Life in the Open Ocean

Life in the Open Ocean
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Life in the Open Ocean Life in the Open Ocean: The Biology of Pelagic Species provides in-depth coverage of the different marine animal groups that form the communities inhabiting the ocean’s pelagic realm. This comprehensive resource explores the physical environment, foraging strategies, energetics, locomotion, sensory mechanisms, global and vertical distributions, special adaptations, and other characteristics of a wide array of marine taxa. Bringing together the most recent information available in a single volume, authors Joseph J. Torres and Thomas G. Bailey cover the Cnidaria (stinging jellies), the ctenophores (comb jellies), pelagic nemerteans, pelagic annelids, crustaceans, cephalopods and pelagic gastropods, invertebrate chordates, as well as micronektonic and larger fishes such as sharks, tunas, mackerels, and mahi-mahi. Detailed chapters on each pelagic group describe internal and external anatomy, classification and history, feeding and digestion, bioluminescent systems and their function, reproduction and development, respiration, excretion, nervous systems, and more. The first book of its kind to address all of the major animal groups comprising both the swimmers and drifters of the open sea, this important resource: Explains how different animals have adapted to live in the open-ocean environment Covers all sensory mechanisms of animals living in the pelagic habitat, including photoreception, mechanoreception, and chemoreception Treats the diverse micronekton assemblage as a community Includes a thorough introduction to the physical oceanography and properties of water in the pelagic realm Life in the Open Ocean: The Biology of Pelagic Species is an excellent senior-level undergraduate and graduate textbook for courses in biology and biological oceanography, and a valuable reference for all those with interest in open-ocean biology.

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Joseph J. Torres. Life in the Open Ocean

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

Life in the Open Ocean. The Biology of Pelagic Species

Preface

Acknowledgments

1 Physics and the Physical Environment

The Vastness of the Open Ocean

The Properties of Water

Density

Viscosity

Reynolds Number

Drag

Temperature

The Oceans and Ocean Basins

Ocean Circulation

Surface Currents: Ocean Gyres and Geostrophic Flow

Coriolis Force

Ekman Transport

Ocean Gyres and Geostrophic Flow

Upwelling

Deep‐Ocean Circulation

Water Masses

Oxygen

Pressure

Sound

Light

Absorption and Scattering

Traditional Depth Zones in the Ocean

Concluding Thoughts

References

2 Physiological Accommodation to Environmental Challenges

Temperature

Terms

Temperature Effects on Survival: The Tolerance Polygon

Temperature Effects on Rate Processes – The Q10 Approximation

Patterns of Thermal Acclimation

Climatic Adaptation in Ectotherms

Temperature Compensation via Changes in Enzyme Concentration: The Quantitative Strategy for Short‐term Change

Compensation via Changes in Enzyme Quality – Isozymes, Allozymes, and Temperature Adaptation

What Properties of Enzymes Can Be Changed?

Lipids and Temperature

A Membrane Primer

Pressure

Early Work

Later Work

Whole Animal Work

Molecular Mechanisms of Adaptation to Pressure

Pressure and Membranes

Oxygen

Box 2.1 An Aside on Units

Severity of Oxygen Minima, “Dead Zones,” and the Intertidal

Adaptations to Oxygen Minima. The Aerobic Strategy

Salinity

Depth

Concluding Thoughts

References

3 The Cnidaria. Introduction

Classification. History

Classification Schemes

Phylum Cnidaria. Subphylum Medusozoa

Subphylum Anthozoa

Subphylum Myxozoa

The Hydromedusae. Morphology Basics

Morphological Detail and Life Histories

The Scyphomedusae. Basic Characteristics

Morphological Detail and Life Histories. General

Coronatae

Semaeostomae

Rhizostomae

The Cubomedusae

Foraging Strategies

General Considerations

The Cnidae

Venoms

Interaction with Prey

Direct Interception

Encounter Zone

Anthomedusae

Leptomedusae

Trachymedusae

Narcomedusae

Scyphomedusae

The Model

Swimming and Hunting Behavior

Water Flow and Swimming

Attraction Between Predator and Prey

Diets, Feeding Rates, and Impacts on Prey Populations

Rogue Hydroids: Predatory Polyps in the Midwater

Feeding in the Cubomedusae

Locomotion

The Mesoglea

Nerve Nets and Nervous Control of Swimming

Senses and Sensory Mechanisms

The Siphonophores

Terminology and Affinities of Siphonophore “Persons”

Whole Animal Organization

Life Histories

The Siphonophore Conundrum

Feeding. Fishing Behavior

Digestion

Diets and Selectivity

Ecological Importance

Locomotion

Buoyancy

Vertical Distribution

Diurnal Vertical Migration

Geographical Distribution

Organization and Sensory Mechanisms

Epithelial Conduction vs. Neural Conduction

The Cnidaria Formerly Known as Chondrophora. Classification

Characteristics of the Chondrophoran Medusa

Evolution Within the Chondrophora

Feeding in the Chondrophora

Locomotion

References

4 The Ctenophora. Introduction

Classification. History

Classification Schemes

Ctenophore Basics

Morphology. Cydippida

Lobata

Cestida

Beroida

Platyctenida

Ganeshida

Thalassocalycida

Foraging Strategies. General Considerations

Interaction with Prey. The Cydippids

The Lobates

The Cestids

The Beroids

The Platyctenids

The Ganeshids

The Thalassocalycids

Specialists

Diet, Feeding Rates, and Impacts on Prey Populations

Ctenophores as Invasive Species

Digestion

Nerves and Sense Organs: Coordination and Conduction

Locomotion

Distribution

Ctenophores and Evolution

References

5 The Nemertea. Introduction

Classification. History

Of Germ Layers and Body Cavities

Hydrostatic Skeletons

Classification

Morphology

Proboscis Apparatus

The Pelagic Body Form and Locomotion

Color

Nervous System

Sense Organs

Circulatory System

Excretory System

Digestive System

Reproduction

Development

Foraging Strategies

Vertical and Geographic Distributions

References

6 The Annelida. Introduction

History

Classification

Box 6.1 Useful Terms

Phylum Annelida

Class Polychaeta

Subclass Echiura

Subclass Errantia

Class Clitellata

Subclass Oligochaeta

Subclass Hirudinea

The Pelagic Polychaetes

Polychaete Subclass Errantia

Order Phyllodocida

Polychaete Subclass Sedentaria

Order Terebellida

Morphology. General

External Anatomy

The Head Region

The Trunk or Metastomial Region

Internal Anatomy

Excretory System

Pelagic Species

The Nervous System

Sense Organs

Circulatory System

Pelagic Species

Gas Exchange

Pelagic Species

Digestive System

Reproduction

Epitoky

Synchronicity

The Pelagic Species. Tomopteris

Alciopini and Lopadorrynchidae

Development

Locomotion

Foraging Strategies

The Hunters

Diets

The Suspension Feeders

Poeobiidae, Chaetopteridae

Distributions. Geographical

Vertical

Bioluminescence

References

7 The Crustacea. Introduction

Arthropod Classification. History

Subphylum Crustacea

Subphylum Hexapoda

Subphylum Myriapoda

Subphylum Chelicerata

Panarthropoda Phyla. Phylum Onychophora

Phylum Tardigrada

Synopsis of Universal Arthropod Characteristics

The Crustacea

Characteristics

Classification

Subphylum Crustacea. Class Remipedia

Class Cephalocarida

Class Branchiopoda

Class Copepoda

Class Thecostraca

Class Tantulocarida

Class Mystacocarida

Class Branchiura

Class Pentastomida

Class Ostracoda

Class Malacostraca

Subclass Phyllocarida

Subclass Hoplocarida

Subclass Eumalacostraca

Superorder Eucarida

Order Euphausiacea

Order Amphionidacea

Order Decapoda

Superorder Peracarida

Order Mysida

Order Lophogastrida

Order Isopoda

Order Amphipoda

Order Cumacea

Order Tanaidacea

Order Mictacea

Order Spelaeogriphacea

Order Stygiomysida

Order Thermosbaenacea

Superorder Syncarida

Crustacean Systems. Integument and Molting. Integument

Molting

Molt Cycle Stages

Hormonal Control

Joints and Appendages. Joints

Appendages

Excretory System

Extra‐renal Mechanisms

How the System Works

Nitrogen Excretion

The Nervous System and Sensory Mechanisms. The Central Nervous System

Sensory Modalities

Photoreception

Mechanoreception

Chemoreception

Circulatory and Respiratory Systems

Circulation and Oxygen Transport in the Blood: Hemocyanin

Digestive System

Basic Development

The Micronektonic Crustacea

The Pelagic Eucarida. Order Euphausiacea. Introduction

History

Classification

Euphausiid Anatomy

Nervous System

Sensory Mechanisms

Bioluminescence and Photophores

Circulatory, Respiratory, and Excretory Systems

Digestive System

Development

Gonad Development and Spawning

Food and Feeding

Geographical Distribution

Vertical Distribution and Vertical Migration

Ecological Factors

Acoustics, Species Composition, Light, and Layer Movement

Adaptive Significance: The Environment

Adaptive Significance of Vertical Migration

Order Decapoda. Introduction

History

Decapod Anatomy

Nervous System

Sensory Mechanisms

Bioluminescence and Photophores

Functions of Bioluminescence

Circulatory, Respiratory, and Excretory Systems

Digestive System

Development

Reproduction

Dendrobranchs

Carideans

Vertical Distribution

Food and Feeding

Geographic Distribution

Infraorder Anomura; Superfamily Galatheoidea; Family Munididae; Genera Pleuroncodes, Munida, and Cervimunida. The Pelagic Squat Lobsters

Order Amphionidacea

Introduction and History

Development and Natural History

Superorder Peracarida

Orders Lophogastrida and Mysida. Introduction

History

Anatomy

Nervous System

Sensory Mechanisms

Bioluminescence

Circulatory, Respiratory, and Excretory Systems

Digestive System

Reproduction and Development

Life History of a Lophogastrid

Food and Feeding

Locomotion

Vertical Distributions

Geographic Distributions

Order Amphipoda. Introduction

History

Classification

Anatomy

Nervous System

Sensory Mechanisms

Bioluminescence

Circulatory, Respiratory, and Excretory Systems

Digestive System

Reproduction and Development

Food and Feeding

Amphipods and Gelatinous Zooplankton

Geographic Distributions

Vertical Distributions

Cameo Players

References

8 The Mollusca. Introduction

Classification. History

The Pelagic Molluscs

Phylum Mollusca. Class Caudofoveata

Class Solenogastres

Class Monoplacophora

Class Polyplacophora

Class Scaphopoda

Class Bivalvia

Class Gastropoda

Class Cephalopoda

Body Organization

The Gastropoda. Classification

Class Gastropoda. Classification Below Subclass Given for Pelagic Species Only

Gastropod Systems and Structures

The Digestive Tract

Circulation

Respiration

Excretion

Shell Formation

The Nervous System

Sensory Mechanisms

The Pelagic Gastropods: Anatomy and Habits. The Janthinid Snails

Food and Feeding

Reproduction and Development

The Heteropods

Locomotion

Food and Feeding

Feeding Chronology

Vertical Distributions and Migrations

Abundance and Geographic Distribution

Reproduction and Development

The Pteropods

Order Pteropoda

Euthecosomata. Anatomy

Locomotion

Food and Feeding

Reproduction and Development

Pteropods, pH, and the Carbonate Cycle

Gymnosomata. Anatomy and Feeding

Locomotion

Reproduction

Superfamily Hydromyloidea

Pteropod Distributions

Vertical Migrations

The Nudibranchs

Family Glaucidae

Family Fionidae

Family Phylliroidae

The Cephalopoda

Classification

Basic Anatomy of the Major Cephalopod Groups

General

The Nautilida

The Sepiida and Spirulida

The Myopsida and Oegopsida

The Octopodiformes

Cephalopod Systems. Feeding and Digestion

The Digestive Tract

Movement of Food

Circulation

Gas Exchange

Excretion

Nervous System and Sensory Mechanisms

The Brain

The Giant Fiber System

Sensory Mechanisms

Touch, Taste, and Smell

Mechanoreception: Equilibrium and Acceleration

Hearing

Vision

Color

Bioluminescence

Locomotion and Buoyancy

Buoyancy

Life Histories

Nautiloids

Coleoids

Reproduction and Development

Vertical Distribution and Migration

Geographic Distribution

References

9 The Chordata. Introduction

Deuterostomes and the Phylogenetic Toolkit

Classification

Subphylum Tunicata. Class Ascidiacea (2935)

Class Appendicularia (68)

Class Thaliacea (78)

Basic Anatomy and Life History. The Ascidians

The Pyrosomes

The Salps

The Doliolids

The Appendicularia

The Appendicularian House

Tunicate Systems. Locomotion and Buoyancy

Pyrosomes

Salps

Doliolids

Appendicularia

Nervous Systems and Sensory Mechanisms

Pyrosomes

Doliolids

Salps

Appendicularia

Gas Exchange, Circulation, and Excretion

Trophic Role

Bioluminescence

Predators, Parasites and Other Interactions

Geographic and Vertical Distributions

References

10 The Fishes. Introduction

The Deep‐Sea Groups

A Brief History of Fishes. The Jawless Fishes

The Jawed Fishes. Teleosts

Elasmobranchs

Holocephali

The Classes of Living Fishes. Class Myxini

Class Petromyzonti

Class Elasmobranchii

Class Holocephali

Class Coelacanthi

Class Dipneusti

Class Cladistii

Class Actinopterygii

Subclass Chondrostei

Subclass Holostei

Subclass Teleostei

Cohort Elopomorpha

Cohort Otocephala

Cohort Euteleostei

Superorder Osmeromorpha

Superorder Ateleopodimorpha

Superorder Cyclosquamata

Superorder Scopelomorpha

Superorder Lamprimorpha

Superorder Paracanthopterygii

Superorder Acanthopterygii

Fish Systems. Basic Anatomy. External Features and Terms

Skull and Skeleton

Feeding and Digestion. Food Acquisition, the Three Dominant Modes: Ramming, Sucking, and Biting. Ram Feeding

Suction Feeding

Biting

Food Sorting: The “Pharyngeal Jaws”

Digestion: The Alimentary Canal

Circulation, Respiration, and Excretion. Circulation

Elasmobranch and Teleost Hearts

Arteries, Veins, and Continuity

Gas‐Exchange in the Teleosts and Elasmobranchs

Unidirectional Flow and Countercurrent Exchange: Maximizing the Concentration Gradient

Blood and Oxygen at the Respiratory Surface. Blood Basics

Red Cell Modifications

CO2 Transport–Far Different from O2

The Bohr and Haldane Effects

Secretion of Gases into the Swimbladder of Fishes

Nitrogen Excretion

Ammonia Excretion

Urea

Osmotic and Ionic Regulation. Introduction

Osmosis and Diffusion

The gills

Esophagus

Stomach

Intestine

Locomotion

Musculature

Red and White Muscle

Drag and Swimming Costs

Swimming Costs

Maximum Swim Speeds

Endothermy

Warm‐Brained Billfishes

Swimming in Mesopelagic Fishes

Buoyancy

The Nervous System. Anatomy and Basics

The Brain

Cranial Nerves

The Spinal Cord

Sensory Mechanisms. Sensory Modalities

Photoreception

The Choroid Body

Photoreceptors: Rods and Cones

Light and Vision

Visual and Morphological Adaptations to Deep‐Sea Life

Tubular Eyes and Retinal Adaptations

Mechanoreception

Free‐Standing Neuromasts and the Lateral Line

The Inner Ear and Sound Reception. Structure

Balance, Acceleration, and Angular Momentum

Hearing

Electroreception

Chemoreception: Olfaction and Gustation

Camouflage, Bioluminescence, Photophores. Camouflage

Bioluminescence and Photophores

References

11 Communities. Introduction

The Gulf of Mexico

The Northern California Current

The Antarctic

System Comparisons

The Decapods and Mysids

The Euphausiids

The Myctophids

Non‐myctophid Fishes

The Cephalopods

Gelatinous Zooplankton and Amphipods

Concluding Observations

Physical and Biological Factors that Change. Mean Annual Temperature

Seasonal Cycling

Annual Production

Current Patterns

References

12 Energetics. Introduction

A Model Energy Budget

Digestibility of Biomolecules

Energy Value of Biomolecules

Measuring Metabolic Rate

Oxygen Consumption Rate‐Modifying Factors

Activity

Experimental Protocol

Routine Metabolic Rate

Animal Size as a Modifier of Metabolism

Life History Strategies

Metabolism and Composition of Pelagic Species

Metabolism of Euphausiids, Decapods, Mysids, and Amphipods

Proximate Composition of Pelagic Decapods, Mysids, and Euphausiids

Terminology

Trends with Depth of Occurrence

Seasonal Changes

Trends Across Systems

Metabolism of Mesopelagic Fishes

Proximate Composition of Pelagic Fishes

Trends with Depth of Occurrence

Trends Across Systems

Energy and Life History in the Midwater Fauna

Midwater Fishes

The Cephalopods

Pteropods, Nemerteans, Annelids, Salps, and Pyrosomes

The Cnidaria and Ctenophora

Conclusions

References

Appendix A Classification of the Chordata

References

Glossary

Index

a

b

c

d

e

f

g

h

i

j

k

l

m

n

o

p

q

r

s

t

u

v

w

x

z

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Joseph J. Torres

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where I0 is the intensity of the incident light, I x is the intensity after traveling through distance x, and α λ is the attenuation coefficient, a function of wavelength.

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