froG

Charlotte Sleigh

Animal series
Frog
 Animal
Series editor: Jonathan Burt

Already published
Ant Charlotte Sleigh · Ape John Sorenson · Bear Robert E. Bieder
Bee Claire Preston · Camel Robert Irwin · Cat Katharine M. Rogers
Chicken Annie Potts · Cockroach Marion Copeland · Cow Hannah Velten
Crow Boria Sax · Dog Susan McHugh · Donkey Jill Bough
Duck Victoria de Rijke · Eel Richard Schweid · Elephant Daniel Wylie
Falcon Helen Macdonald · Fly Steven Connor · Fox Martin Wallen
Frog Charlotte Sleigh · Giraffe Mark Williams · Hare Simon Carnell
Horse Elaine Walker · Hyena Mikita Brottman · Kangaroo John Simons
Lion Deirdre Jackson · Lobster Richard J. King · Moose Kevin Jackson
Mosquito Richard Jones · Otter Daniel Allen · Owl Desmond Morris
Oyster Rebecca Stott · Parrot Paul Carter · Peacock Christine E. Jackson
Penguin Stephen Martin · Pig Brett Mizelle · Pigeon Barbara Allen
Rat Jonathan Burt · Rhinoceros Kelly Enright · Salmon Peter Coates
Shark Dean Crawford · Snail Peter Williams · Snake Drake Stutesman
Sparrow Kim Todd · Spider Katja and Sergiusz Michalski · Swan Peter Young
Tiger Susie Green · Tortoise Peter Young · Trout James Owen
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Frog

Charlotte Sleigh

reaktion books
For Nick, who has no need of metamorphosing,
thank you very much

Published by
reaktion books ltd
33 Great Sutton Street
London ec1v 0dx, uk
www.reaktionbooks.co.uk

First published 2012

Copyright © Charlotte Sleigh 2012

All rights reserved

No part of this publication may be reproduced, stored in a retrieval

system or transmitted, in any form or by any means, electronic,
mechanical, photocopying, recording or otherwise without the prior
permission of the publishers.

Printed and bound in China by C&C Offset Printing Co., Ltd

British Library Cataloguing in Publication Data

Sleigh, Charlotte.
Frog. – (Animal)
1. Frogs 2. Frogs in literature.
3. Frogs as laboratory animals.
4. Frogs in art.
I. Title II. Series
597.8'9-dc23

isbn 978 1 86189 920 0

 Contents

Introduction 7
1 Just a Kiss 31
2 Warts and All 55
3 Them, Us and Frogs 77
4 Under the Knife 95
5 Evolution on Fast-forward 116
6 Of Frogs and Fruitfulness 140
7 Jumped Up 161

Timeline 186
References 188
Select Bibliography 196
Associations and Websites 199
Acknowledgements 202
Photo Acknowledgements 203
Index 205
 Introduction

Frogs are slippery in more ways than one.

Above all, they are liminal creatures: animals of the in-
between. They are the largest creatures to undergo such complete
and dramatic metamorphosis, transforming from an aquatic
to a terrestrial existence in the process. Even as adults many
species are still partially creatures of the water, but they are not
formed like other aquatic vertebrates. Apart from their close
cousins the newts, frogs are the only underwater swimmers
with legs; they are also among the very few vertebrates not to
possess tails. Add these features to their front-facing eyes, their
slim waists and hairless bodies, and you have a beast that is
curiously human in form.
It was perhaps the doubtful nature of frogs that led the
natural historian Ulisse Aldrovandi (1522–1605) to include them
so prominently in his museum of curiosities and monsters: a
frog with the tail of a lizard, and another with a full set of
teeth carefully planted inside its mouth. Cabinets such as Aldro-
vandi’s were a part of the Renaissance drive to order nature,
and frogs have remained a problematic grouping ever since, as
liminal in their classification as in their biology. A classroom
study conducted in New Zealand found that the frog was not
even necessarily an animal. Only about half of fifteen-year-olds
classed it as such; for the rest, animality was tacitly associated

7
 Like other early
moderns, Edward
Topsell classified
frogs with snakes
in his History of
Serpents (1608).

with mammalian characteristics. Being furless and aquatic, the

frog did not count.1
The challenge of frog classification is a particular favourite of
anthropologists. An influential paper of 1979 suggested that
when non-Westerners develop zoological taxonomies they get
to frogs comparatively late in the process and struggle with how
to categorize them. The paper’s model suggested that there are
three basic life forms – fish, snake and bird – that are classified
early on in the development of a language. mammal and wug
(worm and bug), it was claimed, follow on later; and frogs, when Frontispiece to
August Johan
speakers finally get round to them, have to be crow barred into Roesel von
one of these.2 However, other studies – largely conducted in Rosenhof, Historia
Naturalis Ranarum
New Guinea – have shown that frogs are a lot more significant Nostratium (1758).
to forest dwellers than these researchers anticipated, intimately The Latin motto
from Virgil
related with each language group’s specific ecology, lifestyle and
translates as ‘the
culture. frog is, in fact, often labelled before mammal or wug greatest wonder
in the development of a language, and the detailed categor- may be derived
from observation
izations of frogs produced by New Guineans can be remarkably of the slightest
close to the taxonomies generated by Western scientists.3 things’; the
crumbling monu-
One could argue that the anthropologists’ rather elementary ment, juxtaposed
and universalist model for the development of the category frog with vivacious
amphibians,
in 1979 actually captured their own heritage rather better than reasserts the point
that of the New Guineans. The anthropologists’ two options for in pictorial form.

9
This undated
engraving from
the Stuart era
underlines the
often negative
reputation of
batrachians. A
sinful courtier in
Whitehall is
compared to a
toad and found –
astonishingly – to
be the more loath-
some of the two.

including frogs, wug and mammal, were both leftover groups:

neither fish, snake nor bird. Frogs have long been just such
hard-to-fit animals for Europeans, grouped variously with
insects and serpents until early modern times. In 1608 Edward
Topsell put them with serpents: ‘By serpents we understand in
this discourse all venomous Beasts, whether creeping without
legs . . . or with legs . . . or more neerly compacted bodies, as
Toads, Spiders and Bees.’4
The frog’s appearance in a French dictionary of 1694 saw it
defined as ‘an insect that commonly lives in marshes’. The
great comparative anatomist of the nineteenth century, Georges
Cuvier (1769–1832), made the batrachians (frogs and toads)
the fourth order of reptiles; the class Reptilia remained the
taxonomic home of the frogs until the mid- to late nineteenth
century. Batrachians were defined by their deficits in relation to
the other reptiles: their lack of fully formed ribs, of scales, of a
pharynx, and of external organs of generation.
To this very day the discipline of herpetology – literally, the
study of creeping animals – retains the scientific grouping
together of reptiles and amphibians in a way that would seem

10
very odd for, say, specialists in birds and mammals. In scientific
literature and at academic conferences, frogs continue to sit
alongside lizards and snakes. This grouping reflects – even if it
no longer shares – a medieval tradition that saw these creatures
as imperfect, if not downright satanic. The frog and toad were
not so far from the devilish serpent himself.
The category ‘frog’ is itself subdivided different ways by
different cultures. Europeans are consistent in distinguishing
frogs and toads, the latter being considered to have drier,
wartier skin; to have a flatter profile; and to walk rather than
hopping as frogs do. For Topsell, the toad was the ‘most noble
kinde of Frog, most venomous and remarkable for courage and
strength’, but in general (and even elsewhere in Topsell’s History
of Serpents) Europeans have regarded toads with more disgust
and suspicion than frogs – and have maintained separate words
for them.5 The Germanic root of ‘frog’ seems to come from an
early Indo-European verb meaning ‘to hop’, while Romance
languages trace their frog names to the Latin rana (frog). Both

An early modern
English frogge:
detail of hanging
embroidered by
Mary Queen of
Scots and Bess of
Hardwick, c. 1570.

11
Frogs of the genus
Hyla, from Johann
Baptist de Spix’s
Animalia Nova sive
Species Novae
Testudinum et
Ranarum (1824).
The genus consists
entirely of tree
frogs, one of the
most widely
recognized sub-
categories of frog
around the world.

French and German trace their words for toad to a root referring
to their rough skin, although the Latin bufo (toad) is related to
a Germanic verb meaning ‘to puff ’. (The English ‘toad’ is a
mysteriously isolated linguistic stump all on its own.) However,
the derivation of rana complicates the idea of a linguistic separ-
ation of frogs and toads. Although sometimes considered to be
onomatopoeic, it is more likely a corruption of the Greek frýnos,
which means ‘toad’. In other parts of the world, where both
ecology and culture are different, the batrachians break down
in different ways. Japanese, for example, has the same root
word for both frogs and toads. Other languages have three or

12
even more basic categories, with tree frogs especially often dis-
tinguished from marshy frogs.
Current biology gives frogs the status of a taxonomic order,
the Anura. The Anura are themselves split three ways into
sub-orders that comprise 38 families in all. Every frog is an
anuran, and every anuran is a frog. Biologists do not recognize
any biological distinction between frogs and toads; most of
the anuran families contain within them some species that
are informally designated ‘frog’ and others as ‘toad’. Only the
family Bufonidae contains ‘toads’ alone. This book follows scien-
tists’ lead in treating all the anurans, frogs and toads, together.
Modern amphibians (anurans, newts and salamanders, and
primitive caecilians) seem to have developed from the
Temnospondyli, an order of primitive and sometimes gigantic
amphibia that lived approximately 250 to 300 million years
ago. However, for a long time scientists could not find an inter-
mediate organism, from which modern frogs and salamanders
diverged, among the Temnospondyli, and some suggested that
a different order altogether was likely to be their point of common
ancestry. In 2006 scientists at the University of Calgary claimed
to have settled the dispute by finding the frog and salamander’s
common ancestor in a Texan temnospondyl fossil named
Gerobatrachus hottoni (Hotton’s Old Frog). This ‘frogamander’ is,
at 11.5 cm, about the size of a modern frog. It shares features of
both modern-day frogs and salamanders, although at least one
specialist in amphibian fossils has cautioned that it may be
rather too close to a modern frog to count as a true ‘missing link’.
G. hottoni is about 300 million years old, and if it is an ancestor
of both salamanders and frogs, then it suggests that the two
orders separated about 240 to 275 million years ago. If, as most
seem to agree, this is the case, then it leaves another unsolved
puzzle, since it represents a much more recent evolutionary

13
 divergence than the date suggested by molecular drift (a meas-
ure of ancestral similarity that is produced by the dna-copying
errors that accumulate naturally over time). This disagreement
will need to be reconciled in some way. Meanwhile, the earliest
generally agreed true frog, a partial fossil from Argentina, is
roughly 200 million years old. It has been designated as Vieraella
herbsti, and is very small – about 33 mm from top to rump.
Since the Jurassic period, frogs have colonized most of the
Earth. At present over 6,000 extant species are known, slightly
more than the maximum (human) capacity of the Albert Hall,
or twice that of Carnegie Hall. Around 800 new species have
been described in the last five years alone. The only places not
inhabited by frogs are the globe’s extreme north and south, and
the hottest desert areas of Africa and Arabia. Remote oceanic
islands are also uncolonized by anurans.

Fossilized frog
of the genus
Palaeobatrachus
from the Tertiary
period (130–35
million years ago).

14
 The water holding
frog (Litoria
platycephala)
of Australia
can outlast long
periods of drought.

Despite their association with water, anurans are able to

live in some remarkably dry environments. The water holding
frog, Litoria platycephala, inhabits some of the driest and most
inhospitable areas of Australia. It survives underground, buried
alone with nothing but a bladder full of water to keep it going.
In the darkness of the earth it accumulates a cocoon of sloughed-
off skin that becomes almost waterproof, reducing the amount
of energy that it uses as well as retaining moisture. When rain
comes, the frogs emerge to feed and breed. Ideally the rains
are an annual event, but if necessary L. platycephala can survive
inside the earth for up to five years. They lay their eggs in the
temporary puddles produced by rain, and like those of other dry-
living species, their tadpoles develop very quickly to beat the
returning drought. Similarly, spadefoot toads of the southern us
(family Scaphiopodidae) have extraordinarily quick develop-
ment times, as little as seven days from larva to toadlet. No one

15
A Couch’s
spadefoot toad
(Scaphiopus
couchii) of the
southern us; its
extraordinarily
quick maturation
from tadpole to
frog enables it to
beat fast-drying
conditions.

The North
American wood
frog (Rana sylvatica)
survives sub-zero
conditions by
allowing itself
partially to freeze.
knows how the tiny toads migrate and survive the extreme heat
of the Californian desert after their puddles disappear.
By contrast, the widely distributed North American wood
frog, Rana sylvatica, can survive sub-zero temperatures. Its abil-
ity to hibernate is nothing special among frogs, but uniquely it
is able to tolerate the partial freezing (up to about one-third)
of its own body in the process. It uses urea and specially made
glucose to reduce the propensity of its body fluid to crystallize,
thus protecting its cells from rupture.
Unlike model children, frogs are often heard but not seen.
Researchers working with the Karam of the New Guinea
Highlands noted that frogs were ‘extremely numerous . . . and
almost constantly heard’ in their environment. In this context,
frogs become integral to the human soundscape. When frogs
begin to pipe in the evening, the Karam say ‘gl agl agp en amnwno!’
– ‘when the nyingle-nyangle calls it’s time to go home’.6 Aris-
tophanes exploited the frogs’ sound to create a novel chorus in
his play of that name. ‘Brekekekéx-koáx-koáx’ they chant as
Dionysus rows across to Hades. The only way Dionysus can
deal with his aural irritation is to join in. The nature writer
John Burroughs, by contrast, thought the sound a lovely one,
auguring the close of winter. ‘Blessings on thy warty head:/
No bird could do it better’, he concluded in ‘The Song of the
Toad’ (1906).
In many languages, the names of frogs are onomatopoeic:
even the English term, when it has its original final syllable
restored – frogga – sounds distinctly batrachian. Ouauouarons
are Cajun bullfrogs, and Aramaic dialects render frogs as aqruqe
or aqruqta; the list of frog-sounding frogs goes on and on.
Frogs almost always make their characteristic species sound
– from a peep to a deep burp – for reasons that have to do
with reproduction. The commonest of these is the male’s call to

17
 attract a mate, often performed en masse. These calls can be heard
up to a mile away; the frog’s vocal sac expands like bubblegum
to amplify the sound. Other common calls are made with the
mouth closed: ‘get off!’ (when a male is accidentally mounted by
another male); ‘go away!’ (when territory is infringed); and ‘eek!’
– a distress call emitted when a frog is taken by a predator, and
which may possibly startle or distract the latter into letting go.
The order Anura used to be called the Salienta in honour of
frogs’ other noted characteristic, their capacity to jump. The
name was a good one, because even species that have lost the
ability (or need) for this form of locomotion retain the morph-
ological features that make it possible. The bones of the hind-
and fore legs, separate in other vertebrates, are fused in the frog
for extra strength, both in pushing off and in absorbing the
shock of landing. The muscles associated with these bones have
become hypertrophied through evolution, and ongoing research
is exploring the possibility that the tendons are also used like

The Australian
rocket frog (Litoria
nasuta) is named
in honour of its
projectile power.

18
 The giant leaf frog
(Phyllomedusa
bicolor) is among
those with the
ability to adhere to
trees and leaves.
The eggs are
placed in a leaf
nest high above
the ground; after
8–10 days the
newly hatched
tadpoles fall into
the pond below.
From J. G. Wood’s
Illustrated Natural
History: Reptiles,
Fishes, Molluscs
(1863).

springs to store and release energy for jumping. The Australian

rocket frog, Litoria nasuta, seems to hold the record in this
respect, achieving distances of over two metres. This might not
seem so much until one considers that the frog itself is only 5.5
cm long.
Underwater, such leg movements are translated into a kick,
generally supplemented by foot webbing that increases push-off
against the water. Tree frogs, however, have different require-
ments, needing to stick to leaves and stems as they make their
ascent through the forest canopy. The Australian green tree
frog, Litoria caerulea, makes the nature of the challenge clear;
it is not a tiny, delicate species like so many tree frogs but rather
a dumpy and large one, about 10 cm long. Its gravity-defying

19
behaviour is therefore especially in need of explanation. Most
tree frogs have a chunky jigsaw of cells on their toe pads which
can be wedged into the microscopic chinks of the surface of
ascent, a kind of cellular rock-climbing. In addition, frogs produce
mucus that sticks their skin to the plant by adhesive molecular
forces, just like a wet tissue on glass. Tree frogs constantly repos-
ition their toes to maximize the mucus contact between skin
and plant, but – surprisingly – when touched they never feel
sticky. Under a powerful microscope the apparently flat cells of
L. caerulea are revealed to be covered in tightly packed ‘nano-
pillars’, each with a small dimple in the end. No one yet knows
the exact function of the nanopillars, but they may increase the
surface area for adhesion and provide friction against slippage.
Besides the whole-body jump, the other famously fast part
of the frog is the tongue, flipping dramatically out of its mouth
to capture a passing fly while the frog itself sits in nonchalant
immobility. Different species use different mechanisms to pull
the trick off. Some actually shorten their tongue as it is pro-
tracted, while others lengthen it through the inertial or muscular
forces associated with the uncoiling action. Unlike other species
which unfurl their tongues, the African pig-nosed frog, Hemisus
marmoratus, telescopes its tongue outwards, doubling its length
in the process. Lobes on the tongue’s tip grip the prey in pre-
hensile fashion.
The frog’s tongue grabs insects and other invertebrates.
Larger species will also take eggs, small vertebrates and even
other frogs; only a very small handful of frogs are herbivorous.
Tadpoles, on the other hand, have complex mouthparts to allow
their specialist filter-feeding on algae. Some tadpoles are also
cannibalistic, and contrary to what one might imagine it is often
the faster developers that get gobbled up by those still in the
larval stage.

20
 An illustration of
the frog’s fast
tongue in action,
from Roesel von
Rosenhof’s Historia
Naturalis Ranarum
Nostratium (1758).

Tadpoles, as is well-known, have gills to breathe as they

develop. Adult frogs have very simple lungs, with no diaphragm
or rib attachment of pulmonary muscles to assist their intake of
air. One species, Barbourula kalimantanensis, does without lungs
altogether, breathing entirely as other species do partly: through
its skin. In order to absorb oxygen, frogs’ skin must be kept
moist; as an unavoidable side effect it is permeable to many
other molecules and must be kept away from salts. Yet despite
their inefficient uptake of oxygen, frogs are lively and energetic
creatures, qualities that usually depend upon having a lot of
oxygen to burn. They manage this by having a strict separation
of oxygenated and deoxygenated blood in the heart and around
the body, so that what oxygen they do have does not simply dif-
fuse away to oxygen-poor tissue.

21
The Australian The skin of different frog species also contains a vast range of
green tree frog
(Litoria caerulea)
pharmacologically significant compounds, whether manufac-
is one of many tured by the frogs themselves or ingested in their diet (the
frogs from which scientific jury is still out). The family Bufonidae manufactures
pharmacologically
active compounds poison in the parotoid glands just beneath the eyes, but most
have been frogs produce their chemicals in smaller glands distributed
obtained – in
this case, thanks across their backs. Since the 1960s there has been a scientific rush
to a vomiting cat. to find, understand, synthesize and patent these compounds.
The poisonous One of the earliest scientific investigations on batrachian phar-
Southern
Corroboree frog
macology was prompted by a cat. Robert Endean, an expert on
(Pseudophryne the toxins of marine organisms, noticed that his cat enjoyed
corroboree) is
eating frogs but always vomited after eating the species Litoria
one of the most
endangered caerulea. After extracting the relevant compound from dried
native animals frogs’ skins, it was found to be a polypeptide that could produce
in Australia.
a significant and sustained fall in blood pressure, but unfor-
tunately, as the cat could have told them, it also had the side
effect of producing vomiting and diarrhoea. The team named the
compound caerulein for the frog; science has not recorded the
The most potent
of all known toxic name of the cat. Caerulein is now used to stimulate gut activity
frogs: the golden where this property has been damaged, and there are also
poison frog (Phyllo-
bates terribilis) of moves afoot to use it in the treatment of schizophrenia, since a
Colombia. related compound appears to be involved in the neurochemistry

22
of anxiety. Other frog compounds under investigation include
antibiotics, hallucinogens, anti-tumour agents, anti-inflamma-
tories, analgesics, adhesives, spermicides and mosquito repellents.
These days, scientists are able to extract these chemicals without
killing the frog. Instead, electricity is used to stimulate secretion,
but the rarity of many of the species involved makes even their
temporary capture an ethically problematic action.
Coloured frogs typically advertise (or in a few cases, bluff )
their poisonous properties so that predators leave them alone.

24
Alas for the frogs, however, some predators have developed Another species of
highly toxic frog:
tolerance of even their worst weaponry. Humans are not among the dyeing dart
these tough predators, and members of the family Dendro- frog, Dendrobates
batidae of Central and South America are collectively known tinctorius of north-
eastern South
as poison dart frogs for their use by indigenous peoples. Of America. This
these, the golden poison frog, Phyllobates terribilis, is the most blue morph was
previously thought
toxic. It is enough for the Chocó and Cofán Colombians to rub to be a species
their dart tips against the animal’s skin to co-opt its neurotoxic in its own right,
but D. tinctorius
properties to lethal effect, stopping the heart of their victim. It is now agreed to
has been estimated that the chemicals contained within a single be polymorphous
(having more than
frog would be enough to kill ten adult humans, but the toxin may one colour variant).
also have potential as a topical painkiller if used correctly.
It has served Colombian indigenes well to know of their
batrachian neighbours’ toxicity, and frogs are in fact generally
well-understood around the world. Their metamorphic life cycle
is known by most, perhaps all, cultures that are in contact with
them. Tadpoles often have affectionate names of their own; in
the us and to a lesser extent the uk they are known as pollywogs
(or head-wiggles).
The frog’s existence is most precarious at the tadpole stage
of life. Defenceless and comparatively slow-moving, it, along
with its swarm of siblings, is easily hoovered up by predators.
Accordingly, frogs have evolved a variety of methods to try and
better the chances of their offspring. One of these is sheer
numbers: the more tadpoles, the greater the chance of one
sur viving to replace its parent. However, many species have
instead evolved specialist care behaviours in both male and
female parents. Females of the now-extinct gastric-brooding
frogs of Australia (genus Rheobatrachus) managed to suppress
the motions and digestive fluids of their stomachs, swallowing
their tadpoles and allowing them to develop inside until they
were ready to be spat out. The Chilean Darwin’s frog (Rhinoderma

25
 darwinii) performs a similar feat using its vocal sac. The male
‘eats’ the eggs when they are close to hatching, and keeps them
in the sac for up to 70 days, nourishing them with viscous secre-
tions. Other species develop sacs in the skin along their side or
back in which to shelter their young. The fully aquatic Surinam
toad (Pipa pipa), for example, performs a whirling dance of
copulation in the water so that the male can press the fertilized
eggs into the female’s back. Here they sink into the skin, emerging
three to four months later as toadlets. Male toads of the genus
Alytes do not sacrifice their complexion in this way, but tangle
the egg strings around their feet and take regular dips to keep
them moist.
Many species will transport their young to good, watery
locations, either because the area is otherwise dry and water-
transient, or because it is advantageous to find a pool with no
predators present. In the latter case, the pool (typically a pocket
of water inside a plant) is also free of nutrients for the tadpole,
and so food must be provided. The strawberry poison frog
(Dendrobates pumilio) of Central America is one species that

A Lehmann’s
poison frog
(Dendrobates
lehmanni) at Cali
Zoo, Valle del
Cauca, Colombia.

26
 The Surinam toad,
now known as Pipa
pipa, shown here
in J. G. Wood’s
Illustrated Natural
History: Reptiles,
Fishes, Molluscs
(1863).

treats its offspring in this manner. After the eggs are fertilized,
the D. pumilio male tends the clutch on land, periodically urinat-
ing on them to keep them moist. When they have hatched, the
female carries the tadpoles one by one to pockets of water in the
forest plants, laying unfertilized eggs for them to eat. Up to six
tadpoles per brood can be successfully raised to maturity in this
way by a diligent pair of parents.
Tadpoles grow and develop gradually, but the final stage of
metamorphosis to young frog is astonishingly quick and total,
as little as 24 hours. The totality and co-ordination of the change
is necessary to prevent the dangerous situation of the tadpole-
frog being stranded in between forms, only half-adapted to its
environment and helpless to the forces of natural selection. The
filtering, scraping mouthparts must be transformed into preda-
tory jaws, and the long, spiral gut necessary to digest algae gives

27
An early from-life
illustration of
a frog: Jan Ven
Kessel the Elder
(1626–79), Dead
Tree Frog.

way to a predator’s short digestive tract. The skin thickens and

develops glands to keep the frog moist on land. Lungs are
present from early in the tadpole’s life, but the pouch that pre-
viously contained the jaws must make way for the front legs.
Nerves and senses all undergo dramatic change; the eyes
develop massively to detect moving prey, and eyelids form to
protect the eyes on land. Only the tail remains for a while as a
sign of the frog’s former life, physiologically unimportant com-

28
pared to other changes and eventually reabsorbed by the body.
Metamorphosis is a remarkable feat, as yet only partially under-
stood by scientists.
It is no wonder that philosophers, natural historians and
scientists have long been intrigued by batrachians.7 The study
of frogs in the seventeenth and eighteenth centuries can be
classed as natural history; comparative anatomy – a major
component of evolutionary study – informed frog research in
the late eighteenth and nineteenth centuries. Frogs went on to
become subjects for physiologists and developmental biolo-
gists in the nineteenth and twentieth centuries respectively,
and today the majority of herpetological publications concern
anuran ecology.

An unidentified
blue and yellow
frog from an early
herpetological
work: Joannes
Albertus Schlosser,
De Lacerta
Amboinensi (1768).

29
 Even within science, the frog is a creature of mythological
dimensions. The frog on the scientist’s bench is not a neutral tool
of investigation, but comes sticky with culture: with the qualities
and myths that are attributed to it. Such connotations started
out in the early modern period as co-extensive with the myths of
general knowledge, but as time went on science evolved a myth-
ology of its own for the frog. Thus, as the following chapters
recount, the frog has been a scientific embodiment of theologi-
cal humility; a bundle of nerves; a bag of hormones; and an
exemplar of evolution’s extraordinary force. In the laboratory, it
has been a creature both of vivaciousness and death.
The psychoanalyst Ernest Jones noted that the items most sig-
nificant in myth and the unconscious often come in ambivalent
form, auguring at once opposite qualities and significances.8 So
it is with the frog. The frog, as we shall see, stands for fecundity
and sterility; for laughter and loss; for fortune and hopelessness;
and for enlightenment and devilry.
The frog slips continually from our grasp, moving between
its twin realms and trailing our fantasies in its watery wake.

30
1 Just a Kiss

Who would not desire the frog’s powers of total transforma-

tion? Who would not wish to metamorphose from the realm of
frustration and subordination to another world altogether: from
the murky pond to the clear, bright air?
Hindu philosophical teaching in the Upanishad scriptures is
one of the most ancient textual sources to suggest that frogs have
something to teach us about metamorphosis of the self. The
Mandukya Upanishad, also known as ‘the frog’, describes attain-
ment of the very highest state of transcendence and the mystic
syllable ‘aum’. Some of the Mandukya’s many devotees explain
its name in relation to anuran locomotion – jumping, not walk-
ing. This scripture, they say, teaches that spiritual transition from
one state of consciousness to another can only be achieved as a
complete leap, like a frog’s. Other commentators have instead
likened the stillness of the yogi to that of the frog, which at rest
manifests its vitality only by the gentle, rhythmic movements of
its throat. Either way, the frog is co-opted by the human medita-
tor as an agent of metamorphosis.
Bufotenine may be a quicker route to enlightenment than
meditation, but it too comes from frogs; it is the chemical res-
ponsible for making you high when you smoke a toad.1 There
was a flurry of interest in the drug during the 1990s, with users
claiming that it gave a gentle hallucinogenic rush and law-makers

31
declaring its dangers. In the southwestern states of the us, users Illustration
of tadpole
were apparently milking the glands of the Sonoran desert toad, metamorphosis
Bufo alvarius, for their secretions and drying them on car from Roesel von
windscreens prior to smoking them in a pipe. (Most of this Rosenhof’s Historia
Naturalis Ranarum
information, however, came from a single, somewhat unreli- Nostratium (1758).
able, source: a Californian arrested by narcotics agents together
with his four toads, Hans, Franz, Peter and Brian.) Newspaper
articles at the time reported that people were licking toads to get
high, but this was apparently dangerous as well as inaccurate;
besides bufotenine, the glands also contain toxic substances
that can only be safely destroyed by smoking. In Queensland,
Australia, the authorities responded to the menace by declaring
bufotenine a controlled substance and banning the possession
of toad slime.
As frogs move from water to earth, so they perhaps signal
the human possibility of moving from earth to air: in short,

The cane toad

(Bufo marinus) has
been suspected
of being used for
narcotic purposes
in Queensland,
Australia.

33
 resurrection. The earliest Christians in Egypt used the frog as
a symbol of resurrection, their icon referencing the local frog
goddess Heket, who helped Osiris rise from the dead. The early
modern naturalist Jan Swammerdam also offered a reflection
on the similarity of frog and human transformation, albeit in
more doleful vein:

A frog eyes up a The infant-man, who lived before in the water of the
trompe l’oeil fly for
consumption in amnion, now breathes the vital air, which rushes into
this study by Hans his lungs, and dilates and extends them. But . . . his
Hoffman, late
16th century, oil
appendage of misfortunes and trouble, like the tail of
on parchment. the frog, yet adheres for a long time to him, for he is full

34
 Ed Hill, ‘Ah,
You Dear Toad,
I Will Have a Kiss’,
20th century, print.

of misery, and is born in tears; and it is very long before

he comes to maturity of understanding, and full growth
of body.2

The remainder of this chapter discusses in more detail two

important transformations mediated by the frog in philosophy
and folklore: from lead to gold, and from brute to royalty. If we
could but hitch a ride with the frog, we could master the trans-
formative secrets of alchemy or the kiss that yields a prince.
For hundreds of years, metamorphosis was above all studied
in the guise of alchemy. Practitioners of the secret art sought
the elixir that would turn base metal into gold and bring health
and long life to its discoverer. Frogs, the premier animals of
metamorphosis, were never far away, and the toad was a major
element of alchemy’s iconography, featuring on the title page of
Elias Ashmole’s influential Theatrum Chemicum Britannicum
(1652). ‘The Vision’, a poem attributed to the revered Yorkshire
alchemist George Ripley (1415–1490), similarly emphasized the

35
batrachian element of alchemy in late medieval and early
modern Europe. The poem begins:

A Toade full rudde I saw did drinke the juce of grapes so

fast,
Till over charged with the broth, his bowells all to brast
[burst].

The historian Jennifer Rampling demonstrates that Samuel

Norton, who first discovered and translated the poem from Latin
in 1573, glossed it as a fairly straightforward chemical recipe. 3
According to Norton, the ruddy toad stood for red lead, which,
placed in strong wine vinegar, would absorb the liquid until it
became reduced to prime matter. This prima materia was the
fundamental, malleable stuff of creation, worked upon and
perfected by alchemists just as Christ had perfected the human
soul. The toad, turning into the prima materia, was the very origin
of the alchemical process of metamorphosis.
Many alchemists shared Ripley’s (or Norton’s) association
of the toad with the first stage of the alchemical process. One
reason for the association was the appearance of the chem-
istry. Seething and digesting in the flask, the matter appeared
black; and black, in terms of animal symbolism, was a toad. The
alchemical toad’s connection with putrefaction, sin (especially
female) and blackness all indicated the fallen state of matter
that must be perfected or saved by the alchemist.
During the late sixteenth century, Ripley’s poetic writings
on alchemy gave rise to around twenty illustrated scrolls. The
scrolls, in which Ripley himself had no involvement, were sub-
stantial pieces, up to six metres long, expensive and precious;
one was commissioned by the well-known polymath and
occultist John Dee. The Ripley scrolls are all slightly different,

36
 This detail from
a Ripley Scroll
(late 16th century)
shows a toad
in the centre of
the alchemical
alembic, and also
at the top right
of its smaller,
contained
roundels.

complex in their iconology, and comprise references to multiple

sources. One common feature, however, is that toads appear
frequently and prominently.
On a typical Ripley scroll the first depiction of the black toad
comes at the top, contained within a hermetic vessel apparently
held by Hermes Trismegestus himself, the mythical father of
alchemy. Beneath this toad, within the vessel, are eight roundels
linked to a central circle, each showing alchemists producing
homunculi (miniature humanoids) in their vessels. The roundel
in the one o’clock position, labelled prima materia, contains an
Adam and Eve-like pair of figures, related to the Sun and

37
Moon; a very small toad can just be made out on the belly of the
female Luna.
The generation of prima materia is retold lower down the
manuscript, where a woman (labelled ‘spiritus’) holds onto a
male child (‘anima’) who is almost falling from her. What is
most striking about this woman is the green and tail-like thing
emerging from between her buttocks. Scholars of the scroll have
described this as a dragon’s tail and it certainly does resemble
the tail of the dragon further down the scroll. The image likely
refers in part to the legend of Melusine, the woman whose lower
half sometimes turned into a snake’s, and who eventually trans-
formed completely into a dragon. Although Rampling is sceptical
that the similarity was intended,4 the dragon’s tail also looks
strikingly like a ribbon of toadspawn being laid. Given the
medieval slippage between serpents and batrachians, the con-
nection would not seem impossible; indeed, the webbed feet of
the woman might suggest it. A second verse on the scroll (also
ascribed to Ripley) likewise suggests that the outcome of the
mystical alchemical union, via a serpent form, is a toad:

Then Earth on Fire shalbe put,

And Water with Air shalbe knit,
Thus ye shall go to Putrefaccion,
And bring the Serpent to reduction.
First he shalbe Black as any Crow,
And downe in his Den shall lye full lowe:
Iswel’d as a Toade that lyeth on ground . . .

Further still down the scroll, there is a dragon spitting out a

toad (although in some other scrolls it looks more as though the
toad is being swallowed). This toad is labelled with the caption
‘the tayming venome’, which associates the toad not just with

38
the first stage, but also with the whole narrative trajectory of
‘The Vision’. Although Ripley’s poem is in part a practical recipe,
there is also a strong Eucharistic tinge to it, focused by the toad.
Upon receiving wine, the toad’s sin-like poison causes him grief
and pain, and is then purged from his body. After this, the toad
turns black and dies in his cave and, akin to the phenomenon
of resurrection, goes through a series of colour changes as part
of the alchemical process. Ripley claims to extract a venom at the
end of this procedure which, like God’s judgement, can either
kill or save depending on circumstances.
This secret treasure, contained in the toad and extracted by
the magus, bore considerable resemblance to a trope of general
folklore: the precious stone that was thought to be concealed
within the heads of old toads. Variously described as a jewel
or a panacea, the stone is mentioned by Shakespeare in As You
Like It:

Sweet are the uses of adversity;

Which like the toad, ugly and venomous,
Wears yet a precious jewel in his head.

Edward Fenton’s Secrete Wonders of Nature (1569) was among

many sources to agree that ‘there is founde in the heades of old
and great toades, a stone which they call Borax or Stelon . . . of
power to repulse poysons’. Intriguingly, Chinese mythology
includes a similar fable; the three-legged toad of the holy man
Liu Hai produced a pearl in the night which, if ingested, would
turn a man into a saint. The power of the toad stone was remark-
ably like the power of the philosopher’s stone, though both,
alas, were sadly elusive.
Just as the toadish prima materia contained a nugget of per-
fection, so alchemists figured part of their process as an allegory

39
The Grimms’
Frog King story is
much celebrated,
including on this
1966 German
Bundespost stamp.

in which the King’s son went down into the sea, but retained a
spark within him and was restored to life. ‘Be turned unto me
with all your heart and do not cast me aside because I am black
and swarthy . . . and the waters have covered my face’, begins
one such parable, aligning the King’s son with both the colour
and the aqueous setting of the toad. Another account promises:
‘Whosoever will free me from the waters and lead me to dry
land, him will I prosper with everlasting riches.’5 Such parables
are tantalizingly close to tales of ‘The Frog King’.
No one knows the age or provenance of the frog king tale
before it was collected and written down by the Brothers Grimm.
Sometimes a king, sometimes a prince, the batrachian of their
story is now a staple of childhood fiction and popular culture.
Traditionally, it is the very first of the tales in the Grimms’ col-
lection. One might not think that the story requires repeating,
but its plot, as it first appeared to English-speaking audiences
in 1823, may not be quite as is commonly remembered. A young
princess drops her golden ball into a deep well, from which a
frog promises to retrieve it on condition that she grants him

40
favours – to let him live with her, to eat from her plate – and
culminating with the right to sleep in her bed. Thinking that
he will not be able to leave the well to claim his reward, she
assents, only to have him turn up in the middle of dinner and
remind her of her promise. The girl’s father, the King, urges her
to keep her word. Thus she goes through the list of actions,
and is relieved to find the frog gone in the morning. The same
thing happens the second night, and the third, and on the third

Arthur Rackham,
illustration to the
Brothers Grimm’s
‘Frog Prince’, 1909.

41
 morning she awakes to find a handsome prince at the foot of her
bed, who reveals that he had been enchanted by a spiteful fairy,
but that her actions have now undone the spell. And off they go
to live happily ever after.
One immediately notices a striking fact: contrary to popular
belief, the princess does not kiss the frog. In the initially unpub-
lished transcription of 1810, there is another variant. The princess
gingerly carries the frog up to her room on the first and only
night of his visitation, and puts him in the corner of her room.
When he insists on being put in her bed, she finally picks him
up and flings him against the wall in disgust; when he falls
down, he has made his transformation to handsome prince. It’s

An early
20th-century
German postcard
illustrating a
scene from
‘The Frog King’.

42
 The Frog King is an
international fable,
commemorated in
its gender-inverted
Russian form as the
Tsarevna Frog, or
Frog Princess.

a long way from a kiss. In the first published version of the tale
(1812), the princess’s violent action is removed by the Grimms,
and replaced with the three-night visitation better known in
English; however, later published versions in German and
English reverted to the frog being flung against the wall.
It would be futile to seek for a truly original version of the
tale. The Brothers Grimm collected two similar tales which were
conflated by the first English translator, and numerous variants
have since been gathered from around the world. Most feature
not a princess but an ordinary girl from a poor family; a Breton
version has, instead of a girl, a poor lad who is importuned by a

43
female frog. In many versions, the girl goes to get not a golden
ball but a drink from the well; often, she has two sisters who
reject the water that has been made murky by the frog. Several
accounts tell how the girl has been sent to collect water in a sieve
from the ‘well of the world’s end’; the frog’s demands come in
return for giving her the handy hint that she should first line
the sieve with moss. In tales from Scotland and the north of
England, the frog begs to be put out of his misery if he cannot
share the girl’s bed, and it is decapitation that breaks the spell.6
One problem with all these stories is that they were gathered
after the Brothers Grimm had published theirs. Indeed, the
Grimms’ book must have been in large part what inspired folk-
lorists to collect them. Surely the writers, and perhaps even the
tellers, had their tales inflected by the Grimms’. Inspired by the
story, they were on the look-out for frogs; they may have been
more inclined to hear the frog elements in other folklore, and
to bend frog tales to the plot they had already read.
A Sri Lankan story, gathered in 1910, is nevertheless quite
different from The Frog King, and does not suit modern Euro-
pean plot expectations at all. A widow bears a frog for a son,
and this frog goes off to retrieve the jewelled golden cock from
a local ogress, for which feat the King has promised half his
kingdom. Setting out on his quest, the frog turns into a hand-
some man, and armed with magic charms from three other
nearby kings, he gets the cock and defeats both the ogress and
her daughter. Returning home, he turns back into a frog and the
jewelled cock disappears. Understandably disappointed at this
turn of events – everything seemed to be going so well! – he
expires from grief.7
A Korean tale follows the European pattern more closely; an
elderly couple adopt a giant frog who brings them great wealth;
he demands to be married to a powerful neighbour’s beautiful

44
daughter, which feat at last they pull off, thanks in part to the
tradition of covering a bride’s eyes until the wedding night. At
this point, the bride takes a knife to the frog, ostensibly at his own
request but not without a certain willingness too. She thereby
releases a silk-clad prince, son of the King of the Stars.8 In a
lengthy Chinese story, there is a nice twist on the Grimms’ tale:
the frog prince finally tricks the Emperor – who would not keep
his promise to let his daughter marry him – into taking on his
frog skin. The former frog lives on as a handsome man, now
Emperor in his father-in-law’s place.9
Despite the Grimms’ claims to write things down just as
they had been told them, ‘The Frog King’ underwent ideological
changes and expansion in their successive revisions over several
decades. Since then, the story has continued to go through all
sorts of fashions and phases in its interpretation. Jack Zipes
writes confidently that the tale ‘underline[s] morals in keeping
with the Protestant ethic and a patriarchal notion of sex roles’.10
‘The male is [their] reward’, Zipes comments of the Grimms’
rare heroines, highlighting how the princess settles down to a
life of domesticity under male governance. Similarly, the folk-
lorist Lutz Röhrich points out how later Grimm versions make
more of the girl’s father’s authority in forcing her to accede to the
frog’s demands. Feminists have understandably taken issue
with this, just as they have with the supposed happy ending of
the tale. Who wants to give up playing with a golden ball and
become a wifely possession of a king, however handsome? One
jokey re-take on the story thus culminates instead with the girl,
proudly single, enjoying sautéed frogs’ legs. Although feminist
critiques of the patriarchal father are in one sense valid, patri-
archy does not appear to be central to early nineteenth-century
versions of the tale, because it is usually a mother or widow who
insists on the girl’s acquiescence, out of a sense of honour in

45
sticking to one’s word. Women, of course, reinforce patriarchy
just as much as men, but in this nineteenth-century tale at least
the frog’s ‘frogness’ trumped its masculinity; it was not to be
obeyed on the strength of the latter quality.
Just as the king becomes more insistent in some versions of
the tale, so does the princess become correspondingly more
resistant. Most critics of recent years are agreed on the sexual
element of the tale that is indicated by this resistance: that it
portrays ‘crises related to thoughts of marrying or the wedding
night’.11 The princess is a young woman on the verge of sexual
maturity and marriage, and the frog’s requests escalate in intimacy
to the unmistakeable level of sharing her bed. In fact, the pre-
publication version of the Grimms’ tale had only the explicitly
sexual request to sleep with her; the brothers appear to have
inserted the other two to tone things down. (The psychoanalyst
Bruno Bettelheim cites a still earlier version in which the girl
must kiss the frog while it lies by her side in bed, and then sleeps
with him for three whole weeks before his transformation.)
Having taken out the violence in the first published version –
replacing the wall-throwing with three nights of bed-sharing
– the brothers appeared to have thought better of the latter and
reverted to the violence as morally preferable to sex.
These changes reflect an omnipresent anxiety about the
nature of ‘The Frog Prince’; it has frequently been described as
an erotic tale that is not really meant for children at all. The
psychoanalyst Ernest Jones offered the most famous and prob-
ably the original such interpretation in 1928:

[T]he frog is in the unconscious a constant symbol of the

male organ when viewed with disgust. So . . . the story
represents the maiden’s gradual overcoming of her aver-
sion to intimacy with this part of the body.12

46
 The critic Karen Rowe goes one stage further, explaining
animal bridegrooms as a manifestation of the Electra complex
(an alternative version of Freud’s Oedipus, in which the girl
desires her father). According to Rowe, the animal plays a sub-
stitute’s role within the tale, onto which the girl’s unspeakable
love may be safely displaced.13 If Ernest Jones had written more
than a single paragraph on the story, he might perhaps have
gone in the same direction as Rowe. Jones emphasizes elsewhere
that hollow items in stories are images of the vagina or womb,
but does not dwell on the fact – surprisingly, considering his
interpretative framework – that the frog is already in the well
at the beginning of the tale. Whose is the well? Could it be the
mother’s? Could discovery of the parents’ sexual intimacy, and
jealousy of it disguised as horror, be the prompt that sets the girl
on her quest? If only Jones had spent more time on the story.
Bruno Bettelheim (building on the work of Joseph Campbell
and Carl Jung) was comfortable both with the sexual nature of
the tale and its childish audience.14 To him, fairy tales in general,
and tales of animal bridegrooms in particular, were a healthy
way of helping children work through issues that could not
and should not be presented to them overtly. Repression was
less pathological for Bettelheim than for Freud: it was simply a
normal life-stage that needed to be out-grown. Fairy tales were,
for him, a means for the child to do this out-growing: a way to
learn about the stages of selfhood, identity, maturity before they
were encountered.
Bettelheim is one of very few critics (apart from Jones and
his confident aside) who engages the question of why a frog in
particular works so well as a bestial groom. For one thing, the
transformative nature of the frog’s life cycle echoes his child-
to-adult treatment of fairy tales. More importantly, however,
Bettelheim concludes that the frog is distinguished as an animal

47
that inspires disgust but not genuine fear – that is, fear of the
truly life-threatening. As the Grimms wrote: ‘She was afraid of
the cold frog, which she did not like to touch, and which was now
to sleep in her pretty, clean little bed.’ A Bantu fable cited by
Bettelheim has a girl kiss a crocodile to effect his transformation
– an entirely more challenging proposition. Earlier versions of
‘The Frog King’, in which the creature is a snake or a dragon, have
indeed slipped from the collective memory, suggesting that these
more fearsome creatures no longer suit our post-medieval con-
cerns about sexuality. (Such frightening creatures were generally
princesses in disguise rather than princes, in keeping with the
medieval identification of sexual sin with womankind.)15 Focusing
the signified of batrachian disgust still more specifically, Bettel-
heim notes the phallic propensity of the frog to ‘puff up’.
In composing the song ‘Kiss That Frog’ (1993), Peter Gabriel
seems to have read Bettelheim, or else has pulled the erotic themes
of the frog from the communal ether. As far I know, he is the
only person to put together the kissing of popular myth with the
phallic frog of academic criticism, working them in the form of a
sexually knowing song that is at once funny and seductive. ‘He’s
all puffed up . . . kiss it better . . .’. Contributors to an online dis-
cussion board on the topic (songfacts.com) are mostly persuaded
that the song is at least partly about oral sex, except for one
poster from Massachusetts who insists against all interpretative
odds: ‘After Listening to it thouroly I Beleave its About Princess
Dianna and Prince Charles and how she was so beautiful and he
Ugly and mean to her [sic]’.
Bettelheim’s pedagogical role for ‘The Frog King’ is by no
means unique. Educators and agitators of all sorts have re-
written and re-interpreted the story to suit their ends. One does
not fully appreciate the Grimms’ lightness of touch until one
reads the well-meaning efforts of many of these imitators. Geoff

48
Dench, for example, uses the story to interrogate the ‘problem
of men’ in post-feminist society, noting the ‘social peripherality
of the frog’s location’ (by which he means the woods). ‘Women
want men to be responsible like themselves’, he concludes: to
move away from the marginal, superficial world of ‘frog culture’
and become real princes.16 Barbara G. Walker’s version in Fem-
inist Fairy Tales (1996) has a female frog protagonist who kisses
a male, becomes human, and then spends her happy-ever-after
life on her own. Female frogs, Walker notes solemnly in her intro-
duction, make a good role model for feminists as they are larger
than their males. My personal favourite, however, is J. F. Konrad’s
1981 tale ‘Der Lustfrosch’ (the lecherous frog). At first concerned
only with seduction, the frog sees the disgust and suffering that
he thereby causes his inamorata and ceases in his efforts. Recog-
nizing his newfound humanity, the princess begins to treat him
with compassion. It’s not so much the tale as its raison d’être
which is enjoyable for its earnestness. Konrad explains:

The revision has been conceived with a view toward

adolescents and is intended to help in connection with
other media and texts to free sex and eros from egotism,
exploitation, and boasting, and also from inhibition,
besmirching and smut and so enable them to form a part-
nership that lives from consideration and fulfilment.17

Despite applauding Konrad’s aim, one cannot help but find

oneself wishing for a little exploitation after all; perhaps even –
whisper it – smut.
The oeuvre of the Disney studio is emphatically not the
place to go looking for smut. Its animation The Princess and the
Frog (2009) is a typically clean-cut rendering of mythology, in
this case of the Grimms’ tale. It is also, as the title suggests, an

49
instantiation of a trend that has been going on for some time,
namely a shift in interest from the frog prince to the princess –
or rather the girl, as she starts out. In a limited sense therefore
the film echoes Walker’s tale, entitled ‘The Frog Princess’,
although the female focus of the Disney version is some way
from women’s liberation as it is recognized by serious feminists.
Disney’s film blends the plot from the Brothers Grimm tale
with a then-recently published novel for young adults entitled,
like Walker’s tale, The Frog Princess (E. D. Baker, 2002). In this
version, a girl kisses a frog in the hopes of becoming a princess
but instead, and unhappily, finds herself transformed into a frog.
The film too was originally to be known as The Frog Princess;
Disney’s pre-release market research, however, suggested that
the proposed title was considered by some to be a slur against
French people and it was changed accordingly.
The movie was made at a time when Disney’s fortunes were
failing. The computer-generated animations of Pixar and Dream-
Works were all the rage, while Disney had suffered a series of
flops with its recent traditional animations. In 2004, Disney
closed its 2d studios, but failed to find great success with its first
3d animation (Chicken Little, 2005) either. Meanwhile, however,
Disney princess merchandise – launched without much market
research in 2001 – had achieved astonishing commercial success.
Little girls apparently couldn’t get enough of the princessy theme.
The annual value of sales based on the established characters of
Cinderella et al. was estimated at around $3bn in 2006 ($4bn
by 2010); anyone with half a business brain could see the chance
for one more sure-fire, big hit. So, when Disney bought Pixar in
2006, they re-opened the traditional animation studio. There
was one big princess story still to go.
The Princess and the Frog very gently shakes up the recipe
for Disney fairy stories, most obviously by having the studio’s

50
 Still from Lotte
Reiniger’s silhou-
ette animation The
Frog Prince (1954).

first-ever black princess character, and also slightly rewriting the

Grimms’ tale. Turned into a frog by a voodoo spell, Prince Naveen
is on the look-out for a princess – he knows the kissing deal – and
mistakes the waitress Tiana, in a borrowed costume, for being
what he needs. On being kissed, however, she too is transformed
into a frog and together the pair escape to the Louisiana swamp
where the voodoo queen Mama Odie lives; eventually, and
despite many complications, Odie succeeds in guiding them to
break the spell.
Tiana is more independent-minded than her princess pre-
decessors. Her idea of a happy ever after – which of course she
achieves – is to run her own business. Anika Noni Rose, the actor
who voiced Tiana, stated: ‘She’s a strong woman who doesn’t
need anyone to do things for her . . . She wants to do things for
herself.’ Nevertheless, love and marriage are an essential part
of the resolution. Naveen is transformed through his inter-
actions with Tiana, becoming less selfish and vain until he
metamorphoses into the perfect groom. (Just like real-life elite
European-American marriages of the early twentieth-century

51
 era in which the tale is set, the transaction buys him money and
her a pedigree.)
The sense that the frog is simply waiting to be exploited for
his royal connections has its roots in the beautiful silhouette
animation of the tale made by Lotte Reiniger in 1954. This
widely disseminated film sets in place what is now a key feature
of batrachian iconology, the crown on the frog. This headgear
is a major clue for Tiana (or rather the film’s viewers – somehow
she seems to miss it) that this is not your average sort of frog.
The frog is
ruefully cele- The coronet on Reiniger’s frog is identical to the princess’s, fore-
brated as a telling their ultimate compatibility. Indeed, Reiniger’s tale, like
feature of the
contemporary its twenty-first-century successors, is about marriage from the
dating game, as outset. The girl’s father, the King, says the golden ball will bring
this image from
a dating advice
royal marriage to its owner and throws it to his three daughters
website shows. to see which one can catch it.

52
 The title-shift from the Grimms’ story to the movie (from the Still from the
Disney film The
frog’s identity to the girl’s) thus changes to a focus on the female Princess and the
narrative – the girl’s ambition and desire to secure a happy end- Frog (dir. John
ing for herself. Like the heroine of Baker’s story, Tiana aims, in Musker and Ron
Clements, 2009).
kissing her frog, not at marrying a prince but rather at becoming
a princess. Maybe the writer on the Peter Gabriel site was right:
the aspirational ‘Tiana’ is not so far from ‘Diana’, after all.
How did we get from decapitation to a kiss? Today, in the
West, sex is no longer so feared and dreaded that a symbolic act
as severe as decapitation is required for us to capitulate to our
desires. In today’s account of the Grimms’ tale, the aim is only
secondarily to find love. Primarily it is about becoming a princess
– an act of self-realization whose final stage is, thanks to the
frog, an erasure of the processes that have earned ‘success’ and

53
a validation of that success (fame, celebrity, being worth it) as if
it had been assumed by right, like a royal inheritance. The frog
groom is no longer significant in himself, but only insofar as he
enables the girl’s metamorphosis. The chief purpose of the fairy
tale is, as the uk wedding business of the same name states, to
be ‘princess for a day’.

54
2 Warts and All

Carl Linnaeus’ view of the Amphibia was not, by today’s standards,

what you would call a model of objectivity. In his authoritative
edition of the Systema Naturae (1758), he wrote:

These foul and loathsome animals are distinguished by

a heart with a single ventricle and a single auricle, doubt-
ful lungs and a double penis. Most amphibia are abhorrent
because of their cold body, pale colour, cartilaginous
skeleton, filthy skin, fierce aspect, calculating eye, offen-
sive smell, harsh voice, squalid habitation, and terrible
venom; and so their Creator has not exerted his powers
[to create] many of them.1

Most of these characteristics can be found in traditional

representations of frogs. Their imperfect physiology, epitom-
ized by their need to metamorphose, made them theologically
dubious from the Middle Ages and into the early modern period.
The mystifying double penis – for of course frogs have none,
let alone two – can probably be explained by the lumping
together of amphibians and reptiles in Linnaeus’ day; the latter
do indeed possess this remarkable anatomical feature. The
routine grouping of amphibians and reptiles put frogs in the
same group as that satanic creature, the snake. Frogs’ and toads’

55
Frogs have
been popularly
associated with
naughty boys,
although here
the teacher
appears to
turn the tables.

association with damp places, and their supposed venomous-

ness, have also long counted against them. To Linnaeus’ list of
reasons for abhorrence we might also add the unpredictable
nature of batrachians’ movement, a quality that frogs share
with the suddenly scuttling spider. It’s the unexpected ‘boing’
of a frog, guaranteed to raise a shriek, that makes them such
a satisfying surprise resident of the teacher’s desk. The horiz-
ontal slit-form of the pupil visible in some frogs and toads is
also for some reason disconcerting, a characteristic shared
with that other demonic creature, the goat. And, lastly, the squat-
ting posture of the frog perhaps reminds us humans of our
own in defecation.

56
 Of frogs and toads, the latter definitely have the worse press.
Milton had Satan himself ‘squat like a Toad, close at the eare of
Eve’ in Paradise Lost. The natural philosopher Lazzaro Spallan-
zani apologized for writing about toads in his Dissertations;
although the batrachian was not devilish, nevertheless ‘[n]ice
and fastidious persons may perhaps be disgusted at the frequent
mention of an animal so loathsome . . . disagreeable, nauseous
&c.’.2 The toad’s well-known (and entirely spurious) propensity
to cause warts apparently requires refutation to this day in every
factual account of the animals aimed at a general audience.
Toads have been almost universally considered to be venom-
ous. The very word ‘toady’, a sycophant, refers to this property.
In the early modern period, a toady was the sidekick of a char-
latan, employed to eat (or pretend to eat) toads, thus enabling
his master to exhibit his skill in expelling poison. Edward

Bufo cornutus
(now reclassified
as Ceratophrys
cornuta), complete
with horns and
leering mouth,
appeared to
confirm the toad’s
devilish reputation.
From François
Marie Daudin’s
Histoire Naturelle
des Rainettes, des
Grenouilles et des
Crapauds (1803).

57
Topsell, in his History of Serpents, gave considerable attention
to the venom of frogs and toads. Among the effects of frog
poisoning were a swelling-up of the body and pallidness; bad
breath and difficulty in breathing; an ‘involuntary profusion
of seed’; and general dullness and restlessness. Even toad-spit
on the scalp, noted Topsell, was enough to cause baldness. For
Boccaccio, the breath was enough to cause mischief. In the
Decameron a couple dies after rubbing their gums with sage
leaves; a toad is later found among the roots of the plant, infect-
ing it with its poisonous exhalations. Truly, the venom of the toad
was something to be feared, although sometimes good fortune
might save one from apparently certain toad-related death. In a
‘noted story’ of 1617, a country-dweller awoke from a nap to find
that a ‘Toad [had] fixed himself upon the mouth & outside of
[his] lips’. So firmly was the toad clamped on that violence was
needed to remove it; yet this could not be attempted lest the
toad was thereby induced to spit his poison ‘which he uses as
his offensive & defensive weapon’. Fortunately, a nearby spider,
being the natural enemy of the toad, provided the solution. Upon
being stung twice by this beast, the toad ‘swelled & fell dead’.3
The very potency of frog venom also made it an exception-
ally powerful medicine, and Topsell gave numerous recipes for
its use. Here he was largely inspired by Pliny the Elder, whose
32nd book of the Natural History was positively stuffed with
frog remedies. Frogs’ blood, fat, flesh, eyes, heart, liver, gall,
entrails, legs and sperm were all efficacious. They could be used
as powders, distillations, broth and infusions; they were good
for leprosy, scabs, poisoning, dropsie, coughs, gout and much
more. Buried in the garden (as for example in Thomas Hyll’s
1572 Arte of Gardeninge), a toad would drive away pests.
The noted early nineteenth-century savant Georges Cuvier
announced that toads in fact contained no venom, but in 1851

58
and 1852, two French philosophers were reported still to be
attempting to sort fact from fiction, experimenting with poison
extracted from the skin of toads. They succeeded in partially
paralysing a ‘little African tortoise’ and in killing a goldfinch.
The correspondent reporting their experiments to The Zoologist
noted that Messieurs Gratiolet and Cloez were at this time ‘occu-
pied in collecting a large amount of toad-venom’, but ominously
enough no information was forthcoming on their intended use
for this chemical weaponry.4
The notion of the unpalatable nature of frogs goes back a long
way. In Levitical law, the frog was proclaimed by implication to
be unclean. ‘These shall ye eat of all that are in the waters: what-
soever hath fins and scales in the waters, in the seas, and in the
rivers, them shall ye eat . . . Whatsoever hath no fins nor scales
in the waters, that shall be an abomination unto you’ (Leviticus
11:9–12). However, the chief damage to the frog’s reputation
occurred thanks to Moses and his repeated attempts to encour-
age Pharaoh in the matter of Israelite emigration (Exodus 8:2–4):

And if thou refuse to let them go [Moses warns Pharaoh],

behold, I will smite all thy borders with frogs:
And the river shall bring forth frogs abundantly, which
shall go up and come into thine house, and into thy
bedchamber, and upon thy bed, and into the house of
thy servants, and upon thy people, and into thine
ovens, and into thy kneadingtroughs:
And the frogs shall come up both on thee, and upon thy
people, and upon all thy servants.

And so it proves to be. The thought of kneading down on

bread and squashing frogs instead, or finding them in bed and
in the oven, is really rather unpleasant. Pharaoh’s magicians

59
Alluding to the
Mosaic plague,
frogs fall from the
sky in this poster
for the film
Magnolia (dir.
Paul Thomas
Anderson, 1999).
 Johannes Jacob
Scheuchzer,
Exodus: The Plague
of Frogs, from
Physica Sacra,
engraving, 1731–5.

are able to duplicate Moses’ trick, but alas cannot perform the
altogether more useful feat of making them disappear. When
Moses does so instead, things are little better: the people ‘gath-
ered them together upon heaps: and the land stank’ (8:14).
The biblical plague has provoked numerous later reports
of frogs falling as precipitation. The phenomenon is one of
those widely known ‘facts’ that is actually rather hard to find in
well-evidenced form, beyond meteorologists agreeing that in

61
Unclean spirits like principle a small tornado might pick up and then dump small
frogs come out of
the mouths of the
creatures such as frogs. The film Magnolia (1999) ends with a
dragon, the beast downpour of frogs, a surprising conclusion to an otherwise real-
and the false istic film. Many viewers claim to find references to the biblical
prophet in this
English glossed story hidden throughout the movie. Elsewhere, the phenomenon
Apocalypse (Book is purely proverbial; in Polish and Romanian, it is not cats and
of Revelation) of
1260. dogs, but rather frogs that fall from the sky during heavy rain.
Tainted by the Torah with pagan magic, frogs went on to
appear in the book of Revelation (16:12–14) as evil spirits:

And the sixth angel poured out his vial upon the great
river Euphrates; and the water thereof was dried up,
that the way of the kings of the east might be prepared.
And I saw three unclean spirits like frogs come out of

62
 the mouth of the dragon, and out of the mouth of the
beast, and out of the mouth of the false prophet.
For they are the spirits of devils, working miracles . . .

The power of frogs (and toads, and newts) to work miracles has
been widely believed ever since such ancient writings, at least
in European mythology. Perhaps it is also their metamorphic
nature that makes them such potent agents of transformation.
Shakespeare’s witches knew well the devilish virtues of frogs
and toads:

First Witch: Round about the cauldron go;

In the poison’d entrails throw.
Toad, that under cold stone
Days and nights has thirty one
Swelter’d venom sleeping got,
Boil thou first i’ the charmed pot.

Second Witch: . . . Eye of newt and toe of frog,

Wool of bat and tongue of dog,

All: . . . Double, double toil and trouble;

Fire burn, and cauldron bubble.

In Faust, too, Mephistopheles includes frogs and toads in

his recipe to help a blonde with her troublesome freckles. Such
potions were apparently not just in use in fiction. The historian
Deborah Willis tells how, on Lammas Eve 1590, a group of 40
witches set out to kill the King with a poison concocted from
‘toad, adderskin and other vile materials’.5 In 1608, Edward
Topsell noted that witches preferred toad venom as the means
by which to poison their husbands.

63
Two witches
smoke their pipes
by the fire with a
toad at their feet
in this British
engraving (1720).

Frogs and toads were not just the ingredients in satanic

brews; they were also the active assistants of witches, one form
of the demonic sidekicks known as familiars. Mephistopheles
describes how they do his bidding; he is ‘The master of the rats
and mice/ Of flies and frogs, of bugs and lice’. The anthropo-
logist Margaret Murray noted that witches generally kept their
frog and toad familiars in a pot or box and fed them on magi-
cally significant foods. Fifteenth-century witch-hunters recounted
a tale of a witch who placed the Eucharistic bread in a pot with
her toad, while Murray highlighted witches’ frequent use of
breast milk to nurture their batrachians. The demonic nature
of this phenomenon forced Michael Maier to explain in his
alchemical text Atalanta Fugiens (1617) that his emblem of a
woman suckling a toad was not to be taken literally. ‘It is indeed
a thing ominous for a Toad to be born of Woman’, he clarified,

64
and you can’t really argue with that. Beliefs about batrachian
familiars were not just in the currency of fable; in 1645, for
example, a witch found to have an imp in the form of a frog was
hanged in Cambridge.6
It was in the context and period of such hangings that the
eminent English physician William Harvey (1578–1657) is com-
monly supposed to have disproved the existence of demonic
toads. According to the story, at some time in the 1630s Harvey
disguised himself as a wizard and talked his way into a witch’s
dwelling, where he asked to see her toad familiar. He then sent
her off to buy some ale, during which time he dissected the
toad. Upon opening its belly, he found that it had real milk
inside, and concluded that this was no magical beast but
merely, in the splendid phrase of the tale’s author, ‘an arrant
naturall toad’.

A woman gives
birth to a toad
before her child,
from Jacob van
Maerlant’s Der
Naturen Bloeme
(c. 1350).

65
 The witch proved sadly recalcitrant when it came to accept-
ing this experimental disproof of her powers. Far from
capitulating to Harvey’s demonstration, she ‘threw downe the
pitcher of ale, [and] flew like a tigris at his face’. Harvey was then
forced to try the decidedly non-scientific methods of bribery
and finally blackmail to get her to accept his conclusions, reveal-
ing that he was the King’s physician and threatening to have her
prosecuted and hanged.
As historian Cathy Gere explains, the witch was not the
only one to doubt Harvey’s proof. The anecdote, she argues, is
as problematic as such other scientific myths as Newton and
the apple.7 James Long, who wrote down the events in 1686 on
the basis of conversations with Harvey, himself judged that
this was a ‘weake experiment’. Gere highlights how Long cari-
catures the tenuousness of Harvey’s argument: that ‘the toad
. . . had really eaten milk and not in appearance only, [and
that] therefore there are no witches’. Long’s counter-argument
(‘Spirits have recourse to toades . . . at set times . . . but doe
not exert them constantly’) seems to boil down to the idea that
the witch might have a natural toad (therefore anatomically
normal) that was sometimes possessed by an exterior spirit.
Conceivably Long’s argument implied that the toad would
appear internally normal even at the times that it was thus
possessed: that there was no difference between its inside and
outside in this respect. Gere’s account shows that Harvey by
no means established a definitively rational view of the frog.
‘Scientific’ and ‘superstitious’ views of batrachians continued
to co-exist through the early modern period.
Edward Topsell’s account of frogs (reprinted the year after
Harvey’s death) is a perfect illustration of Gere’s argument. In it,
Topsell is most concerned to distinguish ‘natural’ and ‘magical’
properties of the frog. The latter have no place in true learning;

66
he dismisses one instance with the robust judgement: ‘this is
as true as [that] a shoulder of Mutton worn in one’s Hat healeth
the tooth-ach’. Yet, to the modern reader, Topsell’s examples of
the natural properties of frogs are scarcely any different from
their magical ones. Witches, for example, don’t make spells
with them; they just use their venom. Physicians, on the other
hand, will find their healing frog broths work best if the
patient doesn’t know what he is drinking. Topsell’s distinc-
tions between nature and magic are not at root metaphysical
but rather theological. Action at a distance, and through the
hidden sympathies and powers of natural kinds, are all a part
of his world.
Let us backtrack in time to discover why natural philosophy,
or natural magic, made the toad a likely vehicle for malign spir-
itual influence. The answers lie in medieval theology, itself
drawing on the work of the Ancient Greeks. Aristotle noted that
some animals were generated spontaneously by putrefaction
– as, for example, maggots and flies arise from a corpse. When
these creatures then reproduced normally, he observed that
they produced imperfect forms, meaning that they had to meta-
morphose to reach their final, adult form. Thomas Aquinas
(1225–1274) gave the question of corruption and putrefaction a
theological spin; the body’s tendency to undergo this process was
in contrast to the incorruptibility of the soul. (Plotinus [204/5–
270] had in fact used the Greek term ‘amphibian’ to describe
this duality, contrasting the rational and animal elements of
human nature.) The Thomist theology of the corruptible body
was encapsulated by Ecclesiasticus 10:11: ‘For when a man is
dead, he shall inherit creeping things, beasts and worms.’ When
one added the Aristotelian metamorphosis of the frog to its
negative biblical connotations, it became a certain candidate
for generation by putrefaction.

67
 Frogs and toads also played a role in more general accounts
of abominable birth and generation. English folklore in partic-
ular is rich with tales of toads incubating birds; one particular
combination (toad and chicken) results in the monstrous
basilisk.8 Medieval tales of frogs and toads discovered sealed
within stones or rock persisted well into the nineteenth century.
Although nineteenth-century commentators attempted to give
such accounts a non-magical explanation, the stories them-
selves were still regarded as utterly credible for the most part.
Their origin and persistent appeal lay in their apparent evidence
for the spontaneous generation of batrachians.
Malleus Maleficarum, the well-known 1486 treatise on witch-
craft, synthesized Aristotelian and Thomist learning in order
to tackle the question of whether witches could create animals
(or turn people into animals).9 Witches, its authors found, could
indeed perform these feats, but only using Aristotle’s imperfect
animals – such as the frog (along with mice and serpents). Witches
could essentially harness the process of putrefaction to produce
these animals, as could (or so he claimed) the seventeenth-
century Jesuit scholar Athanasius Kircher, using ‘fertile dust’.
Transmutation – the changing of one creature into another –
was dealt with by the Malleus in a similar manner. Magicians
could counterfeit God’s ability to transform the essence of
matter, altering the superficial properties of humans into those
of animals. Claiming the authority of Aquinas, the Malleus
authors explained that frogs were a particularly suitable choice
for the changing of humans; their externally scattered semen
could easily be gathered and used by magicians to effect trans-
mutation, to the same end result (the counterfeit creation of a
Sigimondo Fanti, frog) that would be produced by the semen through putrefac-
Triompho di
Fortuna, 1526,
tion. The authors of the Malleus noted that witches sometimes
engraving. did not even create the superficial properties of a frog when they

68
magicked a human, but merely their appearance. However,
being only apparently changed into a frog was no less serious
than being changed in reality since everyone – even the injured
party – would truly and completely now perceive a frog in the
victim’s place. In such theological complexities we see the
essence of Long’s dispute with Harvey. There was no reason why
a demonically derived toad should not be perceived as normal,
even under the skin.
Edward Topsell was still bothered by the question of gen-
eration by putrefaction in the seventeenth century. His over-
arching programme in The History of Serpents was to show that
all creatures gave ‘testimony of God’ by demonstrating that they
were created by him. However, by the time he got to describing
frogs, this conviction seems to have palled. He observed that
their organs were ‘corrupted’, and gave a standard Aristotelian
account of how ‘some of them [were] engendered by carnal
copulation, and [some] of the slime and rottennesse of the
earth’. In other words, he was back to generation by putrefac-
tion. Tales of women giving birth to frogs were particularly
problematic for Topsell, for they introduced a perturbing blur-
ring of the corrupt frog body and the God-resembling human
body. He papered over the issue in one case-study by assuring
the reader of the woman’s good recovery; and in general he
excused such cases by pointing out that they tended to befall
Roman Catholics. The implication was that frog-birth was a
result of their worshipping the whore of Babylon and her
unclean, batrachian spirits.
Frogs born of putrefaction also took their place in medieval
tomb decoration. During the fourteenth century, idealized
images of the deceased began to be replaced by gruesome
images of the dead and decaying body, the ‘creeping things’ of
Ecclesiasticus.10 These corpses covered by frogs and snakes

70
The deceased
lovers of an
unknown 15th-
century artist
show the use of
frogs and toads in
memento mori-type
iconography.
became especially popular in Germany and Austria during the
fifteenth and sixteenth centuries; a particularly good example
remains in La Sarraz, Vaud (Switzerland). The wealthy de la
Sarra family commissioned their tomb around the late four-
teenth century and it includes a full-size, three-dimensional
sculpture of a naked male corpse covered with snakes or worms;
there is one frog on each eye, two on the mouth, and one on
the genitals.
Commissioned by the rich, such iconology displayed
appropriately humble confession of the sinful body prior to
judgement. The creeping things that emerged by putrefaction
from the body were like the sins that lurked within it. As Kath-
leen Cohen has noted, there were many examples of this trope
at work in medieval German teaching. One thirteenth-century
allegory told of a woman who would not confess to a particular
sin; an angel appeared to her with a beautiful child, whose back
revealed itself to be infested with toads and worms. The angel
explained to the woman that this was the woman’s condition
while she remained unshriven. A similar image archetype, ‘The
Tempter’ consisted of a young man, handsome from the front
but writhing with frogs and snakes behind. A thirteenth-century
monk from Heilsbronn dubbed the body ‘Krotsack, Madensack’
(sack of toads, sack of maggots), and another preacher of the
same era called sins ‘Kröten des Teufel’ (toads of the devil).
Besides its connection to devilish putrefaction, the frog of
the devil was largely to blame because of its posture, squatting
on its victim like a demon. It was particularly associated with
avarice. In the early seventeenth century, Edward Topsell des-
cribed how toads would spend the winter clawing earth into
their mouths to sustain them; their paws were constantly filled
with earth, as though they were afraid they would run out, and
in this they resembled the avaricious man who feared dying in

72
 Avarice is
portrayed as a
frog in this print
made by Jakob
von der Heyden
(1590–1645).

penury. Thus George Pencz, in his sixteenth-century engravings

of sin, used the frog to symbolize avarice, punished by being
placed in cauldrons of boiling oil.
For poet Philip Larkin, (1922–1985) the squatting toad was
the curse of work. ‘Why should I let the toad work/ Squat on
my life?’ he complained in 1954 in ‘Toads’; ‘Six days of the
week it soils/ With its sickening poison’. However, Larkin found
that the devil-may-care attitude necessary to scorn pensions and
shoes – the things that work buys – was lacking in him. He
identified this protestant ethic as a second toad, or ‘something
sufficiently toad-like/ [that] Squats in me, too’. ‘Toads Revisited’
(1962) reaches the same conclusion. Resigning himself to con-
tinued labours, Larkin invites his demon: ‘Give me your arm, old
toad;/ Help me down Cemetery Road.’ In the summer of 2010,
Larkin’s home city of Hull celebrated his life with appropriate
glum wit, commissioning a series of 40 giant, brightly coloured
toad sculptures that were scattered through its streets. Suitably
toad-like, many local politicians squatted on the project

73
Philip Larkin’s
‘toad, work’ was
commemorated
in a witty and
colourful series
of outdoor toad
sculptures in Hull
in 2010. This one
is decorated as the
poet himself.

(named ‘Plague of Toads’), complaining at its squandering of

public money.
In his novel Under The Frog (1992), Tibor Fischer refers to the
Hungarian expression that summarizes the unpleasant experi-
ence of being squatted on by a batrachian. In its full version,
‘under the frog’s arse at the bottom of the coal pit’, it describes a
situation that is as bad as it can possibly be. Although unpleas-
ant characteristics of frogs – including their tendency to inhabit
the marshy borders of Hades – go back to antiquity, it was in the
Middle Ages that they were truly cemented as icons of ill. How-
ever, an important question remains: were the actual creatures

74
regarded as evil, or were they simply used as a representation
thereof? The historian Sophie Page writes that the real animals
of the lived medieval experience were cognitively distinct from
their existence as emblematic and symbolic creatures.11 In real
life, non-threatening animals at least were generally perceived
as wholesome creations of God. Only in mythology and religious
fable did they reflect or personify the devil. The reason for this
separation of the real and the symbolic, she argues, was that
theologians had been compelled to counter the Cathar heresy
– which preached the inherent evil of the body, including the
animal body – by asserting the goodness of beasts. However,
crushing the Cathar heresy seems a long way from seventeenth-
century England, and we have seen evidence to complicate Page’s
assessment. In 1608, Edward Topsell was still concerned with
the putrid generation of frogs, and linked it to their treatment
as primarily noxious animals. The story of William Harvey

One of the most

horrific frogs ever
must be the
monstrous toad
of Pan’s Labyrinth
(dir. Guillermo
del Toro, 2006).

75
This affectionate and the toad suggests that the possibility of physical evil was
19th-century
Japanese print of
considered seriously enough for it to be worth telling in the
a toad reminds us late seventeenth century (even if his encounter didn’t actually
that toads are not happen in reality). Hence, we wind up back where we started,
intrinsically uncon-
genial creatures as with Linnaeus’s surprising commentary on the animal. Much of
European tradition his scientific representation of the ‘foul and loathsome’ frog
holds (School of
Hokusai). remains with us to this very day.

76
3 Them, Us and Frogs

In 1970, when many Californian students were busy dropping

acid, a young biologist named Richard Wassersug was persuading
his postgraduate colleagues at Berkeley to eat tadpoles instead. It
was all in the cause of science; his hypothesis was that highly
visible species of tadpole must taste bad in order to put off pred-
ators, while those that are well-hidden do not need this additional
defence. Duly following Wassersug’s instructions to suck, bite,
chew and rinse, the students gave taste-ratings for various species
and substantiated his theory.1 The arresting quality of the ex-
periment gained Wassersug an Ig Nobel Prize, rewarding him
for ‘improbable research’ that made people ‘laugh and then
think’. As Wassersug’s award indicates, the eating of frogs is an
attention-grabbing act, beset with cultural anxiety across many
parts of the globe. This chapter explores how such anxiety is just
a part – albeit a major one – of a bigger phenomenon: the use of
frogs to make generalizations about nationality and ethnicity.
Examples range from China to India, but most especially con-
cern frogs in France and Australia.
Uneasiness about the consumption of frogs is closely asso-
ciated with the difficulties that we have already seen in categor-
izing anurans; in anthropological terms, the categorization as
edible or inedible, clean or unclean, is a fundamental one. We
might expect creatures that are problematic and variable in

77
A pair of fresh
frog’s legs pre-
pared for cooking.

their categorization – like frogs – to display the same properties

in relation to their presumed edibility, and indeed this is borne
out by observation. In southern China, for example, frogs are
divided into three sorts: ha-ma, wa and toads, and different
cultures in the region variously encourage, permit or revile the
eating of each of these. In Sichuan during the Tang dynasty,
ha-ma frogs were captured whilst copulating and popped on

78
the dinner plate; they were also reared from tadpoles for con-
sumption. Nearby outsiders objected to these customs as
‘barbarian’. Wa frogs, on the other hand, were widely consumed
across southern China.2 Similarly, the 1891 census of Andhra
Pradesh named the Kappala sub-group of the local Yanadi
people as ‘frog-eaters’ and distinguished them from the ‘non-
frog-eater’ sub-group. An inhabitant of the region reported that
The European
the Yanadis of the North Arcot district – the ‘non-frog-eaters’ – edible frog.
did not even permit the Kappala to touch their pots, so unclean Although it is
was the latter’s habit of batrachian consumption.3 often given as
the species Rana
The British and Americans are famously obsessed with the esculenta, it is
alleged passion of the French for consuming frogs’ legs. The habit actually a fertile
hybrid of the pool
just does not seem right to Britons, even to many of those who frog (Pelophylax
otherwise consider themselves as adventurous eaters. The Scot- lessonae) and
the marsh frog
tish naturalist John Claudius Loudon (1783–1843), for example, (Pelophylax
suggested several foreign animals that might be cultivated for ridibundus).

79
the pot on the estates of the gentry, but stopped short at the
Edible Frog (Rana esculenta). Although it was a common meal
in Europe, he averred that ‘there [were] few Englishmen who
have eaten a fricassé of the thighs of this animal in France or Italy
[that] would wish to do so again’.4 In 1860, the Athenæum came
out in favour of frogs. ‘There is no reason’, it remarked, ‘why we
should eschew frogs, and relish turtle’. But notwithstanding
this logic, The Epicure’s Year Book for 1868 admitted that, alas, ‘the
poorest [English] man would disdain to eat a single pair’ of
frogs’ legs.5 A few cookery books from the same era attempted
to convince the English household to essay the dish, but without
much hope.
Even the great chef Marie-Antonin Carême (1784–1833) con-
ceded that the frog was not to everyone’s taste. And, strangely,
even for this fan of the grenouille it seemed necessary to point
out those whose consumption of the animal was beyond the
pale. Carême unfavourably compared the Viennese appetite
for the whole animal with the preference of the French for the
thighs alone, implying that there was something distasteful
about the former. Carême wrote a number of recipes for frogs’
legs, frying them, and especially making them into soups and
bouillons (one recipe calls for six dozen thighs – quite a
demanding requirement).6
Although frogs were to become signifiers of the fashionable
new haute cuisine by the late nineteenth century, Carême’s early
nineteenth-century dishes were very much inspired by the
frog’s traditional health-giving properties; his recommendation
of frog bouillon for curing consumptive coughs reached all the
way back to the remedies of Pliny the Elder. References to French
frog-eating date back to the Middle Ages, but another cele-
brated chef, Georges Auguste Escoffier (1846–1935), traced their
widespread consumption to a sixteenth-century entrepreneur

80
from the Auvergne region who made his fortune selling them
to Parisians. (The fact that this astute merchant only recom-
mended the legs for eating supposedly meant that he sold even
more.) Escoffier was apparently peeved at the refusal of the
English to countenance eating his national delicacy and so, in
1908, he planned a batrachian deception on no less a figure than
the Prince of Wales. Engaged to prepare a buffet for a grande
soirée at the Savoy, he included cuisses de nymphes à l’Aurore
(thighs of dawn nymphs) among the many dishes on offer to the
guests, revealing only afterwards their true nature. Amused and
impressed, Prince Edward ordered some more nymphs a few
days later. Escoffier was apparently chided for the suggestive-
ness of his dish’s title, but he stood fast (excusing himself with
a slightly different version of the story, whereby a beautiful
hostess had asked him to disguise the nature of the dish to spare
her guests’ sensibilities). Nymphes à l’Aurore – without their sala-
cious cuisses – live on in the latest edition of Escoffier’s Guide
Culinaire. To prepare them, poach the frogs’ legs in white wine,
and set them in champagne-flavoured fish aspic. Délicieuses.7
Thus the haute cuisine frog has an international history, cre-
ated as a gourmet dish in England by the French. The historical
complexities of sibling nationalism – the cultural politics of envy
and suspicion – have led to the edible frog’s strange dual status
as a dish that is both desired and reviled, its national hybridity
unacknowledged on either side of the Channel. Since Escoffier’s
time, many great French chefs at home and abroad have con-
tinued to create and refine dishes involving frogs. La Grenouille
in New York (f. 1962) pays homage to the ingredient in its name
– a signifier of its sophistication and, by association with French
cuisine, its excellence. Much as in the early nineteenth century,
fashionable Anglo-American eaters today esteem the frog, while
the popular taste remains somewhat more sceptical.

81
 La Grenouille’s chef imports his frogs’ legs all the way from
France, but down in French-influenced New Orleans, where frogs
are a far more popular dish than up north, the American bull-
frog, Rana catesbeiana, substitutes for the European R. esculenta
on the dinner table. However, the biggest exporter of frogs’ legs
today is Indonesia, which along with other East Asian countries
uses the Indian bullfrog, Hoplobatrachus tigerinus, widely in its
cuisine.8 Indonesia alone exported over four million kilograms
of frogs’ legs in 2009.9 Breeders prevent the meat from tasting
too gamey for Western palates by feeding the frogs a bland diet,
preventing their natural consumption of bugs and vertebrates.
What is at the root of Anglo-American disgust at the notion
of eating frogs? Fish are just as slimy, after all. Following Eco’s
definition of ugliness, a ‘lack of equilibrium, in the organic rela-
tionship between the parts of the whole’, we might suppose that
it is the inharmonious combination of limbs – associated with
mammals – and slime – associated with fish and the primordial
generally – that is to blame.10 My hunch is that Anglo-American
ideas about proper eating are also reliant on a tacit, internalized
version of the great chain of nature. Proper foodstuffs are found
further down the chain, not across it. Thus, most of the meats
consumed in Britain and America are naturally (or thought so
to be) herbivorous. The thought of eating another meat-eater
is wrong; eating a herbivorous frog is one thing, but eating an
Indian bullfrog that has just eaten a mouse – a mammal, like us!
– is abhorrent. And if eating across the chain of nature is wrong,
then cannibalism is off the scale. The zoologist Edward Bartlett
(1836–1908) was horrified to see an American bullfrog (R. cates-
beiana) in his keeping consume several of his European cousins,
the ‘pretty fire-bellied toad’ (Bombinator igneus). ‘Our Yankee
frog commenced immediately to swallow “alive and kicking” the
unfortunate Saxony toads’, he reported. Adding comic horror

82
to the situation was Bartlett’s impression that the frog’s dis-
tended jaw reminded him of ‘kind mothers who say to the baby,
“I could eat you, you darling”’.11 It is an amusing story, but I
confess that I would baulk somewhat at seeing this and then
eating the bullfrog – an edible species – in question. It may be
that eating too far down the chain of nature is also taboo; hence,
even frogs with more harmless diets are, like insects, too far
removed from mammals to be considered for edibility. (Actually,
although Rana esculenta does not have a reputation for grotesque
consumption like R. catesbeiana, scientists have found that it
does possess cannibalistic tendencies.)
Contrary to popular belief, the Anglo-American tendency to
refer to the French as frogs is not, or not entirely, due to their
presumed eating habits. The story starts out entirely differently,
in the Middle Ages. Alas, as historian Michael Randall explains,
the French seem to have brought the slur upon themselves by
unwisely – and possibly uniquely – attempting to forge an affirm-
ative identification with batrachians.12
At the birth of the French legend, the toad had its usual evil
characteristics. A popular fourteenth-century history told how
King Clovis, prior to his conversion to Christianity, had three
toads on his coat of arms. Thanks to a miraculous interven-
tion, these males bestes were transformed overnight into pure,
Christian lilies (fleurs de lis), marking the beginning of Clovis’s
successful Christian reign. During the sixteenth century, how-
ever, ideals of kingship changed. It became important to recog-
nize the king’s absolute power: to show that Clovis had exhibited
sovereign quality even before his conversion. He could not be an
unwise man who became a good king thanks only to a miracle.
Thus, the representation of the toad had to change; Clovis must
have chosen it, even before conversion, for a good reason. ‘It is in
the nature of the toad’, explained one sixteenth-century historian,

83
Frogs at the
Latona Fountain
in the gardens
of the Palace
of Versailles.

‘to find always for himself the best, the most fertile, and the
most useful place. And it is for this reason the king of France
possesses and occupies the best, the most fertile and the most
useful land in the world’. Another sixteenth-century historian
substituted a tree frog for the toad on Clovis’s pre-conversion
coat of arms – slightly less problematic, less demonic, in terms
of its connotations for Clovis and French notions of kingship.
For a while, the pr job performed on the toad-king by six-
teenth-century French historians seemed to hold, until after
about 50 or 100 years, when they began to downplay or even
deny the legend altogether. Unfortunately, however, the story
was out – to the delight of the foes of France. It has been sug-
gested that frogs were used in England to lampoon the French
as early as 1580, when the folk song ‘Frog Went A-courting’
was popularized: supposedly it was a satirical account of the
attempt by Francis, Duke of Anjou, to woo Elizabeth i. By the
seventeenth century, for certain, enemies of the French were
exploiting their toadish connections with glee. The Flemish
were particularly merciless in this respect. They began to call

84
the French toads and their king the ‘descendant of toads’; Dutch
histories of France deliberately mis-drew the contemporary
French coat of arms with toads on it. Edward Topsell, in his
pro-Protestant History of Serpents, also repeated the legend of
Clovis to the detriment of the French.
Historian David Bindman has discovered that during the
later seventeenth and eighteenth centuries, the French were
turning the slur the other way; along with the English they
began portraying the Dutch as frogs, for the ostensible reason
that they lived in a low-lying and swampy land.13 Dutch peas-
ants in particular were pictured in anuran form, as fat, lazy and
cunning. One particularly sophisticated satire of 1672 showed
‘Holland . . . as a huge horse turd’ and the Dutch as ‘frogs who
grow from maggots feeding on it’.
Following the Glorious Revolution of 1688, when the Dutch
William iii took the English throne, inhabitants of the Nether-
lands dropped out of the satirical sights of the British. Their place
was taken by the Catholic French, and by those foppish English-
men and women who imitated them. A crucial failing of the
French was their insincere and fancy sense of taste, characterized
above all by their preference for silly little servings of food
rather than good platefuls of ‘honest’ beef: ‘In foreign vests the
gaudy Fops may shine,/ And on dissected frogs politely dine.’
As the eighteenth century went on, the eating of frogs
‘became a sign less of luxury and affected delicacy than of
poverty’. Hogarth’s 1756 engraving The Invasion showed thin
French soldiers mustering in Calais, dreaming of England’s rich
foods while preparing a paltry kebab of frogs for themselves.
Similarly, Isaac Cruikshank’s etching French Happiness, English
Misery (1793) compared fat Englishmen enjoying their meat
with a group of scrawny, post-Revolutionary Frenchmen grasp-
ing at a single frog. Soon after Cruikshank made his drawing,

85
the Napoleonic Wars produced an absolute proliferation of James Gillray,
‘Amsterdam
English representations of the impoverished, frog-eating French. in a Dam’d
To Bindman’s focus on poverty we could also add that frogs Predicament; or,
were a signifier of Catholicism. Frogs were recommended both The Last Scene
of the Republican
as a health-giving sustenance freely available to the poor invalid, Pantomime’, 1787,
and as a meat that might be eaten during Lent, since they were hand-coloured
etching. The
closer to fish than to other quadrupeds. They had little blood, Dutch were widely
and much phlegm, and were unlikely to arouse the passions.14 caricatured as frogs
in the seventeenth
By the nineteenth century, with Dutch power in abeyance, century, as this
the French resumed their batrachian characterization and print shows.
completed their metamorphosis from frog-eaters to frogs pure
and simple. Political allegories concerning frogs had a lot to
do with this transition. In Aesop’s fable, the frogs demand a
king but later come to regret it when he – a crane or a crocodile,
depending on the version told – begins to eat them. The prop-
ensity of the French to revolt – and like Aesop’s subjects, to
end up with a worse regime than they had before – therefore
completed their identity with frogs in the light of this tale. 15
There is another Aesopian frog connection, ‘The Ox and the
Frog’, in which the frog attempts to puff himself up to the size
of the former, until he unfortunately bursts. Victorian British
satirists recounted this fable, casting the ox as a British bull,
and Louis viii, pretender to the English throne, as the fatally
conceited frog.16
In more recent history, the French have been joined by
other nationalities in their derogatory association with the Isaac Cruikshank’s
etching French
frog. During the Second World War, British propaganda por- Happiness, English
trayed the Japanese as toads. More recent British enemies have Misery of 1793
is an early
been in the Middle East; as the British public assimilated the example of the
notion of the previously demonized Iraqis becoming allies in Anglo-American
association of
2006, the Daily Mail newspaper reflected the difficulty of that frogs with
transition by reporting that ‘Iraqi soldiers bit the heads off the French.

87
 A cartoon in the
British Independent
newspaper (2003)
gives the French-
frog trope another
twist by having
then British Foreign
Secretary, Jack
Straw, as President
Bush’s toady, and
therefore more
despicable even
than the frog.

frogs . . . at a ceremony to transfer Najaf province from us to

Iraqi control’. The Iraqis’ execution of this repugnant batra-
chian ritual signalled their doubtfulness even when in an official
state of alliance. Some three years earlier, shortly before the
commencement of the war, the Independent newspaper gave
another twist on the politicized frog in its cartoon ‘Toadies say:
hop off frogs!’ The French Foreign Minister had come out against
the proposed invasion of Iraq but his British equivalent, Jack
Straw, followed the hawkish us line in condemning the French
stance. In Dave Brown’s cartoon, the frogness of the French is
ironically referenced by portraying Straw as a toad. As a toad(y)
of Bush, it suggests, Straw is more despicable than the much-
despised French frogs.
When frogs get out of place, as ‘invasive species’, they become
a focus for these issues of nation, ethnicity and identity. The
Aesop’s fable of
best-known example of this phenomenon is the cane toad (Bufo the crane and
marinus) in Australia, a large species whose most substantial frogs, illustrated
in a 19th-century
recorded example weighed in at 2.65 kg.17 In 1935, the Queens- wood engraving
land scientist Reginald Mungomery collected 102 cane toads after Gustave Doré.

89
from an experimental colony in Honolulu to take back home.
He had seen the toads demonstrated at a conference in Puerto
Rico, controlling beetle pests of sugar cane; Mungomery hoped
that they would do the same for the Australian Greyback Cane
Beetle. Warning bells should, arguably, have rung when he
bagged his first 50 in only one hour. This was too easy; the
toads had already spread all over the island and begun to plague
local gardens.
Back in Australia, the imported specimens produced 42,000
more adult toads for release within one year. Local farmers were
initially anxious for their land to be selected for toad settlement,
but before very long it became apparent that the toads were of
no use for their intended purpose. Some blamed this on the
fact that the greyback beetle, unlike the Puerto Rican beetle, did
not return regularly to ground where the toads could get at
them. Subsequent commentators have argued that the reduc-
tion in Puerto Rican beetles during the toad trial was actually a
coincidence, and that unusual weather conditions – not the
toads – were the cause of what in any case turned out to be a
temporary reduction in beetle numbers.
Human and ecological concerns about the toads’ release were
expressed as far back as the year of the policy’s introduction,
and faced with these the Australian government temporarily
halted their release. Mungomery’s bullish insistence won
through, however, and the ban was soon reversed. Within a very
short space of time, the critics’ concerns were vindicated, and
the disastrous effects of the toads began to become evident. B.
marinus feeds voraciously, destroying native insect, bird, mam-
malian and reptilian life. It is highly noxious and the few native
snakes that attempt to prey on it are poisoned. (Meat ants have
recently been discovered to be a notable exception to this rule,
and appear to predate effectively on young toads.) Mungomery

90
himself remained unrepentant, continuing to view his task with
almost megalomaniac zeal.
Cane toads continue to spread inexorably across Australia
from their original introduction point in the northeast, at a rate
of over 100 km per year. They are proving suited to all environ-
ments, including semi-arid areas and in water of up to 15 per
cent salinity, and now occupy well over a million square kilo-
metres of land. Some ecologists have questioned the extent to
which B. marinus poses a threat to biodiversity; some habitats
and some species, it seems, are more damaged by the toads than
others. Nevertheless, millions of Australian dollars have been
spent trying to halt the toads’ progress and to eradicate them
where they have already arrived.
With typical Aussie bravado, some Australians have adopted
the cane toad as a kind of mascot. In general, however, they are
loathed. Attracted by garbage and lights (that in turn attract
insects), large numbers of toads swagger into gardens and
houses. Pets that investigate them too closely will die of the
potent venom in their skin. Unavoidable on the highways,
they burst under car wheels and splatter their guts across the
roads. Ecologists and geographers, as well as historians of sci-
ence, have noted the nature of the military and nationalistic
language that is invariably used to articulate this disgust and
dismay: a vocabulary of ‘invasion’, and of ‘alien’ and ‘native’
species. Critics point out the implicitly xenophobic connotations
of this language, and also the illogic of treating humans and
animals in different ways when it comes to discussing migration
and colonization. All organisms once had to colonize the area
in which they now live, and many in their evolutionary past
have come some distance to do so, or (like the toad with its
human vector) have done so in conjunction with other organ-
isms. So why pick out recent colonists as somehow unnatural?

91
 In the case of the cane toad in Australia, this critical reading
seems plausible. This is an instance of an immigrant popula-
tion – and one with a poor record in regard to the indigenous
population at that – getting extremely upset about another
immigrant invader. Australian politics has more recently also
been exercised with human immigrants; in 1992, a policy of
mandatory detention of unauthorized immigration arrivals was
established, and in 1997, the far-right One Nation Party made
anti-immigration policies central to its manifesto. So, one might
be tempted to posit that what we have here is a projection onto
the toads, on the part of the white colonists, of a selfish concern
that no one else should usurp their colonization. Alternatively,
it might conceivably be read as a displacement of the colonists’
guilt at having done much the same thing as the toad.
Or could it just be that the toads are really and simply nasty?
I must confess frankly that I would be horrified and disgusted
to have an evening on the patio spoiled by giant hopping toads,
or worse still by encountering them indoors. Counting in favour
of actual nastiness, one might note that even those with impec-
cable liberal qualities find the toads repulsive; although in
keeping with their world-view they may class the phenomenon
not so much as invasion as an example of the ‘McDonaldization
of the biosphere’. On the other hand, as Tim Low reminds us,
Australians are ignorant of the vast majority of introduced
species to their country, many of which have proved consider-
ably more harmful than the toad. Other introduced species are
‘nice’ and have been given a kind of honorary native status: olives,
honeybees and trout, to name but three. Thus, the toad is doing
some kind of special representative work.
What is it about toads, as opposed to other animals? Find-
ing six woodlice in the house would not bother me at all. Finding
half a dozen feral cats would be perturbing but not revolting.

92
We could here consider a genuine historical counterfactual along
similar lines; in 1902, moles were proposed for introduction
into Australia as a means to control insect pests. Had this been
followed through, and even if they had disrupted Australian
ecologies, it is hard to imagine them provoking such disgust.
Irritation, perhaps, but not disgust. As in the case of eating frogs,
Eco’s definition of formal ugliness – as disequilibrium in the
natural pattern – again works to a certain extent as an explana-
tion for the toads’ effect. In this instance, the toad represents a
disruption in the expected number of batrachians, and also in
size compared to the European norm. The latter is also true of The American
the American bullfrog, which has recently been identified as a bullfrog (Rana
catesbeiana) is a
problem in the uk. ‘The bullies of the frog world [have] crossed recent invasive
the Atlantic’, warned the Independent in 2000, at just the time species in the uk.

93
that anti-American sentiment was running high among Brits. In
some senses, then, there are inherent or at least long-established
attributes of the frog that make cane toads and American bull-
frogs such targets of loathing – slime, warts, poison and so on.
However, their being out of place is a significant factor, and it
seems probable that they are the focus for some xenophobic
sentiment. As in all cases where frogs are used to demarcate
between ‘them and us’, their unpleasant features – both natural
and cultural – have become mutually reinforcing.

94
4 Under the Knife

In 1847, the young surgeon Hermann von Helmholtz was, by his

own account, ‘waiting impatiently for the spring’. One might be
forgiven for thinking that he had love on his mind; his engage-
ment was to be announced only a month or two later. But, in
fact, he was thinking about frogs. Specifically, he was waiting for
the frogs of Potsdam to emerge from their winter hiding places
and venture forth to breed by the ponds. At that moment he
would be able to take his collecting jars and capture them in
their dozens before transporting them back to his laboratory.
Helmholtz had some nasty things in store for the little creatures;
they were, in his own words, ‘those old martyrs of science’.
Helmholtz was neither the first nor the last person to experi-
ment on frogs. One of the earliest hi-tech pieces of science,
Robert Boyle’s air pump (1659), was used for the purpose. It
comprised a chamber that could be emptied of air, creating a
perfect vacuum, and almost immediately experimenters tried
frogs and toads (among other animals) to see how long they
could survive inside.
‘Hey, let’s try it with a frog!’ The same experimental method
quickly occurred to twentieth- and twenty-first-century scientists
working with high field magnets, a contemporary example of
apparatus as impressive and complex as Boyle’s was in its day.
Inside the device, the atoms of the animal act as tiny magnets

95
Frogs were among
the earliest experi-
mental subjects for
X-rays (undated).

and are repelled by the powerful magnet of the machine. In a

lightweight organism, this force of repulsion is sufficient to
counteract gravity and the frog levitates freely as though float-
ing and turning in space. ‘The small frog looked comfortable
inside the magnet and, afterwards, happily joined its fellow
frogs in a biology department’, reported Dutch experimenters.1
Frogs and tadpoles have also been sent into space. Initial Soviet
attempts to raise tadpoles in space – which seem to have been at
the speculative end of science – were only ‘marginally success-
ful’. Other attempts have been more focused in purpose. In 1970,
nasa launched two American bullfrogs in their ‘Orbiting Frog
Otolith’ to understand how the human inner ear might respond

96
 An experimental
frog levitates in a
high field magnet.

to long periods of weightlessness. More recent experiments

have aimed to understand how tadpoles deal with the water-
based gravitational challenges of real life. The scientists behind
these latest experiments wonder whether tadpoles can ‘learn’ to
live in zero gravity.
A great deal of early anatomical and physiological work with
animals was inspired by a desire to understand the human body,
and so frogs, much farther removed from us than mammals,
were not initially an especially popular choice for experimenta-
tion. Perhaps the earliest researcher to make significant use of
frogs was Jan Swammerdam (1637–1680), a Dutchman who is
best known today for his microscopy and anatomy.

97
 Swammerdam put frogs in what is, from today’s perspective,
a surprising category. They formed a part of his Natural History
of Insects (1669). This was later expanded as The Bible of Nature
(c. 1679).2 In it, Swammerdam aimed to show that ‘all God’s
works are governed by the same rules’.3 Swammerdam used
frogs as an in-between animal, standing midway between
insects and the ‘larger, or sanguiferous animals’, to show that all
were part of the same pattern of creation.
Swammerdam’s first staging post in this strategy was to show
that the development of insect larvae and tadpoles proceeded
in the same way. Swammerdam was an extraordinarily talented
dissector and microscopist, and his skills were well suited to the
task. Yet demonstrating that frogs developed according to the
same law as insects was not enough, for it merely reduced the
frogs – never a high-status animal in any case – to that of insects.
So, Swammerdam next took a clever turn in his Bible of Nature,
relating the frogs to the higher animals. Until this time, research
on muscular movements had been largely carried out on cats,

One of the
frogs used in
an embryology
experiment on
the Spacelab-J
flight launched by
the Space Shuttle
Endeavor, 1992.

98
 Jan Swammerdam
was one of the
first natural
philosophers
to experiment
extensively on
frogs. From
Swammerdam’s
Bible of Nature
(1679; published
1737).

dogs and poultry. Typically, their nerves were exposed and

stimulated with all manner of things: pins, scalpels, heat and
caustic or ‘acrid liquors’. What Swammerdam did was to try
these experiments instead upon frogs. Swammerdam found that
the frog’s body was not required whole to produce the effect. A
nerve-muscle preparation dissected out of a frog, when touched
with scissors or other instruments, also strongly contracted.
Swammerdam was quite explicit that ‘experiments on the
particular motion of the muscles in the Frog . . . may be also, in
general, applied to all the motions of the muscles in Men and
Brutes’. One might perhaps think that this was a rather radical
conclusion – placing humans in the same category as animals –
but Swammerdam drew a devout lesson from his findings. The
frog’s assistance in Godly reflection was wonderfully indicated
by its ‘wonderful copulation’, in which the male ‘most beauti-
fully joins his toes between one another, in the same manner as
people do their fingers at prayers’.4 From the insect to the frog,

99
and from the frog to the human; the whole of God’s creation
was now joined in one seamless, law-like continuity.
In the eighteenth century, the Italian priest and investigator
Lazzaro Spallanzani (1729–1799) continued the quest into under-
standing the body, employing an extraordinary and inventive
cruelty to frogs to do so. He cut off their legs to see if they would
re-grow (they did, if the specimens were young enough); he tried
different ways of killing them by suffocation and a pin through
the brain (they all worked); and abused male toads and frogs
during the act of mating to see if he could induce them to let go
(even decapitation would not do the trick).
Spallanzani’s heartless science was a long way from certain
frog studies carried out by the British traveller and naturalist
Robert Townson (1762–1827). During a visit to central Europe in
1793, Townson acquired a female tree frog (Rana arborea), which
he then proceeded to keep in a little box in his pocket. He named
her Musidora, from the Greek ‘gift of the muses’: presumably a
reference to her musical utterances, common to the species as
a whole. As historian Christopher Plumb notes, the sounds of
the beautiful little frog – ‘much like a blunt file against a piece
of steel’ – were more pleasing to some than others.5 Plumb
describes how Townson cared for his little frog, keeping her in
a glass of water by the window of his Göttingen study. In this
warm room she lived through three winters without the hiber-
nation usual to frogs, and grew tame enough to sit in Townson’s
hand, where he fed her on flies. Townson’s scientific observa-
tions on amphibians made numerous reference to Musidora by
name, and it was with great sorrow that he announced in one
such publication:

I am exceedingly sorry to [inform] Musidora’s friends (for

she had many) that she is no more. She sickened soon

100
 Robert Townson’s
pet Musidora
(c. 1791–4) was
a European tree
frog like this (Hyla
arborea, then
known as Rana
arborea).

after she reached Great Britain, and died in the night of

25th of June 1794.6

Yet, notwithstanding his fondness for Musidora, Townson

conducted crueller experiments on her conspecifics, of which
he kept up to 80 in his basement. He was inspired in his
researches by the fact that nobody appeared to have understood

101
frog respiration, merely – and inappropriately – generalizing its
nature from other animals. In a strange kind of way, even his
cruel experiments were perhaps bred of a fondness, or rather
respect for, the frog.
The best known of all eighteenth-century frogs belonged to
Spallanzani’s contemporary, the anatomist Luigi Galvani (1737–
1798). There are several variants on the myth that tells the story
of Galvani, the lightning and the frog. The most touching ver-
sion has Galvani in his kitchen, preparing a meal of frogs’ legs
for his wife Lucia, unwell in bed. As he chops the skinned legs,
he accidentally touches his knife across from the muscle to the
nerve, provoking the leg to kick. Another version has Galvani
cutting frogs up for experiment, when distant lightning causes
the legs to kick; another still has Lucia back in the picture (this
time healthy) and noticing for herself the electrical machine
nearby that is the cause of the leg’s kicking.
Most people are so excited to get on to the electricity part of
the story that they forget to ask what the frogs were doing on
Galvani’s bench in the first place. Vivisecting a frog to the point
where it would respond to electricity was no simple task, and
certainly not something that could be done casually or accident-
ally. By the mid-eighteenth century, it had become commonplace
to ascribe the nervous stimulation of muscle, such as Swammer-
dam had observed, to the action of electricity. The phenomenon
of electricity had been identified earlier in the century, and in
addition to its theoretical role in muscular movement, it was
associated with medical practices whereby patients received
shocks intended to improve or restore their health. Yet despite
these theories and practices, there was little by way of substan-
tial evidence that the two were connected: that ‘animal spirits’
were actually electrical in nature. Thus it was in the late 1770s that
Galvani began, in somewhat the same spirit as Swammerdam,

102
 Engraved
illustration of
Galvani’s frogs’
legs for electrical
experimentation
(1793).

to explore the irritability of nerves, and to attempt to forge a

direct connection between nerves and electricity.
Because the frog’s muscle contracted when it was connected
to the nerve by metal, Galvani considered that his hypothesis
– that frogs contained animal electricity – was confirmed. The
electrical fluid existed in a state of disequilibrium (condensed
in the nerves, and rarefied in muscle), and was conveyed from
one region to the other by the metal, producing a contraction.
Galvani’s conclusion was, however, met with disbelief by the
physicist Alessandro Volta (1745–1827). Volta attempted to
repeat Galvani’s experiment, and claimed that he could do so
only if he used a bridge that was composed of two metals; it
was the metals that were causing the jerk, not the frog. Volta’s
ultimate rebuttal was to create a pile of moist metallic bodies,
making an electric battery from two different types of metal –
no frog was needed. This was widely considered, by physicists
at least, to put paid to Galvani’s ideas. Whereas Galvani treated
the organic body as a topic in its own right, Volta treated the

103
Jean Noel Halle
and Alexander
von Humboldt
experimenting
on frogs (1798).
The rabbits may
get a lucky escape
thanks to the
sacrificial frog.

frog as a piece of physical apparatus, and an unreliable one at

that. Inorganic equipment was much more decisive for himself
and his followers.
Despite the ‘disproof ’ of Volta, many dozens if not hundreds
of frogs were used in electrical experiments by physiologists
over the next century, and the man who really started the trend
was the German Johannes Müller (1801–1858). Müller was trying
to confirm the work of François Magendie (1783–1855), who had
claimed to show that nerves from different roots did different
things, those joining the spine at the front controlling movement,
and those joining it at the back relaying sensation (which in
an experimental context meant pain). Magendie’s experiments
were carried out on live dogs, but when Müller tried replicating
them on rabbits, he found that the creatures’ general distress
was so great that it was impossible to discern the results of

104
stimulating particular nerves. So, at some time in the 1820s or
early 1830s, Müller hit upon the idea of using frogs instead.
Like Swammerdam, Müller found it comparatively easy to
expose the spinal cord, and similarly noted the frog’s ‘tenacious’
grip on life, meaning that its nerves and muscles continued to
respond to irritation for a long time on the laboratory bench –
up to 30 hours, compared with fifteen minutes in cats and dogs.
Müller was so pleased with his results that he encouraged aspir-
ing young physiologists to go and do likewise. His students, the
friends Emil Du Bois-Reymond (1818–1896) and Hermann von
Helmholtz (1821–1894), took him at his word.
Physically, the frog was indispensable to the questions these
men wanted to explore. Helmholtz, for example, wanted to find
out whether muscles observed the law of the conservation of
energy: when they contracted, did all the heat generated come
from their own, internal chemical reactions? To do this, he had
to find a muscle that would work in isolation from its normal
blood supply, otherwise the results would be muddied by the
muscle burning oxygen from incoming blood. Only the mus-
cles of cold-blooded animals could survive in this condition
for any length of time, making the frog an ideal animal to use.
Meanwhile, for Du Bois-Reymond the frog’s leg was simply the
most sensitive galvanometer that he could find. He almost drove
himself crazy trying to create an artificial device that responded
to a current as low as the frog’s leg did, eventually succeeding
by making an unimaginably tedious 24,160 turns in a thin
wire coil.
Working with frogs presented many practical challenges.
Even when dead, the frogs made certain demands of the physi-
ologists in order to go on working for them. Historian Laura
Otis describes all of this wonderfully well.7 There were no dedi-
cated laboratories for the work of the physiologists, and so Du

105
Bois-Reymond had to experiment in his room at his parents’ house,
and at the Anatomical Museum in Berlin. His parents’ neighbours
objected to his filling the house with animals for experiment,
and dogs were absolutely out of the question (suggesting, per-
haps, a more pragmatic reason why he switched to anurans).
Frogs were just about acceptable, although one wonders what
they thought of the numbers; by autumn 1841, 100 frogs were
sharing his room.
Obtaining the frogs was difficult. Du Bois-Reymond depended
upon a supply sent to him by train from Leipzig, if necessary
bribing the conductor to make sure they reached him before
they expired. Events conspired to disrupt his supply; while the
rest of Berlin was in revolutionary upheaval in April 1848, Du
Bois-Reymond complained to his supplier about the ‘lack of
frogs’. Looking after the frogs was also tricky. During four suc-
cessive winters in the 1840s, Du Bois-Reymond lost 10–50 per
cent of his frogs to disease. On another occasion, all his frogs
froze to death, due to the thoughtless absence of his servant. A
final challenge was presented by the need to transport the frogs
and their corpses between the Anatomical Museum and his
parents’ house. Du Bois-Reymond had to come and go between
the two frequently, carrying ‘a sack of frogs and ice’ in the heat
of the day.
The odd thing is that although Müller and his disciples
were utterly dependent on frogs for their research, forced to
confront them in all their bodily reality, they did their best to
deny their animality – to reduce them to mere instruments – in
the story of their science. As Otis explains, the careers of these
men were embedded in debates about vitalism. Vitalism, the
idea that there was something scientifically special about living
bodies, was associated with eighteenth-century romantic philo-
sophy, and tended to be espoused by the politically conservative.

106
Anti-vitalists, who tended to be politically progressive, insisted
that the physical processes of living organisms were no different
to those found in a test tube. As anti-vitalists, Du Bois-Reymond
and Helmholtz had to distance themselves from their animals
– to make it clear that their science was not specially the study
of life.
In his account of Müller’s life, Du Bois-Reymond made his
master’s story one of death and resurrection, a messianic triumph
over the ‘dragon’ of vitalism. He ascribed Müller’s 1827 break-
down to the heroic but draining self-experimentation that he
had been carrying out, trying his best to defeat the dragon.
Müller’s adoption of the frog, ‘forgotten’ since Galvani’s day,
was the turning point according to Du Bois-Reymond. The evil
dragon was slain, thanks to the batrachian victim sacrificed in
Müller’s new experiments. (Or, as in the evolution of the frog
king story, the medieval dragon was replaced with the harmless
nineteenth-century frog.) The sacrificed frog now had to sur-
render for a second time, disappearing from the science so as
not to give the impression that the latter was intrinsically rooted
in the study of living things: to prove that the science was not
vitalist. And Du Bois-Reymond absolutely had to lose the jokey
title ascribed to him by his friends at the Anatomy Museum,
‘the frog doctor’. Thus the experiments of Du Bois-Reymond
and Helmholtz now take their place in retrospective histories of
both physiology and physics, but the frogness of their frogs has
been almost completely lost. The anuran has been reduced to a
mere instrument.
Meanwhile, in Britain there was a growing sense of unease
about animal experimentation. In its founding statement of
1824, the spca (Society for the Prevention of Cruelty to Animals)
singled out vivisection as a practice to which it was opposed.
One should not overstate the extent of this moral outrage, but

107
nevertheless criticism was present, targeted particularly at foreign
researchers. In response to this groundswell of opinion, British
physiologists began to choose cold-blooded animals for their
work whenever they could; intuitively, they were less distress-
ing to work on.
Among these physiologists was Marshall Hall (1790–1857),
whose scientific aims included proving that the brain was the
centre of all sensation, and that only insensate reflexes existed
in decerebrated frogs. Hall had a number of prominent critics
who insisted that consciousness extended into the spinal cord,
and combined with the general public distaste at animal exper-
imentation this prompted Hall to defend his practices. Hall
emphasized that the use of vivisection should be minimized,
and that where necessary it should always be carried out on
the lowest possible type of creature. He suggested that frogs
were especially good animals to use; their low position on the
zoological scale was proved by the fact that they continued to
exhibit the phenomena of life after mutilation – even decapita-
tion. As lower animals, they were also less able to feel pain.
Concerns about animal vivisection were coming to a head in
1870 when Darwin’s great publicist, T. H. Huxley (1825–1895),
borrowed Hall’s frog in his lecture ‘Has a Frog a Soul?’ Huxley’s
short answer to his own question was: no. To get there, he drew
on Hall’s experiments. Huxley’s ad absurdum argument described
how ever-smaller parts of the frog could be dissected out of the
animal but would still respond to stimuli. Either metaphysical
possibility – that the soul, too, was chopped up in this process,
or that it somehow hung in the ether connecting the separate
parts of the frog – was ridiculous.
It appears rather odd that Huxley should be debating the
matter of the soul as late as 1870, by which time it had completely
dropped off the agenda of serious physiological research – a straw

108
man. Moreover, Huxley signally failed to address a much more
important contemporary question in his lecture, namely, whether
the frog had consciousness. This was a significant absence, for just
at this time anti-vivisectionists were arguing that animals’ con-
sciousness of pain made them morally unacceptable subjects for
live experimentation. The 1876 Cruelty to Animals Act turned
upon precisely the question of animal consciousness, insisting
that all vivisected vertebrates should be anaesthetized to prevent
the sensation of pain. No doubt Huxley’s lecture was in large
part, like so much of his lecturing, anti-theist grandstanding.
But by dismissing the question of consciousness and focusing
instead on the ‘enlightenment’ of experiment, it was also an
attempt to assert the right of biologists to continue using live
frogs and other animals in their work.
The best-known experiment with frogs is perhaps also the
cruellest: a frog plunged in boiling water, so it is said, will hop
out, but a frog placed in a vessel of cold water and heated slowly
will stay put and perish. This factoid serves many purposes.
Herpetologist Whit Gibbons of the University of Georgia recalls
hearing a Mississippi Baptist preach on the topic, using it as
an example of how ‘gradual habituation to a devilish situation
leads to acceptance of an even worse one’. Al Gore used the meta-
phor in similar fashion in his documentary An Inconvenient
Truth, to suggest that humans might go on accepting gradual
climate change until it is too late. In September 2009, Fox News’s
Glenn Beck employed the frog in another of its frequently used
senses, to capture the perceived bit-by-bit erosion of civil liberties
by the federal government. A video of him apparently placing a
frog in a pot of boiling water to illustrate this point was swiftly
placed on the Internet, forcing him to explain in a subsequent
show that the experiment was faked. Beck evidently enjoyed
‘frog gate’, as he called it, which for him was further illustration

109
of the foolish liberal sensitivities that he was trying to highlight
in the first place.
The ‘myth’ has, in fact, rather strong historical roots in
reality. In the 1870s, boiling frogs was a popular pastime for
physiologists, although they were not always precisely agreed
on why the activity was a useful one. Upon reading accounts of
these experiments, the fact that immediately strikes one is that,
in most of them, the frogs have had their brains removed. Thus,
Friedrich Goltz reported in 1869:

A normal frog if immersed in water which is gradually

heated, speedily becomes violent in his attempts to
escape. In striking contrast to this phenomenon is the
behaviour of the brainless frog, which . . . sits motionless
until it is dead from the excessive heat.8

To the modern reader, this observation appears to be one of

Swiftian asininity. A frog lacking its brain is hardly thinking
straight. To make sense of why the result seemed significant, we
need to understand what the frog had become for physiologists.
We have already seen an example of how the frog was consid-
ered as a set of qualities that could then be projected in their
purest form onto another carrier; for the early modern Edward
Topsell, frogs were a collection of unpleasant characteristics, and
the ultimate frog was a toad, in the sense that it possessed all
the key qualities of the frog to a purer and greater degree. Now,
for nineteenth-century physiologists, the frog had become a
piece of mechanical apparatus for answering a series of physio-
logical questions about nerves and muscles. Their über-frog was
one in which the brain had been removed, allowing its nervous
responses to be seen still more plainly and in their most auto-
matic state, uncomplicated by higher-level cerebral processes.

110
As one researcher put it, ‘the frog . . . is now admitted to be to a
great extent a reflex mechanism; . . . the brainless frog [is] a
much more perfect reflex apparatus than the normal one’. 9
This conception of the frog was so well-established that at
least two independent lines of batrachian-cooking experiment
were commenced without their authors knowing about one
another’s work.
Goltz’s discovery was surprising to his contemporary phys-
iologists for at least two reasons. The one that concerned the
American William Sedgwick was to do with the general prop-
erties of nerves and muscles at different temperatures; Sedgwick
expected a period of increased activity in the frog’s nervous
system before heat rigor set in, but Goltz did not report this.
Others were interested that decerebrated frogs showed no
response to heat in particular, since they displayed quite co-
ordinated and complex responses to other types of stimuli.
Still another approach to the phenomenon came from the
German physiologist A. Heinzmann, who speculated that the
dying frog demonstrated a general property of nerves. He sug-
gested that if the intensity of nerve stimulation were started
small enough, and increased gradually enough, then it might be
increased in intensity until it destroyed the nerve without ever
provoking a muscular reaction (because the muscle was waiting
for some kind of sudden step-change in the nervous action before
it responded). He found that even a normal frog would stay put
if the water were heated slowly enough.
In his 1897 book, The New Psychology, Edward Wheeler Scrip-
ture reported Heinzmann’s findings and added more: ‘It has
been found possible in 51⁄2 hours to actually crush a frog’s foot
without a sign that the pressure was felt by screwing down a
button at the rate of 0.03 mm per minute.’ Rather chillingly, he
added: ‘If a frog can be crushed or boiled without any evidence

111
A common frog
(Rana temporaria)
dissected to show
various anatomical
features, in George
Rolleston’s text-
book Forms of
Animal Life (1888).
Frogs have been
a mainstay of
laboratory teach-
ing since the late
19th century.

that he has noticed it, it is at least an interesting question of

what can be accomplished in this direction with human beings.’10
Leading biologists of the present day are anxious to dispel the
‘myth’ of the boiled frog. The Curator of Reptiles and Amphib-
ians at the Smithsonian’s National Museum of Natural History
in Washington, dc, asserts with admirable firmness: ‘Well that’s,
may I say, bullshit.’11 These debunkers generally agree that frogs
are way too jumpy to stay put anywhere, least of all in quite such
an unpleasant environment. Yet at least two nineteenth-century
experimenters really did think they had performed the feat of
live-boiling, and it is unlikely that they were lying or totally
incompetent. Perhaps the frogs were already dead when they
were put in the water, or died some other way before it really

112
 One of several
illustrations of
frog dissections
in Roesel von
Rosenhof’s Historia
Naturalis Ranarum
Nostratium (1758).

heated up. Or perhaps they were placed in vessels from which

it was actually not possible to jump. Another more subtle possi-
bility comes in relation to the precise theories that experimenters
were trying to prove. Their definitions of life and death were
based upon particular understandings of nerve and muscle,
and it may be that in looking for ‘heat rigor’ they were not
thinking about life and death in the everyday sense. Hence, the
decerebrated frog, which an ordinary person would definitely
consider dead, was not dead for them.
This anxiety about the definition of death continues to leach
into related anxieties about frog cruelty. When is a frog dead?
When does dissection become vivisection? The reactive frog
that was once celebrated for its tenacious hold on life must now,

113
Martin
Kippenberger’s
Feet First (Prima
i piedi, 1990)
captures anurans’
ubiquitous role
as fall-guy and
sacrificial victim.

for reasons of humaneness, be defined as dead. Decerebrated

frogs are still used extensively by university-level students and
lecturers in physiology: dead enough to feel no pain, but alive
enough to respond to it. The preparation of these frogs is
known as pithing; a needle is inserted near the top of the spinal
column and moved around inside the skull so as to destroy the
brain. The technique was developed as a means of slaughtering
animals for consumption, but began to be used on frogs in the
nineteenth century. It remains in use today, and has produced a

114
seam of guilty humour about the frog’s being ‘pithed off ’ about
its laboratory treatment.
In general, the scientific use of frogs, whether alive or dead,
still fails to provoke as much criticism as the use of mammals,
and anecdotal evidence suggests that herpetologists are not
significant targets for animal rights activists. In 2008, Animal
Aid protested that uk statistics for 2006 showed ‘12,459 amphib-
ians were used in 20,616 procedures’, and that 17,728 of these
were without anaesthesia, even though ‘amphibians are sent-
ient beings capable of feeling stress and pain’.12 The story raised
little interest, however, and it is worth noting that procedures
performed without anaesthetic may be done in this way pre-
cisely because they are quite harmless and pain-free for the
frogs. Ecological herpetologists, who by and large care for frogs
more than it is fashionable for a scientist to admit, complain
that experimental restrictions are actually too tight. uk law, they
say, prevents them from carrying out quite innocuous experi-
ments (such as trying tadpoles in different kinds of pond water)
by imposing disproportionate and onerous paperwork upon
the process.
To date, six Nobel Prizes in physiology and medicine have
resulted from investigations based exclusively on frogs, and there
have been perhaps twice this number in which frogs played some
role. Two Ig Nobel Prizes have also rewarded frogs. Whether used
kindly or cruelly, frogs have been indispensable to a variety of
experimental traditions throughout modern history.

115
5 Evolution on Fast-forward

In 1844, Robert Chambers – best known today for his dictionary

– contemplated evolution by considering the frog. He imagined
how frogspawn might appear to a mayfly during its brief 24 hours
of life; it would have no idea of the drastic changes that were
going to unfurl in their little, wriggly bodies, long after the mayfly
itself was dead:

Suppose that an ephemeron, hovering over a pool for its

one April day of life, were capable of observing the fry of
the frog in the water below. In its aged afternoon, having
seen no change upon them for such a long time, it would
be little qualified to conceive that the external branchiae
[gills] of these creatures were to decay, and be replaced
by internal lungs, that feet were to be developed, the tail
erased, and the animal then to become a denizen of
the land.1

Human beings, suggested Chambers, were in no less a state of

ignorance when it came to nature around them: what appeared
to be fixed species were, in the grand scheme of things, in a
process of transformation from one form to another. Then as
now, tadpoles and frogs are an almost irresistible image for the
process so controversially described by Chambers. With gills

116
 Maria Sibylla
Merian’s 17th-
century rendering
of frog metamor-
phosis was drawn
long before the
frog’s development
came to function
as a metaphor
for the process of
evolution itself.

that turn into lungs, and legs that bud and grow to propel them
from the water, it is as though they fast-forward through the
slow evolutionary exodus from sea to land.
Of course, the frog is not in any literal sense a transitional
organism between fish and mammals, but in the minds of many
it has stood in that intermediary place. Older views of nature
described a ‘great chain’ of lower to higher organisms; in 1849,
for example, the Swiss-born American zoologist Louis Agassiz
claimed that the development of the frog placed it between the
fish and the reptile. When the great chain metaphor was given
an evolutionary twist, the frog retained its place. Darwin, writing
his On the Origin of Species (1859), knew that dissections of fish

117
and land animals by the comparative anatomist Richard Owen
had already demonstrated that the swim bladders of fish were
the ‘same’ as the lungs of land-living animals. Thus Darwin took
it for granted that vertebrate life had begun in the water and
subsequently moved onto the land. Frogs in this scheme too
were ‘between’ fish and reptiles.
The very class Amphibia appears to owe its existence to evo-
lutionary theory. Before Darwin, amphibians and reptiles were
lumped together in a single class. In the authoritative edition of
his Systema Naturae (1758), Linnaeus named the Amphibia as
one of his four classes of vertebrates (alongside birds, mammals
and fish). However, within the Amphibia, frogs and newts
were grouped in the order of reptiles. In 1798, the great French
comparative anatomist Georges Cuvier divided the vertebrates
essentially the same way, but called the class that included frogs
by the name of Linnaeus’ order: ‘reptiles’. Even Lamarck, an evo-
lutionist and opponent of Cuvier, followed suit. Thus, although
The Swiss
the word ‘amphibian’ was in use, it did not carry the imperative physiognomist and
force of categorization that it does today. clergyman Johann
Caspar Lavater
There were just a couple of people who distinguished am- created his series
phibians as a meaningful category before Darwinian evolution ‘From Frog
to Apollo’ to
was widely accepted, but the person to put the taxonomic dis- demonstrate
tinctiveness of the Amphibia in the textbooks did so with an human uniqueness,
in the light
evolutionary twist: Ernst Haeckel. Haeckel (1834–1919) was, of recent trends
according to the historian of biology Erik Nordenskiöld (1935), to place humans
‘the chief source of the world’s knowledge of Darwinism’, but amongst animals.
The frog repre-
his methodology and style were distinctive. Whereas English sented the lowliest
evolution had been built substantially on comparative ana- animal in his
version of the
tomy and palaeontology, Haeckel added a third type of evidence: chain of being.
embryology. ‘The facts of embryology alone would be sufficient Coloured etchings
by Christian von
to solve the question of man’s position in nature’, he wrote boldly Mechel after
in his History of Creation (1868).2 Lavater, 1797.

119
Haeckel’s frogs in
his Kunstformen
der Natur (1904).
Haeckel’s account
of evolution
cemented frogs
in their status in
between water
and land animals.

Haeckel’s theory of embryos was known as ‘recapitulation’,

or the ‘biogenetic law’. In essence, this meant that the evolu-
tionary process that had led upward from primitive organisms
to man was echoed in the development of embryos. Embryos
passed through primitive states resembling the embryos of their
ancestors before reaching the higher form of their species. One
phase of the embryological development of man, wrote Haeckel,
‘presents us with a change of the fish-like being into a kind of
amphibious animal. At a later period the mammal . . . develops
out of the amphibian’. Hammering the point home, he went on:

120
 it is in precisely the same succession [as we see in the
embryo] that we also see the ancestors of man, and of the
higher mammals, appear one after the other in the earth’s
history; first fishes, then amphibians, later the lower, and
at last the higher mammals.3

Thus amphibians had a particular place in the embryological

evidence for evolution.
What made amphibian embryos important from an evolu-
tionary point of view was, according to Haeckel, their lack of an
amnion. The amnion, a membrane that creates a sac surround-
ing the developing foetus, is present in birds, mammals and
the class that we now call reptiles. The fact that frogs’ eggs (like
fishes’) do not possess an amnion put them, for Haeckel, below
the others, and made them distinct from their usual classmates
the reptiles. Haeckel considered the non-amniotic amphibians
as the progenitors of the two branches of amniotic animals:
reptiles, subsequently giving rise to birds, and mammals. In the
light of this, Haeckel expanded the classes of vertebrates beyond
the standard number of four, making the Batrachians (his
preferred name for the Amphibia) a class in their own right.
Batrachians were the top of the non-amniotes, the closest of their
kind to the amniotes in terms of evolutionary development. By
the same logic of recapitulation, frogs were at the top of the
Batrachians, the very pivotal organism between Haeckel’s two
fundamental types of vertebrate, amniote and non-amniote.
Thus, the cognitive category Amphibia – as opposed to its more
inclusive forerunner, reptiles – was constructed with evolution
in mind. Before then, it made no sense to separate frogs, toads
and newts from lizards, but now that the transition from water
to land was scientifically significant, it made sense to consider the
animals that embodied the possibility of the change. Haeckel’s

121
A simplified version
of Haeckel’s
schematic hierarchy
of extant verte-
brates, 1868. The
three columns
represent, from
L–R, main class,
class and subclass.
Batrachians occupy
the intermediary
position between
the two main
classes.

evolutionary construction of the category spread rapidly, giving

impetus to many projects of research.
Haeckel’s one-time student Wilhelm Roux (1850–1924) was
seduced by the topic of embryos, and his powers of self-promo-
tion have ensured that his achievements now take a prominent
place in the history of embryology. Roux was not as interested
in the grand evolutionary questions as his mentor, but rather
in the immediate causes of embryonic development. How did
embryos ‘know’ what pattern to follow? Were the forces inter-
nal, or did they come from the environment? The frog embryo,
which was already in experimental use by his colleagues at the
University of Breslau, was a good choice for finding out. Unlike
mammalian embryos, hidden away in the womb, the frog is fully
visible throughout its development.
Edward Pflüger (1829–1910), professor of physiology at
Breslau, was one person already using the frog to answer those
internal versus external development questions, generally find-
ing that the two interacted to determine how the frog turned

122
 A phylogenetic
(evolutionary
history) version
of Haeckel’s
hierarchy, showing
batrachians as the
ancestors of all
land vertebrates,
1868.

out. In one experiment he rotated frogs’ eggs on a slowly spin-

ning wheel, and found that the axis along which the egg divided
and developed was altered by the absence of a consistent gravita-
tional force.
Roux wondered about a similar set of questions. Did each cell
have its own internal developmental drive (‘self-differentiation’),
or did cells somehow ‘talk’ to one another (‘dependent differ-
entiation’)? Roux caricatured Pflüger as a believer in dependent

123
Half-embryos of
a frog, created by
Wilhelm Roux,
1890.

differentiation and set him up as the intellectual straw man that

his own experiments would destroy. In the most famous of these,
Roux carefully poked a hot needle into one half of a two-cell frog
embryo, killing one cell (or ‘blastomere’). If development were
dependent upon the environment (the position he ascribed to
Pflüger), the remaining cell should compensate for its neigh-
bour’s loss and produce a normal embryo. If development were
independent as Roux claimed, the lone cell should continue
regardless to produce a half-embryo.
With great fanfare, Roux reported in 1888 that he had
indeed produced a (short-lived) half-tadpole. Roux was right
and Pflüger was wrong. This was Roux’s story, at any rate. Hans
Driesch (1867–1941), another of Haeckel’s former pupils then at
the Naples Marine Biological Station, tried a similar experiment
with sea urchin embryos and got the opposite result: normal,
albeit small, embryos. Driesch rather generously explained away
the apparent contradiction: development probably was internal
as Roux claimed, but some regulatory ability was available to

124
compensate when things went wrong. Driesch’s experiment
had provoked this compensatory mechanism; Roux’s would
have done too, but he had probably not succeeded in killing the
blastomere completely. Driesch’s experiment worked because
instead of stabbing one cell of the two-cell embryo, he physically
shook it apart from its twin.
Roux’s disagreement with Driesch highlights an issue that
we have already seen in chapter Four, namely what happens
when scientists start to downplay the frogness of their experi-
mental subjects and instead treat the frog as a standard organism.
Roux was quite explicit about this; he thought that develop-
ment was a mechanical system and that studying one organism
would yield results relevant to all. But as the urchin showed,
this was not true, or at least compelled the experimenter to
come up with a reason why the test showing the ‘wrong’ result
was invalid.
Most accounts of Roux’s fabled experiment fail to convey its
practical difficulties. It was actually very hard to carry out; the
cells were hard to puncture, and once punctured, the remaining
cell was hard to keep alive (only about 20 per cent made it).
Most early twentieth-century amphibian experimenters did
not consider the basic fact that pure water is toxic to embryos,
drawing out their vital salts through osmosis. Moreover, in
some of Roux’s trials, a normal embryo did develop, and in
these instances he was forced to come up with compensating
explanations to show that the experiment had gone wrong.
Similar challenges afflicted the embryological work of Hans
Spemann (1869–1941) at the University of Würzburg. Spemann
was also working at the puzzle of self- versus dependent dif-
ferentiation, having switched to the frog’s close cousin, the
salamander, for his experiments. Taking a hair from his baby
daughter, with the most dextrous patience he tied it around the

125
embryo of a two-celled salamander embryo. Cloned frogs, or
artificial twins, can supposedly be created at home using a tech-
nique much the same as Spemann’s. The spawn need to be
very fresh, as the technique must be done at the two-cell stage.
This can be seen, in large-spawned species, against a white back-
ground and under a magnifying glass. A hair loop can be used
to cut the cells along their join, and the separated cells should
now, in theory, develop into two identical tadpoles.
Spemann, however, tightened the noose without proceeding
all the way to a cut. He found that by this method he could
make the embryo grow into twin, conjoined specimens, and
the experiment was repeated with frogs to the same effect. But
a slight repositioning of the noose would result in a single larva
on one side of the hair, and a blob of meaningless tissue on the
other. Why should this be? What made one part of the embryo
turn into one thing, and another into something completely
different? Spemann turned to the eyes of frog embryos to try
to find out, asking what would happen if one moved a bit of
embryonic tissue; would it turn into tissue appropriate to the
original or the new location?
In a series of experiments, Spemann excised the eye anlages
– the cells that would turn into the lens – from frog embryos
and transplanted them elsewhere in the embryo. If self-differ-
entiating, they should develop into lenses (so-called ‘free lenses’)
wherever they were transplanted. If dependent in their differ-
entiation, they should not. In 1901, Spemann concluded that the
lens did not develop unless it was in contact with the tissue that
normally neighboured it. But results coming in from Czech and
American biologists claimed that free lenses were possible.
These results had been obtained using Rana esculenta and R.
palustris, rather than the R. fusca used by Spemann, and when
Spemann switched to R. esculenta, he too obtained free lenses.

126
 These results have become the stuff of scientific myth. With
many twists and turns of method and interpretation, Spemann’s
frogs came to form the basis of today’s embryological discipline,
developmental biology. But in the process, the frogs have been
stripped of their species specificity. How many of today’s devel-
opmental biologists know that the species described by their
hero, ‘Rana fusca’, is not even a real species, but rather a local
variant of the European common frog, Rana temporaria? Just as
in Roux’s case, it takes work to learn to use a frog, and more work
still to strip it of its frogness and transform it into a standard
organism that works the same as all other species.
By contrast to Spemann’s success story, Haeckel’s legacy was
to prove tragic for the Austrian zoologist Paul Kammerer (1880–
1926), whose attempts to demonstrate developmental evolution
in action concluded in a self-administered gunshot to the head.
Unlike Roux, and perhaps Spemann, Kammerer cared a great
deal about the frogness of his organisms and this know-how,

A recent example
of an experiment
inspired by early
amphibian embry-
ology; a tadpole of
the western clawed
frog, Xenopus
tropicalis, has
been twinned
by transplanting
a Spemann’s
organizer from
another tadpole
into its belly.

127
Table of normal
development of
Rana fusca roesel
(probably Rana
temporaria) by
Friedrich Kopsch,
1952.

re-cast as lack of objectivity by his enemies, was a partial cause of

his downfall. Kammerer was inspired, and ultimately destroyed,
by that most tantalizing feature of evolutionary theory; it was,
as Chambers had realized, impossible to demonstrate within a
human lifespan. (The American T. H. Morgan claimed in the
1920s that his colonies of fruit flies provided such evidence, but

128
many were not convinced and ascribed the flies’ mutations
merely to their unnatural life within Morgan’s lab.) For those
raised on the Haeckellian notion of recapitulation, there was a
powerful appeal in the evolutionary metaphor of amphibian
development; it seemed to say something about the course of
evolution. Thus Kammerer set about trying to modify the phys-
ical characteristics of Amphibia, and, moreover, to investigate
whether such modifications could be inherited by their offspring.
In short, he was trying to speed up evolution, using the organ-
ism that already seemed to do so.
Kammerer was employed in the old exhibition hall of the
Viennese Prater amusement park, recently converted into a state-
of-the-art vivarium: a research facility equipped to maintain
temperature and humidity exactly as required for its many
zoological specimens. During the years before the First World
War, Kammerer was able to set up complex, multi-generational
experiments at the vivarium, inducing heritable terrestrial char-
acteristics in aquatic salamanders and aquatic characteristics in
terrestrial ones. Of all his research, Kammerer was most pleased
with his salamander experiments (along with those using the
sea squirt Ciona), but his most famous – or rather infamous –
experiments involved the midwife toad, Alytes obstetricans.
Like most frogs and toads, the males of these species sit on
top of the female while mating and hold on for several days,
but unlike their aquatic cousins they have no need of rough
patches (nuptial pads) on their thumbs to stop them slipping
off. Kammerer induced his normally terrestrial specimens to
breed in the water, then rescued and reared the fertilized eggs
that sank to the bottom. By these means, Kammerer claimed to
induce the appearance of nuptial pads in the next generation of
midwife toads. Astoundingly, these pads appeared to be passed
onto their offspring.

129
The midwife toad
(Alytes obstetricans),
subject of Paul
Kammerer’s ill-
fated experiments
in evolution.

Although Kammerer’s science now seems extraordinary, it

had its echoes in both fact and fiction of the time. As historian
Sander Gliboff has discovered, Czech experimenters had been
working on salamanders as early as 1876, trying to see whether
environmental conditions could alter their pigmentation. Other
work at the Vienna vivarium concerned the regeneration and
transplantation of limbs in salamanders, newts and frogs. The
scientific significance of amphibians at this time is further illus-
trated by the fact that a small clutch of evolutionary, amphibious
sci-fi appeared in the 1920s and ’30s: Mikhail Bulgakov’s The
Fatal Eggs (1924–6); Alexander Belyaev’s The Amphibian (1928);
Karel Čapek’s War with the Newts (1936); and the Soviet film
Salamandra (1928).
Bulgakov – a trained doctor – set his blackly comic frog novel
somewhere very like the Vienna vivarium. The Fatal Eggs centres
upon Professor Persikov, a scientist working at the Zoological

130
Institute in Moscow who accidentally stumbles upon a red ray The Mallorcan
midwife toad
which bequeaths enhanced life-powers upon the organisms (Alytes muletensis)
that it touches. The first organisms to receive these powers are has been a happier
amoebae, and observing their ‘furious reproduction’ and rapid research topic
for herpetologist
maturation, Persikov – an amphibian specialist – immediately Richard Griffiths.
thinks to try the ray upon frogs. The results are remarkable: The species
was thought
extinct but was
In the course of two days thousands of tadpoles hatched rediscovered in
the late 1970s;
from the grains of roe. But as if that were not enough, in the since then captive
course of a single day the tadpoles grew extraordinarily breeding has been
achieved and
into frogs so vicious and gluttonous that half of them were understanding
gobbled up on the spot by the other half. And then those of its ecological
that remained alive began to spawn in no time at all, and requirements
greatly improved.
in two days, by now without any ray, bred a new genera-
tion, and a quite innumerable one at that.4

131
 But before he can experiment any further, Persikov’s equip-
ment is requisitioned by the State. The country’s chickens have
all died in a mysterious plague, and the authorities want to
replace them by breeding up superior stock from imported
German eggs. Unfortunately, however, the hens’ eggs have been
accidentally replaced by snakes’ eggs, and under the influence
of the ray these emerge as monstrous, people-eating prodigies.
The story is in large part political satire, a tale of humans
fatally transformed by the red ray of Marxist-Leninism. The
batrachian nature of the people, involuntary subjects of politi-
cal experiment, is evidenced by their frequently ‘croaking’ voices,
especially on the telephone. Urban crowds pullulate like frog-
spawn; the military and civilian masses are snake-like forces that
eventually crush Persikov and his assistants in just the manner
that the constricting serpents kill their first victims. On a simpler
level, however, frogs were for Bulgakov – just as they were for
Kammerer – the natural experimental organism of evolution.
In August 1926, after years of doubt over Kammerer’s results,
the American herpetologist G. K. Noble travelled to Vienna to
observe the sole surviving specimen of midwife toad. He returned
to New York, and reported bluntly in the journal Nature that the
pads had been faked; black ink had been injected under the skin
to produce them. One month later, Kammerer shot himself. Most
commentators have taken Kammerer’s suicide as an admission
of guilt, but his sympathetic biographer Arthur Koestler gives
alternative explanations, ranging from his difficult private life
to confusion and embarrassment over the fraud; Koestler sug-
gests that the ink injection might even have been perpetrated
without Kammerer’s knowledge by anti-communist saboteurs.
Kammerer’s reputation as a ‘wizard with lizards’ counted at
least as much against him as it did for him, for it made it very
difficult for others to replicate his results, or even feel inclined

132
to try. Even picking the eggs out of the water and rearing them
to adulthood was a painstaking task. Kammerer’s experiments
also lasted many years. The whole point was to see whether
altered characteristics were heritable, requiring generations of
breeding to find out. (Moreover, the nuptial pads were only
present during the breeding season, so killing a precious speci-
men to preserve evidence – as his critics demanded – meant
one less for breeding.) Part of the criticism of Kammerer can
therefore be ascribed to an emerging split between biology and
natural history: between those who prescribed a lengthy
engagement with the animal in its own right, and those who
believed they knew animals through more general laboratory
or statistical methods.
Commentary on the midwife toad episode peaked in the
mid-twentieth century, and writers were more or less unanimous
in painting Kammerer’s guiding theory as a fundamentally silly
one (hence necessitating fraud for its demonstration). The
word most often used for Kammerer’s work is ‘Lamarckian’, by
which commentators have meant an anti-Darwinian biology
of acquired inheritance – antelopes that purposefully evolve
into giraffes by stretching their necks ever higher for leaves. In
fact, central and Eastern European biologists such as Kammerer
and his boss Hans Przibram counted themselves firmly as
Darwinians.5 However, their Darwinism was refracted through
the lens of Haeckel. Bulgakov’s description of the ray-zapped
tadpoles reveals just such a hybrid understanding of evo-
lution: there is competition among members of the same
species (‘half of them were gobbled up . . . by the other half ’),
but also changed bodily features that persist in the organisms’
inheritance (‘without any ray [they] bred a new generation’).
This Haeckellian version of evolution was a mis-match with the
Anglo-American tradition.

133
 Kammerer was, in fact, sympathetic to the critical suggestion
that his experiments might not demonstrate forward evolution.
It was quite possible that instead he was inducing regression:
stimulating the display of ancestral characteristics that were no
longer necessary for the species in normal life. He was in good
company, as Gliboff has found; some German biologists of the
1920s suggested that creatures could regress to a primitive form,
endowed with more potency and flexibility than their usual,
highly evolved variety. It was even possible that this bodily
plasticity was itself a heritable characteristic: that a creature
which had regressed to its more potent form made it easier for
its offspring to do the same. Hence the research at the vivarium
regarding amphibians’ regeneration of missing limbs. In this
context, Kammerer’s work does not look like an attempt to
fast-forward Lamarckian evolution as his critics have claimed,
but rather to demonstrate the recapitulatory framework of
Haeckellian evolution.
Such youthful flexibility in the adult amphibian suggested
something biologically special, and indeed in the 1920s a theory
about it was developed. It was called paedomorphosis, and was
announced by its proposer as a ‘re-statement’ of Haeckel’s bio-
genetic law. Paedomorphosis described a phenomenon whereby
adult creatures displayed features normally associated with the
juvenile of the species. The phenomenon was most dramatically
displayed by animals whose juvenile and adult characteristics were
normally separated by a Rubicon of complete metamorphosis;
once again, amphibians were in the scientific limelight. Nature
contains several species of Amphibia that look like over-sized
tadpoles but can nevertheless breed, of which the best known
is probably the axolotl. Having identified the thyroid gland as
the thing that controlled metamorphosis, in 1919 scientists
experimented to see if frogs too could be induced to breed in

134
their tadpole state. By removing the thyroid anlage in embryos
of the wood frog, Rana sylvatica, they obtained giant larvae, two
to three times normal size. The tadpoles never matured, but
scientists could not get them to breed in their larval form.
In his retirement, Spemann made his most ‘fantastical’ sug-
gestion yet: that an entire frog might be generated by implanting
the early embryo nucleus from one egg into another, unfertilized
one. Would the organizer principle be able to work on an egg that
was not primed for development? In 1952, American researchers
Robert Briggs and Thomas King managed just this feat using
the high school lab favourite, the northern leopard frog, Rana
pipiens, raising tadpoles from unfertilized eggs that had had
an embryonic nucleus inserted. Briggs and King found that the
earlier the developmental stage of the donor embryo, the greater
the chance of success – the development of an apparently normal
tadpole. Nuclei removed from embryos that had matured so
far as to develop a nervous system were not successful at all. At
Oxford in the late 1950s and early 1960s, John Gurdon (1933–)
attempted to replicate the feat with later-stage nuclei. Gurdon
used a different species of frog – Xenopus laevis – and inserted
nuclei from fully developed tadpole cells into eggs with their
nuclei removed. Gurdon’s eggs went on to develop, each one pro-
ducing a larva genetically identical to its donor. However, they
died without metamorphosing into adult frogs, just like all
attempts ever since – and no one knows why.
If I were a member of Xenopus laevis rather than Homo sapiens,
I might point to Gurdon’s period as the great era in which my
species commenced its colonization of the globe. There were
just a few breeding colonies of these creatures in Europe at the
beginning of the twentieth century, but by its end the species
had become the experimental organism par excellence for devel-
opmental biologists around the world, and had even been sent

135
 into space.6 X. laevis was popularized by Lancelot Hogben
(1895–1975), the British socialist who, having moved to South
Africa, found it such a ‘godsend’ for research that he named his
house after it, and brought a colony of the species back with
him when he returned to the uk in 1930. Hogben’s research con-
tinued the hormonal strand of frog biology that had begun with
the thyroid induction or suppression of metamorphosis, and
in London his team began using X. laevis to try and develop a
human pregnancy test. At that time, the only method involved
injecting rodents with the urine of possibly pregnant women,
then killing the mouse or rabbit to see the state of their ovaries.
The frogs, however, were such sensitive registers of human
chorionic gonadotropin (a hormone produced by pregnant
women) that it was not necessary to kill them to confirm the
result. Instead, they themselves would ovulate only eight to
twelve hours after a positive injection, laying their eggs for all
to see. For the next 30 years, hospitals around the world kept

The African clawed

frog (Xenopus
laevis), favoured
subject for
hormonal and
developmental
laboratory
investigation.

136
 Xenopus laevis was
the most reliable
test for pregnancy
in the 1940s and
’50s.

colonies of X. laevis in their basements, ready to confirm doubt-

ful pregnancies. Excess specimens were sometimes released into
the wild, and occasionally they escaped, forming feral colonies.
Ironically, it was very difficult to breed the frogs in captivity,
but gradually their keepers discovered how they could take
advantage of the hormonal injections to produce actual breed-
ing – as opposed to just egg-laying – at will.
Once it possessed the convenient quality of breeding on
demand, Xenopus rapidly became a favoured animal for all
sorts of research. Using frogs elicited few pangs of conscience,
and now this tough species could be easily procured without
resorting to Helmholtz’s annual frog-hunting frenzy. But X.
laevis also has properties that ought to count against its use;

137
slow to mature, it has an anomalous pattern of development and
a complex genetics, with four copies of each gene rather than
the usual two. Tradition is, however, its own justification, for
learning to breed and keep a reliable colony of usable animals
requires a considerable investment of time and skill. Once scien-
tists have learned how to breed and look after a species, it makes
sense to stick with it. When it comes to molecular biology and
genetics, it is absolutely essential that everyone is talking about
the same species. A ‘model organism’ needs to be like a stan-
dardized instrument that everyone understands and knows
how to calibrate. Hence scientists have by and large stuck with
X. laevis. By about 1990, publications on X. laevis in develop-
mental biology outnumbered those on other amphibians by a
factor of approximately 50 to one, although these days its cousin
X. tropicalis (which possesses the standard dual-copy of genes)
is increasingly used for genetic research. X. laevis remains a
favourite for molecular biology; scientists are using it today to
find ways of replacing damaged tissue – such as heart, brain or
blood cells – in humans.
The fantasy of regenerating humans by recourse to the frog’s
body is one that the 1920s sci-fi writers would have recognized
instantly. Just like theirs, it keys into a popular discourse of
evolution: of taking the species on to the next level, or of the
natural flexibility and potency contained within the meta-
morphosing body. As such, it has something in common with
Haeckel’s biogenetic law. Central to the conceptualization of
amphibians, this ‘law’ retained a remarkably persistent intuitive
appeal long after all its specific claims had been overridden. A
sex education manual of 1917 asserted: ‘Every frog which reaches
maturity repeats again the story of his race in his earlier stages.’7
Even in 1977, the biologist Stephen Jay Gould claimed that most
of his colleagues still secretly thought there was ‘something to it’.8

138
When Richard Wassersug began his long career in frog studies
around the same time, he found that Haeckellian perspectives
were still shaping batrachian classification. There were, he recalls,
‘two conflicting phylogenies [evolutionary family trees] . . . one
based on adult anurans and the other on tadpoles’. To find out
which of these was right, Wassersug began looking in detail at
tadpoles. His work focused on their complex mouthparts, and
caused him to realize that frogs were extremely specialized and
well-adapted to life in their larval form. They were not, as adult-
based phylogenies implied, merely ‘“fish” or imperfect frogs’.
Such phylogenies were implicitly Haeckellian in form, while
Wassersug’s work was, in his own words, ‘a non Haeckellian
line of inquiry’.9
Haeckel is rather like the disreputable uncle at the family
party of developmental biologists, and recapitulation is today a
dirty word in science. Whether scientists like it or not, Haeckel’s
frogs have inspired many branches of developmental science
through the nineteenth and twentieth centuries. Thanks to
Haeckel’s way of seeing things, the frog’s emergence from water
to land has become emblematic of a whole chapter in the evo-
lution of life. Its embodiment of life’s power, waiting to unfold,
has placed it at the heart of both successful and unsuccessful
science. The frog has been a vivacious embryo, a chemically sen-
sitive collection of developing cells: even, perhaps, a source of
human regeneration.

139
6 Of Frogs and Fruitfulness

Swarming and flickering in a shallow pond, tadpoles are a most

vivid symbol of nature’s irrepressible urge to reproduce. Frogs can
appear in vast numbers overnight, a phenomenon of synchro-
nized reproduction that is known by biologists as ‘explosive
breeding’. If a frog laid its eggs on its own, the whole lot might
be eaten by a predator. But if every frog oviposits at once there
is a good chance that, although many eggs will be eaten, some
eggs from any given individual frog will survive: hence its selec-
tive advantage. The Ancient Egyptian tadpole hieroglyphic
commemorated this astonishing fecundity of the frog, standing
for the otherwise unthinkably large number of 100,000.
The appearance of the frogs that produce and fertilize all these
eggs can be no less astonishing. All sorts of strange explanations
have been advanced for the sudden mass appearance of frogs in
nature. A Victorian traveller to Panama reported:

Toads and other frog-like animals are most numerous

during the wet season . . . So prodigious is their number
after rain, that the popular prejudice is that the drops of
rain are changed into toads; and even the more learned
maintain, that the eggs of this animal are raised with the
vapour from the adjoining swamps, and being conveyed
to the city by the rains, are there hatched . . . After a night

140
 Frogs mating
(either the pool
frog, Pelophylax
lessonae, or
the marsh frog,
Pelophylax
ridibundus),
illustrated in Roesel
von Rosenhof’s
Historia Naturalis
Ranarum
Nostratium (1758).

of rain, the streets are almost covered with them, and it

is impossible to walk without crushing some.1

Frogs emerge thus en masse in order to mate. George Orwell

found the sight a deeply appealing one, ascribing to the male
(perhaps wistfully?) a ‘phase of intense sexiness’. Orwell found
that ‘if you offer him . . . your finger, he will cling to it with sur-
prising strength and take a long time to discover that it is not a
female toad’.2 This triumph of urge over precision has frequent

141
 unfortunate results. Females may drown in the iron grip of their
mates, and even passing fish may be harmed or killed if they fall
into the male’s clutches. The sight of a pond heaving with randy
frogs, rolling about in couples, threesomes and even foursomes,
has given the Anura a powerful life with human fertility and sex.
Recently, in European culture at least, the emphasis seems
to be on the mass mating rather than the spawning: the sex, not
the fertility. The most striking recent example treats the anurans
as did Orwell: as pure sex, rather than reproduction. It occurs
in Patrick Süskind’s 1985 novel Perfume, whose anti-hero is the
The massed mating strikingly named Grenouille (frog). Grenouille murders women
of frogs in spring-
time has made for their scent, and on the day that he is due to be executed for
them a symbol his crimes, the scent of the women – including that of his
of unrestrained
sexuality for
intended final victim – overpowers the crowd and changes the
many observers. scene from one of public execution to a giant batrachian orgy.

142
 This unsettling
detail from
Bosch’s Garden
of Earthly Delights
(1490–1510) places
batrachian sexuality
in a decidedly
negative theo-
logical light.

There is also a limited amount of evidence connecting earlier

European culture with anurans as agents of fecundity,3 although
such links may originate in the characteristic swelling-up of
frogs and toads – reminiscent of the fruitful womb – rather than
their breeding behaviour. The sixteenth-century philosopher
and physician Paracelsus called frogspawn ‘the sperm of the
world’. The links with fertility are there in a negative sense too;
if used correctly, frogs can actually prevent it. In England, a
woman that places a frog in her mouth and spits three times
will not conceive that year.4 In the 1950s, the deputy of China’s
National People’s Congress famously exhorted women to swal-
low live tadpoles as a means to restrict their fertility; when an
experiment was carried out to test the efficacy of this traditional
peasant contraceptive, it performed disastrously and the advice
was retracted.5
In lands that are reliant on periodic flooding for agriculture,
the appearance of frogs coincides with that of the water that

143
Ancient Egyptian
depiction of Seti i
(reg. c. 1294–1279
bce) making an
offering to the
frog goddess
Heket.

brings life to the crops; the frogspawn is a sign that the people
too will eat and reproduce for another year. No wonder, then,
that in Ancient Egypt – dependent upon the flooding Nile for
its agriculture – the goddess of fertility and birth took the form
of a frog. Her name was Heket; midwives were her servants, and
women often kept amulets in her shape to protect them during
childbirth. (In a somewhat similar vein, Pliny the Elder later
recommended that a stick which had been used to get a frog
out of a snake would speed up labour; unfortunately, he neg-
lected to mention where such a stick might be found or precisely
how it should be deployed.) Statuettes of Heket exist from as
far back as c. 3000 bce; she appears in a papyrus myth of the
Middle Kingdom (c. 1600 bce) and there is evidence of cults in
her honour in both Middle and Upper Egypt. Although she was
most often pictured as a frog-headed woman, Heket was also
portrayed as a whole frog, or even as a frog atop a phallus. This
last image makes clear Heket’s connection not just with birth but
also with fertility, a connection that was reinforced by the cults

144
 Although many
Ancient Egyptian
women kept Heket
frog amulets to
bring them luck
in childbirth, this
travertine example
(2950–2770 bce)
is thought to have
been kept in a
temple due to its
relatively large size
(approximately
15cm in each
dimension).

that considered her to be a consort of Khnum. Khnum was god

of the Nile’s source, and moulder of children’s bodies from the
silty clay that washed in each year with the floods. Together, the
floodwater and its frogs brought humans to birth.
Dating from a similar period to the Heket papyrus, the
Manduka Sukta (frog hymn) of the Hindu Rigveda praises the
life-giving monsoon in batrachian form:

Green and Spotty have vouchsafed us treasure.

The Frogs who give us cows in hundreds lengthen our
lives in this most fertilizing season.

The words of Brahmans, who have waited until the appointed

time to fulfil their religious vows, are rather sweetly compared
to the sound of the frogs greeting the rain – for which they have

145
also patiently waited. A few hundred years later, Europeans, too,
caught onto the connection between frogs and rain. European
tree frogs (then classified Rana arborea, now Hyla arborea) reliably
strike up their chorus before rainfall, causing them to be dubbed
‘living barometers’.6 In April 1789, the British New Lady’s Maga-
zine described a more convenient device ‘lately discovered at
Paris’ (although probably originating in Germany): a bottle with
water and earth in the bottom; a ladder ascending from them;
and a tree frog. The frog would perch at the top of the ladder as
long as fair weather could be expected to last, but would return
to the water when rain was due. Any English lady attempting to
build this herself would have struggled, since Rana arborea is not
native to Britain and the instruction to ‘take one of those small
green frogs which are found in hedges’ would have resulted in a
long, long search.
Many Native American peoples enshrine in their traditional
tales the role of the frog in bringing rain and fertility. Several
tribes share versions of a myth about a giant monster, Aglebamu,
who swallowed the water source of the people. In some versions,
he is like a giant frog, and in others he is turned into a frog by the
hero Glooskap. Either way, he is made to release the water that
the people need.
A Navajo tale tells a different story about how Frog brought
This frog (likely
water to the people. After Coyote stole fire, it is said, he acciden- Hyla arborea)
tally set the mountain alight, and First Woman could not get climbs a glass
vessel in Roesel
water from the spring, the river or the lake to put it out. The last
von Rosenhof’s
place she could think to try was the swamp. Frog agreed to help Historia Naturalis
and soaked up a tremendous quantity of water in his porous Ranarum
Nostratium (1758),
coat; Crane carried him over the mountain where he squeezed it reminiscent of the
all back out and extinguished the fire. Frog and Crane made the frog barometers
popular in late
woman promise to take no more water from their swamp, for it 18th-century
was now nearly dry. The mist that rose up from the water on the Europe.

147
Many Native
American groups
revere frogs for
their connection
with life-giving
water. This
woollen shaman’s
tunic, decorated
with frogs, is from
the Gitskan people
(one of whose four
clans is Frog).

A golden frog
pendant of
unknown
symbolism,
from the Coclé
culture (modern-
day Panama),
600–800 ce.
mountain became clouds full of rain, and the tale thus teaches
that the frog brings the rain. When you hear the frog calling, the
story explains, you should try and catch him and tie him up in the
cornfield, so that he will bring rain to the crops.7 In a similar
vein, the Zuni tribe of New Mexico traditionally create fertility
fetishes in the form of frogs and bury them near water sources,
in order to ensure their continued production.
Modern Europeans, no less than Egyptians and Native Ameri-
cans, have turned to the frog in order to understand fertility and
generation. There is a strong and pervasive cultural memory
in Britain and the us that sex education in the mid-twentieth
century was somehow conveyed – albeit euphemistically and
unsuccessfully – through the use of frogs. Carol Ann Duffy’s poem
‘In Mrs Tilscher’s Class’ (1990) captures the child’s unspeakable
puzzlement at nature and its sexual mysteries, all embodied
within the classroom frog:

Over the Easter term the inky tadpoles changed

from commas into exclamation marks . . .
. . . A rough boy
told you how you were born. You kicked him, but stared
at your parents, appalled, when you got back home.
. . . You asked her
how you were born and Mrs Tilscher smiled,
then turned away . . .

‘Our teacher was . . . clearly embarrassed’, recalls one former

pupil in a bbc web article. ‘We . . . never progressed, in five years,
beyond the lifecycle of a frog.’ When one of her class fell preg-
nant, the teacher made a remark that revealed her somewhat
unrealistic pedagogical expectations: ‘I can’t understand it, she
attended my classes.’ Obviously children were supposed to make

149
the connections for themselves, but as the blogged recollections
of a former Catholic schoolgirl make clear, these were not always
biologically sound:

It started in grade four with the ancient movies that

showed a female frog leaving her eggs in the stream to be
fertilized by the male frog at a later time. So needless to say
I thought that babies . . . happened in your sleep (or a
stream) until I was at least in grade seven.

The visual similarity between tadpoles and sperm viewed

under the microscope (remarked upon at least as far back as 1678)
has perhaps served to strengthen the uneasy conviction that the
frogs are somehow supposed to be our teachers when it comes
to the facts of life. However, the specific source of the remem-
bered connection between frogs and sexual education is probably
two-fold. Dissecting frogs has long been a ubiquitous feature of
classroom biology, and the sexual organs noted as a part of this
process; in their feverish state of not-knowing, children have
tried to connect up what they see in the frog with what they want
to find out about humans. Secondly, as Duffy’s poem describes,
tadpoles have commonly been kept in the class and observed
as they metamorphose into frogs through the summer term.
In this guise, they illustrate a ‘cycle of life’ which, as historian
Julian Carter argues, was central to sex education during the
inter-war years.8
In this pedagogical model, which still resonates today,
teachers treated the bigger pattern of nature’s replenishment
and avoided a focus upon the sexual act itself. The frog was a
perfect choice for this teaching method. With its metamorphic
life cycle it made a beautiful point about life’s grand story, and
with its external method of fertilization it avoided certain

150
embarrassing features of human reproduction. Frogs took their
place in a hierarchy of animal forms – the ‘ladder of back-boned
animals’ – that was used to teach about sex. The reproductive
differences between these levels of animal were not defined, as
one might expect, in the mechanics of congress of egg and
sperm, but rather in the quality of parenting. ‘The Mother Fish
Neglects Her Babies’, explained one American pamphlet, plac-
ing her firmly at the bottom of the hierarchy that peaked with
caring human mothers. Batrachians were little better, at least
according to the narrator of a British sex education film of 1932.
‘Reptiles [sic] are . . . careless parents’, he intoned. ‘The frog
abandons her eggs . . . and the tadpoles developing within never
know the meaning of a mother’s care.’9 However, according to
most, frogs took their place in the explanatory framework
somewhere between feckless fish and loving mammals. Unlike
the former, they at least bequeathed their eggs some protective
jelly before swimming off.
The British sex educator Cyril Bibby (1914–1987) was not one
to wrap up the facts of life in euphemisms of parenthood, but
even he made special pedagogical use of the frog in the animal
stories that he wrote for the New Pioneer magazine in the late
1930s. With the possible exception of the earthworms, Freda and
Freddie are the most explicit of Bibby’s alliterative protagonists:

The male frog lies on top of the female, grasping her under
the armpits with his thumbs. Mrs Frog gives out into the
water her eggs and Mr Frog emits the male cells, or sperm.
Each egg joins with a sperm, and that is the start of a new
frog . . . Freddie had little pads on his thumbs which Freda
did not have, and he used these in holding tightly to her
while they were mating . . .10

151
 Freddie and Freda are close enough to humans for the sexual
comparison to be made, but far enough away to remain decent.
It is not just sex educators, but also sex researchers who have
made extensive use of frogs. Besides being extremely numerous
in nature, frogs’ eggs are noteworthy for their large size and
easy visibility, and this was a convenient feature for the early
modern philosophers who attempted to unravel the mysteries
of reproduction. There were many controversies in the modern
period concerning the role of eggs and male fluid in the gener-
ation of life, and the priest and natural philosopher Lazzaro
Spallanzani (1729–1799) was prominent among those using the
frog to explore them.11 Spallanzani was disinclined to trust the
work of one Professor Menzius, who supposed that the male
frog’s seed was emitted from his toe, and somehow penetrated
through the female’s thorax to effect fertilization. Another piece
of doubtful research came from a Monsieur Gautier, who claimed

Lazzaro
Spallanzani’s
research on
the sex life of
frogs (among
other topics) is
celebrated in this
statue of 1888 in
his birthplace,
Scandiano.

152
that the frog foetus actually came from the male: little mini-
tadpoles that ‘fed’ upon the female egg and grew. Spallanzani
was dismissive of this ‘pretended discovery’ for various reasons,
not least that he found Gautier’s ‘worms’ in the female frog’s
bladder as well as the male.12 However, Spallanzani was per-
suaded by the experiments of Jan Swammerdam that frogs’ eggs
were fertilized by the male outside the female’s body. Carefully
observing and dissecting fertilized and unfertilized eggs, Spal-
lanzani found that the two were absolutely identical. In 1767,
Spallanzani concluded that the tadpole existed in the egg before
fertilization, wanting only the ‘fecundating liquid of the male to
unfold [itself]’.13 On the basis of this discovery, Spallanzani pro-
ceeded to promote frogs up the great chain of nature, considering
them to be viviparous, like mammals, rather than oviparous like
the lower creatures.
The discovery of the tadpole-within-the-egg provoked a
cascade of further questions in Spallanzani’s mind, and over the
following decade he was able to devise experiments to go where
his thoughts led him. Borrowing the idea of one of his corres-
pondents, Spallanzani constructed some little trousers of waxed
taffeta which he proceeded to fit on his male frogs; thus clad in
a sort of whole-body prophylactic, the frogs did not succeed in
fathering tadpoles by the females that they mounted. Unlike his
correspondent, Spallanzani succeeded in finding droplets of
the males’ wasted liquor inside their waterproof breeches. He
scraped it up and dabbed it on ripe eggs harvested from inside
a female, and thus, in 1777, the first artificial fertilization of any
animal was achieved. Vanquishing Gautier’s crazy ideas once
and for all (or so he hoped), Spallanzani found this worked
both with and without the worms in the seminal fluid. He also
thought that electricity might work instead of seminal fluid in
stimulating the development of the egg, but found that it did

153
Scenes such as
this (illustrated
here in Roesel von
Rosenhof’s Historia
Naturalis Ranarum
Nostratium) caused
Spallanzani to
conclude that
‘nothing can be
more salacious
than the male
toad’.

not. Nor did other bodily fluids from the frog, or seminal fluids
from other species.
Spallanzani also found himself drawing conclusions about
the sexual behaviour of batrachians. Noting that several natural-
ists believed frogs and toads embraced one another for a biblical
period of forty days, he found that the period was actually
shorter than this, although still measured in days rather than
hours. Observing the toad’s fierce embrace, lasting beyond the

154
female’s emission of eggs and resisting removal at all costs, Spal-
lanzani judged that ‘nothing can be more salacious than the male
[toad]’.14 (However, he qualified this by advising that the toad
might be put off his ‘nuptials’ if too obviously observed by the
experimenter.) ‘Brutes are not exempt from jealousy’, Spallan-
zani ruminated; perhaps the toad was worried that if he quit too
soon, another might get in on the act.
Spallanzani’s sexual toad, fecund and grasping, is remarkably
similar to the Chinese frog Ch’an Chu, a prosperity symbol and
traditional bringer of luck. Perhaps the similarity is not so surpris-
ing, for wealth and procreation are universally and tightly linked
in human culture. The happy ending always brings fortune,
marriage and the birth of children, all of which can be symbol-
ized by the frog. Figurines of Ch’an Chu depict him squatting
possessively atop his pile of old coins. More money is threaded
on the strings emerging from his mouth and draped over his
three-legged body, while his malevolent red eyes dare you to steal
the coin that he holds in his open mouth. In terms of cultural
significance, Ch’an Chu is currently most significant within the
globalized phenomenon of Feng Shui and its internet sales. Sup-
posedly essential in any home or business that wants to thrive,
he should be placed in the living room of a home or the main
room of a business, diagonally opposite the main entrance.
Sometimes the toad (or frog – the term is used interchangeably
for Ch’an Chu) has the Buddha on his back for good measure.
There is some doubt as to whether the powers of Ch’an Chu
lie in attracting new money or merely in protecting what you
already have. Most of the many Feng Shui websites which will
sell you a statuette err on the side of optimism and claim that
it will do both. Feng Shui expert, Lillian Too, suggests that pur-
chasing up to nine frogs from her online store (at $49.99 each,
a total of almost $450) will ‘invite more wealth to the house’.

155
An example of the
‘traditional’ lucky
Feng Shui toad,
Ch’an Chu.

Participants in Feng Shui web forums have unfortunately found

some of Too’s instructions ambiguous, and there is considerable
debate about, for example, the direction in which the frogs
should face for failsafe financial gain. On balance, it would seem
safe to say that the best way to obtain money from lucky frogs
is by selling them.
There are so many frog myths in southern China that it is
extremely difficult to disentangle the origins of this contempo-
rary good luck charm. It seems to be based on at least two stories,
but has deeper roots still in a multitude of southern Chinese
batrachian myths and customs. Frogs have traditionally been
eaten, deified and sacrificed in southern China. In Guangdong
and Jiangxi, wa frog cults were widespread and included versions
of the frog king tale. One magical frog helped to repair a burst
dyke in 1115 ce and was rewarded with a temple; frogs were also
sacrificed or put into earth altars as part of a rain ceremony. (The

156
custom persisted in the form of placing three-legged toad figures
in old latrines.)
On the more sinister side, the dark aspect of the human soul
was depicted as a ha-ma frog. A tradition from the Thai border of
southern China also has a negative connotation for batrachians,
ranking the toad as one of five venomous creatures. However, this
property can be turned to positive use by placing the animal (or
a representation of it) in key locations on the fifth day of the
fifth month, thus driving out disease and evil. A frog broth may
also be consumed around this time, and drawing on the ground
with a dried toad’s foot will cause water to start flowing there.15
One obvious source story for the Feng Shui frog concerns an
envious wife who stole the elixir of immortality and fled with it
to the Moon. Here, she was transformed into a toad with three
legs – just like Ch’an Chu – representing three phases of the
moon.16 The three-legged toad in the Moon dates back at least
to the poet Li Bai (701–762 ce), and the outline of a toad can still,
it is claimed, be seen there at night. A tradition of eating frogs in
the fifth month appears to connect this lunar frog with a role in
fertility. Held at Full Moon during the mating period of the
frogs, this festival of frog-eating celebrates the sexual congress of
Heaven and Earth.
A second well-known Chinese frog story concerns the immor-
tal holy man Liu Hai, and in its different versions his toad Ch’an
Chu takes on different qualities. According to some, the toad is
his consort and helps him travel instantaneously to new loca-
tions. However, the toad also has a propensity to sulk and hide
down wells and from time to time Liu Hai must lure him out
with a string of coins. In other versions, the toad in the well is
actually evil, emanating lethal vapours, and is tempted out with
the coins so that he can be killed by Liu Hai. In still another
account, Liu Hai’s toad turns into a girl and marries him.

157
A 20th-century
woodcut of the
immortal Liu Hai
sporting with his
three-legged toad.
 Thus the three legs of the Feng Shui frog connect it with the
Moon, with protection from disease and perhaps with enhanced
fertility. The string of coins connects it with Liu Hai (who, inci-
dentally, is a Taoist figure, making the Buddha on Ch’an Chu’s
back an incongruous addition). Whichever version of the Liu
Hai story we take, the toad’s avarice is not a particularly admir-
able quality – even the means of his undoing – and the use of
the toad for luck is not an easy fit with the logic of Western
iconography. At least one Chinese specialist gives a similarly
negative account of the motif, stating that the toad in the Moon
symbolizes the unattainable while Liu Hai’s toad represents
how money can lure men to their destruction.17 Nor, it would
seem, is the tradition of using Ch’an Chu’s statuette for luck so
ancient as sellers of Feng Shui accoutrements routinely claim.
Three-legged toad figurines pre-date the Liu Hai story by many
hundreds of years, and during the Han dynasty were often carved
in jade,18 but figurines of ‘Liu Hai sporting with the toad’ do not
seem to emerge in significant numbers until the eighteenth
century. Such figures, which were indeed considered auspicious,
became more common in the nineteenth century. However, the
form in which they are now sold is shorn of Liu Hai. This is a very
recent phenomenon; I can find no evidence of the present-day
format, showing Ch’an Chu alone, before the late twentieth cen-
tury. It would appear to be a significant change from the logic
of luck which required both Liu Hai and his toad to be present
in earlier carvings. It is almost as though the tale of Liu Hai is too
distracting or irrelevant for contemporary Feng Shui purchasers,
many of whom are not of Chinese background. And yet, it is not
quite that simple, for the tale of Liu Hai (or a short, bastardized
version thereof ) is generally presented on Feng Shui websites to
add authenticity to the object. Despite its problematic icon-
ology and doubtful authenticity, the lucky money frog has now

159
taken its place in the marketplace of consumable ‘spirituality’, a
result of the general fondness of Westerners for frogs and the
allure of Chinese exoticism, combined with the frog’s ancient
association with fruitfulness of all kinds.

160
7 Jumped Up

Be kind and tender to the Frog,

And do not call him names,
As ‘Slimy skin,’ or ‘Polly-wog,’
Or likewise ‘Ugly James,’
Or ‘Gape-a-grin,’ or ‘Toad-gone-wrong,’
Or ‘Billy Bandy-knees’:
The Frog is justly sensitive
To epithets like these.

Hilaire Belloc’s 1896 poem ‘The Frog’ works by cutting slyly

across its own admonition. Despite advising against the calling
of names, it then goes on to give an unnecessarily long list of
suggestions. It seems as though the poet has ceased giving exam-
ples after ‘Ugly James’, but he unexpectedly continues, with just
a bit too much enthusiasm and inventiveness. At the poem’s
end, Belloc concedes with pretended reluctance that those
who stick up for frogs are ‘extremely rare’. The verse is, contrary
to its stated intent, an invitation to laugh at batrachians. Even
herpetologists, those professional students of the frog, have been
forced to confront the unscientific but seemingly unavoidable
question: why are frogs so funny? In a serious research paper,
three of them conclude that frogs’ short, squat bodies, their ‘sur-
prising saltatory gait’ and their harmlessness are the source of

161
Seisei Kyôsai, Circle
of Frogs Dressed
in Lotus Leaves,
c. 1879, ink and
colour on paper.
The humorousness
inherent in frogs
is brought out
in this Japanese
illustration,
showing them
capering about.

humour. One might plausibly add bug-eyes and explosive croaks

to this list, and even theorize that recent frog humour is in part
a guilty normalization of anurans’ role in scientific experiment.
However, one can only agree with the authors’ conclusion that
collectively, these characteristics render them ‘both bizarre and
benign’: beasts of ‘intrinsic whimsy’.1 Some species even have
an upwardly curving mouth that makes them look as though
they are sharing the joke.
The comedy value of frogs dates back to at least the sixth
century bce, when Aesop told unflattering fables at their expense.
The proud frog, attempting to imitate the mighty ox, inflates
itself until it bursts. The frogs that persistently importune Zeus
for a king are eventually rewarded with a crane that eats them
up. In another piece of frog humour from the ancient world, the
Batrachomyomachia savours the ridiculousness of the frogs.
This, the first known comic epic (c. fifth century bce), tells a tale
of deadly revenge and war . . . of frogs and mice. The frog king
starts the whole thing off by giving a mouse a lift across the river

162
 This illustration
of ‘The Battle of
Frogs and Mice’
was made for John
Ogilby’s Fables of
Aesop (1665). The
tale is not usually
attributed to
Aesop.

and carelessly allowing him to drown en route. When the mice

rise up in armed response, the gods refuse to intervene. Instead,
they sit back and enjoy the altercation, finding it doubly funny
because the two equally insignificant parties take it so seriously.
The humour of the Batrachomyomachia is in some ways not so
very far from Mark Twain’s story, ‘The Celebrated Jumping Frog
of Calaveras County’ (1865). A large part of Twain’s comedy lies
in the mise en scène: the narrator has been set up by his friend to
suffer the interminable anecdotes of Simon Wheeler, propping
up the bar in the dilapidated Angels hotel. Wheeler himself
is suspiciously batrachian; fat and bald, he tells his stories with
a comic earnestness: ‘Far from his imagining that there was

163
anything ridiculous or funny about his story, he regarded it as a
really important matter.’ Wheeler’s tale (when he gets round to
it) concerns an inveterate gambler, Smiley, who catches and
trains a frog to be a magnificent jumper. ‘Smiley said all a frog
wanted was education, and he could do ’most anything’, reports
Wheeler. When a stranger comes into town, Smiley bets him
that his frog can out-jump any other. The stranger agrees to the
bet, but while Smiley is off catching a frog for his adversary, he
sneakily fills Smiley’s prize jumper with leaden quail shot.
When the race begins, Smiley’s frog remains ‘planted as solid as
a church’, and after the stranger has hot-footed it with his $40,
Smiley continues to scratch his head. ‘I wonder if there ain’t
something the matter with him’, he ponders of his frog. ‘He
’pears to look mighty baggy, somehow.’
Some time later, Twain came upon a translation of the tale
into French, and took it upon himself to translate it back into
English, aiming to show the ‘odious . . . bad grammar’ that the
French version had imposed upon his story. The translation is
nowhere near as bad as Twain makes out in his humorous
attempt to ‘claw’ it back to ‘civilized language’. For example, he
is either ignorant, or pretends to be ignorant, of such basic
French constructions as the ‘ne . . . pas’ double form of negative.
Twain doggedly mistranslates each instance to highlight what
he regards as nothing more than grammatical recalcitrance on
the part of the French: ‘Eh bien! I no saw not that that frog had
nothing of better than each frog.’ American readers of the mid-
to late nineteenth century were used to the connection of frogs In this Japanese
and the French, and so this linguistic parody – the frog’s own frog battle, the
anurans’ opponents
version – adds another dimension to Twain’s original story. are snakes.
However, a little prying into us journals of the nineteenth Woodblock print
by Utagawa
century suggests that when Americans heard of the French as Yoshitsuya
frogs or frog-eaters, it was usually in the context of the English. (1822–66).

165
A competitor
urges his frog
on at the annual
re-creation of
Twain’s contest in
Calaveras County.

Either it was an English author making the connection (notably

Charles Dickens, who was widely read in the us), or else the
slur was made by an English character. With this in mind, a
seemingly trivial part of Twain’s tale attains fresh significance.
Part of Wheeler’s lead-up to the frog story concerns Smiley’s
dog, also the object of his owner’s bets. This dog is a bulldog,
and its modus operandi in fights is to take firm hold of its oppo-
nent’s hind leg until the latter gives up. The bulldog is eventually
beaten by a trickster who enters a two-legged canine into the
ring (the back legs have been lost in an accident with a circular
saw). Not knowing where to bite, Smiley’s dog gives up, loses the
fight and dies of shame. The dog is a bulldog – the variety most
closely associated with the English. Its inescapably English
nature is highlighted by Twain, who claims that the French trans-
lator, not knowing what it would even be in his own language,
renders it as ‘bouledogue’. Thus in the palimpsest version of the

166
tale, English-French-English, both the English bulldog and the
French frog are bested by American wit.
Twain’s story continues to celebrate American identity in its
eponymous county, and is annually re-enacted (without the
lead shot, which would be poisonous to the frogs). The contest
started in 1928 and, notwithstanding a rift and the commence-
ment of a rival frog-jump in 2007, is still going strong. The record
for the longest jump has been held since 1986 by Lee Giudici,
whose champion frog Rosie the Ribiter managed 6.55 m. There
is a $5,000 prize awaiting the person whose frog can ever beat
this distance.
If one had to choose a single word that encapsulates the
comic appeal of frogs, it would undoubtedly be ‘hapless’. And
the frog that best performs the role is Kenneth Grahame’s Mr
Toad, of the novel, The Wind in the Willows (1908). Although the
word ‘hapless’ only occurs twice in the text, it is forever associ-
ated with the impulsive and vainglorious batrachian, cruising
for one of his numerous falls just like Aesop’s exploding frog.
Fearing for his arrest after stealing a motor car, Toad utters his
memorable lament:

Another popular
recreation of
batrachian
locomotion: The
Country Diversion
of Leap-frog,
etching after
Hayman, c. 1743.

167
‘A luckless bull-frog
lost his voice while
talking in his sleep,
and now he’ll
never fish it out –
his voice it is so
deep.’ A version
of the perenially
unfortunate
batrachian from
Harper’s Round
Table, 1896.

O, what a fool I have been! What did I want to go strutting

about the country for, singing conceited songs . . . instead
of hiding till nightfall and slipping home quietly by back
ways! O hapless Toad! O ill-fated animal!

‘Hapless’ is funny because it implies that some grand,

malevolent scheme of fortune is to blame, whereas in fact Toad’s
(relatively trivial) adversities arise entirely through defects in
his own character: from his frogness, in fact. The critic Seth Lerer
points out that Toad’s words echo Milton’s in Paradise Lost: ‘O
much deceiv’d, much failing, hapless Eve’, again highlighting
the preposterousness of Toad’s self-regard.2
Mr Toad shares his overweening character with the hero of
Beatrix Potter’s The Tale of Mr Jeremy Fisher (1906). At the begin-
ning, we are told that Jeremy can get away with what children
generally cannot; he ‘liked getting his feet wet’ but ‘nobody
ever scolded him’. But, as fabular frogs are inclined to do, Mr
Fisher over-reaches himself – in his case by going out to catch
fish. Potter, as a keen naturalist, would have known that fish
form no part of the common frog’s diet; Rana temporaria sticks

168
to invertebrates such as insects, slugs and worms. Mr Fisher
adds conspicuous consumption to his sins; he plans to invite his
two important-sounding acquaintances to dinner, but only if he
catches at least five of the fish. This is the opposite of feeding
the 5,000; here, only an excess of catering will do.
As a child, I remember being vaguely but unpleasantly dis-
concerted by Mr Fisher’s dinky, white-clad feet. Now I realize that
they lack webs; however, their insertion into galoshes forms an
important part of the tale, for it is only due to proper rain-wear
that Fisher escapes the trout that swallows him. Disliking the
taste of his macintosh, the trout spits him back out. The fish
does, however, swallow Fisher’s galoshes, and for a few pages
the frog displays naturalistic feet (although interestingly these
are often modestly concealed in Potter’s illustrations by water
or foliage). When Fisher’s friends arrive at the end of the book,
the feet are once again tucked away inside spats and improb-
ably tiny shoes. Potter finishes the tale with Fisher sharing a
meal of insects with the tortoise and the newt: ‘roasted grass-
hopper with lady-bird sauce; which frogs consider a beautiful
treat; but I think it must have been nasty!’ Jeremy Fisher has
(just about) learned his place; he is permitted to have fun and
be free so long as he obeys certain rules; he must wear galoshes
and raincoat and must not aspire to inappropriate dining. His
pictures summon up the image of a butler, and he may be
unconsciously based in part upon Lewis Carroll’s frog footman.
His frogness is strictly disciplined within a particular frame-
work of class and childhood.
Arnold Lobel’s delightful Frog and Toad books span the
1970s, but with their poignant humour and characterization
they recollect Kenneth Grahame’s gentler touch. ‘One day in
summer, Frog was not feeling well’, begins one:

169
Queen Alice
conversing with an
old and comic
frog. Wood
engraving after
Tenniel’s drawings
for Through the
Looking Glass
(1872).

‘Toad said, ‘Frog, you are looking quite green.’

‘But I always look green,’ said Frog. ‘I am a frog.’3
The stories are whimsical but never cloying, and have
something of a Zen-like quality to them. Toad is a slightly more
complex character, more given to grumpiness and melancholy,
but also more resourceful than Frog. Frog’s blitheness, however,
often carries the day more successfully than Toad’s deliberation.

170
‘A Swim’ demonstrates Lobel’s straight-edged technique. Frog
and Toad go for a swim, and Toad insists on wearing his bathing
suit. He makes Frog promise not to look at him until he is in the
water: ‘because I look funny in my bathing suit’, he explains. By
the time they are ready to get back out, a curious crowd has
gathered to see Toad, contrarily attracted by Frog’s requests that
they should go away and not look at his friend:

‘If Toad looks funny in his bathing suit,’ said the snake,
‘then I, for one, want to see him.’
‘Me too,’ said a field mouse.
‘I have not seen anything funny for a long time.’4

Arnold Lobel’s
delightful (but
inevitably comic)
Frog and Toad.

171
 Lobel avoids the clichéd children’s book ending, whereby
Toad’s bathing suit somehow becomes invaluable, or the crea-
tures learn that laughter at others’ expense is wrong. Instead, it
all turns out just as Toad predicted when he eventually starts to
feel the cold and is obliged to emerge from the water:

The tortoise laughed.

The lizards laughed.
The snake laughed.
The field mouse laughed,
and Frog laughed.
‘What are you laughing at, Frog?’ said Toad.
‘I am laughing at you, Toad,’ said Frog, ‘Because you do
look funny in your bathing suit.’
‘Of course I do,’ said Toad. Then he picked up his clothes
and went home.5

Somehow, Toad’s dignified response maintains the innocence

of the laughter; the resolution denies a too-neat justice, but also
keeps the world clean of cruelty. Frog and Toad are funny, but
never diminished by Lobel.
Meanwhile, frogs have also been much in evidence in film,
tv and Internet media. A frog named Flip was the hero of the
first ever colour sound cartoon, Fiddlesticks (1930). The animators
never quite seemed sure how to render a frog in cartoon form,
having a particular problem with whether or not shoulders (and
teeth) should be present. His character in earlier cartoons is
similarly vague, and in Fiddlesticks his actions were confined to
generalized capering. In successive films, Flip became less and
less frog-like, and his character became a little more defined
as a down-and-out, doing such unheroic things as spying on a
woman in the shower and taking opium. In 1933, he disappeared

172
 The apparently
‘juvenile’ element
of the frog’s
brand-image
did not dissuade
Budweiser from
using it to advertise
their beer in a
popular campaign
of the 1990s.

altogether, but some twenty years later another singing and

dancing frog took his place in the cartoon world. Michigan J.
Frog debuted in 1955, in the Looney Tunes cartoon One Froggy
Evening (described by Steven Spielberg as ‘the Citizen Kane of
animated film’). Michigan switches the classic frog joke into the
human realm; in this tale it is a man that is haplessly ambitious.
Discovering the remarkable performing frog, he plans to make
his fortune out of showing him. But the frog remains resolutely
silent when anyone else is present, and so his master’s greed
and hubris are punished. Despite remaining off-screen for 40
years, Michigan was revived as the mascot of the wb Television
Network in 1995, only to be retired a decade later for giving an
overly juvenile edge to the brand.
The year 1955 was also the year that Jim Henson’s Kermit the
Frog made his first appearance. Henson stitched the prototype

173
Arguably the most
famous frog ever:
Jim Henson’s
Kermit (b. 1955).

from his mother’s overcoat and two ping-pong balls; at first he

was indeterminately herpetological, and only over a decade or
more did he gradually metamorphose into something unam-
biguously froggish. In Sesame Street, Kermit shared something
of Mr Toad’s overweening ambition, a frustrated know-it-all
constantly bothered by the antics all about him. In The Muppet
Show, he kept order and steered skilfully clear of Miss Piggy’s
attentions. Of all Henson’s puppets, Kermit appears to have been
the creature closest to his maker’s heart. Henson voiced Kermit,
compared his lot in life to his own, and used him as the logo of
his company. He retained control of the character after rights to
the others were sold on. But despite Kermit’s many positive
attributes, only the cruellest parent would today choose the
once-common name – now forever linked with the frog – for
their son. Like his namesake, such a boy would most certainly
find that ‘it’s not easy being green’.
The protagonist of Frogger was a hapless frog par excellence.
This 1981 arcade game, which quickly became available for the

174
earliest home computers, featured a frog that players had to navi-
gate to safety across a multi-lane highway and a dangerous river.
Alas, real frogs and toads face the challenge of roads that lie
between their elevated feeding zones and lower-lying breed-
ing grounds. Occasionally – and notably in the uk – these have
toad-crossings at particularly vulnerable points; elsewhere, toads
have to be content with road signs warning of their presence.
These unintentionally humorous (and, one suspects, ineffective)
signs perfectly echo the poignant comic appeal of Frogger.
It is a horrible irony that frogs are, in terms of ecological
threat, amongst the most hapless of all animals today. Over the
past 30 years, habitat destruction, pollution, collection, uv-b
radiation, climate change and pandemics have all contributed
to massive declines and extinctions of numerous species: a ‘per-
fect storm’ for the frog, according to many scientists. In 2004, it
was estimated that more than 120 species had become extinct
since the 1980s; today, that figure is likely to be even higher.
Malcolm McCallum of Texas a&m University puts the current

A close view of
a Wyoming toad
(Bufo baxteri or
Anaxyrus baxteri),
victim of the
‘perfect storm’
that is leading
to extinction for
many anurans.
The species
suffered a sharp
decline during
the 1970s and has
been extinct in
the wild since
1991.

175
This Woodland
Still-life with a Frog
is an unusually
early representa-
tion of a frog
within its habitat
– although its
elements may be
symbolic rather
than naturalistic.
(Johann-Adalbert
Angermeyer, 1736,
oil on copper.)

rate of frog extinction at an unimaginable 25,000–45,000 times

the natural background level; others agree that over one-third
of species are currently threatened. This figure was produced
by over 500 scientists working together on the Global Amph-
ibian Assessment in 2004, updated in 2006 and 2008.
Since 1989, frogs have commonly been described as ‘bio-
logical indicators’ of ecological health, a somewhat vague term

176
 The Solomon
Island leaf frog
(Ceratobatrachus
guentheri) seems
to be a survivor
of environmental
degradation,
flourishing in the
ravaged forests of
Papua New Guinea
and its recent
urban gardens.

that is generally taken to mean that their numbers are sensitive

to habitat contamination in general, and also that they reflect
population levels in other types of animals (so that if frogs
decline, we can assume that mammals and trees have declined
to a comparable degree).6 Two reasons are commonly given for
the frog’s role as environmental bellwether: that it inhabits both
water and land, so is exposed to a particularly wide range of
environmental stresses; and that its absorbent skin makes it
sensitive to any pollutants in its habitat. Pesticides, for exam-
ple, have been implicated in frogs’ compromised immune
systems and reduced ability to breed – even sterility.7 The
British-based herpetologists Trevor Beebee and Richard Griffiths
are cautious about the generalized notion of frogs as biological
indicators, however, pointing out that some are actually ‘tough
as old boots’ and able to take advantage of changed habitats.
The collection of frogs – by scientists and traders – has had
a serious impact on frog numbers. Unusual or attractive species
such as members of the Madagascan genus Mantella are
particularly at risk, being taken in unsustainable numbers from

177
The Panamanian the wild. These days, the understandable desire to buy and keep
golden frog
(Atelopus zeteki)
gem-like, colourful frogs can be harmlessly satisfied by the
is a critically Apple app, Pocket Frogs (2010). This game successfully taps into
endangered toad, many features of the urge to collect: aesthetic appeal, rarity,
now probably
extinct in the ownership, care, breeding – and perhaps showing off – and has
wild. Only after it proved a huge success. Alas, no such virtual alternative exists
had been allowed
to die out did the for the eating of frogs. The consumption of frogs poses at least
Panamanian a two-fold threat to their continued existence; firstly by their
government
decide to enshrine
straightforward destruction for eating, and secondly by the
it as a national introduction of edible or farmed species – notably bullfrogs –
symbol,
commemorated
which out-compete or even predate upon native anurans.
annually. Quite besides all these other problems – and by far the most
immediate threat to frogs at the present time – is a disease known
as chytridiomycosis, caused by the fungus Batrachochytrium
dendrobatidis (bd). Spores of bd invade the skin of anurans,

178
breaking down their cells to steal nutrients, and eventually pro-
ducing spores which can either re-infect the host or spread to
further victims. The skin of infected frogs is visibly damaged,
and can no longer properly absorb water or oxygen. The animals
become lethargic and unresponsive to environmental stress. This frog pendant,
3rd–8th century ce
Infected frogs are most easily spotted due to their odd sitting from modern-day
posture, with legs trailing helplessly out behind them rather Peru, perhaps
than tucked under the body in a neat squat. reflects modern
the urge to collect
Unexplained groups of dead and dying frogs were first and own perfect
spotted in Queensland in 1993; five years later, bd was identified little frogs –
whether living or
as the cause of this mysterious disease. Since then, chytridio- as App ‘pocket
mycosis has devastated frog populations around the globe. frogs’.
Around 30 per cent of species have been affected to date; once
bd is present in a population, 80 per cent of individuals can be
expected to be dead within a year. This is an average figure;
sometimes the annihilation is total. Herpetologists are some-
times in the distressing and tragic position of describing new
species from the dead specimens that they find; discovery and
extinction bookend their descriptions. The Australian researcher

The Caribbean
giant ditch frog or
mountain chicken
frog (Leptodactylus
fallax) – one of the
world’s largest
species – has been
brought to the
point of extinction
partly by human
consumption.

179
 Lee Berger, who along with colleagues discovered the role of bd
in chytridiomycosis, states that the disease is producing ‘the
most spectacular loss of vertebrate biodiversity due to disease in
recorded history’. Other herpetologists simply report weeping
as they return to their study pools and find them empty.
Is bd a new pathogen, or is it an old one that has recently
increased its virulence? Or is the problem that it has spread
beyond an original host population that had good immunity,
and into an unprotected wider world? Xenopus laevis, the African
clawed frog, is one species that can be infected by bd but has
low morbidity. Preserved specimens from the early twentieth
century, held in European laboratories, have been tested for the
disease and one from 1938 was found to be infected. This has
prompted suspicion that the global trade in X. laevis for preg-
nancy testing helped to spread the fungus far and wide. There
is a similar concern over the American bullfrog, which also has
high resistance to bd and also has been introduced around the
globe – for eating, in this case. The spores of bd can travel only
very short distances – 1–2 cm – and so it seems likely that the

A frog
suffering from
chytridiomycosis.

180
 A Chinese
pharmacist displays
her collection of
dried frogs and
lizards. Trade in
biological materials
such as these
poses a direct
threat to some
species,
and moreover
increases the
global transmission
of infective agents
such as Bd.

unnatural proximity of frogs in human keeping has had a sig-

nificant role to play in the spread of the disease. Other possible
human factors to blame for the pandemic include the use of
pesticides that further reduce the resistance of anurans to bd, and
changed climates that favour the fungus more than previously.
The discipline of herpetology is today more focused on con-
servation than perhaps any other animal specialism. The First
World Congress of Herpetology (1989) in Canterbury, uk, was a
turning point; scientists compared notes and realized that all
of them were seeing major declines in the species they were

181
 studying. From then on, anxiety about survival became key to
the discipline. Today, herpetologists are understandably concen-
trating most of their resources on combating chytridiomycosis,
but with little success to date. There has been some suggestion
that coating frogs’ skin with a certain bacterium will afford pro-
tection from the disease; painstaking treatments with anti-fungal
agents have also had some positive results. Neither of these ideas,
however, seems like a realistic treatment for wild populations
in the near future. Meanwhile, a programme of breeding in
captivity has begun, a Noah’s ark approach to the desperately sad
possibility that many wild populations might simply disappear.
A rather sci-fi backstop, the Amphibian Ark Biobanking Advisory
Committee was formed in 2008. This body explores strategies
for cryopreserving cells from frogs and other Amphibia, in the
hopes that they might one day be resurrected and re-released. As
if the effects of chytridiomycosis were not depressing enough,
some herpetologists warn that too much focus upon it will allow
other issues such as deforestation and climate change – over
which we might have more control – to continue apace. Scientists
at the Amphibian Conservation Summit of 2005 worked out a
five-year action plan to address all aspects of conservation,
This beautiful
budgeted at $409 m.
Dendropsophus The best chance of mobilizing help on this kind of financial
ebraccatus
illustrates the
scale requires tapping in to some kind of popular concern and
limited value understanding of biodiversity issues. Rainforests seem as though
of artificial they might be a good place to start. Rainforests have long been
conservation
without agree- a focus of concern for biodiversity conservation, and in more
ment on ecosys- recent years ecologists have also emphasized the value of these
tem protection; it
is impossible to ‘lungs of the planet’ in regulating the Earth’s climate. Rainforests
breed in captivity. are actually rather lacking in the large, charismatic species that
Thankfully the
species is not yet
are typically used to excite popular interest in biodiversity issues,
endangered. but there are plenty of beautiful and unusual frogs to take their

182
place. Indeed, these have now come to stand for the plants and
animals of the world’s threatened tropical forests. The Rainforest
Alliance (f. 1986) uses an image of a frog on its accreditation mark
for responsibly produced goods, such as coffee. ‘The little green
frog is your assurance that goods and services are produced in a
socially, economically and environmentally sustainable way’,
the Alliance explains on its website. At the time of writing (early
2011), it states that Mars Inc. has undertaken to earn the frog’s
seal of approval for their Galaxy chocolate bars in the short
term, and for all their chocolate products by 2020.
This is a start, but it will take considerably more than bars
of chocolate to protect the frog in its latest and most dangerous
metamorphosis – into a creature of the warming world that it
shares with humans. Each species that dies out is a grievous loss,
for what Swammerdam wrote 350 years ago remains true to this
day: ‘There is a much greater number of miracles, and natural
secrets in the Frog, than any one hath ever before thought of
or discovered.’8

184
Timeline of the Frog
200 million years ago c. 3000 bce 405 bce c. 1st or 2nd century ce

Earliest confirmed The Egyptian god- Aristophanes The Mandukya

modern frog lives in dess Heket is com- writes The Frogs Upanishad praises the
Argentina memorated in frog spiritual lesson of the
figurines frog

Late 1770s 1812 1865 1869

Galvani begins his The Brothers Grimm ‘The Celebrated Live frogs are
experiments on publish ‘The Frog Jumping Frog of placed in cold
frogs’ legs King’ for the first Calaveras County’ is water and boiled
time; it appears in published by Mark slowly by Friedrich
English in 1823 Twain Goltz

1952 1960s 1961 1986

Robert Briggs and Thomas King Bioprospectors Soviet Union Rosie the Ribiter
create a tadpole with the nucleus begin to hunt out launches the first sets the world
from one egg and the outer mat- valuable pharma- frogs into space record for length of
ter from another ceuticals in frogs frog-jump at 6.55 m
c. 750 1115 1590 1777

Poet Li Bai describes A magic frog mends Forty witches Lazzaro Spallanzani
the three-legged toad a breached dyke in attempt to kill the achieves the first-ever
in the Moon southern China and English King with a artificial fertilized
is rewarded with a poison concocted organism using frogs
temple dedicated to it from toads

1908 1930 1935 1950s

Auguste Escoffier Xenopus laevis is Bufo marinus is National People’s
launches the frog as introduced to introduced to Congress of China
haute cuisine for the Europe as breeding Australia encourages women
English by sneaking colonies for research; to swallow tadpoles
it into the Prince of Flip the Frog stars in as contraception
Wales’s banquet the first-ever colour
sound cartoon

1990s 2004 2009

Toad smoking becomes Global Amphibian Disney makes ‘The Frog

fashionable; Queensland Assessment estimates that King’ into a film – The
authorities ban possession 120 of c. 6,000 frog species Princess and the Frog
of toad slime have become extinct since
the 1980s; 32 per cent of
frog species are in danger
of extinction
 References

introduction

1 Beverley F. Bell, ‘When Is an Animal, Not an Animal?’, Journal of

Biological Education, xv (1981), pp. 213–18.
2 C. H. Brown, ‘Folk Zoological Life-Forms: Their Universality and
Growth’, American Anthropologist, lxxxi (1979), pp. 791–817.
3 Ralph N. H. Bulmer and M. J. Tyler, ‘Karam Classification of
Frogs’, Journal of the Polynesian Society, lxxvii (1968), pp. 333–85;
Peter D. Dwyer and David C. Hyndman, ‘“Frog” and “Lizard”:
Additional Life-Forms from Papua New Guinea’, American
Anthropologist, lxxxv (1983), pp. 890–96.
4 Edward Topsell, The History of Serpents; Or, the Second Book of
Living Creatures, appended to The History of Four-Footed Beasts
(London, 1658), p. 597.
5 Topsell, History of Serpents, p. 726. The early Christian text
Physiologus (c. 2nd century ce) appears to be the sole exception to
this moral ordering of frogs and toads. Its author compares the
‘frog of the dry place’ with ‘fine abstinent men’, and the water
frogs with those driven by their desires. Michael J. Curley, trans.,
Physiologus: A Medieval Book of Nature Lore (Austin, tx, and
London, 1979), pp. 60–61.
6 Ralph N. H. Bulmer and Michael J. Tyler, ‘Karam Classification of
Frogs’, Journal of the Polynesian Society, lxxvii (1968), pp. 333–85.
7 Kenneth R. Porter gives an overview of his discipline’s history and
key publications in the introduction to Herpetology (Philadelphia,
London and Toronto, 1972).

188
8 Ernest Jones, Essays in Applied Psychoanalysis (London, 1951), vol.
ii, p. 94.

1 just a kiss

1 R. P. Chamakura, ‘Bufotenine – A Hallucinogen in Ancient

Snuff Powders of South America and a Drug of Abuse on the
Streets of New York City’, Forensic Science Review, vi (1994),
pp. 1–18.
2 John [Jan] Swammerdam, The Book of Nature, or, The History of
Insects (London, 1758), part ii, p. 105.
3 See Jennifer Rampling, ‘Establishing the Canon: George Ripley
and His Alchemical Sources’, Ambix, lv (2008), pp. 189–208.
The source on which I have drawn here is Rampling’s unpublished
PhD thesis, University of Cambridge (2009).
4 Personal communication.
5 Carl Jung, Psychology and Alchemy (London, 1968), p. 327.
6 J. F. Campbell, Popular Tales of the West Highlands: Orally Collected
(Edinburgh, 1860), vol. ii, no. 33, pp. 130–32.
7 H. Parker, Village Folk Tales of Ceylon [1910] (Whitefish, mt, 2003),
vol. i, pp. 67–72.
8 William Elliot Griffis, The Unmannerly Tiger, and Other Korean
Tales (New York, 1911), pp. 112–25.
9 Folk Tales from China, 3rd series (Peking, 1958), pp. 74–82.
10 Jack Zipes, The Brothers Grimm: From Enchanted Forests to the
Modern World (New York and London, 1988), p. 15.
11 James M. McGlathery, Grimms’ Fairy Tales: A History of Criticism
on a Popular Classic (Columbia, sc, 1993), p. 63.
12 Ernest Jones, Essays in Applied Psycho-Analysis (London, 1951),
vol. ii, p. 16.
13 McGlathery, Grimms’ Fairy Tales, p. 62.
14 Bruno Bettelheim, The Uses of Enchantment: The Meanings and
Importance of Fairy Tales (London and New York, 1991), p. 283.
15 Marina Warner, From the Beast to the Blonde: On Fairy Tales and
Their Tellers (London, 1995), pp. 288–90.

189
16 Geoff Dench, The Frog, The Prince, and the Problem of Men
(London, 1994), p. 251.
17 Jack Zipes, ‘The Struggle for the Grimms’ Throne: The Legacy of
the Grimms’ Tales in the frg and gdr since 1945’, in The Reception
of Grimms’ Fairy Tales: Responses, Reactions, Revisions, ed. Donald
Haase (Detroit, mi, 1993), pp. 167–206; p. 190.

2 warts and all

1 Translated in Kenneth R. Porter, Herpetology (Philadelphia,

London and Toronto, 1972), p. 2.
2 Lazzaro Spallanzani, Dissertations Relative to the Natural History
of Animals and Vegetables [1780] (London, 1789), vol. ii, p. 50.
3 Michael Maierus, Atalanta Fugiens (Oppenheim, 1617), 5th
discourse. Topsell in his History of Serpents gives the story as
concerning a monk, and originating from Erasmus; the toad
requires biting three times.
4 Edward Newman, The Zoologist: A Popular Miscellany of Natural
History, x (London, 1852), p. 3,658.
5 Deborah Willis, Malevolent Nurture: Witch-hunting and Maternal
Power in Early Modern England (Ithaca, ny, 1995), p. 126. In purely
practical terms, they would not have succeeded.
6 Thomas Wright, Narratives of Sorcery and Magic, from the Most
Authentic Sources (London, 1851), p. 308.
7 Cathy Gere, ‘William Harvey’s Weak Experiment: The
Archaeology of an Anecdote’, History Workshop Journal, li (2001),
pp. 19–36.
8 Edward A. Armstrong, The Folklore of Birds: An Enquiry into the
Origin and Distribution of Some Magico-Religious Traditions, 2nd
edn (New York, 1970), pp. 192–5.
9 Heinrich Kramer and Jacob Sprenger, Malleus Maleficarum:
Or, The Hammer of Witches (www.forgottenbooks.org, 2008),
pp. 198–201.
10 Kathleen Cohen, Metamorphosis of a Death Symbol: The Transi Tomb
in the Late Middle Ages and the Renaissance (Berkeley, ca, 1973).

190
11 Sophie Page, ‘Good Creation and Demonic Illusions: The
Medieval Universe of Creatures’, in A Cultural History of Animals
in the Medieval Age, ed. Brigitte Resl (Oxford and New York,
2007), pp. 27–58.

3 them, us and frogs

1 Richard Wassersug, ‘On the Comparative Palatability of Some

Dry-season Tadpoles from Costa Rica’, American Midland
Naturalist, lxxxvi (1971), pp. 101–9.
2 Wolfram Eberhard, The Local Cultures of South and East China
(Leiden, 1968), pp. 202–3.
3 Anonymous, ‘Frog-Eating’, The Journal of American Folklore, xv
(1902), p. 190. Edgar Thurston and K. Rangachari, Castes and
Tribes of Southern India (Madras, 1909), vol. vii.
4 John Claudius Loudon, An Encyclopedia of Agriculture, vol. ii,
p. 1,057.
5 William B. Jerrold, ed., The Epicure’s Year Book and Table
Companion for 1869 (London, 1869), p. 167 and Jerrold, ed.,
The Epicure’s . . . 1868 (London, 1868), pp. 158 and 195–6.
6 Marie Antonin Carême and Armand Plumerey, L’Art de la cuisine
française au dix-neuvième siècle, vol. v (Paris, 2005), pp. 255–9.
7 Kenneth James, Escoffier: The King of Chefs (London, 2002),
pp. 135–54.
8 Shane Mitchell, ‘Wild and Refined’, Saveur, 128 (2010). At www.
saveur.com.
9 un comtrade, at http://comtrade.un.org.
10 Umberto Eco, On Ugliness, trans. Alistair McEwen (London,
2007), p. 19.
11 Edward Bartlett, Wild Animals in Captivity (London, 1899),
pp. 200–1.
12 The following is drawn from Michael Randall, ‘On the Evolution
of Toads in the French Renaissance’, Renaissance Quarterly, 57
(2004), pp. 126–64.
13 David Bindman, ‘How the French Became Frogs: English

191
 Caricature and Stereotypes of Nations’, in The European Print and
Cultural Transfer in the Eighteenth and Nineteenth Centuries, ed.
Philippe Kaenel and Rolf Reichardt (Zürich, 2007), pp. 423–36.
The whole of the section on Dutch and French caricature is drawn
from Bindman’s research.
14 Philippe Hecquet, Traité des dispenses du carême (Paris, 1709),
pp. 164–5.
15 Bindman, ‘How the French Became Frogs’.
16 Gilbert Abbott A’Beckett and John Leech, The Comic History of
England (London, 1847), p. 116.
17 Tim Low, Feral Future: The Untold Story of Australia’s Exotic
Invaders (Chicago, il, 2002), pp. 46–54.

4 under the knife

1 High Field Magnet Laboratory, Radboud University Nijmegen,

‘The Frog that Learned to Fly’, at www.ru.nl.
2 The Bible of Nature was completed in manuscript form shortly
before Swammerdam’s death but was not published until 1737
in Dutch and 1758 in English, for which its title was translated
as The Book of Nature.
3 John [Jan] Swammerdam, The Book of Nature, or, The History of
Insects (London, 1758), part ii, p. 135.
4 Ibid., part ii, p. 111.
5 The European Magazine, and London Review, lxix (1816), p. 214.
6 Robert Townson, Tracts and Observations in Natural History and
Physiology (London, 1799), p. 115.
7 Laura Otis, Müller’s Lab (Oxford and New York, 2007).
8 William Thompson Sedgwick, ‘On Variations of Reflex
Excitability in the Frog, Induced by Changes of Temperature’,
in Studies from the Biological Laboratory, vol. ii, ed. Newell Martin
(Baltimore, md, 1883), pp. 385–410; 388–9.
9 Sedgwick, ‘On Variations of Reflex Excitability in the Frog’, p. 386.
10 E. W. Scripture, The New Psychology (London, 1897), pp. 300–1.
11 Fast Company, ‘Next Time, What Say We Boil a Consultant’, Fast

192
 Company, 31 October 1995, at www.fastcompany.com.
12 Animal Aid, ‘Endangered Frogs – the Vivisection Connection’
(posted 2008), at www.animalaid.org.uk.

5 evolution on fast-forward

1 Robert Chambers, Vestiges of the Natural History of Creation

(London, 1844), pp. 210–11.
2 Ernst Haeckel, The History of Creation [1868] (New York, 1880),
vol. i, p. 294.
3 Haeckel, History of Creation, vol. i, pp. 310–11.
4 Mikhail Bulgakov, The Fatal Eggs (London, 2005), p. 17.
5 For the story of Russian and Soviet Darwinism, see Mark B. Adams,
‘Sergei Chetverikov, the Kol’tsov Institute, and Evolutionary
Biology’, in The Evolutionary Synthesis: Perspectives on the
Unification of Biology, ed. Ernst Mayr and William B. Provine
(Cambridge, ma, 1988), pp. 242–78.
6 This account of the history of Xenopus laevis is taken from John
B. Gurdon and Nick Hopwood, ‘The Introduction of Xenopus
laevis into Developmental Biology: Of Empire, Pregnancy Testing
and Ribosomal Genes’, International Journal of Developmental
Biology, xliv (2000), pp. 43–50.
7 Bertha Chapman Cady and Vernon Mosher Cady, The Way Life
Begins: An Introduction to Sex Education [1917], at
Generalbooks.net (accessed 2010), p. 15.
8 Stephen Jay Gould, Ontogeny and Phylogeny (Cambridge, ma,
1977), p. 1.
9 Personal communication.

6 of frogs and fruitfulness

1 Edward Newman, The Zoologist: A Popular Miscellany of Natural

History, x (London, 1852), p. 3316.
2 George Orwell, ‘Some Thoughts on the Common Toad’, Tribune
(1946).

193
 3 See Sulochana R. Asirvatham et al., eds, Between Magic and
Religion: Interdisciplinary Studies in Ancient Mediterranean Religion
and Society (Lanham, md, 2001), pp. 181–6.
4 Trevor Beebee and Richard Griffiths, Amphibians and Reptiles:
A Natural History of the British Herpetofauna (London, 2000), p. 21.
5 Leo A. Orleans, ‘Birth Control: Reversal or Postponement?’,
The China Quarterly, iii (1960), pp. 59–70; p. 67.
6 Edward Polehampton, Gallery of Nature and Art (London, 1821),
vol. v, p. 561.
7 Franc Johnson Newcomb, Navaho Folk Tales (Albuquerque, nm,
1990) pp. 151–61.
8 Julian B. Carter, ‘Birds, Bees, and Venereal Disease: Toward an
Intellectual History of Sex Education’, Journal of the History of
Sexuality, x (2001), pp. 213–49.
9 Mary Field, dir., The Mystery of Marriage (uk, 1932).
10 Red Squirrel [Cyril Bibby], ‘The Story of Mr and Mrs Frog’, New
Pioneer (August 1939), pp. 118–19.
11 Lazzaro Spallanzani, Dissertations Relative to the Natural History
of Animals and Vegetables (London, 1789), vol. ii.
12 Ibid., vol. ii, pp. 114–18.
13 Lazzaro Spallanzani, An Essay on Animal Reproductions (London,
1769), p. 43. Spallanzani dates his discovery to 1767 in
Dissertations, vol. ii, p. 144.
14 Spallanzani, Dissertations, vol. ii, pp. 42–3.
15 Wolfram Eberhard, The Local Cultures of South and East China
(Leiden, 1968), pp. 155 and 159.
16 E.T.C. Werner, A Dictionary of Chinese Mythology (New York,
1961); Werner, Myths and Legends of China (New York, 1976),
pp. 125–6 and 128.
17 Charles Alfred Speed Williams, Outlines of Chinese Symbolism and
Art Motives [1941] (New York, 1976), p. 403. However, like other
sources, Williams states that ‘Liu Hai sporting with the toad’ is
a figure of good luck.
18 Patricia Bjaaland Welch, Chinese Art: A Guide to Motifs and Visual
Imagery (North Clarendon, vt, 2008), p. 106.

194
7 jumped up

1 Michael J. Tyler et al., ‘How Frogs and Humans Interact:

Influences beyond Habitat Destruction, Epidemics and Global
Warming’, Applied Herpetology, iv (2007), pp. 1–18.
2 Kenneth Grahame, The Wind in the Willows: An Annotated Edition,
ed. Seth Lerer (Cambridge, ma, 2009), p. 216.
3 Arnold Lobel, Frog and Toad Are Friends [1970] (Surrey, 1978),
p. 16.
4 Ibid., pp. 46–7.
5 Ibid., pp. 51–2.
6 Trevor Beebee and Richard Griffiths, ‘The Amphibian Decline
Crisis: A Watershed for Conservation Biology?’, Biological
Conservation, cxxv (2005), pp. 271–85.
7 University of California, ‘Pesticide Atrazine Can Turn Male Frogs
into Females’, (1 March 2010), at www.universityofcalifornia.edu.
8 John [Jan] Swammerdam, The Book of Nature (London, 1758), part
ii, p. 105.

195
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Kingdom’, Journal of Near Eastern Studies, lviii (1999), pp. 107–16
Beebee, Trevor J. C., and Richard A. Griffiths, ‘The Amphibian
Decline Crisis: A Watershed for Conservation Biology?’, Biological
Conservation, cxxv (2005), pp. 271–85
Beetschen, Jean-Claude, ‘How Did Urodele Embryos Come into
Prominence as a Model System?’, International Journal of
Developmental Biology, xl (1996), pp. 629–36
Boll, Valerie, Autour du Couple Ambigu Crapaud-Grenouille: Recherches
Ethnozoologiques (Paris, 2000)
Bowler, Peter, ‘Fins and Limbs and Fins into Limbs: The Historical
Context, 1840–1940’, in Fins into Limbs: Evolution, Development,
and Transformation, ed. Brian Keith Hall (Chicago, il, 2007),
pp. 7–14
Bulmer, Ralph N. H., and Michael J. Tyler, ‘Karam Classification of
Frogs’, Journal of the Polynesian Society, lxxvii (1968), pp. 333–85
Cobb, Matthew, The Egg and Sperm Race: The Seventeenth-century
Scientists Who Unlocked the Secrets of Sex and Growth (London,
2006)
DeGraaff, Robert M., The Book of the Toad: A Natural and Magical
History of Toad-Human Relations (Rochester, vt, 1991)
Dubois, Alain, ‘The Higher Nomenclature of Recent Amphibians’,
Alytes, xxii (2004), pp. 1–14
Eberhard, Wolfram, The Local Cultures of South and East China
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Erspamer, Vittorio, ‘Bioactive Secretions of the Amphibian
Integument’, in Amphibian Biology, vol. i: The Integument, ed.
H. Heatwole and G. T. Barthalmus (Chipping Norton, nsw, 1994)
Gascon, Claude, et al., eds, Amphibian Conservation Action Plan.
Proceedings of the iucn/ssc Amphibian Conservation Summit 2005
(Gland, Switzerland, 2007)
Gere, Cathy, ‘William Harvey’s Weak Experiment: The Archaeology
of an Anecdote’, History Workshop Journal, li (2001), pp. 19–36
Gilbert, Scott F., ed., A Conceptual History of Modern Embryology
(Baltimore, md, and London, 1991); see especially chapters by
Maienschein and Saha
Gliboff, Sander, ‘“Protoplasm . . . Is Soft Wax in Our Hands”: Paul
Kammerer and the Art of Biological Transformation’, Endeavour,
xxix (2005), pp. 162–7
Gould, Stephen Jay, Ontogeny and Phylogeny (Cambridge, ma, 1977)
Gurdon, John B., and Nick Hopwood, ‘The Introduction of Xenopus
laevis into Developmental Biology: Of Empire, Pregnancy Testing
and Ribosomal Genes’, International Journal of Developmental
Biology, xliv (2000), pp. 43–50
Hamburger, Jeffrey, ‘Bosch’s “Conjuror”: An Attack on Magic and
Sacramental Heresy’, Simiolus: Netherlands Quarterly for the History
of Art, xiv (1984), pp. 5–23
Holmes, Frederic L., ‘The Old Martyr of Science: The Frog in
Experimental Physiology’, Journal of the History of Biology, xxvi
(1993), pp. 311–28
Lewis, Mark, dir., Cane Toads: An Unnatural History (Australia, 1988);
there is also a follow-up film, Cane Toads: The Conquest (2010)
Mattison, Chris, Frogs and Toads (London, 2011)
McDiarmid, Roy W., and Ronald Altig, Tadpoles: The Biology of
Anuran Larvae (Chicago, il, 1999)
Newton, Giles, ‘Why the Frog?’, in The Human Genome (on the
Wellcome Trust website, http://genome.wellcome.ac.uk, 2004)
Nyhart, Lynn K., Biology Takes Form: Animal Morphology and the
German Universities, 1800–1900 (Chicago, il, and London, 1995)
Otis, Laura, Müller’s Lab (Oxford and New York, 2007)

197
Pera, Marcello, The Ambiguous Frog: The Galvani-Volta Controversy on
Animal Electricity, trans. Jonathan Mandelbaum (Princeton, nj,
1992)
Richards, Robert, The Tragic Sense of Life: Ernst Haeckel and the
Struggle over Evolutionary Thought (Chicago, il, and London, 2008)
Rosenhof, August Johann Rösel von, Historia naturalis Ranarum nos-
tratium (Nuremburg, 1758), online at http://num-scd-ulp.u-stras-
bg.fr
Sleigh, Charlotte, ‘Plastic Body, Permanent Body: Czech
Representations of Corporeality in the Early Twentieth Century’,
Studies in History and Philosophy of Biological and Biomedical
Sciences, xl (2009), pp. 241–55
Swammerdam, John [Jan], The Book of Nature, or, The History of Insects
(London, 1758), at http://digital.lib.usu.edu
Tyler, Michael J., et al., ‘How Frogs and Humans Interact: Influences
Beyond Habitat Destruction, Epidemics and Global Warming’,
Applied Herpetology, iv (2007), pp. 1–18
Wells, Kentwood D., The Ecology and Behavior of Amphibians
(Chicago, il, 2007)

198
 Associations and Websites

associations

Scientists who study frogs are members of herpetological associa-

tions, of which there are too many around the world to list. The Society
for the Study of Amphibians and Reptiles (f. 1958; www.ssarherps.
org) is the largest international herpetological society, a not-for-profit
organization established to advance research, conservation and edu-
cation concerning amphibians and reptiles. The Herpetologists’
League (f. 1946; www.herpetologistsleague.org/en) is another major
international organization, and the American Society of Ichthyologists
and Herpetologists (f. c. 1913; www.asih.org) also has international
reach. In Europe, the Societas Europaea Herpetologica (f. 1979; http://
t-ad.net/ishbh) links up herpetologists including, in Great Britain,
the British Herpetological Society (f. 1947; www.thebhs.org) and the
Amphibian and Reptile Conservation trust (f. 2009; www.arc-trust.
org).

Members of all these associations, and more, meet every three to five
years at the World Congress of Herpetology (f. 1982; www.worldcon
gressofherpetology.org). The International Society for the History and
Bibliography of Herpetology (f. 1998; http://t-ad.net/ishbh) keeps track
of the discipline and its publications.

The International Herpetological Society (http://international-herpeto

logical-society.org) is a club for amateur breeders and keepers. Australia
appears to be unique in having (at least) three amateur societies devoted

199
specifically to anurans: the Frog and Tadpole Study Group of NSW, the
Queensland Frog Society, and the Victorian Frog Group.

websites

AmphibiaWeb is an online system that provides access to information

on amphibian declines, conservation, natural history and taxonomy.
It is a very rich and frequently-updated source of knowledge about
anurans, compiled at the University of California, Berkeley.
http://amphibiaweb.org

The Amphibian Specialist Group, supports a global web of partners

aiming to achieve shared, strategic amphibian conservation goals,
including a search for ‘lost’ species.
www.amphibians.org

The amphibians section of the IUCN Red List is dedicated to amphibian

species that are currently highly threatened.
http://www.iucnredlist.org

The Biodiversity Heritage Library has photographic online versions of

many original works mentioned in Frog. These include books by Charles
Darwin, Richard Owen, Wilhelm Roux (in German), Ernst Haeckel
(German and English), Lazzaro Spallanzani (English) and many more.
www.biodiversitylibrary.org

D. L. Ashliman, ‘Frog Kings’, (last updated 2008), has fourteen different

versions of the Frog King tale from around the world, along with further
links.
www.pitt.edu/~dash/frog.html#taylor

The Whole Frog Project is a rich and detailed virtual frog dissection
resource.
http://froggy.lbl.gov

200
The Amphibian Specialist Group, supports a global web of partners
aiming to achieve shared, strategic amphibian conservation goals,
including a search for ‘lost’ species.
www.amphibians.org

The amphibians section of the IUCN Red List is dedicated to amphibian

species that are currently highly threatened.
http://www.iucnredlist.org

The Biodiversity Heritage Library has photographic online versions of

many original works mentioned in Frog. These include books by Charles
Darwin, Richard Owen, Wilhelm Roux (in German), Ernst Haeckel
(German and English), Lazzaro Spallanzani (English) and many more.
www.biodiversitylibrary.org

D. L. Ashliman, ‘Frog Kings’, (last updated 2008), has fourteen different

versions of the Frog King tale from around the world, along with further
links.
www.pitt.edu/~dash/frog.html#taylor

The Whole Frog Project is a rich and detailed virtual frog dissection
resource.
http://froggy.lbl.gov

201
 Acknowledgements

I am grateful to the following people for generously sharing their

expertise and contributing gems of anuran knowledge:
Alixe Bovey, Matthew Cobb, Stefan Goebel, Richard Griffiths, Lesley
Hall, Diane Heath, Mubariz Hussain, Dunstan Lowe, Sophie Page, Julie
Peakman, Neil Pemberton, Christopher Plumb, Jenny Rampling,
Victoria Resnick, John-Paul Riordan, Janine Rogers, Gill Sinclair, Paul
Sleigh, Crosbie Smith, Nick Thurston, Richard Wassersug.
Thank you most especially to Alice White, who did such a wonderful
job researching and obtaining the pictures for Frog.

202
 Photo Acknowledgements

The author and the publishers wish to express their thanks to the below
sources of illustrative material and/or permission to reproduce it.

Andrey: p. 43; The Art Institute of Chicago: p. 146 bottom (Wirt D.

Walker Fund Income 1969.792); Steven Barker: p. 142; Sergio Barbierei:
p. 152; Forrest Brem. From open access creative commons article:
Gewin V (2008) Riders of aModern-Day Ark. PLoS Biol 6(1): e24.doi:
10.1371/journal.pbio.0060024: p. 180; British Cartoon Archive, University
of Kent: p. 89; British Library, London: pp. 62, 158; © The Trustees of
the British Museum: pp. 73, 86, 163, 164, 167; Jeremy Brooks (jeremy
brooks.net): p. 173; Canterbury Cathedral Archives: p. 99; Phil Carter:
p. 166; Cleveland Museum of Art, Ohio: pp. 145 (Andrew R. and Martha
Holden Jennings Fund 1976.5), 179 top right (Gift of W. J. Gordon
1955.375); © Depositphotos: p. 156 (Yuliya Krzhevka); Fine Arts Museums
of San Francisco: p. 35; Christian Fisher: p. 130; Froggydarb: pp. 18, 33;
Getty Images: pp. 22 right, 26, 175; Brian Gratwicke: pp. 177, 178; R. A.
Griffiths: p. 131; Peter Halasz: p. 136; High Field Magnet Laboratory,
Radboud University, Nijmegen, The Netherlands: p. 97; Used with
permission of HarperCollins Publishers: p. 167; Jeremy Hubert: p. 84;
Chris Irie: p. 144; Istockphoto: p. 6 (Paul Tessier); © Estate Martin
Kippenberger, Galerie Gisela Capitain, Cologne: p. 114; Christoph
Leeb: p. 101; Library of Congress: pp. 103, 168; LiquidGhoul: p. 22 left;
The Lord of the Allaurs: p. 24; nasa: p. 98; National Library of Med-
icine, Betheseda, Maryland: p. 181; National Library of the Netherlands:
p. 65; Minneapolis College of Art and Design Collection: p. 42; The

203
Minneapolis Institute of Arts Collection: p. 117 (The Ethel Morrison van
Derlip Fund 66.25.171); Des Musées de Strasbourg: p. 71; Princeton
University Library: p. 37; Rex Features: pp. 16 top (Design Pics Inc), 23
(Chris Martin Bahr), 53 (c. Walt Disney/Everett/Rex Features), 78
(MonkeyBusiness Images), 79 (Bernard Caselein/Nature Picture Library),
93 (MonkeyBusiness Images); Professor David S. Richard, Susquehanna
University: p. 112; Science Photo Library: pp. 104 (Sheila Terry), 137
(National Museum of Heath and Science); Dr Tony Shaw: p. 74;
Courtesy of Special Collections, University of Houston Library: p. 9;
Tnarg 12345: p. 15; University of California, San Diego: p. 146 top; Tim
Vickers: p. 179 bottom; Kevin Walsh: p. 14; Warburg Institute, London:
pp. 11, 34, 61, 69, 176; Wellcome Library, London: pp. 10, 64, 118; Andrea
E Wills: p. 127; W-van: p. 16 bottom; Zoological Society of London: pp.
12, 19, 21, 27, 29, 32, 57, 113, 128, 141, 146, 154.

204
 Index

Aesop 87, 88, 162, 167 plague of frogs 59, 60, 61, 61
Agassiz, Louis 117 Bindman, David 85
alchemy 35, 36, 37, 37, 38, 39, 64 Boccaccio 58
Aldrovandi, Ulisse 7 Bombina bombina (European
Alytes 26 fire-bellied toad) 82
Alytes muletensis (Mallorcan Bosch, Hieronymus 143
midwife toad) 131 Briggs, Robert 135
Alytes obstetricans (midwife toad) 129, Budweiser 173
130, 132 Bufo alvarius (Sonoran desert toad) 33
Amphibia 13, 55, 67, 119, 128, 134, 182 Bufo baxteri (Wyoming toad) 175
anthropology 9, 77 Bufo marinus (cane toad) 33
Anura, definition of 13, 18 as invasive species 89–94
Aquinas, Thomas 67, 68 Bufonidae (family of ‘true toads’) 13, 22
Aristophanes 17 Bulgakov, Mikhail 130–32
Aristotle 67, 68, 70 Burroughs, John 17
artificial fertilization 135, 153
Ashmole, Elias 35 Caribbean 179
Atelopus zeteki (Panamanian golden Carroll, Lewis 169, 170
frog) 178 Carter, Julian 150
Australia 15, 19, 25, 33, 77, 89–92, 179 Ceratobatrachus guentheri (Solomon
avarice, toad as symbol of 72–3, 73 Island leaf frog) 177
Ceratophrys cornuta (Surinam horned
Barbourula kalimantanensis (Bornean frog) 57
flat-headed frog) 21 Ch’an Chu 155, 156, 157, 159
barometers, frogs as 146, 147 Chambers, Robert 116, 128
basilisk 68 Chile 25
Batrachomyomachia 162–3, 163 China 39, 77, 78, 156, 157, 158, 181
Belloc, Hilaire 161 Christian symbolism and frogs 34, 36,
Bibby, Cyril 151 39, 99, 114
Bible 59, 62, 62, 63, 67, 70 chytridiomycosis 178–80, 180, 181–2

205
classification 7, 9, 10, 13, 119, 121 frog as transitional organism 117,
difference between frog and toad 119, 120, 120, 121, 122, 122, 138
11, 13, 57, 110 experiments on frogs 95–115, 96, 104
related to edibility 77 dissection 112, 113, 113, 114, 150
climbing 19, 20 physiology 97, 104–8, 122
collection of frogs 177–9
Colombia 23, 25, 26 feng shui see Ch’an Chu
comic value of frogs 115, 161, 162, 162, fertility 143, 144, 145, 147, 148, 149, 160
171, 172, 173 prevention of 143
comparative anatomy 29 Fiddlesticks (Flip the frog) 172
consumption of frogs by humans 45, Fischer, Tibor, Under the Frog 74
77–83, 78, 79, 178, 181 Fisher, Jeremy 168–9
Cruikshank, Isaac 85, 86 France 77
Cuvier, Georges 10, 58 French as frogs 79, 83, 84, 85, 86, 87,
165, 166
Darwin, Charles 117, 119, 133 Frogger 174–5
decline and extinction 175–7, 175, 178, ‘Frog King, The’ 40–43, 40, 41, 42
178–82, 183, 184 Frog Prince, The (Lotte Reiniger) 43–9,
Dendrobates lehmanni (Lehmann’s 51, 52, 107
poison frog) 26 Princess and the Frog, The 49–54, 53
Dendrobates pumilio (strawberry poi- ‘Frog went a-courting’ 84
son frog) 26, 27
Dendrobates tinctorius (dyeing dart Gabriel, Peter 48, 53
frog) 24 Galvani, Luigi 102, 103, 103, 107
Dendropsophus ebraccatus gastric brooding 25, 26
(hourglass tree frog) 183 Gere, Cathy 66
developmental biology 29 Gerobatrachus hottoni (Hotton’s old
diet 20, 27–8, 34 frog or ‘frogamander’) 13
distribution 14 Gliboff, Sander 130, 134
Driesch, Hans 124, 125 Goethe 63, 64
Du Bois-Reymond, Emil 105, 106, 107 Gould, Stephen Jay 138
Duffy, Carol Ann 149 Griffiths, Richard 131
Dutch as frogs 85, 86 Gurdon, John 135

Eco, Umberto 82, 93 Haeckel, Ernst 119, 120, 120–22, 122,

ecology 15, 29, 175, 176, 177, 177 123, 123, 127, 129, 133, 134, 138, 139
Egypt 140, 144, 144, 145, 145 Hall, Marshall 108
embryology 122, 123, 124, 125, 126, haplessness 167, 168, 168, 173
127, 128 Harvey, William 65, 66, 70, 75
Endean, Robert 22 Heket 34, 144, 144, 145, 145
etymology of frogs 11, 12, 13, 17 Helmholtz, Hermann von 95, 105,
evolution 13, 14 107, 137

206
Hemisus marmoratus (African pig- Magendie, François 104
nosed frog) 20 Magnolia 60, 62
herpetology 10, 29, 115, 161, 179, 181, 182 Manduka Suka 145
hibernation 16, 17 Mandukya Upanishad 31
Hogarth, William 85 Mantella 177
Hogben, Lancelot 136 Mary, Queen of Scots 11
Hoplobatrachus tigerinus (Indian bull- mating 18, 141, 141–3, 151, 154, 154, 155
frog) 82 Merian, Maria Sibylla 117
Huxley, Thomas 108, 109 metamorphosis 7, 27, 29, 31, 32, 35,
Hyla (genus of tree frogs) 12 36, 63, 67, 117, 134
Hyla arborea (European tree frog) 100, as metaphor for evolution 116,
101, 146 117, 118, 128, 129, 138
Milton, John 57, 168
India 77, 79 Müller, Johannes 104, 105, 106, 107
Indonesia 82 Mungomery, Reginald 89, 90
Iraq 87, 89, 89
Native Americans 147, 148, 149
Japan 12, 76, 87, 162, 164 New Guinea 9, 17
Jones, Ernest 30, 46, 47 New Zealand 7
jumping 18, 56, 161, 163, 165–7, 167 newts 13, 130
Nobel and Ig Nobel Prizes 77, 115
Kammerer, Paul 127–30, 132–4
Kermit 173–4, 174 One Froggy Evening 173
King, Thomas 135 Orwell, George 141, 142
Kippenberger, Martin 114 Otis, Laura 105
Kircher, Athanasius 68 Owen, Richard 119
kiss 35, 42, 43, 51, 52, 53
paedomorphosis 134
Larkin, Philip 73, 74, 74 Page, Sophie 75
Leptodactylus fallax (giant ditch frog) Palaeobatrachus 14
179 Pan’s Labyrinth 75
Linnaeus, Carl 55, 76, 119 Panama 140, 148, 178
Litoria caerulea (Australian green tree Paracelsus 143
frog) 19, 20, 22, 22 Peru 179
Litoria nasuta (Australian rocket frog) Pflüger, Edward 122, 123, 124
18, 19 pharmacological properties of frogs
Litoria platycephala 15, 15 24, 31, 33, 58, 80, 181
Liu Hai 39, 157, 158, 159 Phyllobates terribilis (golden
loathsomeness 10, 55, 56, 75, 75, 76, poison frog) 23, 25
92, 94 Phyllomedusa bicolor (giant leaf frog) 19
Lobel, Arnold, Frog and Toad 169–72, physiology 29
171 Pipa pipa (Surinam toad) 26, 27

207
Pliny the Elder 58, 80, 144 Spemann, Hans 125–7, 135
Plumb, Christopher 100 spiders 58
Pocket Frogs 178 squatting 56, 72, 73, 74, 161
poison 22, 23, 24, 57–9, 63, 157 stone in head of toads 39
Pseudophryne corroboree (southern cor- stone, frogs inside 68
roboree frog) 22 Süskind, Patrick 142
putrefaction, generation of frogs by Swammerdam, Jan 34, 97, 98, 99, 99,
67, 68, 70, 71, 72, 75 102, 153, 184

rain, frogs falling as 60, 61, 62, 140 tadpoles 20, 21, 25, 26, 27, 77, 79, 97,
Rampling, Jennifer 36, 38 98, 124, 126, 139
Rana catesbeiana (American bullfrog) altered development of 134–5
17, 82, 83, 93, 93, 180 as hieroglyphic 140
Rana esculenta (edible frog) 79, 80, 82, in sex education 149, 150, 151
83, 126 Temnospondyli 13
Rana fusca 126, 127, 128 Toad, Mr 167, 168, 174
Rana palustris 126 toady 57, 89
Rana sylvatica 16, 17, 135 tongue 20
Rana temporaria (European Topsell, Edward 9, 10, 11, 66, 67, 70,
common frog) 112, 168 72, 75, 85, 110
Randall, Michael 83 tree frogs 13, 19, 20, 28, 29
regeneration 130, 134, 138 Twain, Mark 163, 165–7
Renaissance 7
respiration 21, 102 usa 15, 16, 17, 33
Rheobatrachus 25
Rhinoderma darwinii (Darwin’s frog) Vieraella herbsti 14
25, 26 vitalism 106, 107
Ripley, George 35–9 Volta, Alessandro, 103, 104
Rosenhof, August Johan Roesel von 8,
21, 32, 113, 141, 146 warts 57
Roux, Wilhelm 122–3, 124, 124–5, 127 Wassersug, Richard 77, 139
Willis, Deborah 63
salamanders 13, 130 witchcraft 63, 64, 64–8, 70
Scaphiopodidae 15, 16 women give birth to frogs 64, 65, 70
Scaphiopus couchii (Couch’s spadefoot
toad) 16 Xenopus laevis (African clawed frog)
sex education 149–51 135, 136, 138, 180
Shakespeare 39, 63 cloning 135
snakes 9, 10, 11, 38, 55, 70, 71, 72, 132, in pregnancy testing 136–7, 137
144, 164 Xenopus tropicalis (western clawed
sound of frogs 17, 18, 100 frog) 127, 138
Spallanzani, Lazzaro 57, 100, 152, 152–5

208