Origins of diversity in Southeast Asia

Nadia Santodomingo1, Kenneth Johnson1, Willem Renema2, and the Throughflow Project

1 Natural History Museum. Palaeontology Department. Cromwell Road. SW7 5BD. London. [email protected] 2 Netherlands Centre for Biodiversity Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands.

Fossil AZ-coral genera and first occurrences Beyond the palaeontology This table shows the current knowledge of AZ-coral taxa, based on inventories of recent species (Cairns and Zibrowius, 1997) and fossil coral Evolution: First occurrences of fossil collections at NCB Naturalis and the Natural History Museum. A total of 51 fossil AZ-coral species have been recognised, from which 30 are can be used for the calibration of molecular extinct and 21 are present in modern fauna. The Conosmilia is the only one extinct. clocks in phylogenetic analysis. The first occurrences of the different AZ-coral genera are illustrated in the graph below. Climate archivals: Preservation can be excellent allowing geochemical analysis 1mm to understand the palaeoenviroments. Detail of calicular margin of Caryophyllia sp. Modern species: 212 spp. 3 5 1 1 3 2 14 6 2 7 4* 1 3 1 2 1 1 11 2 2 2 13 1 15 18 6 2 2* TF 526 (Bengalon) Fossil record Extant: 21 spp. 1 1 1 1 1 2 1 1 1 1 1 2 1 4 1 1 1 1 2 1 1 2 1 2 3 2 1 1 2 1 1 1 2 3 2 Extinct: 30 spp. Museum collections Age (Ma) 0 40 80 120 EPOCH AGE Ma HOLOCENE Y 0.01 TER- 0 A CALABRIAN NAR U PLEISTOCENE 1.8

Q Late occurrence of common genera (e.g. GELASIAN 2.6 PIACENZIAN 3.6 ) PLIOCENE ZANCLEAN Madracis, Astrangia) that are known from 5.3

) -5 MESSINIAN earlier records in adjacent areas. 7.2 Coen. ( L TORTONIAN Cyathelia Odon. *Madracis -10 11.6 ( SERRAVALLIAN Endopachys Most first occurrences of AZ-coral genera 13.8 M LANGHIAN Conotrochus -15 are from the Early to Middle Miocene. ) ) 16.0 Astrangia ) MIOCENE

NEOGENE BURDIGALIAN Stephanocyathus Acin. Aplo. ( E 20.4 ( -20

AQUITANIAN Acan. ( 23.0 Stephanocyathus Paracyathus Madrepora -25 Paucity of AZ-corals or “Oligocene gap”?

CHATTIAN Placotrochides Lophelia L Deltocyathus ropidocyathus ) Cyathotrochus

Phyllangia Some recent studies in the area include T *Heteropsammia

28.4 Anthemiphyllia runcatoflabellum Caryophyllia *Heterocyathus Dendrophyllia

T earlier records of Lophelia and few other

Flabellum -30 E rochocyathus RUPELIAN Conosmilia Bath. T Caryophyllia ( species. OLIGOCENE

33.9 Stephanocyathus L PRIABONIAN -35 37.2 ALEOGENE

P BARTONIAN -40 40.4 The Eocene is represented by a single EOCENE rochocyathus

LUTETIAN Fungiacyathus T Notocyathus Balanophyllia locality in Java, containing 4 species. M -45 CARYOPHYLLIIDAE FLABELLIDAE DENDROPHYLLIIDAE POCI FUNG RHIZANGIIDAE OCULINIDAE ANTE TURB

A B D F H I K N AZ-corals show the same pattern as shallow Z-corals: most first occurrences are from the Early to Middle Miocene. This supports the idea that plate tectonics and other regional processes (climatic events) are the main factors involved in the diversification of this area. G L No fossil record is known for the basal clades of Scleractinians in this region (Fam. Micrabaciidae). C E More than half of known fossil AZ-coral species became extinct, suggesting high rates of origination. J M O In combination with parallel studies on shallow water ecosystems, these new data provide insights to the origins of the high diversity in this region.

A. Madracis myriaster, B. Fungiacyathus (Bathyactis) eocaenica, C. Astrangia (Coenangia) polygonalis, D-E. Madrepora sp., F. Caryophyllia sp. G. Deltocyathus sp., H. Heterocyathus aequicostatus, ct-scan reconstruction, I. Paracyathus cf. javana,, J. Stephanocyathus (Odontocyathus) sp., K. Notocyathus viola, L. Truncatoflabellum variabile, M. patens, N. Dendrophyllia rutteni, O. Balanophyllia javaensis (?). Scale bar= 5mm in B-D,F,H-O; scale bar= 1mm in E; scale bar= 2.5 mm. Images: A-C, K-O from Leloux and Renema, 2007.

Historical collections Biological and Geological context Most fossil corals were collected by geologist employed by the Dutch Government, during Southeast Asia hosts the maximum centre of marine diversity for both azooxanthellate (AZ-) and their explorations of mineral resources in Indonesia (Netherlands East Indies). First collections zooxanthellate (Z-) corals. Various hypotheses have been proposed to explain this pattern (centre of were made by Junghuhn (1809-1864) and J.K.L. Martin (1851-1942), followed by Wanner, originiation, accumulation, survival), for which fossils could offer key information. However, the fossil Gerth, Verbeek, Rutten, Leupold and Umbgrove, with last expeditions by the end of the 40's. record is still markedly undersampled. Monographies of this fossil coral fauna include Martin (1880, 1917), Felix (1913,1921), Gerth (1921-1933) and Umbgrove (1926-1950). Azooxanthellate coral richness Coral richness We are currently examining the identifications and taxonomic status of these collections.

92

157

79 Coral 138 Triangle 83 The Throughflow Expeditions 116 137 99 76 The Throughflow network includes five major topics: palaeoceanography, palaeocology, stratigraphy, Cairns, 2007 Bellwood et al.,2005 geochemistry and biodiversity. Researchers of seven European institutions work in close collaboration with the Geological Survey in Indonesia. >70 >80 >100 species >0 >100 >200 >300 >400 >500 >600 species EAST Evidence from palaeontological and molecular studies suggests that the Miocene was an important KALIMANTAN period for diversification in the region. Reef development during the Cenozoic has been linked to Sangkulirang major global climatic changes, such as the Middle Miocene Climatic Optimum, when corals reached its acme, followed by their decrease during the gradual ocean cooling after this period. Bontang This global pattern is mainly 4 Study area based on studies of the 0 Samarinda Mahakan δ18O (‰) 110°E 115°E 120°E 125°E 130°E 135°E Delta Mediterranean and ene ene Plio- c Caribbean fossil fauna. They 5°N ITF Today Sampling in outcrop TF530 Pacific Ocean Pliocene (Sangkulirang Bay) present different trajectories Pleistoc 0 50 100 Km e Mindanao Eddy 10

n Makassar Strait

e SULAWESI ene suggesting that regional c Borneo

o Halmahera Eddy i 0° Halm Maluku ahera factors might also play an Mioc e M l Seram

d a i~n N

d Two expeditions were carried out in Nov-Dec 2010 and Jun-Jul 2011 important control on the coral a

i 5°S Sulawesi L akassar

limatic Optimum M M 20 development. Plate tectonic C Java resulting in the collection of about 3 tons of coral specimens of

DewakangSill Banda dynamics and consequent Java Flores Million years old). Although this extensive sampling was focussed on 10°S constraint of the Indonesian ene shallow marine ecosystems, some deep-water facies were also studied. Timor

Lombok Timor 10 Throughflow Current (ITF) 30 ITF AZ-corals have been found in 8 of the 49 outcrops so far. Oligoc Indian Ocean Australia during the Miocene could Gordon, 2005 have favoured the 0 4 8 5 mm diversification in the region. Ice-free temperature (°C) Zachos et al., 2008 Caryophyllia sp. TF 530 (Sangkuliran Bay)

Acknowledgements Thanks to all the members of the Throughflow team. To Brian Rosen, Jill Darrell, Lindsey Douglas, and the V-factor Team (NHM). To Alex Ball and Thomasz Goral (SEM at NHM)