Human Anatomy & Physiology

Laboratory Manual, Fetal Pig Version

13th Edition (eBook PDF)
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Your time is valuable. Make the most
of your time inside and outside the lab.

To help you manage your time inside and outside the A&P lab classroom, this
best-selling manual works hand-in-hand with Mastering A&P, the leading online
homework and learning program for A&P. This edition features dozens of new,
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expanded set of pre-lab videos, dissection videos, and more.
M09_MARI6358_13_SE_C09.indd Page 115 11/6/17 3:17 PM f-0035 /203/PH03335/9780134806358_MARIEB/MARIEB_HUMAN_ANATOMY_AND_PHYSIOLOGY_LAB_MANUAL1 ...

9
E X E R C I S E
The Axial Skeleton NEW! Mastering
A&P study tools
are highlighted on the
first page of each lab
Learning Outcomes Go to Mastering A&P™ > Study exercise, along with a
Area to improve your performance
▶▶ Name the three parts of the axial skeleton. in A&P Lab. photo preview of a
▶▶ Identify the bones of the axial skeleton, either by examining disarticulated
bones or by pointing them out on an articulated skeleton or skull, and
related pre-lab video,
name the important bone markings on each. image from Practice
▶▶ Name and describe the different types of vertebrae.

▶▶ Discuss the importance of intervertebral discs and spinal curvatures.

Anatomy Lab 3.1 (PAL),
▶▶ Identify three abnormal spinal curvatures. or animation.
▶▶ List the components of the thoracic cage.

▶▶ Identify the bones of the fetal skull by examining an articulated skull or

image. > Lab Tools > Bone & Dissection
▶▶ Define fontanelle, and discuss the function and fate of fontanelles. Videos
▶▶ Discuss important differences between the fetal and adult skulls. Instructors may assign new NEW! Mastering
Building Vocabulary coaching
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Instructors may assign these and other Pre-Lab
Pre-Lab Quiz Quiz questions using Mastering A&P™
Labeling activities, related bone videos
and coaching activities, Practice including NEW
1. The axial skeleton can be divided into the skull, the vertebral column,
Anatomy Lab Practical questions (PAL),
and more using the Mastering A&P™ Building Vocabulary
and the:
a. thoracic cage c. hip bones
Item Library.
Coaching Activities,
b. femur
2. Eight bones make up the
d. humerus
, which encloses and protects Materials are signaled at
the brain. ▶▶ Intact skull and Beauchene skull appropriate points
a. cranium b. face c. skull ▶▶ X-ray images of individuals with scoliosis,

3. The vertebrae articulate with the corresponding ribs. lordosis, and kyphosis (if available) throughout the manual
a. cervical c. spinal ▶▶ Articulated skeleton, articulated vertebral

b. lumbar d. thoracic column, removable intervertebral discs to help you connect the
4. The , commonly referred to as the breastbone, is a flat
bone formed by the fusion of three bones: the manubrium, the body,
▶▶ Isolated cervical, thoracic, and lumbar
vertebrae, sacrum, and coccyx exercises to relevant
and the xiphoid process. ▶▶ Isolated fetal skull
assignments that can
a. coccyx b. sacrum c. sternum
5. A fontanelle: be auto-graded in
a. is found only in the fetal skull
b. is a fibrous membrane Mastering A&P.
c. allows for compression of the skull during birth
d. all of the above

T he axial skeleton (the green portion of Figure 8.1 on p. 104) can be divided into
three parts: the skull, the vertebral column, and the thoracic cage. This division
of the skeleton forms the longitudinal axis of the body and protects the brain,
spinal cord, heart, and lungs.

115

See p. 115

i
Acknowledgments
Continued thanks to our colleagues and friends at Pearson uncommon grace and skill, including Janet Vail, production
who collaborated with us on this edition, especially Editor-in- coordinator; David Novak, art and photo coordinator; Kristin
Chief Serina Beauparlant, Acquisitions Editor Lauren Harp, Piljay, photo researcher; Gary Hespenheide, interior and cover
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Media Producers Kimberly Twardochleb and Lauren Chen. We Last but not least, we wish to extend our sincere thanks to
also thank the Pearson Sales and Marketing team for their the many A&P students who have circulated through our lab
work in supporting instructors and students, especially Senior classrooms and have used this lab manual over the years—you
A&P Specialist Derek Perrigo and Director of Product Mar- continue to inspire us every day! As always, we welcome your
keting Allison Rona. feedback and suggestions for future editions.
Special thanks go out to Amanda Kaufmann for her lead-
ership and expertise in producing the 18 pre-lab videos that
support this edition, and to Mike Mullins of BIOPAC®, who
helped us update the instructions for consistency with the
upgraded software.
We’re also grateful to Michele Mangelli and her superb
production team, who continue to cross every hurdle with Elaine N. Marieb & Lori A. Smith

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x
 A Pre-Lab video is available in
Contents Mastering A&P™ for selected activities.

THE HUMAN BODY: AN ORIENTATION

5 The Cell: Transport Mechanisms
1 The Language of Anatomy 1 and Cell Permeability 51
1 Locating Body Regions 3 1 Observing Diffusion of Dye Through
Agar Gel 53
2 Practicing Using Correct Anatomical
Terminology 4 2 Observing Diffusion of Dye Through Water 54
3 Observing Sectioned Specimens 6 3 Investigating Diffusion and Osmosis Through
Nonliving Membranes 54
4 Identifying Organs in the Abdominopelvic
Cavity 9 4 Observing Osmometer Results 56
5 Locating Abdominopelvic Surface Regions 10 5 Investigating Diffusion and Osmosis Through
Living Membranes 56
Review Sheet 11
6 Observing the Process of Filtration 59
2 Organ Systems Overview 15 7 Observing Phagocytosis 60
Review Sheet 61
1 Observing External Structures 17
2 Examining the Oral Cavity 17
3 Opening the Ventral Body Cavity 17 HISTOLOGY: BASIC TISSUES
4 Examining the Ventral Body Cavity 18 OF THE BODY
5 Examining the Human Torso Model 22
Review Sheet 23
6 Classification of Tissues 65
1 Examining Epithelial Tissue Under the
Microscope 67
THE MICROSCOPE AND ITS USES 2 Examining Connective Tissue Under the
Microscope 73
3 The Microscope 25 3 Examining Nervous Tissue Under the
Microscope 79
1 Identifying the Parts of a Microscope 26
2 Viewing Objects Through the Microscope 27 4 Examining Muscle Tissue Under the
Microscope 80
3 Estimating the Diameter of the Microscope
Review Sheet 83
Field 30
4 Perceiving Depth 31
5 Preparing and Observing a Wet Mount 31 THE INTEGUMENTARY SYSTEM
Review Sheet 33
7 The Integumentary System 89
THE CELL 1 Locating Structures on a Skin Model 90
2 Identifying Nail Structures 93
4 The Cell: Anatomy and Division 37 3 Comparing Hairy and Relatively Hair-Free
Skin Microscopically 94
1 Identifying Parts of a Cell 38
4 Differentiating Sebaceous and Sweat Glands
2 Identifying Components of a Plasma Microscopically 96
Membrane 39
5 Plotting the Distribution of Sweat Glands 96
3 Locating Organelles 40
6 Taking and Identifying Inked Fingerprints 97
4 Examining the Cell Model 40
Review Sheet 99
5 Observing Various Cell Structures 41
6 Identifying the Mitotic Stages 43
7 “Chenille Stick” Mitosis 43
Review Sheet 47

xi
xii Contents

THE SKELETAL SYSTEM THE MUSCULAR SYSTEM

8 Overview of the Skeleton: Classification
and Structure of Bones and
Cartilages 103
1 Examining a Long Bone 107
2 Examining the Effects of Heat and Hydrochloric
Acid on Bones 108
3 Examining the Microscopic Structure of Compact
Bone 109
4 Examining the Osteogenic Epiphyseal Plate 110
Review Sheet 111
9 The Axial Skeleton 115
1 Identifying the Bones of the Skull 116
Group Challenge Odd Bone Out 125
2 Palpating Skull Markings 126
3 Examining Spinal Curvatures 127
4 Examining Vertebral Structure 131
5 Examining the Relationship Between Ribs and
Vertebrae 133
6 Examining a Fetal Skull 134
Review Sheet 135
12 Microscopic Anatomy and Organization
of Skeletal Muscle 183
1 Examining Skeletal Muscle Cell Anatomy 186
2 Observing the Histological Structure of a
Skeletal Muscle 186
3 Studying the Structure of a Neuromuscular
Junction 188
Review Sheet 189
13 Gross Anatomy of the Muscular
System 193
1 Identifying Head and Neck Muscles 195
2 Identifying Muscles of the Trunk 195
3 Identifying Muscles of the Upper Limb 209
4 Identifying Muscles of the Lower Limb 214
5 Review of Human Musculature 218
6 Making a Muscle Painting 220
Review Sheet 223
14 Skeletal Muscle Physiology: Frogs
and Human Subjects 231

10 The Appendicular Skeleton 143 1 Observing Muscle Fiber Contraction 232

2 Inducing Contraction in the Frog Gastrocnemius
1 Examining and Identifying Bones of the Muscle 234
Appendicular Skeleton 143 3 Demonstrating Muscle Fatigue in Humans 239
2 Palpating the Surface Anatomy of the Pectoral BIOPAC 4 Electromyography in a Human Subject Using
®

Girdle and the Upper Limb 146 BIOPAC® 240

3 Observing Pelvic Articulations 149
4 Comparing Male and Female Pelves 151
5 Palpating the Surface Anatomy of the Pelvic
Girdle and Lower Limb 155
6 Constructing a Skeleton 156
Review Sheet 157
11 Articulations and Body
Movements 165
1 Identifying Fibrous Joints 166
2 Identifying Cartilaginous Joints 166
3 Examining Synovial Joint Structure 168
4 Demonstrating the Importance of Friction-
Reducing Structures 168
5 Demonstrating Movements of Synovial
Joints 170
6 Demonstrating Actions at the Hip Joint 173
7 Demonstrating Actions at the Knee Joint 173
8 Demonstrating Actions at the Shoulder
Joint 175
9 Examining the Action at the TMJ 176
Review Sheet 179
Review Sheet 247
THE NERVOUS SYSTEM
15 Histology of Nervous Tissue 251
1 Identifying Parts of a Neuron 254
2 Studying the Microscopic Structure of Selected
Neurons 256
3 Examining the Microscopic Structure of a
Nerve 258
Review Sheet 259
16 Neurophysiology of Nerve Impulses:
Frog Subjects 263
1 Stimulating the Nerve 266
Instructors may download two additional
lab activities: Inhibiting the Nerve and
Visualizing the Compound Action Potential
with an Oscilloscope. Instructors, please go to
Mastering A&P™ > Instructor Resources >
Additional Resources > Additional Exercises

Review Sheet 267

 Contents xiii

5 Demonstrating Adaptation of Temperature

17 Gross Anatomy of the Brain and Cranial Receptors 347
Nerves 269 6 Demonstrating the Phenomenon of Referred
Pain 348
1 Identifying External Brain Structures 271 Review Sheet 349
2 Identifying Internal Brain Structures 273
3 Identifying and Testing the Cranial Nerves 278 23 Special Senses: Anatomy of the Visual
Group Challenge Odd (Cranial) Nerve Out 286 System 351
Review Sheet 287
1 Identifying Accessory Eye Structures 353
18 Electroencephalography 293 2 Identifying Internal Structures of the Eye 353
3 Studying the Microscopic Anatomy of the
1 Observing Brain Wave Patterns Using an Retina 354
Oscilloscope or Physiograph 294 4 Predicting the Effects of Visual Pathway
BIOPAC 2 Electroencephalography Using BIOPAC® 295 Lesions 358
®

Review Sheet 299 Review Sheet 359

19 The Spinal Cord and Spinal Nerves
1 Identifying Structures of the Spinal Cord 302
2 Identifying Spinal Cord Tracts 305
3 Identifying the Major Nerve Plexuses and
Peripheral Nerves 312
Review Sheet 313
20 The Autonomic Nervous System 317
1 Locating the Sympathetic Trunk 318
2 Comparing Sympathetic and Parasympathetic
Effects 320
BIOPAC 3 Exploring the Galvanic Skin Response
®
301 24 Special Senses: Visual Tests and
Experiments 363
1 Demonstrating the Blind Spot 364
2 Determining Near Point of Vision 365
3 Testing Visual Acuity 366
4 Testing for Astigmatism 366
5 Testing for Color Blindness 367
6 Testing for Depth Perception 367
7 Demonstrating Reflex Activity of Intrinsic and
Extrinsic Eye Muscles 368
8 Conducting an Ophthalmoscopic
Examination 369

(Electrodermal Activity) Within a Polygraph Using Review Sheet 371

BIOPAC® 320
Review Sheet 327 25 Special Senses: Hearing and
Equilibrium 373
21 Human Reflex Physiology 329
1 Identifying Structures of the Ear 374
1 Initiating Stretch Reflexes 331 2 Examining the Ear with an Otoscope
2 Initiating the Crossed-Extensor Reflex 333 (Optional) 376
3 Initiating the Plantar Reflex 333 3 Examining the Microscopic Structure of the
Cochlea 377
4 Initiating the Corneal Reflex 334
4 Conducting Laboratory Tests of Hearing 378
5 Initiating the Gag Reflex 334
5 Audiometry Testing 380
6 Initiating Pupillary Reflexes 334
6 Examining the Microscopic Structure of the Crista
7 Initiating the Ciliospinal Reflex 335
Ampullaris 381
8 Initiating the Salivary Reflex 335
7 Conducting Laboratory Tests on Equilibrium 382
9 Testing Reaction Time for Intrinsic and Learned
Review Sheet 385
Reflexes 336
BIOPAC 10 Measuring Reaction Time Using BIOPAC 337
26
®
®

Special Senses: Olfaction and Taste 389

Review Sheet 339
1 Microscopic Examination of the Olfactory
22 General Sensation 343 Epithelium 391
2 Microscopic Examination of Taste Buds 392
1 Studying the Structure of Selected Sensory
3 Stimulating Taste Buds 392
Receptors 345
4 Examining the Combined Effects of Smell,
2 Determining the Two-Point Threshold 346
Texture, and Temperature on Taste 392
3 Testing Tactile Localization 347
5 Assessing the Importance of Taste and Olfaction
4 Demonstrating Adaptation of Touch in Odor Identification 394
Receptors 347
6 Demonstrating Olfactory Adaptation 394
Review Sheet 395
xiv Contents

THE ENDOCRINE SYSTEM

32 Anatomy of Blood Vessels 461
27 Functional Anatomy of the Endocrine 1 Examining the Microscopic Structure of Arteries
Glands 397 and Veins 463
2 Locating Arteries on an Anatomical Chart or
1 Identifying the Endocrine Organs 398
Model 470
2 Examining the Microscopic Structure of Endocrine
3 Identifying the Systemic Veins 474
Glands 402
4 Identifying Vessels of the Pulmonary
Group Challenge Odd Hormone Out 404
Circulation 475
Review Sheet 405
Group Challenge Fix the Blood Trace 476
5 Tracing the Pathway of Fetal Blood Flow 476
28 Endocrine Wet Labs and Human
6 Tracing the Hepatic Portal Circulation 478
Metabolism 409
Review Sheet 479
1 Determining the Effect of Pituitary Hormones on
the Ovary 410 33 Human Cardiovascular Physiology:
2 Observing the Effects of Hyperinsulinism 411 Blood Pressure and Pulse
Group Challenge Thyroid Hormone Case Determinations 485
Studies 412
Review Sheet 413 1 Auscultating Heart Sounds 488
2 Palpating Superficial Pulse Points 489
BIOPAC 3 Measuring Pulse Using BIOPAC® 490
®

THE CIRCULATORY SYSTEM 4 Taking an Apical Pulse 492

29 Blood 415 5 Using a Sphygmomanometer to Measure Arterial

1 Determining the Physical Characteristics
of Plasma 418
2 Examining the Formed Elements of Blood
Microscopically 418
3 Conducting a Differential WBC Count 421
4 Determining the Hematocrit 422
5 Determining Hemoglobin Concentration 424
6 Determining Coagulation Time 425
7 Typing for ABO and Rh Blood Groups 426
8 Observing Demonstration Slides 428
9 Measuring Plasma Cholesterol
Blood Pressure Indirectly 492
6 Estimating Venous Pressure 493
7 Observing the Effect of Various Factors on Blood
Pressure and Heart Rate 494
8 Examining the Effect of Local Chemical and
Physical Factors on Skin Color 496
Review Sheet 499
34 Frog Cardiovascular Physiology 505
1 Investigating the Automaticity and Rhythmicity
of Heart Muscle 506
BIOPAC 2 Recording Baseline Frog Heart Activity 508
®

Concentration 428 3 Investigating the Refractory Period of Cardiac

Review Sheet 429 Muscle Using the Physiograph 511
BIOPAC 4 Assessing Physical and Chemical Modifiers of
®

30 Anatomy of the Heart 435 Heart Rate 511

5 Investigating the Effect of Various Factors on the
1 Using the Heart Model to Study Heart Microcirculation 513
Anatomy 438
Review Sheet 515
2 Tracing the Path of Blood Through the
Heart 439
3 Using the Heart Model to Study Cardiac 35 The Lymphatic System and Immune
Circulation 440 Response 519
4 Examining Cardiac Muscle Tissue Anatomy 441 1 Identifying the Organs of the Lymphatic
Review Sheet 445 System 521
2 Studying the Microscopic Anatomy of a Lymph
31 Conduction System of the Heart and Node, the Spleen, and a Tonsil 522
Electrocardiography 449 Group Challenge Compare and Contrast Lymphoid
Organs and Tissues 524
1A Recording ECGs Using a Standard ECG 3 Using the Ouchterlony Technique to Identify
Apparatus 453 Antigens 525
BIOPAC 1B Electrocardiography Using BIOPAC® 454
®

Review Sheet 527

Review Sheet 459
 Contents xv

THE RESPIRATORY SYSTEM 4 Reporting Results and Conclusions 595

Group Challenge Odd Enzyme Out 595
36 Anatomy of the Respiratory 5 Observing Movements and Sounds of the
System 531 Digestive System 596
6 Viewing Segmental and Peristaltic
1 Identifying Respiratory System Organs 537 Movements 597
2 Demonstrating Lung Inflation in a Sheep Review Sheet 599
Pluck 537
3 Examining Prepared Slides of Trachea and Lung
Tissue 537 THE URINARY SYSTEM
Review Sheet 539
40 Anatomy of the Urinary System 603
37 Respiratory System Physiology 543 1 Identifying Urinary System Organs 605
1 Operating the Model Lung 544 2 Studying Nephron Structure 608
2 Auscultating Respiratory Sounds 546 3 Studying Bladder Structure 611
3 Measuring Respiratory Volumes Using Group Challenge Urinary System Sequencing 612
Spirometers 547 Review Sheet 613
4 Measuring the FVC and FEV1 553
BIOPAC 5 Measuring Respiratory Volumes Using
®

BIOPAC® 553
41 Urinalysis 617

6 Visualizing Respiratory Variations 557 1 Analyzing Urine Samples 619

7 Demonstrating the Reaction Between Carbon 2 Analyzing Urine Sediment Microscopically
Dioxide (in Exhaled Air) and Water 559 (Optional) 622
8 Observing the Operation of Standard Review Sheet 623
Buffers 560
9 Exploring the Operation of the Carbonic Acid–
Bicarbonate Buffer System 560 THE REPRODUCTIVE SYSTEM,
Review Sheet 561 DEVELOPMENT, AND HEREDITY
42 Anatomy of the Reproductive
THE DIGESTIVE SYSTEM System 625

38 Anatomy of the Digestive System 567 1 Identifying Male Reproductive Organs 626
2 Penis 629
1 Identifying Alimentary Canal Organs 569 3 Seminal Gland 630
2 Studying the Histologic Structure of the Stomach 4 Epididymis 630
and the Esophagus-Stomach Junction 573
5 Identifying Female Reproductive Organs 630
3 Observing the Histologic Structure of the Small
Intestine 576 6 Wall of the Uterus 633
4 Examining the Histologic Structure of the Large 7 Uterine Tube 633
Intestine 578 Review Sheet 635
5 Identifying Types of Teeth 579
6 Studying Microscopic Tooth Anatomy 580 43 Physiology of Reproduction:
7 Examining Salivary Gland Tissue 580 Gametogenesis and the Female
8 Examining the Histology of the Liver 581 Cycles 641
Review Sheet 583
1 Identifying Meiotic Phases and Structures 643
2
39 Digestive System Processes: Chemical
3
Examining Events of Spermatogenesis 644
Examining Meiotic Events Microscopically 645
and Physical 589
4 Examining Oogenesis in the Ovary 646
1 Assessing Starch Digestion by Salivary 5 Comparing and Contrasting Oogenesis and
Amylase 590 Spermatogenesis 646
2 Assessing Protein Digestion by Trypsin 593 6 Observing Histological Changes in the
3 Demonstrating the Emulsification Action of Bile Endometrium During the Menstrual Cycle 648
and Assessing Fat Digestion by Lipase 594 Review Sheet 651
xvi Contents

44 Survey of Embryonic Development 655 PIG DISSECTION EXERCISES

1 Microscopic Study of Sea Urchin
Development 656
1 Dissection and Identification of Fetal Pig
Muscles 695
2 Examining the Stages of Human
Development 656 2 Dissection of the Spinal Cord and Spinal
3 Identifying Fetal Structures 659 Nerves of the Fetal Pig 711
4 Studying Placental Structure 660
Review Sheet 661
3 Identification of Selected Endocrine Organs
of the Fetal Pig 715

45 Principles of Heredity 665 4 Dissection of Blood Vessels and Main

Lymphatic Ducts of the Fetal Pig 719
1 Working Out Crosses Involving Dominant and
Recessive Genes 667 5 Dissection of the Respiratory System of
2 Working Out Crosses Involving Incomplete the Fetal Pig 729
Dominance 667
3 Working Out Crosses Involving Sex-Linked 6 Dissection of the Digestive System of
Inheritance 668 the Fetal Pig 733
4 Exploring Probability 669
5 Using Blood Type to Explore Phenotypes and
7 Dissection of the Urinary System of
the Fetal Pig 739
Genotypes 670
6 Using Agarose Gel Electrophoresis to Identify 8 Dissection of the Reproductive System of
Normal Hemoglobin, Sickle Cell Anemia, and the Fetal Pig 743
Sickle Cell Trait 670
Review Sheet 673
PHYSIOEX TM 9.1 COMPUTER SIMULATIONS

SURFACE ANATOMY
1 Cell Transport Mechanisms and
Permeability PEx-3

46 Surface Anatomy Roundup 677

2 Skeletal Muscle Physiology PEx-17
1 Palpating Landmarks of the Head 678
2 Palpating Landmarks of the Neck 680 3 Neurophysiology of Nerve Impulses PEx-35
3 Palpating Landmarks of the Trunk 682
4 Palpating Landmarks of the Abdomen 684 4 Endocrine System Physiology PEx-59
5
6
Palpating Landmarks of the Upper Limb 686
Palpating Landmarks of the Lower Limb 689
5 Cardiovascular Dynamics PEx-75

Review Sheet 693

6 Cardiovascular Physiology PEx-93

7 Respiratory System Mechanics PEx-105

8 Chemical and Physical Processes of

Digestion PEx-119

9 Renal System Physiology PEx-131

10 Acid-Base Balance PEx-149

11 Blood Analysis PEx-161

12 Serological Testing PEx-177

Credits C-1
Index I-1
 Anatomy and Physiology
Laboratory Safety Guidelines*
1. Upon entering the laboratory, locate exits, fire extinguisher, fire blanket, chemical shower,
eyewash station, first aid kit, containers for broken glass, and materials for cleaning up spills.
2. Do not eat, drink, smoke, handle contact lenses, store food, or apply cosmetics or lip balm in
the laboratory. Restrain long hair, loose clothing, and dangling jewelry.
3. Students who are pregnant, are taking immunosuppressive drugs, or have any other medical
conditions (e.g., diabetes, immunological defect) that might necessitate special precautions in
the laboratory must inform the instructor immediately.
4. Wearing contact lenses in the laboratory is inadvisable because they do not provide eye
protection and may trap material on the surface of the eye. Soft contact lenses may absorb
volatile chemicals. If possible, wear regular eyeglasses instead.
5. Use safety glasses in all experiments involving liquids, aerosols, vapors, and gases.
6. Decontaminate work surfaces at the beginning and end of every lab period, using a
commercially prepared disinfectant or 10% bleach solution. After labs involving dissection of
preserved material, use hot soapy water or disinfectant.
7. Keep all liquids away from the edge of the lab bench to avoid spills. Clean up spills of viable
materials using disinfectant or 10% bleach solution.
8. Properly label glassware and slides.
9. Use mechanical pipetting devices; mouth pipetting is prohibited.
10. Wear disposable gloves when handling blood and other body fluids, mucous membranes, and
nonintact skin, and when touching items or surfaces soiled with blood or other body fluids.
Change gloves between procedures. Wash hands immediately after removing gloves. (Note: Cover
open cuts or scrapes with a sterile bandage before donning gloves.)
11. Place glassware and plasticware contaminated by blood and other body fluids in a disposable
autoclave bag for decontamination by autoclaving, or place them directly into a 10% bleach
solution before reuse or disposal. Place disposable materials such as gloves, mouthpieces,
swabs, and toothpicks that have come into contact with body fluids into a disposable autoclave
bag, and decontaminate before disposal.
12. To help prevent contamination by needlestick injuries, use only disposable needles and lancets. Do
not bend the needles and lancets. Needles and lancets should be placed promptly in a labeled,
puncture-resistant, leakproof container and decontaminated, preferably by autoclaving.
13. Do not leave heat sources unattended.
14. Report all spills or accidents, no matter how minor, to the instructor.
15. Never work alone in the laboratory.
16. Remove protective clothing before leaving the laboratory.

*Adapted from:
Biosafety in Microbiological and Biomedical Laboratories (BMBL), Fifth Edition. 2007. U.S. Government Printing Office. Washington,
D.C. www.cdc.gov/od/OHS/biosfty/bmbl5/bmbl5toc.htm
Centers for Disease Control. 1996. “Universal Precautions for Prevention of Transmission of HIV and Other Bloodborne Infec-
tions.” Washington, D.C. www.cdc.gov/ncidod/dhqp/bp_universal_precautions.html
Johnson, Ted, and Christine Case. 2010. Laboratory Experiments in Microbiology, Ninth Edition. San Francisco: Pearson Benja-
min Cummings.
School Chemistry Laboratory Safety Guide. 2006. U.S. Consumer Product Safety Commission. Bethesda, MD. www.cpsc.gov/
CPSCPUB/PUBS/NIOSH2007107.pdf
The Metric System
Metric to English English to metric
Measurement Unit and abbreviation Metric equivalent conversion factor conversion factor

Length 1 kilometer (km) 5 1000 (103) meters 1 km 5 0.62 mile 1 mile 5 1.61 km
1 meter (m) 5 100 (102) centimeters 1 m 5 1.09 yards 1 yard 5 0.914 m
5 1000 millimeters 1 m 5 3.28 feet 1 foot 5 0.305 m
1 m 5 39.37 inches 1 foot 5 30.5 cm
1 centimeter (cm) 5 0.01 (1022) meter 1 cm 5 0.394 inch 1 inch 5 2.54 cm
1 mm 5 0.039 inch
1 millimeter (mm) 5 0.001 (1023) meter
1 micrometer (μm) 5 0.000001 (1026) meter
[formerly micron (μ)]
1 nanometer (nm) 5 0.000000001 (1029)
[formerly millimicron meter
(mμ)]
1 angstrom (Å) 5 0.0000000001
(10210) meter

Area 1 square meter (m2) 5 10,000 square 1 m2 5 1.1960 square 1 square yard 5
centimeters yards 0.8361 m2
1 m2 5 10.764 square 1 square foot 5
feet 0.0929 m2
1 square centimeter 5 100 square 1 cm2 5 0.155 square 1 square inch 5
(cm2) millimeters inch 6.4516 cm2

Mass 1 metric ton (t) 5 1000 kilograms 1 t 5 1.103 ton 1 ton 5 0.907 t
1 kilogram (kg) 5 1000 grams 1 kg 5 2.205 pounds 1 pound 5 0.4536 kg
1 gram (g) 5 1000 milligrams 1 g 5 0.0353 ounce 1 ounce 5 28.35 g
1 g 5 15.432 grains
1 milligram (mg) 5 0.001 gram 1 mg 5 approx. 0.015
grain
1 microgram (μg) 5 0.000001 gram
Volume 1 cubic meter (m ) 3
5 1,000,000 cubic 1 m3 5 1.3080 cubic 1 cubic yard 5
(solids) centimeters yards 0.7646 m3
1 m3 5 35.315 cubic 1 cubic foot 5
feet 0.0283 m3

1 cubic centimeter 5 0.000001 cubic meter 1 cm3 5 0.0610 cubic 1 cubic inch 5
(cm3 or cc) 5 1 milliliter inch 16.387 cm3
1 cubic millimeter (mm3) 5 0.000000001 cubic meter
Volume 1 kiloliter (kl or kL) 5 1000 liters 1 kL 5 264.17 gallons 1 gallon 5 3.785 L
(liquids 1 liter (l or L) 5 1000 milliliters 1 L 5 0.264 gallons 1 quart 5 0.946 L
and gases) 1 L 5 1.057 quarts
1 milliliter (ml or mL) 5 0.001 liter 1 ml 5 0.034 fluid 1 quart 5 946 ml
5 1 cubic centimeter ounce 1 pint 5 473 ml
1 ml 5 approx. 14 1 fluid ounce 5
teaspoon 29.57 ml
1 ml 5 approx. 15–16 1 teaspoon 5 approx.
drops (gtt.) 5 ml
1 microliter (μl or μL) 5 0.000001 liter
Time 1 second (s or sec) 5 601 minute
1 millisecond (ms or msec) 5 0.001 second
Temperature Degrees Celsius (°C) °F 5 95 (°C) 1 32 °C 5 59 (°F 2 32)
 1
E X E R C I S E
The Language
of Anatomy

Learning Outcomes Go to Mastering A&P™ > Study

Area to improve your performance
▶▶ Describe the anatomical position, and explain its importance. in A&P Lab.
▶▶ Use proper anatomical terminology to describe body regions, orientation
and direction, and body planes.
▶▶ Name the body cavities, and indicate the important organs in each.

▶▶ Name and describe the serous membranes of the ventral body cavities.

▶▶ Identify the abdominopelvic quadrants and regions on a torso model or

image.

Instructors may assign these and other Pre-Lab

Pre-Lab Quiz Quiz questions using Mastering A&P™
Instructors may assign new
Building Vocabulary coaching
1. Circle True or False. In anatomical position, the body is lying down.
activities, Pre-Lab Quiz questions, Art
2. Circle the correct underlined term. With regard to surface anatomy, Labeling activities, and more using the
abdominal / axial refers to the structures along the center line of the Mastering A&P™ Item Library.
body.
3. The term superficial refers to a structure that is:
a. attached near the trunk of the body Materials
b. toward or at the body surface ▶▶ Human torso model (dissectible)
c. toward the head ▶▶ Human skeleton
d. toward the midline ▶▶ Demonstration: sectioned and labeled
4. The ________ plane runs longitudinally and divides the body into kidneys (three separate kidneys uncut
right and left sides. or cut so that [a] entire, [b] transverse
a. frontal c. transverse sectional, and [c] longitudinal sectional
b. sagittal d. ventral views are visible)
5. Circle the correct underlined terms. The dorsal body cavity can be ▶▶ Gelatin-spaghetti molds

divided into the cranial / thoracic cavity, which contains the brain, and ▶▶ Scalpel

the sural / vertebral cavity, which contains the spinal cord.

A
student new to any science is often overwhelmed at first by the terminol-
ogy used in that subject. The study of anatomy is no exception. But with-
out specialized terminology, confusion is inevitable. For example, what
do over, on top of, above, and behind mean in reference to the human body?
Anatomists have an accepted set of reference terms that are universally under-
stood. These allow body structures to be located and identified precisely with a
minimum of words.
This exercise presents some of the most important anatomical terminology used
to describe the body and introduces you to basic concepts of gross anatomy, the
study of body structures visible to the naked eye.

1
 2 Exercise 1

Anatomical Position
When anatomists or doctors refer to specific areas of the human important to remember that “left” and “right” refer to the sides
body, the picture they keep in mind is a universally accepted of the individual, not the observer.
standard position called the anatomical position. In the ana-
• Assume the anatomical position. The hands are held unnatu-
tomical position, the human body is erect, with the feet only
slightly apart, head and toes pointed forward, and arms hanging rally forward rather than hanging with palms toward the thighs.
at the sides with palms facing forward (Figure 1.1a). It is also Check the box when you have completed this task.
1
Cephalic (head) Cephalic
Frontal Otic
Orbital Occipital
Nasal
Buccal
Upper limb
Oral Cervical (neck)
Acromial
Mental
Brachial
Cervical Antecubital
Thoracic Olecranal Back (dorsal)
Sternal Antebrachial Scapular
Axillary Carpal
Mammary Vertebral
Abdominal
Umbilical Lumbar

Pelvic Manus (hand) Sacral

Inguinal Palmar
Pollex Gluteal
Digital
Perineal
(between anus
Lower limb and external
Coxal genitalia)
Pubic
(genital) Femoral
Patellar
Popliteal
Crural
Sural
Fibular or peroneal

Thorax
Pedal (foot)
Abdomen Tarsal
Back (Dorsum) Calcaneal
Digital
Plantar
Hallux

(a) Anterior/Ventral (b) Posterior/Dorsal

Figure 1.1 Anatomical position and regional terms. Heels are raised to illustrate
Instructors may assign this figure as an Art
the plantar surface of the foot, which is actually on the inferior surface of the body.
Labeling Activity using Mastering A&P™

Regional Anatomy
The body is divided into two main regions, the axial and
appendicular regions. The axial region includes the head,
neck, and trunk; it runs along the vertical axis of the body.
The appendicular region includes the limbs, which are also
called the ­appendages or extremities. The body is also divided
up into smaller regions within those two main divisions.
Table 1.1 summarizes the body regions that are illustrated
in Figure 1.1.
 The Language of Anatomy 3

Table 1.1 Regions of the Human Body (Figure 1.1)

Region Description Region Description

Abdominal Located below the ribs and Nasal Nose
above the hips
Acromial Point of the shoulder Occipital Back of the head
Antebrachial Forearm Olecranal Back of the elbow
Antecubital Anterior surface of the elbow Oral Mouth
Axillary Armpit Orbital Bony eye socket 1
Brachial Arm (upper portion of the Otic Ear
upper limb)
Buccal Cheek Palmar Palm of the hand
Calcaneal Heel of the foot Patellar Kneecap
Carpal Wrist Pedal Foot
Cephalic Head Pelvic Pelvis
Cervical Neck Perineal Between the anus and the
external genitalia
Coxal Hip Plantar Sole of the foot
Crural Leg Pollex Thumb
Digital Fingers or toes Popliteal Back of the knee
Femoral Thigh Pubic Genital
Fibular Side of the leg Sacral Posterior region between the
(peroneal) hip bones
Frontal Forehead Scapular Shoulder blade
Gluteal Buttocks Sternal Breastbone
Hallux Great toe Sural Calf
Inguinal Groin Tarsal Ankle
Lumbar Lower back Thoracic Chest
Mammary Breast Umbilical Naval
Manus Hand Vertebral Spine
Mental Chin

Activity 1
Locating Body Regions
Locate the anterior and posterior body regions on yourself,
your lab partner, and a human torso model.

Directional Terms
Study the terms below, referring to Figure 1.2 for a visual aid.
Notice that certain terms have different meanings, depending
on whether they refer to a four-legged animal (quadruped) or
to a human (biped).
Superior/inferior (above/below): These terms refer to place-
ment of a structure along the long axis of the body. The nose,
for example, is superior to the mouth, and the abdomen is
inferior to the chest.
Anterior/posterior (front/back): In humans, the most anterior
structures are those that are most forward—the face, chest, and
abdomen. Posterior structures are those toward the backside of
the body. For instance, the spine is posterior to the heart.
Medial/lateral (toward the midline/away from the midline or
median plane): The sternum (breastbone) is medial to the ribs;
the ear is lateral to the nose.
The terms of position just described assume the person is
in the anatomical position. The next four term pairs are more
absolute. They apply in any body position, and they consistently
have the same meaning in all vertebrate animals.
 4 Exercise 1

Cephalad (cranial)/caudal (toward the head/toward the tail): In
humans, these terms are used interchangeably with superior
and inferior, but in four-legged animals they are synonymous
with anterior and posterior, respectively.
Ventral/dorsal (belly side/backside): These terms are used
chiefly in discussing the comparative anatomy of animals,
assuming the animal is standing. In humans, the terms ventral
and dorsal are used interchangeably with the terms anterior
and posterior, but in four-legged animals, ventral and dorsal are
1 synonymous with inferior and superior, respectively.
Proximal/distal (nearer the trunk or attached end/farther from
the trunk or point of attachment): These terms are used primar-
ily to locate various areas of the body limbs. For example, the
fingers are distal to the elbow; the knee is proximal to the toes.
However, these terms may also be used to indicate regions
(closer to or farther from the head) of internal tubular organs.
Superficial (external)/deep (internal) (toward or at the body
surface/away from the body surface): For example, the skin is
superficial to the skeletal muscles, and the lungs are deep to
the rib cage.

Superior (cephalad)

Posterior Anterior
(dorsal) (ventral)

Proximal Superior (dorsal)

Posterior Anterior
(caudal) (cephalad)

Distal

(a) Inferior (caudal) (b) Inferior (ventral)

Figure 1.2 Directional terms. (a) With reference to a human. (b) With reference
Instructors may assign this figure as an Art
to a four-legged animal.
Labeling Activity using Mastering A&P™

Activity 2
Practicing Using Correct Anatomical Terminology
Use a human torso model, a human skeleton, or your own body 3. The femoral region is ________________ to the tarsal
to practice using the regional and directional terminology. region. (proximal or distal)

1. The popliteal region is ________________. (anterior or 4. The bones are ________________ to the skin. (superficial or
­posterior) deep)

2. The acromial region is ________________ to the otic region.

(medial or lateral)
 The Language of Anatomy 5

Body Planes and Sections

The body is three-dimensional, and in order to observe its or line called a plane. A section is named for the plane
internal structures, it is often necessary to make a section, along which it is cut. Anatomists commonly refer to three
or cut. When the section is made through the body wall or planes (Figure 1.3), or sections, that lie at right angles to one
through an organ, it is made along an imaginary surface another.

(a) Median (midsagittal) plane (b) Frontal (coronal) plane (c) Transverse plane

Vertebral Right Left

column lung Heart lung Liver Aorta Pancreas Spleen

Rectum Intestines Liver Stomach Spleen Subcutaneous Spinal

fat layer cord

Figure 1.3 Planes of the body with corresponding magnetic resonance

Instructors may assign this figure as an Art
imaging (MRI) scans. Note the transverse section is an inferior view. Labeling Activity using Mastering A&P™
 6 Exercise 1

Sagittal plane: A sagittal plane runs longitudinally and divides

the body into right and left parts. If it divides the body into
equal parts, right down the midline of the body, it is called a
median, or midsagittal, plane.
Frontal plane: Sometimes called a coronal plane, the frontal
plane is a longitudinal plane that divides the body (or an organ)
into anterior and posterior parts.
Transverse plane: A transverse plane runs horizontally, divid-
ing the body into superior and inferior parts. When organs are
1 ­sectioned along the transverse plane, the sections are com-
monly called cross sections. (a) Transverse (cross)
section
On microscope slides, the abbreviation for a longitudinal
section (sagittal or frontal) is l.s. Cross sections are abbreviated
x.s. or c.s.
A median or frontal plane section of any nonspherical
object, be it a banana or a body organ, provides quite a different
view from a cross section (Figure 1.4).

Activity 3
Observing Sectioned Specimens
1. Go to the demonstration area and observe the transversely
and longitudinally cut organ specimens (kidneys).
2. After completing instruction 1, obtain a gelatin-spaghetti (b) Median
mold and a scalpel, and take them to your laboratory bench. section
(Essentially, this is just cooked spaghetti added to warm gelatin,
which is then allowed to gel.)
3. Cut through the gelatin-spaghetti mold along any plane,
and examine the cut surfaces. You should see spaghetti strands (c) Frontal sections
that have been cut transversely (x.s.) and some cut longitudi-
nally (a median section).
Figure 1.4 Objects can look odd when viewed in section.
4. Draw the appearance of each of these spaghetti sections This banana has been sectioned in three different planes (a–c),
below, and verify the accuracy of your section identifications and only in one of these planes (b) is it easily recognized as
with your instructor. a banana. If one cannot recognize a sectioned organ, it is
possible to reconstruct its shape from a series of successive
cuts, as from the three serial sections in (c).

Transverse section Median section

 The Language of Anatomy 7

Body Cavities
The axial region of the body has two large cavities that pro-
vide different degrees of protection to the organs within them
(Figure 1.5).
Dorsal Body Cavity
The dorsal body cavity can be subdivided into the cranial ­cavity,
which lies within the rigid skull and encases the brain, and the
vertebral (or spinal) cavity, which runs through the bony verte-
bral column to enclose the delicate spinal cord.
Ventral Body Cavity
Like the dorsal cavity, the ventral body cavity is subdivided.
The superior thoracic cavity is separated from the rest of the
ventral cavity by the dome-shaped diaphragm. The heart and
lungs, located in the thoracic cavity, are protected by the bony
rib cage. The cavity inferior to the diaphragm is referred to
as the abdominopelvic cavity. Although there is no further­
physical separation of the ventral cavity, some describe the
abdominopelvic cavity as two areas: a superior abdominal
­cavity, the area that houses the stomach, intestines, liver, and
other organs, and an inferior pelvic cavity, the region that is
partially enclosed by the bony pelvis and contains the repro-
ductive organs, bladder, and rectum.
1
Serous Membranes of the Ventral Body Cavity
The walls of the ventral body cavity and the outer surfaces of
the organs it contains are covered with a very thin, double-
layered membrane called the serosa, or serous membrane.
The part of the membrane lining the cavity walls is referred
to as the parietal serosa, and it is continuous with a similar
membrane, the visceral serosa, covering the external surface
of the organs within the cavity. These membranes produce a
thin lubricating fluid that allows the visceral organs to slide
over one another or to rub against the body wall with minimal

Cranial
Cranial cavity cavity
(contains brain)

Vertebral
cavity

Pleural
Dorsal Thoracic cavity
body cavity
cavity Mediastinum
(contains
heart and
lungs) Pericardial
cavity

Vertebral cavity
(contains spinal Diaphragm Ventral body
cord) cavity
(thoracic and
Abdominal cavity abdominopelvic
(contains digestive Abdomino- cavities)
organs) pelvic
cavity

Pelvic cavity
(contains urinary
Dorsal body cavity bladder, reproductive
Ventral body cavity organs, and rectum)

(a) Lateral view (b) Anterior view

Figure 1.5 Dorsal and ventral body cavities and their subdivisions.
Instructors may assign this figure as an Art Labeling
Activity using Mastering A&P™
 8 Exercise 1

Lung Ribs Heart

Parietal pleura
Parietal
pericardium
Pleural cavity Pericardial cavity
with serous fluid with serous fluid

Visceral pleura Visceral

1
pericardium
Diaphragm

(a) Serosae associated with the lungs: pleura (b) Serosae associated with the heart: pericardium

Anterior Visceral
peritoneum

Liver Peritoneal Outer balloon wall

cavity (with (comparable to parietal serosa)
serous fluid)
Air (comparable to serous cavity)
Stomach
Parietal Inner balloon wall
peritoneum Kidney (comparable to visceral serosa)
(retroperitoneal)

Wall of
Posterior
body trunk
(c) Serosae associated with the abdominal viscera: peritoneum (d) Model of the serous membranes and serous cavity

Figure 1.6 Serous membranes of the ventral body cavities.

friction. Serous membranes also compartmentalize the vari-
ous organs to prevent infection in one organ from spreading
to others.
The specific names of the serous membranes depend on
the structures they surround. The serosa lining the abdominal
c­avity and covering its organs is the peritoneum, the serosa
enclosing the lungs is the pleura, and the serosa around the
heart is the pericardium (Figure 1.6). A fist pushed into a limp
balloon demonstrates the relationship between the visceral and
parietal serosae (Figure 1.6d).
 The Language of Anatomy 9

Activity 4
Identifying Organs in the
Abdominopelvic Cavity
Examine the human torso model to respond to the following
questions.

Name two organs found in the left upper quadrant.

1
Right upper Left upper ________________ and ________________
quadrant quadrant
(RUQ) (LUQ) Name two organs found in the right lower quadrant.

________________ and ________________

Right lower Left lower
quadrant quadrant What organ (Figure 1.7) is divided into identical halves by
(RLQ) (LLQ)
the median plane? ________________

Figure 1.7 Abdominopelvic quadrants. Superficial organs

are shown in each quadrant.

Abdominopelvic Quadrants and Regions
Because the abdominopelvic cavity is quite large and contains
many organs, it is helpful to divide it up into smaller areas for
discussion or study.
Most physicians and nurses use a scheme that divides the
abdominal surface and the abdominopelvic cavity into four
approximately equal regions called quadrants. These quadrants
are named according to their relative position—that is, right
upper quadrant, right lower quadrant, left upper quadrant, and
left lower quadrant (Figure 1.7). Note that the terms left and
right refer to the left and right side of the body in the figure, not
the left and right side of the art on the page.
A different scheme commonly used by anatomists divides
the abdominal surface and abdominopelvic cavity into nine
separate regions by four planes (Figure 1.8). As you read

Liver Diaphragm
Right Left
Epigastric Spleen
hypochondriac hypochondriac
region Gallbladder Stomach
region region

Ascending colon of Transverse colon

Right Left of large intestine
Umbilical large intestine
lateral (lumbar) lateral (lumbar)
region region region Small intestine Descending colon
of large intestine
Cecum Initial part of
Right inguinal Pubic Left inguinal
(iliac) (hypogastric) (iliac) sigmoid colon
Appendix
region region region
Urinary bladder

(a) (b)

Figure 1.8 Abdominopelvic regions. Nine regions delineated by four planes. (a) The
Instructors may assign this figure
superior horizontal plane is just inferior to the ribs; the inferior horizontal plane is at the
as an Art Labeling Activity using
superior aspect of the hip bones. The vertical planes are just medial to the nipples.
Mastering A&P™
(b) Superficial organs are shown in each region.
Another random document with
no related content on Scribd:
which the spermatozoa are packed; they are often very large and assume
characteristic shapes, especially in the Decapoda.
The spermatozoa show a great variety of structure, but they conform to
two chief types—the filiform, which are provided with a long whip-like
flagellum; and the amoeboid, which are furnished with radiating
pseudopodia, and are much slower in their movements. The amoeboid
spermatozoa of some of the Decapoda contain in the cell-body a peculiar
chitinous capsule, and Koltzoff[12] has observed that when the
spermatozoon has settled upon the surface of the egg the chitinous
capsule becomes suddenly exceedingly hygroscopic, swells up, and
explodes, driving the head of the spermatozoon into the egg. We cannot
enter here into a description of the embryological changes by which the
egg is converted into the adult form. Crustacean eggs as a whole contain a
large quantity of yolk, but in some forms total segmentation occurs in the
early stages, which is converted later into the pyramidal type, i.e. the
blastomeres are arranged round the edge, and the yolk in the centre is
only partly segmented to correspond with them. The eggs during the early
stages of development are in almost all cases (except Branchiura, p. 77,
and Anaspides, p. 116) carried about by the female either in a brood-
pouch (Branchiopoda, Ostracoda, Cirripedia, Phyllocarida, Peracarida),
or agglutinated to the hind legs or some other part of the body (Copepoda,
Eucarida), or in a chamber formed from the maxillipedes (Stomatopoda).
Development may be direct, without a complicated metamorphosis, or
indirect, the larva hatching out in a form totally different to the adult
state, and attaining the latter by a series of transformations and moults.
The various larval forms will be described under the headings of the
several orders.
The respiratory organs are typically branchiae, i.e. branched
filamentous or foliaceous processes of the body-surface through which
the blood circulates, and is brought into close relation with the oxygen
dissolved in the water. In most of the smaller Entomostraca no special
branchiae are present, the interchange of gases taking place over the
whole body-surface; but in the Malacostraca the gills may reach a high
degree of specialisation. They are usually attached to the bases of the
thoracic limbs (“podobranchiae”), to the body-wall at the bases of these
limbs, often in two series (“arthrobranchiae”), and to the body-wall some
way above the limb-articulations (“pleurobranchiae”). In an ideal scheme
each thoracic appendage beginning with the first maxillipede would
possess a podobranch, two arthrobranchs, and a pleurobranch, but the
full complement of gills is never present, various members of the series
being suppressed in the various orders, and thus giving rise to “branchial
formulae” typical of the different groups.
After this brief survey of Crustacean organisation we may be able to
form an opinion upon the position of the Crustacea relative to other
Arthropoda, and upon the question debated some time ago in the pages of
Natural Science[13] whether the Arthropoda constitute a natural group.
The Crustacea plainly agree with all the other Arthropoda in the
possession of a rigid exoskeleton segmented into a number of somites, in
the possession of jointed appendages metamerically repeated, some of
which are modified to act as jaws; they further agree in the general
correspondence of the number of segments of which the body is
primitively composed; the condition of the body-cavity or haemocoel is
also similar in the adult state. An apparently fundamental difference is
found in the entire absence during development of a segmented coelom,
but since this organ breaks down and is much reduced in all adult
Arthropods, it is not difficult to believe that its actual formation in the
embryo as a distinct structure might have been secondarily suppressed in
Crustacea.
The method of breathing by gills is paralleled by the respiratory
structures found in Limulus and Scorpions; the transition, if it occurred,
from branchiae to tracheae cannot, it is true, be traced, but the separation
of Arthropods into phyletically distinct groups of Tracheata and
Branchiata on this single characteristic is inadmissible. On the whole the
Crustacea may be considered as Arthropods whose progenitors are to be
sought for among the Trilobita, from whose near relations also probably
sprang Limulus and the Arachnids.
 CHAPTER II
CRUSTACEA (CONTINUED): ENTOMOSTRACA
—BRANCHIOPODA—PHYLLOPODA—
CLADOCERA—WATER-FLEAS

SUB-CLASS I.—ENTOMOSTRACA.

The Entomostraca are mostly small Crustacea in which the

segmentation of the body behind the head is very variable, both in regard
to the number of segments and the kind of differentiation exhibited by
those segments and their appendages. An unpaired simple eye, known as
the Nauplius eye from its universal presence in that larval form, often
persists in the adult, and though lateral compound eyes may be present
they are rarely borne on movable stalks. In the adult the excretory gland
(“shell-gland”) opens on the second maxillary segment, but in the larval
state or early stages of development a second antennary gland may also
be present, which disappears in the adult. The liver usually points
forwards, and is simple and saccular in structure, and the stomach is not
complicated by the formation of a gastric mill. With the exception of most
Cladocera and Ostracoda the young hatch out in the Nauplius state.

Order I. Branchiopoda.[14]

The Branchiopods are of small or moderate size, with flattened and

lobate post-cephalic limbs, and with functional gnathobases. Median and
lateral eyes are nearly always present. The labrum is large, and the second
maxillae are small or absent in the adult.
Branchiopods are found in every part of the world; a few are marine,
but the great majority are confined to inland lakes and ponds, or to
slowly-moving streams. The fresh waters, from the smallest pools to the
largest lakes, often swarm with them, as do those streams which flow so
slowly that the creatures can obtain occasional shelter among vegetation
along the sides and bottom without being swept away, while even rivers of
considerable swiftness contain some Cladocera. Several Branchiopods are
found in the brackish waters of estuaries, and some occur in lakes and
pools so salt that no other Crustacea, and few other animals of any kind,
can live in them. The great majority swim about with the back
downwards, collecting food in the ventral groove between their post-oral
limbs, and driving it forwards, towards the mouth, by movements of the
gnathobases (p. 10). The food collected in this way consists largely of
suspended organic mud, together with Diatoms and other Algae, and
Infusoria; the larger kinds, however, are capable of gnawing objects of
considerable size, Apus being said to nibble the softer insect larvae, and
even tadpoles. Many Cladocera (e.g. Daphnia, Simocephalus) may be
seen to sink to the bottom of an aquarium, with the ventral surface
downwards, and to collect mud, or even to devour the dead bodies of their
fellows, while Leptodora is said to feed upon living Copepods, which it
catches by means of its antennae.
The Branchiopoda fall naturally into two Sub-orders, the Phyllopoda
including a series of long-bodied forms, with at least ten pairs of post-
cephalic limbs, and the Cladocera with shorter bodies and not more
than six pairs of post-cephalic limbs.

Sub-Order 1. Phyllopoda.

The Phyllopoda include a series of genera which differ greatly in

appearance, owing to differences in the development of the carapace,
which are curiously correlated with differences in the position of the eyes.
Except in these points, the three families which the sub-order contains
are so much alike that they may conveniently be described together.
In the Branchipodidae the carapace is practically absent, being
represented only by the slight backward projection on each side of the
head which contains the kidney (Fig. 2); the paired eyes are supported on
mobile stalks, and project freely, one on either side of the head.
In the Apodidae[15] the head is broad and depressed, the ventral side
being nearly flat, the dorsal surface convex; the hinder margin of the head
is indicated dorsally by a transverse cervical ridge, bounded by two
grooves, behind which the carapace projects backwards as a great shield,
covering at least half the body, but attached only to the back of the head.
In Lepidurus productus the head and carapace together form an oval
expansion, deeply emarginate at the hinder, narrower end, the sides of
the emargination being toothed. The carapace has a strong median keel.
The kidneys project into the space between the folds of skin which form
the carapace, and their coils can be seen on each side, the terminal part of
each kidney-tube entering the head to open at the base of the second
maxilla. In all Branchiopoda with a well-developed carapace the kidney is
enclosed in it in this way, whence the older anatomists speak of it as the
“shell-gland.”
 Fig. 2.—Chirocephalus diaphanus, female, × 5, Sussex. D.O, Dorsal
organ; H, heart; Ov, ovary; U, uterus; V, external generative opening.

Associated with the development of the carapace, in this and in the next
family, is a remarkable condition of the lateral eyes, which are sessile on
the dorsal surface of the head, and near the middle line, the median eye
being slightly in front of them. During embryonic life a fold of skin grows
over all three eyes, so that a chamber is formed over them, which
communicates with the exterior by a small pore in front.
In the Limnadiidae the body is laterally compressed, and the carapace
is so large that at least the post-cephalic part of the body, and generally
the head also, can be enclosed within it.
In Limnetis (Fig. 3) the dorsal
surface of the head is bent
downwards and is much
compressed, the carapace being
attached to it only for a short
distance near the dorsal middle line.
The sides of the carapace are bent
downwards, and their margins can
be pulled together by a transverse
adductor muscle, so that the whole
structure forms an ovoid or
spheroidal case, from which the
head projects in front, while the rest
Fig. 3.—Limnetis brachyura, × 15. (After
of the body is entirely contained G. O. Sars.)
within it. When the adductor muscle
is relaxed the edges of the carapace
gape slightly, like the valves of a Lamellibranch shell, and food-particles
are drawn through the opening thus formed into the ventral groove by the
movements of the thoracic feet, locomotion being chiefly effected by the
rowing action of the second antennae, as in the Cladocera, to which all the
Limnadiidae present strong resemblances in their method of locomotion,
in the condition of the carapace, and in the form of the telson.
In Limnadia and Estheria the carapace projects not only backwards
from the point of attachment to the head, but also forwards, so that the
head can be enclosed by it, together with the rest of the body.
In all these genera the carapace is flexible along the middle dorsal line;
in Estheria especially the softening of the dorsal cuticle goes so far that a
definite hinge-line is formed, and this, together with the deposition of the
lateral cuticle in lines concentrically arranged round a projecting umbo,
gives the carapace a strong superficial likeness to a Lamellibranch shell,
for which it is said to be frequently mistaken by collectors.
The eyes of the Limnadiidae are enclosed in a chamber formed by a
growth of skin over them, as in Apodidae, but the pore by which this
chamber communicates with the exterior is even more minute than in
Apus. The paired eyes are so close together that they may touch
(Limnadia, Estheria) or fuse (Limnetis); they are farther back than in the
Apodidae, while the ventral curvature of the head causes the median eye
to lie below them. In all these points the eyes of the Limnadiidae are
intermediate between those of Apus and those of the Cladocera.
Dorsal Organ.—A structure very characteristic of adult Phyllopods is
the “dorsal organ” (Figs. 2, 5, D.O), whose function is in many cases
obscure. It is always a patch of modified cephalic ectoderm, supplied by a
nerve from the anterior ventral lobe of the brain on each side; but its
characters, and apparent function, differ in different forms. In the
Branchipodidae the dorsal organ is a circular patch, far forward on the
surface of the head (Figs. 2, 5, D.O). Its cells are arranged in groups,
which remind one of the retinulae in a compound eye; each cell contains a
solid concretion, and the concretions of a group may be so placed as to
look like a badly-formed rhabdom. Claus,[16] who first called attention to
this structure in the Branchipodidae, regarded it as a sense-organ. In
Apodidae the dorsal organ is an oval patch of columnar ectoderm,
immediately behind the eyes; it is slightly raised above the surrounding
skin, and is covered by a very delicate cuticle (with an opening to the
exterior?), and below it is a mass of connective tissue permeated by blood;
Bernard has suggested that it is an excretory organ.
Most Limnadiidae resemble the Cladocera in the possession of a
“dorsal organ” quite distinct from the above; in Limnetis and Estheria it
has the form of a small pit, lined by an apparently glandular ectoderm,
and this is its condition in many Cladocera; in Limnadia lenticularis it is
a patch of glandular epithelium on a raised papilla. Limnadia has been
observed to anchor itself to foreign objects by pressing its dorsal organ
against them, and many Cladocera do the same thing; Sida crystallina,
for example, will remain for hours attached by its dorsal organ to a
waterweed or to the side of an aquarium. Structures resembling a dorsal
organ occur in the larvae of many other Crustacea, but the presence of
this organ in the adult is confined to Branchiopods, and indeed in many
Cladocera it disappears before maturity. It is certain that the sensory and
adhesive types of dorsal organ are not homologous, especially as
rudimentary sense-organs may exist on the head of Cladocera together
with the adhesive organ.
The telson differs considerably in the different genera. In the
Branchipodidae[17] the anus opens directly backwards; and the telson
carries two flattened backwardly directed plates, one on each side of the
anus, the margins of each plate being fringed with plumose setae. In
Artemia the anal plates are rarely as large as in Branchipus, and never
have their margins completely fringed with setae; in A. salina from
Western Europe, and in A. fertilis (Fig. 4, A) from the Great Salt Lake of
Utah, there is a variable number of setae round the apical half of each
lobe, but in specimens of A. salina from Western Siberia the number of
setae may be very small, or they may be absent; in the closely allied A.
urmiana from Persia the anal lobes are well developed in the male, each
lobe bearing a single terminal hair, but they are altogether absent in the
female. Schmankewitch and Bateson have shown that there is a certain
relation between the salinity of the water in which Artemia salina occurs
and the condition of the anal lobes, specimens from denser waters having
on the whole fewer setae; the relation is, however, evidently very complex,
and further evidence is wanted before any more definite statements can
be made.
 Fig. 4.—A, Ventral view of the anal region in Artemia fertilis, from the
Great Salt Lake; B, ventral view of the telson and neighbouring parts of
Lepidurus productus; C, side view of the telson and left anal lobe of
Estheria (sp.?).

In the Apodidae the anal lobes have the form of two-jointed cirri, often
of considerable length; in Apus the anus is terminal, but in Lepidurus
(Fig. 4, B) the dorsal part of the telson is prolonged backwards, so as to
form a plate, on the ventral face of which the anus opens, much as in the
Malacostraca.
In the Limnadiidae (Fig. 4, C) the telson is laterally compressed and
produced, on each side of the anus, into a flattened, upwardly curved
process, sharply pointed posteriorly, and often serrate; the anal lobes are
represented by two stout curved spines, while in place of the dorsal
prolongation of Lepidurus we find two long plumose setae above the
anus. In the characters of the telson and anal lobes, as in those of the
head, the Limnadiidae approximate to the Cladocera. In Limnetis
brachyura the ventral face of the telson is produced into a plate
projecting backwards below the anus, in a manner which has no exact
parallel among other Crustacea.
The appendages of the Phyllopoda are fairly uniform in character,
except those affected by the sexual dimorphism, which is usually great.
 Fig. 5.—Chirocephalus diaphanus, male. Side view of head, showing
the large second antenna, A2, with its appendage Ap, above which is
seen the filiform first antenna; D.O, dorsal organ; E1, median eye.

Of the cephalic appendages, the first antennae are generally small, and
are never biramous; in Branchipus and its allies they are simple unjointed
rods, in some species of Artemia they are three-jointed, in Apus they are
feebly divided into two joints, while in Estheria they are many-jointed.
The second antennae are the principal organs of locomotion in the
Limnadiidae, where they are large and biramous; in all other Phyllopoda
they are uniramous in the female, being either unjointed triangular plates
as in Chirocephalus (Fig. 2), or minute vestigial filaments as in Apus, in
which genus Zaddach, Huxley, and Claus have all failed to find any trace
of a second antenna in some females. In the male Branchipodidae the
second antennae are modified to form claspers, by which the female is
seized, the various degrees of complication which these claspers exhibit
affording convenient generic characters. In Branchinecta each second
antenna is a thick, three-jointed rod, the last joint forming a claw, while
the second joint is serrate on its inner margin; in Branchipus the base is
much thickened, and bears on its inner side a large filament (perhaps
represented by the proximal tubercle of Branchinecta and Artemia),
which looks like an extra antenna. In Streptocephalus the terminal joint
of the antenna is bifid, and there is a basal filament like that of
Branchipus; in Chirocephalus
diaphanus (Figs. 5, 6) the main
branch of the antenna consists of
two large joints, the terminal joint
being a strong claw with a serrated
process at its base, while the
proximal joint bears two
appendages on its inner side; one of
these is a small, subconical tubercle,
the second is more complicated,
consisting of a main stem and five
outgrowths. The main stem is
many-jointed and flexible, its basal
joint being longer than the others,
and bearing on its outer side a large,
triangular, membranous appendage,
and four soft cylindrical
appendages, the main stem and its
appendages being beset with
curious tubercles, ending in short
spines, whose structure is not
understood. Except during the act of
copulation this remarkable Fig. 6.—Chirocephalus diaphanus.
Second antenna of male, uncoiled.
apparatus is coiled on the inner side
of the antennary claw, the jointed
stem being so coiled that it is often compared to the coiled proboscis of a
butterfly, and the triangular membrane folded like a fan beside it, so that
much of the organ is concealed, and the general appearance of the head is
that shown in Fig. 5. During copulation, the whole structure is widely
extended.
The males of Artemia (Fig. 7) have the second antenna two-jointed, the
basal joint bearing an inner tubercle, the terminal joint being flattened
and bluntly pointed, its outer margin provided with a membranous
outgrowth. In A. fertilis the breadth of the second joint varies greatly, the
narrower forms presenting a certain remote resemblance to
Branchinecta. In the males of Polyartemia the second antennae have a
remarkable branched form not easily comparable with that found in other
Branchipodidae.
The cephalic jaws are fairly uniform throughout the order. The
mandibles have an undivided molar surface, and no palp; the first maxilla
is very generally a triangular plate, with a setose biting edge; mandibles
 and maxillae are covered by the
labrum. The second maxilla
generally lies outside the chamber
formed by the labrum, and is a
simple oval plate, with or without a
special process for the duct of the
kidney.
The thoracic limbs, in front of the
genital segments, are not as a rule
differentiated into anterior
Fig. 7.—Artemia fertilis. Front view of the maxillipedes and posterior
head of a male, showing the large second locomotive appendages, as in higher
antennae, A.2; A.1, first antennae. forms; we have seen, however, that
all these limbs take part in the
prehension of food, and except in
the Limnadiidae they all assist in locomotion. One of the middle thoracic
legs of Artemia (Fig. 8, A) has a flattened stem, with seven processes on
its inner, and two on its outer margin. The gnathobase (gn) is large, and
fringed with long plumose setae, each of which is jointed; this is followed
by four smaller “endites” (or processes on the median side), and then by
two larger ones, the terminal endite (the sixth, excluding the gnathobase)
being very mobile and attached to the main stem by a definite joint. On
the outer side are two processes; a proximal “bract,” a flat plate with
crenate edges, partly divided by a constriction into two, and a distal
process, cylindrical and vascular, called by Sars and others the
“epipodite.” In other Branchipodidae we have essentially the same
condition, except that the fifth endite often becomes much larger than in
Artemia, throwing the terminal endite well over to the outer edge of the
limb; such a shift as this, continued farther, might well lead to the
condition found in the Limnadiidae, or Apodidae, where the lobe which
seems to represent the terminal endite of Artemia is entirely on the outer
border of the limb, forming what most writers have called the exopodite
(Lankester’s “flabellum”).[18] In the two last-named families the basal
exite or bract of the Branchipodidae does not appear to be represented.
 Fig. 8.—A, Thoracic limb of Chirocephalus diaphanus; B, prehensile
thoracic limb of male Estheria. gn, Gnathobase; 1–6, the more distal
endites.

The limbs of the Apodidae are remarkable in two ways; those in front of
the genital opening (very constantly ten pairs) are not so nearly alike as in
most genera of the sub-order, the first two pairs especially having the axis
definitely jointed, while the endites are elongated and antenniform;
further, while the first eleven segments bear each a single pair of limbs, as
is usual among Crustacea, many of the post-genital segments bear several
pairs; thus in Apus cancriformis there are thirty-two post-cephalic
segments in front of the telson, the first eleven having each one pair of
limbs, while the next seventeen have fifty-two pairs between them, the
last four segments having none.
In all the Phyllopoda some of the post-cephalic limbs are modified for
reproductive purposes; in the Branchipodidae the last two pairs (the 12th
and 13th generally, the 20th and 21st in Polyartemia) are so modified in
both sexes. In the female these appendages fuse at an early period of
larval life, and surround the median opening of the generative duct (Fig.
2); in the male the two pairs also fuse, but traces of the limbs are left as
eversible processes round the paired openings of the vasa deferentia.
 In the other families, one or more limbs of the female are adapted for
carrying or supporting the eggs. In the Apodidae the appendages of the
eleventh segment have the exopodite in the form of a rounded,
watchglass-shaped plate, fitting over a similarly shaped process of the
axis of the limb, so that a lens-shaped box is formed, into which the eggs
pass from the oviduct. In Limnadiidae the eggs are carried in masses
between the body and the carapace, and are kept in position by special
elongations of the exopodites of two or three legs, either those near the
middle of the thorax (Estheria, Limnadia), or at its posterior end
(Limnetis). In female Limnetis the last thoracic segments bear two
remarkable lateral plates, which apparently also help to support the eggs.
In the male Limnadiidae, the first (Limnetis) or the first two thoracic feet
(Limnadia, Estheria) are prehensile (Fig. 8, B).
Alimentary Canal.—The mouth of the Phyllopoda is overhung by the
large labrum, so that a kind of atrium is formed, outside the mouth itself,
in which mastication is performed; numerous unicellular glands, opening
on the oral face of the labrum, pour their secretion into the atrial
chamber, and may be called salivary, though the nature of their secretion
is not known. The mouth has commonly two swollen and setose lips,
running longitudinally forwards from the bases of the first maxillae, and
often wrapping round the blades of the mandibles. It leads into a vertical
oesophagus, which opens into a small globular stomach, lying entirely
within the head; the terminal part of the oesophagus is slightly
invaginated into the stomach, so that a valvular ring is formed at the
junction of the two. The stomach opens widely behind into a straight
intestine, which runs backwards to about the level of the telson, where it
joins a short rectum, leading to the terminal or ventral anus. The stomach
and intestine are lined by a columnar epithelium, and covered by a thin
network of circularly arranged muscle-fibres; the rectum has a flatter
epithelium, and radial muscles pass from it to the body-wall, so that it can
be dilated. The only special digestive glands are two branched glandular
tubes, situated entirely within the head, which open into the stomach by
large ducts, one on each side. In Chirocephalus the gastric glands are
fairly small and simple; in the Apodidae their branches are more complex
and form a considerable mass, filling all that portion of the head which is
not occupied by the nervous system and the muscles. Backwardly directed
gastric glands, like those of the higher Crustacea, are not found in
Branchiopods; both forms occur together in the genus Nebalia, but with
this exception the forwardly-directed glands are peculiar to
Branchiopods.
 Heart.—In Branchipus and its allies, and in Artemia, the heart
extends from the first thoracic segment to the penultimate segment of the
body, and is provided with eighteen pairs of lateral openings, one pair in
every segment through which it passes except the last; it is widely open at
its hinder end, and is prolonged in front for a short distance as a cephalic
aorta, the rest of the blood-spaces being lacunar.
In most, at least, of the other Branchiopods, the heart is closed behind
and is shortened; in Apus and Lepidurus it only extends through the first
eleven post-cephalic segments, while in the Limnadiidae it is shorter still,
the heart of Limnetis passing through four segments only. In all cases
there is a pair of lateral openings in every segment traversed by the heart.
The blood of the Branchipodidae and Apodidae contains dissolved
haemoglobin, the quantity present being so small as to give but a faint
colour to the blood in Branchipus, while Artemia has rather more, and
the blood of Apus is very red. The only other Crustacea in which the blood
contains haemoglobin are the Copepods of the genus Lernanthropus,[19]
so that the appearance of this substance is as irregular and inexplicable in
Crustacea as in Chaetopods and Molluscs.
The nervous system of Branchipus may be described as an
illustration of the condition prevailing in the group. The brain consists of
two closely united ganglia, in each of which three main regions may be
distinguished; a ventral anterior lobe, a dorsal anterior lobe, and a
posterior lobe. The ventral anterior lobes give off nerves to the median
eye, to the dorsal organ, and to a pair of curious sense-organs,
comparable with the larval sense-knobs of many higher forms, situated
one on each side of the median eye; in late larvae Claus describes the
terminal apparatus of each frontal sense-organ as a single large
hypodermic cell; W. K. Spencer[20] has lately described several terminal
cells, containing peculiar chitinous bodies, in the adult. The homologous
sense-organs of Limnetis are apparently olfactory. The dorsal anterior
lobes give off the large nerves to the lateral eyes, while the posterior lobes
supply the first antennae. The oesophageal connectives have a coating of
ganglion-cells, and some of these form the ganglion of the second
antenna, the nerve to this appendage leaving the connective just behind
the brain. The post-oral nerve-cords are widely separate, each of them
dilating into a ganglion opposite every appendage, the two ganglia being
connected by two transverse commissures. The ganglia of the three
cephalic jaws, so often fused in the higher Crustacea, are here perfectly
distinct. Closely connected with each thoracic ganglion is a remarkable
unicellular gland, opening to the exterior near the middle ventral line; it
is conceivable that these cells may be properly compared with the larval
nephridia of a Chaetopod,[21] but no evidence in support of such a
comparison has yet been adduced.
Behind the genital segments, where there are no limbs, the nerve-cords
run backwards without dilating into segmental ganglia, except in the
anterior two abdominal segments where small ganglionic enlargements
occur. In Apodidae, on the other hand, those segments which carry more
than one pair of appendages have as many pairs of ganglia, united by
transverse commissures, as they have limbs.
A stomatogastric nervous system exists in Apus, where a nerve arises
on each side from the first post-oral commissure, and runs forward to join
its fellow of the opposite side on the anterior wall of the oesophagus.
From the loop so formed a larger median and a series of smaller lateral
nerves pass to the wall of the alimentary canal. A second nerve to the
oesophagus is given off from the mandibular ganglion of each side.
Reproductive Organs.—In Chirocephalus the ovaries (Fig. 2, Ov)
are hollow epithelial tubes, lying one on each side of the alimentary canal,
and extending from the sixth abdominal segment forwards to the level of
the genital opening; at this point the two ovaries are continuous with
ducts, which bend sharply downwards and open into the single uterus
contained within the projecting egg-pouch and opening to the exterior at
the apex of that organ. Short diverticula of the walls of the uterus receive
the ducts of groups of unicellular glands, the bodies of which contain a
peculiar opaque secretion, said to form the eggshells. In Apodidae the
ovaries are similar in structure, but they are much larger and branch in a
complex manner, while each ovary opens to the exterior independently of
the other in the eleventh post-cephalic segment; nothing like the median
uterus of the Branchipodidae being formed. The epithelium of the ovarian
tubes proliferates, and groups of cells are formed; one becoming an ovum,
the others being nutrient cells like those which will be more fully
described in the Cladocera.
In Chirocephalus the testes are tubes similar in shape and position to
the ovaries, each communicating in front with a short vas deferens, which
dilates into a vesicula seminalis on its way to the eversible penis; an
essentially similar arrangement is found in all Branchipodidae, but in
Apodidae and Limnadiidae there is no penis.
All the Branchiopoda are dioecious,[22] and many are parthenogenetic.
Among Branchipodidae Artemia is the only genus known to be
parthenogenetic, but parthenogenesis is common in all Apodidae, while
the males of several species of Limnadia are still unknown, although the
females are sometimes exceedingly common. In Artemia, generations in
which the males are about as numerous as the females seem to alternate
fairly quickly with others which contain only parthenogenetic females; in
Apus males are rarely abundant, and often absent for long periods; during
five consecutive years von Siebold failed to discover a male in a locality in
Bavaria, though he examined many thousands of individuals; near
Breslau he found on one occasion about 11 per cent of males (114 in 1026),
but in a subsequent year he found less than 1 per cent; the greatest
recorded percentage of males is that observed by Lubbock in 1863, when
he found 33 males among 72 individuals taken near Rouen.
The eggs of most genera can resist prolonged periods of desiccation,
and indeed it seems necessary for the development of many species that
the eggs should be first dried and afterwards placed in water. Many eggs
(e.g. of Chirocephalus diaphanus and Branchipus stagnalis) float when
placed in water after desiccation, the development taking place at the
surface of the water.
Habitat.—All the Phyllopoda, except Artemia, are confined to
stagnant shallow waters, especially to such ponds as are formed during
spring rains, and dry up during the summer. In waters of this kind the
species of Branchipus, Apus, etc., develop rapidly, and produce great
numbers of eggs, which are left in the dried mud at the bottom after
evaporation of the water, where they remain quiescent until a fresh rainy
season. The mud from the beds of such temporary pools often contains
large numbers of eggs, which may be carried by wind, on the legs of birds,
and by other means, to considerable distances. Many exotic species have
been made known to European naturalists by their power of hatching out
when mud brought home by travellers is placed in water. The water of
stagnant pools quickly dissolves a certain quantity of solid matter from
the soil, and often receives dissolved solids through surface drainage from
the neighbouring land; such salts may remain as the water evaporates, so
that the water which remains after evaporation has proceeded for some
time may be very sensibly denser than that in which the Branchiopods
were hatched; these creatures must therefore be able to endure a
considerable increase in the salinity of the surrounding waters during the
course of their lives. My friend Mr. W. W. Fisher points out that the
plants present in such a pond would often precipitate the carbonate of
lime, so that this might be removed as evaporation went on, but that
chlorides would probably remain in solution; from analyses which Mr.
Fisher has been kind enough to make for me, it is seen that this happened
in a small aquarium in my laboratory, in which Chirocephalus diaphanus
lived for four months. In April, mud from the dry bed of a pond, known to
contain eggs of Chirocephalus, was placed in this aquarium in Oxford,
and water was added from the tap. Oxford tap-water contains about 0·3
grm. salts per litre, the chlorine being equivalent to 0·023 grm. NaCl.
Water was added from time to time during May and June, but in July
evaporation was allowed to proceed unchecked. At the end of July there
was about half the original volume of water, the Chirocephalus being still
active; the residue contained 0·96 grm. dissolved solids per litre, with
chlorine equal to 0·19 grm. NaCl, so that the percentage of chlorides was
about eight times the initial percentage, but there were only three and a
fifth times the original amount of total solid matter in solution, the
carbonate of lime having precipitated as a visible film.
Some species of Branchipus (e.g. B. spinosus, M. Edw.) and of Estheria
(E. macgillivrayi, Baird, E. gubernator, Klutzinger) occur in salt pools,
but Artemia flourishes in waters beside whose salinity that endured by
any other Branchiopod is insignificant. In the South of Europe, Artemia
salina may be found in swarms, as it used to be found in Dorsetshire, in
the shallow brine-pans from which salt is commercially prepared; Rathke
quotes an analysis showing that a pool in the Crimea contained living
Artemia when the salts in solution were 271 grms. per litre, and the water
was said to have the colour and consistency of beer.
The behaviour of the animals in the water differs a little; in normal
feeding all the species swim with the back downwards, as has already
been said; the Branchipodidae rarely settle on the ground, or on foreign
objects, but the Apodidae occasionally wriggle along the bottom on their
ventral surface, and Estheria burrows in mud.
The greater number of species are found in pools in flat, low-lying
regions, and many appear to be especially abundant near the sea; Apus
cancriformis has, however, been found in Armenia at 10,000 feet above
sea level.
Wells and underground waters do not generally contain Phyllopods;
but a species of Branchipus and one of Limnetis, both blind, have been
described from the caves of Carniola.
One of the many puzzles presented by these creatures is the erratic way
in which they are scattered through the regions they inhabit; a single
small pond, a few yards or less in diameter, may be the only place within
many miles in which a given species can be found; in this pond it may,
however, appear regularly season after season for some time, and then
suddenly vanish.
Geographically, the Phyllopoda are cosmopolitan, representatives of
every family and of some genera (e.g. Streptocephalus, Lepidurus,
Estheria) being found in every one of the great zoological regions, though
a few aberrant genera are of limited range, thus Polyartemia is known
only from the northern Palaearctic and Nearctic regions,
Thamnocephalus only from the Central United States. The genus Artemia
is not at present known in Australia.[23] The only recorded British species
are Chirocephalus diaphanus, Artemia salina, and Apus cancriformis,[24]
but other continental islands, for example the West Indian group, are
better supplied. The distribution of the species is very imperfectly known,
but on the whole every main zoological region seems to have its own
peculiar species, which do not pass beyond its boundaries. Branchinecta
paludosa and Lepidurus glacialis are circumpolar, both occurring in
Norway, in Lapland, in Greenland, and in Arctic North America; but with
these exceptions the Palaearctic and Nearctic species seem to be distinct.
The European species Apus cancriformis occurs in Algiers, but the
relations between the species of Northern Africa as a whole and those of
Southern Europe on the one hand, or of Central and Southern Africa on
the other, have yet to be worked out.
The soft-bodied Branchipodidae are not known in the fossil condition;
[25]
an Apus, closely related to the modern A. cancriformis, has been
found in the Trias, but the most numerous remains have been left, as
might be expected, by the hard-shelled Limnadiidae; carapaces, closely
resembling those of the modern Estheria, are known in beds of all ages
from the Devonian period to recent times; these carapaces are in several
cases associated with fossils of an apparently marine type. None of the
fossil species differ in any important characters from those now living, so
that the Phyllopoda have existed in practically their present form for an
enormously long period; this fact, and the evidence that species of
existing genera were at one time marine, explain the wide distribution of
animals at present restricted to a remarkably limited range of
environmental conditions.

Summary of the Characters of the Genera.

Sub-Order Phyllopoda.—Branchiopoda with an elongated body,

provided with at least ten pairs of post-cephalic limbs, the heart
extending through four or more thoracic segments, and having at
least four pairs of ostia.
Fam. 1. Branchipodidae.[26]—Carapace rudimentary, eyes stalked;
the second antennae flat and unjointed in the female, jointed and
prehensile in the male; female generative opening single; telson not
laterally compressed, bearing two flattened lobes, or none. The heart
extending through the thorax and the greater part of the abdomen.
A. Eleven pairs of praegenital ambulatory limbs.
a. Abdomen of six well-formed segments and a telson; anal
lobes well formed, their margins setose.
Branchinecta, Verrill—Second antennae of ♂ without
lateral appendages; ovisac of ♀ elongated. B. paludosa,
O. F. Müll.—Circumpolar.
Branchiopodopsis, G. O. Sars[27]—Second antennae of ♂ as
in Branchinecta; ovisac of ♀ short. B. hodgsoni, G. O.
Sars—Cape of Good Hope.
Branchipus, Schaeffer—Second antennae of ♂ with simple
internal filamentous appendage. B. stagnalis, Linn.—
Central Europe.
Streptocephalus, Baird—Second antennae of ♂ 3–jointed,
the last joint bifid; an external filamentous appendage. S.
torvicornis, Wagn., Poland.
Chirocephalus, Prévost—Second antennae of ♂ 3–jointed,
with a jointed internal appendage, which bears secondary
processes, four cylindrical and one lamellar. C.
diaphanus, Prévost (Fig. 2, p. 20).—Britain, Central
Europe.
b. Abdominal segments five or fewer, and a telson. Anal lobes
small or 0, sparsely or not at all setose.
Artemia, Leach—Second antennae of ♂ without
filamentous appendage, 2–jointed, the second joint
lamellar. A. salina, Linn.—Brine pools of the Palaearctic
region.
c. Hinder abdominal segments united with telson to form a fin;
anal lobes absent.
Thamnocephalus, Packard—Head with a branched median
process of unknown nature. Only species T. platyurus,
Packard—Kansas, U.S.A.
B. Nineteen pairs of praegenital ambulatory limbs.
 Polyartemia, Fischer—Second antennae of ♂ forcipate;
ovisac of ♀ very short. Only species P. forcipata, Fisch.
Fam. 2. Apodidae.[28]—Carapace well developed as a depressed
shield, covering at least half the body. Eyes sessile, covered; no male
clasping organs; anal lobes long, jointed cirri.
Apus, Scopoli—Telson not produced backwards over the
anus; endites of first thoracic limb very long. A.
cancriformis, Schaeffer—Britain, Europe, Algiers, Tunis.
A. australiensis, Central Australia.
Lepidurus, Leach—Telson produced backwards to form a
plate above the anus; endites of first thoracic limb short.
L. productus, Bosc.—Central Europe. L. viridis, Southern
Australia, New Zealand, L. patagonicus, Bergh,
Argentines.
Fam. 3. Limnadiidae.—Body compressed; carapace in the form
of a bivalve shell, the two halves capable of adduction by means of a
strong transverse muscle; second antennae biramous, alike in both
sexes; in the male, the first or the first and second thoracic limbs
prehensile; telson laterally compressed.
A. Only the first thoracic limbs prehensile in the male; the carapace
spheroidal, without lines of growth; head not included within the
carapace-chamber.
Limnetis, Lovén—Compound eyes fused; anal spines
absent; ambulatory limbs 10–12. L. brachyura, O. F.
Müll (Fig. 3, p. 21).—Norway, Central Europe.
B. The first and second thoracic limbs prehensile in the male;
carapace distinctly bivalve, enclosing the head, with concentric
lines of growth round a more or less prominent umbo.
Eulimnadia, Packard—Carapace narrowly ovate, with few
(4–5) lines of growth. E. mauritani, Guérin—Mauritius.
E. texana, Packard—Texas, Kansas.
Limnadia, Brongniart—Carapace broadly ovate, with
numerous lines of growth, without distinct umbones; L.
lenticularis, Linn.—Northern and Central Europe.
Estheria, Rüppell—Carapace with well-marked umbones
and numerous lines of growth, oval; E. tetraceros,
Kryneki—Central Europe.