Andrea Polanco Fernández Dynamics of the continental slope demersal fish community in the Colombian Caribbean – Deep-sea research in the Caribbean DYNAMICS OF THE CONTINENTAL SLOPE DEMERSAL FISH COMMUNITY IN THE COLOMBIAN CARIBBEAN Deep-sea research in the Caribbean by Andrea Polanco F. A Dissertation Submitted to the DEPARTMENT OF MINES (Universidad Nacional de Colombia) and the DEPARTMENT OF BIOLOGY & CHEMISTRY (Justus Liebig University Giessen, Germany) in Fulfillment of the Requirements for obtaining the Degree of DOCTOR IN MARINE SCIENCES at UNIVERSIDAD NACIONAL DE COLOMBIA (UNal) and DOCTOR RER. NAT. at THE JUSTUS-LIEBIG-UNIVERSITY GIESSEN (UniGiessen) 2015 Deans: Prof. Dr. John William Branch Bedoya (Unal) Prof. Dr. Holger Zorn (UniGiessen) Advisors: Prof. Dr. Arturo Acero Pizarro (Unal) Prof. Dr. Thomas Wilke (UniGiessen) Andrea Polanco F. (2014) Dynamics of the continental slope demersal fish community in the Colombian Caribbean - Deep-sea research in the Caribbean. This dissertation has been submitted in fulfillment of the requirements for a cotutelled advanced degree at the Universidad Nacional de Colombia (UNal) and the Justus Liebig University of Giessen adviced by Professor Arturo Acero (UNal) and Professor Thomas Wilke (UniGiessen). A mi familia y al mar… mi vida! To my family and to the sea….. my life! Después de esto, jamás volveré a mirar el mar de la misma manera… Ahora, como un pez en el agua… rodeado de inmensidad y libertad. After this, I will never look again the sea in the same way Now, as a fish in the sea… surounded of inmensity and freedom. TABLE OF CONTENTS TABLE OF CONTENTS .................................................................................................................................. I TABLE OF FIGURES .................................................................................................................................... III LIST OF TABLES .......................................................................................................................................... IV LIST OF ANNEXES ....................................................................................................................................... V ACKNOWLEDGEMENTS ............................................................................................................................. VI ABSTRACT .................................................................................................................................................... 7 RESUMEN ..................................................................................................................................................... 9 1. RATIONALE ........................................................................................................................................ 11 2. INTRODUCTION ................................................................................................................................. 13 3. OBJECTIVES AND HYPOTHESES .................................................................................................... 17 4. STUDY AREA ..................................................................................................................................... 18 5. CHAPTER I: Comprehensive assessment of Colombian Caribbean deep-sea demersal fish diversity below 200 m ................................................................................................................................................. 22 5.1. INTRODUCTION .......................................................................................................................... 22 5.2. MATERIALS AND METHODS ..................................................................................................... 24 5.3. RESULTS .................................................................................................................................... 26 5.4. DISCUSSION ............................................................................................................................... 32 6. CHAPTER II: Structure patterns of the deep-sea demersal fish fauna of the continental slope of the Colombian Caribbean .................................................................................................................................. 36 6.1. INTRODUCTION .......................................................................................................................... 36 6.2. MATERIALS AND METHODS ..................................................................................................... 39 6.2.1. Sampling strategy ................................................................................................................ 39 6.2.2. Multivariate statistical analyses ............................................................................................ 41 6.3. RESULTS .................................................................................................................................... 43 6.3.1. Distribution patterns ............................................................................................................. 43 6.3.2. Species composition and indicator species .......................................................................... 46 6.4. DISCUSSION ............................................................................................................................... 50 I 7. CHAPTER III: Spatial description of demersal fish biodiversity patterns within the Colombian Caribbean continental slope. ........................................................................................................................ 55 7.1. INTRODUCTION .......................................................................................................................... 55 7.2. MATERIALS AND METHODS ..................................................................................................... 57 7.3. RESULTS .................................................................................................................................... 60 7.4. DISCUSSION ............................................................................................................................... 66 8. CONCLUSIONS .................................................................................................................................. 72 9. BIBLIOGRAPHY .................................................................................................................................. 74 10. APPENDIX ....................................................................................................................................... 91 11. RELATED PUBLICATIONS ........................................................................................................... 175 12. CV ANDREA POLANCO FERNÁNDEZ ......................................................................................... 188 II TABLE OF FIGURES Figure 1. Area covered by this study. The dot lines and the black line indicate the bathymetric range in the continental upper slope of Colombian Caribbean sea (200-1000 m). .......................................................... 20 Figure 2. Major submarines basins, ridges and surface currents in the region. .................................................. 21 Figure 3. Deep bottoms trawls made by the different foreign and national cruises in the Colombian Caribbean Sea territorial waters. ....................................................................................................................................... 24 Figure 4. Cumulative number of deep demersal fish species found by the different cruises that collected samples in the Colombian Caribbean region until today. ............................................................................. 26 Figure 5. Holotype of Eptatretus aceroi Polanco F. and Fernholm from the Colombian Caribbean................... 31 Figure 6. INVEMAR’s research vessel B/I Ancón .................................................................................................... 39 Figure 7. Sampling method: sieving process (>500 mm). ...................................................................................... 40 Figure 8. Six discrete assemblages associated to the bathymetric and geographic distributions: 1) USa yellow; USb green; USc dark green; 2) MSa dark blue; MSb cyan; MSc dark cyan. ................................... 44 Figure 9. (A) Groups obtained by multivariate regression tree analysis based on abundance data. (B) Ordinal representation (NMDS) of the data including an ordisurf function indicating the depth where the sites are located. Identified groups are represented by different colors: MSa dark blue; MSb cyan; MSc dark cyan; USa yellow; USb green; USc dark green. ............................................................................................. 45 Figure 10. Dominant fish orders in terms of number of species (≥7%) included in the different fish assemblages of the Caribbean coast of Colombia. ....................................................................................... 47 Figure 11. Grid of 10 minutes per side used to spatialize the main cruise collection samplings points in the Colombian Caribbean territorial waters. ......................................................................................................... 57 Figure 12. Corresponding color value for the largest species range obtained in the square area is assigned. 58 Figure 13. Overlaying process of the different spatial analysis scenarios of the four main cruises with colletions of deep-sea demersal fish fauna. ................................................................................................... 58 Figure 14. Map of the Colombian Caribbean showing the distribution of areas with the highest values of deep- sea fish species in a grid of 10 minutes per side collected by the R/V Oregon. ......................................... 60 Figure 15. Map of the Colombian Caribbean showing the distribution of the areas with the highest values of deep-sea fish species in a grid of 10 minutes per side collected by the R/V Oregon II. ............................ 61 Figure 16. Map of the Colombian Caribbean showing the distribution of the areas with the highest values of deep-sea fish species in a grid of 10 minutes per side collected by the R/V Pillsbury. ............................. 62 Figure 17. Map of the Colombian Caribbean showing the distribution of the areas with highest values of deep- sea fish species in a grid of 10 minutes per side collected by the R/V . ...................................................... 63 Figure 18. Map of the Colombian Caribbean showing the distribution of the largest numbers of deep-sea fish species in a grid of 10 minutes per side overlaying the four main cruises with collections in the study area. ................................................................................................................................................................... 64 III Figure 19. Benthic terrain model of the Colombian Caribbean showing the distribution of the areas with the highest values of deep-sea fish species in a grid of 10 minutes per side and some of the geomorphological features detected around. ................................................................................................ 67 Figure 20. Image of the paleogeography and litofacies of the Guajira Basin. Evidencing the limestone sucesion present in the Continenatl shelf break and upper slope in offshore waters of El Cabo de la Vela. Take from the Seismic Atlas of Colombia (Cediel et al. 1998). ............................................................ 68 Figure 21. Image of the paleogeography and litofacies of the Cays of San Andres and Providencia basin. Evidencing the Bioclastic limestone sucesion present in the offshore waters of the Cays basin. Take from the Seismic Atlas of Colombia (Cediel et al. 1998). .............................................................................. 70 LIST OF TABLES Table 1. Different foreign and national cruises with collections in the Colombian Caribbean Sea territorial waters. ............................................................................................................................................................... 27 Table 2. Deep-sea fishes by numbers of families and species in the North Atlantic, New England, Gulf of Mexico, Caribbean region, and Colombian Caribbean slope. The number of deep-sea fish families and species include the mesopelagic fish fauna. ................................................................................................. 28 Table 3. Total number of species, genera and families of each order present below 200 m depth in the Colombian Caribbean, Caribbean Basin (Polanco and Acero in prep), and in the Gulf of Mexico. Gulf of Mexico data extracted from McEachran (2009). ............................................................................................. 29 Table 4. Total number of deep-sea demersal fish species, genera and families present below 200 m depth in the Colombian Caribbean, the Caribbean Basin, and the Gulf of Mexico excluding the orders Stomiiformes and Myctophiformes. The New England data are also presented, including the benthopelagic species living between 200 and 1000 m depth. ..................................................................... 30 Table 5. Indicator species characterizing the obtained groups with the highest association values based in species mean abundances and its frequencies of occurrence in the group. ............................................. 48 IV LIST OF ANNEXES Annex 1. Annotated species list ............................................................................................................................... 91 Annex 2. Summary information about sampling stations along the Colombian Caribbean continental slope including station number, latitude, longitude, depth, landscape (dt: delta; cs: submarine canyon; es: escarpment; fc: apron; pl: soft continental shelf; ms: plateau ), water mass (AIS: Subantartic Intermediate Water; APC: Deep-sea Caribbean Water), Temperature (T°), Salinity (S), Oxygen (O2), NA – Non available data. ......................................................................................................................................... 159 Annex 3. List of deep-sea demersal fish species caught in the Colombian Caribbean slope indicating their presence or absence among the different obtained assemblages. ........................................................... 161 Annex 4. Deep-sea demersal stations sampled in the Colombian Caribbean by the four main cruises (R/V Oregon I, R/V Oregon II, R/V Pillsbury and R/V ) that collected in the territorial waters. Coordinates and total number of species .................................................................................................................................. 168 V ACKNOWLEDGEMENTS Financial and logistic aid for the accomplishment of this work came mainly from the Center of Excellence in Marine Sciences – CEMARIN and the Marine and Coastal Research Institute – INVEMAR. Thanks to Lynne Parenti from the Smithsonian Institution - National Museum of Natural History (USNM); John Lundberg and Mark H. Sabaj-Pérez from the Academy of Natural Sciences of Drexel University (ANSP); Larry Page, Rob Robins, and Mary Brown from the Florida Museum of Natural History (UF) who supported my aplications to the awards (Short Term Visitor Travel Grant of the Smithsonian Institution; Böhlke Memorial Endowment Fund Award), or housing me for long periods of time making me feel like home. To the fish curators, collection managers, and staff of the different fish collections that I had visited during the study. Thank to Lynne Parenti, Jeff Williams, Carole Baldwin, Jeff Clayton, Sandra Raredon, Jerry Finch and Kris Murphy (USNM); John Lundberg, Mark H. Sabaj-Pérez and Kyle R. Luckenbill (ANSP); George V. Lauder, Jr., Karsten Hartel, and Andrew Williston from the Museum of Comparative Zoology of Harvard (MCZ); Larry Page, Rob Robins, and Mary Brown (UF) and all the persons in those lab who helped during the revision of the material. In my home institution I want to thank Francisco A. Arias Isaza and David Alonso Carvajal for their support to finish this process. To my Colombian advisor Profesor Arturo Acero Pizarro for being my academic life advisor, to Profesor Thomas Wilke, my German advisor, for unconditionally supporting my work. To Professor Bernd Werding for believing in my passion for marine fishes, support my application to the CEMARIN program and be around all the time and Torsten Hauffe who help me with the ecological analysis. I am deeply grateful to all the scientists and crews of the different research vessels that collected the study material. To the Colombian Marine Natural History Museum – INVEMAR personnel, all those that have gone out of the Institute and those who are still there: G. Navas, A. Gracia, Patricia Latig, L.S. Mejía, N. Ardila, A. Roa, L.M. Saavedra, G. Borrero, N. Cruz, N. Santodomingo, M. Benavides, D. Báez, A. Bermúdez, M. Díaz, P. Flórez, E. Montoya, J. Medellín, M. Garrido, N. Suárez, C. Cedeño, A. Merchán, F. Dorado, V. Yepes, J. M. Gutiérrez, Marisosl Santos and Martha Vides for their help and contributions during this study. To Sandra Raredon, who assisted with images and edition of the USNM material and Sergio Silva for helping with the edition of ANSP and MCZ images. To the laboratory of Geographic Information of INVEMAR (Nancy Barreto and Alberto Gutiérrez) for helping with the maps included in this manuscript. Special thanks to Henry, my father, who printed in my soul an example of life and gave me the tools and the perfect sentences to walk along this experience with courage. To my lovely mother Libia and my brother Sergio who support me and take my hand at all times. To my Tere, my Flower, my Artur, Caroli, Charlie, Sergio, Nats to be there as far as a phone call or a text message. To my close student partners JuanFe, Corvin, Gustav, Meli, and Carlitos for all the good moments. To my Büro partners Paulinette, Kim, Diana who cried and laughed with me during my months in Giessen. To my fishes and the sea being my source of inspiration. To all of those that helped me without hesitance during this unforgettable time, thanks for your support, understanding, friendship, and love. VI
Description: