SG124 003-264 (renema) 15-01-2007 15:48 Pagina 1 VRIJE UNIVERSITEIT LARGER FORAMINFERA AS MARINE ENVIRONMENTAL INDICATORS ACADEMISCH PROEFSCHRIFT TER VERKRIJGING VAN DE GRAAD VAN DOCTOR AAN DE VRIJE UNIVERSITEIT AMSTERDAM, OP GEZAG VAN DE RECTOR MAGNIFICUS PROF. DR. T. SMINIA, IN HET OPENBAAR TE VERDEDIGEN TEN OVERSTAAN VAN DE PROMOTIECOMMISSIE VAN DE FACULTEIT DER AARD- EN LEVENSWETENSCHAPPEN OP MAANDAG 9 DECEMBER 2002 OM 10.45 UUR IN HET AUDITORIUM VAN DE UNIVERSITEIT, DE BOELELAAN 1105 DOOR WILLEM RENEMA GEBOREN TE APELDOORN SG124 003-264 (renema) 15-01-2007 15:48 Pagina 2 Promotor: prof.dr. J.E. van Hinte Copromotor: dr. S.R. Troelstra dr. C.F. Winkler Prins SG124 003-264 (renema) 15-01-2007 15:48 Pagina 3 Larger foraminifera as marine environmental indicators Willem Renema Renema, W. Larger foraminifera as marine environmental indicators. — Scripta Geol., 124: 1-260, 114 figs., 13 pls; Leiden, November 2002. Willem Renema, Nationaal Natuurhistorisch Museum, PO Box 9517, 2300 RA Leiden, The Netherlands; [email protected] Key words — Foraminifera, Spermonde Shelf, Sulawesi, Cabilao, Philippines, Recent, Palaeogene, diversity. Contents 1. General introduction and abstract ................................................................................................... 3 2. Larger benthic foraminifera and their distribution patterns on the Spermonde Shelf, South Sulawesi ................................................................................................... 9 - Renema, W., Hoeksema, B.W. & Hinte, J.E. van: Zoologische Verhandelingen Leiden 334 (2001): 115-150. 3. Larger foraminifera distribution on a mesotrophic carbonate shelf in SW Sulawesi (Indonesia) .................................................................................................................... 40 - Renema, W. & Troelstra, S. R.: Palaeogeography, Palaeoclimatology, Palaeoecology 175 (2001): 125-146. 4. Thanatofacies of larger benthic foraminifera on the Spermonde Shelf (Southwest Sulawesi, Indonesia) ................................................................................................... 57 5. Larger foraminifera from Cabilao Island (Bohol, Philippines) ........................................ 88 6. Palaeogene nummulitids (Foraminiferida) from the Indonesian Archipelago: a review .................................................................................................................................................... 110 - Renema, W., Racey, A. & Lunt, P.: accepted in Cainozoic Research 2 (2) (2002) 7. Palaeogene larger foraminiferal generic diversity in southeast Asia as compared to the western Tethys .................................................................................................. 166 8. Synthesis .................................................................................................................................................. 189 9. References ............................................................................................................................................... 213 10. Groot foraminiferen als milieu-indicatoren ........................................................................... 226 11. Dankwoord ............................................................................................................................................ 235 12. Curriculum vitae ................................................................................................................................. 236 1. General introduction and abstract “Larger benthic foraminifera” are large, single celled organisms living under simi- lar conditions as zooxanthellate corals, in shalllow tropical seas. Just like other reef associated taxa the diversity of larger benthic foraminfera is highest in the border region between the Pacific and Indian Ocean, the Indo-West Pacific (IWP). Since foraminifera have calcareous tests and occur in huge numbers, they fossilise easily. Other taxa on which stratigraphical schemes have been built, e.g., planktonic foraminifera and nannoplankton, are not as abundant in tropical shallow marine car- SG124 003-264 (renema) 15-01-2007 15:48 Pagina 4 4 Renema. Larger foraminifera as marine environmental indicators. Scripta Geol., 124 (2002) bonates and in these settings larger foraminfera have been used for stratigraphical purposes. The Recent distribution of larger foraminifera has mainly been studied in pristine areas with a deep photic zone. Distribution of larger foraminfera in such areas proba- bly differs from those on mesotrophic carbonate shelfs. The latter are also more likely to preserve and dominate carbonate production during the Cenozoic in tropical south- east Asia (Wilson, 2002). The present thesis aims at: - Describing the Recent distribution over various reefs and carbonate platform. - Identifying the most important parameters that determine the occurrence of species of larger benthic foraminifera. - Determining whether the distribution of empty tests of larger benthic foraminifera reflects the distribution of the living protists, and determining whether the occur- rence of larger benthic foraminifera can be used for palaeoenvironmental analysis. - Use these models to interpret the fossil record. Two main groups of foraminifera, differing in test structure, house symbionts, lamellar perforate foraminifera and imperforates. Hyaline shells have a higher trans- parency because the calcite crystals have their longest crystallographic axis perpendic- ular to the outer test wall. Transparency of the test is much lower in imperforate foraminifera, in which the crystals are randomly oriented with an outerlayer with the crystal axis parallel to the outer test wall (Haynes, 1965, 1981; Hallock, 1999). In order to profit from photosynthesising endosymbionts, the symbionts need to be irradiated through the test wall. In this introduction I will first give a summary of the parameters determining larg- er benthic foraminiferal distribution over reefs and carbonate platforms and their strategies to cope with these conditions. The second part of the introduction compris- es of a brief summary of the chapters. The material is deposited in the National Museum of Natural History (Leiden, The Netherlands) with registration numbers prefixed RGM. Parameters The distribution of larger benthic symbiont-bearing foraminifera is determined by a complex set of often inter-related parameters Fig. 1.1 summarises these parameters and the way they interact with foraminifera. Two parameters limit the geographical distribution of larger benthic foraminifera occurrence, these are temperature and nutrient availability, of which the latter is affected by the first. Housing symbionts is only profitable in conditions where organic matter is concentrated in particles (Hallock, 1981c). The host collects and digests par- ticulate organic matter, and the symbiont feeds on the waste product of the host. Excess energy produced by the symbiont is again used by the host (Hallock, 1981c). Laboratory experiments show that 10-90% of the energy of larger symbiont-bearing foraminifera is provided by the symbionts (ter Kuile, 1991). The role of photosynthesis in calcification is not clear. Ter Kuile (1991) holds that 2- photosynthesis consumes CO2, producing CO3 and thus enhances calcification. SG124 003-264 (renema) 15-01-2007 15:48 Pagina 5 Renema. Larger foraminifera as marine environmental indicators. Scripta Geol., 124 (2002) 5 Environment Nutrient availabilty transparency Depth Temperature Terrestrial influx Depth light intensity Stabilty Hydrodynamic Energy Substrate Type Animal symbiont type shell structure shell morphology life history Fig. 1.1. Schematic representation of the environmentalparameters influencing the distribution of larg- er foraminifera and properties of the foraminfera to deal with these conditions. McConnaughley (1989) and McConaughley & Whelan (1997) have proposed the reverse interpretation, though. They postulate that lack of CO2 limits photosynthesis in warm, shallow aquatic environments and that calcification provides protons that make CO2 readily available (Hallock, 2001). Carbon cycling in perforate foraminifera supplies 10% of the carbonate incorporat- ed in the skeleton, while feeding is mainly used as a source for nutrients (ter Kuile et al., 1987; ter Kuile & Erez, 1987; ter Kuile, 1991). In imperforate foraminifera the car- bon uptake for calcification and photosynthesis occurs in two separate flows, and thus one does not affect the other (ter Kuile, 1991). The dependence on light for their sym- bionts limits larger benthic foraminifera to the photic zone. In warm water, the metabolic rate of larger foraminifera is higher than in cool water. In invertebrates metabolic rate doubles with a 10 degrees increase in tempera- ture. Thus it is more likely to find sufficiently oligotrophic conditions in tropical than in temperate seas. Larger symbiont-bearing foraminifera are restricted to a climatic belt limited by the 16°C isotherm in the coldest month (Langer & Hottinger, 2000). In most studies on the distribution of larger foraminifera depth is the most impor- tant directly measurable environmental parameter. However, depth only affects the distribution of foraminifera indirectly (Fig. 1.1). At first, light intensity decreases with depth dependent on the transparency of the water column. Transparency in its turn is influenced by nutrient availability and terrestrial influx. In areas with high terrestrial SG124 003-264 (renema) 15-01-2007 15:48 Pagina