PSAMMOLITTORAL MARINE TARDIGRADA FROM COLEMAN BEACH, CALIFORNIA by Albert Carranza A thesis submitted to Sonoma State University in partial fulfillment of the requirements for the degree of MASTER OF ARTS in Biology 0.Y..1-(-?:1-( fJh__ Date Copyright 1996 By Albert Carranza ii AUTHORIZATION FOR REPRODUCTION OF MASTER'S THESIS I grant permission for the reproduction of the thesis in its entirety, without further authorization from me, on the condition that the person or agency requesting reproduction absorb the cost and provide proper acknowledgement of authorship. DATE: _____. I L...../ ...!J{' ___ Signature Street Adress City, State, lip iii PSAMMOLITTORAL MARINE TARDIGRADA FROM COLEMAN BEACH, CALIFORNIA Thesis by Albert Carranza ABSTRACT Purpose of the Study: Most of what is known about marine tardigrades is based on research conducted in Europe and along the east coast of the United States. Prior to this study only two cursory surveys of marine sand-dwelling taxa had been conducted along the coast of California. In each case a new species was described, but the number of species recovered from any single beach did not exceed three, giving the impression that species diversity along the California coast is unusually low. This study is a survey of the taxonomic composition, intertidal distribution, and temporal variation of marine interstitial tardigrades from Coleman Beach, Sonoma County, California. Procedure: Sand cores were collected on March 23, July 23, and November 3, 1994 at Coleman Beach, Sonoma County, California. These were treated as representative samples for spring, summer, and fall respectively. The cores were taken along an 1 8 meter transect within the upper intertidal area (mean lower low water to mean higher high water) in 3.5cm diameter by 10cm depth (1 OOcm3) sections to a maximum depth of 60cm. Findings: Eight species representing three arthrotardigrade families were encountered. Two of these are new records for the west coast of North America and two others are new to science. In the family, Batillipedidae Batillipes tridentatus were the most abundant in the spring and B. sonomacus n. sp. were most abundant in fall. iv Halechinischus remanei was the single most common member of the family Halechiniscidae and the most abundant species in summer samples. H. californiensis n. Spa occurred along with its congener in very low abundance. The Halechiniscidae was also reprsented byTanarctus arborspinosus in summer and T. ramazzotti in fall. Stygarctus spinifer and S. bradypus comprised the family Stygarctidae. The halechiniscids and stygarctids typicaly occurred in the deepest (40-60cm) cores at mid to high tide level. Conclusions: The presence of seven different species of tardigrades in the summer samples alone establishes that species diversity along the California coast is as rich as that found elsewhere. The distribution patterns observed during this study conform to the findings in both Europe and the eastern United States. Seasonal variation patterns suggest an alternation in abundance between congeneric species at Coleman Beach and may be the out come of interspecific competition. Chair: MA Program: Biology Sonoma State University Date: ~j1., dft v ACKNOWLEDGEMENTS I would like to thank Dr. Leland W. Pollock of Drew University for providing literature, advice and species confirmations. I also thank Dr. Diane R. Nelson of East Tennessee State University for her advice and for providing some essential literature. I am most grateful to Dr. Colin O. Hermans for his support and guidance throughout this project. For their assistance in the preparation of this manuscript, wish to thank Dr. Colin O. Hermans, Dr. Chris K. Kjeldsen and Marilyn Cannon. I also wish to thank the following people for their support, assistance and inspiration; Dr. Jean Renaud-Mornant, Dr. Mathew James, Dr. Joe H. Brumbaugh, Dr. John Hopkirk, Sam Horton, Ron Dudash, Rosemary Saunders, Jeff Hughey, William F. Samaras, Ann Hill, Walter Brelje, Ben Race, Wilford E. Dickinson and Linda M. Dickinson. This work is dedicated to Megan and our daughter, Hannah. vi LIST OF TABLES TABLE 1 . Salinity, temperature and beach slope readings at Coleman Beach...... a ................................. , ••••••••••••••••••••••••••••" .................. JI ••• 1 5 2. Cumulative frequency series assesment of sediment............... 16 3. Morphometry of BatilJipes tridentatus and B. sonomacus n.sp ......................................................"................................ 24 4. Morphometry of Halechiniscidae from Coleman Beach .............. 35 5. Morphomometry of Stygarctus spinifer and S. bradypus from Coleman Beach ....................................................41 6. Arthrotardigrada recovered during Spring 1 994 .........................43 7. Arthrotardigrada recovered during Summer 1994 ......................46 8. Arthrotardigrada recovered during Fall 1994...............................49 vii LIST OF FIGURES Figure 1 . Feeding structures of the tardigrada ................................................2 2. Schematic representation of the cephalic appendages of an arthrotardigrade ..............................................................................4 3. Foot Morphology of the Heterotardigrada ........................................5 4. Diagramatic representation of eutardigrade claws ....................6 5. Cumulative frequency curve for sediment assessment data ....................................................................................1 6 6. Intertidal distribution and abundance, Spring 1994..................43 7. Vertical distribution within substratum at 10m from MLLW , Spring 1994 ..............................................................................44 8. Horizontal (intertidal) distribution of Tardigrada Summer 1994 ...........................................................................................,. ... 4 7 9. Distribution and abundance of Tardigrada in samples collected Summer 1994......................................................................47 10. Horizontal distribution of Tardigrada observed during Fall 1994.................................................................... "......... t ........................ 50 11 . Distribution and abundance of specimens in samples collected Fall 1994..... ".. ,..................".....,.............................. 11." ••"..............50 12. Distribution of Batillipes tridentatus and B. sonomacus Fall 1994...................."."."... ""' ......".......... ,, ......................................................51 13. Relative Temporal Abundance of all Tardigrade Taxa..............52 viii LIST OF PLATES List of abbreviations ..............................................................................................65 Plate 1 Batillipes sonomacus. n. sp...............................................................66 2 Batillipes tridentatus Pollock, 1989 ...........................................67 3 Halechiniscus remanei remanei Schulz, 1955 ..........................68 4 Halechinscus californiensis n. sp...................................................69 5 Tanarctus arborspinosus Lindgren, 1971 ....................................70 6 Tanarctus ramazzotti Renaud-Mornant, 1975 .......................... 71 7 Stygarctus spinifer Hiruta, 1985...................................................72 8 Stygarctus bradypus Schulz, 1951 ................................................73 ix TABLE OF CONTENTS List of Tables....................................•........... ,......................... ,........................ wvii List of Figures..................................................................................................viii List of Plates ........•......... ~ ........................................... ,.......................................... 1x Chapter I INTRODUCTION ...............................................................................................1 General Description of the Phylum ................................................. 1 Systematics of the Phylum ................................................................3 Ecology of the Phylum .........................................................................6 Phylogenetic Affinities .........................................................................8 Marine Tardigrada in Western North America .............................9 I I MATERIALS AND METHODS........................................,............................... iII •••• ~ 12 III RESULTS..... "....... "............... "' .......... ,,. ......... " ...... ,.................................. ,..... 15 Physica I Parameters ....."......,...".".......... "' ................................"................,'" 15 Systematic Account ..•...."" ....... ,.............,...,...,', .. "' ......,......,",.....•.....,......, 1 7 Batillipes sonomacus n. sp.........................................................19 Batillipes tridentatus Pollock, 1989......................................22 Halechiniscus remanei remane; Schulz, 1955.....................27 Ha/echinscus californiensis n. sp.............................................28 Tanarctus arb orspin os us Lindgren, 1971 .............................32 Tanarctus ramazzotti Renaud-Mornant, 1975 .....................33 Stygarctus spinifer Hiruta, 1985 .............................................37 Stygarctus bradypus Schulz, 1951 ...........................................38 Distribution and Temporal Abundance ..........................................42