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 Example 4.3—Pressure Drop in a Packed-Bed Reactor
4.5 Filtration
4.6 Fluidization
4.7 Dynamics of a Bubble-Cap Distillation Column
4.8 Cyclone Separators
4.9 Sedimentation
4.10 Dimensional Analysis
Example 4.4—Thickness of the Laminar Sublayer
Problems for Chapter 4
PART II—MICROSCOPIC FLUID MECHANICS
CHAPTER 5—DIFFERENTIAL EQUATIONS OF FLUID
MECHANICS
5.1 Introduction to Vector Analysis
5.2 Vector Operations
Example 5.1—The Gradient of a Scalar
Example 5.2—The Divergence of a Vector
Example 5.3—An Alternative to the Differential Element
Example 5.4—The Curl of a Vector
Example 5.5—The Laplacian of a Scalar
5.3 Other Coordinate Systems
5.4 The Convective Derivative
5.5 Differential Mass Balance
Example 5.6—Physical Interpretation of the Net Rate of Mass
Outflow
Example 5.7—Alternative Derivation of the Continuity Equation
5.6 Differential Momentum Balances
5.7 Newtonian Stress Components in Cartesian Coordinates
Example 5.8—Constant-Viscosity Momentum Balances in
Terms of Velocity Gradients
Example 5.9—Vector Form of Variable-Viscosity Momentum
Balance
Problems for Chapter 5
CHAPTER 6—SOLUTION OF VISCOUS-FLOW PROBLEMS

8
 6.1 Introduction
6.2 Solution of the Equations of Motion in Rectangular Coordinates
Example 6.1—Flow Between Parallel Plates
6.3 Alternative Solution Using a Shell Balance
Example 6.2—Shell Balance for Flow Between Parallel Plates
Example 6.3—Film Flow on a Moving Substrate
Example 6.4—Transient Viscous Diffusion of Momentum
(COMSOL)
6.4 Poiseuille and Couette Flows in Polymer Processing
Example 6.5—The Single-Screw Extruder
Example 6.6—Flow Patterns in a Screw Extruder (COMSOL)
6.5 Solution of the Equations of Motion in Cylindrical Coordinates
Example 6.7—Flow Through an Annular Die
Example 6.8—Spinning a Polymeric Fiber
6.6 Solution of the Equations of Motion in Spherical Coordinates
Example 6.9—Analysis of a Cone-and-Plate Rheometer
Problems for Chapter 6
CHAPTER 7—LAPLACE’S EQUATION, IRROTATIONAL AND
POROUS-MEDIA FLOWS
7.1 Introduction
7.2 Rotational and Irrotational Flows
Example 7.1—Forced and Free Vortices
7.3 Steady Two-Dimensional Irrotational Flow
7.4 Physical Interpretation of the Stream Function
7.5 Examples of Planar Irrotational Flow
Example 7.2—Stagnation Flow
Example 7.3—Combination of a Uniform Stream and a Line
Sink (C)
Example 7.4—Flow Patterns in a Lake (COMSOL)
7.6 Axially Symmetric Irrotational Flow
7.7 Uniform Streams and Point Sources
7.8 Doublets and Flow Past a Sphere
7.9 Single-Phase Flow in a Porous Medium

9
 Example 7.5—Underground Flow of Water
7.10 Two-Phase Flow in Porous Media
7.11 Wave Motion in Deep Water
Problems for Chapter 7
CHAPTER 8—BOUNDARY-LAYER AND OTHER NEARLY
UNIDIRECTIONAL FLOWS
8.1 Introduction
8.2 Simplified Treatment of Laminar Flow Past a Flat Plate
Example 8.1—Flow in an Air Intake (C)
8.3 Simplification of the Equations of Motion
8.4 Blasius Solution for Boundary-Layer Flow
8.5 Turbulent Boundary Layers
Example 8.2—Laminar and Turbulent Boundary Layers
Compared
8.6 Dimensional Analysis of the Boundary-Layer Problem
8.7 Boundary-Layer Separation
Example 8.3—Boundary-Layer Flow Between Parallel Plates
(COMSOL)
Example 8.4—Entrance Region for Laminar Flow Between Flat
Plates
8.8 The Lubrication Approximation
Example 8.5—Flow in a Lubricated Bearing (COMSOL)
8.9 Polymer Processing by Calendering
Example 8.6—Pressure Distribution in a Calendered Sheet
8.10 Thin Films and Surface Tension
Problems for Chapter 8
CHAPTER 9—TURBULENT FLOW
9.1 Introduction
Example 9.1—Numerical Illustration of a Reynolds Stress Term
9.2 Physical Interpretation of the Reynolds Stresses
9.3 Mixing-Length Theory
9.4 Determination of Eddy Kinematic Viscosity and Mixing Length
9.5 Velocity Profiles Based on Mixing-Length Theory

10
 Example 9.2—Investigation of the von Kármán Hypothesis
9.6 The Universal Velocity Profile for Smooth Pipes
9.7 Friction Factor in Terms of Reynolds Number for Smooth Pipes
Example 9.3—Expression for the Mean Velocity
9.8 Thickness of the Laminar Sublayer
9.9 Velocity Profiles and Friction Factor for Rough Pipe
9.10 Blasius-Type Law and the Power-Law Velocity Profile
9.11 A Correlation for the Reynolds Stresses
9.12 Computation of Turbulence by the k−ε Method
Example 9.4—Flow Through an Orifice Plate (COMSOL)
Example 9.5—Turbulent Flow in an Obstructed U-Duct
(COMSOL)
9.13 Analogies Between Momentum and Heat Transfer
Example 9.6—Evaluation of the Momentum/Heat-Transfer
Analogies
9.14 Turbulent Jets
Problems for Chapter 9
CHAPTER 10—BUBBLE MOTION, TWO-PHASE FLOW, AND
FLUIDIZATION
10.1 Introduction
10.2 Rise of Bubbles in Unconfined Liquids
Example 10.1—Rise Velocity of Single Bubbles
10.3 Pressure Drop and Void Fraction in Horizontal Pipes
Example 10.2—Two-Phase Flow in a Horizontal Pipe
10.4 Two-Phase Flow in Vertical Pipes
Example 10.3—Limits of Bubble Flow
Example 10.4—Performance of a Gas-Lift Pump
Example 10.5—Two-Phase Flow in a Vertical Pipe
10.5 Flooding
10.6 Introduction to Fluidization
10.7 Bubble Mechanics
10.8 Bubbles in Aggregatively Fluidized Beds
Example 10.6—Fluidized Bed with Reaction (C)

11
 Problems for Chapter 10
CHAPTER 11—NON-NEWTONIAN FLUIDS
11.1 Introduction
11.2 Classification of Non-Newtonian Fluids
11.3 Constitutive Equations for Inelastic Viscous Fluids
Example 11.1—Pipe Flow of a Power-Law Fluid
Example 11.2—Pipe Flow of a Bingham Plastic
Example 11.3—Non-Newtonian Flow in a Die (COMSOL)
11.4 Constitutive Equations for Viscoelastic Fluids
11.5 Response to Oscillatory Shear
11.6 Characterization of the Rheological Properties of Fluids
Example 11.4—Proof of the Rabinowitsch Equation
Example 11.5—Working Equation for a Coaxial-Cylinder
Rheometer: Newtonian Fluid
Problems for Chapter 11
CHAPTER 12—MICROFLUIDICS AND ELECTROKINETIC FLOW
EFFECTS
12.1 Introduction
12.2 Physics of Microscale Fluid Mechanics
12.3 Pressure-Driven Flow Through Microscale Tubes
Example 12.1—Calculation of Reynolds Numbers
12.4 Mixing, Transport, and Dispersion
12.5 Species, Energy, and Charge Transport
12.6 The Electrical Double Layer and Electrokinetic Phenomena
Example 12.2—Relative Magnitudes of Electroosmotic and
Pressure-Driven Flows
Example 12.3—Electroosmotic Flow Around a Particle
Example 12.4—Electroosmosis in a Microchannel (COMSOL)
Example 12.5—Electroosmotic Switching in a Branched
Microchannel (COMSOL)
12.7 Measuring the Zeta Potential
Example 12.6—Magnitude of Typical Streaming Potentials
12.8 Electroviscosity

12
 12.9 Particle and Macromolecule Motion in Microfluidic Channels
Example 12.7—Gravitational and Magnetic Settling of Assay
Beads
Problems for Chapter 12
CHAPTER 13—AN INTRODUCTION TO COMPUTATIONAL
FLUID DYNAMICS AND ANSYS FLUENT
13.1 Introduction and Motivation
13.2 Numerical Methods
13.3 Learning CFD by Using ANSYS Fluent
13.4 Practical CFD Examples
Example 13.1—Fluent: Developing Flow in a Pipe Entrance
Region
Example 13.2—Fluent: Pipe Flow Through a Sudden Expansion
Example 13.3—Fluent: A Two-Dimensional Mixing Junction
Example 13.4—Fluent: Flow over a Cylinder
References for Chapter 13
CHAPTER 14—COMSOL MULTIPHYSICS FOR SOLVING FLUID
MECHANICS PROBLEMS
14.1 COMSOL Multiphysics—An Overview
14.2 The Steps for Solving Problems in COMSOL
14.3 How to Run COMSOL
Example 14.1—Flow in a Porous Medium with an Impervious
Hole (COMSOL)
Example 14.2—Drawing a Complex Shape (COMSOL)
14.4 Variables, Constants, Expressions, and Units
14.5 Boundary Conditions
14.6 Variables Used by COMSOL
14.7 Wall Functions in Turbulent-Flow Problems
14.8 Streamline Plotting in COMSOL
14.9 Special COMSOL Features Used in the Examples
14.10 Drawing Tools
14.11 Fluid Mechanics Problems Solvable by COMSOL
14.12 Conclusion—Problems and Learning Tools

13
APPENDIX A: USEFUL MATHEMATICAL RELATIONSHIPS
APPENDIX B: ANSWERS TO THE TRUE/FALSE ASSERTIONS
APPENDIX C: SOME VECTOR AND TENSOR OPERATIONS
GENERAL INDEX
COMSOL MULTIPHYSICS INDEX
THE AUTHORS

14
Preface

THIS text has evolved from a need for a single volume that embraces a
very wide range of topics in fluid mechanics. The material consists of two
parts—four chapters on macroscopic or relatively large-scale phenomena,
followed by 10 chapters on microscopic or relatively small-scale
phenomena. Throughout, I have tried to keep in mind topics of industrial
importance to the chemical engineer. The scheme is summarized in the
following list of chapters.
Part I—Macroscopic Fluid Mechanics

1. Introduction to Fluid Mechanics

3. Fluid Friction in Pipes
2. Mass, Energy, and Momentum
4. Flow in Chemical Balances Engineering Equipment
Part II—Microscopic Fluid Mechanics

5. Differential Equations of Fluid

6. Solution of Viscous-Flow Problems Electrokinetic Flow Effects
7. Laplace’s Equation, Irrotational
8. Boundary-Layer and Other Dynamics and Fluent Nearly
Unidirectional Flows
9. Turbulent Flow
10. Bubble Motion, Two-Phase Flow, and Fluidization
11. Non-Newtonian Fluids Mechanics
12. Microfluidics and
13. An Introduction to and Porous-Media Flows Computational Fluid
14. COMSOL Multiphysics for Solving Fluid Mechanics Problems
In our experience, an undergraduate fluid mechanics course can be
based on Part I plus selected parts of Part II, and a graduate course can be
based on much of Part II, supplemented perhaps by additional material on
topics such as approximate methods and stability.
Third edition. I have attempted to bring the book up to date by the
major addition of Chapters 12, 13, and 14—one on microfluidics and two
on CFD (computational fluid dynamics). The choice of software for the
CFD presented a difficulty; for various reasons, I selected ANSYS Fluent

15
and COMSOL Multiphysics, but there was no intention of “promoting”
these in favor of other excellent CFD programs. The use of CFD examples
in the classroom really makes the subject come “alive,” because the
previous restrictive necessities of “nice” geometries and constant physical
properties, etc., can now be lifted. Chapter 9, on turbulence, has also been
extensively rewritten; here again, CFD allows us to venture beyond the
usual flow in a pipe or between parallel plates and to investigate further
practical situations such as turbulent mixing and recirculating flows.
Example problems. There is an average of about six completely
worked examples in each chapter, including several involving COMSOL
(dispersed throughout Part II) and Fluent (all in Chapter 13). The end of
each example is marked by a small square: □. All the COMSOL examples
have been run with Version 5.2a, both on a Mac Book Pro computer and
on Linux and Windows platforms; those using other releases of COMSOL
may encounter slightly different windows than those reproduced here. The
format for each COMSOL example is: (a) problem statement, (b) details of
COMSOL implementation, (c) results, and (d) discussion.
The numerous end-of-chapter problems have been classified roughly as
easy (E), moderate (M), or difficult/lengthy (D). The University of
Cambridge has given permission, kindly endorsed by Professor J.F.
Davidson, F.R.S., for several of their chemical engineering examination
problems to be reproduced in original or modified form, and these have
been given the additional designation of “(C).”
Acknowledgments. I gratefully acknowledge the valuable contributions
of my former Michigan colleague Stacy Birmingham (non-Newtonian
fluids), Brian Kirby of Cornell University (microfluidics), and Chi-Yang
Cheng of ANSYS, Inc. (CFD). My former doctoral student and good
friend Kevin Ellwood has been enormously helpful with this third edition.
Although I wrote most of the original examples and Chapter 14 (in
COMSOL 3.2), Kevin has supplied all the necessary expertise to rewrite
them in the longer and more comprehensive COMSOL 5.2a.
I have had much help from many people at COMSOL Inc. and
COMSOL AB. On their part, it has been a great cooperative effort all the
way, involving the COMSOL Development, Licensing, and Applications
teams, and I am very grateful to all for their assistance. At ANSYS, Inc.,
Chi-Yang Cheng was ideally suited for writing and updating the chapter
on Computational Fluid Dynamics and Fluent.
I have appreciated the assistance of several other friends and colleagues,
including Nitin Anturkar, Mark Burns, John Ellis, Scott Fogler, Tom
Grindley, Amy Horvath, Leenaporn Jongpaiboonkit, Lisa Keyser, Ronald

16
Larson, Susan Montgomery, Sunitha Nagrath, Michael Solomon, Sandra
Swisher, Robert Ziff, my wife, Mary Ann Gibson Wilkes, and the late
Stuart Churchill, Kartic Khilar, Donald Nicklin, Margaret Sansom, and
Rasin Tek. I also drew much inspiration from my many students and
friends at the University of Michigan and Chulalongkorn University in
Bangkok. Others are acknowledged in specific literature citations.
Also exceptionally helpful, with prompt attention for this third edition,
were the Prentice Hall editing and production team, to whom I extend my
gratitude: Kathleen Karcher, Carol Lallier, Laura Lewin, Julie Nahil, and
Dana Wilson.
Further information. The website http://fmche.engin.umich.edu is
maintained as a “bulletin board” for giving additional information about
the book—hints for problem solutions, errata, how to contact the authors,
etc.—as proves desirable. My own Internet address is [email protected].
The text was composed on an old but very faithful Power Macintosh G5
computer using the TEXtures “typesetting” program. Eleven-point type
was used for the majority of the text. Most of the figures were constructed
using MacDraw Pro, Excel, and Kaleida-Graph.
Professor Terence Fox, to whose memory this book is dedicated, was a
Cambridge engineering graduate who worked from 1933 to 1937 at
Imperial Chemical Industries Ltd., Billingham, Yorkshire. Returning to
Cambridge, he taught engineering from 1937 to 1946 before being selected
to lead the Department of Chemical Engineering at the University of
Cambridge during its formative years after the end of World War II. As a
scholar and a gentleman, Fox was a shy but exceptionally brilliant person
who had great insight into what was important and who quickly brought
the department to a preeminent position, which it still maintains. He
succeeded in combining an industrial perspective with intellectual rigor.
Fox relinquished the leadership of the department in 1959, after he had
secured a permanent new building for it (carefully designed in part by
himself).1
1 The department—now Chemical Engineering and Biotechnology—has just
(2017) moved to a new building on the West Cambridge site.

17
 T.R.C. Fox

Fox was instrumental in bringing an outstanding cast of faculty

members into the department during my student years there—Stan Sellers,
Kenneth Denbigh, John Davidson, Peter Danckwerts, Denys Armstrong,
and Peter Gray. He also kindly accepted me in 1956 as a junior faculty
member, and I spent four good years in the Cambridge University
Department of Chemical Engineering. Danckwerts subsequently wrote an
appreciation2 of Fox’s talents, saying, with almost complete accuracy:
“Fox instigated no research and published nothing.” How times have
changed—today, unless he were known personally, his résumé would
probably be cast aside and he would stand little chance of being hired, let
alone of receiving tenure! However, his lectures, meticulously written
handouts, enthusiasm, genius, and friendship were a great inspiration to
me, and I have much pleasure in acknowledging his positive impact on my
career.
2 P.V. Danckwerts, “Chemical engineering comes to Cambridge,” The
Cambridge Review , pp. 53–55, February 28, 1983.
James O. Wilkes
August 18, 2017

18
 Part I: Macroscopic Fluid
Mechanics
Some Greek Letters

α alpha
β beta
γ, Γ gamma
δ, Δ delta
ε, ε epsilon
ζ zeta
η eta
θ, ϑ, Θ theta
ι iota
κ kappa
λ, Λ lambda
μ mu
ν nu
ξ, Ξ xi
º omicron
π, Π pi
ρ, ϱ rho
σ, ς, Σ sigma
τ tau
υ, Υ upsilon
φ, ϕ, Φ phi

19
χ chi
ψ, Ψ psi
ω, Ω omega

20
Chapter 1. Introduction to Fluid Mechanics

1.1 Fluid Mechanics in Chemical Engineering

A knowledge of fluid mechanics is essential for the chemical engineer
because the majority of chemical-processing operations are conducted
either partly or totally in the fluid phase. Examples of such operations
abound in the biochemical, chemical, energy, fermentation, materials,
mining, petroleum, pharmaceuticals, polymer, and waste-processing
industries.
There are two principal reasons for placing such an emphasis on fluids.
First, at typical operating conditions, an enormous number of materials
normally exist as gases or liquids or can be transformed into such phases.
Second, it is usually more efficient and cost-effective to work with fluids
in contrast to solids. Even some operations with solids can be conducted in
a quasi-fluidlike manner; examples are the fluidized-bed catalytic refining
of hydrocarbons and the long-distance pipelining of coal particles using
water as the agitating and transporting medium.
Although there is inevitably a significant amount of theoretical
development, almost all the material in this book has some application to
chemical processing and other important practical situations. Throughout,
we shall endeavor to present an understanding of the physical behavior
involved; only then is it really possible to comprehend the accompanying
theory and equations.

1.2 General Concepts of a Fluid

We must begin by responding to the question, “What is a fluid?”
Broadly speaking, a fluid is a substance that will deform continuously
when it is subjected to a tangential or shear force, much as a similar type
of force is exerted when a water-skier skims over the surface of a lake or
butter is spread on a slice of bread. The rate at which the fluid deforms
continuously depends not only on the magnitude of the applied force but
also on a property of the fluid called its viscosity or resistance to
deformation and flow. Solids will also deform when sheared, but a
position of equilibrium is soon reached in which elastic forces induced by
the deformation of the solid exactly counterbalance the applied shear
force, and further deformation ceases.
A simple apparatus for shearing a fluid is shown in Fig. 1.1. The fluid is
contained between two concentric cylinders; the outer cylinder is

21
stationary, and the inner one (of radius R) is rotated steadily with an
angular velocity ω. This shearing motion of a fluid can continue
indefinitely, provided that a source of energy—supplied by means of a
torque here—is available for rotating the inner cylinder. The diagram also
shows the resulting velocity profile; note that the velocity in the direction
of rotation varies from the peripheral velocity Rω of the inner cylinder
down to zero at the outer stationary cylinder, these representing typical no-
slip conditions at both locations. However, if the intervening space is filled
with a solid—even one with obvious elasticity, such as rubber—only a
limited rotation will be possible before a position of equilibrium is
reached, unless, of course, the torque is so high that slip occurs between
the rubber and the cylinder.

Fig. 1.1 Shearing of a fluid.

There are various classes of fluids. Those that behave according to nice
and obvious simple laws, such as water, oil, and air, are generally called
Newtonian fluids. These fluids exhibit constant viscosity but, under typical
processing conditions, virtually no elasticity. Fortunately, a very large
number of fluids of interest to the chemical engineer exhibit Newtonian
behavior, which will be assumed throughout the book, except in Chapter
11, which is devoted to the study of non-Newtonian fluids.
A fluid whose viscosity is not constant (but depends, for example, on
the intensity to which it is being sheared), or which exhibits significant
elasticity, is termed non-Newtonian. For example, several polymeric
materials subject to deformation can “remember” their recent molecular
configurations, and in attempting to recover their recent states, they will

22
exhibit elasticity in addition to viscosity. Other fluids, such as drilling mud
and toothpaste, behave essentially as solids and will not flow when subject
to small shear forces, but will flow readily under the influence of high
shear forces.
Fluids can also be broadly classified into two main categories—liquids
and gases. Liquids are characterized by relatively high densities and
viscosities, with molecules close together; their volumes tend to remain
constant, roughly independent of pressure, temperature, or the size of the
vessels containing them. Gases, on the other hand, have relatively low
densities and viscosities, with molecules far apart; generally, they will
compleetely fill the container in which they are placed. However, these
two states—liquid and gaseous—represent but the two extreme ends of a
continuous spectrum of possibilities.
The situation is readily illustrated by considering a fluid that is initially
a gas at point G on the pressure/temperature diagram shown in Fig. 1.2. By
increasing the pressure, and perhaps lowering the temperature, the vapor-
pressure curve is soon reached and crossed, and the fluid condenses and
apparently becomes a liquid at point L. By continuously adjusting the
pressure and temperature so that the clockwise path is followed, and
circumnavigating the critical point C in the process, the fluid is returned to
G, where it is presumably once more a gas. But where does the transition
from liquid at L to gas at G occur? The answer is at no single point, but
rather that the change is a continuous and gradual one, through a whole
spectrum of intermediate states.

Fig. 1.2 When does a liquid become a gas?

1.3 Stresses, Pressure, Velocity, and the Basic Laws

Stresses. The concept of a force should be readily apparent. In fluid
mechanics, a force per unit area, called a stress, is usually found to be a
more convenient and versatile quantity than the force itself. Further, when

23
considering a specific surface, there are two types of stresses that are
particularly important.
1. The first type of stress, shown in Fig. 1.3(a), acts perpendicularly to
the surface and is therefore called a normal stress; it will be tensile or
compressive, depending on whether it tends to stretch or to compress the
fluid on which it acts. The normal stress equals F/A, where F is the normal
force and A is the area of the surface on which it acts. The dotted outlines
show the volume changes caused by deformation. In fluid mechanics,
pressure is usually the most important type of compressive stress, and will
shortly be discussed in more detail.

Fig. 1.3(a) Tensile and compressive normal stresses F/A, acting on a

cylinder, causing elongation and shrinkage, respectively.
2. The second type of stress, shown in Fig. 1.3(b), acts tangentially to
the surface; it is called a shear stress τ and equals F/A, where F is the
tangential force and A is the area on which it acts. Shear stress is
transmitted through a fluid by interaction of the molecules with one
another. A knowledge of the shear stress is very important when studying
the flow of viscous Newtonian fluids. For a given rate of deformation,
measured by the time derivative dγ/dt of the small angle of deformation γ,
the shear stress τ is directly proportional to the viscosity of the fluid (see
Fig. 1.3(b)).

Fig. 1.3(b) Shear stress τ = F/A, acting on a rectangular parallelepiped,

shown in cross section, causing a deformation measured by the angle γ
(whose magnitude is exaggerated here).

24
 Pressure. In virtually all hydrostatic situations—those involving fluids
at rest—the fluid molecules are in a state of compression. For example, for
the swimming pool whose cross section is depicted in Fig. 1.4, this
compression at a typical point P is caused by the downward gravitational
weight of the water above point P. The degree of compression is measured
by a scalar, p—the pressure.

Fig. 1.4 (a) Balloon submerged in a swimming pool; (b) enlarged view
of the compressed balloon, with pressure forces acting on it.
A small inflated spherical balloon pulled down from the surface and
tethered at the bottom by a weight will still retain its spherical shape (apart
from a small distortion at the point of the tether), but will be diminished in
size, as in Fig. 1.4(a). It is apparent that there must be forces acting
normally inward on the surface of the balloon and that these must
essentially be uniform for the shape to remain spherical, as in Fig. 1.4(b).
Although the pressure p is a scalar, it typically appears in tandem with
an area A (assumed small enough so that the pressure is uniform over it).
By definition of pressure, the surface experiences a normal compressive
force F = pA. Thus, pressure has units of a force per unit area—the same
as a stress.
The value of the pressure at a point is independent of the orientation of
any area associated with it, as can be deduced with reference to a
differentially small wedge-shaped element of the fluid, shown in Fig. 1.5.

25
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 Fig. 174.—Development of wings in Caloptenus spretus: the upper row
gives a lateral view of the thoracic segments, and the lower row a
dorsal view of these segments; 1, second instar; 2, third instar; 3,
fourth instar; 4, fifth instar. (After Riley.) t, tegmen; w, wing.

Fig. 175.—Caloptenus spretus. North America. A, Newly hatched,

much magnified; B, adult, natural size. (After Riley.)

Riley and Packard have given an account[221] of some parts of the

post-embryonic development of the Rocky Mountain Locust, which
enables us to form a satisfactory conception of the stages of
development of the wings. Fig. 175, A, represents the first instar, the
young locust, just emerged from the egg and colourless. Fig. 174
shows some of the subsequent stages of development of the wings,
the upper line of figures giving a profile view of the thoracic
segments, and the lower line showing their dorsal aspects; 1 shows
the condition of the parts in the second instar, the chief difference
from the first instar being the development of colour; in the third
instar there is an evident slight development of the future alar
organs, exhibited chiefly in the outgrowth and lobing of the free
posterior angles of the meso- and metanota, as shown in Fig. 174, 2.
After the third moult there is a great difference; the instar then
disclosed—the fourth—has undergone a considerable change in the
position of the meso- and metathoraces, which are thrust forward
under the pronotum; this has become more enlarged and hood-like
(Fig. 174, 3); at the same time the wing-rudiments have become free
and detached, the metathoracic pair being the larger, and
overlapping the other pair. The fifth instar (Fig. 174, 4) differs but
little from the fourth, except in the larger size of the pronotum and
wing-rudiments. The sixth—shown in Fig. 175, B—is the perfect
Insect, with the alar organs free and large, the prothorax much
changed in form, the colour different. From the above it will be seen
that the chief changes occurred at the third and fifth ecdyses, after
each of which a considerable difference in the form of the Insect was
revealed. In the first three instars the sexes can scarcely be
distinguished, in the fourth they are quite distinct, and in the fifth
coupling is possible, though usually it does not occur till the final
stage is attained.

The discovery that Orthoptera change their colours in the course of

their development, and even after they have become adult, is
important, not only from a physiological point of view, but because it
throws some light on the questions as to the number of species and
the geographical distribution of the migratory locusts, as to which
there has existed a great confusion.

The Acridiidae are considered to be exclusively vegetable feeders,

each individual consuming a very large quantity of food. The mode in
which the female deposits her eggs has been described by Riley,
[222] and is now widely known, his figures having been frequently
reproduced. The female has no elongate ovipositor, but possesses
instead some hard gonapophyses suitable for digging purposes; with
these she excavates a hole in the ground, and then deposits the
eggs, together with a quantity of fluid, in the hole. She prefers hard
and compact soil to that which is loose, and when the operation is
completed but little trace is left of it. The fluid deposited with the eggs
hardens and forms a protection to them, corresponding to the more
definite capsules of the cursorial Orthoptera.

The details of the process of oviposition and of the escape of the

young from their imprisonment are of much interest. According to
Künckel d'Herculais[223] the young Stauronotus maroccanus
escapes from the capsule by putting into action an ampulla formed
by the membrane between the head and the thorax; this ampulla is
supposed to be dilated by fluid from the body cavity, and is
maintained in the swollen condition by the Insect accumulating air in
the crop beneath it. In order to dislodge the lid of the capsule, six or
seven of the young ones inside combine their efforts to push it off by
means of their ampullae. The ampulla subsequently serves as a sort
of reservoir, by the aid of which the Insect can diminish other parts of
the body, and after emergence from the capsule, penetrate cracks in
the earth so as to reach the surface. Immediately after doing this the
young Stauronotus moults, the skin it casts being called by Künckel
an amnios. The cervical ampulla reappears at subsequent moults,
and enables the Insect to burst its skin and emerge from it.

The process is apparently different in Caloptenus spretus, which,

according to Riley, ruptures the egg-shell and works its way out by
the action of the spines at the apex of the tibiae. This latter Insect
when it emerges moults a pellicle, which Riley considers to be part of
the embryonic membranes.

Riley states that a female of Caloptenus spretus makes several egg-

masses. Its period of ovipositing extends over about 62 days, the
number of egg-masses being four and the total number of eggs
deposited about 100. The French naturalists have recently observed
a similar fact in Algeria, and have ascertained that one of the
migratory locusts—Schistocerca peregrina—may make a deposit of
eggs at more than one of the places it may alight on during its
migration.
It has been ascertained that the eggs of Acridiidae are very nutritious
and afford sustenance to a number of Insects, some of which indeed
appear to find in them their sole means of subsistence. Beetles of
the family Cantharidae frequent the localities where the eggs are laid
and deposit their eggs in the egg-masses of the Orthoptera, which
may thus be entirely devoured. Two-winged flies of the family
Bombyliidae also avail themselves of these eggs for food, and a mite
is said to be very destructive to them in North America. Besides
being thus destroyed in enormous quantities by Insects, they are
eaten by various birds and by some mammals.

Most of the Insects called locusts in popular language are members

of the family Acridiidae, of which there are in different parts of the
world very many species, probably 2000 being already known. To
only a few of these can the term Locust be correctly applied. A locust
is a species of grasshopper that occasionally increases greatly in
number, and that moves about in swarms to seek fresh food. There
are many Orthoptera that occasionally greatly increase in numbers,
and that then extend their usual area more or less; and some
Acridiidae multiply locally to a great extent—very often for one or two
seasons only,—and are then called locusts. The true migratory
locusts are species that have gregarious habits strongly developed,
and that move over considerable distances in swarms. Of these
there are but few species, although we hear of their swarms in many
parts of the world.

The migratory locusts do much more damage than the endemic

species. In countries that are liable to their visitations they have a
great influence on the prosperity of the inhabitants, for they appear
suddenly on a spot in huge swarms, which, in the space of a few
hours, clear off all the vegetable food that can be eaten, leaving no
green thing for beast or man. It is difficult for those who have not
witnessed a serious invasion to realise the magnitude of the event.
Large swarms consist of an almost incalculable number of
individuals. A writer in Nature[224] states that a flight of locusts that
passed over the Red Sea in November 1889 was 2000 square miles
in extent, and he estimated its weight at 42,850 millions of tons, each
locust weighing 1⁄16 of an ounce. A second similar, perhaps even
larger, flight was seen passing in the same direction the next day.
That such an estimate may be no exaggeration is rendered probable
by other testimony. From official accounts of locusts in Cyprus we
find that in 1881,[225] up to the end of October, 1,600,000,000 egg-
cases had been that season collected and destroyed, each case
containing a considerable number of eggs. By the end of the season
the weight of the eggs collected and made away with amounted to
over 1300 tons, and, notwithstanding this, no less than
5,076,000,000 egg-cases were, it is believed, deposited in the island
in 1883.

When we realise the enormous number of individuals of which a

large swarm of locusts may consist we can see that famine is only a
too probable sequence, and that pestilence may follow—as it often
has done—from the decomposition of the bodies of the dead Insects.
This latter result is said to have occurred on some occasions from
locusts flying in a mass into the sea, and their dead bodies being
afterwards washed ashore.

Locust swarms do not visit the districts that are subject to their
invasions every year, but, as a rule, only after intervals of a
considerable number of years. It has been satisfactorily ascertained
that in both Algeria and North America large swarms occur usually
only at considerable intervals. In North America Riley thought[226]
the average period was about eleven years. In Algeria the first
invasion that occurred after the occupation of the country by the
French was in 1845, the second in 1864, the third in 1866, since
which 1874 and 1891 have been years of invasion. These breaks
seem at first strange, for it would be supposed that as locusts have
great powers of increase, when once they were established in any
spot in large numbers, there would be a constant production of
superfluous individuals which would have to migrate as regularly as
is the case with swarms of bees. The irregularity seems to depend
on three facts: viz. that the increase of locusts is kept in check by
parasitic Insects; that the eggs may remain more than one year in
the ground and yet hatch out when a favourable season occurs; and
that the migratory instinct is only effective when great numbers of
superfluous individuals are produced.

It is not known that the parasites have any power of remaining in

abeyance as the locust eggs may do; and the bird destroyers of the
locusts may greatly diminish in numbers during a year when the
Insects are not numerous; so that a disproportion of numbers
between the locusts and their destroyers may arise, and for a time
the locusts may increase rapidly, while the parasites are much
inferior to them in numbers. If there should come a year when very
few of the locusts hatch, then the next year there will be very few
parasites, and if there should then be a large hatching of locusts
from eggs that have remained in abeyance, the parasites will not be
present in sufficient quantity to keep the destructive Insects in check;
consequently the next year the increase in number of the locusts
may be so great as to give rise to a swarm.

It is well established that locusts of the migratory species exist in

countries without giving rise to swarms, or causing any serious
injuries; thus Pachytylus cinerascens—perhaps the most important
of the migratory locusts—is always present in various localities in
Belgium, and does not give rise to swarms. When migration of
locusts does occur it is attended by remarkable manifestations of
instinct. Although several generations may elapse without a
migration, it is believed that the locusts when they migrate do so in
the direction taken by predecessors. Their movements are to a large
extent dependent on the wind, and it is said that they make trial
flights to ascertain its direction. When on the wing probably very little
muscular effort is necessary. Their bodies contain elastic air sacs in
communication with the tracheae, and at the time of flight it may be
presumed that the body is comparatively empty, food being wanting,
and the internal organs of reproduction, which occupy a large space
when in activity, yet undeveloped, hence the sacs have full room for
expansion, as explained on p. 283. Thus the Insects exert but little
effort in their aerial movements, and are, it is believed, chiefly borne
by the wind. Should this become unfavourable it is said that they
alight and wait for a change.

The most obscure point in the natural history of the migratory locusts
appears to be their disappearance from a spot they have invaded. A
swarm will alight on a locality, deposit there a number of eggs, and
then move on. But after a lapse of a season or two there will be few
or none of the species present in the spot invaded. This appears to
be partly due to the young locusts dying for want of food after
hatching; but in other cases they again migrate after growth to the
land of their ancestors. The latter fact is most remarkable, but it has
been ascertained by the U.S. Entomological Commission that these
return swarms do occur.

Fig. 176.—Portions of body of Caloptenus spretus to show some of the

air-sacs. (Modified from Packard.) A, Dorsal aspect of anterior
parts; B, lateral aspect of posterior parts of body; a, enlargements
of tracheae in head; b, pair of large sacs in thorax; c, sacs on the
tracheal trunks of abdomen; s, spiracles.

In South Africa it would appear that the movements of the migratory

locusts are frequently made before the Insects have acquired their
wings. Mrs. Barber, in an account of "Locusts and Locust-Birds in
South Africa,"[227] has illustrated many points in the Natural History
of these Insects. The South African species manifests the gregarious
and migratory disposition when the individuals are quite young, so
that they travel in flocks on foot, and are called by the Dutch
"Voetgangers." After hatching, the various families of young
amalgamate, so that enormous numbers come together. Having
denuded the neighbourhood of all its food-supplies, they move off in
search of fresh crops and pastures new. They take advantage of
roads, and sometimes a good many miles will be traversed in a day;
they proceed by means of short leaps, rapidly repeated. When the
"Voetgangers" are thus returning northwards towards the lands in the
interior from which their progenitors departed, no obstacles can stay
their course. Forests or rivers may intervene, diverting them for a
while from their line of march, but they succeed ultimately in
continuing their journey to the interior.

The manner in which these wingless locusts occasionally cross

broad rivers is interesting, as it has some bearing on the difficult
question of the possibility of winged locusts crossing seas of
considerable width. Mrs. Barber refers to an instance that took place
on the Vaal River in the spring of the year 1871, shortly after the
discovery of the Diamond-fields. The country was at that time
swarming with young locusts; every blade of grass was cleared off
by them. One day a vast swarm of the "Voetgangers" made their
appearance on the banks of the Vaal River; they appeared to be in
search of a spot for crossing, which they could not find, the river
being somewhat swollen. For several days the locusts travelled up
the stream; in the course of doing this they paused for some time at
an abrupt bend in the river where a number of rocks were cropping
out, as if in doubt whether to attempt a passage at this place. They,
however, passed on, as if with the hope of finding a better ford; in
this apparently they were disappointed, for three days afterwards
they returned to the same bend of the river, and there plunged in
vast multitudes into the stream, where, assisted by a favourable
current and the sedges and water-plants which grew upon the
projecting rocks, they managed to effect a crossing, though great
numbers were drowned and carried away by the flooded river. Mrs.
Barber adds that "Voetgangers" have been known to attempt the
passage of the Orange River when it was several hundred yards in
breadth, pouring their vast swarms into the flooded stream
regardless of the consequences, until they became heaped upon
each other in large bodies. As the living mass in the water
accumulated, some portions of it were swept away by the strong
current from the bank to which they were clinging, and as the living
locusts tightly grasped each other and held together, they became
floating islands, the individuals continually hopping and creeping
over each other as they drifted away. Whether any of the locust-
islands succeeded in reaching the opposite bank is unknown;
probably some of them were drifted on land again. They are by no
means rapid swimmers; they do not perish easily in the water when
in masses, their habit of continually changing places and hopping
and creeping round and round upon each other being very
advantageous as a means of preservation. It is a common practice
for the young locusts to form a bridge over a moderately broad
stream by plunging indiscriminately into it and holding on to each
other, grappling like drowning men at sticks or straws, or, in fact,
anything that comes within their reach, and that will assist in floating
them; meanwhile those from behind are eagerly pushing forward
over the bodies of those that are already in the stream and hurrying
on to the front, until at length by this process they reach the opposite
bank of the river; thus a floating mass of living locusts is stretched
across the stream, forming a bridge over which the whole swarm
passes. In this manner few, comparatively speaking, are drowned,
because the same individuals do not remain in the water during the
whole of the time occupied by the swarm in crossing, the Insects
continually changing places with each other; those that are beneath
are endeavouring to reach the surface by climbing over others, whilst
those above them are, in their turn, being forced below. Locusts are
exceedingly tenacious of life, remaining under water for a
considerable time without injury. An apparently drowned locust will
revive beneath the warm rays of the sun, if by chance it reaches the
bank or is cast on shore. Mrs. Barber relates an interesting case
where the instinct of the "Voetgangers" was at fault, they plunging
into a river from a steep sandy bank, only to find another similar
sandy precipice on the other side. On this they could gain no footing,
and all perished in the stream, where they putrefied, and caused the
death of the fish, which floated likewise on the surface; so powerful
were the effluvia produced that no one was able to approach the
river.
Locusts are able to travel considerable distances, though how far is
quite uncertain. Accounts vary as to their moving by night. It has,
however, been recently proved that they do travel at night, but it is
not ascertained how long they can remain in the air without
descending. The ocean is undoubtedly a source of destruction to
many swarms; nevertheless, they traverse seas of considerable
width. They have been known to reach the Balearic Islands, and
Scudder gives[228] a well-authenticated case of the occurrence of a
swarm at sea. On the 2nd of November 1865 a ship on the voyage
from Bordeaux to Boston, when 1200 miles from the nearest land,
was invaded by a swarm of locusts, the air and the sails of the ship
being filled with them for two days. The species proved to be
Acridium (Schistocerca) peregrinum. This is an extraordinary case,
for locusts do not fly with rapidity, being, indeed, as we have
remarked, chiefly carried by the wind. Possibly some species may
occasionally rest on the water at night, proceeding somewhat after
the fashion of the "Voetgangers" when passing over rivers as
described by Mrs. Barber. In Sir Hans Sloane's history of Jamaica an
account of an occurrence of this kind is given on the authority of
Colonel Needham, who states that in 1649 locusts devastated the
island of Tenerife, that they were seen to come from Africa when the
wind was blowing thence, that they flew as far as they could, then
alighted on the water, one on the other, till they made a heap as big
as the greatest ship, and that the next day, being refreshed by the
sun, they took flight again and landed in clouds at Tenerife. De
Saussure says[229] that the great oceans are, as a rule, impassable
barriers, and that not a species of the tribe Oedipodides has passed
from the Old World to the New. It is, however, possible that Acridium
peregrinum, of the tribe Acridiides, may have originally been an
inhabitant of America, and have passed from thence to the Old
World.
 Fig. 177.—European migratory locust, Pachytylus cinerascens ♀.

The species of Acridiidae that have been ascertained to be migratory

are not numerous.[230] The most abundant and widely distributed of
them is Pachytylus cinerascens (Fig. 177), which has invaded a
large part of the Eastern hemisphere, extending from the Atlantic
Ocean to China. It exists in numerous spots in the Oriental region
and the Asiatic Archipelago, and even in New Zealand. It is the
commoner of the locusts of Europe. Its congener, P. migratorius, is
much less widely distributed, its migrations being, according to de
Saussure, limited to Turkestan and Eastern Europe. A third species,
P. migratorioides, inhabits Eastern Africa, and a variety of it is the
"Yolala" or locust of Madagascar. Mr. Distant has informed the writer
that this migratory locust is found in South Africa. P. (Oedaleus)
marmoratus has almost as wide a distribution in the Eastern
hemisphere as P. cinerascens, except that it is more exclusively
tropical; it is thus excluded from New Zealand. P. (Oedaleus)
nigrofasciatus has a more northern distribution than its congener, but
has extended to Africa and the Asiatic Archipelago. This Insect is so
variable that the distinctions of its races from other species of the
same genus are not yet clear. All the above-mentioned locusts
belong to the tribe Oedipodides. Acridium peregrinum, now more
frequently called Schistocerca peregrina, belongs to the tribe
Acridiides. It is a large locust (Fig. 84), and has a wide distribution. It
is the chief species in North Africa, and is probably the locust of the
plagues of Egypt mentioned in the book of Exodus. It is also,
according to Cotes,[231] the chief locust of North-West India. In this
latter country Pachytylus cinerascens and some other species also
occur. With the exception of S. peregrina, the species of the genus
Schistocerca are confined to the New World. In North America
locusts are more usually called grasshoppers. Several species of the
genus Caloptenus are injurious in that country, but the chief
migratory species is C. spretus (Fig. 175). This genus belongs to
Acridiides. A large locust, Schistocerca americana, is also migratory
to a small extent in the United States. In South America other
species of Schistocerca are migratory; it is not known how many
there may be, and it is possible that one or more may prove to be the
S. peregrina of the Old World. A Chilian species, according to Mr. E.
C. Reed,[232] exhibits distinctions of colour similar to those that have
been observed in S. peregrina in Algeria.

Fig. 178.—Cephalocoema lineata, female, × ⅔. S. America. (After

Brunner.)

In Britain we are now exempt from the ravages of locusts, though

swarms are said to have visited England in 1693 and 1748.
Individuals of the migratory species are, however, still occasionally
met with in England and the south of Scotland. P. cinerascens has
been recorded from Kerry in Ireland, but erroneously, the Insect
found being Mecostethus grossus (Fig. 173). According to Miss
Ormerod,[233] large locusts are imported to this country in fodder in
considerable numbers, but are usually dead; living individuals are,
however, sometimes found among the others. In 1869 living
specimens of Schistocerca peregrina were found in various parts of
the country, having, in all probability, arrived here by crossing the
German Ocean. Pachytylus cinerascens has also, it is believed,
occurred here, the specimens that have been recorded at different
times under the name of P. migratorius being more probably the
former species.
Although the majority of the very large number of species included in
Acridiidae are recognised with ease from their family likeness as
belonging to the group, yet there are others that present an unusual
aspect. This is specially the case with the members of the small
tribes Tettigides, Proscopides, and Pneumorides, and with some of
the apterous forms of the Oedipodides. The tribe Proscopides (Fig.
178, Cephalocoema lineata, female) includes some of the most
curious of the Acridiidae. Breitenbach gives[234] a brief account of
the habits of certain species which he met with near Porto Alegre in
South America. On a stony hill there was some grass which, by
several months' exposure to the sun's rays, had become withered
and brown. Apparently no live thing was to seen on this hillock
except the ubiquitous ants, but after a while he noticed some
"lightning-like" movements, which he found were due to specimens
of Proscopia. The Insects exactly resemble the withered vegetation
amongst which they sit, and when alarmed seek safety with a
lengthy and most rapid leap. When attention was thus directed to
them he found the Insects were really abundant, and was often able
to secure fifty specimens on a single afternoon. These Insects bear a
great general resemblance to the Phasmides, but there is no
evidence at present to show that the two kinds of Insects live in
company, as is the case with so many of the Insects that resemble
one another in appearance. Although the linear form and the
elongation of the body are common to the stick-Insects and the
Proscopides, yet this structure is due to the growth of different parts
in the two families. In the Phasmidae the prothorax is small, the
mesothorax elongate, while in the Proscopides the reverse is the
case. The elongation of the head is very curious in these Insects; the
mouth is not thus brought any nearer to the front, but is placed on
the under side of the head, quite close to the thorax. The tribe
Tryxalides contains Insects (Fig. 165) that approach the Proscopides
in the form of the head and other characters. In most cases the
sexes of the Proscopides differ from one another so strongly that it is
difficult to recognise them as being of the same species. Usually
both sexes are entirely apterous, but the Chilian genus Astroma
exhibits a remarkable exception and an almost unique condition of
the alar organs, the mesonotum being in each sex entirely destitute
of such appendages, while the female has on the metanotum
rudiments of wings which are absent in the male.

Fig. 179.—Tettix bipunctatus. Britain. A, The Insect magnified; B, part

of the middle of the body; a, prolongation of pronotum; b, tegmen;
c, wing.

The tribe Tettigides is a very extensive group of small Acridiidae, in

which the pronotum extends backwards as a hood and covers the
body, the tegmina and wings being more or less modified. In our
British species (Fig. 179) this condition does not greatly modify the
appearance of the Insect, but in many exotic species (Fig. 180) the
hood assumes remarkable developments, so that the Insects have
no longer the appearance of Orthoptera. It would be impossible,
without the aid of many figures, to give an idea of the variety of forms
assumed by this prothoracic expansion. It is a repetition of what
occurs in the Order Hemiptera, where the prothoracic hoods of the
Membracides exhibit a similar, though even more extraordinary,
series of monstrous forms. So great is the general similarity of the
two groups that when the genus Xerophyllum (Fig. 180, A) was for
the first time described, it was treated by the describer as being a
bug instead of a grasshopper. This genus includes several species
from Africa. The curious Cladonotus (Fig. 180, B) is a native of
Ceylon, where it is said to live in sandy meadows, after the fashion
of our indigenous species of Tettix (Fig. 179). Very little is known as
to the habits of these curious Tettigides, but it has been ascertained
that some of the genus Scelimena are amphibious, and do not
hesitate to enter the water and swim about there; indeed it is said
that they prefer plants growing under water as food. This habit has
been observed both in Ceylon and the Himalayas. The species are
said to have the hind legs provided with dilated foliaceous
appendages useful for swimming.
 Fig. 180.—Tettigides: A, Xerophyllum simile Fairm.; B, Cladonotus
humbertianus. (After Bolivar.)

Fig. 181.—A, Mastax (Erianthus) guttatus, male. Sumatra. (After

Westwood.) B, profile; C, front of head.

The tribe Mastacides includes thirty or forty species of Acridiidae

with short antennae and vertical head (Fig. 181, Mastax guttatus);
they are apparently all rare and little known, but are widely
distributed in the tropics of the Old and New Worlds. Nothing
whatever seems to be known of their habits or of their development.

The tribe Pneumorides includes a still smaller number of species of

very aberrant and remarkable grasshoppers, of large size, with short
antennae, and with the pronotum prolonged and hood-like; they are
peculiar to South Africa. Although amongst the most remarkable of
Insects, we are not able to give any information as to their habits. It
would appear from the form of their legs that they have but little
power of hopping. The species of which we figure the female (Fig.
182) is very remarkable from the difference in colour of the sexes.
The female is so extravagantly coloured that she has been said to
look as if "got up" for a fancy-dress ball. She is of a gay green, with
pearly white marks, each of which is surrounded by an edging of
magenta; the white marks are very numerous, especially on the
parts of the body not shown in our figure; the face has magenta
patches and a large number of tiny pearly-white tubercles, each of
which, when placed on a green part, is surrounded by a little ring of
mauve colour. Though the female is certainly one of the most
remarkably coloured of Insects, her consort is of a modest, almost
unadorned green colour, and is considerably different in form. He is,
however, provided with a musical apparatus, which it is possible may
be a means of pleasing his gorgeous but dumb spouse. It consists of
a series of ridges placed on each side of the inflated abdomen,
which, as we have previously (p. 200) remarked, has every
appearance of being inflated with the result of improving its
resonance.

Fig. 182.—Pneumora scutellaris, female. South Africa.

The Pyrgomorphides[235] is a small tribe of about 120 described

species, two of which are found in the south of Europe (Fig. 183,
Pyrgomorpha grylloides). The tribe includes a number of large and
curious Insects, among them the species of Phymateus and Petasia,
with peculiar excrescences on the pronotum and vivid colours on
some parts of the body or its appendages, which are apparently
common Insects in South Africa.

The tribe Tryxalides includes a great many species of grasshoppers.

In them the front of the head joins the upper part at an acute angle
(Figs. 165 and 173). This tribe and the Acridiides are the most
numerous in species of the family. To the latter belong most of the
migratory locusts of the New World (Fig. 175, Caloptenus spretus). A
Spanish species of this tribe, Euprepocnemis plorans, though
provided with well-developed wings, possesses the remarkable habit
of seeking shelter by jumping into the water and attaching itself
below the surface to the stems of plants.

Fig. 183.—Pyrgomorpha grylloides. South Europe. (After Fischer.)

Fig. 184.—Xiphocera (Hoplolopha) asina. S. Africa. (After de

Saussure.)

The tribe Pamphagides[236] includes some 200 species, found

chiefly in Africa and the arid regions near the Mediterranean Sea.
They are mostly apterous forms, and this circumstance has,
according to de Saussure, exercised a marked influence on the
geographical distribution of the species. Although the tribe consists
chiefly of apterous forms, several species possess well-developed
wings; sometimes this is the case of the male but not of the female.
Some of the species are highly modified for a desert life, and exhibit
a great variation in the colour of the individuals in conformity with the
tint of the soil they inhabit. Xiphocera asina (Fig. 184) is thought by
Péringuey to be the prey of the extraordinary South African tiger-
beetles of the genus Manticora.

We have already mentioned the tribe Oedipodides[237] as including

most of the species of migratory locusts of the Old World. Some
striking cases of variation in colour occur amongst the winged
Oedipodides. In certain species the hind wings may be either blue or
rosaceous in colour; it is thought that the latter is the tint natural in
the species, and that it is due to the mixture of a red pigment with the
pale blue colour of the wing; hence the blue-coloured wings are
analogous to cases of albinism. But the most remarkable fact is that
this colour difference is correlative with locality. Brunner von
Wattenwyl says[238] that the blue variety of Oe. variabilis occurs only
in a few localities in Europe—he mentions Vienna and Sarepta,—
and that where it occurs not a single red example can be met with.
Similar phenomena occur in other species in both Europe and North
America, and L. Bruner has suggested[239] that the phenomena in
the latter country are correlative with climatic conditions.

The group Eremobiens, a subdivision of Oedipodides, includes some

of the most interesting forms of Acridiidae. Its members have several
modes of stridulation. Cuculligera flexuosa and other of the winged
forms, according to Pantel,[240] produce sounds by the friction of the
middle tibia against the wing, both of these parts being specially
modified for the purpose in the male sex. The most peculiar
members of the Eremobiens are some very large Insects, modified
to an extraordinary extent for a sedentary life in deserts and arid
places. Trimen says[241] that a South African species, Trachypetra
bufo, which lives amongst stones, is so coloured that he had much
difficulty in detecting it, and that he noticed in certain spots, often
only a few square yards in extent, where the stones lying on the
ground were darker, lighter, or more mottled than usual, that the
individuals of the grasshopper were of a similar colour to the stones.
 Fig. 185.—Methone anderssoni, female. S. Africa. a, Front of head; b,
posterior leg; c, d, front and hind feet. (c and d magnified, the
others natural size.)

The Insect referred to by Trimen is, we believe, the Batrachotettix

whiti of de Saussure. In this species the alar organs are completely
absent, and the pronotum forms a sort of hood that protects the base
of the hind body. Some of the desert Eremobiens vary so much that
the differences found among individuals of the same species are
said by Brunner and de Saussure to be so great as to affect even the
generic characters, and give rise to the idea of an "uncompleted
species-formation."

Fig. 186.—Portions of middle of the body and hind leg of Methone

anderssoni ♂ : a, femur; b, an inferior fold; c, rattling-plate; d,
striated surface; e, the adjoining sculpture; f, grooved portion of
tegmen. The part e is really, like d, a portion of the second
 abdominal segment, not of the third, as might be supposed from
the figure.

Methone anderssoni, an inhabitant of the Karoo Desert of South

Africa, is one of the largest of the Acridiidae. A female of this species
is represented of the natural size in Fig. 185. This Insect is
remarkable on account of the complex organs for producing sound,
and for the great modification of the posterior legs (Fig. 185, b),
which do not possess locomotive functions, but serve as a portion of
the sound-producing apparatus, and as organs for protecting the
sides of the body. This Insect is said to be very efficient in making a
noise. The sexes differ considerably in their sound-producing
organs, a portion of which are present in the female as well as in the
male (Fig. 186). Connected with the first abdominal segment, but
extending backwards on the second, there is a peculiar swelling
bearing two or three strongly raised chitinous folds (Fig. 186, c).
When the leg is rotated these folds are struck by some peg-like
projections situate on the inner face of the base of the femur, and a
considerable noise is thus produced. The pegs cannot be seen in
our figure. This apparatus is equally well developed in female and
male. On the second abdominal segment, immediately behind the
creaking folds we have described, there is a prominent area, densely
and finely striated (Fig. 186, d): this is rubbed by some fine asperities
on the inner part of the femur near its base. Sound is produced by
this friction on the striated surface, the sculpture of which is abruptly
contrasted with that of the contiguous parts: these structures seem
to be somewhat better developed in the male than they are in the
female, and to be phonetic, at any rate in the former sex. The male
has the rudimentary tegmina (Fig. 186, f) much longer than they are
in the female (Fig. 185), and their prolonged part is deeply grooved,
so as to give rise to strong ridges, over which plays the edge of the
denticulate and serrate femur. There is nothing to correspond to this
in the female, and friction over the surface of this part of the male
produces a different and louder sound. There can be little doubt that
this is a phonetic structure peculiar to the male. It approximates in
situation to the sound-producing apparatus of the males of the
Stenobothri and other Acridiidae. Methone anderssoni has large