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 Fractional Integral
Transforms

Fractional Integral Transforms: Theory and Applications presents over twenty-five

integral transforms, many of which have never before been collected in one single vol-
ume. Some transforms are classic, such as the Laplace, Fourier, etc, and some are more
recent, such as the Fractional Fourier, Gyrator, Linear Canonical, Special Affine Fourier
Transforms, as well as, continuous Wavelet, Ridgelet, and Shearlet transforms.

The book provides an overview of the theory of fractional integral transforms with ex-
amples of such transforms, before delving deeper into the study of important fractional
transforms, including the fractional Fourier transform. Applications of fractional inte-
gral transforms in signal processing and optics are highlighted. The book’s format has
been designed to make it easy for the readers to extract the essential information they
need to learn about the fundamental properties of each transform. Supporting proof
and explanations are given throughout.

Features
•  Brings together integral transforms never before collected into a single volume
•  A useful resource on fractional integral transforms for researchers and graduate stu-
dents in mathematical analysis, applied mathematics, physics and engineering
•  Written in an accessible style with detailed proofs and emphasis on providing the
reader with an easy access to the essential properties of important fractional integral
transforms
Ahmed I. Zayed is a Professor of Mathematics at the Department of Mathematical Sci-
ences, DePaul University, Chicago, and was the Chair of the department for 20 years,
from 2001 until 2021. His research interests varied over the years starting with general-
ized functions and distributions to sampling theory, applied harmonic analysis, special
functions and integral transforms. He has published two books and edited seven re-
search monographs. He has written 22 book chapters, published 118 research articles,
and reviewed 173 publications for the Mathematical Review and 81 for the Zentralb-
latt für Mathematik (zbMath). He has served on the Editorial Boards of 22 scientific
research journals and has refereed over 200 research papers submitted to prestigious
journals, among them are IEEE, SIAM, Amer. Math. Soc., Math Physics, and Optical
Soc. Journals.
Taylor & Francis
Taylor & Francis Group
http://taylorandfrancis.com
Fractional Integral
Transforms
Theory and Applications

Ahmed I. Zayed
DePaul University, USA
Designed cover image: Ahmed I. Zayed
First edition published 2024
by CRC Press
6000 Broken Sound Parkway NW, Suite 300, Boca Raton, FL 33487-2742

and by CRC Press

4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN

CRC Press is an imprint of Taylor & Francis Group, LLC

© 2024 Ahmed I. Zayed

Reasonable efforts have been made to publish reliable data and information, but the author and publisher
cannot assume responsibility for the validity of all materials or the consequences of their use. The authors
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Library of Congress Cataloging-in-Publication Data

Names: Zayed, Ahmed I., author.

Title: Fractional integral transforms : theory and applications / Ahmed I.
Zayed, DePaul University, USA.
Description: First edition. | Boca Raton, FL : C&H/CRC Press, 2024. |
Includes bibliographical references and index.
Identifiers: LCCN 2023046154 (print) | LCCN 2023046155 (ebook) | ISBN
9780367543877 (hardback) | ISBN 9781003089353 (paperback) | ISBN
9780367544485 (ebook)
Subjects: LCSH: Integral transforms. | Transformations (Mathematics)
Classification: LCC QA432 .Z39 2024 (print) | LCC QA432 (ebook) | DDC
519.2/3--dc23/eng/20231031
LC record available at https://lccn.loc.gov/2023046154
LC ebook record available at https://lccn.loc.gov/2023046155

ISBN: 978-0-367-54387-7 (hbk)

ISBN: 978-0-367-54448-5 (pbk)
ISBN: 978-1-003-08935-3 (ebk)

DOI: 10.1201/9781003089353

Typeset in TeXGyreTermes-Regular font

by KnowledgeWorks Global Ltd.

Publisher’s note: This book has been prepared from camera-ready copy provided by the authors
 Dedication

To

My wife Elena and my daughter Nora for their love and support
Taylor & Francis
Taylor & Francis Group
http://taylorandfrancis.com
Contents

Preface xiii

Chapter 1  Introduction and Preliminaries 1

1.1 NOTATION 1
1.2 SPECIAL FUNCTIONS AND ORTHOGONAL POLYNOMIALS 4
1.2.1 The Gamma Function 4
1.2.2 The Beta Function 5
1.2.3 The Hermite Polynomials Hn (x) 6
1.2.4 The Laguerre Polynomials Lαn (x)(α > – 1) 7
(α,β)
1.2.5 The Jacobi Polynomials Pn (x)(α, β > – 1) 8
1.2.6 The Bessel Functions 9
1.2.7 The Mittag–Leffler function 11
1.2.8 The Hypergeometric and q-Hypergeometric Functions 12
1.3 NON-ORTHOGONAL BASES AND FRAMES IN A HILBERT SPACE 13
1.3.1 Non-orthogonal Bases and Frames 13
1.3.2 Reproducing-Kernel Hilbert Spaces 16
1.4 SHIFT-INVARIANT SPACES 17
1.5 GENERALIZED FUNCTIONS AND DISTRIBUTIONS 19
1.5.1 Testing-Function Spaces and Their Duals 19
1.5.2 Spaces of Generalized Functions 20
1.5.3 A Special Type of Generalized Functions 21
1.6 SAMPLING AND THE PALEY-WIENER SPACE 22
1.7 POISSON SUMMATION FORMULA 26
1.8 UNCRTAINTY PRINCIPLE 27

Chapter 2  Integral Transformations 28

2.1 INTRODUCTION AND BRIEF HISTORY 28

2.2 WHAT IS AN INTEGRAL TRANSFORM? 29

vii
viii  Contents

2.3 EXAMPLES OF INTEGRAL TRANSFORMS 29

2.3.1 One-Dimensional Integral Transforms 29
2.3.2 Higher Dimensional Transforms 33
2.3.3 Special Cases of Higher Dimensional Transforms 34
2.4 GENERAL PROPERTIES OF INTEGRAL TRANSFORMATIONS 36
2.5 WHY INTEGRAL TRANSFORMS? 39

Chapter 3  Fractional Integral Transforms 41

3.1 INTRODUCTION 41
3.2 PRELUDE TO FRACTIONAL INTEGRAL TRANSFORMS 45
3.2.1 The Fractional Fourier Transform 45
3.2.2 The Fractional Hankel Transform 48
3.3 GENERAL CONSTRUCTION OF FRACTIONAL INTEGRAL
TRANSFORMS 50
3.3.1 Examples of the General Construction 52
3.3.2 Fractional Integral Transforms Associated With the Jacobi
Polynomials 56
3.4 FRACTIONAL DERIVATIVES AND INTEGRALS VERSUS
FRACTIONAL INTEGRAL TRANSFORMS 58
3.5 OTHER FRACTIONAL INTEGRAL TRANSFORMS 61

Chapter 4  The Fractional Fourier Transform (FrFT) 62

4.1 HISTORICAL OVERVIEW 62

4.2 PRELIMINARIES 64
4.3 OPERATIONAL CALCULUS 68
4.3.1 Convolution Theorem 75
4.3.2 Poisson Summation Formula for the Fractional Fourier Transform 79
4.3.3 Sampling Theorem for the Fractional Fourier Transform 80
4.3.4 The Wigner Distribution 82
4.4 THE FRACTIONAL HILBERT TRANSFORM 85
4.5 FRACTIONAL TIME-FREQUENCY REPRESENTATIONS 88
4.5.1 Fractional Wigner Distributions 89
4.5.2 Fractional Time and Frequency Shifts 92
4.5.3 The Fractional Cross-Ambiguity Function 93
4.5.4 Fractional Windowed (Sliding-Window)-Fourier Transform 98
 Contents  ix

4.6 UNCERTAINTY PRINCIPLE FOR THE FRACTIONAL FOURIER

TRANSFORM 99
4.7 FRACTIONAL FOURIER TRANSFORM OF GENERALIZED
FUNCTIONS 100
4.7.1 The Embedding Method 101
4.7.2 The Space of Boehmians 102
4.7.3 The Algebraic Method 104
4.8 APPLICATIONS OF THE FRACTIONAL FOURIER TRANSFORM 106

Chapter 5  Shift-Invariant and Sampling Spaces of the Fractional

Fourier Transform 108

5.1 INTRODUCTION 108

5.2 BASIC DEFINITIONS 108
5.3 DISCRETE FRACTIONAL FOURIER TRANSFORM AND
CONVOLUTION 113
5.3.1 Discrete Fractional Fourier Transform 113
5.3.2 Fractional Convolution 114
5.4 SHIFT-INVARIANCE IN THE FRFT DOMAIN 115
5.5 THE FRACTIONAL ZAK TRANSFORM 119
5.6 APPLICATIONS: FRACTIONAL DELAY FILTERING 121

Chapter 6  Two-Dimensional Coupled Fractional Fourier Transform

(CFrFT) 122

6.1 INTRODUCTION 122

6.2 FRACTIONAL FOURIER TRANSFORM IN HIGHER DIMENSIONS 124
6.2.1 The Direct Product Representation 124
6.2.2 Metaplectic Representation 125
6.3 THE TWO-DIMENSIONAL FRACTIONAL FOURIER TRANSFORM 126
6.3.1 Complex Hermite Polynomials 126
6.3.2 Integral Representation of the Two-Dimensional Fractional
Fourier Transform 128
6.3.3 Inversion Formula 129
6.3.4 Examples 131
6.4 ADDITIVE PROPERTY 132
6.5 CONVOLUTION THEOREM 137
6.6 POISSON SUMMATION FORMULA 138
x  Contents

6.7 A SPACE OF BANDLIMITED SIGNALS AND ITS SAMPLING

THEOREM 142
6.7.1 Space of Bandlimited Signals 142
6.7.2 Sampling Theorems 143
6.7.3 Examples and Sampling Points Configuration 146
6.8 THE COUPLED FRACTIONAL FOURIER TRANSFORM OF
GENERALIZED FUNCTIONS 148
6.9 THE GYRATOR TRANSFORM 152
6.9.1 Motivation and Definitions 152
6.9.2 Elementary Properties of the Gyrator Transform 155

Chapter 7  The Two-Dimensional Fractional Fourier Transform and

The Wigner Distribution 160

7.1 INTRODUCTION 160

7.2 THE WIGNER DISTRIBUTION 160
7.3 FOUR-DIMENSIONAL ROTATIONS 161
7.4 THE FOUR-DIMENSIONAL WIGNER DISTRIBUTION 163
7.5 THE FOUR-DIMENSIONAL WIGNER DISTRIBUTION AND THE
COUPLED FRACTIONAL FOURIER TRANSFORM 165

Chapter 8  Short-Time Coupled Fractional Fourier Transform and

Uncertainty Relations 172

8.1 INTRODUCTION AND NOTATION 172

8.2 PROPERTIES OF THE COUPLED FRACTIONAL FOURIER
TRANSFORM 173
8.3 CONVOLUTION AND EXTENSION OF THE COUPLED
FRACTIONAL FOURIER TRANSFORM 176
8.4 SHORT-TIME COUPLED FRACTIONAL FOURIER TRANSFORM 178
8.5 PROPERTIES OF THE SHORT-TIME COUPLED FRACTIONAL
FOURIER TRANSFORM 180
8.6 UNCERTAINTY PRINCIPLE 182

Chapter 9  The Linear Canonical Transform (LCT) 184

9.1 INTRODUCTION AND HISTORICAL OVERVIEW 184

9.2 DEFINITIONS AND SPECIAL CASES OF THE LINEAR CANONICAL
TRANSFORM 184
 Contents  xi

9.3 PROPERTIES OF THE LINEAR CANONICAL TRANSFORM 187

9.3.1 Basic Properties 187
9.3.2 Convolution Theorems 190
9.3.3 Additive Property of the Linear Canonical Transform 191
9.3.4 Sampling Theorem 193
9.3.5 Eigenfunctions and Eigenvalues 195
9.4 THE METAPLECTIC REPRESENTATION AND CONVOLUTION 197
9.5 ELEMENTARY PROPERTIES OF THE METAPLECTIC
TRANSFORMATIONS 199
9.6 TWO-DIMENSIONAL SAMPLING THEOREM FOR THE LINEAR
CANONICAL TRANSFORM 201
9.6.1 Two-Dimensional LCT in Polar Coordinates 201
9.6.2 Sampling Theorem for LCT 203

Chapter 10  The Special Affine Fourier Transform (SAFT) 209

10.1 INTRODUCTION AND HISTORICAL REMARKS 209

10.2 DEFINITIONS 209
10.3 THE OFFSET LINEAR CANONICAL TRANSFORM 210
10.4 ELEMENTARY PROPERTIES OF THE SPECIAL AFFINE FOURIER
TRANSFORM 213
10.5 POISSON SUMMATION FORMULA FOR SAFT 215
10.6 CONVOLUTION AND PRODUCT THEOREMS FOR SPECIAL
AFFINE FOURIER TRANSFORM 218
10.6.1 Modulation and Convolution Operations 218
10.6.2 Convolution Theorem 218
10.6.3 Product Theorem 220
10.7 SHIFT-INVARIANT SPACES FOR THE SPECIAL AFFINE FOURIER
TRANSFORM 221
10.7.1 Preliminaries 221
10.7.2 Discrete Special Affine Fourier Transform 222
10.7.3 Riesz Basis for Shift-Invariant Spaces in the SAFT Domain 223
10.8 ZAK TRANSFORM ASSOCIATED WITH THE SAFT 225
10.9 SHANNON’S SAMPLING THEOREM AND THE SAFT:
REINTERPRETATION, EXTENSION AND APPLICATIONS 228

Appendix 233

Bibliography 235

Index 261
Taylor & Francis
Taylor & Francis Group
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Preface

Integral transforms have existed for two centuries and their historical roots go back to the
work of the French mathematicians Pierre Simon Laplace and Joseph Fourier whose work
led to the formation of the two seminal integral transforms that were later named after them:
the Laplace and Fourier transforms. Integral transforms have proved to be a useful tool in
solving many problems in mathematics, applied mathematics, physics and engineering.
Hundreds, if not thousands, of articles and books have been written about them.
Although the subject of integral transforms is considered classic with a long history,
it has not ceased to grow. Many novel transforms with a plethora of practical applications
have been introduced in the last three decades, such as the linear canonical transforms,
wavelets and shearlets transforms.
In 1980 an off-shoot of the subject of integral transforms started to emerge in the work
of Victor Namias who introduced a new integral transform which he called fractional order
Fourier transform and which he used to solve problems in quantum mechanics. Namias’s
idea of a fractional Fourier transform had appeared implicitly in earlier work by J. Wiener
in 1927 and E. U. Condon in 1937, but it was Namias who developed it explicitly and was
the first to use the phrase fractional transform.
At the beginning, Namias’s work received little attention and the one that was received
was mainly from mathematicians, such as A. C. McBride, F. H. Kerr and D. Mustard.
Almost a decade later, at the beginning of the 1990s, the subject started to take shape and
receive more attention when engineers and physicists found numerous practical applications
of Namias’s transform in signal processing and optics. Some of the pioneers in this field
were L. B. Almeida, M. Kutay, A. Lohmann, D. Mendlovic, H. M. Ozaktas and Z. Zalevsky,
just to mention a few.
After the publication of his paper on the fractional Fourier transform, Namias extended
his work to another integral transform, namely the Hankel transform, and he called the new
transform fractional Hankel transform. Namias’s work opened a window on new possibilities
of extending the fractional transform idea to other integral transforms. Currently there is
a slew of fractional integral transforms, such as fractional Hilbert, fractional Stockwell,
fractional wavelets, fractional Radon transforms, etc. It is no exaggeration to say that
nowadays, practically every integral transform has a fractional version. However, so far
most of them have not shown to render any useful practical applications to compete with
the fractional Fourier transform.
The notion of fractions existed in mathematics for thousands of years as part of the
counting systems used by the Ancient Egyptians, Babylonians and Indian Mayans. The term
fractional appeared in classical mathematical analysis in the work of Riemann on fractional
derivatives and fractional integrals. There is a connection between fractional derivatives

xiii
xiv  Preface

and fractional integrals on the one hand and fractional integral transforms on the other
hand, which will be discussed in Chapter 3 of this volume.
The fractional Fourier transform has distinguished itself among other fractional integral
transforms of being a special case of a more general class of integral transforms that arose
in quantum mechanics, time-frequency representations and abstract harmonic analysis. The
fractional Fourier transform is a special case of a class of integral transforms known as the
Linear Canonical Transforms which may be viewed as a group of unitary transformations
acting on the Hilbert space of all square-integrable functions on the real line L2 (R). Their
action on that Hilbert space is represented by the Metaplectic group.
The idea of this book came to me when some of my colleagues and students, who were
interested in learning the rudiments of the subject, asked me for references. I then realized
that although there are many books in the literature on integral transforms, there is a very
limited number on fractional integral transforms. The first and most noticeable one is the
book by Ozaktas, Zalevsky and Kutay, The fractional Fourier transform with Applications
in Optics and Signal Processing, (2001) which is more than twenty years old. Since the
publication of that book, new results and advances in the field have taken place which
deserved to be compiled in a publication. Another point of departure from the former book
is that the former focused mainly on the fractional Fourier transform and its applications,
while this one deals with the general topic of fractional integral transforms and then delves
into the study of important fractional transforms, including the fractional Fourier transform.
The book is designed to give the reader an overview of the subject from its infancy until
the present state of affairs. It is written at a level that a graduate student in mathematics
will find accessible. The book is not meant to be a textbook with exercises and class
activities, nevertheless, I provided detailed proofs of results that a student can easily follow.
I envisioned this book to be a supplementary reference for a course on integral transforms.
The first five chapters should also be accessible to scientists with some basic knowledge of
mathematical analysis and could be used as an introductory course on fractional integral
transforms, in general, and on the fractional Fourier transform, in particular. I hope this
book will be useful to scientists in different fields. In fact, more than fifty percent of
the references cited in the bibliography were written by physicists and engineers and are
published in engineering and optics research journals.
The book is organized as follows: To make the book self-sufficient, I included in Chapter
1 the preliminary material and basic results on special functions, functional analysis and
harmonic analysis that will be used later. Some special topics, like sampling theorems, Pois-
son summation formula, generalized functions and distributions and uncertainty principle,
which are discussed in later chapters, were also included in Chapter 1 but without proofs.
However, references where the interested reader can find the proofs are given. Chapter 2 is
an introduction to the general topic of integral transforms and its history. It includes a list of
more than 25 integral transforms, some are more than hundred years old, like the Laplace
transform, and some are recent, like the Gyrator transform.
The general concept of fractional integral transforms is presented in Chapter 3 with a
number of examples of such transforms, such as fractional Fourier, fractional Hankel and
fractional Jacobi transforms. The connection between fractional derivatives and fractional
integrals on the one hand and fractional integral transforms on the other hand is discussed
in this chapter.
 Preface  xv

Chapter 4 focuses on the fractional Fourier transform and its basic properties. It may
be viewed as a tour of the fractional Fourier transform’s journey; it begins by a historical
overview of the development of the transform, then proceeds to unwrap some of its unique
properties and finally ends with one of its basic applications in optics. This is the longest
chapter in the book because in addition to the discussion of the basic properties of the
transform, it explores how the fractional Fourier, fractional Hilbert, fractional Wigner and
fractional ambiguity transforms intertwine.
Chapter 5 discusses shift-invariant and sampling spaces in the setting of the fractional
Fourier transform. This discussion leads to the introduction of a discrete fractional Fourier
transform, fractional convolution structure and fractional Zak transform. Chapter 6 intro-
duces the coupled fractional Fourier transform which is a novel extension of the fractional
Fourier transform to two dimensions. The chapter discusses the unique properties of this
transform, in particular, its sampling theorem and the associated sampling points config-
urations. The chapter is concluded by a discussion of a relatively new integral transform
that is intimately related to the two-dimensional fractional Fourier transform, and which is
called the Gyrator transform. The Gyrator transform, which was also discovered in optics
in the year 2000, is obtained from the two-dimensional fractional Fourier by coordinate
rotations.
The relationship between the two-dimensional coupled fractional Fourier transform
and the four-dimensional Wigner distribution is presented in Chapter 7. This relationship
inevitably leads to the study of four-dimensional rotations which is not as well known
as three-dimensional rotations. To make the material accessible to junior researchers, a
detailed discussion of four-dimensional rotations is presented.
Chapter 8 contains more properties of the coupled fractional Fourier transform, such
as its extension, convolution and uncertainty relations. It also introduces the short-time
coupled fractional Fourier transform and its basic properties.
Chapter 9 is an introduction to the class of linear canonical transforms which contains
the fractional Fourier transform as a special case. Several properties of the linear canonical
transform, such as its convolution, sampling theorems and metaplectic group representation
are presented. The metaplectic representation of the linear canonical transform facilitates the
extension of the transform to higher dimensions and makes the derivation of its properties
managable.
Chapter 10, which is the last chapter of the book, introduces the Special Affine Fourier
Transform and another variant of it, known as the offset linear canonical transform. The
special affine Fourier transform is a generalization of the linear canonical transform and
is the most general inhomogeneous, lossless linear mapping in phase space. Elementary
properties, Poisson summation formula, Zak transform and sampling theorems for the
special affine Fourier transform are presented in this chapter.
The book ends with a bibliography that contains a wholesome list of references. Al-
though there are thousands of articles written on these topics, it was impractical to include
most of them. I have limited my choices to those references that are closely related to the
topics presented in this volume. I apologize to the authors whose work I have missed. The
majority of the references are published in engineering and optics research journals which
attests to the fact that the subject of fractional integral transforms is not only relevant to
mathematicians, but to engineers and physicists as well.
xvi  Preface

I strived to make some chapters self-contained. To this end, I had to restate results
presented in previous chapters and provide proofs with significant level of detail. This
might have come at the expense of brevity and elegancy. To appreciate the development of
any mathematical topic, one ought to know its history and how it started. To help the reader
achieve that, I have included a brief historical introduction to each of the main integral
transforms presented in this volume. Inspite of concerted efforts of everyone involved in
this book project, one may expect to find some typographic errors which I hope will be few
and obvious and will not cause any distraction.
Some of the work presented in this volume is a result of the author’s own research and
publications, either individually or in collaboration over the years with several colleagues,
of whom the most recent are Professors Ayush Bhandari, Firdous Shah, Azhar Tantary and
Rajakumar Roopkumar. My collaboration with Professor Bhandari began when he was an
undergraduate student at Nanyang Technological University, Singapore. But ironically, I
have never met either Professors Shah, Tantary or Roopkumar, but thanks to the internet
that made our collaboration possible.
This project took more time than I anticipated because most of the work done on it took
place while I was chairing a large mathematics department with more than 70 full-time
and part-time instructors, having other university administrative duties and dealing with
restricted working conditions under Covid-19 lockdown.
I take this opportunity to acknowledge the encouragement and guidance I have received
over the years from Professors Paul Butzer, Johen Benedetto and Gilbert Walter.
Finally, I would like to express my appreciation to the staff of CRC Press, Taylor &
Francis Group, in particular, to Mansi Kabra and Kumar Shashi for their support throughout
the production process.
12  Fractional Integral Transforms: Theory and Applications

where the integration path, c, is a loop that starts and ends at −∞ and encircles the disc
|t| ≤ |z|1/a in the positive sense. Moreover, it satisfies the relations
Z ∞
1
e−t tb−1 Ea,b (ta z) dt = ,
0 1−z
and m
d

[z b−1 Ea,b (z a )] = z b−m−1 Ea,b−m (z a ).
dz

1.2.8 The Hypergeometric and q -Hypergeometric Functions

Recall the Pochhammer symbol

(a)0 = 1, (a)n = a(a + 1)(a + 2) · · · (a + n − 1), n = 1, 2, 3, . . . .

Or
Γ(a + n)
(a)n = , n = 0, 1, 2, . . . .
Γ(a)
The hypergeometric function [107]
∞
X (a1 )n · · · (ap )n z n
p Fq (ar ; bt ; z) = ,
n=0
(b1 )n · · · (bq )n n!

which converges for all z if p ≤ q, diverges (except for z = 0) if p > q + 1, and converges
for |z| < 1 if p = q + 1.
We use the notation
n
(1 − aq k−1 ), |q| < 1, n = 1, 2, . . . , ∞,
Y
(a; q)0 = 1, (a; q)n =
k=1
m
Y
(a1 , . . . , am ; q)n = (al ; q)n ,
l=1
(a; q)∞
(a; q)α = .
(aq α , q)∞

Note that
(q a ; q)n
lim = (a)n = a(a + 1) · · · (a + n − 1);
q→1− (1 − q)n

hence for a = 1 the limit is n!.

The symbol r+1 φr stands for the q-hypergeometric function
∞
X (a1 , . . . , ar+1 ; q)n n
r+1 φr (a1 , . . . , ar+1 ; b1 , . . . , br ; q, z) = z .
n=0
(q, b1 , . . . , br ; q)n
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small specimens sit on, or run over the big individuals, and even
nestle under them without their displaying the least resentment. The
common cockroach is a rather amusing pet, as the creatures
occasionally assume most comical attitudes, especially when
cleaning their limbs; this they do somewhat after the fashion of cats,
extending the head as far as they can in the desired direction, and
then passing a leg or antenna through the mouth; or they comb other
parts of the body with the spines on the legs, sometimes twisting and
distorting themselves considerably in order to reach some not very
accessible part of the body.

There is very little information extant as to the domestic Blattidae

found in parts of the world outside Europe, but it seems that there
are numerous species that prefer the dwellings of man, even though
they only tolerate the owners. Belt says[159] "the cockroaches that
infest the houses of the tropics are very wary, as they have
numerous enemies—birds, rats, scorpions, and spiders; their long
trembling antennae are ever stretched out, vibrating as if feeling the
very texture of the air around them; and their long legs quickly take
them out of danger. Sometimes I tried to chase one of them up to a
corner where on a wall a large cockroach-eating spider stood
motionless looking out for his prey; the cockroach would rush away
from me in the greatest fear, but as soon as it came within a foot of
its mortal foe nothing would force it onwards, but back it would
double, facing all the danger from me rather than advance nearer to
its natural enemy." To this we may add that cockroaches are the
natural prey of the fossorial Hymenoptera of the group Ampulicides,
and that these wasps sometimes enter houses in search of the
Insects.

Fig. 127.—Nocticola simoni. A, male; A1, tegmen and rudiment of wing;

A2, front of head; B, female. The cerci are broken, in B the right
one is restored in outline. (After Bolivar.)

We have already noticed the considerable difference that exists in

many cases between the sexes of the same species. This is
sometimes carried to such an extent that nothing but direct
observation could make us believe that the males and females are of
one kin. Fig. 118 (p. 220) shows a case of this kind. Though the
young as a rule are excessively similar to the adults, yet this is by no
means invariably the case. In some of the more amply winged forms,
such as Blabera, the young is about as different from the adult as the
female of Heterogamia is from its male. In Blattidae it is always the
case—so far as is yet known—that when there is a difference as
regards the alar organs between the two sexes, it is the male that
has these structures most developed, and this even when they can
be of little or no use for purposes of flight.

Among the most interesting forms of the family are the two species
of the genus Nocticola, recently discovered by M. Simon in caves in
the Philippine Islands.[160] They are amongst the smallest of the
Orthoptera, the male being scarcely ⅛ of an inch long. In the larval
state of N. simoni the ocular organs exist as three ocelli, or facets,
on each side of the head, and in the perfect state the number is
increased somewhat, as shown in Fig. 127, A2. In the second
species of the genus the female is quite blind (the male being still
undiscovered). The fenestræ in Nocticola are absent; the tegmina
and wings are totally wanting in the female (Fig. 127, B), but are
present in a very peculiar condition in the male (Fig. 127, A1). There
are other anomalies in the structure of these cavernicolous Insects,
the cerci being apparently of peculiar structure, and the spines of the
legs more hair-like than usual. The condition of the eyes is
remarkable; the peculiarity in their development is worthy of study.

Fig. 128.—Corydia petiveriana, with tegmina extended, A; closed, B.

To those who are acquainted with Blattidae only through our

domestic "black beetle" it may seem absurd to talk of elegance in
connexion with cockroaches. Yet there are numerous forms in which
grace and beauty are attained, and some exhibit peculiarities of
ornamentation that are worthy of attention. Corydia petiveriana (Fig.
128) is a common cockroach in East India. It has an effective system
of coloration, the under wings and the sides of the body being vividly
coloured with orange yellow; when the tegmina are closed the upper
surface of the body is of a velvet-black colour, with cream-coloured
marks; these spots are different on the two tegmina, as shown in Fig.
128, A, but are so arranged that when the tegmina are closed (Fig.
128, B) a symmetrical pattern is produced by the combination of the
marks of the two differently spotted tegmina. It is very curious to
notice the great difference in the colour of the part of the right
tegmen that is overlapped by the edge of the left one; this part of the
tegmen being coloured orange yellow in harmony with the wings.
The result of the remarkable differentiation of the colours of the two
tegmina may be summarised by saying that on the right one the
colour of a part is abruptly contrasted with that of the rest of the
organ, so as to share the system of coloration of the under-wings
and body, while the corresponding part of the other tegmen is very
different, and completes the system of symmetrical ornamentation of
the upper surface.

Many other members of the Blattidae have an elegant appearance,

and depart more or less from their fellows in structural characters,
with the result of adding to their graceful appearance; in such cases,
so far as at present known, these Insects are brightly coloured. Thus
Hypnorna amoena (Fig. 129) has the antennae banded in white,
black, and red, while the overlapping part of the tegmina is arranged
so as to bring the line of junction between them nearly straight along
the middle line of the body, and thus produce a more symmetrical
appearance than we find in other cockroaches. The head in this
Insect is not so concealed as usual, and this undoubtedly adds
somewhat to the effective appearance of this cockroach. This
visibility of the front of the head in Hypnorna is not, as would be
supposed, owing to its being less inflexed than usual. On the
contrary, the head is quite as strongly inflexed as it is in other
Blattidae, but the part just at the front of the thorax is unusually
elongate, so that the eyes are exposed and the Insect has a larger
field of vision. This interesting Insect belongs to the tribe
Oxyhaloides [Plectopterinae Sauss.], in which group the most highly
developed folded wings occur.

Fig. 129.—Hypnorna amoena. Central America. Tribe Oxyhaloides.

(After de Saussure.)
The wingless forms never exhibit the grace and elegance possessed
by some of the more active of the winged Blattidae. One of them,
Gromphadorhina portentosa, found in Madagascar (Fig. 130), is a
very robust Insect, and attains a length of 78 millim.—somewhat
more than 3 inches. This Insect has projections on the thorax that
remind us of the horns that exist in some of the Lamellicorn beetles.

Little has been yet written as to the resemblances of Blattidae to

other species of their own family, or to other creatures, but it is
probable that such similarities will be found to prevail to a
considerable extent. W. A. Forbes has called attention[161] to the
larva of a Blattid from Brazil as being remarkable for its superficial
resemblance to an Isopod crustacean. Some of the wingless forms
have a great resemblance to the small rolling-up Myriapods of the
group Glomerides; Pseudoglomeris fornicata, of which we figure the
female (Fig. 131), has received its name from this resemblance. The
females of the S. African genus Derocalymma possess this Glomerid
appearance, and have a peculiar structure of the prothorax,
admitting of a more complete protection of the head. Brunner states
that the wingless kinds of Derocalymma roll themselves up like
wood-lice. In many of the forms of this tribe—Perisphaeriides—the
males are winged, though the females are so like Myriapods.
According to de Saussure[162] the gigantic Megaloblatta rufipes
bears an extreme resemblance in appearance to the large
cockroaches of the genus Blabera.

Fig. 130.—Gromphadorhina portentosa, × ⅔. Tribe Perisphaeriides.

(After Brunner.)

Fig. 131.—Pseudoglomeris fornicata, ♀. Burma. Tribe Perisphaeriides.

(After Brunner.)
Some of the species of Holocompsa remind us strongly of Hemiptera
of the family Capsidae; they have an arrangement of colours similar
to what prevails in that group, and their tegmina and wings which, as
being those of Blattids may be said to be abnormally formed,
resemble in texture and the distribution of the venation those of the
Hemiptera. These Insects are closely allied to Diaphana, of which
genus we have figured a species (Fig. 122).

There is very little evidence on which to base an estimate of the

number of species of Blattidae existing in the world at present.
Probably the number extant in collections may amount to 1000 or
thereabouts, and the total existing in the world may be as many as
5000. The species of Blattidae cannot tolerate cold, and are
consequently only numerous in tropical regions. Europe possesses
about twenty species, and in Britain there are only three that are truly
native; these are all small Insects belonging to the genus Ectobia,
and living out of doors, amongst leaves, under bushes, and in
various other places. We have, however, several other species that
have been introduced by the agency of man, and these all live under
cover, where there is artificial warmth and they are protected from
the inclemencies of the winter season. The commonest of these
forms is Stilopyga orientalis, the "black beetle" of our kitchens and
bakehouses. This Insect is said to have been brought to Europe from
"Asia" about 200 years ago, but the evidence as to its introduction,
and as to the country of which it is really a native, is very slight. It is
indeed said[163] that S. orientalis has been found in peat in
Schleswig-Holstein. Periplaneta americana is a larger Insect, and is
common in some places; it is apparently the species that is most
usually found on board ships, where it sometimes multiplies
enormously, and entirely devours stores of farinaceous food to which
it obtains access: it is known that sometimes a box or barrel
supposed to contain biscuits, on being opened is found to have its
edible contents entirely replaced by a mass of living cockroaches.
Fortunately Periplaneta americana has not spread widely in this
country, though it is found in great numbers in limited localities; one
of the best known of which is the Zoological Gardens in the Regent's
Park at London. Periplaneta australasiae is very similar to P.
americana, but has a yellow mark on the shoulder of each tegmen.
This has obtained a footing in some of the glass-houses in the
Botanic Gardens at Cambridge and Kew; and it is said to be fairly
well established in Belfast. Another of our introduced domestic
cockroaches is Phyllodromia germanica, a much smaller Insect than
the others we have mentioned. It has only established itself at a few
places in this country, but it is extremely abundant in some parts of
Northern and Eastern Europe. It has been increasing in numbers in
Vienna, where, according to Brunner, it is displacing Stilopyga
orientalis. In addition to these, Rhyparobia maderae and species of
the genus Blabera have been met with in our docks, and are
possibly always to be found there. They are Insects of much larger
size than those we have mentioned. We figure the alar organs of one
of these species of Blabera of the natural size: the species in this
genus are extremely similar to one another. Blaberae are known in
the West Indies as drummers, it being supposed that they make a
noise at night,[164] but details in confirmation of this statement are
wanting.

Fig. 132.—Alar organs of Blabera sp. A, tegmen; B, wing.

It is a remarkable fact that no satisfactory reasons can be assigned

for the prevalence of one rather than another of these domestic
cockroaches in particular localities. It does not seem to depend at all
on size, or on the period of development, for the three species
Stilopyga orientalis, Periplaneta americana, and Phyllodromia
germanica, which are the most abundant, differ much in these
respects, and replace one another in particular localities, so that it
does not appear that any one is gaining a permanent or widespread
superiority as compared with another. There are, however, no
sufficient records on these points, and further investigation may
reveal facts of which we are at present ignorant, and which will throw
some light on this subject. We may remark that Mr. Brindley has
found it more difficult to obtain hatching of the young from the egg-
capsules of Periplaneta americana and Phyllodromia germanica at
Cambridge, than from those of Stilopyga orientalis.

Although much work has been done on the embryology of Blattidae,

the subject is still very incomplete. The recent memoirs of
Cholodkovsky[165] on Phyllodromia germanica contain so much of
general interest as to the development of the external parts of the
body that we may briefly allude to them. The earliest appearance of
segmentation appears to be due to the centralisation of numerous
cells round certain points in the ventral plate. The segmentation of
the anterior parts is first distinct, and the appearance of the
appendages of the body takes place in regular order from before
backwards, the antennae appearing first; the mandibles, however,
become distinct only subsequent to the maxillae and thoracic
appendages. There are in the course of the development
appendages to each segment of the body (he counts eleven
abdominal segments); the cerci develop in a similar manner to the
antennae; the first pair of abdominal appendages—at first similar to
the others—afterwards assume a peculiar stalked form. The
abdominal appendages subsequently disappear, with the exception
of the ninth pair, which form the ventral styles, and the eleventh pair,
which become the cerci. The last ventral segment is said to be
formed by the union of the tenth and eleventh embryonic ventral
segments.

Fig. 133.—A, Tegmen(?) of Palaeoblattina douvillei; B, of Etoblattina

manebachensis. (After Brauer and Scudder.)
As regards their Palaeontological forms Blattidae are amongst the
most interesting of Insects, for it is certain that in the Carboniferous
epoch they existed in considerable number and variety. A still earlier
fossil has been found in the Silurian sandstone of Calvados; it
consists of a fragment (Fig. 133, A), looking somewhat like an
imperfect tegmen of a Blattid; it was described by Brongniart under
the name of Palaeoblattina douvillei, and referred by him, with some
doubt, to this family. Brauer has, however, expressed the opinion[166]
that the fragment more probably belonged to an Insect like the mole-
cricket, and in view of this discrepancy of authorities we may be
pardoned for expressing our own opinion to the effect that the relic
has no connexion with the Insecta. The figure given by Scudder[167]
has not, however, so uninsect-like an appearance as that we have
copied from Brauer. Whatever may prove to be the case with regard
to Palaeoblattina, it is certain, as we have already said, that in the
Palaeozoic epoch Insects similar to our existing cockroaches were
abundant, their remains being found in plenty in the coal-measures
both of Europe and North America. Fig. 133, B, shows a fossil
tegmen of Etoblattina manebachensis from the upper Carboniferous
beds of Ilmenau in Germany. It will be noticed that the disposition of
the nervures is very much like that which may be seen in some of
our existing Blattidae (cf. the tegmen of Blabera, Fig. 132, A), the
vena dividens (a) being similarly placed, as is also the mediastinal
vein on the front part of the organ. The numerous carboniferous
Blattidae have been separated as a distinct Order of Insects by
Scudder under the name Palaeoblattariae, but apparently rather on
theoretical grounds than because of any ascertained important
structural distinctions. He also divided the Palaeoblattariae into two
groups, Mylacridae and Blattinariae, the former of which was
supposed to be peculiar to America. Brongniart has, however,
recently discovered that in the Carboniferous deposits of Commentry
in France Mylacridae are as common as in America. This latter
authority also states that some of the females of these fossil
Blattidae are distinguished by the presence of an elongate exserted
organ at the end of the body. He considers this to have been an
ovipositor by which the eggs were deposited in trees or other
receptacles, after a manner that is common in certain Orthoptera at
the present day. If this view be correct these Carboniferous Insects
must have been very different from the Blattidae of our own epoch,
one of whose marked characteristics is the deposition of the eggs in
a capsule formed in the body of the parent.

In the strata of the secondary epoch remains of Blattidae have also

been discovered in both Europe and America, in Oolitic, Liassic, and
Triassic deposits. From the Tertiary strata, on the other hand,
comparatively few species have been brought to light. A few have
been discovered preserved in amber.

Fig. 134.—Front leg of Periplaneta australasiae.

The classification of the Blattidae is attended with considerable

difficulty on account of the numerous wingless forms, and of the
extreme difference in the organisation of the two sexes of many
species. It has, however, been brought to a fairly satisfactory state
by the reiterated labours of Brunner von Wattenwyl, and we
reproduce his recently perfected exposition of their characters. His
first division is made by means of a structure which is very easily
observed, viz. whether the femora are armed with spines, as in Fig.
134, or not. The terms used in connexion with the wings and other
parts of the body we have already explained.

Brunner's system is adopted by de Saussure,[168] who, however,

proposes to replace the names Ectobiides and Oxyhaloides by
Anaplectinae and Plectopterinae. He also proposes to apply the
generic name Blatta to the Insect that is now so frequently called
Phyllodromia germanica in zoological works. If that view be adopted,
Brunner's group Phyllodromiides will be called Blattides.

Table of the tribes of Blattidae, after Brunner:—

1. Femora spiny beneath.[169]

2. The last ventral plate of the female large, without valves.
3. Supra-anal lamina of both male and female transverse, narrow.
Wings, when present, furnished with a triangular apical field.
Posterior femora unarmed beneath, or armed with two spines on
the anterior margin. Egg-capsules furnished with a longitudinal
suture. Tribe 1. Ectobiides. [Anaplectinae Saussure.]
3′. Supra-anal lamina of each sex more or less produced, triangular, or
emarginate. Wings, when present, without apical field. Posterior
femora with both edges spiny.
4. Supra-anal lamina of each sex triangular, not notched. Cerci
projecting much beyond this lamina.
5. Pronotum and elytra smooth (i.e. without peculiarity of surface
other than punctuation). The radial nervure of the wing giving
off several parallel branches, pectinate on the anterior margin
(except in the genus Abrodiaeta). Tarsal joints without pads.
Tribe 2. Phyllodromiides. [Blattinae Saussure.]
5′. Pronotum and elytra holosericeous. Radial nervure of the wings
giving off irregular branches on the anterior margin (ulnar vein
many-branched). Tarsal joints furnished with pads. Tribe 3.
Nyctiborides.
4′. Supra-anal lamina of males more or less four-sided, with obtuse
angles, of females broad, rounded, or lobed. Cerci not projecting
beyond the lamina. (Tarsal joints with distinct pads.) Ulnar
nervure of the wings giving off parallel branches towards the vena
dividens. Tribe 4. Epilamprides.
2′. The last ventral plate of the female furnished with valves. Tribe 5.
Periplanetides.[170] (Fig. 119, Periplaneta australasiae.)
1′. Femora unarmed beneath. (In the tribe Panesthiides the anterior femora
are frequently armed with two spines.)
 2. Supra-anal lamina of each sex more or less produced, posterior margin
notched.
3. A distinct pad between the claws. Tribe 6. Panchlorides.
3′. No pad between the claws, or only an excessively small one.
4. Wings with a folded fan-like anal field. Pronotum smooth. Tribe 7.
Blaberides. (Fig. 132, Blabera sp. wings.)
4′. Anal field of the wing with a single fold. Pronotum more or less
pilose. Tribe 8. Corydiides. (Fig. 128, Corydia petiveriana. Fig.
118, Heterogamia aegyptiaca.)
2′. Supra-anal lamina of each sex, short, transverse, posterior margin
straight or rounded.
3. Subgenital lamina of the male somewhat produced, furnished with a
single style. Tarsal claws with a distinct pad (except in the genus
Paranauphoeta).
4. Anterior portion of the wings pointed, either the apical field of the
wing very much produced, or the wings twice as long as the
tegmina, folded in repose. Tribe 9. Oxyhaloides. [Plectopterinae
Saussure.] (Fig. 129, Hypnorna amoena.)
4′. Anterior portion of wing, when present, rounded, with no apical
field. Tribe 10. Perisphaeriides. (Fig. 130, Gromphadorhina
portentosa; Fig. 131, Pseudoglomeris fornicata.)
3′. Subgenital lamina of males extremely small, without styles. No pad
between claws. Tribe 11. Panesthiides.

To the above tribes another one—Geoscapheusides—has been

recently added by Tepper,[171] for an extraordinary Australian Insect
of fossorial habits, with front legs formed somewhat like those of
Gryllotalpa.

CHAPTER X

ORTHOPTERA CONTINUED—MANTIDAE—SOOTHSAYERS
 Fam. IV. Mantidae—Soothsayers or Praying Insects.

Orthoptera with exserted but deflexed head and elongate

prothorax, the first pair of legs largely developed, raptorial, the
coxae elongate, free, femora and tibiae armed with spines:
second and third pair of legs simple and similar; the tarsi five-
jointed, without a pad (arolium) between the claws; a pair of
jointed cerci near the extremity of the body.

The Mantidae are an extensive family of Orthoptera, showing

extreme variety in the shapes and outlines of the body, and
characterised by the very remarkable front legs; the function of these
legs being to seize and hold their prey, which consists of living
Insects, Mantidae being carnivorous and highly voracious.

The labium is deeply divided, each half exhibiting a very near

approach to the structure of a maxilla; there is a large membranous
lingua reposing on the inner face of the lower lip. The head is quite
free from the thorax, its front part being deflexed, and even
somewhat inflexed, so that the mouth is directed downwards and
somewhat backwards: it is very mobile, being connected to the
thorax by a comparatively slender neck, which is, however,
concealed by the pronotum. There are two large, prominent eyes,
the antennae are frequently very slender, but they sometimes differ
according to sex, and in some genera are pectinate in the male; just
above and between their insertion are three ocelli placed in a
triangle, two above, one below; between the antennae and the
clypeus there is an interval called the scutellar space. In some forms
of Mantidae the head assumes most extraordinary shapes; the eyes
may become elongate and horn-like; there may be a projection
between them bearing the ocelli, and attaining occasionally a great
length; the scutellar space also may have a remarkable
development, the whole thus forming a peculiar ornamental
structure, as in Fig. 136.
 Fig. 135.—Deroplatys sarawaca, female. Borneo. (After Westwood.)

The prothorax is elongate, but there are a few genera, e.g.

Eremiaphila, in which it is exceptionally short, and there are several
others in which the elongate form is more or less masked by
foliaceous expansions of the sides. The pronotum shows near the
front a transverse depression or seam, which marks the position of
an internal chitinous ridge. The anterior legs are inserted near the
front of the prosternum, which extends less far forwards than the
pronotum does; the posterior part of the prosternum is very elongate,
and is completely separated from the anterior part by the base of the
coxae and the membranes attached to them; the pronotum and
sternum are closely connected at the sides till near the posterior part
where they diverge, the space so formed being occupied by a
membrane in which the prothoracic stigma is situated. The
mesothorax is as long as broad, and the front wings are attached to
the whole length of the sides; the mesosternum is a triangular piece
pointed behind, and bearing very large side-pieces, to the hinder
portion of which the middle coxae are attached; these latter are large
and quite free, and repose on the metasternum which they cover; the
mesothoracic stigma may be detected as a slit situated on a slight
prominence just behind and a little below the membranous hind-
margin of the tegmen. The metathorax differs comparatively little in
size and structure from the mesothorax; the membranous hind wings
are attached to the sides of the notum along nearly the whole length
of the latter. The abdomen is moderately long; in each sex ten dorsal
plates may be detected, and there is a pair of ringed cerci projecting
from beneath the sides of the tenth plate. The number of ventral
plates is more difficult to verify, the first one being much reduced;
eight other plates can be demonstrated in the male and six in the
female.

Fig. 136.—Head of Harpax variegatus, seen from the front.

The anterior legs are formed in a remarkable manner in the

Mantidae, and are, in fact, the most characteristic feature of the
family. Attached near the front of the thorax there is a very long coxa,
to the apex of which is articulated the triangular trochanter; this
bears the elongate femur, which is furnished on its lower face with
sharp spines and teeth; the tibia which follows is much shorter and
smaller than the femur; its lower face bears also an armature of
teeth, and it is so articulated with the femur that it can be completely
closed thereon, its teeth fitting in among those of the femur (Fig. 137,
B); the latter has one or more longer spines overlapping the apical
part of the tibia when contracted. The tarsus is slender, five-jointed,
without pad. The other two pairs of legs are simple; the hinder
usually a little the longer, and in some species that possess powers
of leaping (Ameles), with the femora a little thicker at the base.
 Fig. 137.—Front leg of Empusa pauperata, female: A, with tibia
extended and tarsus wanting; B, more magnified (the basal parts
removed), showing the mode of closure.

The alar organs of the Mantidae are as regards the nervures and
areas fairly similar to those of the Blattidae. The tegmina are usually
narrow, and exhibit three well-marked areas; the one in front or
external (according as the wing is expanded or closed) is the
mediastinal area; it is usually more elongate and occupies a larger
portion of the surface of the tegmen than in Blattidae. The middle
area, forming the larger part of the wing, is occupied by the branches
of the radial and ulnar nervures. The third area, the anal, possesses
a sort of appendage in the form of a small space of a more delicately
membranous nature at the inner part of the base. The tegmina are
often more or less leaf-like in texture and consistence; this character
is as a rule not very marked, but there are a few species with the
tegmina very like foliage, this being more marked in the female; in
some, if not in all, of these cases the mediastinal area is
considerably increased. One tegmen overlaps the other, as in
Blattidae, but to a less extent, and the correlative asymmetry is but
slight: there is frequently a pallid spot close to the main vein on the
principal area, nearer to the base than to the extremity. The hind
wings are more ample than the front, and of much more delicate
consistence; they possess numerous veins converging to the base;
the anterior part of the wing is firmer in consistence, and its veins are
more numerously furcate; there are many more or less distinct
minute cross-veinlets, and an elegant tinting is not infrequent. They
close in a fan-like manner, transverse folding being unknown in the
family.

But little has been written on the internal anatomy of the Mantidae.
Dufour has described only very partially that of M. religiosa. The
salivary glands are largely developed, salivary receptacles exist; the
alimentary canal possesses eight elongate coecal diverticula placed
on the chylific ventricle; there are about one hundred Malphigian
tubules. In each ovary there are about 40 egg-tubes, and they are
joined at their bases in clusters of about half a dozen; each cluster
has a common sinus; these sinuses are placed at intervals along a
tube, which is one of two branches whose union forms the oviduct;
there are a large number of "serific glands" of two kinds in the
female. The testes are unusually complex in their structure.

According to Schindler[172] the Malphigian tubes in Mantis are not

inserted, as usual, at the base of the intestine, but on the intestine
itself at about one-third of its length from the base. There is some
doubt about this observation. Schindler considers the fact, if it be
such, unique.

The eggs of the Mantidae are deposited in a singular manner: the

female, placing the extremity of the body against a twig or stone,
emits some foam-like matter in which the eggs are contained. This
substance dries and forms the ootheca; whilst attaining a sufficient
consistence it is maintained in position by the extremity of the body
and the tips of the elytra, and it is shaped and fashioned by these
parts. The eggs are not, as might be supposed, distributed at
random through the case, but are lodged in symmetrically-arranged
chambers, though how these chambers come into existence by the
aid of so simple a mode of construction does not appear. The
capsule is hard; it quite conceals the eggs, which might very
naturally be supposed to be efficiently protected by their covering:
this does not, however, appear to be the case, as it is recorded that
they are subject to the attacks of Hymenopterous parasites. The time
that elapses after the eggs are laid and before they hatch varies
greatly according to circumstances. In France, Mantis religiosa
deposits its eggs in September, but they do not hatch until the
following June; while in E. India the young of another species of
Mantis emerge from the eggs about twenty days after these have
been deposited. Trimen has recorded some particulars as to the
formation of its egg-case by a Mantis in S. Africa. This specimen
constructed four nests of eggs at intervals of about a fortnight, and
Trimen states that the four were "as nearly as possible of the same
size and of precisely similar shape." He also describes its mode of
feeding, and says that it was fond of house-flies, and would eat
"blue-bottles," i.e. Musca vomitoria, but if while eating one of the
latter a house-fly were introduced, the "blue-bottle" was generally
dropped, even though it might be in process of being devoured. The
young have to escape from the chambers in which they are confined
in these egg-cases; they do so in a most curious manner; not by the
use of the feet, but by means of spines directed backwards on the
cerci and legs, so that when the body is agitated advance is made in
only one direction. The eggs last deposited are said to be the first to
hatch. On reaching the exterior the young Mantids do not fall to the
ground, but remain suspended, after the manner of spiders, to the
ootheca by means of two threads attached to the extremities of the
cerci; in this strange position they remain for some days until the first
change of skin is effected, after which they commence the activity of
their predatory life.

Fig. 138.—Egg-case of Mantis with young escaping: A, the case with

young in their position of suspension; B, cerci magnified, showing
the suspensory threads. (After Brongniart.)

Dr. Pagenstecher has given an account[173] of the development of

Mantis religiosa, from which it would appear that the statements of
Fischer and others as to the number of moults are erroneous, owing
to the earliest stages not having been observed. When the young
Mantis emerges from the egg it bears little resemblance to the future
Insect, but looks more like a tiny pupa; the front legs, that will
afterwards become so remarkable, are short and not different from
the others, and the head is in a curious mummy-like state, with the
mouth-parts undeveloped and is inflexed on the breast: there are, he
says, nine abdominal segments. The first ecdysis soon takes place
and the creature is thereafter recognisable as a young Mantis.
Pagenstecher's specimens at first would only eat Aphididae, but at a
later stage of the development they devoured other Insects greedily:
the number of ecdyses is seven or eight. The ocelli appear for the
first time when the wing rudiments do so; the number of joints in the
antennae increases at each moult. Dr. Pagenstecher considers that
this Insect undergoes its chief metamorphosis immediately after
leaving the egg, the earlier condition existing apparently to fit the
Insect for escaping from the egg-case. In the immature stage of the
Mantidae the alar organs appear (Fig. 139) as adjuncts of the sides
of the meso- and meta-notum, projecting backwards and very deeply
furrowed and ribbed in a wing-like manner. According to
Pagenstecher, this wing-like appearance only commences in the fifth
stadium, but he has not given particulars of the conditions of these
parts in the preceding instars. According to de Saussure[174] the
wings of the females of some species remain permanently in this
undeveloped or nymphal state.

Fig. 139.—Tegmina (t) and wings (w) of immature Mantis.

Fig. 140.—Iris oratoria, female. South Europe. Natural size.

The Mantidae, as a rule, have a quiet unobtrusive mien, and were it

not for their formidable front legs would look the picture of
innocence; they, however, hold these legs in such manner as to
greatly detract from the forbidding appearance thereof, stretching
them out only partially so as to give rise to an appearance of
supplication or prayer;[175] this effect is increased by their holding
themselves in a semi-erect position, standing on the hind and middle
legs with the upper parts of the body directed somewhat forwards,
hence they are called by various names indicating prayer or
supplication, and it is said that in some countries they are considered
sacred. Some of the older writers went so far as to say that a Mantis
would indicate the road a child should take by stretching out one of
its arms in the right direction. The traveller Burchell, speaking of a
species since described by Westwood under the name of
Tarachodes lucubrans, says: "I have become acquainted with a new
species of Mantis, whose presence became afterwards sufficiently
familiar to me by its never failing, on calm warm evenings, to pay me
a visit as I was writing my journal, and sometimes to interrupt my
lucubrations by putting out the lamp. All the Mantis tribe are very
remarkable Insects; and this one, whose dusky sober colouring well
suits the obscurity of night, is certainly so, by the very late hours it
keeps. It often settled on my book, or on the press where I was
writing, and remained still, as if considering some affair of
importance, with an appearance of intelligence which had a
wonderful effect in withholding my hand from doing it harm. Although
hundreds have flown within my power, I never took more than five. I
have given to this curious little creature the name of Mantis
lucubrans; and having no doubt that he will introduce himself to
every traveller who comes into this country [Southern Africa] in the
months of November and December, I beg to recommend him as a
harmless little companion, and entreat that kindness and mercy may
be shown to him." This appearance of innocence and quietness
must have struck all who have seen these Insects alive;
nevertheless, it is of the most deceptive character, for the creature's
activity consists of a series of wholesale massacres carried on day
after day, the number of victims it sacrifices being enormous. The
Mantis does not even spare its own kind; it is well known that the
female not unfrequently devours its own mate. A very different
picture to that of Burchell has been drawn by Potts, who observed
the habits of a species in New Zealand.[176] He informs us that when
about making an attack it approaches its intended prey with slow,
deliberate movements, its anterior limbs folded in an innocent