Fossil BehaviorCompendium

Fossil BehaviorCompendium

Arthur J. Boucot George O. Poinar, Jr.

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Library of Congress CataloginginPublication Data

Boucot, A. J. (Arthur James), 1924-Fossil behavior compendium / Arthur J. Boucot, George O. Poinar, Jr.

p. cm.Includes bibliographical references and index.ISBN 978-1-4398-1058-3 (hardcover : alk. paper)1. Animals, Fossil. 2. Animal behavior--Evolution. 3. Pathogenic microorganisms. I. Poinar, George O. II. Title.

QE770.B68 2010591.5--dc22 2009047092

Visit the Taylor & Francis Web site athttp://www.taylorandfrancis.comand the CRC Press Web site athttp://www.crcpress.com

vContentsPreface .......................................................................................................................................................................................xxiAcknowledgments ..................................................................................................................................................................xxiiiAuthors ..................................................................................................................................................................................... xxvIntroduction ...........................................................................................................................................................................xxvii

1Chapter Functional Morphology ........................................................................................................................................ 1

Summary ................................................................................................................................................................. 2

2Chapter Specialized, Potentially Interacting Biological Substrates ................................................................................ 3

2A. Marine Invertebrate Benthos ........................................................................................................................ 32Aa. Coral Barnacles ............................................................................................................................... 32Ab. Whale Barnacles.............................................................................................................................. 32Ac. Platyceratids .................................................................................................................................... 32Ad. Mangrove Oysters ........................................................................................................................... 42Ae. Productid Brachiopod Spines .......................................................................................................... 42Af. Phosphannulus on Paleozoic Crinoid Stems .................................................................................. 42Ag. Paleozoic Host-Specific, Pit-Forming Crinoid Epizoans ................................................................ 42Ah. HydroidSerpulid Relationship ....................................................................................................... 52Ai. CrinoidTabulate Coral Relationship .............................................................................................. 52Aj. SipunculidCoral Towing ............................................................................................................... 52Ak. Polydorid Mud Blisters in Bivalves ................................................................................................. 52Al. Boring Bivalves and Corals ............................................................................................................. 62Am. Lepadomorph Barnacles and Eurypterids, Other Substrates, and Balanoids ................................. 6

Lepadomorphs ................................................................................................................................. 6Balanomorphs .................................................................................................................................. 7Verrucomorphs ................................................................................................................................ 8

*2An. AuloporaLieoclema Association ................................................................................................... 8*2Ao. Soft-Bodied Marine Algal Substrates for Shelly Organisms .......................................................... 8*2Ap. Trypanopora and Torquaysalpinx ................................................................................................... 9*2Aq. StromatoporoidCoral Intergrowths ............................................................................................. 10*2Ar. Foraminiferal Consortium............................................................................................................. 10*2As. Decapod Inquilinism within Ammonite Shells ............................................................................ 10*2At. MicropolychaeteScleractinian Relationship ................................................................................ 10*2Au. EpizoanSponge Relationship ....................................................................................................... 10*2Av. Epibionts ........................................................................................................................................ 10*2Aw. Umbrophilic Brachiopods ............................................................................................................. 10*2Ax. MeekoporellaCrinoid Relationship ............................................................................................. 11*2Ay. BrachiopodOrthoceroid Relationship ......................................................................................... 11*2Az. Helicosalpinx................................................................................................................................. 11*2Aza. Lumbrineris flabellicolaScleractinian Relationship ................................................................... 11*2Azb. CarapusHolothurian Relationship .............................................................................................. 12*2Azc. Sphenia Nestling ........................................................................................................................... 12

Summary ............................................................................................................................................................... 12

3Chapter Mutualism ............................................................................................................................................................ 15

3a. Mycorrhiza ................................................................................................................................................. 153b. Zooxanthellae ............................................................................................................................................. 153c. Vermiforichnus and Other Epibionts .......................................................................................................... 153d. HalecostomeInoceramid Relationship ..................................................................................................... 15

vi Contents

*3e. Pinnotherid Crabs ....................................................................................................................................... 15*3f. Fig Wasps ................................................................................................................................................... 16*3g. Tube Worms, Bivalves, and Rhynchonellid Brachiopods from Deep Sea Vents, plus Crustaceans .......... 16

Hot Vents .................................................................................................................................................... 16Barite Deposit Vents ................................................................................................................................... 17Cold Seeps .................................................................................................................................................. 17Whale-Fall Communities ............................................................................................................................ 18Wood-Fall Communities ............................................................................................................................. 18Summary ..................................................................................................................................................... 19

*3h. CoralBryozoan Association ...................................................................................................................... 19*3i. BacillusBee Relationship ......................................................................................................................... 19*3j. TerebratuloidBryozoan Relationship ........................................................................................................ 20*3k. Ant and Symbiotic Scale Insect .................................................................................................................. 20*3l. Riodinidae ButterflyAnt Symbiosis .......................................................................................................... 20*3m. Acarodomatia ............................................................................................................................................. 21*3n. Lichens........................................................................................................................................................ 21*3o. Herbivory among Tetrapods ....................................................................................................................... 21*3p. AzollaAnabaena Symbiosis ...................................................................................................................... 21*3q. Luminescent Fishes .................................................................................................................................... 22*3r. Fossil Flatus: Indirect Evidence of Intestinal Microbes ............................................................................. 22

George O. Poinar, Jr.*3s. Termite and Cockroach Gut Mutualists ..................................................................................................... 25

Summary ............................................................................................................................................................... 25

4Chapter HostParasite and HostParasitoid Relationships and Disease ..................................................................... 27

4A. AnimalAnimal Relationships ................................................................................................................... 274AI. Marine ........................................................................................................................................... 27Invertebrates ............................................................................................................................................... 27

4AIa. CopepodFish ................................................................................................................ 274AIb. Bopyrid IsopodDecapod .............................................................................................. 274AIc. PearlBivalveRayTrematode or Cestode Relationship ............................................ 284AId. Sea Urchins and Parasitic Snails ................................................................................... 294AIe. Sea Urchins and Parasitic Crustaceans .......................................................................... 294AIf. Myzostomid Annelids Parasitic on Crinoids ................................................................. 294AIg. Articulate Brachiopod Mantle Dwellers ........................................................................ 304AIh. Graptolite Tubothecae .................................................................................................... 304AIi. Echinoid SpinesGastropod .......................................................................................... 304AIj. Zieglers Blisters ............................................................................................................ 31

*4AIk. BivalveTrematode Pit-Forming Relationship .............................................................. 31*4AIl. Trilobite Swellings and Borings .................................................................................... 33*4AIm. Octocoral and Ascothoracican Barnacle ....................................................................... 33*4AIn. Ammonoid and Belemnite Paleopathology ................................................................... 33*4AIo. RhizocephalanDecapod Parasitism ............................................................................. 34*4AIp. Parasitic Coralliophilidae (Gastropoda) ........................................................................ 34*4AIq. Trapeziidae Crabs and Scleractinians ............................................................................ 35*4AIr. Hohlenkehle ................................................................................................................... 35*4AIs. Abnormal Echinoid Plates ............................................................................................. 35*4AIt. Bald-Sea-Urchin Disease ............................................................................................... 35*4AIu. EulimaHolothurian ...................................................................................................... 35*4AIv. Foraminiferal Teratologies ............................................................................................ 35*4AIw. SharkIsopod ................................................................................................................. 36*4AIx. ForaminiferEchinoid ................................................................................................... 36Summary ....................................................................................................................................... 36

Vertebrates .................................................................................................................................................. 36*4AIxa. Avascular Necrosis ........................................................................................................ 36*4AIxb. Dipnoan and Chondrichthyan Dentition and Jaw Injuries ............................................. 37

Contents vii

*4AIxc. Schmorls Nodes ............................................................................................................ 37*4AIxd. Osteoporosis in Pleistocene Deer from Crete ................................................................ 37*4AIxe. Pterodactyloid Pathologies ............................................................................................ 37

4AII. Freshwater ..................................................................................................................................... 374AIIa. Unionids, Actinopterygia, and Glochidia ...................................................................... 37

*4AIIb. Ilial Frog Tumors ........................................................................................................... 37*4AIIc. Late Devonian Fish ........................................................................................................ 37

4AIII. Terrestrial ...................................................................................................................................... 384AIIIa. NematodePlanthopper, NematodeDipteran, and NematodeAnt ............................. 38

*4AIIIb. HairwormInsect ........................................................................................................... 394AIIIc. Lice and Mammals (Nits) and Birds ............................................................................. 394AIIId. Ticks and Mites as Micropredators and Potential Disease Vectors .............................. 404AIIIe. Mosquitoes as Micropredators and Potential Disease Vectors ...................................... 434AIIIf. Tsetse Flies as Micropredators and Disease Vectors ..................................................... 444AIIIg. Parasitoid WaspInsect Hosts ........................................................................................ 444AIIIh. Fleas as Micropredators and Disease Vectors ............................................................... 454AIIIi. Parasitoid Wasps and Parasitic FliesSpiders ............................................................... 454AIIIj. Human Disease .............................................................................................................. 46

*4AIIIk. Parasitic Insects Other than Wasps and Flies ................................................................ 48*4AIIIl. Trichurids and Caviomorph Rodents ............................................................................. 48*4AIIIm. ProtozoanDeer Relationship ........................................................................................ 48*4AIIIn. HyaenidNematode Relationship .................................................................................. 48*4AIIIo. LizardParasitic Nematode Relationship ...................................................................... 48*4AIIIp. Vertebrate Pathology Other than Human ...................................................................... 48*4AIIIq. Tyrannosaur Pathology .................................................................................................. 48*4AIIIr. Tetrapod Osteomyelitis .................................................................................................. 48*4AIIIs. Co-Ossified Vertebrae in Mosasaurs and Whales ......................................................... 48*4AIIIt. LizardTick Relationship ............................................................................................... 48*4AIIIu. Hyperdisease in North American Mammoths............................................................... 48*4AIIIv. Osteochondroma ............................................................................................................ 49*4AIIIw. Mycobacteriosis in a Pliocene Kangaroo ...................................................................... 50*4AIIIx. Inflammatory Arthritis (Spondyloarthropathy) ............................................................. 50*4AIIIy. Vertebrate Intestinal Parasites ....................................................................................... 50*4AIIIz. Parasite Eggs on a Bird Feather ..................................................................................... 50*4AIIIza. Trematode Eggs ............................................................................................................. 50*4AIIIzb. SpiderMermithid Relationship..................................................................................... 50*4AIIIzc. Pentastomida .................................................................................................................. 51*4AIIIzd. Strepsipteran Parasitism ................................................................................................ 51*4AIIIze. Phorid DipteranAllantonematid Nematode ................................................................. 51*4AIIIzf. Insect VectorTrypanosome Relationship ..................................................................... 52*4AIIIzg. Viruses in Biting Midges and Sand Flies ...................................................................... 52*4AIIIzh. Hypermetamorphosis ..................................................................................................... 52*4AIIIzi. Gregarine Infections in Insects ...................................................................................... 54

George O. Poinar, Jr.*4AIIIzj. Association between Fly Planidium and Mites ............................................................. 56

Summary ..................................................................................................................................................... 564B. AnimalPlant Relationship ........................................................................................................................ 56

4Ba. Hickory AphidLeaves and AphidPlant Relationships............................................................... 584Bb. Arthropod Leaf Miners ................................................................................................................. 584Bc. Leaf Galls ...................................................................................................................................... 584Bd. Stem and Petiole Galls .................................................................................................................. 594Be. Cone Galls ..................................................................................................................................... 604Bf. Scale Insects .................................................................................................................................. 60

*4Bg. Acorn Galls ................................................................................................................................... 60*4Bh. Seed and Spore Boring .................................................................................................................. 60*4Bi. Bark Beetle Mycangia ................................................................................................................... 60*4Bj. Hispine BeetleGinger Grazing .................................................................................................... 61

viii Contents

*4Bk. AraucariaBeetle Relationship and Araucaria Seed Cone Damage ............................................ 61*4Bl. NematodePlant Relationship ....................................................................................................... 61

Summary ..................................................................................................................................................... 614C. MycotaPlant Relationships ....................................................................................................................... 61

FungusPlant, FungusAlga, FungusFungus, and FungusAnimal Fossil Associations ......................... 63Symbiotic Associations .............................................................................................................................. 63

FungusPlant ................................................................................................................................. 63Parasitic Associations ................................................................................................................................. 63

FungusPlant ................................................................................................................................. 63FungusFungus ............................................................................................................................. 64FungusPlant Saprophytic Associations ....................................................................................... 64FungusAnimal Saprophytic and Parasitic Associations .............................................................. 65

Resinicolous Fungi ..................................................................................................................................... 674D. PlantPlant Relationships ........................................................................................................................... 67

4Da. Arceuthobium (Dwarf Mistletoe) .................................................................................................. 674E. MycotaAnimal Relationships ................................................................................................................... 67

*4Ea. Bark LouseFungus ...................................................................................................................... 67*4Eb. CaddisflyMold ............................................................................................................................. 68

Notes on the Origins and Evolution of Bacillus in Relation to Insect Parasitism ................................................. 68George O. Poinar, Jr.

Introduction ................................................................................................................................................ 68Antiquity of Bacteria .................................................................................................................................. 68

Origins in a Marine Environment .................................................................................................. 68Invasion of the Terrestrial Environments ....................................................................................... 68Origin of the Genus Bacillus and Early Associations with Invertebrates...................................... 68Associations of Bacillus with Insects ............................................................................................ 69Symbiotic Associations with Bacillus ........................................................................................... 69Parasitic Associations with Bacillus .............................................................................................. 70Associations of Bacillus with Vertebrates ..................................................................................... 70Specializations of Bacillus in Relation to Toxin Production ......................................................... 70

5Chapter Density and Spacing ............................................................................................................................................ 73

5a. Benthic Shell Examples .............................................................................................................................. 74Oysters ....................................................................................................................................................... 74Brachiopods ................................................................................................................................................ 74Ophiuroids, Brittle Stars ......................................................................................................................... 74Echinoids Sea Urchins and Edrioasteroids ............................................................................................. 75Balanomorph and Lepadomorph Barnacles ............................................................................................... 75Decapods .................................................................................................................................................... 75

5b. Belemnite Shoals ........................................................................................................................................ 755c. Crane Fly and Fungus Gnat Swarms: Insect Swarms ................................................................................ 765d. Shrimp Schools ........................................................................................................................................... 765e. Fish Schools ................................................................................................................................................ 765f. Dinosaur Herds ........................................................................................................................................... 765g. Mammalian Herds ...................................................................................................................................... 77

*5h. Dicynodont Herds: Mammal-Like Reptiles ............................................................................................... 77*5i. Diapsid Aggregation: Reptile ..................................................................................................................... 77*5j. Pterosaur Colony ........................................................................................................................................ 77*5k. Acridid Aggregation: Grasshoppers ........................................................................................................... 77*5l. Mass Moth Migration ................................................................................................................................. 77*5m. Ant Imago Swarms ..................................................................................................................................... 77*5n. Termite Swarms .......................................................................................................................................... 77*5o. Platypodid Swarms ..................................................................................................................................... 77*5p. Cryptic Trilobite Behavior .......................................................................................................................... 77*5q. Juvenile Millipede Aggregation ................................................................................................................. 78

Summary ............................................................................................................................................................... 78

Contents ix

6Chapter Predation and Feeding Behaviors ...................................................................................................................... 79

6A. Marine ........................................................................................................................................................ 796AI. Invertebrate.................................................................................................................................... 79

6AIa. NaticidMuricidCassid Borehole Position and Boring: Gastropods ........................... 796AIb. CrabsMollusks and GastropodBivalve ...................................................................... 826AIc. Echinoid Lantern Scratches: Aristotles Lantern Grazing Traces ................................. 836Aid. Chiton and Gastropod Radular Grazing Traces ............................................................ 846AIe. Nematode Predation on Foraminifera ........................................................................... 846AIf. Starfish Feeding on Mollusks ........................................................................................ 846AIg. Position of Boreholes in Ostracodes .............................................................................. 846AIh. CruzianaTeichichnusHalopoa Community and CruzianaTeichichnus

Nutritional Relationship ................................................................................................. 846AIi. Octopus Boreholes ......................................................................................................... 846AIj. Capulid Gastropods as Commensals on Bivalves ......................................................... 856Aik. SquidFish ..................................................................................................................... 856AIl. JuliidaeCaulerpa Relation ........................................................................................... 856AIm. Scaphopod Feeding on Foraminifera ............................................................................ 85

*6AIn. Paleozoic Predation on Gastropods ............................................................................... 85*6AIo. Stomatopod Predation on Gastropods ........................................................................... 86*6AIp. Boreholes in Brachiopods and Predation on Brachiopods in General ......................... 86*6AIq. Possible Ophiuroid, Brittle Star Predation .................................................................... 89*6AIr. Ammonite Feeding ........................................................................................................ 89*6AIs. Graptolite Predation ....................................................................................................... 89*6AIt. Predation on Echinoids .................................................................................................. 89*6AIu. Ostenocaris Predation or Scavenging ........................................................................... 90*6AIv. Invertebrate Predation on Ammonoids and Nautiloids ................................................. 90*6AIw. Predation on Bryozoans ................................................................................................. 90*6AIx. Invertebrate Predation on Trilobites .............................................................................. 90*6AIy. Predation on Crinoids .................................................................................................... 90*6AIz. Potential Cephalopod Predation of Lobsters ................................................................. 90*6AIza. Large Abalones (Haliotis) and Coldwater Kelps ........................................................... 91*6AIzb. Conus ............................................................................................................................. 91*6AIzc. Boreholes in Hederellid Bryozoans ............................................................................... 91*6AIzd. Predation on Mobergella ............................................................................................... 92*6AIze. Ostracode Scavenging ................................................................................................... 92*6AIzf. Predation on Dacryoconarids ........................................................................................ 92*6AIzg. Loosely Attached, Limpet-Like Foraminifer ................................................................ 92

Summary ....................................................................................................................................... 926AII. Vertebrate ...................................................................................................................................... 92

6AIIa. Ray Holes ....................................................................................................................... 926AIIb. Vertebrate Tooth Puncture Marks and Potential

Invertebrate-Correlated Shell Injuries and Gut Contents .............................................. 926AIIc. ArthrodireCtenacanth Shark ....................................................................................... 956AIId. Shark Feeding ................................................................................................................ 956AIIe. Vertebrate Predation on Cephalopods ........................................................................... 96

*6AIIf. Crocodilian Turtle Feeding ........................................................................................... 96*6AIIg. Plankton Feeding ........................................................................................................... 96*6AIIh. AllosaurusStegosaurus Relationship ........................................................................... 98*6AIIi. Branchiosaur Feeding .................................................................................................... 98Summary ....................................................................................................................................... 98

6B. Freshwater................................................................................................................................................... 986Ba. Carboniferous Scorpion Decomposition ....................................................................................... 986Bb. Predation Marks on Estheriids ...................................................................................................... 98

*6Bc. Ciliate Feeding .............................................................................................................................. 99*6Bd. Crayfish Predation ......................................................................................................................... 99*6Be. Unionid Predation ......................................................................................................................... 99

x Contents

6C. Terrestrial ................................................................................................................................................... 996CI. Invertebrates .................................................................................................................................. 99

6CIa. Spider Webs, Spinnerets, and Bundled Prey ................................................................. 996CIb. Dung Beetles ................................................................................................................ 1006CIc. Flesh-Eating Insects .................................................................................................... 1006CId. Reduviid BugAnts .......................................................................................................1016CIe. Gardening Ants: Leafcutter Ants and Bees ..................................................................1016CIf. Xyelidae Feeding ..........................................................................................................101

*6CIg. Petioles with Cavities Containing Coprolites .............................................................. 102*6CIh. Reduviid Bug Using Resin and Stingless Bee with Resin and Pollen ......................... 102*6CIi. Protorthopteran Spore Feeding.................................................................................... 103*6CIj. Palm Flowers with Microlepidopteran Coprolites Containing Palm Pollen ............... 103*6CIk. Beetle Containing Pollen ............................................................................................. 103*6CIl. Praying Mantis Attacked by Ants................................................................................ 103*6CIm. Whip Scorpion and Insect Prey ................................................................................... 103*6CIn. Plant-Feeding Snail ...................................................................................................... 103*6CIo. Dolichopodid Fly with an Enchytraeid Worm Fragment ............................................ 103*6CIp. Coccid Salivary Sheaths .............................................................................................. 103*6CIq. Elaterid Feeding ........................................................................................................... 103*6CIr. Insect Mouthparts ........................................................................................................ 103*6CIs. Oribatid Mite Feeding ................................................................................................. 103*6CIt. Insect Herbivory .......................................................................................................... 104*6CIu. AntPseudoscorpion Relationship ............................................................................... 105*6CIv. Blood-Feeding Dipterans ............................................................................................. 105*6CIw. Piercing and Sucking ................................................................................................... 105*6CIx. Empidid Fly and a Chironomid ................................................................................... 106*6CIy. Phorid Fly Attacked by an Insect Larva ...................................................................... 106*6CIz. Ground Sloth DungSciarid Larvae ............................................................................ 106Summary ..................................................................................................................................... 106

6CII. Vertebrates ................................................................................................................................... 1066CIIa. Fighting Dinosaurs ................................................................................................... 1066CIIb. Hadrosaurian Dinosaur Diet ........................................................................................ 1076CIIc. Owl Pellets ................................................................................................................... 1076CIId. Felid Activities ............................................................................................................. 1086CIIe. Sloth Diets.................................................................................................................... 1086CIIf. Bite Marks on Fossil Nuts and Mammal Bones .......................................................... 1086CIIg. Insectivorous Bats ........................................................................................................ 1096CIIh. Pangolin Feeding on Ants and Termites.......................................................................1106CIIi. Beaver Wood Cutting and Beaver-Gnawed Mastodon Molars .....................................110

*6CIIj. Venom-Conducting Reptilian Teeth .............................................................................110*6CIIk. Long-Fingered, Mammalian Insect Seekers ................................................................110*6CIIl. Frozen Pleistocene Mammals .......................................................................................111*6CIIm. Mammoth Diet ..............................................................................................................111*6CIIn. Propalaeotherium Stomach Contents ...........................................................................111*6CIIo. Insectivore Diets ...........................................................................................................111*6CIIp. Piciform Bird with Stomach Contents ..........................................................................111*6CIIq. Eurotamandua Feeding ................................................................................................111*6CIIr. Rodent, Horse, and Even-Toed Ungulate Feeding ........................................................111*6CIIs. Diprotodon and Thylacoleo ..........................................................................................111*6CIIt. Velociraptorine Feeding on a Pterosaur .......................................................................112*6CIIu. Early Cretaceous Seed-Eating Bird from China ..........................................................112*6CIIv. Late PleistoceneHolocene Caprinid Diet....................................................................112*6CIIw. Jurassic Salamander Diet ..............................................................................................113*6CIIx. Copes Rule and Hypercarnivory ..................................................................................113*6CIIy. Ursid Activities .............................................................................................................113*6CIIz. TurtleCeltis Feeding ...................................................................................................113

Contents xi

*6CIIza. Eocene Mammalian PredatorPrey Example ..............................................................113Summary ......................................................................................................................................113

6D. Marine, Freshwater, and Terrestrial ..........................................................................................................1136Da. Vertebrates Swallowing Other Vertebrates ..................................................................................1136Db. Gastrolith-Mediated Digestion? ...................................................................................................115

*6Dc. Petalodontid Gut Contents ............................................................................................................116*6Dd. MosasaurShark ...........................................................................................................................116*6De. Crocodilian Mammal Feeding .....................................................................................................116

Summary ....................................................................................................................................................1166E. Fungal ........................................................................................................................................................117

*6Ea. Nematophagous Fungi ..................................................................................................................117

Chapter 7 Communication ..................................................................................................................................................119

Auditory ...............................................................................................................................................................119Vision .................................................................................................................................................................. 120Luminous Organs ................................................................................................................................................ 120Chemoreceptors and Tactile Organs .................................................................................................................... 120Electrical Organs ................................................................................................................................................. 120Lateral Line Organs ............................................................................................................................................. 121

8Chapter Trace Fossils and Their Formers ..................................................................................................................... 123

8A. Marine ...................................................................................................................................................... 1238Aa. Limuloid Trails ............................................................................................................................ 1238Ab. Ophiomorpha and Callianassa: Crustacea ................................................................................. 1248Ac. Decapod Trails ............................................................................................................................ 1248Ad. Crab Burrows .............................................................................................................................. 1248Ae. Echinoid Burrows and Traces ..................................................................................................... 1248Af. Ophiuroid Resting Traces ............................................................................................................ 1248Ag. Orthoceroid Traces ...................................................................................................................... 124

*8Ah. Fish Scraping ............................................................................................................................... 124*8Ai. Macroboring into Hard Substrates .............................................................................................. 125*8Aj. Bivalve Trace Former .................................................................................................................. 125*8Ak. Isopod Traces .............................................................................................................................. 125*8Al. Tellinoidean Bivalve Trace .......................................................................................................... 125*8Am. Sabia Pits ..................................................................................................................................... 125*8An. Limpet Traces .............................................................................................................................. 125*8Ao. Anomid Bivalve Traces ............................................................................................................... 125

Summary ................................................................................................................................................... 1258B. Freshwater................................................................................................................................................. 125

8Ba. Caddisfly Cases ........................................................................................................................... 1258Bb. Lungfish Burrows ........................................................................................................................ 127

*8Bc. Crayfish Burrows ......................................................................................................................... 128*8Bd. Fish Traces................................................................................................................................... 128Summary ................................................................................................................................................... 128

8C. Terrestrial ................................................................................................................................................. 1288CI. Invertebrate.................................................................................................................................. 128

8CIa. Mud Wasp Nests .......................................................................................................... 1288CIb. Leaf-Cutting Bees ........................................................................................................ 1288CIc. Mining Hymenopterans ............................................................................................... 1288CId. Aleyrodidae Pupal Case .............................................................................................. 130

*8CIe. Coleopteran Pupal Chambers and Possible Scarabid Beetle Burrows ........................ 130*8CIf. Caterpillar Coprolites Misidentified as Araliaceae Fruits ........................................... 130*8CIg. Neuropteroid Cocoon ................................................................................................... 130*8CIh. Chrysomelid Larval Case ............................................................................................ 130

xii Contents

*8CIi. Earthworm Burrows .................................................................................................... 130Summary ..................................................................................................................................... 130

8CII. Vertebrate .....................................................................................................................................1318CIIa. Vertebrate Tracks ..........................................................................................................1318CIIb. Daemonelix .................................................................................................................. 1328CIIc. Dabble Marks and Accompanying Tracks, Plus Associated Skeletal Material .......... 1328CIId. Pocket Mouse and Kangaroo Rat Burrows .................................................................. 132

*8CIIe. Cicioniiformes-Like Tracks ......................................................................................... 132*8CIIf. Artiodactyl Tracks ....................................................................................................... 132*8CIIg. Human Footprints ........................................................................................................ 132Summary ..................................................................................................................................... 132

9Chapter Specialized Substrates ....................................................................................................................................... 133

9A. Marine ...................................................................................................................................................... 1339Aa. Shell-Boring Fungi and Algae ..................................................................................................... 1339Ab. Clionid Boring Sponges .............................................................................................................. 1339Ac. BryozoanSnailHermit Crab Complex and HydrozoanGastropod Complex ......................... 1339Ad. Helicotaphrichnus: Trace Fossil ................................................................................................. 1359Ae. ThylacusLarger Gastropod Relationship .................................................................................. 1359Af. Rock- and Wood-Boring Bivalves ............................................................................................... 135

Rock Borings ............................................................................................................................... 135Teredinid and Pholad Bored Wood .............................................................................................. 135

9Ag. Acrothoracican Barnacles ........................................................................................................... 136*9Ah. Acrothoracican BarnacleHermit Crab Shell ............................................................................. 137*9Ai. Arachnostega ............................................................................................................................... 137*9Aj. Algal and Fungal Paleozoic Microborings in Corals .................................................................. 137*9Ak. Limpets and Bone Substrates ...................................................................................................... 137*9Al. Limpet Depressions in Ammonites ............................................................................................. 137*9Am. Asteriastoma cretaceum Breton, 1992 ........................................................................................ 137*9An. Savazzis Leaning Tower of Pisa Morphology ........................................................................ 137*9Ao. Hard Substrates ........................................................................................................................ 138

Cornulites .................................................................................................................................... 139*9Ap. Dendroid Graptolite Substrates ....................................................................................................141*9Aq. PygmaeoconusHyolithid ............................................................................................................141*9Ar. Meiofauna .....................................................................................................................................141

Summary ....................................................................................................................................................1419B. Terrestrial ..................................................................................................................................................141

9Ba. Beetle Boring in Wood .................................................................................................................1419Bc. Termite Borings in Wood ............................................................................................................ 1429Bd. Wood-Boring Bees .......................................................................................................................143

*9Bf. Wood-Boring Mites ......................................................................................................................143Summary ....................................................................................................................................................143

1Chapter 0 Sexual Behavior ................................................................................................................................................. 145

10A. Terrestrial ................................................................................................................................................. 14510AI. Plants ........................................................................................................................................... 145

10AIa. Reproduction in Lower Plants ..................................................................................... 14510AII. Invertebrates ................................................................................................................................ 146

10AIIa. Spider Sperm Pumps and Copulation .......................................................................... 14610AIIb. Mating Insects ..............................................................................................................147Side-by-Side Mating ................................................................................................................... 150Eggs, Oviposition, and Maternal Care in Amber ........................................................................ 152George O. Poinar, Jr.10AIIc. Other Evidence of Insect Egg Laying .......................................................................... 15810AIId. Spider Cocoons, Eggs, and Spiderlings ....................................................................... 159

Contents xiii

10AIIe. Caterpillars and Dipteran Larvae and Nymphs: Immature Insect Stages ................... 15910AIIf. Beetle Eggs Deposited on a Leaf ..................................................................................161

*10AIIg. Cockroach Ootheca ......................................................................................................161*10AIIh. Mate Guarding in Gerrids ............................................................................................161*10AIIi. Phasmida Eggs ..............................................................................................................161*10AIIj. Odonata Eggs Laid on Leaves ......................................................................................161*10AIIk. Coleopteran Pupal Cases ............................................................................................. 162*10AIIl. Collembolan Sperm and Insect Spermatophores......................................................... 162*10AIIm. Mosquito and Biting Midge Mating Swarms (Leks) ................................................... 162*10AIIn. Isopod with Young ....................................................................................................... 162*10AIIo. Oviposition Notches..................................................................................................... 162*10AIIp. Opilione Sexual Organs ............................................................................................... 162*10AIIq. Platyhelminth Eggs ...................................................................................................... 162

Summary ................................................................................................................................................... 16310B. Aquatic Invertebrates................................................................................................................................ 163

10Ba. Dimorphism and Brood Care in Ostracodes ............................................................................... 16310Bb. Crepidulid Gastropod Sex Changes ............................................................................................ 16410Bc. Ammonoid Egg Sacs ................................................................................................................... 16410Bd. Trilobite and Crab Clusters ......................................................................................................... 16410Be. Gastropod Eggs and Bivalve Brood ............................................................................................ 164

*10Bf. Patagonian Oyster Reproduction ................................................................................................. 164*10Bg. Probable Hirudinean and Earthworm Cocoons .......................................................................... 165*10Bh. Ctenophore Gonads ..................................................................................................................... 165*10Bi. Graptolite Regeneration ........................................................................................................... 165*10Bj. Crab Larvae ................................................................................................................................. 166*10Bk. Cladoceran Eggs .......................................................................................................................... 166*10Bl. Estherian Crustacean Egg Brooding ........................................................................................... 166*10Bm. Foraminiferal Plastogamy ........................................................................................................... 166*10Bn. Argonaut Paper Nautilus Egg Cases ........................................................................................... 166*10Bo. Cupuladrid Bryozoan Reproduction ........................................................................................... 166*10Bp. Copulating Gastropods ................................................................................................................ 166Summary ................................................................................................................................................... 166

10C. Vertebrates ................................................................................................................................................ 16710Ca. Vertebrate Eggs and Egg Cases ................................................................................................... 167

Chondrichthyan Cartilaginous Fish ......................................................................................... 167Amphibian ................................................................................................................................... 168Reptilian ...................................................................................................................................... 168Dinosaur ...................................................................................................................................... 168Pterosaur ...................................................................................................................................... 169Lizards ......................................................................................................................................... 169Gecko ........................................................................................................................................... 169Turtles .......................................................................................................................................... 169Crocodilian .................................................................................................................................. 169Bird .............................................................................................................................................. 169Uncertain ..................................................................................................................................... 169

10Cb. Claspers and Pregnant Chondrichthyans .................................................................................... 16910Cc. Anuran Pollux, Egg-Laying, and Larval Amphibians ................................................................ 16910Cd. Nesting and Parental Care among Dinosaurs and Crocodilians ..................................................17010Ce. Ichthyosaur and Mosasaur Birth Delivery Attitude .....................................................................17010Cf. Possible Nursing ...........................................................................................................................17110Cg. Otarioid Seal Rookery ..................................................................................................................17110Ch. Internal Fertilization in Placental Mammals (Bacula and Pregnant Females) ............................17110Ci. Fighting Phytosaurs ......................................................................................................................17110Cj. Fish Nests .................................................................................................................................... 172

*10Ck. Viviparity or Ovoviviparity in Fishes ......................................................................................... 172Summary ................................................................................................................................................... 172

xiv Contents

1Chapter 1 Parental Care ......................................................................................................................................................175

*Invertebrates .......................................................................................................................................................175*Vertebrates ..........................................................................................................................................................176

1Chapter 2 Depth Behavior .................................................................................................................................................. 177

*12a. Heteropods ................................................................................................................................................ 177*12b. Paleoecologic Dissonance ........................................................................................................................ 177*12c. Pelagic Trilobite Depth Selection ..............................................................................................................178*12d. Depth Distributions .................................................................................................................................. 179

1Chapter 3 Phoresy ................................................................................................................................................................181

13a. Pseudoscorpions ........................................................................................................................................18113b. Mites with a Midge and with a Bark Beetle and Other Insects................................................................ 182

*13c. Macrochelid Mites and Drosophilid Flies ................................................................................................ 182*13d. Mammalian Hair Epizoochory ................................................................................................................ 182*13e. Female Fig Wasps and Nematodes ........................................................................................................... 183*13f. Swinging Springtails: Phoretic Behavior in Fossil Collembola ............................................................... 183

George O. Poinar, Jr.Summary ............................................................................................................................................................. 184

1Chapter 4 Defense ................................................................................................................................................................ 185

14a. Operculate Gastropods ............................................................................................................................. 18514b. Serpulid Worm Operculae ........................................................................................................................ 18514c. Cephalopod Ink Sacs ................................................................................................................................ 18514d. Camouflage ............................................................................................................................................... 186

Cephalopods ............................................................................................................................................. 187Echinoderms ............................................................................................................................................. 187Brachiopods .............................................................................................................................................. 187Bivalvia ..................................................................................................................................................... 187Gastropods ................................................................................................................................................ 187Monoplacophorans ................................................................................................................................... 187Lobopodians ............................................................................................................................................. 187Trilobites ................................................................................................................................................... 187Hyolithids ................................................................................................................................................. 187Insects ....................................................................................................................................................... 187Bird Feathers ............................................................................................................................................ 187Fish ........................................................................................................................................................... 188Amphibia .................................................................................................................................................. 188Reptilia ..................................................................................................................................................... 188Cases of Camouflage in Amber ................................................................................................................ 188George O. Poinar, Jr.Summary ....................................................................................................................................................191

14e. Autotomy ...................................................................................................................................................191Echinoderms ..............................................................................................................................................191Non-Insect Arthropods ............................................................................................................................. 192Insects ....................................................................................................................................................... 192Vertebrates ................................................................................................................................................ 192Behavioral Implications from Lizards Preserved in Amber ..................................................................... 193Summary ................................................................................................................................................... 194

14f. Enrollment ................................................................................................................................................ 19414g. Spines and Thorns .................................................................................................................................... 19414h. Belemnite Swimming and Other Cephalopods ........................................................................................ 195

*14i. Trilobites Sheltered within Nautiloid Shells and Crustaceans within Ammonites .................................. 195*14j. Stingray Spines and Other Venomous Fish Spines .................................................................................. 195*14k. Onychophoran Slime Secretion ................................................................................................................ 195

Contents xv

*14l. Soldier Beetle ........................................................................................................................................... 19614m. Cryptorhynch Weevil Locking Mechanism ............................................................................................. 196Summary ............................................................................................................................................................. 196

1Chapter 5 Carrier Shells ..................................................................................................................................................... 199

1Chapter 6 Pollination Ecology ............................................................................................................................................ 201

1Chapter 7 Social Insects ...................................................................................................................................................... 203

17a. Stratigraphic Ranges................................................................................................................................. 203Wasps ........................................................................................................................................................ 203Ants ........................................................................................................................................................... 203Mesozoic Bees .......................................................................................................................................... 203

Synopsis of Fossil Bees ............................................................................................................... 203Termites .................................................................................................................................................... 205

17b. Nest Building ............................................................................................................................................ 205Termite Frass ............................................................................................................................................ 206

17c. Workers Carrying Larvae and Pupae ....................................................................................................... 20617d. Fungus-Gardening Ants ........................................................................................................................... 20617e. Scale and Ant Relation ............................................................................................................................. 20617f. Trophallaxis .............................................................................................................................................. 20617g. Termite Nasutes ........................................................................................................................................ 20717h. Wasp Nest Controversy ............................................................................................................................ 207

*17i. Termite Bugs............................................................................................................................................. 207*17j. Ant Mimic ................................................................................................................................................ 207*17k. Termite Larvae ......................................................................................................................................... 207*17l. Termite Nest Associates ........................................................................................................................... 207*17m. Ant Nest Beetles ....................................................................................................................................... 207*17n. Army Ant and Prey .................................................................................................................................. 207*17o. Replete Ant ............................................................................................................................................... 207*17p. Orchid Bees .............................................................................................................................................. 207*17q. Bee Pollen Feeding ................................................................................................................................... 208*17r. Weaver Ants ............................................................................................................................................. 208

Summary ............................................................................................................................................................. 208

1Chapter 8 Long-Range Migration ..................................................................................................................................... 209

1Chapter 9 Molting ................................................................................................................................................................211

19a. Trilobites ....................................................................................................................................................21119b. Decapods ...................................................................................................................................................21119c. Insects ........................................................................................................................................................211

2Chapter 0 Sensitive Plants .................................................................................................................................................. 213

2Chapter 1 Reptilian and Mammalian Burrows and Dens ............................................................................................... 215

Mammalian .......................................................................................................................................................... 215Reptilian .............................................................................................................................................................. 215Amphibian ............................................................................................................................................................216

2Chapter 2 Vertebrate Endocranial Casts ...........................................................................................................................217

22a. Mammals and Pterosaurs ..........................................................................................................................21722b. Hominid Handedness ................................................................................................................................217

*22c. Hominids ...................................................................................................................................................217

xvi Contents

2Chapter 3 Preening ..............................................................................................................................................................219

2Chapter 4 Grain-Size Selectors .......................................................................................................................................... 221

24b. Magnetite Ballast Grains in Sand Dollars (Weight Belts) .................................................................... 221

*2Chapter 5 The Seagrass Community Complex .............................................................................................................. 223

*2Chapter 6 Shelter .............................................................................................................................................................. 225

*2Chapter 7 Flying and Gliding Vertebrates ..................................................................................................................... 227

*2Chapter 8 Possible GeneticDevelopmental Defects...................................................................................................... 229

*28a. Transposed Bivalve Hinge Lines and Double Siphonal Grooves in a Gastropod ................................. 229*28b. Rhinoceratid Tooth and Other Possible Genetic Defects ...................................................................... 229

*2Chapter 9 Teratologies ...................................................................................................................................................... 231

*Marine Invertebrates ......................................................................................................................................... 231*29a. Trilobites ................................................................................................................................................ 231*29b. Bivalves .................................................................................................................................................. 231*29c. Brachiopods ........................................................................................................................................... 232*29d. Echinoids ............................................................................................................................................... 232*Terrestrial Invertebrates .................................................................................................................................... 233

*3Chapter 0 Disease .............................................................................................................................................................. 235

*30a. Metastatic Cancer in the Jurassic .......................................................................................................... 235*30b. Diseased Elephant Upper Jaw and Deformed Teeth.............................................................................. 235*30c. Bone Fractures ....................................................................................................................................... 235*30d. Ceratopsian Stress Fracture ................................................................................................................... 236

*3Chapter 1 Marine Molluscan Larval Types and Their Behavior ................................................................................. 237

*3Chapter 2 Competition Involving Bryozoans ................................................................................................................. 239

*3Chapter 3 Lost Behaviors and Their Vestigial Evidence .......................................................................................... 241

*3Chapter 4 Stunting ............................................................................................................................................................ 243

*3Chapter 5 Oceanic vs. Neritic .......................................................................................................................................... 245

*3Chapter 6 Human Behavior .....................................................................