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Burrowing Techniques, Behaviors, and Trace Morphologies of Extant Larval to Adult

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Burrowing Techniques, Behaviors, and Trace Morphologies of Extant Larval to Adult Burrowing Techniques, Behaviors, and Trace Morphologies of Extant Larval to Adult Beetles A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science Joseph J. Wislocki April 2021 © 2021 Joseph J. Wislocki. All Rights Reserved. 2 This thesis titled Burrowing Techniques, Behaviors, and Trace Morphologies of Extant Larval to Adult Beetles by JOSEPH J. WISLOCKI has been approved for the Department of Geological Sciences and the College of Arts and Sciences by Daniel I. Hembree Professor of Geological Sciences Florenz Plassmann Dean, College of Arts and Sciences 3 Abstract WISLOCKI, JOSEPH J, M.S., April 2021, Geological Sciences Burrowing Techniques, Behaviors, and Trace Morphologies of Extant Larval to Adult Beetles Director of Thesis: Daniel I. Hembree Studies of the relationship between extant trace makers, known environmental conditions, and the morphology of their biogenic structures allow for the interpretation of continental ichnofossils. This study examined the burrowing techniques, behaviors, and trace morphologies of three extant species of burrowing beetles, Tenebrio molitor, Zophobas morio, and Phyllophaga sp., from their larval to adult life stages under normal and stressed environmental conditions in a laboratory setting. Tenebrio molitor and Z. morio burrowed using their mandibles to compact the substrate, while Phyllophaga sp., burrowed by excavation and backfilling. The three species primary behaviors were locomotion, mobile deposit feeding, intermittent resting, and pupation. Larvae burrows of T. molitor and Z. morio included open boxworks, while Phyllophaga sp. larvae generated elongate backfilled burrows which terminated in an open chamber. All three species created ovoid to ellipsoidal chambers when preparing for pupation. During their adult stage, T. molitor and Z. morio created conical traces and chambers, while Phyllophaga sp. produced loosely backfilled burrows. The environmental stresses tested were related to sediment sand and water content as well as sediment compactness. Higher trace abundance was produced in sediments with decreased sand content, increased water content, and low compactness, although trace morphologies did not change. Highly 4 compacted substrates had little activity, but distinct trace morphologies. The total level of bioturbation, quantified with the ichnofabric index, produced by multiple specimens of each species in large enclosures filled with layered sediment varied from 1 (T. molitor and Phyllophaga sp.) to 2-5 (Z. morio). Quantitative analyses of the quantitative properties of the different traces showed that, despite having similar morphologies, the traces produced by the three species were dissimilar, but also showed variation within species. Understanding extant traces of beetles can help in the recognition of their ichnofossils when body fossils are not found, which can allow for improved interpretations of paleoecosystems and paleoenvironments in continental settings. 5 Dedication This is dedicated to my friends, family, supportive peers, and my professors both near and far. Geology is a pathway to many abilities, and I am glad you showed me the way. 6 Acknowledgments I would like to thank the Geologic Society of America for the Graduate Student Research Grant. I would like to thank the Ohio University Geological Science Alumni for their Graduate Research Grant. I would like to thank Ceara Purcell, Mary Reichle, and Rachel Wawrzynski for assisting me in field collection and laboratory setup. I would like to thank my wonderful committee of Dr. Dan Hembree, Dr. Alycia Stigall, and Dr. Xizhen Schenk for their support. I would like Dr. Dan Hembree for being an amazing advisor, and all-around great person. 7 Table of Contents Page Abstract ................................................................................................................................3 Dedication ............................................................................................................................5 Acknowledgments................................................................................................................6 List of Tables .......................................................................................................................9 List of Figures ....................................................................................................................10 Chapter 1: Introduction ......................................................................................................11 Chapter 2: Background ......................................................................................................14 Extant and Fossil Beetles .............................................................................................14 Ichnology and Neoichnology of Beetles ......................................................................20 Chapter 3: Methods ............................................................................................................25 Experiment 1 ................................................................................................................31 Experiment 2 ................................................................................................................31 Experiment 3 ................................................................................................................31 Experiment 4 ................................................................................................................32 Experiment 5 ................................................................................................................32 Experiment 6 ................................................................................................................32 Burrow Analysis ..........................................................................................................33 Comparison with Ichnofossils......................................................................................35 Chapter 4: Results ..............................................................................................................39 Burrow Techniques ......................................................................................................39 Larval Stage ...........................................................................................................39 Adult Stage.............................................................................................................41 Behavior .......................................................................................................................42 Larval Stage ...........................................................................................................42 Pupal Stage.............................................................................................................44 Adult Stage.............................................................................................................45 Burrow Morphology ....................................................................................................47 8 Larval Stage ...........................................................................................................47 Pupal Stage.............................................................................................................48 Adult Stage.............................................................................................................51 Environmental Controls ...............................................................................................53 Experiment 3 ..........................................................................................................53 Experiment 4 ..........................................................................................................56 Experiment 5 ..........................................................................................................59 Quantitative Analysis ...................................................................................................62 Comparison of Traces Between species ................................................................62 Comparison of Traces Between Produced in Increased Sand vs Decreased Sand Substrate .................................................................................................................63 Comparison of Traces Produced in Increased Moisture vs Decreased moisture conditions ...............................................................................................................64 Comparison of traces produced in firm vs loose substrate ....................................66 Comparison of extant traces with comparable ichnogenera ..................................68 Bioturbation and Ichnofabric .......................................................................................72 Chapter 5: Discussion ........................................................................................................80 Controls on Beetle Trace Morphology ........................................................................80 Behaviors ...............................................................................................................80 Body Morphology ..................................................................................................83 Specific Trace Maker .............................................................................................85
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