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BIOLOGY AND CULTURE
OF PORTUNID CRABS
OF WORLD SEAS
Biology and Ecology of Marine Life
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ABOUT THE AUTHOR
Series Author:
Ramasamy Santhanam, PhD
Former Dean, Fisheries College and Research Institute,
Tamil Nadu Veterinary and Animal Sciences University,
Thoothukkudi-628 008, India
Email: rsanthaanamin@yahoo.co.in
1. Introduction ................................................................................................ 1
2. Biology and Ecology of Portunid Crabs ................................................... 9
3. Profile of Portunid Crabs (Family: Portunidae) ................................... 23
4. Aquaculture of Portunid Crabs ............................................................ 273
5. Nutritional Values of Portunid Crabs .................................................. 307
6. Biomedical, Food, and Industrial Applications of
Portunid Crab Wastes............................................................................ 329
7. Diseases and Parasites of Portunid Crabs ........................................... 349
References ................................................................................................ 375
Index ......................................................................................................... 389
LIST OF ABBREVIATIONS
INTRODUCTION
CONTENTS
People living on the coast have depended on marine resources for many
thousands of years. Millions earn their living in commercial fisheries,
and many people still depend on seafood as their primary source of pro-
tein. The very high catch rates of modern industrialized fishing fleets
have lead to widespread effects on fished species, non-target species,
and marine habitats. Further, overfishing continues to be a major threat
to marine ecosystems and vulnerable species throughout the world.
Although dwarfed by fisheries for finfish, crab fisheries are important in
2 Biology and Culture of Portunid Crabs of World Seas
Crabmeat, like fish, has been reported as a nutritious food, low in calories
and fat but high in protein and omega-3 fatty acids, which help prevent
heart disease. These fatty acids are especially important for women who
are pregnant or breastfeeding because they are needed for their child’s
developing nervous system. Omega-3s, which are higher in crabmeat than
Introduction 5
The crabmeat is very tasty and nourishing. Crab curry is a reputed cure for
asthma. Similarly, the soup made from the swimming crabs Portunus san
quinolentus and Portunus pelagicus are commonly used by the people just
after recovery from malaria and typhoid. Scylla serrata serves as a cure
for diarrhea and dysentery. All the edible crabmeats are rich in vitamins
and are good for colds, asthma, eosinophilia, primary complex, wheezing,
etc. They are also believed to stimulate brain cells. Crab shells are also
used nowadays for preparing suture threads. Ointment prepared from crab
shells is known to heal wounds quickly and avoids scars.
All the portunid crabs are indirectly important as they play a dominant
role in the marine food web. The faeces of these crabs consist of carbon,
nitrogen, phosphorus and trace metals which form a rich food for other
consumers of the marine ecosystems. The crabs and their larvae are con-
sumed by many predators and omnivorous fishes, and they play a vital
role in the transfer of energy through the food chain. Thus they are also of
immense help in recycling nutrients which enhance the richness of the soil
by “ploughing”.
Portunid crabs have been associated with religion since first century.
Romans worshiped the crab as Sea God and named the crab as Neptune.
The crab Charybdis feriatus was the symbol of cross by Christians from
coastal areas of India. They do not normally eat this crab and let it free to
6 Biology and Culture of Portunid Crabs of World Seas
swim in the sea after fishing. Portunus sanguinolentus, which has three
reddish round spots each encircled by whitish ring, is considered to be the
manifestation of Lord Siva who has three eyes. The abnormal growth of
body cells (due to cancer/tumor) resembles the nesting ground of Cancer
spp. So the generic name is used to denote this killer disease.
Many highly active biocompounds have recently been isolated from por-
tunid crabs associated with antimicrobial, antileukemic, anticoagulant and
cardio active properties. A few species of portunid crabs have been labeled
as ‘Pests’ as they are causing damage to coastal plants and culture ponds.
However, the smaller intertidal crabs which are not of direct economic
value have been used in the preparation of high-energy yielding, cheaper
artificial pellet feeds for the aquaculture of edible varieties of seafood.
Portunid crabs are always the chosen as test animals since their tolerance
level is very wide. They have been used as tools in the elucidation of
physiological mechanisms, fertilization, regeneration and cell association
and mechanisms of drug action.
At present, there is no ban on fishing immature and the berried crabs and
the minimum size at capture is not implemented in many countries. As
a conservation measure, the only possibility is to educate fishermen to
Introduction 7
release the juvenile, berried and soft crabs to the sea while they are alive.
The governments should take necessary steps to implement ban during
peak spawning seasons to prevent indiscriminate fishing. The best method
to ensure a sustainable fishery throughout the year as well as to improve
the quality of the yield is to ban fishing and marketing of undersized and
berried crabs.
KEYWORDS
• aquaculture
• commercial species
• conservation
• indirect economic value
• nutritional and health value
• socio-cultural and scientific value
CHAPTER 2
CONTENTS
2.1.1 HABITAT
The various portunid species live in a variety of habitats, from soft sub-
strates like mudflats and sandflats to harder bottoms like coral reefs and
rubble. Adult habitats range from near shore estuarine to offshore open
water and deep benthic environments. Most portunid species inhabit
inshore habitats as adults and move to more offshore areas such as mouths
of inlets and open waters to spawn.
2.1.2 DISTRIBUTION
Portunid crabs are ubiquitous around the world and the majority of these
crabs are present in the Pacific and Indian Oceans.
10 Biology and Culture of Portunid Crabs of World Seas
Carapace: It is the shell covering the body that provides rigidity and
protective covering. It is made of chitin and is the part of exoskeleton (hard
outer covering) that covers the head and thorax (center) of the crab. Cara-
pace of portunid crabs may be hexagonal—transversely ovate to trans-
versely hexagonal or almost circular. Dorsal surface is generally relatively
flat, and is often ridged or granulose. Front is broad and its margin is usually
multidentate. There are 5 to 9 teeth on each anterolateral margin. Posterolat-
eral margins are usually distinctly converging. The antero-lateral margins of
certain common genera of portunid crabs are given in the figure.
Scylla Portunus
12 Biology and Culture of Portunid Crabs of World Seas
Thalamita Charybdis
Carapace of certain portunid genera
Gills: These are the places of respiration and filtration and are consisting
of many plume-like filaments arranged around a central axis. There are
eight gills on each side of a crab’s body.
Heart: It is the pump of the circulatory system. It is broad in size and
located in the lower center part of the body.
Hepatopancreas (midgut gland): It is an extremely large organ
with several functions, including the secretion of digestive enzymes and
absorption and storage of digested food. It fills most of the area around the
stomach, depending on its contents of food reserves and water.
Intestine: It is the portion of the digestive system through which
digested food passes.
Stomach: This is the organ of the digestive system that breaks down
swallowed particles of food. It is lined with small hard plates and projec-
tions which aid digestion.
Testes: These are the parts of the male reproductive system, located on
top of the hepatopancreas on either side of the stomach.
Cartilage: It encases muscles that help in movement of the legs. The
muscles are the edible portions of the crab.
14 Biology and Culture of Portunid Crabs of World Seas
Portunids include filter feeders, sand cleansers, mud, plant, and carrion
feeders, predators, commensals, and parasites. The diet of portunids var-
ies between the species ranging from slow-moving prey such as snails
and annelids to fast-moving ones such as fish and shrimps. The crab uses
its mouthparts to chop the food into small pieces and then the gastric mill
ossicles further reduce the food to fragments. They are all opportunistic
omnivores with a preference for animal prey, but within that framework
only rarely feed on more mobile prey such as fish and prawns. Crusta-
ceans have been observed to constitute the most favored food item in
Portunus pelagicus diet, followed by molluscs and fish. The presence of
detritus (80%) in the stomachs of this species suggests that these crabs are
also detritivorous, consuming both fresh and decaying flesh of all kinds
of animals.
2.2.3 BEHAVIOR
Portunid crabs are particularly active at night, but are often also out and
about during the day. Besides the large adults, small juvenile swimming
crabs are also hidden among the seagrass and seaweed, and other nooks
and crannies. These active crabs may possess all kinds of colors. Some
can react fiercely by waving spiny pincers. If disturbed, swimming crabs
often fearlessly wave their pincers menacingly. This is not an idle threat.
When people come too close, this crab might just give a good nip that
draws blood.
formed inside, and the old shell becomes loosened. As the crab begins to
molt, the carapace is lifted, revealing a gap between it and the abdomen.
The shell splits on either side of the under surface, and the posterior part
of the crab begins to protrude. Over a period of a few minutes, the crab
completes the molting process, gradually backing out of the old shell.
Immediately after molting, the shell (carapace) is thin, soft and wrinkled.
Subsequently, the new cuticle becomes stretched, and the crab is consid-
erably larger than before the molt. Growth in size of the crab results from
the copious intake of water after molting. The number of times that a
crab molts during its lifetime, and the length of time between molts, vary
among species and are influenced by factors such as temperature and the
amount of food available.
Crabs often lose one or more legs during a lifetime (unless they have
reached a stage where they no longer molt) and are able to re place these
with new limbs. Most crabs have the ability to drop an injured leg. This
process is referred to as autotomy, which is the severance of an appendage
at a preformed breakage point as the result of a reflex action. When a leg
is severed at this breakage point, the resulting opening is closed by a valve,
and there is very little loss of blood from the stump. It has also been shown
16 Biology and Culture of Portunid Crabs of World Seas
As in other crabs, in portunid crabs, sexes are separate and sexes can be
distinguished from the shape of the abdomen. In males the abdomen is
narrow, inverted ‘T’ shaped and in addition mature males have larger and
broader chelae. The first and second abdominal appendages (pleopods)
are highly modified to form an intromittent copulatory organ. Females, on
the other hand, have a broad abdomen which is conical or oval in shape
(according to the stage of maturity) and bear four pairs of pleopods. Many
species of portunid crabs show sexual dimorphism, with males being
larger, smaller, or possessing special or enlarged structures. In some spe-
cies, the females are the larger. Most commonly, males have proportion-
ately much larger chelipeds or chelae. Males always have only two pairs
of gonopods (uniramous swimmerets or pleopods), which are specially
modified for copulation (most crabs have internal fertilization). The first
gonopod is a highly modified pleopod which has been folded or rolled
longitudinally to form a cylindrical tube. The form of the first gonopod
varies from broad to very slender, straight to sinuous, and even recurved.
All day long I have been wandering about through the tombs of the
kings who ruled Egypt three or four thousand years ago. I have gone
into the subterranean chambers which the Pharaohs dug out of the
solid rocks for their burial vaults, and I have visited the tombs of
kings older than they. The last resting places of more than fifty of
these monarchs of early Egypt have been discovered, and the work
is still going on. Some of the best work of excavation all along the
Nile valley is being done by Americans. While at Cairo I found the
money of Harvard College and the Boston Museum uncovering the
cemeteries of the nabobs and paupers who were buried at the time
of King Cheops under the shadow of the Great Pyramid of Gizeh.
The Egyptian Exploration Fund, which is supported by Great Britain,
Canada, and the United States, has a small army of workmen
operating near Luxor, the University of Pennsylvania has made
important discoveries, and a large part of the uncovering of the
valley in which these royal tombs lie has been done by the
Americans.
The Egyptologists of the Metropolitan Museum in New York and
Lord Carnarvon of England are responsible for some of the most
remarkable finds of this generation. During my trip of to-day I met a
young archæologist, in charge of the American operations, who
showed me through the tombs of the kings and explained the
symbols and pictures on the walls. I went to that part of the valley
where the excavation is now going on and took pictures of a gang of
one hundred and fifty Egyptian men and boys who are working there.
Let me describe the place that the ancient Egyptian monarchs
selected for their burials, the Valley of the Kings. They wanted to
hide their remains so that posterity could never find them, and to
cover them so that future generations would have no idea that they
and their treasures lay beneath. Our cemeteries are chosen for the
beauty of their surroundings. We like to turn up our toes to the
daisies and to have leafy trees whisper a requiem over our heads.
The old Egyptian kings wanted to lie under the sterile desert waste
and chose a region about as far up the Nile valley as Cleveland is
inland from the Atlantic, and fully six miles back from the fertile strip
on which their people lived. I can imagine no place more dreary. At
this point the Nile is walled on the west by limestone mountains. As
far as the moisture reaches, the valley is the greenest of green, but
beyond lies a desert as brown as any part of the Sahara. There is
not a blade of grass, nor a sprig of vegetation of any kind. There is
nothing but sand and arid mountains, the latter almost as ragged in
outline as the wildest parts of the Rockies. Some of their stony sides
are built up in great precipices while in other places there are fort-like
bluffs and similar convulsions of nature.
Rameses II, the greatest egoist of Egyptian history, covered his dominions with
his monuments and inscriptions. Standing against the colossal leg of this statue is
the figure of his sister, Nefertari, who was also his favourite wife.
Hatshepsut, the Queen Elizabeth of Egypt, reserved for herself the best space in
the splendid temple-tomb at Deir-el-Bahari, tucking away in small quarters the
bodies of her male relatives. A brother later retaliated by removing her name from
the inscriptions.
Every great temple in ancient Egypt had its sacred lake, where the worshippers
performed their ablutions and the religious processions of boats took place. The
banks of this lake at Karnak were originally lined with smooth-cut stone.
To visit this valley one first comes to Luxor, which is very nearly on
the site of Old Thebes, the capital of Egypt in the days of its most
brilliant past. The ancient city lay on both sides of the Nile, but Luxor
is on the east bank. Crossing the river in a ferry boat, I rode for an
hour or more through the desert before I came into the Valley of the
Kings. My donkey boy was a good one and his donkeys were young.
His name was Joseph, and the brute I bestrode was called
“Gingerbread.”
We traversed green fields, winding in and out along the canals,
until we came to the desert and entered a gorge walled with rocks of
yellow limestone and a conglomerate mixture of flint and limestone
of curious formation. The gorge shows evidences of having been cut
out by some mighty stream of the past. There are masses of débris
along the sides, and the way is rough except on the road which has
been made by the explorers.
Looking at the valley from the Nile one would not suppose it to be
anything other than a desert ravine, so I did not at first realize that it
was a cemetery. There are neither gravestones nor monuments, for
the kings obliterated every sign that might indicate their burial
places. They dug out great chambers under the bed of this dried-up
river and built cisterns for their proper drainage, but when they had
finished they did all they could to make the spot look as it was in
nature. For this reason their tombs remained for ages untouched and
unknown.
From time to time, however, one or another was discovered.
Strabo, the Greek geographer, who was alive when Christ was born,
speaks of forty of them as being worthy of a visit, and others are
mentioned by subsequent writers. Later they were again lost, and
not until in our generation when some Arabs began to sell curious
antiquities was it learned that the tombs had been rediscovered and
were being rifled by these vandals. The archæologists then went to
work on their explorations which resulted in the opening up of tomb
after tomb, until we now have what might almost be called a
subterranean city of the dead in the heart of the desert.
The tombs are nothing like our burial vaults. They are large rooms
cut out of the solid rock, with walls straight and smooth. They are
reached by many steps, going down inclined planes until they bring
one far below the surface of the valley and deep under the
mountains. Each king had his own tomb, which he decorated with
sketches and paintings representing the life of his time and the
achievements of his reign. The ceilings are beautiful. From some of
them the figures of gods and goddesses look down upon us. Others
are decorated with geometric designs in beautiful colours. In some,
men and women are carved in bas-relief out of the solid rock and
then coloured. Many of the scenes are religious, so that from them
the Egyptologist is able to learn what the people of that day believed.
The carvings show, too, how they lived when our remotest ancestors
were savages in the wilds of Europe and Asia.
The Americans have had remarkably good luck in their finds. One
of them was the tomb of the parents of Queen Tiy in which all the
objects were in as good condition as if they had been in a house just
closed for the summer. There were armchairs beautifully carved and
decorated with gold. The cushion on one of them was stuffed with
down and covered with linen perfectly preserved. In another part of
the chamber were two beds decorated with gold, while a light chariot
stood in a corner. But most wonderful of all was the discovery in this
tomb of a jar of honey, still liquid and still fragrant after thirty-three
hundred years.
In some of the tombs I saw the massive stone boxes in which lay
the mummies of the dead kings. I measured one ten feet long, six
feet wide, and eight feet high. It was hollowed out of a block of
granite, and would weigh many tons. That mighty burial casket was
cut out of the quarries of Aswan far above here, on the banks of the
Nile. It must have been brought down the river on a barge and
carried to this place. When it was finally on the ground it had to be
lowered into the vault. All these feats were done without modern
machinery. As I went through the tombs I saw several such caskets,
and the archæologist who guided me showed me the holes in the
stone walls of the entrance ways where beams had been put across
in order that ropes might be used to prevent these stone masses
from sliding too far when let down. It is a difficult job for us to handle
safes. One of these stone boxes would weigh as much as several
safes, yet the old Egyptians moved them about as they pleased.
Indeed, I venture to say that the civil engineers of the Pharaohs
could teach us much. All through this region there are enormous
monuments which it would puzzle the engineers of to-day to handle.
For instance, there are the Colossi of Memnon, the two mammoth
stone statues that sit upon pedestals in the Nile valley within a few
miles of where I am writing. Each is as high as a six-story building,
and the stone pedestals rise thirteen feet above the ground. As I
rode by them on my way home from the Valley of the Kings I climbed
up and ran a tape measure over their legs. Each leg is nineteen feet
from sole to knee. The feet are each over three yards in length, so
long that one would fill the box of a farm wagon from end to end, and
so wide that it could hardly be fitted within it. Each arm from finger
tips to elbow measures five yards, and the middle finger of each
hand is a yard and a half long. As I stood beside the pedestal, with
my feet on Gingerbread’s saddle, I could not reach the top.
These two colossal figures sit side by side on the edge of the Nile
valley with the desert mountains at their backs. They were set up in
honour of an Egyptian king who lived more than thirty-five centuries
ago. The temple he constructed behind them has now entirely
disappeared. The statues overlook green fields, and as I gazed at
the giant shapes I thought how they had watched the people sowing
and reaping through all these centuries.
Not far from these monuments are the ruins of the temple of
Rameses II, according to some authorities the Pharaoh who “would
not let the people go.” Among them I saw the remains of a statue of
that old king, once part of a structure at least sixty feet high. There is
no granite nearer here than in the quarries of Aswan, so this mighty
monument must have been cut there and brought down the Nile to
Thebes, a distance of one hundred and thirty-five miles.
Consider the obelisks which the Egyptians made at those quarries
and carried down the Nile to Thebes, to Cairo, and to Alexandria.
There are two of them still at this place. You may see them in the
great Temple of Karnak, which is not more than a twenty-minute walk
from Luxor. They weigh something like four hundred tons each, and
if they were broken up and loaded upon wagons it would take one
thousand six hundred horses to haul them. Each is a single block of
granite, and each was carried in that shape to this place. There are
inscriptions on the Deir-el-Bahari Temple here which show that these
two shafts were dug out of the quarries, covered with hieroglyphic
carvings, brought here, and put up all in the space of seven months.
I doubt whether our engineers could do such a job as quickly or as
well.
We thought it a wonderful work to bring the Alexandria obelisk
from Egypt to New York in the hold of a steamer. To load it a hole
had to be cut in the bow of the vessel and the pillar dragged through.
The Egyptian obelisk at Paris was carried across the Mediterranean
on a barge, while that which now stands in London was taken there
in an iron watertight cylinder which was shipped to Alexandria in
pieces and built around the column as it lay upon the shore. When
the great stone was thoroughly encased, the whole was rolled into
the sea and thus towed to London. After the huge monoliths were
landed, the modern engineers had great trouble to get them where
they wanted them. The New York obelisk was rolled along upon iron
balls running in iron grooves laid down for the purpose, while that of
London was hauled over greased ways to the place where it now
stands on the banks of the Thames.
The oldest temple of Egypt by five hundred years was unearthed
here by the agents of the Egyptian Exploration Fund. This lies near
the famous temple of Deir-el-Bahari, and in a valley which is a
branch of that of the tombs of the kings. When I visited it to-day the
excavators were at work, and the men in charge told me they had
great hopes of making valuable discoveries. It was with the
American representative of the Exploration Fund, that I went over the
temple. I met him at the little one-story house which forms the
laboratory and home of the foreign explorers, and had a chat with
the other members as to the progress of the work. A number of
specialists from Canada, England, and the United States, supported
by the fund, are superintending the Egyptians, who do the hard
labour. They have quite an army of men at work and have been
successful. Of what they find one half goes to the museum at Cairo
and the rest to the countries which subscribe to the fund in
proportion to the amount of their subscriptions. The chief money
from America has come from Boston, New York, Baltimore, and
Washington, so that our share of what is now being unearthed will go
to the museums of those cities.
More famous than this ancient temple itself is its shrine of the cow
goddess, Hathor, from which the noted statue was excavated by the
Egyptian Exploration Fund and taken to Cairo. I saw the place
whence it came and talked to the men who dug it out of the earth.
The statue, which is life-sized, is a perfect likeness of a beautiful cow
carved out of stone. It is reddish-brown in colour, with spots shaped
like a four-leaved clover. Traces still remain of the gold that once
covered the head, neck, and horns. The head is crowned with lotus
flowers and lotus stalks hang down each side the neck almost to the
ground. Beneath the head stands the dead king whom Hathor
protects, while the living king, whom she nourishes, kneels beneath
her form. That image was probably worshipped at the time the
Israelites were working in the valley of the Nile, and it may have
been after one like her that they modelled their calf of gold.
Near the site of this oldest temple are the ruins of the great temple
of Hatshepsut, the Queen Elizabeth of Egypt, who ruled fifteen
hundred years before Christ was born. Her epitaph says that “Egypt
was made to labour with bowed head for her.” The temple is really a
tomb-chapel in memory of the royalties buried there—her father, her
two brothers, and herself. Hatshepsut took most of the space,
however, and put the bodies of her male relatives into as small
quarters as she could. She called her temple “most splendid of all”
and covered its walls with engravings and paintings showing her
principal acts. Hers is a long record of kingly deeds. She discarded
the dress of a woman, wore the crown, attached an artificial beard to
her chin, and let it be known that she liked to be addressed as His
Majesty by her courtiers and subjects. The New Woman is
apparently as old as civilization itself!
It was the work of Americans, again, that unearthed here the tomb
of the first great pacifist, Pharaoh Akhnaton, who reigned from 1375
to 1358 b.c. When he came to the throne Egypt, in the height of her
power, was mistress of the chief parts of the civilized world. But the
country was then ridden by the priesthood of Amon with its hosts of
gods and its degraded worship. According to the inscriptions which
have been deciphered young Akhnaton defied the priests of Amon
and declared his belief in one God, a “tender and merciful Father
and Mother of all that He had made,” the “Lord of Love,” the
“Comforter of them that weep.” It is thought that he was the Pharaoh
in Egypt when the Children of Israel came into the land and that the
One Hundred and Fourth Psalm in our Bible was written by him. He
did not believe that warfare or military conquests were consistent
with his creed and when revolts broke out in his Syrian provinces he
refused to fight, though his soldiers tried desperately hard to hold the
different people of his empire faithful to their king.
Breaking entirely with the priests, Akhnaton left Thebes and set up
his capital at Aton, one hundred and sixty miles south of Cairo on the
eastern bank of the Nile. He died at the age of twenty-eight, leaving
only daughters to succeed him. They reëstablished the court at
Thebes, the city of Aton was abandoned, and its temples and
palaces were left to crumble and decay.
I had thought of the Pharaoh who forced the Israelites to make
bricks without straw as living at Memphis, near where Cairo now
stands. The truth is, he had a great city there, but his capital and
favourite home was at Thebes, over four hundred and fifty miles
farther up the Nile valley. Thebes was one of the greatest cities of
antiquity. It covered almost as much ground as Paris does now and
is said to have had more than a million people. The metropolis had
walls so thick that chariots drawn by half-a-dozen horses abreast
could easily pass as they galloped along them. It had one hundred
gates, and temples and residences which were the wonder of the
world. Some of the houses were five stories high, the skyscrapers of
those days. The riches of Thebes were increased by the successful
wars which the kings waged with other nations. The monarchs of
that day had mighty armies of infantry and cavalry. Some of the