Diversity and Paleoecology of Miocene Coral-Associated Mollusks from East Kalimantan (Indonesia)

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Diversity and paleoecology of Miocene coral-associated mollusks from East Kalimantan (Indonesia)

Article in Palaios · February 2015 DOI: 10.2110/palo.2013.124

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DIVERSITY AND PALEOECOLOGY OF MIOCENE CORAL-ASSOCIATED MOLLUSKS FROM EAST KALIMANTAN (INDONESIA)

ARIES KUSWORO,1 SONJA REICH,2 FRANK P. WESSELINGH,2 NADIEZHDA SANTODOMINGO,3 KENNETH G. JOHNSON,3 3 2 JONATHAN A. TODD, AND WILLEM RENEMA 1Pusat Survei Geologi, Jl. Diponegoro 57, Bandung 40122, Indonesia 2Naturalis Biodiversity Center, Department of Geology, P.O. Box 9517, 2300 RA Leiden, The Netherlands 3Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW& BD, UK e-mail: [email protected]

ABSTRACT: This study is a preliminary assessment of an extremely diverse Tortonian (late Miocene) mollusk assemblage from a coral carpet environment preserved at Bontang (East Kalimantan, Indonesia). Even though coral-associated aragonitic faunas are rarely well preserved, the composition of the assemblage described here can be used to address the following questions: (1) How do the mollusk assemblages in coral habitats differ from other habitats, and (2) What is the effect of sampling on estimates of taxon richness? The mollusk assemblage is dominated by predatory snails and includes typical modern coral- associated taxa such as the gastropod Coralliophila and the bivalve Tridacna. Our investigation implies that adequate documentation of Cenozoic mollusk diversity in the Indo-Pacific is even more challenging than previously expected as very large samples are required to capture species richness. Further assessments of fossil faunas from coral-dominated habitats will be required to provide insight to development of Indo-Pacific biodiversity through time.

INTRODUCTION and (2) What is the effect of sampling on taxon richness estimates? To achieve these goals we reconstruct the paleoenvironment of the locality, The Indo-West Pacific (IWP), extending from the Red Sea and East document the mollusk assemblage in terms of taxon richness and feeding Africa to the Central Pacific, is the largest of four major oceanic guild composition, and compare our results to published faunas from biogeographic regions, and the one holding the highest taxon diversity elsewhere in the IWP. (Ekman 1934; Briggs 1974; Paulay 1997). Within the IWP, the center of maximum marine biodiversity (species richness) is located in the Indo- Malayan region where local species richness peaks in and around coral GEOLOGICAL SETTING AND STUDY LOCALITY habitats (Hoeksema 2007; Renema et al. 2008). A large variety of The Miocene outcrops of Bontang are located on the eastern side of the organisms, including corals, fish, mollusks, crustaceans, and echinoderms Kutai Basin, the southeasternmost extension of the Sunda platform on contribute to the high diversity in the region (Bellwood et al. 2005; the east coast of Kalimantan, Indonesia. The Kutai Basin is confined by Hoeksema 2007; Renema et al. 2008). Documentation of the fossil record the Sangkulirang fault zone in the north, the Adang fault zone in the is required in order to understand the ecological and environmental south, the Kalimantan High in the west, and to the east the Makassar context of the origin of the biodiversity hotspot, as well as its Strait. The formation of the Kutai Basin initiated during the Eocene development through time (Renema et al. 2008). Although the sampling (Moss and Chambers 1999). Subsequently the basin developed in various of ancient coral reef–associated habitats is necessary to document levels phases during the Cenozoic, including major changes of the basin of biodiversity, this is frequently difficult: coral facies often suffer strong architecture and depositional systems. During the late Neogene and diagenesis, compromising the preservation of associated organisms, Quaternary the basin development was regressive, a result of sediment fill, especially those with aragonitic hard-parts, such as mollusks (Wright et inversion, and uplift (Allen and Chambers 1998). In Miocene times the al. 2003). This results in highly biased mollusk assemblages from reefal study area was located in a coastal zone that received considerable input environments that are largely made up of taxa with more diagenetically of terrigenous clastic sediments (Cibaj 2009; Marshall et al. 2015). In spite resistant and largely calcitic shells, such as oysters and pectinids (Wright of relatively turbid water conditions, rich coral communities could et al. 2003, and references therein; Santodomingo et al. 2015). develop in the region (Wilson 2005; Novak et al. 2015; Santodomingo In this study an unusually well preserved association of ramose corals et al. 2015). and mollusks from upper Miocene deposits in Bontang (East Kaliman- The sampling locality TF 102 (0.16821u N, 117.44350u E) is an tan, Indonesia) is presented. The material provides the opportunity to abandoned quarry on the north side of the northern entrance road to investigate a rare Neogene coral-associated mollusk assemblage from the Bontang (Fig. 1). The locality represents the upper part of a marine IWP biodiversity hotspot. The aim of this study is to reconstruct the interval of about 150 cm of fossiliferous, coarsening-upward, silty clay to paleoenvironment and to characterize the mollusk assemblage in terms of fine sandy silt (Fig. 1). Below this interval gray-blue clay with dispersed diversity and ecology. Specific questions that we address include: (1) How fossiliferous lenses occurs. Above the marine interval lies a non- do the mollusk assemblages in coral habitats differ from other habitats, fossiliferous undulating clay bed of approximately 160 cm thickness

Published Online: February 2015 Copyright E 2015, SEPM (Society for Sedimentary Geology) 0883-1351/15/030-116/$03.00 PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 117

FIG. 1.—Overview over locality TF 102, Bontang, East Kalimantan. A) Location in the Bontang area. B) Situation sketch of coral patches on the floor of abandoned quarry TF 102. C) Lithological column of the TF 102 quarry section: a 5 clay, b 5 silt, c 5 sand, d 5 organic matter, e 5 shells, f 5 corals. The asterisk denotes the sampled interval. Height in meters. followed by approximately three meters of barren fine sandstones. Those sediments represent the basal part of an overlying fluvial unit. Biostratigraphic data (large benthic foraminifera and calcareous nannoplankton) suggest that the TF 102 locality and other fossiliferous localities nearby are assigned a Tortonian age (Renema et al. 2015). Strontium isotope data from scleractinian corals and giant clams (Tridacninae) from six localities in the Bontang area (including TF 102) show ages ranging from 8.24 Ma to 9.75 Ma (61 Ma) with an average of 9.4 6 0.2 Ma, matching a Tortonian biostratigraphic age (Renema et al. 2015).

MATERIAL AND METHODS FIG. 2.—Images of the surface of a coral-mollusk patch on the floor of the TF Fossil mollusk and coral assemblages were collected from discrete 102 quarry. A) Coral fragments and mollusks; a larger muricid gastropod is visible patches of fragmented ramose corals, weathering out on the approxi- in the center. B) Melongena in the center; the smaller gastropod shell in front is a mately horizontal surface of a bedding plane of marine silts and clays Coralliophila. C) A comparatively large strombid shell on the left. (Figs. 1, 2). Although these patches had been rainwashed, leading to some concentration of the fauna, specimens showed no signs of lateral larger mollusks that might indicate partial dissolution of aragonite. transport. In addition, many of the thinner-shelled mollusks were whole, Therefore we could largely discount compositional distortion through suggesting patches had been exposed on the quarry floor for only a short atmospheric weathering of the recovered assemblage. The size and shape period of time. This is confirmed by the lack of any surficial features on of each coral patch was measured along a line transect of 10 m length. A 118 A. KUSWORO ET AL. PALAIOS

TABLE 1.—Mollusk data and feeding ecology. For abbreviations of feeding guilds see Figure 6.

Taxon FG SR50 SR51 SR52 SR53 SR54 Total Diodora sp. 1 H 2 2 1 2 2 9 Stomatolina sp. 2 H 0 0 0 1 0 1 Gibbula leupoldi Beets, 1941 H 5 2 2 11 4 24 ?Gibbula sp. 1 H 0 1 0 2 0 3 Trochidae indet. sp. 1 H 0 1 0 0 0 1 Angaria aff. spaerula (Kiener 1873) sensu Beets, 1942 H 0 0 1 0 0 1 Bothropoma sp. 1 H 1 0 0 0 0 1 Smaragdia semari Beets, 1941 H 6 1 2 5 1 15 Cerithium sp. 3 H 3 2 0 13 7 25 Cerithium sp. 7 H 24 21 4 48 10 107 Colina sp. 1 H 7 5 2 0 0 14 Rhinoclavis sp. 1 s.l. H 0 0 0 0 1 1 Diala semistriata s.l. (Philippi 1849) H 2 4 1 10 9 26 Modulus praeangerensis Martin, 1905 H 0 0 0 4 1 5 ?Potamididae indet. sp. 1 H 0 2 0 0 0 2 Finella sp. 3 H 1 0 0 0 0 1 Ampullina s.l. sp. 2 H 1 0 0 0 0 1 Ampullospira sp. 1 H 2 2 0 1 0 5 Ampullospira sp. 2 H 3 0 0 2 1 6 Plesiotrochus sp. 6 H 14 13 1 18 8 54 ?Capulus sp. 2 SU 0 0 0 0 1 1 Eatoniella s.l. sp. 3 H 1 0 0 0 0 1 Natica helvacea Lamarck, 1822 sensu Beets, 1941 CP 2 4 0 4 8 18 Rissoina (Phosinella)sp.2 H 5 2 0 8 2 17 Rissoina indet. H 0 0 0 0 1 1 Rissolina sp. 2 H 3 1 1 4 0 9 Cyclostremiscus novemcarinatus (Melvill 1906) sensu Beets, 1986 H 0 0 0 0 1 1 Canarium unifasciatum s.l. (Martin 1884) H 14 3 1 7 8 33 Strombus s.l. sp. 2 H 0 0 0 1 2 3 Strombus s.l. sp. 3 H 0 0 0 0 1 1 Strombus s.l. sp. 4 H 2 0 0 0 0 2 Varicospira sp. 1 H 0 0 0 1 0 1 Terebellum sp. H 1 1 0 0 1 3 Cymatium (Reticutriton)sp.1 CP 0 0 1 0 0 1 Cymatium (Turritriton)cf.tenuiliratum (Lischke 1873) CP 0 1 0 0 0 1 Ranellidae indet. sp. 2 CP 1 0 0 0 0 1 Eratoena sp. 1 CB 0 0 0 0 1 1 Epitonium sp. 3 CB 0 0 0 0 1 1 Triphora s.l. sp. 3/Triphora indet. [Beets] CB 3 2 2 5 3 15 Seila sp. 1 CB 1 0 0 1 0 2 ?Buccinidae indet. sp. 1 CP 1 1 0 1 0 3 Euplica sp. 1 CP 1 0 0 4 0 5 Mitrella s.l. cf. gembacana (Martin 1921) CP 4 1 1 5 6 17 Mitrella s.l. njalindungensis sensu (Beets 1941) non (Martin 1921) CP 2 1 1 4 2 10 Zafra sp. 1 CP 14 10 2 23 10 59 Mitrella s.l. sp. 1 CP 2 0 0 2 0 4 Mitrella s.l. sp. 2 CP 0 0 0 1 0 1 Mitrella s.l. sp. 3 CP 0 0 0 0 1 1 Mitrella s.l. sp. 4 CP 0 0 0 0 1 1 ?Pyreneola sp. 1 CP 1 0 0 0 0 1 Peristernia beberiana sensu Beets, 1941 non Martin, 1921 CP 2 0 1 7 1 11 Chicoreus sp. 1 CP 0 2 0 4 0 6 Chicoreus sp. 2 CP 1 0 0 0 0 1 Coralliophila aff. clathrata (A. Adams 1854) CB 3 3 2 7 1 16 Coralliophila sp. 2 CB 2 0 0 0 0 2 Coralliophila sp. 3 CB 5 1 0 1 3 10 ?Muricidae indet. sp. 1 CP 0 0 0 2 0 2 Melongena sp. 1 CP 0 0 1 0 0 1 Nassarius sp. 2 CP 19 9 7 13 7 55 Nassarius sp. 3 CP 7 5 2 16 1 31 Nassarius sp. 4 CP 0 0 0 2 0 2 Nassarius sp. 5 CP 7 4 1 19 4 35 Vexillum sp. 9 CP 4 1 1 8 2 16 Vexillum sp. 11 CP 5 6 1 5 4 21 Vexillum sp. 12 CP 4 5 0 0 0 9 Vexillum sp. 14 CP 4 0 1 5 1 11 Vexillum sp. 15 CP 1 0 0 0 0 1 Vexillum sp. 16 CP 0 1 0 0 0 1 PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 119

TABLE 1.—Continued.

Taxon FG SR50 SR51 SR52 SR53 SR54 Total Vexillum sp. 17 CP 0 0 0 1 0 1 Cystiscus sp. 3 CB 2 0 0 9 0 11 Granulina menkrawitensis (Beets 1986) CB 0 0 0 0 1 1 Granulina sp. 2 CB 0 0 0 1 0 1 Cryptospira sp. 1 CB 3 0 0 2 0 5 Cryptospira sp. 2 CB 1 0 0 0 0 1 Dentimargo sp. 1 CB 4 6 1 7 5 23 Marginellidae indet. CP 0 2 1 0 0 3 Mitra sp. 1 CP 0 0 0 1 1 2 Mitra sp. 2 CP 2 0 0 0 0 2 Mitra sp. 4 CP 0 1 0 0 0 1 Mitra sp. 5 CP 0 0 0 1 0 1 Olivella sp. 2 CP 3 0 0 3 2 8 Dendroconus odengensis s.l. (Martin 1895) CP 0 1 0 1 0 2 Conidae indet. sp. 3 CP 2 1 0 2 0 5 Conidae indet. sp. 4 CP 1 0 0 3 0 4 Conidae indet. sp. 5 CP 3 0 0 0 0 3 Conidae indet. sp. 6 CP 1 0 0 0 0 1 Hemilienardia sp. 1 CP 1 0 0 1 1 3 Lienardia sp. 2 CP 1 0 0 0 3 4 Clathurellidae indet. sp. 1 CP 1 1 0 2 0 4 Clathurellidae indet. sp. 2 CP 1 0 0 0 0 1 Eucithara sp. 5 CP 2 1 1 5 3 12 Eucithara sp. 6 CP 4 1 2 4 2 13 Eucithara sp. 7 CP 0 0 0 1 0 1 Raphitomidae indet. sp. 3 CP 1 0 0 1 0 2 Raphitomidae indet. sp. 4 CP 0 2 0 0 0 2 Raphitomidae indet. sp. 5 CP 0 0 1 0 1 2 Tomopleura sp. 1 CP 1 0 0 0 0 1 Iredalea sp. 1 CP 2 8 5 12 11 38 Tylotiella sp. 2 CP 0 0 0 3 1 4 Drilliidae indet. sp. 2 CP 0 0 0 0 2 2 Inquisitor sp. 3 CP 0 0 0 1 0 1 Pseudomelatomidae indet. sp. 4 CP 0 1 0 0 0 1 Pseudomelatomidae indet. sp. 5 CP 0 0 0 0 1 1 Pseudomelatomidae indet. sp. 6 CP 0 0 0 0 1 1 Strictispira sp. 2 CP 2 0 2 2 2 8 Strictispira sp. 3 CP 0 0 0 1 0 1 Lophiotoma sp. 1 CP 23 15 1 30 9 78 Terebra s.l. sp. 2 CB 0 1 0 0 0 1 Adelphotectonica karikalensis (Cossmann 1910) sensu (Beets 1941) CB 0 1 0 0 1 2 Linopyrga sp. 1 CB 0 1 0 1 0 2 Odostomiinae indet. sp. 1 CB 0 0 0 1 0 1 Pyramidella sp. 2 CB 0 0 0 0 1 1 ?Tibersyrnola sp. 1 CB 0 1 0 1 0 2 Pyramidelloidea indet. sp. 1 CB 1 0 0 0 0 1 Pyramidelloidea indet. sp. 2 CB 0 0 0 1 0 1 Pyramidelloidea indet. sp. 3 CB 0 0 0 1 0 1 Talahabia cf. dentifera Martin, 1921 CP 1 1 0 0 2 4 Cylichna s.l. sp. 2 CP 0 0 0 0 2 2 Cylichna s.l. sp. 3 CP 1 0 0 0 0 1 Pupa sp. 2 CP 0 1 0 0 0 1 Gastropoda indet. spp. - 17 20 4 59 13 113 Nucula sp. 1 SU 0 0 0 0 1 1 Brachidontes sp. 1 SU 0 0 0 1 1 2 Modiolus sp. 1 SU 0 0 0 1 0 1 Acar sp. 1 SU 2 0 0 1 2 5 Anadara sp. 3 SU 13 6 1 13 8 41 ?Barbatia sp. 1 SU 0 0 0 1 1 2 Arcopsis sculptilis sensu Beets, 1941 SU 18 7 2 19 7 53 Pteria s.l. sp. 1 SU 1 0 0 2 1 4 ?Malleus s.l. sp. 1 SU 0 1 0 1 0 2 Dendostrea sp. SU 11 14 4 26 11 66 Lopha sp. 1 SU 0 0 0 0 2 2 Ostreidae indet. SU 1 3 1 5 5 15 Mimachlamys aff. menkrawitensis (Beets 1941) SU 3 2 1 6 1 13 Chlamys s.l. sp. 1 SU 0 1 1 2 1 5 Spondylus sp. 2 SU 0 0 0 0 1 1 120 A. KUSWORO ET AL. PALAIOS

TABLE 1.—Continued.

Taxon FG SR50 SR51 SR52 SR53 SR54 Total Plicatula sp. 1 SU 0 0 1 0 0 1 Cardiolucina sp. 2 CD 2 1 0 1 2 6 Cardita s.l. sp. 3 SU 2 1 0 2 1 6 Trachycardium denticostulatum (Beets 1941) SU 0 1 0 1 0 2 Vasticardium sp. 1 SU 0 0 0 1 0 1 Cardiidae indet. sp. 6 SU 0 0 1 1 0 2 Cardiidae indet. sp. 7 SU 2 0 3 4 2 11 Tridacna (Chametrachea) mbalavuana Ladd, 1934 SU 1 0 0 0 0 1 Chama sp. 1 ? SU 3 3 2 15 7 30 Tellina s.l. sp. 3 D 0 0 1 0 0 1 Tellina s.l. sp. 4 D 0 0 0 0 1 1 Tellina s.l. sp. 5 D 0 0 0 1 1 2 Tellina s.l. sp. 6 D 0 0 0 1 0 1 Circe ickeae sensu (Beets 1941) non Martin, 1922 SU 2 2 0 4 4 12 Dosinia sp. 1 SU 0 0 0 2 0 2 Gafrarium sp. 1 SU 3 1 1 3 0 8 ?Timoclea sp. 1 SU 0 1 0 1 0 2 ?Timoclea sp. 2 SU 0 0 0 1 0 1 Veneridae indet. sp. 3 SU 1 0 0 0 0 1 Veneridae indet. sp. 4 SU 0 0 0 2 0 2 Veneridae indet. sp. 5 SU 0 0 0 1 0 1 Corbula solidula Hinds, 1843 sensu Beets 1941 SU 4 2 1 4 0 11 Corbula sp. 1 SU 1 0 0 0 0 1 Corbula sp. 2 SU 0 0 0 0 1 1 Pholadidae indet. sp. 1 B 0 0 0 0 1 1 Polyplacophora indet. sp. 1 H 0 0 0 0 1 1 total specimens 344 232 78 558 246 1458 total species 84 67 46 97 80 163

total of five coral patches, each with an area of about 1–3 m2, were corals by Johnson et al. (2015). Specimens identified as Acropora were studied (Fig. 1). classified into morphological species groups sensu Wallace (1999). The The first set of samples was obtained by handpicking every mollusk coral material is deposited in the collections of the Natural History seen at the surface of each coral patch. One additional handpicked surface Museum, London, UK (indicated by NHMUK numbers). According to collection includes larger, well-preserved coral colonies, selected to the state of preservation and the available taxonomic frameworks, both complement the taxonomic inventory. Subsequently, two bulk sediment mollusks and corals were identified to the lowest possible rank (species). samples were collected from each patch, resulting in a total of 10 bulk Otherwise, taxa were left in open nomenclature indicating their genus or samples with weights ranging from 3 to 6 kg. One of each of the paired family. Because part of the assemblages could not be identified below samples (numbered for each site: SR50 to SR54) was used to study the family level, the term taxon is used in this study rather than species. In taxonomic and ecological composition of the mollusk assemblage. The addition to the taxonomic classification, feeding guilds of mollusk taxa second sample of each pair (numbered for each site: NS50 to NS54) was were defined as outlined in Reich et al. (2014). Coral abundances were used to study the composition of the coral assemblage. estimated based on number of colony fragments as well as on weight of Bulk samples used for the study of mollusks were treated following the material (dry weight in grams) of each taxon per coral patch. procedures outlined in Reich et al. (2014), assessing all mollusk remains The sampling adequacy was explored using sample rarefaction, also retained on a sieve with a mesh of 1 mm. Counting of specimens was providing an estimate of mollusk diversity. Analysis was performed using likewise done as described in Reich et al. (2014). Taxonomic identifica- PAST (Paleontological Statistics; Hammer et al. 2001). We tested for tions follow publications on fossil faunas of Indonesia (Beets 1941, 1986; statistical differences between the samples using Raup-Crick dissimilar- Leloux and Wesselingh 2009; Reich et al. 2014) as well as descriptions of ities. Raup-Crick dissimilarities were calculated to assess how taxonomic modern Indo-Pacific faunas (e.g., Okutani 2000; Poppe 2008a, 2008b, composition varied among the assemblages, using the vegan package in 2008c, 2010). Specimens that could not be identified due to their poor the R statistical programming environment (Oksanen et al. 2013; R Core preservation are labeled as Gastropoda or Bivalvia indet. spp. and are Team 2013). The method is specifically designed for cases where sampling excluded from statistical analyses. The mollusk material is housed in the is incomplete and uneven among sites, because Raup-Crick dissimilarities collection of Naturalis Biodiversity Center, Leiden, The Netherlands are based on the probability that the compared sampling units have non- (indicated by RGM numbers). identical compositions (Raup and Crick 1979). Two versions of Raup- Bulk samples used for the study of corals were soaked in water, Crick dissimilarities were applied. The first is based on analytic results of washed, and sieved. The size fraction retained on a sieve mesh of 3.5 mm a hypergeometric distribution to find probabilities with the underlying was studied. Identifications of corals are based on macro- and assumption that all species occurrences are equally probable. The second micromorphological features of colonies and corallites, following is based on permutation results that allow sampling probabilities to be taxonomic accounts of Indonesian fossil faunas (Gerth 1921, 1923; proportional to species frequencies. Raup-Crick dissimilarity for almost Umbgrove 1929; mostly illustrated by Leloux and Renema 2007). Taxon identical assemblages is close to zero. Raup-Crick dissimilarity for two names were assigned according to the recent revision of Indo-Pacific fossil assemblages with only a few shared species is close to 1. PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 121

RESULTS bivalves are much more abundant and make up almost 80% of the bivalve Taxonomic Composition of the Mollusk Assemblage assemblage in terms of specimens. This is due to the presence of a few very abundant taxa, such as Dendostrea and Arcopsis (Fig. 3B, C). The A total number of 178 identified mollusk taxa were obtained from the most abundant herbivorous/detritivorous gastropods are Cerithium, samples (Table 1). The five surface collections contained on average 31 Plesiotrochus, and Canarium unifasciatum (Fig. 3G–I, K). taxa (ranging from 17 to 54). The five bulk sediment samples contained The obligate seagrass-feeding gastropod Smaragdia (Rueda et al. 2009; on average 70 taxa (ranging from 45 to 96). The overall most abundant Unabia 2011) is present in low numbers. Browsing carnivores are the least taxa are illustrated in Figure 3. Twenty-four species were present only abundant gastropod feeding guild present. The guild is more prominent in surface-collected samples, whereas 102 species only occurred in bulk in species richness than in abundance. The most common taxon is samples. Gastropods represent the most abundant and most species-rich the marginellid Dentimargo, followed by the coral feeder Coralliophila mollusk group in all samples. They make up more than 70% in terms of (Fig. 3L), and Triphora, an ectoparasite on sponges. The most species- specimens and species numbers in bulk samples. Bivalves make up around rich, though not very abundant, group included in the latter guild are the a quarter of the assemblage. A single polyplacophoran plate was found. ectoparasitic pyramidelloids. Infaunal, deposit-feeding bivalves include Taxon abundance is highly skewed, with few abundant and many rare tellinids and chemosymbiotic deposit feeders, the latter represented by a taxa. species of Cardiolucina (Fig. 3D). Both families contribute little to the abundance and species richness of mollusks from the sampled units Sample Size and Dissimilarity (, 1% to 2%). The rarefaction curves show undersaturation for all samples, especially for the hand-picked surface collections (Fig. 4). Because rarefaction Taxonomic Composition of the Coral Assemblage curves unambiguously showed that the surface-picked collections were Nineteen coral taxa were identified from the bulk sediment samples. too small to provide enough data to accurately characterize assemblage Material from handpicked surface collections was better preserved and composition and taxon richness (Fig. 4), we excluded them from further included larger colonies, but did not yield any additional taxa (Table 3). analyses. Taxon abundances measured as weight of colonies indicate that corals Raup-Crick sample dissimilarity of all five bulk samples using equal are mainly represented by ramose forms (67.5%) with subordinate probability ranges from 0 to 0.18 (mean 5 0.02, Table 2A). Using columnar forms (21.5%). The most abundant ramose corals are taxa in abundance-based dissimilarities, sample SR50 differs from almost the genus Dictyaraea (50.2%; Fig. 5A–C). Of these, three taxa could be all other samples apart from sample SR52 (Table 2B). Raup- Dictyaraea Dictyaraea Crick dissimilarity between SR50 and SR54 is highest. A moderate identified as sp. 1 (29.6%), sp. 2 (19.2%), and dissimilarity is seen between SR50 and samples SR51 and 53. The only Dictyaraea micrantha var. spinosa (1.4%). Other abundant ramose other samples that display a moderate dissimilarity are samples SR51 corals are Alveopora sp. (13.2%), Seriatopora irregularis (2.6%), and and 54. Acropora sp. aspera group (1.5%). Columnar forms include Platygyra The most abundant mollusk taxon in bulk samples is a medium-sized sp. (9.6%) and Goniopora sp. (5.9%). Other coral forms make up less (,10 mm) cerithiid (Fig. 3G). The second most abundant taxon is a than 10% of the total assemblage. They include among others a free-living comparatively large (,25 mm) turrid (Lophiotoma sp. 1, Fig. 3R). The mushroom coral (Fungia sp.), fragments of small massive Oulophyllia sp., next most abundant taxa in bulk samples are represented by the genera and the flabello-meandroid Trachyphyllia sp. No significant difference was Dendostrea, Zafra, Nassarius, Plesiotrochus, Arcopsis, Anadara, and seen when abundances were estimated as number of colony fragments. Iredalea (Fig. 3). The five most abundant taxa in the surface samples are Lophiotoma sp. 1, Corallliophila aff. clathrata, Natica helvacea, DISCUSSION Cryptospira sp. 1, and Euplica sp. 1. These represent the 2nd, 22nd, 19th, 47th, and 48th most important species in the bulk samples. Paleoenvironment Corals are abundant at the sampling locality, but they form small Ecological Composition of the Mollusk Assemblage thickets, evidently following the seafloor morphology instead of forming Feeding ecology was reconstructed for the assemblage obtained from build-up structures. We therefore refer to the paleoenvironment as a coral adding all bulk-sample patches together (Table 1). The assemblage is carpet sensu Riegl and Piller (2000). dominated by predatory and scavenging carnivorous gastropods, both in The dominance of ramose forms in the coral assemblage is interpreted terms of species numbers (55%) and in abundance (48%). The most as a response to high sedimentation rates and high turbidity in the water numerous carnivore, Lophiotoma sp.1(Fig.3G),belongstothe column, because ramose colonies are more likely to overcome potential superfamily Conoidea, which is represented by a total of 25 taxa (193 burial and suffocation caused by constant or episodic sediment discharge specimens) in the sampled assemblage as a whole. The likewise abundant (Sanders and Baron-Szabo 2005; Reuter et al. 2012). Among the ramose Iredalea (Fig. 3G) is a conoidean as well. Both genera were formerly corals present, Dictyaraea species are the most abundant in our samples. lumped together in the Turridae, the members of which commonly prey Dictyaraea is an extinct genus of the family Poritidae, with its oldest fossil on polychaetes (Beesley et al. 1998; Bouchet et al. 2011). Nassariidae are occurrence in the Oligocene of France (Chevalier 1956). The genus is also common, represented by four species (123 specimens; Fig. 3N–P), commonly reported from Miocene fossil coral faunas of Indonesia (Gerth two of which are among the ten most abundant taxa. Members of the 1923; Umbgrove 1929; Johnson et al. 2015) and became extinct in the family largely feed on carrion, although they may shift facultatively to Pleistocene (Johnson et al. 2015). Modern representatives of the family plant matter (Brown 1969). Suspension feeders and herbivores (including Poritidae in reefal habitats include the genera Porites and Goniopora. detritivores) typically make up about a quarter of the assemblage. The Both genera include species that are common in relatively calm waters latter group is slightly more prominent (28%) in the abundance dataset. and that tolerate comparatively high input of sediments into their habitat Suspension feeders are represented by bivalves, apart from a single (Stafford-Smith and Ormond 1992). Assuming Dictyaraea shares similar capulid gastropod. Within the suspension-feeding bivalves, shallow ecologies with its modern counterparts, it is inferred that these sediment- infaunal as well as epifaunal taxa are present. Whereas infaunal and tolerant coral assemblages occupied calm, shallow waters influenced by epifaunal bivalves are represented by the same number of taxa, epifaunal high siliclastic input habitats in the Miocene of East Kalimantan. 122 A. KUSWORO ET AL. PALAIOS

FIG. 3.—Common mollusk taxa in the TF 102 fauna, and some characteristic ecological indicator species. All scale bars 5 1 mm. A) Anadara sp. 3, RGM 793.987, W 9.9 mm. B) Arcopsis ? sculptilis, RGM 793.986, W 5.0 mm. C) Dendostrea sp. 1, RGM 793.965, L 18 mm. D) Cardiolucina sp. 2, RGM 793.967, W 4.4 mm. E) Chama sp. 1, RGM 793.992, H 5.5 mm. F) Gibbula leopoldi, RGM 793.994, H 6.6 mm. G) Cerithium sp. 7, RGM 793.966, H 9.5 mm. H) Plesiotrochus sp. 6, RGM 793.985, H 7.6 mm. I) Cerithium sp. 3, RGM 793.993, H 9.6 mm. J) Diala semistriata s.l., RGM 793.995, H 2.9 mm. K) Canarium unifasciatum s.l., RGM 793.990, H 15.5 mm. L) Coralliophila aff. clathrata, RGM 793.969, H 16.2 mm. M) Zafra sp. 1, RGM 793.983, H 2.9 mm. N) Nassarius sp. 2, RGM 793.984, H. 5.7 mm. O) Nassarius sp. 5, RGM 793.988, H. 4.7 mm. P) Nassarius sp. 3, RGM 793.991, H. 7.1 mm. Q) Iredalea sp. 1, RGM 793.989, H 4.7 mm. R) Lophiotoma sp. 1, RGM 793.968, L 25 mm. PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 123

FIG. 4.—Rarefaction curves. A) Mollusk richness of the TF 102 samples including surface-collected samples (SR45–SR49) and bulk samples (SR50–SR54). B) richness of the entire TF 102 fauna compared to an early Miocene seagrass fauna from Banyunganti (Java, Indonesia; data from Reich et al. 2014).

Fossil mollusks were predominantly observed within the coral patches In addition, some shells of the obligate seagrass-feeding gastropod and not in the sediments in between. This suggests that the occurrence of Smaragdia (Reich et al. 2015) were found. Specimen numbers are very mollusks is associated with the presence of corals in the studied low, but they are equally well preserved as the other taxa (no surface environment. Therefore, we assume that Dictyaraea corals offered a wear, similar coloration). Thus we assume that they were part of the microhabitat that supported the rich mollusk community presented in original life assemblage (e.g., Kidwell and Bosence 1991; Kidwell and this study. The assumed high rates of clay-grade sedimentation probably Flessa 1996; Kidwell 2008). Therefore, we conclude that interspersed provided a preservation window for the aragonitic mollusks. seagrasses or seagrass patches were present in this habitat, as is frequently Close association of components of the mollusk assemblage to the seen in modern coral carpets. once-living coral assemblage is also indicated by the presence of the obligate coral grazer Coralliophila (Beesley et al. 1998). Furthermore, Variation among Samples fragments of the reef-associated bivalve Tridacna were recovered from the quarry floor and in some of the samples. The high abundance of the small The relatively low numbers of specimens in the hand-picked samples and epifaunal ostreid genus Dendostrea may indicate the presence of the observed differences in the composition of the hand-picked versus bulk gorgonians (sea whips, sea fans) in the environment, the preferred samples confirm that the hand-picked samples cannot be used to substrate of modern Dendostrea frons in tropical America (Forbes 1971). meaningfully address taxon richness in mollusk assemblages. Overall, the Although the assemblage is assumed to be largely taphonomically very low Raup-Crick dissimilarities among samples indicate little variation unbiased, we cannot exclude the possibility that the numerical dominance in their taxonomic composition. Therefore, the samples from different of that taxon is due to its higher preservation potential because of its coral patches represent a single coral-associated mollusk community, albeit calcitic shell. Possibly thinner-shelled bivalve taxa were more affected by sampled over a small horizontal distance. This justifies the combined weathering and fragmentation and were therefore not included in the analysis of all five merged bulk samples as a single assemblage. The high sampled assemblages. abundance-based dissimilarity between sample SR54 and samples SR50 and SR51 is probably due to comparatively low abundances of overall very common species, such as Cerithium sp. 7 and Lophiotoma sp. 1 in sample TABLE 2.—Raup-Crick sample dissimilarity. A) Equal probability. SR54 (e.g., 59 specimens of Ceritium sp. 7 in SR51 vs. 13 specimens in B) Proportional to species abundance. SR54). Combining the bulk samples is needed to develop a realistic estimate of overall richness, as the sample size of each of the bulk samples individually (100–500 specimens) is insufficient to produce a realistic A. Patch1 Patch2 Patch3 Patch4 sampling distribution when total richness is at least 178 taxa. Patch2 0.0103 Patch3 0.0001 0.0000 Patch4 0.0109 0.0004 0.0000 Comparison with Other Faunas Patch5 0.1807 0.0391 0.0000 0.0003 Few faunas have been analyzed using a comparable approach and are B. SR50 SR51 SR52 SR53 available for comparison. Even though the total number of observed SR51 0.497 taxa in the early Miocene Banyunganti fauna (Reich et al. 2014) SR52 0.049 0.001 is higher, the sampling curve of the TF 102 fauna is steeper. The SR53 0.545 0.121 0.008 rarefaction curve shows that the overall species richness of the seagrass SR54 0.936 0.704 0.013 0.136 assemblage is lower than that of the coral carpet assemblage (Fig. 4). 124 A. KUSWORO ET AL. PALAIOS

TABLE 3.—Coral taxa abundances and growth forms. Data is presented as the dry weight of each taxon in grams and the number of fragments per sample. Ra 5 ramose, Co 5 columnar, Ma 5 small massive, Pl 5 platy, Fm 5 flabello-meandroid, Fr 5 free-living.

Weight in grams Taxon Form NS50 NS51 NS52 NS53 NS54 Hand picked Total % Dictyaraea sp.1 Ra 95.5 39.5 47.5 139 73 14.5 409 29.6 Dictyaraea sp.2 Ra 121 53.5 19.5 23.5 10 37.5 265 19.2 Dictyaraea micrantha Ra 2.5 5 0.5 6 5 19 1.4 Porites sp. Ra 0.5 0.5 0.5 3.5 5 0.4 Goniopora sp. Ra/Co 5 2.5 5 13.5 0.5 54.5 81 5.9 Seriatopora irregularis Ra 25 0.5 7 3 0.5 0.5 36.5 2.6 Seriatopora hystrix Ra 0.5 0.5 0.5 1.5 0.1 Stylophora sp. Ra 0.5 0.5 0.5 1.5 0.1 Acropora sp. aspera group Ra 21 21 1.5 Alveopora sp. Ra/Co 55 26.5 11.5 28 14 47 182 13.2 Montipora sp. Ra 0.5 12.5 3 2 6 18 1.3 Fungiidae sp. Fr 0.5 2 2.5 0.5 0.5 74 80 5.8 Leptastrea -like Co 13 2 5.5 20.5 1.5 Oulophyllia sp. Ma 0.5 27 27.5 2.0 Platygyra sp. Co 3.5 47 9.5 72 132 9.6 Pectinia sp. Co 0.5 3.5 4 0.3 Trachyphyllia sp. Fm 0.5 3 0.5 0.5 70 74.5 5.4 Pavona sp. Pl 0.5 1 1.5 0.1 Millepora sp. Ra 0.5 0.5 1 0.1 Total 307 181.5 103 269.5 108.5 417 1380.5 Count of fragments Taxon Form NS50 NS51 NS52 NS53 NS54 Hand picked Total % Dictyaraea sp.1 Ra 309 189 108 317 136 11 1070 51.0 Dictyaraea sp.2 Ra 134 123 16 25 8 17 323 15.4 Dictyaraea micrantha Ra 7 14 4 16 3 44 2.1 Porites sp. Ra 6 45 1 24 76 3.6 Goniopora sp. Ra/Co 9 13 3 1 2 11 39 1.9 Seriatopora irregularis Ra 108 10 11 12 4 4 149 7.1 Seriatopora hystrix Ra 4 10 2 16 0.8 Stylophora sp. Ra 1 5 1 7 0.3 Acropora sp. aspera group Ra 52 52 2.5 Alveopora sp. Ra/Co 55 20 21 31 13 9 149 7.1 Montipora sp. Ra 6 38 7 5 2 56 2.7 Fungiidae sp. Fr 2 13 6 3 1 2 27 1.3 Leptastrea -like Co 2 4 2 8 0.4 Oulophyllia sp. Ma 1 2 3 0.1 Platygyra sp. Co 1 21 6 11 39 1.9 Pectinia sp. Co 4 2 6 0.3 Trachyphyllia sp. Fm 5 15 1 2 6 29 1.4 Pavona sp. Pl 1 1 2 0.1 Millepora sp. Ra 1 1 2 0.1 Total 646 554 184 461 172 82 2097

However, it remains unresolved whether the observed difference is based data available suggest a similar pattern to that observed in modern tropical on environmental parameters or is due to the time difference between coral-associated mollusk assemblages. Taylor (1977, 1978) observed that early Miocene (Banyunganti) and late Miocene (TF 102) assemblages. the richness of predatory gastropods is highest in modern reef-associated This stresses the necessity for comparisons of assemblages from the same gastropod assemblages compared to those from other shallow tropical habitat when reconstructing biodiversity over time. To our knowledge, marine environments in his study area at the Maldives. the TF 102 mollusk assemblage is at present the only detailed numerical Because of the similarity of the applied sample processing and counting analysis of a Neogene coral-associated mollusk assemblage from the methods, the feeding guild composition of the TF 102 assemblage can be region, so no comparisons with diversity from other coral-associated directly compared to the composition of the seagrass-associated assemblages can be made. Quantitative data from comparable modern Banyunganti assemblage (Fig. 6; Reich et al. 2014). settings (e.g., Zuschin et al. 2001) were not available for comparison. The Banyunganti assemblage and the assemblage from TF 102 have a These results emphasize first, the need for bulk samples rather than similar composition of feeding guilds when taxon richness is used. hand-picked samples, and second, the large sample sizes needed in However, the feeding guild abundance data discriminate clearly between hyperdiverse faunas. both environments, with the seagrass assemblage dominated by herbivores and the coral carpet assemblage by predators. Suspension Ecological Variation between Environments feeders are less taxon-rich and abundant in the seagrass environment. A The TF 102 mollusk assemblage is dominated both in taxon richness and dominance of herbivorous and detritivorous gastropods is expected for in feeding guild abundance by predatory gastropods. The limited numerical seagrass-associated mollusk communities and has been used previously to PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 125

FIG. 5.—Coral diversity at the TF 102 locality. A) Dictyaraea sp. 1, BMNH AZ6065. B) Dictyaraea sp. 2, BMNH AZ6035. C) Dictyaraea micrantha var. spinosa, BMNH AZ8797. D) Seriatopora irregularis, BMNH AZ8796. E) Alveopora polyacantha, BMNH AZ8788. F) Goniopora sp., BMNH AZ8790. G) Platygyra sp. AZ8786. H) Acropora sp. aspera group, BMNH AZ6006. I) Fungia sp., BMNH AZ8793. identify seagrass meadows in the fossil record (Moulinier and Picard taxonomic impediment is huge. In the current assemblage we were able to 1952; Davies 1970; Brasier 1975; Ivany et al. 1990; Reich et al. 2014). The identify only slightly more than 12% of the present taxa to species level, comparatively low abundance and taxon richness of bivalves in the and several of these identifications remain uncertain. For paleoenviron- Banyunganti assemblage might be attributed to the presence of a dense mental reconstructions, characterizing a fauna by functional groups rhizome mat that inhibited the establishment of an infauna (Davies 1970; instead of a full taxon inventory yields much of the same information James and Bone 2007). However, the majority of filter-feeding bivalves in (Todd et al. 2002). the TF 102 assemblage is represented by epifaunal taxa, and it remains The paleoenvironment of the upper Miocene TF 102 site from Bontang unknown why those taxa are rare to absent in the Banyunganti (East Kalimantan, Indonesia) can be interpreted as a coral carpet that assemblage, although they are often common in modern seagrass developed in calm, shallow waters influenced by high inputs of fine- associations (Mikkelsen et al. 1995; Reich et al. 2014). The data presented grained sediments. Ramose corals of the extinct genus Dictyaraea were here may be applied for further studies on differences in feeding ecologies dominant in these settings and provided a suitable microhabitat for a of mollusk assemblages from different habitats as a possible indicator for diverse mollusk community. The mollusk assemblage associated with the different paleohabitats. TF 102 coral carpet is dominated in taxon richness and abundance by predatory gastropods. The proportion of taxon-based and abundance- based feeding guild composition in the studied coral carpet assemblage is CONCLUSIONS very similar. A Miocene seagrass assemblage from the same region is Our results indicate that in diverse tropical molluscan faunas it is typified by an increase in the proportion of the herbivore/detritivore guild essential to collect abundance data from large bulk sediment samples to in the abundance data when compared to the taxon data. Therefore, be able to make standardized diversity estimates and ecological feeding guild composition of shallow marine tropical mollusk assemblag- characterizations (e.g., Jackson et al. 1999). Furthermore, with over a es might yield a tool for the discrimination of paleohabitats in addition to thousand specimens counted, the rarefaction curve is not saturated, sedimentological or other information, but a lot more data and further implying the need of even larger sample sizes in order to estimate species studies are required to determine the generality of the pattern. richness. When considered individually, the hand-picked samples yield far The coral-associated mollusk assemblage is more diverse than an early too few specimens to be useful in analyses. Another obstacle is that the Miocene seagrass-associated assemblage from the same region; however, 126 A. KUSWORO ET AL. PALAIOS

BOUCHET, P., KANTOR, Y.I., SYSOEV,A.,AND PUILLANDRE, N., 2011, A new operational classification of the Conoidea (Gastropoda): Journal of Molluscan Studies, v. 77, p. 273–308. BRASIER, M.D., 1975, An outline history of seagrass communities: Palaeontology, v. 18, p. 681–702. BRIGGS, S., 1974, Marine Zoogeography: New York, McGraw Hill, 475 p. BROWN, S.C., 1969, The structure and function of the digestive system of the mud snail Nassarius obsoletus (Say): Malacologia, v. 9, p. 447–500. CHEVALIER, J.P., 1956, Les polypiers anthozoaires du Stampien de Gaas (Landes): Bulletin de la Societe´ Histoire Naturelle de Toulouse, v. 90, p. 375–410. CIBAJ, I., 2009, A fluvial series in the Middle Miocene of Kutei Basin: a major shift from Proto-Mahakam shallow marine to the continental environment, in AAPG Hedberg Conference on Variations in Fluvial-Deltaic and Coastal Reservoirs Deposited in Tropical Environments: AAPG (American Association of Petroleum Geologists) Search and Discover Article #90102: Jakarta, Indonesia, p. 1–11. DAVIES, G.R., 1970, Carbonate bank sedimentation, eastern Shark Bay, Western Australia: Memoirs of the American Association of Petroleum Geology, v. 13, p. 85–168. EKMAN, S., 1934, Indo-Westpazifik und Atlanto-Ostpazifik, eine tiergeographische Studie: Zoogeographica, v. 2, p. 320–374. FORBES, M.L., 1971, Habitats and substrates of Ostrea frons, and distinguishing features of early spat: Bulletin of Marine Science, v. 21, p. 613–625. GERTH, H., 1921, Coelenterata. Anthozoa., in Martin K., ed., Die Fossilien von Java Auf Grund Einer Sammlung von Dr. R. D.M. Verbeek Und von Anderen.: Sammlungen des geologischen Reichs-Museums in Leiden. Neue Folge 1(2), p. 387– 445, pl. 55–57. GERTH, H., 1923, Die Anthozoenfauna des Jungtertia¨rs von Borneo [Anthozoa from the upper Tertiary of Borneo]: Sammlungen des Geologischen Reichs-Museums in Leiden, Serie 1, v. 10, p. 37–136, 9 plates. HAMMER, Ø., HARPER, D.A.T., AND RYAN, P.D., 2001, PAST: Paleontological Statistics Software Package for Education and Data Analysis: Palaeontologia Electronica, v. 4, p. 1–9, http://palaeo-electronica.org/2001_1/past/issue1_01.htm. Checked December 2014. HOEKSEMA, B.W., 2007, Delineation of the Indo-Malayan Centre of Maximum Marine FIG. 6.—Comparison of mollusk feeding ecology using species richness (upper Biodiversity: The Coral Triangle, in Renema, W., ed., Biogeography, Time, and Place: diagrams) and abundance data (lower diagrams) of the late Miocene coral carpet Distributions, Barriers, and Islands: Dordrecht, Springer, p. 117–178. fauna from TF 102 (Bontang) and an early Miocene seagrass fauna from IVANY, L.C., PORTELL, R.W., AND JONES, D.S., 1990, Animal-plant relationships and Bayunganti (Java; Reich et al. 2014). H 5 herbivores (including detritivores), CP paleobiogeography of an Eocene seagrass community from Florida: PALAIOS, v. 5, 5 predatory carnivores, CB 5 browsing carnivores, S 5 suspension feeders, D 5 p. 244–258. deposit-feeding bivalves, CD 5 chemosymbiotic deposit feeders. The specimen JACKSON, J.B.C., TODD, J.A., FORTUNATO, H.M., AND JUNG, P., 1999, Diversity and assemblages of Neogene Caribbean Mollusca of lower Central AmericaBulletins number for Bontang TF 102 is higher than reported in Table 1 as unidentified American Paleontology, v. 357, p. 193–230. species of families were included in this analysis. JAMES, N.P., AND BONE, Y., 2007, A late Pliocene-early Pleistocene, inner shelf, subtropical, seagrass-dominated carbonate: Roe Calcarenite, Great Australian Bight, Western Australia: PALAIOS, v. 22, p. 343–359. JOHNSON, K.G., RENEMA, W., ROSEN,B.R.,AND SANTODOMINGO, N., 2015, Old data for old questions: what can the historical collections really tell us about the Neogene it is unknown if this is the result of variation in diversity over time or origins of reef-coral diversity in the Coral Triangle: PALAIOS, v. 30, p. 94–108. between different environments. This indicates the importance of KIDWELL, S.M., 2008, Ecological fidelity of open marine molluscan death assemblages: carefully distinguishing paleohabitats in order to make meaningful effects of post-mortem transportation, shelf health, and taphonomic inertia: Lethaia, comparisons of biodiversity. v. 41, p. 199–217. KIDWELL, S.M., AND BOSENCE, D.W.J., 1991, Taphonomy and time-averaging of marine shelly faunas, in Allison, P.A., and Briggs, D.E.G., eds., Taphonomy: Releasing the ACKNOWLEDGMENTS Data Locked in the Fossil Record: New York, Plenum, p. 115–209. KIDWELL,S.M.,AND FLESSA, K.W., 1996, The quality of the fossil record: populations, species, and communities: Annual Review of Ecology and Systematics, v. 26, p. 269–299. This project is part of the Throughflow project (Marie Curie Initial LELOUX,J.,AND RENEMA, W., 2007, Types and originals of fossil Porifera and Cnidaria Training Network: Grant No. 237922). In the field we were assisted by of Indonesia in Naturalis: NNM (Nationaal Natuurhistorisch Museum) Technical Professor Fauzie Hasibuan, Pak Aseb, Pak Untung, and Sonia Rijani (Pusat Bulletin, v. 10, p. 1–305. Survei Geologi Bandung). Vibor Novak (Naturalis, Leiden) made the TF 102 LELOUX,J.,AND WESSELINGH, F.P., 2009, Types of Cenozoic Mollusca from Java in the lithological column available. Charles Barnard (Naturalis) helped with Martin Collection of Naturalis: NNM (Nationaal Natuurhistorisch Museum) sample processing. J. Darrell and L. Douglas assisted with the curation of Technical Bulletin, v. 11, p. 1–765. coral specimens at the Natural History Museum, London (NHM), and we MARSHALL, N., CIBAJ, I., FRASER, N., HOLBOURN, A., LOGIUDICE, E., MORLEY, R.J., gratefully acknowledge the enthusiastic support of A. Thomas and the NHM NOVAK, V., RENEMA,W.,AND KRIJGSMAN, W., 2015, Integrated stratigraphy and correlation of the Miocene deposits the northern Kutei Basin, E. Kalimantan: ‘‘V-Factor’’ volunteers for exhaustive sample processing. We thank Jill PALAIOS, v. 30, p. 7–25. Leonard-Pingel and an anonymous reviewer for their thoughtful comments. MIKKELSEN, P.M., MIKKELSEN, P.S., AND KARLEN, D.J., 1995, Molluscan biodiversity in the Indian River lagoon, Florida: Bulletin of Marine Science, v. 57, p. 94–127. MOLINIER,R.,AND PICARD, J., 1952, Recherches sur les herbiers de Phane´rogames REFERENCES marines du littoral me´diterraneén français: Annales de l’Institut Oceánographique, v. 27, p. 157–234. ALLEN, G.P., AND CHAMBERS, J.L.C., 1998, Sedimentation in the Modern and Miocene MOSS, S.J., AND CHAMBERS, J.L.C., 1999, Tertiary facies architecture in the Kutai Basin, Mahakam Delta: Indonesian Petroleum Association, Jakarta, Indonesia. 236 p. Kalimantan, Indonesia: Journal of Asian Earth Sciences, v. 17, p. 157–181. BEESLEY, P.L., ROSS, G.J.B., AND WELLS, A.E., 1998, Mollusca: The Southern Synthesis: NOVAK, V., SANTODOMINGO, N., RO¨ SLER, A., DI MARTINO, E., BRAGA, J.C., TAYLOR, Fauna of Australia. Part A. XVI: Melbourne, Australia, CSIRO Publishing. 563 p. P.D., JOHNSON, K.G., AND RENEMA, W., 2013, Environmental reconstruction of a late BEETS, C., 1941, Eine jungmiocane Mollusken-Fauna von der Halbinsel Mangkalihat, Burdigalian (Miocene) patch reef in deltaic deposits (East Kalimantan, Indonesia): Ost-Borneo: Verhandelingen Geologisch Mijnbouwkundig Genootschap Nederland Palaeogeography, Palaeoclimatology, Palaeoecology, v. 374, p. 110–122. and Kolonien, Geologie, v. 13, p. 1–219. OKSANEN, J., BLANCHET, F.G., KINDT, R., LEGENDRE, P., MINCHIN, P.R., O’HARA, R.B., BEETS, C., 1986, Molluscan fauna of the Lower Gelingseh Beds s. str., Sangkulirang SIMPSON, G.L., SOLYMOS, P., STEVENS, M.H.H., AND WAGNER, H., 2013, Vegan area, Kalimantan Timur (East Borneo): Scripta Geologica, v. 82, p.1–82. community ecology package, version 2.0-8, http://vegan.r-forge.r-project.org/. BELLWOOD, D.R., HUGHES, T.P., CONNOLLY, S.R., AND TANNER, J., 2005, Environmental Checked December 2014. and geometric constraints on Indo-Pacific coral reef biodiversity: Ecology Letters, OKUTANI,T.,ED., 2000, Marine Mollusks in Japan: Tokai, Tokai University Press, p. 1– v. 8, p. 643–651, doi: 10.1111/j.1461-0248.2005.00763.x. 1173. PALAIOS MIOCENE INDONESIAN CORAL-ASSOCIATED MOLLUSKS 127

PAULAY, G., 1997, Diversity and distribution of reef organisms, in Birkeland, C.E., ed., SANDERS,D.,AND BARON-SZABO, R.C., 2005, Scleractinian assemblages under sediment Life and Death of Coral Reefs: New York, Chapman and Hall, p. 298–353. input: their characteristics and relation to the nutrient input concept: Palaeogeo- POPPE, G.T., 2008a, Philippine Marine Mollusks, Vol. 1 (Gastropoda—Part I): graphy, Palaeoclimatology, Palaeoecology, v. 216, p. 139–181. Conchbooks, p. 1–759. SANTODOMINGO, N., NOVAK, V., PRETKOVIC, V., MARSHALL,N.,DI MARTINO, E., LO POPPE, G.T., 2008b, Philippine Marine Mollusks, Vol. 2 (Gastropoda—Part II): GIUDICE CAPELLI, E., RO¨ SLER, A., REICH, S., BRAGA, J.C., RENEMA,W.,AND JOHNSON, Conchbooks, p. 1–848. K.G., 2015, A diverse patch reef from turbid waters in the middle Miocene (East POPPE, G.T., 2008c, Philippine Marine Mollusks, Vol. 3 (Gastropoda—Part III): Kalimantan, Indonesia): PALAIOS, v. 30, p. 128–149. Conchbooks, p. 1–702. STAFFORD-SMITH, M.G., AND ORMOND, R.F.G., 1992, Sediment-rejection mechanisms of POPPE, G.T., 2010, Philippine Marine Mollusks, Vol. 4 (Gastropoda—Part IV and 42 species of Australian scleractinian corals: Australian Journal of Marine and Bivalvia—Part I): Conchbooks, p. 1–665. Freshwater Research, v. 43, p. 683–705. RCORE TEAM, 2013, R: A Language and Environment for Statistical Computing, TAYLOR, J.D., 1977, Food and habitats of predatory gastropods on coral reefs: Report version 3.0.1: R Foundation for Statistical Computing, Vienna, Austria, http://www. of the Underwater Association, New Series, v. 2, p. 155–193. R-project.org. Checked December 2014. TAYLOR, J.D., 1978, Habitats and diet of predatory gastropods at Addu Atoll, Maldives: RAUP, D.M., AND CRICK, R.E., 1979, Measurement of faunal similarity in paleontology: Journal of Experimental Marine Biology and Ecology, v. 31, p. 83–103. Journal of Paleontology, v. 53, p. 1213–1227. TODD, J.A., JACKSON, J.B.C., JOHNSON, K., FORTUNATO, H.M., HEITZ, A., ALVAREZ,M. REICH, S., WESSELINGH, F.P., AND RENEMA, W., 2014, A highly diverse molluscan AND JUNG, P., 2002, The ecology of extinction: molluscan feeding and faunal turnover seagrass fauna from the early Burdigalian (early Miocene) of Banyunganti (south- in the Caribbean Neogene, Proceedings of the Royal Society, Biological Sciences, v. central Java, Indonesia): Annalen des Naturhistorischen Museums in Wien, Serie A, v. 116, p. 5–129. 269, p. 571–577. UMBGROVE, J.H.F., 1929, Anthozoa van N.O. Borneo: Wetenschappelijke Mededelingen REICH, S., WARTER, V., WESSELINGH, F.P., ZWAAN, H., RENEMA,W.,AND LOURENS,L., 2015, Paleoecological significance of stable isotope ratios in Miocene tropical shallow van de Dienst van de Mijnbouw in Nederlands Indie¨, v. 9, p. 46–76, pl. 1–5. marine habitats (Indonesia): PALAIOS, v. 30, p. 53–65. UNABIA, C.R.C., 2011, The snail Smaragdia bryanae (Neritopsina, Neritidae) is a RENEMA, W., BELLWOOD, D.R., BRAGA, J.C., BROMFIELD, K., HALL, R., JOHNSON, K.G., specialist herbivore of seagrass Halophila hawaiiana (Alismatidae, Hydrocharitaceae): LUNT, P., MEYER, C.P., MCMONAGLE, L.B., MORLEY, R.J., O’DEA, A., TODD, J.A., Invertebrate Biology, v. 130, p. 100–114. WESSELINGH, F.P., WILSON, M.E.J., AND PANDOLFI, J.M., 2008, Hopping Hotspots: WALLACE, C., 1999, Staghorn Corals of the World: A Revision of the Coral Genus Global Shifts in Marine Biodiversity: Science, v. 321, p. 654–657. Acropora: Collingwood, Victoria, Australia, CSIRO Publishing. RENEMA, W., WARTER, V., NOVAK, V., YOUNG,Y.,AND MARSHALL, N., 2015, Age of WILSON, M.E.J., 2005, Development of equatorial delta-front patch reefs during the Neogene localities in the northern Kutai Basin: PALAIOS, v. 30, p. 26–39. Neogene, Borneo: Journal of Sedimentary Research, v. 75, p. 114–133, doi: 10.2110/ REUTER, M., PILLER, W.E., AND ERHART, C., 2012, A Middle Miocene carbonate jsr.2005.010. platform under silici-volcanoclastic sedimentation stress (Leitha Limestone, Styrian WRIGHT, P., CHERNS,L.,AND HODGES, P., 2003, Missing molluscs: field testing Basin, Austria): depositional environments, sedimentary evolution and palaeoecology: taphonomic loss in the Mesozoic through early large-scale aragonite dissolution: Palaegeography, Palaeoclimatology, Palaeoecology, v. 350–352, p. 198–211. Geology, v. 31, p. 211–214, doi: 10.1130/0091-7613(2003)031,0211. RIEGL,B.M.,AND PILLER, W.E., 2000, Reefs and coral carpets in the northern Red Sea ZUSCHIN, M., HOHENEGGER,J.,AND STEININGER, F.F., 2001, Molluscan assemblages on as models for organism-environment feedback in coral communities and its reflection coral reefs and associated hard substrata in the northern Red Sea: Coral Reefs, v. 20, in growth fabrics: Geological Society, London, Special Publications, v. 178, p. 71–88, p. 107–120. doi: 10.1144/GSL.SP.2000.178.01.06. RUEDA, J.L., SALAS, C., URRA,J.,AND MARINA, P., 2009, Herbivory on Zostera marina by the gastropod Smaragdia viridis: Aquatic Botany, v. 90, p. 253–260. Received 12 November 2013; accepted 26 November 2014.

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