The influence of light attenuation on the biogeomorphology of a marine karst cave: 1 a case study of Puerto Princesa Underground River, Palawan, the Philippines. 2 3 Martin A. Coombes 1* , Emanuela C. La Marca 1 , Larissa A. Naylor 2, Leonardo Piccini 3, Jo De 4 Waele 4 , Francesco Sauro 4 5 1. School of Geography and the Environment, University of Oxford, Oxford, UK. 6 2. School of Geographical and Earth Sciences, University of Glasgow, UK. 7 3. Department of Earth Science, University of Firenze, Italy. 8 4. Department of Biological, Geological and Environmental Sciences, University of Bologna, Italy. 9 *[email protected]10 11 Abstract 12 Karst caves are unique biogeomorphological systems. Cave walls offer habitat for 13 microorganisms which in-turn have a geomorphological role via their involvement in rock 14 weathering, erosion and mineralisation. The attenuation of light with distance into caves is 15 known to affect ecology, but the implications of this for biogeomorphological processes 16 and forms have seldom been examined. Here we describe a semi-quantitative microscopy 17 study comparing the extent, structure, and thickness of biocover and depth of endolithic 18 penetration for the Puerto Princesa Underground River system in Palawan, the 19 Philippines, which is a natural UNESCO World Heritage Site. 20 Organic growth at the entrance of the cave was abundant (100% occurrence) and 21 complex, dominated by phototrophic organisms (green microalgae, diatoms, 22 cyanobacteria, mosses, and lichens). Thickness of this layer was 0.28 ± 0.18 mm with 23 active endolith penetration into the limestone (mean depth = 0.13 ± 0.03 mm). In contrast, 24 phototrophs were rare 50 m into the cave and biofilm cover was significantly thinner (0.01 25 ± 0.01 mm, p < 0.000) and spatially patchy (33% occurrence). Endolithic penetration here 26
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The influence of light attenuation on the biogeomorphology of a marine karst cave: 1
a case study of Puerto Princesa Underground River, Palawan, the Philippines. 2
3
Martin A. Coombes1*, Emanuela C. La Marca1, Larissa A. Naylor2, Leonardo Piccini3, Jo De 4
Waele4, Francesco Sauro4 5
1. School of Geography and the Environment, University of Oxford, Oxford, UK. 6
2. School of Geographical and Earth Sciences, University of Glasgow, UK. 7
3. Department of Earth Science, University of Firenze, Italy. 8
4. Department of Biological, Geological and Environmental Sciences, University of Bologna, Italy. 9
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2004. Distribution of phototrophic biofilms in cavities (Garraf, Spain). Nova 455
Hedwigia, 78, 329-351. 456
Roldàn, M., Hernandez-Mariné, M., 2009. Exploring the secrets of the three-dimensional 457
architecture of phototrophic biofilms in caves. International Journal of Speleology 458
38(1), 41-53. 459
Taylor, M.P., Viles, H.A., 2000. Improving the use of microscopy in the study of 460
weathering: sampling issues. Zeitschrift für Geomorphologie, Supplementbände, 461
120, 145-158. 462
Trudgill, S.T., 1976. The marine erosion of limestones on Aldabra Atoll, Indian Ocean. 463
Zeitschrift für Geomorphologie, Supplementbände, 32, 67-74. 464
Viles, H., 1987. A quantitative scanning electron microscope study of evidence for lichen 465
weathering of limestone, Mendip Hills, Somerset. Earth Surface Processes and 466
Landforms, 12(5), 467-473. 467
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recolonization data from limestone surfaces, Aldabra Atoll, Indian Ocean: 469
implications for biological weathering. Earth Surface Processes and Landforms, 470
25(12), 1355-1370. 471
472
473
Table 1. Rock sampling in relation to distance into the Puerto Princesa Underground River 474
system and height above mean water level. 475
20–30 cm above waterline
40–50 cm above waterline
70–80 cm above waterline
Cave entrance SP0/1 SP0/2 SP0/3 50 m from entrance SP1/1 SP1/2 SP1/3
250 m from entrance SP2/1 SP2/2 SP2/3 476
Figure Captions: 477
Figure 1. Location map of the Saint Paul karst area. Location of the Puerto Princesa 478
Underground River system (PPSE) indicated. 479
Figure 2. Photographs showing: (a) well-developed notches of the Saint Paul karst area; (b) 480
notch development at the cave entrance; (c) core sampling at the cave entrance; (d) core 481
sampling 50 m into the cave, and; (e) core sampling 250 m into the cave (scale bars indicate 482
10 cm). 483
Figure 3. Occurrence (%) of phototrophic groups of microorganisms and invertebrates in 484
light microscope observations of samples taken at three distances from the cave entrance 485
(0 m n = 15, 50 m n = 15, 250 m, n = 13). 486
Figure 4. Varying structure of biofilm on the surface of samples from different locations in 487
the cave: (a) thick and stratified biofilm at the cave entrance; (b and c) complex biofilm 488
with visible filamentous algae at the cave entrance; (d) simple epilithic biofilm 489
characteristic of surfaces 50 m from the entrance, and; (e and f) 250 m from the cave 490
entrance (scale bars = 1 mm). 491
Figure 5. SEM micrographs of top surfaces of samples taken at: (a) 0 m, (b) 50 m, and (c) 492
250 m from the cave entrance (all samples are 40–50 cm above the mean waterline, 493
magnification and scale as shown). 494
Figure 6. Invertebrates present on samples sampled 20–30 cm from the mean waterline: 495
(a) Mollusca (bivalves) at the cave entrance; (b) Mollusca (gastropods) at the cave 496
entrance; (c) Mollusca (bivalves) 250 m from the cave entrance; (d) Arthropod crustacean 497
at the cave entrance (bars = 1 mm). 498
Figure 7. SEM observations of surfaces in cross-section for samples taken at: (a) 0 m 499
from the entrance ([i] thickness of biocover, [ii] zone of bioerosion, [iii] close-up view of in 500
situ filaments penetrating the rock); (b) 50 m from the entrance ([i] possible zone of 501
biochemical alteration); (c) 250 m from the cave entrance ([i] amorphous structures, of 502
possible chemical rather than organic origin) (all samples were from 40–50 cm above the 503
mean waterline, magnification and scale as shown). 504
Figure 8. Indicative thickness of biocover (mean + SD) and depth of active penetration into 505
the substratum (mean + SD) by microorganisms at different distances within the cave (all 506
samples from 40–50 cm above mean waterline, n = 15). 507
508
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Roldàn, M., Clavero, E., Canals, A., Gòmez-Bolea, A., Arino, X., Hernandez-Mariné, M., 2004. Distribution of 587 phototrophic biofilms in cavities (Garraf, Spain). Nova Hedwigia, 78, 329-351. 588
Roldàn, M., Hernandez-Mariné, M., 2009. Exploring the secrets of the three-dimensional architecture of 589 phototrophic biofilms in caves. International Journal of Speleology 38(1), 41-53. 590
Taylor, M.P., Viles, H.A., 2000. Improving the use of microscopy in the study of weathering: sampling issues. 591 Zeitschrift für Geomorphologie, Supplementbände, 120, 145-158. 592
Trudgill, S.T., 1976. The marine erosion of limestones on Aldabra Atoll, Indian Ocean. Zeitschrift für 593 Geomorphologie, Supplementbände, 32, 67-74. 594
Viles, H., 1987. A quantitative scanning electron microscope study of evidence for lichen weathering of 595 limestone, Mendip Hills, Somerset. Earth Surface Processes and Landforms, 12(5), 467-473. 596
Viles, H.A., Spencer, T., Teleki, K., Cox, C., 2000. Observations on 16 years of microfloral recolonization data 597 from limestone surfaces, Aldabra Atoll, Indian Ocean: implications for biological weathering. Earth 598 Surface Processes and Landforms, 25(12), 1355-1370. 599