ) ) ) ) S94246 S68825 sp. ( S84797 sp. (S94204) sp. S94088 S90474 S90473 S90205 ) S90192 S90189 S90035 S90029 S84907 S84905) S84793 cf. balteatum sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. (S84813 sp. ( Clypeomorus Pseudovertagus Cerithium Philippi, 1848 ( (Watson, 1886) ( 1886) (Watson, Cerithidium perparvulum Cerithidium diplaxCerithidium ( 1886) (Watson, Bittium Bittium Bittium Bittium Bittium Bittium Bittium Bittium Bittium Bittium Bittium ? ? Taxa and speciesTaxa registration number 142 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , , , 127 , 136 130 107 ,134 , , , 105, , 105, , 126, 137, 137, , 126, , 139, 140 , 139, , 108, , 108, 123 138

103 129* 102* 102 133*

126*

109* 130, 135, 137 102* 132

103* 106, 138 Offshore , , 156 168 165 , 176* , , 171 150 , , , 159, , 159, , 149, 153 162 175 150,155, 160, 162, 165

156* 154* 164* 150*

166* 160 158 170 South164* North

Midshelf

147, 148147, Stations 112 ,

111

111*

North

Inshore

95, 100*

Central

South

S/H S/H S/H S/H S/H Habitat S/H S/H S/H S/H S/H No. historical taxa

5 0

U U U

H

H H No. survey taxa

KimMoll365 KimMoll253 KimMoll239 KimMoll339 KimMoll271 KimMoll502 KimMoll580 KimMoll568 KimMoll594 KimMoll554 KimMoll616 KimMoll241 KimMoll9 KimMoll9 of =?var. KimMoll502 KimMoll338 KimMoll595 OTU KimMoll241 M M

M ?M ?M Me ?Me M Me Me M ?M ?Me Me Ma Shell size ?Me ) ) ) ) )

) ) ) ) ) ) )

S94451 S94202 S84787 S94066 S94491 S94286 S84537 S84765 S94447 S84975 S94105 ( sp. (S94343 sp. (

) ) ) zebrum sp. ( sp. ( sp. (

sp. ( cf. punctatum S90264) S84979 S84818 S68880 Diala albugo ( 1886) (Watson, Diala semistriata (Philippi, ( 1849) sp. ( sp. ( sp. ( sp. ( Clypeomorus Clypeomorus Cerithium stigmosum Gould, 1861 Cerithium Cerithium Cerithium Cerithium egenum Gould, 1849 ( Cerithium ( 1841 Kiener, Cerithium Cerithium Bruguière, ( 1792 Family: Dialidae Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 143 , 124* , 128, 130, 130, , 128, , 139, 140, 140, , 139, 113

131 145 123, 123, 126 133 144

134*

123*

105, 107, 123, 123, 107, 105, 125, 124*, 132, 130, 128, 133, 134, 136, 143, 144, 145

139*

Offshore 171 153* 174 , , 168 159* 150, South North

176

162*

152, 156*, 164

157

162* 149, 155 176* Midshelf

147*

147

Stations

North

110

Inshore Central

93, 100*

South

91*

S/H S/H Habitat S/H S/H S/H S/H S/H No. historical taxa

U U 3 1

U U 5 0 U U No. survey taxa 4 0

U

KimMoll585 KimMoll524 OTU KimMoll264 KimMoll133 KimMoll407 KimMoll390 KimMoll452 KimMoll578 KimMoll369 KimMoll512 KimMoll295 KimMoll586 KimMoll521 KimMoll390 M M M M

M M M M

M M Shell size

M M M )

)M )

) ) ) ) ) ) S68801 ) ) S68850 S94003 S68917 S94332 S68814 translucida sp. ( S84805 S84640 ) ) S94375 sp. ( cf. cf. goniochila S94414 S68963 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( S94407 S90177 S94075 ) Microliotia Scaliola Scaliola Finella Finella sp. ( sp. ( Sansonia Astrosansonia dautzenbergi ( 1917) (Bavay, ? Styliferina Styliferina (Hedley, 1906) ( 1906) (Hedley, Styliferina ( Adams,A. 1860 Diala Diala sp. ( Family: Scaliolidae Family: Pickworthiidae Taxa and speciesTaxa registration number Family: Litiopidae Family: 144 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 130 , 140 143

127, 14 0 109*,

138* 127* 127* 125, 134 116* 109*, 132 134* 123, 126, 138* 126, 123, 127* 109* 126* 107, 139 *

126*

Offshore , 156 , 170

158

156* 165 152*, 165

156 162

152

South North 167

Midshelf

147

147

Stations

111* 111

113

111

North

Inshore 114* 93*

99, 114

114*

114, 116 111, 113

Central

South

S/H S/H S/H S/H S/H S/H S/H Habitat S/H 11 No. historical taxa 1 U U 40 H U

U U

U

U

U

U

No. survey taxa H 16

1 5 KimMoll215 KimMoll351 KimMoll221 KimMoll27 KimMoll572 KimMoll565 KimMoll446 KimMoll413 KimMoll379 KimMoll220 KimMoll28 KimMoll26 KimMoll481 KimMoll269 KimMoll410 KimMoll25 KimMoll545 OTU KimMoll294 KimMoll547

M M M M M M M M M M M M M M M M

M

Shell size )M )M

) ) ) ) ) ) ) ) ) ) ) ) ) ) S90018 S84729 S90106 S84518 S92559 S90447 S90247 S84516 S90438 S94265 S90364) S90255 S90140 S84734 S84556 S84517 sp. ( ) cf. poppearum sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. (S94307 sp. ( sp. ( sp. ( ) S94139 onophorus Costatophora sp. ( S94136 ?M ? Synthopsis Synthopsis Paraseila Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Jaculator Horologica Horologica Horologica Horologica Cecalupo & Perugia, 2012 Cecalupo & Perugia, 2012 ( Clathropsis Cerithiella sp. ( sp. ( Family: Triphoridae Family: Cerithiopsidae Taxa and speciesTaxa registration number Turritellidae Family: SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 145 136 , 142 , , 125 , 140

102, 145

138*

105* 139* 105*

109*

102* 105* Offshore , 158 156 174

154, 156* 154* 157* 170* 174

South North

Midshelf 147*

148 147

147 147

Stations

113

111*

111 110 North

113*

Inshore

114*

114* 114* Central 95, 97, 100, 97, 95, 114* 114* 114

South

S/H S/H S/H S/H S/H S/H S/H Habitat S/H S/H S/H No. historical taxa U U

U U

H U U

U H No. survey taxa

KimMoll388 KimMoll218 KimMoll573 KimMoll445 KimMoll570 KimMoll581 KimMoll613 KimMoll458 KimMoll328 KimMoll321 KimMoll270 KimMoll216 KimMoll214 KimMoll21 OTU KimMoll217 KimMoll213 KimMoll219 KimMoll343 KimMoll322 ?Me M Me M M M M M ?Me M M M M M Shell size M )M )M )

)M )M

) ) ) ) ) ) ) ) ) ) )

) S94315 S84986 S90363 S84924 S84817 S94288 S85565

S94346 S90168 sp. ( S84519 S84731 S84727 S90433 S84931 S84923 S90054 sp. ( sp. ( cf. decorata S84732 sp. ( cf. atratus lamberti sp. ( sp. ( sp. ( sp. ( cf. ) ) Opimaphora S84730 S84728 (Kosuge, 1962) ( 1962) (Kosuge, Monophorus Metaxia fuscoapicata Thiele, ( 1930 Mastoniaeformis Mastoniaeforis Mastoniaeforis ( (Laseron, 1958) Mastonia (Hervier, 1898) ( 1898) (Hervier, Iniforis progressa ( (Laseron, 1958) Euthymella Euthymella Euthymella Costatophora sp. ( sp. (juvenile) (juvenile) Costatophora sp. ( sp. (juvenile) (juvenile) Costatophora sp. ( Costatophora granifera ( 1894) (Brazier, (Brazier, 1894) ( 1894) (Brazier, Costatophora cf. granifera (Brazier, 1894) ( 1894) (Brazier, Costatophora cf. granifera Coriophora Bouchetriphora ? Taxa and speciesTaxa registration number 146 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

143 132, , 135 136 , 145 , 143

, 144 , 107*, 109

135* 145* 142* 140* 102* 128, 134, 140, 140, 134, 128, 145 109, 127*, 143 147*

140* 147 102*

109 99, 140 , , Offshore 156 168 174 174 , , , , 171, , 171, , , 161 South North 166 176* 156* 153* 156*, 162 158, 161, 161, 158, 163 174 170*

154 157 156 174 156

154*, 174

Midshelf 146 148

147*

146*

Stations North

111*

Inshore Central

South

Habitat S/H S/H S/H S/H S/H S/H S/H S/H S/H S/H S/H No. historical taxa No. survey taxa

U U

U

U

U

U U

H OTU KimMoll23 KimMoll24 KimMoll526 KimMoll576 KimMoll451 KimMoll385 KimMoll454 KimMoll373 KimMoll341 KimMoll463 KimMoll387 KimMoll396 KimMoll386 KimMoll267 KimMoll518 KimMoll340 KimMoll487 KimMoll22 KimMoll291 Shell size M M M M M M M M M M M M M M ?Me M Me

)M ) ) ) ) )

) ) ) ) S90415 )M S84857 S94314 S90454 S90126 S84984 tessellatus S85564 S94285 ) cf. cf. tessellatus cf. episcopalis sp. ( cf. sarcira ) ) ) ) ) ) ) S94063 S90164 S84983 S68823 sp. ( sp. ( sp. ( sp. ( S94007 S94323 S90401 S90198 S90163 S84812 S68937 Subulophora rutilans (Hervier, 1897) ( sp. ( sp. ( Laseron, 1958 ( Laseron, 1958 Opimaphora Ophinophora sp. ( Monophorus ( 1963) (Kosuge, (Kosuge, 1963) ( 1963) (Kosuge, Monophorus (Hervier, 1898) ( 1898) (Hervier, Monophorus Viriola sp. ( sp. ( sp. ( sp. ( sp. ( Viriola Viriola Viriola Viriola cancellataViriola (Hinds, 1843) ( Subulophora Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 147 133* 102 , 139 102

104

128 130*

Offshore , 154 , , , 162 , 151 , 158, , 158, 153* 157 161 149

South North

Midshelf

Stations

North 110

111*

Inshore

Central

116*

93

South

91* 91* 91* 91* 91* 91* 91* 91* 91* 91* 91* 91* 91*

Habitat S/H

S/H No. historical taxa 5 U

2 1 U U U U U U U U U U U U U U U

No. survey taxa 16 KimMoll503 KimMoll307 KimMoll274 KimMoll124 KimMoll123 KimMoll122 KimMoll121 KimMoll120 KimMoll119 KimMoll118 KimMoll117 KimMoll116 KimMoll115 KimMoll114 KimMoll113 KimMoll75 KimMoll168 KimMoll350 OTU M

M M M M M M M M M M M M M M Me

Shell size ) )M

)M ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) S94002 S68881 S68981 S84823 S84631 S84630 S84629 S84628 S84627 S84626 S84625 S84624 S84623 S84622 S84621 S84620 S84578 S84679 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( Plesiotrochus unicinctus Plesiotrochus ( Adams, 1853) (A. sp. ( Epitonium sp. Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Epitonium Cycloscala Cycloscala revoluta ( 1899) (Hedley, Family: Plesiotrochidae Taxa and speciesTaxa registration number Family: Epitoniidae 148 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 142,

, 108, , 108, 145 105 ,126 , 136, 138, 138, , 136, , 100 , 97

123, 123, 127 123, 123, 132, 131, 129, 135 139, 140, 143 102,

128* 107* 124, 126, 130*, 134, 139 105*

132*

, , , , 176 151 157 171 , Offshore 169

, , , , , , , , , 168, , 168, , 154 , 156 152 156 167 165 , 176 175* 170 150 150 156 162 , , 168 , 174, , 160 , , , , , ,171, 174, , , , , , 152, 163 163 149 169 174

149 151 153* 155 157 167, 170 175, 176 153

South North

162 175

153* 153* 153* Midshelf

146

Stations

North

Inshore

114* Central 93

116*

South

91*

S/H S/H Habitat S/H S/H S/H

S/H S/H S/H No. historical taxa 3

U 33 U

No. survey taxa U U 3 1 U

KimMoll532 KimMoll207 KimMoll326 KimMoll12 KimMoll172 KimMoll430 KimMoll287 OTU KimMoll478 KimMoll170 KimMoll465 KimMoll527 KimMoll525 KimMoll528 M M M M M

Ma ?Ma M Ma

Shell size M M M M ) ) ) ) ) )

) ) ) ) )

) S84720 S84848 S90309 S94025 S94001 S92540 S90472 S84548 S94005 ) S94014 S94015 S84681 S84683 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( Separatista Haurakia Haurakia novarensis ( (Frauenfeld, 1867) Haurakia marmorata (Hedley, 1907)(Hedley, ( Alvania ? (Hedley, 1899) ( 1899) (Hedley, cf.Parashiela invisibilis Haurakia sp.Trichamathina ( Capulus cf.Parashiela ambulata Laseron, 1956 ( Laseron, 1956 Haurakia sp.Parashiela ( Haurakia Family: Rissoidae Family: Taxa and speciesTaxa registration number Capulidae Family: SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 149 , 126 , 139, , 139, , , 138, , 138, 105 138 103* , 124, , 124, 135 , 139 , 132, 134, 134, , 132,

124*, 137

102, 138, 140

100*

105* 127, 144 127 135, 140, 143, 145

140, 142*

123, 126*

123*, 124, 124, 123*, 130 102, , , Offshore ,

, 162 153 161 156* , , , 176

176 155 165, 166, 168, 168, 166, 165, 170, 171, 174, 175 165 , 168, 174

149, 151, 152 154, 159, 157,

154*

South North

Midshelf

146

148

Stations

111*

110*

North

Inshore

97* 98*

Central

91*

91*

South 91*

S/H S/H S/H S/H S/H S/H S/H Habitat S/H No. historical taxa U

U

U U

U

U

U U

No. survey taxa KimMoll137 KimMoll485 KimMoll273 KimMoll313 KimMoll357 KimMoll330 KimMoll248 KimMoll148 KimMoll314 KimMoll181 KimMoll569 KimMoll372 KimMoll514 KimMoll305 KimMoll305 ?=KimMoll326 KimMoll508 OTU KimMoll242 M M M M M M M M M M M M

M M M M Shell size ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

S84766 S84903 94319 S94130 S84885 S68897 S84692 S84821 S84657 S84644 S68802 S90125 ) S90038 S84936 S84775 S94287 plicatula sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( cf. sp. ( Rissoina Rissoina Rissoina Rissoina Rissoina Rissoina Rissoina Rissoina Rissoina Gould, 1861 ( 1861 Gould, Rissoina ambigua (Gould, ( 1849) ? Rissoina Apataxia cerithiiformis ( 1887) (Tryon, Parashiela Parashiela sp.Parashiela ( sp.Parashiela ( Parashiela liddellianaParashiela 1907)(Hedley, ( Family: Rissoinidae Family: Taxa and speciesTaxa registration number 150 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

, , , , , 133

133 143 , 138, , , , , 129, , 129, 124, 136 132 140 127 130 , 109, , 109, 143* , , , 128*

134 139 139, 123*, 126, 130

133*

102*, 126, 126, 102*, 140, 138, 130, 144 134, 135, 136 140, 143, 144 145 100, 102, 105 107* 126,

Offshore

168*, 170

176 South North 152

Midshelf

148*

147

147, 148147,

Stations

North

112

100* , Inshore 98

96

95*

Central

91* South

91* S/H S/H S/H S/H S/H Habitat S/H No. historical taxa

5 0 U U U U

U No. survey taxa

U

KimMoll255 KimMoll432 KimMoll405 KimMoll360 KimMoll441 KimMoll347 KimMoll607 KimMoll177 KimMoll507

KimMoll337 OTU KimMoll11 KimMoll135 M M M M

M M M M M M

M

Shell size )M ) ) ) S90340 S84851 sp. ( S84974 S84642 ) ) ) ) ) ) ) sp. ( S90314 S90215 ) S84790 S68921 S90005 S85533 S84688 S68891 Crassatoniella sp. ( sp. ( sp. ( sp. ( ? sp. ( sp. ( sp. ( sp. ( Lironoba Family: Eatoniellidae Superfamily: Rissooidea (unplaced) Pandalosia subulata ( Laseron, 1956 Pandalosia ephamilla ( 1886) (Watson, Family: Zibinidae Taxa and speciesTaxa registration number Family: Lironobidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 151 , , , 135 108 , 128, , 128, , 138 , 132, 145 , 132, , 123, 127*, 127*, 123, 136 134, 128, 143

136*

128* 106 116*, 136 116* 109*, 124, 124, 109*, 133, 128, 126, 138, 139

114* 127* 130

105

Offshore ,

, , , 165 , 157 152 159 , , , , 163, 167 , 154, , 154, , 170, 175 , 170, , 162 164 168 152

161 158, 152

150 149, 152, 153 156 162

South North

Midshelf

148

Stations

111, 113

111

111*

North

Inshore

100*

98, 100 93, 97, 98, 98, 97, 93, 100*,

100* 98

Central

91*

91* South 91* S/H S/H S/H S/H S/H S/H S/H Habitat 1 No. historical taxa

18 U

U

U U U

U U U No. survey taxa 6 0 KimMoll411 KimMoll391 KimMoll247 KimMoll473 KimMoll16 KimMoll15 KimMoll246 KimMoll175 KimMoll244 KimMoll204 KimMoll14 KimMoll377 KimMoll266 KimMoll174 KimMoll173 OTU M

M M M M M M M M M M M M M

Shell size )M ) ) )

) )

) ) ) ) ) S84773 S90178 S84515 S90137 S90249 S84771 S84552 S84551 S84768 S84716 sp. (S84686) sp. (S84685) sp. sp. (S84684) sp. sp. ( sp. (S92514) sp. ( japonicum folini folini cf. cf. sp. ( sp. ( cf. japonicum sp. ( sp. ( cf. sepimentum Anabathron Anabathron Anabathron Caecum Caecum Kisch, ( 1959 Badepigrus Anabathron Caecum Caecum Caecum Habe, 1878 ( 1878 Habe, Anabathron Caecum Caecum Caecum sepimentum Caecum ( Folin,de 1868 Caecum de Folin, 1868 ( Folin,de 1868 Habe, 1878 (S84808 1878 Habe, ? ? ? Family: Caecidae Taxa and speciesTaxa registration number Family: Anabathridae 152 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 142

124, 124, 128* 128*

123*, 124, 139 124, 123*,

123

128, 129* 145* 123*, 128 123*,

128

Offshore

South North 156, 171

151 171* 167*

152

Midshelf

Stations 111*

North

110* 111* 110 , 98, 100*, 98, Inshore

Central

93*, 96

93

91* South 91* 91* 91* 91*

S/H S/H Habitat

S/H S/H No. historical taxa 0

U U U U 16 U

U U 2 1 U No. survey taxa U U U U U U

KimMoll284 KimMoll188 KimMoll474 KimMoll436 KimMoll198 KimMoll194 KimMoll368 KimMoll138 KimMoll176 KimMoll348 OTU KimMoll471 KimMoll455 KimMoll366 KimMoll611 KimMoll603 KimMoll300 KimMoll265 KimMoll245 M M M M

M

M M

M Shell size M M M M M M M M

)M )M ) ) ) )

) ) ) ) ) ) ) ) ) S90320 S92523 S90007 S90117 S84705 ) S84687 S84844 S84699 sp. ( sp. (juvenile) (juvenile) sp. S92509 S90421 S90114 S85551 S85509 S84869 S84807 S84770 sp. ( ) sp. ( sp. ( s sp. ( s sp. sp. ( ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( S84645 Teinostoma Circulu Pseudoliotia Iravadia S84709 Circulus Circulus Circulus ? ? ( ? ? Spirostyliferina lizardensis Bandel, 2006 ( sp. ( ? Hedley, 1902 ( 1902 Hedley, Ctiloceras clathratum Caecum Caecum Caecum Caecum Caecum Caecum Caecum Caecum Family: Tornidae Family: Family: Spirostyliferinidae Family: Iravadiidae Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 153 , , , 143, , 143, 140 , 135 , 106, 109, 109, , 106, , 142, 143*, 142,

109* 144 137 133, 109* 146* 105 126 124, 123, 131 128, 129, 134, 138, 137, 140, 142, 143, 144

137* 128 106

129* , , Offshore

, , 157* 174 170 176 , , , 155, , 161, 162, 162, , 161, South North 160 167 166, 163, 168, 171

149, 151, 152, 152, 151, 149, 153 156 175,

, 148 Midshelf 146

147*

Stations North

111, 113

Inshore Central

97, 100, 114* 100, 97,

South

91* 91* 91

91* 91* 91* 91* 91*

Habitat S/H S/H S/H

S/H No. historical taxa 6

No. survey taxa U U U

H

U U 22 U U U U U U

OTU KimMoll18 KimMoll163 KimMoll158 KimMoll29 KimMoll369 KimMoll389 KimMoll42 KimMoll42 =KimMoll389 KimMoll490 KimMoll160 KimMoll540 KimMoll187 KimMoll203 KimMoll199 KimMoll186 KimMoll200 KimMoll448 M Shell size M M M M M M

M M M ) )M )M )M

)M )M )M ) S94168

) ) ) ) ) S90173 S84524 S68865 S84715 S84710 S84697 S94344 sp. ( sp. ( S94092 S84698 sp. ( sp. (juvenile) (juvenile) sp. sp. ( sp. ( sp. ( sp. (juvenile) (juvenile) sp. S84672 S84667 S84500 sp. ( sp. ( ) ) ) sp. ( sp. ( sp. ( sp. (juvenile) (juvenile) sp. Annulobalcis Hemiliostraca Hemiliostraca S68830 S84669 S84711 Eulima Eulima Lophocochlias parvissimus Lophocochlias (Hedley, 1899) ( 1899) (Hedley, ? Lophocochlias Rubio & Rolán, 2015 ( & Rolán, 2015 Rubio Lophocochlias procerus Lophocochlias Lophocochlias ( ? ? ( Vitrinella Vitrinella Pseudoliotia Pseudoliotia Pseudoliotia Pseudoliotia ( Taxa and speciesTaxa registration number Family: Eulimidae Family: 154 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 136 107* , ,

135* 128*

102, 107, 132, 132, 107, 102, 140, 139, 133, 138*, 145

130* 127* 105 105*

127

Offshore , 151, , 151, 171

167*, 170 150* 162

167, 152

South North

Midshelf

147

Stations

110

110

North 110 111

100, 115, , Inshore

100*

Central

116* 93 116*

91* 91* 91* 91* 91* 91* 91* 91*

South

91* 91

S/H

S/H S/H S/H S/H S/H Habitat S/H S/H S/H No. historical taxa U

U U U 5 4 U U

U U U U U U

No. survey taxa

KimMoll605 KimMoll555 KimMoll450 KimMoll435 KimMoll262 KimMoll164 KimMoll155 KimMoll136 KimMoll462 KimMoll165 KimMoll159 KimMoll157 KimMoll156 KimMoll153 KimMoll480 KimMoll381 KimMoll320 KimMoll290 KimMoll166 KimMoll31 OTU KimMoll30 M M ?Ma Ma Ma

M M M M M M M M M Shell size ) )M )M )M )M )M )M ) ) ) ) ) ) ) ) ) ) S92543 S84916 )MS90147 S84676 S84554 S84553 S84664 S90451 S84674 S84668 S84666 S84665 S84662 sp. ( sp. ( sp. ( sp. (S84855 sp. ( sp. ( sp. ( S90319 S84801 ) ) ) sp. ( sp. (S84889 sp. ( sp. ( S85515 S94203 S90395 sp. ( sp. ( sp. ( Cheilea Cheilea Sticteulima Pyramidelloides mirandus (S84643 Adams, 1861) (A. sp. ( Melanella sp. ( Melanella sp. ( Melanella sp. ( Melanella sp. ( Melanella sp. ( Hemiliostraca Hemiliostraca Hemiliostraca Hemiliostraca Hemiliostraca Hemiliostraca Hemiliostraca Family: Hipponicidae Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 155 , 143 128*, 130, 130, 128*, 134, 137 106, 127, 138, 138, 127, 106, 144

128*, 134

104* 105*

130* 123* 127* 105

105* 109*

Offshore 160 149, 161

159*

152, South North

Midshelf

Stations

110

North 110*

112

114* Inshore

95, 116*

114*

Central

100,

South

91* Habitat S/H S/H S/H No. historical taxa U U 2 1 U 1 3 U 1 4

U U 3 4 U U U

H No. survey taxa U 8 2 KimMoll434 KimMoll169 KimMoll433 KimMoll202 KimMoll331 KimMoll329 KimMoll296 KimMoll523 KimMoll515 KimMoll383 KimMoll319 KimMoll205 KimMoll20 KimMoll19 OTU KimMoll193 M M

?M

Ma

?M ?Ma

M M M M M M M

Shell size )M ) ) )M ) )

)

) ) ) ) ) S84717 ) S84932 S84713 S84950

S84549 S84512 S84865 sp. ( S90151 S94377 S84704 sp ( sp. ( S90317 S84680 ) ) S68997 sp. ( ( sp. ( sp. ( sp. ( cf. cuvieriana cf. gueriniana S90316 S68922 Vermetus Vanikoro Vanikoro Dendropoma Dendropoma sp.Atlanta ( sp.Atlanta ( sp. ( Trivirostra oryza Trivirostra ( (Lamarck, 1810) ? ? ? ? sp. ( ? Vanikoro Macromphalus Vanikoro Vanikoro Vanikoro ( (Recluz, 1844) Vanikoro ( (Recluz, 1844) Family: Atlantidae Family: Family: Velutinidae Family: Triviidae Family: Vermetidae Taxa and speciesTaxa registration number Family: Vanikoridae 156 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 145 , , 144*, 134 127 , 135, , 135, , 126, 145

109 145* 109 137*

142* 144

126

133* 137

Offshore 153*

175*

150, 151

168* 165 169*

176*

167*

South North

Midshelf

147

Stations

110*

110

North

Inshore

Central

91*

South 91*

91* S/H S/H S/H S/H S/H S/H

S/H Habitat S/H S/H S/H

27 No. historical taxa 22 27

U

H H

18

U

1 5

2 U No. survey taxa 1 KimMoll380 KimMoll533 KimMoll162 KimMoll530 KimMoll309 KimMoll37 KimMoll541 KimMoll36 KimMoll609 KimMoll375 KimMoll161 KimMoll538 KimMoll504 KimMoll615 KimMoll190 OTU M Me M Me M M M M Me

Ma M Me

Ma Ma

Ma

Shell size ) ) S94194 ) )

) ) )

) ) )

) ) ) ) S90130

S84701 S94021 S68882 S94498 ) S94095 S94083 S85543 S94028 S84671 S84670 S85527 atkinsoni sp. ( cf. muricata sp. ( cf. sp. ( sp. ( S90408 sp. ( Zafrona Zafra Euplica Euplica Seminella peasei Martens(von & Langkavel, 1871) ( sp.Mitrella ( cf.Mitrella rorida (Reeve, 1859) ( 1859) (Reeve, cf.Mitrella moleculina (S84897(Duclos, 1840) (Tenison-Woods, 1876) ( 1876) (Tenison-Woods, Anachis ? ? sp.Mitrella ( sp.Mitrella ( ? ? Colubraria (Lightfoot, 1786) ( (Lightfoot, 1786) Gyrineum Gyrineum sp. (juvenile) ( (juvenile) sp. Naticarius zonalis ( (Recluz, 1850) Family: Columbellidae Family: Colubrariidae Family: Ranellidae Taxa and speciesTaxa registration number Family: Naticidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 157 142 145*

135 126, 107, 124, 126,107, 130, 140 127,

109 145*

Offshore , 163, , 163,

156* 156* 165

152*

South North

168* 174 151* 151*

166* Midshelf

148

Stations North

110*

111, 112

116* , 95, 100 , , 95, Inshore Central

114

114, 93

South

91*

91*

Habitat

S/H S/H S/H

S/H

S/H S/H S/H S/H S/H No. historical taxa 84 33 U 1

4 3 U 1 8

1 No. survey taxa U

U

U KimMoll453 KimMoll571 KimMoll402 KimMoll298 KimMoll92 KimMoll334 OTU KimMoll93 KimMoll376 KimMoll60 KimMoll558 KimMoll557 KimMoll32 KimMoll552 KimMoll600 Ma

Me M M

M

Me

Shell size M Me M M M M M M ) )

S94258 S84563 ) ) ) )

) ) ) )

2)

S94322 S94306 S85528 S94195 ) S94488 S84596 S84867 ) S94229 S94228 S9419 sp. ( sp. ( cf. neburosa sp. ( sp. ( sp. (S90414) sp. sp. ( sp. ( sp. ( S84595 Conus Etrema Lienardia rubida Lienardia (Hinds, 1843) ( Lienardia lischkeana ( 1904) (Pilsbry, Etrema sp. ( Reticunassa cockburnensis Reticunassa ( 2006) & Dekker, (Kool

Zafrona ( 1860) (Gould, Zafra troglodytes in Souverbie (Souverbie ( 1866) & Montrouzier, Zafra Seminella Zafra Seminella Zafra Family: Conidae Family: Family: Clathurellidae Family: Family: Drilliidae Family: Family: Nassariidae Family: Taxa and speciesTaxa registration number 158 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 142 , , 126, 130, 130, , 126, 105*

145* 130*

140* 127*

130*

105*

140* 107 140*

Offshore , 157 , 165 , , 154, , 154, , 170

151*, 152, 151*, 158

156*

167

156*

166

161 151 156 South North

Midshelf

Stations

North

Inshore

Central

91* 91* 91*

91*

South

S/H S/H S/H

S/H S/H S/H Habitat S/H 9 No. historical taxa

H U

10 U U U H U 5 6

U

U

No. survey taxa 4 0

KimMoll324 KimMoll559 KimMoll467 KimMoll575 KimMoll476 KimMoll90 KimMoll91 KimMoll89 KimMoll519 KimMoll409 KimMoll574 KimMoll477 KimMoll477 ?=KimMoll 293 KimMoll88 KimMoll312 KimMoll464 KimMoll293 OTU Ma M M M M

M M M M M

M M M M M

Shell size ) )M

) ) )

) ) ) ) ) ) ) ) S84926 S84591 ) ) ) sp. ( S92534 S68930 S84902 S94316 S90462 S90487 S94317 S84594 S84592 S68946 S90246 S84593 S94231 S92533 sp. ( cf. minutissima sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( Kernia Exomilus Kernia Hemilienardia ocellata (Jousseaume, 1884) ( ? Rehder, 1980 ( 1980 Rehder, Microdaphne morrisoni Microdaphne ? Guraeus Eucithara Eucithara Eucithara Eucithara Carinapex Pseudodaphnella (Brazier, 1876) ( 1876) (Brazier, Pseudodaphnella barnardi Pseudodaphnella (Garrett, 1873) ( Carinapex Carinapex (Garrett, 1873) ( Carinapex minutissima Family: Raphitomidae Family: Family: Mangeliidae Family: Taxa and speciesTaxa registration number Family: Horaiclavidae SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 159 , 136 140* 109* , , 109, , 109, , , 138, , 138,

109* 109* 105 134* 137 102, 105 127 142*, 143

105, 109* 140* 139

137* 165 Offshore , , 156 152 170 , ,

165* 167

154 , 149*

154, 165 149 174

South North

Midshelf

146*

Stations 111*

North

Inshore

100*

Central

South 91* 91* S/H

S/H S/H S/H S/H

S/H S/H S/H S/H S/H Habitat 72 60 No. historical taxa

U 2

2

H H No. survey taxa 9 2 U U U KimMoll285 KimMoll597 KimMoll35 KimMoll34 KimMoll38 KimMoll41 KimMoll548 KimMoll40 KimMoll371 KimMoll251 KimMoll39 KimMoll520 KimMoll486 OTU KimMoll87 KimMoll86 KimMoll539 ?Ma Ma

Me Me M M

M M M M M M M Shell size

M M ) ) ) ) ) ) ) ) ) ) ) ) ) ) S94085 )M

S94477 ) S84529 sp. ( S94148 S68821 S84846 S84590 S84589 S92561 S90467 S90123 S84778 S84506 S68948 angasi sp. ( cf. sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( cf. coronata cf. parvu s sp. (S84504 sp. (juvenile) ( (juvenile) sp. Leptoconchus peronii (Lamarck, 1818) ( cf. philpoppei Cossignani, 2006 (S84507 Gibberula (Hedley, 1896) ( 1896) (Hedley, Gibberula Cystiscus Cystiscus Cystiscus Cystiscus (Crosse, 1870) ( 1870) (Crosse, Cystiscus Cystiscus Veprecula Veprecula Veprecula Pseudodaphnella Family: Muricidae Strigatella Family: Mitridae Family: (Lamarck, 1811) ( (Lamarck, 1811) Taxa and speciesTaxa registration number Family: Cystiscidae Family: 160 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 139 134

103*

126, 128*, 126, 128*, 144, 140, 134, 145 137 134, 131, 106, 138, 142 103*, 108, 136, 128, 127, 139, 143

128* 128* 128*,

123*, 134 123*, 137*

Offshore 174* 176* , , South North 170

162* 162*

162

156 Midshelf

148

Stations 112 , 112 111 North

110*,

110,

Inshore Central

114*

95, 99, 100* 99, 95,

91*

South

Habitat S/H

S/H S/H

S/H S/H No. historical taxa No. survey taxa

U U 4 5 U U

4 0 U U U U

U U U

9 0 OTU KimMoll537 KimMoll206 KimMoll238 KimMoll306 KimMoll438 KimMoll260 KimMoll240 KimMoll590 KimMoll589 KimMoll429 KimMoll421 KimMoll420 KimMoll127 KimMoll497 KimMoll397 KimMoll534

Shell size

M M M M M M M

)M )M )M )M )M )M )M )M )M S68899 S84762

S94077 S84718 S90200 S84887 S90323 S84799 S84764 ( sp. ( sp. ( ) ) ) ) ) ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( ) S94423 S94421 S90306 S90277 S90276 S84634 Murchisonella S94040 Dentimargo Dentimargo ? Orbitestella Orbitestella Orbitestella Orbitestella sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( Murchisonella ? (Gray in G.B. Sowerby I., 1832) 1832) I., Sowerby in G.B. (Gray ( Eratoena sulcifera Taxa and speciesTaxa registration number Family: Marginellidae Family: Orbitestellidae Family: Murchisonellidae Family: SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 161 , , , , 138 , , 110 , 105* 108 134* 140 136 108, , , , , , , 123, 126 , 123, ,

123, 124, 128* 124, 123, 133 130, 129, 134 108* 133 134* 107 102 106 102* 109* 129, 129, 130, 130, 129, 129, 134, 109 105* 124, 151

126, 127*

124, 126, 130, 130, 126, 124, 139 128*, 130, 134 139* 162 176 Offshore , , , , 161 159 174* , 165, , 165, 162 , 167, 175 , 167, , , 164 , 162 , South North 153,

151 155 167

164 153, 149

165* 164* 175

Midshelf

147, 148147, 146

148*

146

Stations North

Inshore Central

93*

South

Habitat S/H

S/H S/H S/H S/H S/H S/H S/H S/H

S/H No. historical taxa U No. survey taxa

5 0

H

H U

5 2 1 2 7 0 H

U OTU KimMoll425 KimMoll492 KimMoll535 KimMoll493 KimMoll352 KimMoll344 KimMoll318 KimMoll10 KimMoll43 KimMoll46 KimMoll403 KimMoll349 KimMoll416 KimMoll598 KimMoll489 KimMoll431 M

Shell size M M M M M M M M M M

)M )M )M ) )M )M ) ) ) ) ) ) ) ) ) ) S90291 S90437 S94480 S94449 S90311 S84508 S84544 S90211 S90009 S68833 S94042 S68834 S90026 S84988 S84909 sp. ( S90260 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. group) ( sp. ( Bullina confusa Rissoella Rissoella Rissoella Rissoella Rissoella Rissoella Ammonicera Acteocina Acteocina Acteocina ? Rissoella ( Ammonicera Ammonicera Ammonicera Taxa and speciesTaxa registration number Family: Omalogyridae Family: Cylichnidae Family: Aplustridae Family: Rissoellidae Family: 162 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE 139 109*

134, 139* 139*

123*

126, 127 123*, 127, 128 127, 123*, 103*, 128 124, 144

114*, 138

129*, , 155 Offshore , 170* 153

165 165,

149, 162 159, 165, 170 165, 159,

South North

167*

171*

Midshelf

Stations

111*

North 111*

Inshore

93, 95*, 114 95*, 93,

114* Central 116

South

91*

S/H

S/H S/H Habitat S/H

S/H No. historical taxa U U 2 3 U

2 1

U

U U U No. survey taxa 7 7 U 1 8 U 1 0 U 1 3 U

KimMoll47 KimMoll232 KimMoll459 KimMoll456 KimMoll364 KimMoll367 KimMoll384 KimMoll208 KimMoll64 KimMoll412 KimMoll233 OTU KimMoll602 KimMoll191 KimMoll226

KimMoll612 KimMoll468 M

?Ma ?Ma

M M M M M M ?Me Shell size

Ma

M

Ma M M ) )M ) S84755 ) ) ) ) ) ) ) ) ) ) ) ) ) S84536 S90004 s sp. ( s sp. S84743 S68912 S85575 S90115 S84721 S90499 S84702 S84819 S90104 S90440 S90424 sp. ( S85500 S85556 sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( cf. quadridentata sp. ( Philine Philine Berthellina Phaneropthalmu Diacria ( (Blainville, 1821) Diacavolinia ? ? Mnestia Limulatys Limulatys Limulatys Diniatys Haminoea ? Aplysia Ringicula ? Retusa Acteocina Family: Cavoliniidae Family: Philinidae Family: Taxa and speciesTaxa registration number Family: Haminoeidae Family: Aplysiidae Family: Ringiculidae Family: Pleurobranchidae Family: SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 163 , 109 , 144 , 105*, , 105*, , 145 100, 126, 145 126, 100, 103, 126 138*

102* 136,

131* 109*

109* 116* 102 143 124*

Offshore 174 ,

154, 156, 157

South North

Midshelf

Stations

110

North

111

100 , 99 Inshore

100*

Central

93, 97,

100*

91* 91*

91* 91* 91* 91*

91* South

91*

S/H S/H Habitat S/H S/H

S/H S/H

No. historical taxa 25 20 U U

U U U U U

U U

U U 47

No. survey taxa U

1 KimMoll140 KimMoll139 KimMoll461 KimMoll150 KimMoll149 KimMoll131 KimMoll180 KimMoll261 KimMoll44 KimMoll132 KimMoll408 KimMoll13 KimMoll59 KimMoll154 KimMoll257 KimMoll48 OTU KimMoll325 KimMoll325 M M M M M M M M M M M M

Me

Shell size )

)M )M ) )M )M )M ) ) ) ) ) ) )

) )

) ) S84638 S84927 S68813 S84691 S90446 S84659 S84658 S90231 S84562 S84527

S84639 S84992 S84794

sp. ( sp.( S84534 S84800 S84663 S84647 S84646 sp. ( sp. ( sp. ( cf. tenera cf. setoutiensis sp. ( sp. ( sp. ( sp. ( sp.Clathrella ( Brachystomia Miralda Megastomia Megastomia Megastomia (A. Adams, 1860) ( Adams, 1860) (A. (Nomura, 1939) ( 1939) (Nomura, Megastomia Hinemoa ligata ( 1877) (Angas, Hinemoa duplex Hinemoa ( Laseron, 1959 (Hedley, 1907)(Hedley, ( Herewardia kesteveni Herewardia articulata Besla ( 1909) (Hedley, ? ? zea ( 1902) (Hedley, ? ? sp.Rostanga ( Family: Pyramidellidae Taxa and speciesTaxa registration number Family: Discodorididae 164 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE , 136 138 130 , 126*, 143

132, 145

126

128,

128*

128, 143

138 102*

, Offshore , 164 170 , 153 , 176

151*, 149*

159*

156, 158, 168, 175

162*

151

South North Midshelf

Stations

110

111

North

111*

Inshore

116* 98, 100, 116 100, 98, 97* Central 114

100*

91 91* 91* 91*

91* 91* 91* 91* 91*

91* 91 South 91

S/H S/H S/H S/H S/H Habitat S/H No. historical taxa U

H U U U

U

U U U

U

U U

U U No. survey taxa KimMoll544 KimMoll561 KimMoll144 KimMoll147 KimMoll146 KimMoll145 KimMoll584 KimMoll143 KimMoll142 KimMoll179 KimMoll178 KimMoll151 KimMoll428 KimMoll201 KimMoll58 KimMoll71 KimMoll358 KimMoll231 KimMoll272 OTU KimMoll254 M M M M M M M M M M M M M M M M M M M M Shell size

)

) )

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

S90303 S94127 S94374 S84690 S84689 S94233 S94157 S84656 S84655 S84654 S84561 S84712 microscopica S84652 S84651 S94434 S90043 S84993 S84820 S84788 S84660 sp. ( sp. ( sp. ( aff. aff. sp. ( sp. ( sp. ( sp. ( cf. introspecta sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( microscopica Turbonilla ( Laseron, 1959 Pyrgulina Pyrgulina Pyrgulina Paregila Paregila Turbonilla ( Laseron, 1959 Syrnola Odostomia opaca ( 1906) (Hedley, Hedley, 1907 ( 1907 Hedley, Moerchia Miralda umeralis ( 1902 Hedley, Miralda Miralda Miralda Miralda Miralda Taxa and speciesTaxa registration number SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 165 , 138, , 138, 134 ,

109* 109* 128 142* 105* 123, 128

126*

135 137

123* Offshore

150*

South North

Midshelf

Stations

North

112

112*

Inshore

Central

97

97

South 91* 91*

91* 91* 91* 91* 91* 91* 91* 91 Habitat

S/H

S/H S/H S/H No. historical taxa 10 U U 2

U

2 0

3 0

U U U U U U U U No. survey taxa U KimMoll84 KimMoll85 KimMoll45 KimMoll33 KimMoll361 KimMoll311 KimMoll354 KimMoll556 KimMoll543 KimMoll152 KimMoll141 KimMoll134 KimMoll130 KimMoll128 KimMoll126 KimMoll125 OTU KimMoll129 M Ma

M M

M M M

M M M M M M M M M Shell size M ) ) ) ) ) ) ) ) ) ) ) ) ) ) S84588

S84901 S94214 S94118 S84661 S84649 S84641 S84637 S84635 S84633 S84632 S68913 sp. ( S90033 S90127

) ) sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( sp. ( ) exquisita cf. S84523 S84532 S84587 Gadila brycei Gadila 1998 Lamprell & Healy, ( Fissidentalium sp. ( sp. ( Julia ( 1862 Gould, Berthelinia Berthelinia darwini Berthelinia Jensen, 1997 ( Jensen, 1997 Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Turbonilla Family: Dentaliidae Family: Class: Scaphopoda ?Family Family: Juliidae Family: Taxa and speciesTaxa registration number 166 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

SIGNIFICANT RECORDS AND SPECIES (KimMoll80, WAM S84583) (Figure 5E) is recorded for the frst time in the Kimberley (Middelfart The diversity captured in this study includes 2002b). Warrana triangulata Middelfart, 2002 was 102 family or superfamily-level taxa. Twenty- described from single valves from the Joseph one of these familial level taxa were new to the Bonaparte Gulf, NT, and its distribution is here survey area. These and other signifcant records extended to WA (Figure 5F). The range of Warrana and species are discussed below to contextualise brucemarshalli Middelfart, 2002 (Figure 5G) is the fndings. Size category data, condition (live or dead) and all references to images are reported in extended from North West Cape to the northern Table 2. offshore Kimberley.

Class Superfamily Galeommatoidea

Subclass Pteriomorphia The illustrated specimen (KimMoll483, WAM S92587) (Figure 5H) is most likely a new and Superfamily Limopsoidea species. Mylitta sp. (KimMoll531, WAMS 94062) is Family Philobryidae the frst record of any Mylitta from north-western WA and does not match any described Mylitta from Cosa sp. (KimMoll442, WAM S90346) (Figure 5A) Australia. has until now only been recorded from caves in the Ryukyu Islands, Japan, and is still undescribed (see Superfamily Veneroidea Hayami and Kase 1993: 35, fgs 100–102). Family Neoleptonidae Superfamily Pectinoidea Six species (KimMoll69, WAMS90108 (Figure 6A); Family Propeamussidae KimMoll 83, WAMS84586 (Figure 6B); KimMoll106, WAMS84612 (Figure 6C); KimMoll250, WAMS84777 Parvamussium araneum Djikstra, 1991 (Figure 6D); KimMoll426, WAMS90293 (Figure (KimMoll506, WAM S94109) (M) (Figure 5B) has 6E) and KimMoll522, WAMS68990 (Figure 6F) not previously been recorded from Australia. It were found and are most likely new species (P. is most likely that this discovery represents a Middelfart, unpublished revision of Australian Parvamussium new record and not a new species. Neoleptonidae). araneum Djikstra, 1991 was described from Indonesia based on material from 130–300 m Class Gastropoda depth. The specimen is a juvenile, about 2 mm maximum diameter, while the size of the Subclass Indonesian type material is around 9 mm. Superfamily Subclass Heterodonta Family Liotiidae Superfamily Carditoidea Bathyliotina schepmani (Habe, 1953) (KimMoll466, Family Condylocardiidae WAM S90485) (Figure 6G) from station 145/K13 is the frst record of this genus and species from Condylocuna tricosa Middelfart, 2002 (Figure Australia (ALA, AFD). 5C) was described from Qld and synonymised with Condylocuna io (Barsch, 1915) by Graham and Holmes (2004) despite Middelfart (2002) Family Skeneidae comparing C. tricosa with the South African type Lodderena ornata (Olsson and McGinty, 1958) of C. io (United States National Museum (USNM (KimMoll243, WAM S84767) (Figure 7A) is 251043) and stating the dissimilarity. Oliver recorded for the frst time in the central Indo-west and Holmes (2004) did not examine the type Pacifc Ocean. There is doubt whether the Indo- of C. io, and compared it only to material from Rodrigues Island, which is C. tricosa and not C. Pacifc taxon is conspecifc with the Atlantic taxon io. Hence, the synonymy is here revoked and the that comprises the type species (Panama, West distribution of C. tricosa is extended across the Indies) (see Lima et al. 2011 and Rubio et al. 1998). tropical Indo-Pacifc region from Qld to Rodrigues Leucorhynchia caledonica Cro s s e, 1867 Island and the Kimberley region. Crassacuna sp. (KimMoll567, WAM S94284) (Figure 7B) and (KimMoll230, WAM S84747) (Figure 5D) has not Leucorhynchia tricarinata (Melvill and Standen, been recorded before (Middelfart 2002b) and is 1896) (KimMoll579, WAM S94333) (Figure 7C) are likely new. Crassacuna praecalva (Hedley, 1909) both new records for WA. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 167

Family Trochidae Subclass

Alcyna ocellata A. Adams, 1860 (KimMoll370, Superfamily Orbitestelloidea WAM S90121) (Figure 7D) is recorded for the frst Family Orbitestellidae time from Australia (ALA). Four Orbitestella spp. were found in the survey area, all of which are new species, viz., Orbitestella Subclass sp. (KimMoll240, S84764) (Figure 8E), Orbitestella Superfamily Triphoroidea sp. (KimMoll260, S84799) (Figure 8F), Orbitestella sp. (KimMoll306, WAM S84887) (Figure 8G) Family Triphoridae and Orbitestella sp. (KimMoll438, WAM S90323) (Figure 8H). Three OTUs are considered to be related to the corresponding species: Superfamily OTU KimMoll581, (WAM S94346) (Figure 7E) aligns with Mastonia lamberti (Hervier, 1898). Family Omalogyridae OTU KimMoll341, (WAM S84984) (Figure 7F) aligns Five species of Ammonicera (KimMoll43, WAM with Monophorus tessellatus (Kosuge, 1963). S90437 (Figure 9H); KimMoll425, WAM S90291 (Figure 9I); KimMoll431, WAM S90311 (Figure OTU KimMoll388, (WAM S90168) (Figure 7G) aligns 9J); KimMoll489, WAM S94449 (Figure 9K); with Monophorus atratus (Kosuge, 1962). KimMoll598, WAM S94480 (Figure 9L) were found These are new records for WA. in this study and all are new species.

Family Cerithiopsidae Superfamily Rissoelloidea

Paraseila sp. (KimMoll572, WAM S94307) (Figure Family Rissoellidae 7H) is the frst record of the genus in WA. There are only three species of this genus recorded from Previously no species of this family had Australia (ALA, AFD), with records from New been reported from Exmouth to the NT border. South Wales (NSW), Qld and Tasmania. However, at least seven new species were found in this study including Rissoella sp. (confusa group) Superfamily (KimMoll10, WAM S84508) (Figure 9A), Rissoella sp. (KimMoll318, WAM S84909) (Figure 9B), Rissoella Family Epitoniidae sp. (KimMoll344, WAM S84988) (Figure 9C), Rissoella sp. (KimMoll352, WAM S90026) (Figure Cycloscala revoluta (Hedley, 1899) (KimMoll350, 9D), Rissoella sp. (KimMoll492, S68833) (Figure 9E), WAM S68981) (Figure 8A) is a new record for WA, Rissoella sp. (KimMoll493, WAM S68834) (Figure although it is known from the Indo-west Pacifc 9F) and Rissoella sp. (KimMoll535, WAM S94042) Ocean. (Figure 9G).

Superfamily Rissoidea Superfamily Pyramidelloidea

Families Rissoidae and Rissoinidae Family Pyramidellidae

Haurakia marmorata (Hedley, 1907) (KimMoll532, Chrysallida zea (Hedley, 1902) (KimMoll13, WAM WAM S94025) (Figure 8B) and Parashiela liddelliana S84527) (Figure 9N), Herewardia kesteveni (Hedley, (Hedley, 1907) (KimMoll242, WAM S84766) (Figure 1907) (KimMoll408, WAM S90231) (Figure 9O), 8C) are new records for WA. Odostomella opaca (Hedley, 1906) (KimMoll428, WAM S90303) (Figure 9P), Turbonilla microscopica Laseron, 1959 (KimMoll561, WAM S94233) (Figure Superfamily Capuloidea 9Q) and Besla articulata (Hedley, 1909) (KimMoll59, Family Capulidae WAM S84562) (Figure 9R) are all new records for WA. The OTU KimMoll201 (WAM S84712) (Figure Trichamathina sp. (KimMoll287, WAM S84848) 9M) aligns with Morchia introspecta (Hedley, 1907), (Figure 8D) is a new record for Australia which has not previously been recorded from (AFD, ALA). Australia. 168 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

FIGURE 5 New species and records. A) Cosa sp. (KimMoll442, WAM S90346), new species; B) Parvamussium cf. araneum Djikstra, 1991 (KimMoll506, WAM S94109), new record Australia; C) Condylocuna tricosa Middelfart, 2002 (KimMoll310, WAM S84900), new record WA; D) Crassacuna sp. (KimMoll230, WAM S84747), new species; E) Crassacuna praecalva (Hedley, 1909) (KimMoll80, WAM S84583), new record in survey area; F) Warrana triangulata Middelfart, 2002b (KimMoll100, WAM S84604), new WA record; G) Warrana brucemarshalli Middelfart, 2002b (KimMoll76, WAM S84579), new record for Project Area; H) Galeommatoidea (KimMoll483, WAM S92587), new species and genus. Scale bars 500µm. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 169

FIGURE 6 New species and records. A) Neolepton sp. (KimMoll69, WAM S90108), new species; B) ?Neoleptonidae (KimMoll83, WAM S84586), unknown species; C) ?Micropolia sp. (KimMoll106, WAM S84612), new species; D) ?Neoleptonidae (KimMoll250, WAM S84777), unknown species and genus; E) Neolepton sp. (KimMoll426, WAM S90293), new species, no scale; F) ?Micropolia sp. (KimMoll522, WAMS68990), unknown species; G) Bathyliotina schepmani (Habe, 1953) (KimMoll466, WAM S90485), new Australian record. Scale bars 500µm. 170 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

FIGURE 7 New species and records. A) Lodderena ornata (Olsson and McGinty 1958) (KimMoll243, WAM S84767), new Australian record; B) Leucorhynchia caledonica Crosse, 1867 (KimMoll567, WAM S94284), new WA record; C) Leucorhynchia tricarinata Melvill and Standen, 1896) (KimMoll579, WAM S94333), new WA record; D) Alcyna ocellata Adams, 1860 (KimMoll370, WAM S90121), new Australian record; E) Mastonia cf. lamberti (Hervier, 1898) (KimMoll581, WAM S94346) new record for Australia if verifed; F) Monophorus cf. tessellatus (Kosuge, 1963) (KimMoll341, WAM S84984), new record for Australia if verifed; G) Monophorus cf. atratus (Kosuge, 1962) (KimMoll388, WAM S90168) new record for Australia if verifed; H) Paraseila sp. (KimMoll572, WAM S94307), frst record of Paraseila in WA. Scale bars 500µm. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 171

FIGURE 8 New species and records. A) Cycloscala revoluta (Hedley, 1899) (KimMoll350, WAM S68981), new record for WA; B) Haurakia marmorata (Hedley, 1907) (KimMoll532, WAM S94025), new record for WA; C) Parashiela liddelliana (Hedley, 1907) (KimMoll242, WAM S84766), new record for WA; D) Trichamathina sp. (KimMoll287, WAM S84848), new record for Australia; E) Orbitestella sp. (KimMoll240, S84764), new species; F) Orbitestella sp. (KimMoll260, S84799), new species; G) Orbitestella sp. (KimMoll306, WAM S84887), new species; H) Orbitestella sp. (KimMoll438, WAM S90323), new species. Scales 500µm, E–H share the same scale. 172 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

FIGURE 9 New species and records. A) Rissoella sp. (confusa group) (KimMoll10, WAM S84508); B) Rissoella sp. (KimMoll318, WAM S84909); C) Rissoella sp. (KimMoll344, WAM S84988); D) Rissoella sp. (KimMoll352, WAM S90026); E) Rissoella sp. (KimMoll492, S68833); F) Rissoella sp. (KimMoll493, WAM S68834). G) Rissoella sp. (KimMoll535, WAM S94042); H) Ammonicera sp. (KimMoll43, WAM S90437); I) Ammonicera sp. (KimMoll425, WAM S90291); J) Ammonicera sp. (KimMoll431, WAM S90311); K) Ammonicera sp. (KimMoll489, WAM S94449); L) Ammonicera sp. (KimMoll598, WAM S94480); M) Moerchia cf. introspecta Hedley, 1907 (KimMoll201, WAM S84712); N) Chrysallida zea (Hedley, 1902) (KimMoll13, S84527); O) Herewardia kesteveni (Hedley, 1907) (KimMoll408, WAM S90231); P) Odostomella opaca (Hedley, 1906) (KimMoll428, WAM S90303), not to scale; Q) Turbonilla microscopica Laseron, 1959 (KimMoll561, WAM S94233); R) Besla articulata (Hedley, 1909) (KimMoll59, WAM S84562). A–L new species, M–R new records for WA. Scale 500µm, A–L share the same scale. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 173

TAXONOMIC DISCUSSION Superfamily Limoidea

Family Limidae Class Bivalvia Micro-limids represented about half of the Subclass Protobranchia diversity compared to the Kimberley historic Superfamily Nuculoidea mollusc paper (fve in this study versus nine in Willan et al. 2015). Family Nuculidae

The Nuculidae include the world’s smallest Superfamily Pterioidea mollusc measuring only 400 μm (Anders Warén personal communication); see Bergmans (1978) and Families Pteriidae and Malleidae Huber (2010) for recent overviews. Four species were found in this study. This low diversity is likely The sampled pteriids and malleids were mainly due to the focus on hard bottom reef areas during juveniles or spat and sorting was halted due to the the Kimberley survey. diffculty of separating dead from live material at the species level and linking dead shells to live collected material for these families. Subclass Pteriomorphia

Superfamily Arcoidea Superfamily Pectinoidea Family Arcidae Family Propeamussidae Twelve species were found. While some of the The propeamussiids consisted of three species recorded were juvenile arks, this reduced morphospecies. The material of two of them number of species illustrates the bias towards consisted of left valves only, while the third sampling of larger species during earlier surveys and their partial affliation with softer sediments species was represented by a single right valve. (e.g. Anadara lives in soft sediments, as opposed to It is possible that these three morphospecies Barbatia which is found attached with a byssus to represent growth stages of a single inequivalve hard substrates). species (see Cyclopecten ryukyuensis (Hyamy and Kase, 1991 in Okutani 2000: 915, pl. 455, fg. 16), but unless a full specimen with right and left Family Philobryidae valves intact is found, the link between the valves The philobryids are a diverse assemblage of very cannot be made with certainty. These records are small flter feeding pteriomorphs, taxonomically signifcant as their small size rules out any affnity covered in Laseron (1953) and Tevesz (1977). These with Amusium spp., the only closely related taxon very small mussel-shaped bivalves cling to hard reported in the Kimberley historic mollusc paper substrates with a byssus and are particularly (Willan et al. 2015). abundant in caves. They have a characteristic embryonic shell resembling a small hat and are Family Spondylidae direct developers. Seven morphospecies were found in this study, most from offshore stations. Only two juvenile spondyliids were found. In No member of this family was reported in the contrast, Willan et al. (2015) recorded 13 species of Kimberley historic mollusc paper (Willan et al. Spondylus. 2015) and there are no records from northern WA (ALA, AFD). Subclass Heterodonta

Superfamily Mytiloidea Superfamily Carditoidea

Family Mytilidae Family Carditidae

Most of the mytilids recorded were juvenile or The carditiids include some truly microscopic small specimens from a common macro species, species (Table 2), one of which, Cyclocardia sp.: including Septifer cumingii Récluz, 1849 (KimMoll52, KimMoll560, WAM S94232, was found only at WAM S68867). Rowley Shoals stations (see Bryce et al. 2018). 174 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

Family Condylocardiidae (Huber 2015), the superfamily is under revision. There are currently 14 families (Middelfart in prep.), The condylocardiids were represented by the hence molecular systematics are needed to clarify two subfamilies, Condylocardiinae and Cuninae, relationships (Goto et al. 2012; Li et al. 2012). both reviewed by Laseron (1953) and Middelfart (2002a,b). A total of 15 species were found in this study, with seven of the most common observed Superfamilies Ungulinoidea, at Station 91/K12. At least three new species Gastrochaenoidea, Mactroidea (KimMoll230, KimMoll457 and KimMoll501) and Myoidea and possibly two new genera (KimMoll457 and KimMoll502) await formal description. Only species The micro-gastrochaenids were represented KimMoll230 and KimMoll76 were found more than only from inshore stations (110/K12 and 116/K12) once, illustrating the rarity of this group. Only two by live material, while micro-representatives of specimens of two species were taken alive, Warrana mesodesmatids, corbulids, myids, pholadids and brucemarshalli Middelfart, 2002 (KimMoll76) and ungulinids were all dead collected in low numbers ?Crassacuna (KimMoll582). and some are most likely juvenile.

Family Crassatellidae Superfamily Veneroidea

Three crassatellid species were found at offshore Family Neoleptonidae stations (134/K13, 153/K14, and 175/K14) (Table 2) The neoleptonids are under revision (Middelfart and none overlapped at the species level with those recorded in the Kimberley historic mollusc paper in prep.), but include three genera, Neolepton, (Willan et al. 2015). Micropolia and Puysegeria (the latter ?Neoleptonidae Middelfart 2002: 79). KimMoll83 might be a juvenile venerid and KimMoll250 could be affliated with Superfamilies Hiatellolidea, cyamiids, but with a transposed hinge. Hinge Tellinoidea, Lucinoidea, Arcticoidea transposition is known from lower heterodonts and Veneroidea (Cox et al. 1969). While the micro-hiatellids were equal in species numbers to those reported in the Kimberley Class Scaphopoda historic mollusc paper (Willan et al. 2015), the semelids, tellinids, lucinids, trapezids and venerids Lamprell and Healy (1998) revised the Australian were far more prevalent in the Kimberley historic scaphopods. Two species were encountered; Gadila mollusc paper (Willan et al. 2015), because of their brycei Lamprell and Healy, 1998 (KimMoll84, typically large adult size making them conspicuous WAM S84587) and a juvenile Fissidentalium sp. in the feld. (KimMoll84, WAM S84587), both from Station 91/K12. Superfamily Chamoidea Class Polyplacophora Family Chamidae Minute polyplacophorans were found The micro-chamiids were all juvenile. In contrast, the Kimberley historic mollusc paper by Willan et predominantly from offshore stations 109/K12, 127/ al. (2015) recorded fve species of Chama. K13, 136/K13 and 137/K13 (Table 2). All belong in the Leptochitonidae. Superfamily Galeommatoidea Class Gastropoda The galeommatoideans are a challenging superfamily with either a complete lack of Many representatives of this class were characters or a myriad of unique ones (Goto et al. obtained during this project. Table 2 provides 2012), making resolution of relationships diffcult. records of Hipponicidae, Vermetidae, The superfamily was represented by 20 species Triviidae, Velutinidae, Atlantidae, Ranellidae, in this study, mainly montacutids (e.g. Mysella Colubrariidae, Columbellidae, Drillidae, Conidae, spp.). This material represents two new species Murchisonellidae, Aplustridae, Cylichnidae, (KimMoll517 and KimMoll483) and KimMoll517 is Pleurobranchidae, Ringiculidae, Aplysiidae, also a new genus. While there is a recent overview Philinidae, Cavoliniidae, and Discodorididae. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 175

Subclass Patellogastropoda research is needed to clarify conspecifcity. This species formed part of the numerous, possibly Only one patellogastropod limpet species was congeneric, series that also includes the less found, and it was only collected from offshore planispiral Lodderena sp. (KimMoll237, WAM stations 134/K13, 139/K13, 143/K13 and 169/K14 S84760). Two Leucorhynchia spp. (Leucorhynchia (Table 2). caledonica Crosse, 1867 and Leucorhynchia tricarinata Melvill and Standen, 1896) were recovered from Subclass Vetigastropoda the southern offshore stations (154, 156, 161 and 165/K14), and both are new records for WA (ALA). Superfamily Scissurelloidea The trochids and turbinids form complicated assemblages, which are challenging to place as Family Scissurellidae either juvenile macro or true micromolluscs. Although diverse in the 0–30 m depth range However, Alcyna ocellata A. Adams, 1860 is (Geiger 2012, Figure 26), only fve species were signifcant and is recorded for the frst time from identifed in the study. This represents less than Australia (ALA, AFD). 10% of the Indo-Malayan Archipelago’s diversity (sensu Geiger 2012). The most common species Superfamily Sequenzoidea was Scissurella sp. (KimMoll4) identifed from 25 stations, while the less common Sukashitrochus A minute and characteristic seguenzioidean atkinsoni (Tenison-Woods, 1877) (KimMoll404) was species of the genus Brookula (KimMoll415, WAM identifed from fve stations. The least common S90258) was found exclusively offshore. The species, Scissurella sp. (KimMoll577) was only superfamily was not recorded in the Kimberley identifed from one station (Station 156/K14). historic mollusc paper (Willan et al. 2015) as the paper only included molluscan holdings housed Superfamily Fissurelloidea in the WAM and Museum and Art Gallery of the Northern Territory from the Project Area; however, Family Fissurellidae specimens from the Kimberley do exist in other institutions. The fissurellids were mostly juveniles or broken shells of larger species, with KimMoll299 (Puncturella sp.) being the only true micro- Subclass Caenogastropoda fssurellid discovered during the study. Superfamily

Superfamily Haliotoidea Cerithidium perparvulum (Watson, 1886) was one of the most common micromolluscs in the study. KimMoll460 (Ma category) was the only species The dialids were represented by fve species, with of haliotid, represented by juvenile shells from Diala albugo (Watson, 1886) the most common. stations 139/K13 and 150/K14. The litiopids belong in the Cerithioidea. Houbrick (1987) reviewed the anatomical differentiation between genera. There is no extensive review of Superfamily Trochoidea Australian taxa. and family Turbinidae Despite the shallow water focus of this study, The only calliostomatids recovered were from all of the live collected Styliferina cf. goniochila A. Station 91/K12 (Table 2). The six species of liotiids Adams, 1860 and S. cf. translucida (Hedley, 1906) recorded were Austroliotia sp. and Bathyliotina were found in depths of 10 m or more at offshore schepmani (Habe, 1953) from Station 145/K13, stations, where deeper light penetration might have with the latter a new record for Australia. The facilitated growth of algae and seagrass. phasianellid genus Tricolia was represented by fve Members of the Pickworthiidae are close relatives species and Gabrielona was represented by two of the familiar periwinkles often seen on rocky species. Gabrielona sp. (KimMoll494, S68836) might shores and in mangrove trees, but are very small be conspecifc with Gabrielona pisinna Robertson, and live hidden in rubble, cracks and crevices or 1973 once more material is examined. caves in deeper water. Nothing is known about Thirteen species of skeneids were found, with the biology of these . The pickworthiids the planispiral species Lodderena ornata (Olsson and were generally found offshore and while the Indo- McGinty, 1958) recorded for the frst time from the Pacifc Astrosansonia dautzenbergi (Bavay, 1917) is central Indo-west Pacifc. Previously considered noteworthy, Sansonia sp. (KimMoll452) was the an Atlantic species; however, some Red Sea and most prevalent in offshore material. None of the Pacifc records do exist (Lima et al. 2011), and more scaliolids (Scaliola spp.) were encountered alive, 176 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE only the larger and more common Finella sp., a KimMoll388, which aligns with Monophorus cf. genus also recorded in the Kimberley historic atratus (Kosuge, 1962). Monophorus episcopalis mollusc paper (Willan et al. 2015), was found in the (Hervier, 1898) is recorded from northern Australia micromollusc samples. (Chang and Wu 2005 through AFD) and ALA has records from New Caledonia, therefore KimMoll463 is tentatively attributed to this taxon, Superfamily Triphoroidea forming a starting point for exploring this species Members of the Cerithiopsidae are very close through vouchered Australian material. The taxon relatives of the triphorids. Of the 16 species of identifed as Opimaphora cf. sarcira Laseron, 1958a, cerithiopsids, none were found inshore with while resembling that species in coloration and live material only found from offshore stations. shape, differs in having weaker, less nodulose However, new data have revealed this group spirals and a with two spiral ridges does occur in the inshore Kimberley with a single and axial ribs. Subulophora rutilans (Hervier, 1897) record from Bonaparte Archipelago (Middelfart et is most likely a widely distributed Indo-Pacifc al. unpublished). The group is in need of revision, species (ALA, AFD, Marshall 1983) and aligns well with only Laseron (1956a) and the Philippine with KimMoll24. Viriola cancellata (Hinds, 1843) has work by Cecalupo and Perugia (2012) providing an Indo-Pacifc wide distribution and is known taxonomic works useful for identifcation purposes. from the Queensland coast, Cocos (Keeling) Islands Laseron (1958a) was the last researcher to assess (AFD) and Ashmore Reef (ALA, Museum and Art the northern Triphoridae taxonomically. The Gallery of the Northern Territory records) and south Australian triphorid fauna was reviewed by is reported from stations 109/K12, 154/K14 and Marshall (1983) and is better known, and deep- 174/K14. sea species have been recently studied (Fernandes and Pimenta 2017). Forty triphorid species were Superfamily Epitonoidea recorded in this study compared to 11 in the Epitoniids or are circumglobal, Kimberley historic mollusc paper (Willan et al. but particularly diverse in the central Indo-west 2015). This is to be expected given that most species Pacifc (Weil et al. 1999). This study found two are diminutive (Willan et al. 2015). morphospecies in Cycloscala Dall, 1889, including Costatophora granifera (Brazier, 1894) has several the live collected Cycloscala revoluta (KimMoll350). synonyms, two of which are from Thiele’s work Both species were found at inshore stations 91/ on Western Australian molluscs (see Thiele, K12 and 93/K12 and at offshore stations 128/K13 1930 and Albano and Bakker, 2016) and the rest and 130/K13. The entire group are specialised are from NSW (Laseron, 1954). Based on the cnidarian predators, many on the shade-loving material observed, the morphological variation hard coral, Tubastrea. Cycloscala revoluta (Fig. 8A) is substantial and might collapse the various was found living in less than 20 m in a habitat with morphs of Costatophora found in this study caves, vertical coral walls and cnidarians, and was (KimMoll21, KimMoll213, KimMoll217). Iniforis the only live epitoniid micromollusc found during progressa (Laseron, 1958a) was described from the entire study. Garcia (2004) reviewed the genus Christmas Island, with other records from WA, Cycloscala. Eleven additional epitoniid species were while Okutani (2017) described a distribution collected from Station 91/K12. ranging from Japan to northern Australia. The OTU KimMoll581 (WAM S94346) aligns with Mastonia Superfamily lamberti (Hervier, 1898), which has not previously been recorded from Australia (AFD and ALA). The family Plesiotrochidae was mainly Mastoniaeforis decorata (Laseron, 1958a) is also represented by Plesiotrochus unicinctus (A. Adams, described from Christmas Island, but aligns with 1853) from offshore sites, which matches previous taxon KimMoll570 from Station 154/K14. Metaxia records from the same area (ALA). Another possible fuscoapicata Thiele, 1930 is recorded from north of aberrant form was found, but may be a worn Exmouth and off Lake Macleod, WA. The range is specimen (KimMoll503). thus extended northeast to the Kimberley survey area. The taxa KimMoll373 and KimMoll341 align Superfamily Rissoidea with Monophorus tessellatus (Kosuge, 1963), which has not previously been recorded from Australia Generic identifcation of Rissoidae/Rissoinidae and are most likely that species, which follows was facilitated by Ponder (1985) and the family has Okutani (2000, 2017). The same can be said for been addressed (Criscione and Ponder 2013), with SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 177 the rissoids recovered as polyphyletic. The genus Layton et al. (2019) transferred Spirostyliferina Rissoina was given familial status by Criscione lizardensis Bandel, 2006 from the Litiopidae to the and Ponder (2013) based on molecular research. new genus Spirostyliferinidae. Apataxia cerithiiformis (Tryon, 1887) was quite The tornids are represented by 16 species with common offshore and has been recorded previously most records from offshore sites. Lophocochlias from the Kimberley (ALA). Although Haurakia sp. was the most prevalent species encountered. marmorata (Hedley, 1907) was very common at The systematic position of Lophocochlias has been southern offshore stations, no record was found in clarifed for representatives from this study. the historical records (ALA) from WA or NT. The congener H. novarensis (Frauenfeld, 1867) was only Family Tornidae found from northern offshore stations, but appears to be present from the inshore Pilbara (ALA and Genus Lophocochlias Pilsbry, 1921 P. Middelfart, personal observation). Parashiela Haplocochlias (Lophocochlias) Pilsbry, 1921: 377. Type cf. invisibilis (Hedley, 1899) was one of the most species Haplocochlias (Lophocochlias) minutissimus common molluscs, but only found alive offshore. Pilsbry, 1921: 377 (monotypy), syn. of Lophocochlias The congener P. liddelliana (Hedley, 1907) was parvissimus (Hedley, 1899). also recorded offshore, but at fewer stations, and is a new record for WA. Three additional species Aqabarella Alhejoj, Bandel and Al-Najjar, 2016:418. (KimMoll508, KimMoll305 and KimMoll514) also Type species: Aqabarella urdunensis Alhejoj, align with the genus Parasheila. Another common Bandel and Al-Najjar, 2016 (original designation), species, the zebinid, Pandalosia ephamilla (Watson, unavailable name under ICZN Art. 16.4: no 1886), has a unique morphology resembling some explicit fxation of name-bearing types (placed in eulimids and has an Indo-Pacific distribution, Aqabarellidae). known from eastern Australia and the Sahul Bank, Timor Sea (Australian Museum via OZCAM). REMARKS Rissoina ambigua (Gould, 1849) was found alive This genus contains only two living species offshore and is the frst record of this species in the worldwide (AFD). Familial placement is uncertain Kimberley (compare ALA). as data are absent regarding soft anatomy and molecular systematics. Superfamily Capuloidea

Only one capulid was found live, Trichamathina Lophocochlias parvissimus sp., a genus not previously recorded from Australia (Hedley, 1899) (AFD, ALA). Liotia parvissima Hedley, 1899: 554–555. Figure 67. Type locality: Funafuti Atoll. Superfamily Cingulopsoidea Haplocochlias (Lophocochlias) minutissimus Pilsbry, Ponder and Yoo (1978) revised the Australian 1921:377. Type locality: Hawaii (new synonym). Eatoniellidae. Names have not been allocated to the eatoniellids from the study, despite one species REMARKS being exceedingly common offshore. Lophocochlias parvissimus was described from Superfamily Truncatelloidea Funafuti Atoll in the Pacifc Ocean (Hedley, 1899). Despite a review of the genus and the description The anabathrids were uncommon with only of a new species, aligning this taxon with the six species found and only one of them was live type of Lophocochlias (Lophocochlias) minutissimus (Anabathron sp., KimMoll247). The Caecidae is a (Pilsbry, 1921) has not occurred. After comparing common component of the Kimberley smaller- clear images of the type material of L. minutissimus mollusc assemblage and has been revised (Pizzini (ANSP) with L. parvissimus (AMS), no discernible 2013). Eighteen species were found in this study. differences could be found and they have been The most common species, Caecum sepimentum de synonymised here, a situation suspected by Rubio Folin, 1868 , was mostly found offshore, with one and Rolán (2015). The species has been recorded lot identifed from a mid-shelf station (148/K13). from the western and central Pacifc Ocean, as well Two uncommon, dead-collected Iravadiids were as Cocos (Keeling) Islands and Christmas Island in included in the sorted samples. the Indian Ocean. 178 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

Lophocochlias procerus Superfamily Buccinoidea Rubio and Rolán, 2015 Although the Australian fauna is substantial, Lophocochlias procerus Rubio and Rolán, 2015: 113. in excess of 78 species (AFD), only one species Figures 5A–G, 6A–G, 7A–E. Holotype IM-2000- (KimMoll334), represented by a single specimen 28215. Type locality: Gulf of Aqaba. was recorded from each of the stations 114/K12 and 152/K14. This is most likely due to sampling Aqabarella urdunensis Alhejoj, Bandel and Al-Najjar, bias towards hard substrates and macromolluscs. 2016: 418. Plate 6, Figure 6 and Plate 7, Figures KimMoll334 aligns with Reticunassa cockburnensis 1–6. Type locality: Gulf of Aqaba. Unavailable (Kool and Dekker, 2006), but the distribution is name under ICZN Art. 16.4: no explicit fxation of apparently disjunct. This species was originally name-bearing types. described from Cockburn Sound near Perth, southern WA, with a southerly distribution to REMARKS Esperance. It is absent from the mid-coast of WA, and recorded again in the north, from Exmouth Aqabarella pulchella Alhejoj, Bandel and Al-Najjar, northward to the offshore atolls of the Kimberley. 2016 (unaccepted name under IZCN) was placed in the new family Aquabarellidae Alhejoj, Bandel Superfamily Conoidea and Al-Najjar, 2016 (unaccepted name under IZCN). In this paper, the authors suggested a close The polyphyletic ‘’ have been revised and relationship to Pickworthiidae. The placement in the family has now been split into 13 monophyletic Aquabarellidae was discovered by chance, when families (Bouchet et al. 2011). This has a major the frst author was conducting literature review impact on the classification of taxa and cross- for Kimberley Pickworthiidae. The same taxon referencing previous checklists assembled from was earlier described by Rubio and Rolán (2015) as not only WA but Australia as a whole. However, Lophocochlias proceus and placed in the Tornidae. taxa may still be tracked at the genus level or The name and taxonomic placement follows Rubio below. Relevant families for this study include and Rolán (2015) in this study. Clathurellidae, Horaiclavidae, Mageliidae and Raphitomidae. The clathurellids are a small-sized group of non-operculate conoideans. Lienardia Superfamily Eulimoidea rubida (Hinds, 1843) (KimMoll571, WAM S94306) has been recorded from Ashmore Reef (ALA) and Approximately 36 species are known from WA was recorded live in this study from Mermaid Reef, (AFD). Some taxonomic papers for the groups Rowley Shoals (Station 156/K14). The most common include those by Warén (1980a, b, 1981, 1983, 1984). clathurellid was ?Lienardia sp. (KimMoll402, A total of 22 species of eulimids were collected, WAM S94322) from Heywood Shoal (Station 148/ 10 of which were found alive. The most common K14), a mid-shelf rise between Hibernia Reef and eulimid was Pyramidelloides mirandus (A. Adams, Rowley Shoals. Two additional species (KimMoll90, 1861), with its presence in WA consistent with the KimMoll93) were identified from the inshore known distribution (Warén 1983). This species Station 91/K12. was taken live offshore, but dead at inshore Station 91/K12. The Horaiclavidae (Bouchet et al. 2011) is an assemblage of genera that exhibit varying Superfamily morphological traits, but are united by molecular data (Bouchet et al. 2011, p. 293, 296). Two species of Macromphalus were recorded A key is lacking, and identifcation relies on a from the survey area, M. aspersus (Hedley, 1912) few characters, including a stout shell, short and M. aculeatus (Hedley, 1900), but neither match and weak spiral . The Macromphalus sp. (KimMoll205, WAM S84717) most commonly encountered ‘’ was Carinapex found in this study. More work is required to revise minutissima (Garrett, 1875) (KimMoll293) identifed the 20 or so living species recorded worldwide from 11 offshore stations. This occurrence is (P. Middelfart, pers. obs.). consistent with the distributional data in ALA, which indicates a ‘clear water’ distribution off NSW, Superfamily Naticoidea Qld and WA (Ashmore and northern Ningaloo Reef). Carinapex sp. (KimMoll574) is similar to Naticids are sand/mud dwellers and are Ceritoturris theoteles (Melvill and Standen, 1896) underrepresented in this study because of the hard (= theoteles (Melvill and Standen, 1896). It bottom bias of the stations sampled. Two damaged is hard to discern from the illustration by Melvill specimens of separate species (KimMoll190, and Standen (1897), but Iredalea s.s. appears to be KimMoll193) were sorted from Station 91/K12. different to KimMoll574 (compare Oliver 1915: 538. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 179

Plate 11, Figure 34). Morphologically KimMoll574 Superfamily Muricoidea appears similar to C. minutissima with a deep anal sulcus, teleoconch nodules and a monocarinate The group is mainly macroscopic, but a few protoconch. More information on I. theoteles and species meet the Me category, e.g. Aspella. In this justifcation of its placement in Iredalea (placed in study mostly young post-metamorphic juveniles the Drillidae) may change that. One single dead were found and they proved almost impossible specimen of KimMoll464 from Ashmore Reef has to place, although KimMoll285 is likely to be a not been placed, but aligns with the horaiclavids. muricine. A juvenile specimen of Magilus striatus (Rüppell, 1835) (sensu Okutani 2000) (= Leptoconchus The re-ranked mangellids are generally medium peronii (Lamarck, 1818), a coral-inhabiting species, to small shouldered ‘turrids’ with strong axials. The was found from Imperieuse Reef (Station 165/K14). protoconch is very characteristic with a sinusigera . Four species have been identified; two (KimMoll409, KimMoll519) from Ashmore Reef and Subclass Heterobranchia two (KimMoll89, KimMoll91) from Station 91/K12. Superfamily Orbitestelloidea The family Raphitomidae is highly variable and hard to determine morphologically. All the Orbitestellidae in Australia and the Australian species identifed in this group were infrequently territories consists of just over 10 species (Laseron encountered with only Microdaphne morrisoni 1954, 1958, Marshall 1988, Ponder 1990, Beesley Rehder, 1981 (KimMoll575) recorded at two et al. 1998). Only two species of Orbitestella and stations from Clerke and Imperieuse Reefs (Station two of Microdiscula have been described from 156/K14 and 165/K14 respectively). Exomiles sp. tropical Australia and neither was recorded from (KimMoll467) is another oceanic species, only tropical WA until now. All four species recovered found south of Hibernia Reef (Station 145/K13). during this study are new. KimMoll240 was the Veprecula spp. (KimMoll86, KimMoll87) were only most common species in twelve stations, while discovered as singletons at Station 91/K12. KimMoll360 was found in eight stations.

Unassigned superfamily (WoRMS) Superfamily Omalogyroidea (previously ) The Omalogyridae was taxonomically assessed While the supraspecific classification of relatively recently (Sartori and Bieler 2014). There marginellids is now clearer (G.A. and H.R. are 33 species worldwide and 11 in Japan, Papua Coovert 1995), the Australian marginellid and New Guinea and the Maldives. Only two species cysticid fauna is in need of major revision. The are known from Australia, both described from most recent work at the species level in Australia NSW (redescribed by Sartori and Bieler 2014). was by Laseron (1957). Many of the described At present no other species is known from Australian species have been illustrated by Beechey Australia despite the existence of numerous (http://seashellsofnsw.org.au) and subsequently species mentioned in Beesley et al. (1998). Four by Cossignani (2006). A very helpful key to genera species, all of which are new, and one new is provided by G.A. and H.R. Coovert (1995). The record (KimMoll489), have been sorted from this most common species was KimMoll371, which was study. Some of the species are quite common only found from Ashmore Reef. KimMoll41 aligns with KimMoll43 identifed from 16 stations and with Pugnus Hedley, 1896 (type Pugnus parvus KimMoll425 from 12 stations. In contrast, only Heldey, 1896). Only one species of Pugnus (the two specimens of KimMoll598 were found from type) has been recorded in Australia (ALA), and Imperieuse Reef (Station 165/K14). its distribution does not include tropical Australia, so more work is required. There are fve additional species worldwide (WoRMS), with only Pugnus Superfamily Rissoelloidea maesae Roth, 1972 from the Indo-Pacific Ocean. The Rissoellidae was last assessed by Ponder and KimMoll41 cannot currently be assigned to any Yoo (1977) and reviewed in Beesley et al. (1998). species. There are 54 species known worldwide (Caballer et Superfamily Mitroidea al. 2011, 2014) and 14 species from Australia (Ponder and Yoo 1977). However, no species have been The mitrid family is mainly macroscopic and reported from Exmouth, WA, to Torres Strait, Qld. only two species, both represented by single Since Ponder and Yoo (1977), no additional species specimens, were identifed from Heywood Shoal have been added to the Australian fauna. WAM (Station 109/K12). It is possible both species are collections from this study indicate there are at least juvenile Scabricola. seven putative new species. 180 PETER MIDDELFART, LISA KIRKENDALE AND CLAY BRYCE

Superfamily Haminoeoidea 2018), but less than the 864 species (under 4.1 mm) collected at Koumac, New Caledonia (Bouchet Six species were identifed, of which three were et al. 2002). The latter study encompassed the live collected from offshore stations (e.g. Station highest habitat diversity while this present study 111/K12, see Table 2). covered the largest geographic area of 476,000 km2 (Bryce et al. 2018) versus 295 km2 at Koumac Superfamily Pyramidelloidea and 18 km2 on Hawai’i. The diversity of smaller Moerchia introspecta (Hedley, 1907), described molluscs in the Kimberley survey area, including from Masthead Reef, Qld, and subsequently a signifcant component (81%) of micromolluscs (≤5 discovered in Darwin (listed in Laseron (1958), is mm adult size class), is yet to reach an asymptote recorded for the frst time in WA (ALA). Other new (Figure 1). More species will be encountered once WA records are: Odostomella opaca (Hedley, 1906), additional habitat types are surveyed, including previously known only from eastern Australia; soft benthic substrates, seagrass beds, deeper Turbonilla microscopica Laseron, 1959, originally waters, mangroves and host-specifc taxa living described from Darwin; Besla articulata (Hedley, with or on other marine . 1909) originally recorded from Hope Island, Qld; This study has indicated how much diversity Chrysallida zea (Hedley, 1902) originally recorded has been missed by earlier expeditions, which did from northern Qld; Chrysallida kesteveni (Hedley, not target smaller molluscs specifcally (Willan 1907) with its type locality at Masthead Reef, Qld, 2005; Willan et al. 2015). This study highlights was moved to the genus Herewardia Iredale, 1935 by the additional diversity afforded when a smaller Peñas and Rolán (2017) (Rissoidae), but transferred mollusc component is added. Ignoring them leads to Chrysallida (see WoRMS). to an underestimate and incomplete assessment of molluscan diversity in broader biodiversity contexts. Superfamily Oxynooidea

Three species, belonging in Berthelinia and Julia ACKNOWLEDGEMENTS were found in this study. Two species of Berthelinia The WAM respectfully acknowledge the have been described from Australia (Jensen 1993, Traditional Custodians of the Kimberley land 1997), but cannot be identifed on their shells alone and sea country, of Elders past and present, and and require dissection for verification (Kathe in particular the Dambimangari and Wunambal Jensen, personal communication). Gaambera peoples, for collections made on their sea country. DISCUSSION We appreciate comments by Winston Ponder, Daniel Geiger, Richard Willan and Kathe Jensen. This study of smaller molluscs has added We are grateful for the suggestions made by two significantly to the knowledge of molluscan reviewers, both of whom greatly improved this biodiversity and biogeography in the Kimberley manuscript and the editor. We acknowledge Project Area. It has documented 624 species Alison Miller and Mandy Reid for information across 102 families or superfamilies, which includes on Lophocochlias spp. and are grateful to Corey 26 new species and 23 new records for WA and Whisson for databasing this material. Sincere Australia. thanks to Woodside for long standing support Twenty one families were added to the Kimberley in association with the Western Australian historical mollusc inventory published by Willan Museum during the Woodside Collection Project et al. (2015) (Figure 2), and more families could be (Kimberley). added with more collecting and research. The samples included in this survey yielded REFERENCES 1,883 identifed lots representing 505 molluscan AFD (Australian Faunal Directory) (2018). Canberra, morphospecies from ≤5 mm adult size category Australia: http://www.environment.gov.au/ (M), 43 molluscs from >5–10 mm adult size category biodiversity/abrs/online-resources/fauna/afd/home (Me), 68 macromolluscs from the >10 mm adult size ALA (Atlas of Living Australia) (2018). Canberra, category (Ma), and 8 species whose adult size could Australia: https://www.ala.org.au/ not be determined. Albano, P., Sabelli, B. and Bouchet, P. (2011). The challenge of small and rare species in marine The level of micromollusc diversity in the biodiversity surveys: microgastropod diversity in a ≤5 mm adult size class category for Hawai’i is more complex tropical coastal environment. Biodiversity and than the 317–375 species (Campagnari and Geiger Conservation 20: 3223–3237. SHALLOW WATER MICROMOLLUSCS OF THE KIMBERLEY 181

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