Stripped Bare: The Art of Animal Anatomy 9780691181424, 069118142X

A lavishly illustrated compendium of the art and history of animal anatomy from antiquity to today For more than two tho

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Stripped Bare: The Art of Animal Anatomy
 9780691181424, 069118142X

Table of contents :
From Antiquity to the Renaissance
Leonardo da Vinci: Animal as Machine
Albrecht Dürer: The Wonderful Line
Cutting and Cataloguing: 16th–19th centuries
Carlo Ruini: Anatomia del Cavallo
George Stubbs: The Anatomy of the Horse
A Pictorial Menagerie: 16th–19th centuries
Volcher Coiter: De Partibus Similaribus Humani Corporis
Georges Cuvier: Le Règne Animal
Alfred Brehm: Tierleben
Richard Owen: The Anatomy of Vertebrates
Evolution and Development in the 19th Century
Ernst Haeckel: Development of the Embryo, Development of the Race
Edweard Muybridge: Animal Locomotion
Santiago Ramón y Cajal: Textura del Sistema Nervioso del Hombre y de los Vertebrados
The Wonders of Life since 1900
D’Arcy Wentworth Thompson: On Growth and Form

Citation preview

stripped bare the art of animal anatomy

David Bainbridge

stripped bare the art of animal anatomy

David Bainbridge

princeton university press princeton and oxford

To Michael Blooman, For the art, Italy, and new ways of seeing.

Published in 2018 by Princeton University Press 41 William Street Princeton, New Jersey 08540 © 2018 Quarto Publishing plc an imprint of The Quarto Group All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording or any information storage and retrieval system, without prior permission in writing from the publisher. Library of Congress Control Number: 2018935458 ISBN 978-0-691-18142-4 QUAR.SKEL Conceived, edited, and designed by Quarto Publishing plc, an imprint of The Quarto Group Design: Blok Graphic, London Editor: Kate Burkett Art editor: Jackie Palmer Picture researcher: Sara Ayad Publisher: Samantha Warrington Printed in China 10 9 8 7 6 5 4 3 2 1

Contents Introduction Chapter One

Chapter Two

Chapter Three



From Antiquity to the Renaissance


Leonardo da Vinci: Animal as Machine


Albrecht DÜrer: The Wonderful Line



Cutting and Cataloguing: 16th–19th centuries


Carlo Ruini: Anatomia del Cavallo


George Stubbs: The Anatomy of the Horse



A Pictorial Menagerie: 16th–19th centuries


Volcher Coiter: De Partibus Similaribus Humani Corporis

Chapter Four


Georges Cuvier: Le Règne Animal


Alfred Brehm: Tierleben


Richard Owen: The Anatomy of Vertebrates



Evolution and Development in the 19th Century


Ernst Haeckel: Development of the Embryo, Development of the Race Edweard Muybridge: Animal Locomotion

196 206

Santiago RamÓn y Cajal: Textura del Sistema Nervioso del Hombre y de los Vertebrados Chapter Five



The Wonders of Life since 1900


D’Arcy Wentworth Thompson: On Growth and Form






Introduction The study of animal anatomy has led to some of the most striking images ever created.


or two and a half thousand years,

or augmented for artistic impact. Artists

animal bodies have been picked

draw animals because they respect or love

apart to drive arguments in natural

them, or both, and the emotional impact

philosophy, to reinforce dogma, to remind us

of viewing animals cannot be expunged

of death, to horrify, educate, and enthral.

from their work. A flamingo’s skeleton is

I teach comparative anatomy to veterinary

not just an armature of spindly struts, and

students at Cambridge University, so I

a cow’s innards are not just a series of pipes.

spend my days surrounded by the interior

Artists cannot ignore the fact that a rearing

world of animals—skeletons, pickled pots

cobra comes with an instinctual backstory

of viscera, tendons, ligaments, offal. I have

of fear and threat, even when it is reduced

unexpected objects in my office, and many

to a skeleton. In short, there is usually a lot

of them are just begging to be drawn.

more art and emotion in these images than

Some are impressive and some beautiful.

is strictly necessary.

There is much more to animal anatomy

In producing the works in this book,

than describing the structure of animals for

craftsmen extracted a clean, simple truth

those who need to work with them—I would

from dirty, complex objects. Dissection is

call that practical “veterinary anatomy.”

rarely a neat procedure, and many of the

Another reason to create images of animals’

subjects depicted must have been messy

constituent parts is to add to the sum of

and malodorous at the time. Yet here

human scientific knowledge by finding out

they are, abstracted objects of clarity and

how animals work, or why they are dispersed

beauty, cleansed of their mundane filth and

into such a myriad of bewilderingly disparate

presented in woodblock print, pen and ink,

forms; this could be called “comparative

lithograph, oil, or luminous spray paint. My favorite moment in preparing this

anatomy” or even “zootomy.” A further reason to depict animal anatomy is art

book was to discover in my workplace

itself: to exploit animal structure to create

a forgotten sixteenth-century Venetian

works that inspire.

imprint of the Anatomia del Cavallo

These three motivations inform the works in this book to varying extents, but

(“Anatomy of the Horse”), the motherlode of Western animal anatomy by Carlo Ruini

art is always there. Even the most pragmatic, functional images hidden in anatomy textbooks are framed, distorted, simplified,

Synsacrum of a domestic fowl.


(Previous page) Pieter van der Heyden after Pieter Bruegel the Elder, Big Fish Eat Little Fish, 1557.

same mechanical functions. Muscles and tendons radiate and converge like pistons in a strange, medieval war-engine. And guts wriggle like convoluted sewers beneath some steampunk city. Today, we understand what

(see page 54). (For examples of Ruini’s

all these body parts actually do, and, unlike

work, see pages 54–60.) This volume is

many of our predecessors, we also know

in remarkable condition, with stout, firm

that they were not fashioned by a benign

binding, and its crisp, surprisingly modern-

creator, but by the uncaring hand of patient

looking images still look fresh in today’s

natural selection. To realize that the working

jpeg world. Yet among the pictures hides a

animal components depicted in this volume

profile of a horse’s face, veins exposed, its

were not consciously designed is remarkable

eye staring terrified from the plane of the

enough, but it is truly incredible to consider

page as if to remind us that these beautiful

that, instead, a few simple evolutionary rules

creations spring from somewhere dark and

forged all this complexity and variety. This book recounts the intertwined

hidden from genteel view. Images of animal anatomy are also

intellectual and artistic journeys of animal

arresting because they remind us of other

anatomy from antiquity to the present

things. Spars of bone vault and arch like

day. Rather than offering an exhaustive

the pillars and buttresses of a cathedral,

listing, it focuses on the distinctive artistic

and, indeed, they perform precisely the

flavors of the five main overlapping phases of anatomical endeavor. The first phase

Kitâb al-baytara (“Treatise on Hippiatry”). Egyptian manuscript, 1670; Skeleton of a horse.

is largely defined by that which no longer remains: we know that savants from antiquity to the Renaissance drew animal


J. E. V. Boas (1855–1935) and Simon Paulli (1865–), The Elephant’s Head: studies in the comparative anatomy of the organs of the head of the Indian elephant and other mammals.

anatomy, but what remains mainly tells a

drive for artistic and scientific progress

story of charming ignorance and accidental

created the modern world. I imposed one restriction on myself when

survival. Then, from the sixteenth century, the horse was king, and can sometimes

writing, and that is the types of animals I

appear to have been the only animal worth

included. The images are all of vertebrates:

depicting, probably as a result of its practical,

animals defined by having backbones and

financial, and social value, and also, no

which also possess limbs, mouths, eyes, ears,

doubt, because horses are rather handsome.

and faces that make them more emotionally

The third phase, from the seventeenth to the

accessible to the human eye. There are no

nineteenth centuries, reflected an obsession

insects, worms, or molluscs, partly because

with the sheer variety of nature, with artists

I know little about them and partly because

deliberately selecting the most bizarre and

I would argue they are just too alien for us to

obscure creatures they could find as their

relate to emotionally. So, here instead are only vertebrates.

models. Next came the nineteenth century, when God lost control and anatomists first

After all, they look a little like us and seem a

realized that mundane processes generate

bit like us, too. Horses opened like books, the

animal form: sex, inheritance, evolution,

leer of a shark’s eye, the humming loom of

embryonic development, and physics.

the brain—all life is featured here, dissected

The fifth and last phase is the twentieth

and depicted.

century, when everything changed, as new David Bainbridge, Cambridge, 2018

techniques, new ways of thinking, and a 11

chapter 1

before the press From Antiquity to the Renaissance

Hans Hoffmann (c.1530–1591/2; after Albrecht DÜrer, 1524), Left Wing of a Blue Roller, c.1580. Watercolor and gouache on vellum.

For ancient animal art, the medium is as important as the message.

before the press


n the five centuries before the birth of Christ, Greek thinkers such as Democritus, Empedocles, and Aristotle studied the internal structure and function of the body in detail. They even drew parallels between animals and humans in a way not again permitted until the Renaissance. In many ways, Aristotle was the first true anatomist and a comparative anatomist at that—he even wrote a text called The Parts of Animals. So precise and vivid are the anatomical writings of the ancient Greeks that it seems inconceivable they did not also draw diagrams to support their ideas. Yet, frustratingly, no graphic representations of animal structure survive from this time. So, the first true anatomical depiction in European art is not a drawing, but something altogether more robust: a bronze sculpture, small enough to nestle in the hand. Dug from a field in north-western Italy in the late nineteenth century, the Piacenza Liver was probably crafted in the late second century bce. An enigmatic object at first sight, it is, in fact, a miniature representation of the characteristically slablike liver of a ruminant animal, etched with labels written in Etruscan. Of all the entrails “read” by the soothsayers of antiquity, lambs’ livers were the most assiduously interrogated, and therein lies the clue to the Piacenza Liver’s function. Most anatomical art is produced for one of three reasons—for the education of those who work with animals, to illustrate a concept in natural science, or for sheer artistic expression. The Liver is exceptional because it was created for none of these purposes, but existed instead for augury or, more specifically, “hepatoscopy.” The Piacenza Liver qualifies as true anatomical art because it represents a clear attempt at anatomical accuracy. It recreates the bilobed shape of the ruminant liver, its teardrop-shaped gall bladder draped lazily toward the right side and the two distinctive protuberances delving backward into the animal’s intestines. To some extent, the liver simply grows to fill the space left by other viscera, and this is why its shape varies subtly between individual animals—and hence the ancients’ fascination with those variations. The Piacenza Liver is the most striking, but by no means the earliest, example of hepatic art. There are, for example, clay livers from Babylon 14

The Piacenza Liver. Life-sized model of a sheep’s liver covered in Etruscan inscriptions, found at Gossolengo near Piacenza, late 2nd century bce.


from antiquity to the renaissance

inscribed in Akkadian almost two thousand years earlier. Thus, half the history of animal anatomical art might appear to be dominated by strange livers, but this is probably only because bronze and clay survive better than papyrus and vellum. We know the Greeks did draw diagrams—for example, the architectural plans for the great temple at Didyma in modern-day Turkey survive, etched on the monument itself, probably because someone forgot to delete them. Yet little else survives from the classical world. In the second century ad, Claudius Galen described in remarkable detail the internal structure of animals, coining terminology still used today, but no images survive. A Roman statue of an eviscerated sheep sits incongruously in London’s British Museum, as do the wonderful wall reliefs of the royal lion hunts of Nineveh from seven centuries earlier, reveling in the anatomical accuracy of their depiction of pain. However, these isolated examples are without any context of the anatomical understanding that went into their creation. The historical record of animal anatomical art continued in this fragmentary manner throughout the early medieval era. While this is often claimed to be due to proscriptions placed on dissection by the Christian church, this seems unlikely at a time when far more people were experienced in butchering animals than is the case today. The

before the press

Wall panel relief depicting a lion hunt. Neo-Assyrian, from the North Palace at Kouyunjik (Nineveh), Northern Iraq, 645–635 bce.

dearth of Western medieval anatomical art may be due to a lack of scientific inquisitiveness, an assumption that everyone knew what lay inside a pig’s belly, or, most probably, because the graphic media used has simply rotted or crumbled away. Around the world, scattered examples remain—mysterious petroglyphs (carvings or inscription on rocks) in what is now the southwestern United States and Armenian treatises on equine medicine—but not much survives in Europe. If the intellectual lights really did go out during the early European medieval period, then, as was so often the case, a spirit of enquiry persisted instead in the Islamic world. Fifteenth-century images from the Middle East hint at a meticulous reductionism absent from the Christian West, and a sense that previous knowledge could be improved upon. Of course, these depictions appear crude by today’s standards, but they look as if they were made by people who not only wanted to understand how animals work, but had also actually seen the things they were drawing themselves. The same cannot be said of Western European bestiaries. These diligent catalogs of God’s creation were never meant to be anatomy books, but it is the anatomy that exposes the ignorance that underpins them. Jacob van Maerlant’s fourteenth-century elephants, for example, are certainly impressive creatures, but their huge, funnel-like trunks 16

Leonardo da Vinci (1452–1519), A bear’s foot, c.1488–90.


from antiquity to the renaissance

indicate that their portraitist had never actually seen one. Even more cryptic are Maerlant’s various horned beasts (see page 24). The fact that his “antlered fish” bears five legs suggests it is perhaps an unusually blessed, and presumably never-encountered, walrus. It is a cliché to state that Leonardo da Vinci (see page 26) played a pivotal role in the cultural and scientific history of the world, but in the case of the art of animal anatomy, that cliché is true. Suddenly, at the end of the fifteenth century, a genius appears, looks at the world in new ways, creates some artistic landmarks, and, fortunately for us, leaves behind copious notes. Looking at da Vinci’s drawings comparing the anatomy of animals to that of humans is to realize that, as the saying goes, he truly was centuries ahead of his time. For example, it is not clear whether his anatomical study of a bear’s foot was based on an ursine template or a poorly pedicured human one, yet he had the scientific insight to realize that, unlike most quadrupeds, the bear shares our species’ “plantigrade” foot position (that is, it walks on the sole of its foot, with the heel touching the ground). Again and again biological insights spring from the page, even if they are implied by the images rather than explicitly stated in da Vinci’s famously inverted handwriting. In fact, the main problem with his anatomical musings is that there are not enough of them. He is easily distracted, and his notes on animal anatomy are often mixed with or

before the press

Leonardo da Vinci (1452–1519), Rete mirabile, detail from a drawing of the brain and cranium, c.1508–09.

supplanted by speculations on war machines, human flight, or how to capture the drapery or flowing hair of the angel in his next masterpiece. Da Vinci was also prone to two of the most common anatomical errors: assuming that the ancients were correct and mistakenly conflating animal and human structures. In one drawing he inserts a rete mirabile—the “wondrous net” of arteries that is thought to cool the brains of many hoofed mammals—underneath a human brain, where no such net is found. This mistake can be traced back to Galen who evidently did not dissect humans as thoroughly as he claimed, an error compounded when he asserted that this nonhuman structure is the very seat of the soul. Da Vinci obviously never checked Galen’s observation, so he recreated a nonexistent rete in his own beautiful images. A similar error, described later in this chapter (see page 28), sees him showing a human fetus being nutrified via a perfectly drawn bovine placenta. Working at the same time as da Vinci, Albrecht Dürer (see pages 36–38) was an expert in visual animal pathos who also happened to be the greatest draftsman in history. Few have depicted the tragic dignity of the injured and slain as thoroughly as Dürer. So keenly is this felt in his work that it is hard to avoid concluding he had modern ideas about animal sentience and suffering. His Head of a Stag Shot by An Arrow 18

Albrecht Dürer (1471–1528), Head of a stag, pierced by an arrow, 1495 or 1504.


from antiquity to the renaissance

(below) is suffused with melancholy rather than any sense of victory at an animal slain for the pot, while his Injured Hare foreshadows the story of an imminent merciful killing. While da Vinci was a man of dispassionate, almost callous, analysis, Dürer’s lush images speak of a sad empathy. The final image in this chapter—Conrad Gessner’s Cetus ingens or “Huge whale” (see pages 42–43)—is placed there not because it is chronologically the last, but rather because it is a transitional depiction. It shows a scene of utter carnage, in which the Leviathan has beached itself, been hauled ashore, and is now being cut to pieces by a group of enthusiastic Faroe islanders. The print is quite crude, homespun almost, and has all the traits of an event recounted rather than seen. Yet one feature of the beast hints at scientific enlightenment still to come—it is drawn, correctly, with mammary glands, an indication that the artist knew it was a mammal and not a fish. The importance of this realization for future science is not lessened by the fact that the whale erroneously bears the transplanted udder of a sow, rather than the more streamlined mammaries of a true cetacean. Gessner was making a scientific point about a Cetus ingens he had never seen and, in doing so, was signposting the direction his artistic successors were to take.

before the press

Model of a sheep’s liver used for divination in a Babylonian Temple School. Clay, c.2000 bce. Models of livers used for hepatoscopy—the prediction of the future based on the shape of animals’ livers—were produced in West Asia and around the Mediterranean for more than two thousand years. Indeed, they are the earliest surviving artistic representations of animal anatomy. This model shares its bilobed shape, teardrop-shaped gall bladder, and protruding caudate lobe with the Piacenza Liver (see page 15), but its inscriptions are in cuneiform.


The intestines offer the greatest scope for anatomical variation of all the internal organs, because they are relatively mobile. While this apparently writhing mass suggests chaotic churning, its form indicates two particular origins. First, sheep, goats, and cattle have a large intestine with a distinctively spiral portion, the ansa spiralis, which this model closely resembles. Second, the design echoes the labyrinth motif seen in ancient images found throughout a large geographical region, spanning the Atlantic to the Asian Subcontinent.


from antiquity to the renaissance

Model of a sheep’s intestine for extispicin, or the ‐­ rûtu divination, from examination of entrails for B­­­a Mesopotamia, Iraq. Clay, 2nd millennium bce.

before the press

Altar with a slain ram, Roman, 100–200 ce. Marble. This partially eviscerated sheep is a rare example of a surviving classical depiction of animal anatomy. The context of the statue is unknown, especially as it was carved at a time when most people would have been familiar with the process of butchery. The wriggling small intestine, the bulbous rumen, and the margin of the liver may be seen protruding from an abdominal incision.

Bighorn sheep pierced with arrows; Jornada Mogollon petroglyph at Three Rivers, near Tularosa, New Mexico, between 900 and 1400. High in the desert at Three Rivers, a series of low hillocks is covered with stones bearing thousands of petroglyphs: some abstract, some humanoid, and some zoomorphic. Inscribed by the local Mogollon tribes, the reason for their creation is entirely unknown, but some of the animal images include schematic representations of their internal organs—a coincidental echo of some Australian Aboriginal art.


Jacob van Maerlant (1230/35–c.1291), Der Naturen Bloeme. Dutch manuscript from Utrecht or Flanders, c.1350; “Cerius marinus” (“deer of the sea”); Elephant; Horned seafish; Vespertilio (“winged mince,”or bat). From a time when storytelling and spiritual guidance were more important than anatomical accuracy, van Maerlant’s many images are a charming reminder of the state of animal anatomical knowledge. Shown here are some of his more fanciful illustrations, apparently of beasts he had never seen or had not inspected closely. The antlered fish remains a mystery, but the bat and elephant are recognizable, despite the funnellike trunk of the latter—a common error in medieval texts. The fivelegged, tusked, aquatic animal is also problematic. One suspects it may be a walrus, but the superfluity of limbs suggests it is perhaps a lobster.


from antiquity to the renaissance

Sketch with handwritten notes from the Cilician book on the curing of horses, Armenian kingdom of Cilicia, 1295–98. Although little anatomical art survives from medieval Europe, there is considerable evidence that anatomical and veterinary science was thriving in Western Asia. This example comes from Cicilia, at the southeastern coast of Anatolia, and is written in Armenian. This language flourished from the Caucasus to the eastern Mediterranean, often in separate city-states that survived until the end of the Byzantine Empire. Although not an image of internal organs, this is a diagram of the various maladies that are associated with different anatomical regions.


Leonardo da Vinci (1452–1519): Animal as Machine

Leonardo da Vinci’s paintings have come down to us as gems: selfcontained works of genius in which the flowing locks, straining muscles, cascading drapery, and hazy chiaroscuro are so perfect that they obscure the thought and labor that went into producing them. Yet, fortunately and famously, da Vinci made detailed notes (in his famously inverted handwriting) and diagrams about the world around him and how it could be manipulated and processed in his art, and many notebooks still survive in treasured collections around the world. Da Vinci’s understanding of anatomy was crucial to his paintings, but his interest went before the press

further than that—he assiduously sketched structures unlikely to appear in any artwork, such as placentas, and his written notes make clear his interest in the structure of the body for its own sake. So da Vinci’s reinvigoration of the science of anatomy was not just a by-product of his artistic process, but a quest in its own right. He was certainly an intellectual butterfly, flitting between anatomy, mathematics, art, and engineering, but never so much that he could not focus on a single topic. For example, his diagrams of bones and muscles are far more technically accurate than anything that had gone before: they certainly look like drawings made from direct observation of the real thing. His sketch of a horse’s leg (opposite) shows bony and muscular structures that are clearly identifiable to the modern student of comparative anatomy, some of which are unique to this animal. To da Vinci, animal anatomy also served as a resource of visual motifs that he could use elsewhere in his work. The rambling course of the coronary arteries lies somewhere between the dividing forks of the veins of a leaf and the cascading rivulets of an angel’s hair, while bat’s wings are recruited into fantastical flying machines. The term “Renaissance man” is a tired cliché today, but has very real roots in a time when a single individual could know all existing scientific knowledge, and make genuine advances in several different fields of human endeavor.

Leonardo da Vinci (1452–1519), Horse’s leg, 1506–07.



before the press

Leonardo da Vinci (1452–1519), A foetus in the womb, with an ox’s placenta, c.1511. This human child has been magically furnished with a ruminant’s placenta. Had da Vinci inspected a single human placenta, he would have known that the baby’s main nutritive organ is a blood-red, discoid slab. Instead, he goes into great detail illustrating the cotyledonary placenta of an ewe or a cow—divided into tens of separate “mini-placentas.” On the far left, the clambering branches of the uterine vessels track over the surface of a correctly drawn, globular human uterus, yet the rest of the image erroneously depicts multiple regions where maternal and fetal tissues interdigitate. Top right, the maternal and fetal components are shown being peeled apart, just as a vet must do when a ruminant mother retains her placenta after birth.

Leonardo da Vinci (1452–1519), The heart and coronary vessels of an ox, c.1511–13.

One heart looks much like another, and bovine hearts are much easier for the artist to come by than human ones. In da Vinci’s day, the heart was thought to be far more than a pump—the Aristotelian cardiocentric model held that it was the central motive and mental organ of the body. In these sketches, however, the artist seems more interested in the twiglike branching of the coronary arteries.


before the press

Leonardo da Vinci (1452–1519), A horse in profile divided by lines, c.1480. No animal’s proportions have been manipulated as much as the horse’s. A horse’s importance in a scene can range from being part of the background to being its primary focus. Particularly important are horses’ relationships to people—especially those riding them—and a common trick is to reduce the size of the equine head to give a rider more prominence. Also, many artists struggled with the challenges of depicting a cavalry battle in an involving way; that is, allowing the viewer to scan the entire field, yet focus on the expressions on individual human and equine faces. Earlier, Paolo Uccello had used strikingly distorted visual perspectives to allow this in his mid-fifteenth century Battle of San Romano, and it is thought that da Vinci explored these devices further in his now-lost 1505 Battle of Anghiari.


leonardo da vinci: anim al as m achine

Leonardo da Vinci (1452–1519), Study for a mechanical wing with cryptographic notes, c.1493–95. Leonardo was quick to realize that animals’ miraculous abilities are due in part to their distinctive structures and, when he wished to design a machine for some purpose, he would often resort to animal anatomy for inspiration. It is not immediately apparent how birds’ feathers might work, so the Renaissance engineer looked to bats’ wings as a guide. In fact, neither structure works as simply and stably as an artificial aerofoil, so heavier-than-air manned flight had to wait for another four hundred years.


s ee pages 3 2 –3 3

Kitâb al-baytara (“Treatise on Hippiatry”), 15th century; Anatomy of the horse. Islamic art is known for its stylization, in which familiar curved or organic forms are frequently transformed into something more geometric. In this way, Arabic text may be deformed into almost unintelligible, angular calligraphic designs, and naturalistic images are often replaced by tessellating tiles and intricate abstract carvings. Similarly, this image of a horse is spatchcocked in such a way as to be nearly unrecognizable, to show the pure rectangular segments of its trachea and the mazelike course of its neatly folded intestines. In contrast, the lungs and heart receive a relatively

before the press

cursory artistic treatment compared to the prominent male genitals.


Like Leonardo da Vinci, of whom he was a contemporary, Albrecht Dürer was a great Renaissance artistic anatomist. However, while da Vinci’s images have a disinterested incisiveness, Dürer’s possess an inherent warmth—they invite empathy as much as analysis. From an anatomical point of view, bats have the most recognizable anatomy of any of the three groups of flying vertebrates (the others being birds and pterosaurs). In this painting, the elongated equivalents of the middle, ring, and little finger can be seen supporting the membrane of each wing. The “index finger” is smaller and not clearly shown, but the “thumb” is evident as a tiny clawed digit pointing forward from the leading edge of the animal’s wing. Unlike other flying vertebrates, bats’ wings extend to entrap the hind limbs and even the tail.

Kitâb al-baytara (“Treatise on Hippiatry”), c.1670, probably copied from a 15th-century Mamluk manuscript; Horse skeleton It is believed that Islamic medical and veterinary manuscripts frequently drew on the writings of classical authors, some of which no longer exist. However, Islamic veterinary science had its own distinct, questioning flavor. This clearly drawn, if slightly cartoonish, equine (opposite) demonstrates the “points” of the horse and the afflictions that may affect them.


from antiquity to the renaissance

Albrecht Dürer (1471–1528), Bat, 1522.

Albrecht Dürer (1471–1528): The Wonderful Line

Albrecht Dürer was born in Nuremberg to a family of craftsmen and artists, and he is perhaps the artist of animal anatomy for whom craft constituted the greatest part of his genius. Otherwise best known for his engravings and prints, many of his finest anatomical works were instead created in watercolor or pen and ink. Dürer was well traveled and corresponded widely­—he exchanged communications with Leonardo da Vinci, for example. Of all the artists of

before the press

Albrecht Dürer (1471–1528), The Rhinoceros, 1515. In perhaps Dürer’s most famous work of all, this bereft creature has been abducted to a new, and presumably shortened, life away from his natural habitat. Many of the wretched animal’s anatomical details—the scaly legs and the “hornlet” atop the withers, for example—are so wide of the mark, however, that it is doubtful whether Dürer, the great observer, ever actually saw it.


the northern European Renaissance he was most influenced by the works of the Italian masters, and many of their ideas and innovations suffuse his work. Initially, however, his own work was little discussed in Italy, but since that time, he has proved to be one of those rare artists who has never fallen out of fashion. Like many Italian artists, Albrecht Dürer had a wide range of interests—philosophy, geometry, astronomy, and, of course, anatomy— which he introduced into his art. Platonic regular polyhedra appear

Albrecht Dürer (1471–1528), Wing of a Blue Roller, c.1500 or 1512.

this a severed wing not only shows off the skill of the painter, but also tells a story of a small, everyday tragedy. It is worthwhile comparing this image with Hoffmann’s later reinterpretation which forms the title page of this chapter. Both are beautiful, but the brushwork on DÜrer’s original is astounding—every last covert and barb is softly but clearly drawn.

albrecht dÜrer:

A strange, disembodied subject for a painting, one might think, but even isolated like

the wonderful line


unbidden in his calm, reflective works, and wildlife casualties are elevated to the status of masterpieces. His studies of animal anatomy seem less direct, less obvious, than da Vinci’s. Rather than being acquired and slaughtered for his anatomical enquiries, Dürer’s subjects seem to have ambled or fluttered into his field of view, or were maybe simply grazing in a nearby field. His images are very much “northern” images—depictions of

before the press

objects, sometimes melancholy, under the light of cool, gray skies

Albrecht Dürer (1471–1528), A greyhound, c.1500–01. One usually thinks of greyhounds as lean and muscular, but Dürer has softened and warmed this hound in several ways. First, the outline is made up of individual brushstrokes of fur to produce a calm, even slightly diffuse, presence on the paper. Also, rather than being a pounding, bounding animal, the dog is shown at rest, with his limbs irregularly placed in a relaxed manner. In addition, he gazes trustingly up at an unseen master, presumably waiting to receive his next command. For all the emotional content of this image, its anatomical accuracy is formidable, and even today I use it in my clinical examination teaching classes.


and lengthening nights. And yet, paradoxically, the softness and gentleness of his anatomical works are largely created by an immense precision and fluidity of myriad thin lines. To focus on a small region of a painting or drawing by Dürer is to discover a hidden world of beautifully expressive lines. Each is a little artistic opus in itself, and together their expertly flowing torrents combine to tell powerful stories of science and emotion.

albrecht dÜrer: the wonderful line

Albrecht Dürer (1471–1528), Muzzle of an Ox, 1501–5 (later inscribed 1523).


There is a long history of descriptions of rabbits and hares with horns or antlers, from the above image to contemporary claims of the North American “jackalope.” Conrad Gessner was a Swiss naturalist and one of the first to assemble what we would now recognize as a modern catalog of animal species. To a great extent, he was dependent on information from his correspondents across Europe, and this may be whence the idea of this horned hare came. The animal clearly possesses antlers rather than horns, as they are branched, and the partial skull on the left looks like that of a roe deer—the most abundant native small deer in Europe.

Andreas Vesalius (1514–1564), Tabulae anatomicae sex, 1538. One area in which the comparison of animal and human bodies caused confusion was the role of the rete mirabile or “wondrous net” (see page 18). Andreas Vesalius was a key figure in the examination of animal and human anatomy, and shown here is a diagram from his book of “six anatomical tables,” depicting the arteries of the human body in highly schematic form. At the very top of the image, hidden in the crown-of-thorns-like arteries of the head and face, sit ovoid masses that represent a meshwork of vessels underlying the brain, the retia. These arterial plexi are now thought to allow cooler blood to reach the brain, especially in hoofed mammals and desert species. More than a millennium earlier, Galen had described the retia he found residing beneath the animal brain, and even claimed they were the seat of the soul. It was for human anatomists some time after Vesalius to point out that, ironically, our own species is one of those that does not possess a rete.


from antiquity to the renaissance

Conrad Gessner (1516–65), Conradi Gesneri medici Tigurini Historiae animalium liber primus de quadrupedibus viuiparis... 1602; Lepus cornutus (“Horned rabbit”).

before the press

Conrad Gessner (1516–65), Conradi Gesneri medici Tigurini Historiae animalium... 1604; Whale with teats being attacked by mariners. “A huge whale, pushed ashore by a tempest and secured by an anchor, which the inhabitants of the Faroe Islands, habitual fish eaters, cut apart with axes and divide among themselves.” It is doubtful that Gessner ever traveled as far as the Faroe Islands, but this image shows he had heard of some of the distinctive features of whale anatomy. First, cetaceans’ nostrils have migrated from their snouts to the top of their heads to become blowholes. This is illustrated in the image above, although those holes have for some reason been augmented by snorkel-like extensions. Also, Gessner had obviously


from antiquity to the renaissance

heard that the whale, like land mammals, produces milk for its young. Indeed, he clearly realized the importance of modes of reproduction in classifying animals, because the first two of the four volumes of the Historiae Animalium are allocated, respectively, to land animals that give birth to live young (Quadrupedes vivipares) and those that lay eggs (Quadrupedes ovipares). Never having seen a female whale himself, however, Gessner transplants the unwieldy udder of a sow in place of the streamlined mammary glands of the whale. Despite these errors, Gessner’s innovation of classifying animals on the basis of their anatomy or physiology was to come to dominate zoology in later centuries.


chapter 2

THE HORSE STRIPPED BARE Cutting and cataloging: 16th–19th centuries

George Stubbs (1724–1806), A Lion Attacking a Horse, 1762. Oil on canvas.

Of all the beasts, one species reigns supreme in the art of animal anatomy.

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orses were neither the earliest nor the most numerous creatures to be recruited to live alongside humans, but they hold a unique place in human civilization and the human heart. From the Middle Ages until the recent advent of the worldwide small-furry-pet industry, the horse has dominated veterinary comparative anatomy. No one is certain of the origins of the word “veterinary,” but it may have the same etymology as “veteran”—indeed, veterinary medicine was originally reserved for animals of such value that they were permitted to grow old. In contrast, farm animals are usually eaten when young and tender, or once the flow of milk wanes, and are as replaceable as the dogs and cats that guard, retrieve, or hunt for us. Yet horses are different: each represents a valuable investment. Even a foal is a special, vulnerable creature, and rearing it to an age when it can perform its intended purpose is expensive. Horses are kept for practical, everyday reasons, but I think the historical importance of those reasons has been overstated. For the last few thousand years, the main draft animals used in times of both peace and war have been the tractable ox and resilient donkey. In comparison, horses have tended to have higher-status roles: propelling the knight into battle, drawing the carriages of the rich, competing athletically in races, performing quadrupedal acrobatics, or simply existing as talismans of their owners’ wealth and status. In addition, there is something about the anatomy of the horse that seems to appeal to the human eye. Although few of us give them much thought, horses’ bodies are perhaps the most specialized of all the domestic animals. Some animals—such as the bear, with its evenly proportioned, stocky body, which can climb, swim, dig, manipulate, maul, and gambol—are generalists. In contrast, equine anatomy speaks of an evolutionary drive to walk and graze with minimal effort, and to stir to speedy flight when necessary. Horses’ muscles are bunched and tensed at the top end of their limbs, close to the central muscular powerhouse, whereas the lower limbs are spindly, fleshless, and vulnerable. Each leg perches on the fingernail of a lone, bizarrely elongated digit in a manner perfect for fast running, but useless for 46

anything else other than the occasional feisty kick. Yet these alien, hyper-specialized creatures are all around us, inviting the artist to comment on their incongruously familiar anatomical strangeness. Another reason why horses may have dominated so much of animal art is, I believe, that people are sensually conflicted about them. One explanation for this conflict is that they are notoriously capricious creatures—calm, quiet grazers one minute, but provoked to a frenzy of panic or aggression the next. Horses are certainly emotionally demonstrative, with their snorting, whinnying, and wide-eyed nostril flaring, and artists seem to find their prancing, rearing, and cowering irresistible. As we will see, some artists revel in this equine expressiveness by depicting fantasies of horses being ambushed by devilish predators. Furthermore, the relationship between humans and horses has been strangely sexualized over the years. From certain viewpoints 47

c u t t i n g a n d c a t a l o g i n g: 16 t h – 19 t h c e n t u r i e s

Andrew Snape (1644–1708), The Anatomy of an Horse, 1683; The upper part of the windpipe fastened to the os Hyoides.

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Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Horse’s heart.

they appear toned, muscular, and handsome, while from others they look disquietingly curvaceous for a beast: a firm, potent presence to feel between one’s stirrups. In this light, Peter Schaffer’s 1973 play Equus—which explores the pathological relationship between the male protagonist and horses—can be seen not as an isolated perverse statement, but rather as the logical conclusion of the uneasy, centurieslong bond between our two species. For all these reasons, when anatomists step in to flay and dissect horses, it seems a violation so disturbing and compelling that artists cannot resist it. Horses were not the animal that started the story of modern Western anatomical art—instead the honor goes to us humans. In late Renaissance Europe there seems to have been few restrictions on medics dissecting human corpses, and this relative freedom caused a flowering of anatomy led, most of all, by a man from the low countries, Andreas Vesalius. Vesalius worked as a practicing doctor, but after

Andreas Vesalius (1514–1564), De humani corporis fabrica... 1543.


c u t t i n g a n d c a t a l o g i n g: 16 t h – 19 t h c e n t u r i e s


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studying the works of Galen he turned his attention to making his own observations and testing his own ideas about human structure from his new base in Padua. His new anatomy made its greatest impact through the publication of De Humani Corporis Fabrica, “On the Fabric of the Human Body,” in 1543, a book which changed medicine and science forever. Indeed, almost every element of modern biomedicine can trace its ancestry back to Vesalius images. Yet the hero of this story, and probably the most important character in this book, is not Vesalius, but an Italian called Carlo Ruini (see page 54). Before Ruini, the art of animal anatomy was charming and quirky, but also error-strewn and crude—sometimes even cartoonlike. With the publication of Anatomia del Cavallo (“Anatomy of the Horse”) in 1598, Ruini created a book that was surprisingly modern in its clarity, its meticulous attempt at comprehensiveness, and its quest to test and sometimes challenge accepted wisdom. The text is clear and surprisingly accurate, and its images anatomically detailed and emotionally involving. We do not know if the Anatomia was based on previous, now lost, works, but if Ruini created it afresh from the morass of medieval anatomical confusion, then his achievement is truly miraculous. Today’s anatomist can view every plate and know precisely what it depicts without needing to read the text, and will be continually surprised by how much of what we consider to be modern knowledge is clearly explained in this sixteenth-century volume. In addition, Ruini may even have changed the entire course of medicine when he described the valves of the heart in detail, and the idea that blood can only pass in one direction between its chambers became a central tenet of William Harvey’s later work on the circulation of the blood, De Motu Cordis, which is regarded as the fount of modern physiology. Anatomy means, literally, “cutting up,” and Ruini displayed the internal components of the horse both together and separately. Viscera spill, albeit artistically, from incisions in otherwise intact bodies, but also individual organs are cut free to be presented and discussed alone—both literally and figuratively a reductionist process. Many of the horse’s internal organs are distinctively graceful and sleek: Ruini’s horse heart is not the lumpen, globular heart of a pig or ox, but a realistic statuesque cone; the lungs are delicately elongated and sculpted to fit the powerful equine chest; and the liver is displayed with its baroquely scalloped edges. In contrast, the intestines are the tortuous, sacculated coils of the most visceral of nightmares—the horse ferments its food in its large intestine, so this organ is distended out of 50

Wilhelm Ellenberger (1848–1929) and Hermann Baum (1864–1932), Handbuch der vergleichenden Anatomie der Haustiere, 1912; Veins draining the equine face.


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all proportion, and is even further inflated in death by the gaseous emanations of its resident microbes. With Ruini, the die was cast, and the rest of the history of equine anatomy followed his lead. Veterinary texts filled in the gaps in the Anatomia so that surgeons could cure with greater confidence the diseases of those exposed, bony limbs and that precariously writhing gut. The sciences of equine mechanics, genetics, and palaeontology were to dominate animal biology in the future— fortuitously, the horse has the most complete fossil record of any animal on earth. And finally, the urge for horsey artistic expression has always been there, culminating in George Stubbs (see pages 72–77) whose works feature prominently later in this chapter. As we will see, it was Stubbs’ artistic knowledge that helped to make him the greatest animal artist of all. Certainly, his ability to paint a glint in an eye matched even Goya’s.

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Giovanni Battista Ferrari (1584–1655); Trattato utile, necessario ad ogni agricoltore per guarir caualli... 1681; Skeleton of a horse. This diagram from Ferrari’s treatise on veterinary care is remarkably crude. Little distinguishes this horse from any other creature apart from the single digit on each foot. The canine nature of the skull, the unimpressive number of ribs (horses have 18 on each side), and the lack of carpal (wrist) and hock (ankle) bones suggest that the draftsman’s familiarity with the inner workings of the horse was cursory at best.

Gervase Markham (c.1568–1637), Markham’s maister-peece: contayning all knowledge belonging to the smith, farrier, or horseleech, touching the curing of all diseases in horses... 1717; Skeleton of a horse; Circulatory system of a horse. Markham’s aim was a bold one: to write a comprehensive book of horse management, health, and farriery—a masterpiece, no less. His skeleton is an improvement on the representation by Ferrari above, clearly looking like a horse, but it is still simplistic. His diagram of blood vessels is misleading in its certainty and obviously dates from before Harvey’s discovery of the circulation of the blood: the vessels branch and flow beautifully but aimlessly, never ending anywhere in particular. Obviously confident of his insights, five years later he published The English Huswife, Containing the Inward and Outward Virtues Which Ought to Be in a Complete Woman.


c u t t i n g a n d c a t a l o g i n g: 16 t h – 19 t h c e n t u r i e s


Carlo Ruini (1530–1598): Anatomia del Cavallo, 1598

Carlo Ruini was born and died in the beautiful city of Bologna, in Italy. Although little is known of the man himself, it was the posthumous publication of his masterwork, the Anatomia del Cavallo, Infermità, et Suoi Rimedii (“The Anatomy of the Horse, Its diseases, and Their Treatment”) that made him famous. Ruini had a privileged upbringing, and eventually rose to the position of Senator, but it is not thought he received any education as a physician (at

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that time, self-styled human and animal surgeons were a lowly class with no formal training). Historically, the Anatomia is a strange phenomenon— it seems to appear from nowhere. Ruini was known to have been inspired by previous unillustrated treatises on horse management and medicine, and also by the ongoing north Italian renaissance of human anatomy, but the comprehensiveness and accuracy of his work was entirely without precedent. If the core of the book is indeed the work of one man, drawing solely on previous written texts, then it ranks as the greatest achievement in animal anatomy. The text is long, but clear and detailed, yet it is the images that capture the modern imagination. From the very first glance, they make it clear that the Anatomia is a very modern work, and its contents can be measured favorably against contemporary standards of accuracy. Coupled with this is the artistic impact of the many diagrams throughout the book. Even when dissected almost to extinction, every horse retains its nobility, and some of them seem noticeably feisty. They have sturdy, thick horseshoes and piercing stares, and many also have flowing manes that would not look out of place on one of Botticelli’s goddesses. Almost certainly Ruini hired some of the best engravers of his age, but it seems certain that the controlling artistic vision was his. The Anatomia was reprinted many times in its first few decades, including the copy to which I had access during the preparation of this book, and was also translated into German. Perhaps the best indication of its impact is that, for centuries, almost every work on equine anatomy seemed to either plagiarize it or struggle desperately not to.


carlo ruini: a n at o m i a d e l c ava l l o

Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Intestines.


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Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Horse opened up. The ribcage of a supine horse, with its shoes still nailed in place, is opened like a pair of drapes to reveal the wonders beneath. The scalloped lobes of the liver (“C”) sit center stage, characteristically without the gall bladder in this species, and are connected via a large venous trunk to the heart, which is not visible in the image. Much of the rest of this internal view is filled with the diaphragm (“A”) and the large veins that snake across it.


carlo ruini: a n at o m i a d e l c ava l l o

Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Foal. A dead foal, probably removed from a deceased mare rather than itself stillborn, is peeled free of its membranous coverings. Foals are surrounded by two membranes, a pale inner one and a dark, vascular outer one, and the blood vessels branching across the upper figure suggest that this image depicts the latter. The lower image shows the cut stump of the umbilical cord, and also hints at the “hoof slippers”—soft, rounded excrescences of the hoof, which protect membranes and mother should the foal decide to frolic in utero.


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Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Hind limb muscles. The level of detail in this image far surpasses anything that went before it, and a modern veterinary anatomy student could name any of the muscles and tendons depicted here. The image on the right is the view from the side of the animal, the “outer aspect” of the limb, while the image on the left is the view of the animal’s inner thigh and limb. There is even room for the saphenous vein to course down the inner side of the limb, for thin sprigs of nerve to branch over the foot, and for the flowing locks of the tail to cascade groundward.




s ee page 5 9

Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Brain and meninges. The first image in the Anatomia is one of its most striking—two steeds ornately bridled together in death, their heads cleaved open to reveal the level of anatomical accuracy is high—the convolutions of the two cerebral hemispheres are visible in the right-hand animal, and above them lies the cerebellum or “little brain,” its hemispheres, vermis, and flocculus evident to a modern anatomist.

Here Ruini’s equine protagonist strides away, picking his rocky, unshod course through the trees and along a winding path to a distant fortress. Even in this view, subtle details are visible: the fine muscles that draw back the angle of the lips, the lobulated parotid salivary gland at the base of the ear, and the bifurcation of a particular tendon as it reaches the foot. Ruini’s images are exactly as simple or ornate as they need to be—details are emphasized, but the final visual impression is neither sparse nor cluttered.


a n at o m i a d e l c ava l l o

Carlo Ruini (1530–1598), Anatomia del cavallo, infermità, et suoi rimedii, 1618; Muscles viewed from behind.

carlo ruini:

the brain (right) and its meningeal membrane coatings (left). As always,

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Jean Héroard (1551–1628), Hippostologie, c’est a dire Discours des os du cheval, 1599; Forelimb; Hind limb. Héroard’s “Bones of the Horse” is a short volume of 50 pages or so, often overshadowed by Carlo Ruini’s book (which was published the previous year), yet it too represents a genuine attempt at anatomical accuracy. Here are illustrated, in a strangely horizontal position, the bones of the fore- and hind limbs, and “the box or shoe” (hoof) of each. In fact, there are few diagrams in the book, but they focus on the body regions with which animal doctors most often have to deal.


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Jean Héroard (1551–1628), Hippostologie, c’est a dire Discours des os du cheval, 1599; Horse skeleton. Héroard’s horse skeleton suggests a confident and strutting beast. Artistic judgement allows it to retain its hooves even in this reduced state—they presumably looked more fittingly stable than the diminutive phalanx bones that lie inside them.


the horse stripped bare

Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683. Andrew Snape was farrier to King Charles II of England, and his Anatomy of an Horse contains some of the most beautifully striking anatomical images ever made of this species. All the engravings were by N. Yeates, whose line is immensely fluid and subtle, but the improvement over Ruini is technical rather than scientific: almost every image in the book is a copy, albeit a mirror image, of a diagram in Ruini’s.


c u t t i n g a n d c a t a l o g i n g: 16 t h – 19 t h c e n t u r i e s

Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; The brain of an Horse taken out of the skull, with the Optick, Eye Moving and Pathetick Nerves. The observer observed. This unsettling diagram shows the visual and motor connections of the eyes. The optic nerves (“I”) pass back from the eyeballs to partially cross over at the optic chiasm (“H”), named after the Greek letter chi, χ . “E” is the cut stump of the pituitary gland. “P” to “V” are the six extraocular muscles: two to rotate the eye in each of the three dimensions of space.


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Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; Diaphragm and sublumbar. Notwithstanding the fanciful Valentine hearts on its soles, this horse is little more than a fine-lined version of a similar diagram in Carlo Ruini’s work. The ribby chest is shown separated from the abdomen by the diaphragm, perforated by the three holes through which the aorta, oesophagus, and vena cava pass—each drawn on the opposite side to its true position. The structures labeled “G” are the ilio-psoas muscles, better known in their bovine incarnation as fillet steak.

Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; Heart and lungs. The interconnections between the heart and lungs are so intimate— two large arteries and several veins—that these organs are most neatly excised together, following the same logic by which the organs are today often transplanted together in humans. Looked at with the eyes of faith, the lungs might appear like the wings of a cardiac angel, at a time when the heart was thought to be the seat of the soul.


the horse stripped bare

Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; Limb bones. This diagram of the equine forelimb skeleton is perfectly serviceable as a teaching aid for today’s veterinary students. For example, it clearly illustrates the distinctive features of this animal’s bones: knobbles (“N”) at the top end show where the tendon of the biceps muscle runs through a unique double-furrow; the ulna (“F”) fades to nothingness before it reaches the wrist; and the complexities of the fetlock joint (“N,” “P,” “R,” and “V”) are picked apart and displayed.


This representation of a stillborn foal’s fetal membranes is a tragedy put to good use. In the womb the foal is protected by the inner, allantoamniotic membrane (“H”) and nutrified by placental transfer from the mother to the outer, chorioallantoic membrane (“I”). Two umbilical arteries (“G”) run alongside the bladder (“D”) to exit the navel and convey deoxygenated, waste-bearing blood to be refreshed at the placenta. The oxygenated, nutrient-rich blood then returns to the foal through a single umbilical vein (“F”), which disappears into the liver (“A”).


c u t t i n g a n d c a t a l o g i n g: 16 t h – 19 t h c e n t u r i e s

Andrew Snape (1644–1708), The Anatomy of an Horse: containing an exact and full description of the frame, situation, and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; Foal and membranes.

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Charles Parrocel (1688–1752), Skeleton of a horse, drawn after one at the Academy of Science, undated. Parrocel’s squelette (“skeleton”) finds itself an alien in an incongruously everyday setting: an entire articulated skeleton ambling through a litterstrewn farmyard, past a dilapidated barn and rotting farm cart. Look closer, however, and one can see the gray form of a horse’s corpse (the same horse as the one in the foreground, now deboned?) and, far right, the jaunty tail of the dog that has discovered it.

Unknown, Anatomy of the horse, after drawings of the human anatomy compiled by Mansurf ibn Muhammed Ilyas (18th century); Intestinal system and blood vessels. Probably compiled using earlier images from Egypt, this diagram illustrates the continued tendency of anatomical drawings from the Islamic world toward stylization. The horse is shown in a strangely human squatting position, although this may simply reflect an aerial view of the splayed pose of a supine dissected animal. The intestines are depicted as an unnaturally neat, yellow maze and the blood vessels as a sprawling network with no clear origin or destination.


George Stubbs (1724–1806):

The Anatomy of the Horse, 1766 George Stubbs was a painter and engraver who depicted many subjects, but his main legacy is his images of horses. The naturalism of Stubbs’ horses has its roots in his obsession with anatomy. Indeed, at one stage, he spent almost two years in a barn in Lincolnshire, England, drawing dead horses that he had suspended from the roof by hooks. In the days before refrigeration and preservation, this cannot have been a pleasant experience, but Stubbs meticulously peeled

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the horses apart layer by layer until only their bones were left. At each stage of the process, he created a set of three engravings of the carcass viewed obliquely from the front, obliquely from behind, and directly from the side. The images’ level of detail and accuracy is unparalleled, but the animals retain their dignity throughout: they look as if they are standing resolutely, not passively slung from the rafters. The dissection engravings were published as The Anatomy of the Horse in 1766, but had already led to a surge in Stubbs’ popularity as an artist to such a degree that he was regularly hired by the aristocracy to paint their horses as a means of demonstrating their social standing. Horseracing was

George Stubbs (1724–1806), Anatomy of the horse... 1766; Skeletal head of a horse, key.


george stu bbs: the a natomy of the horse

George Stubbs (1724–1806), Finished study for “The Second Anatomical Table of the Skeleton of the Horse,” 1756–58.



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the sport of the super-rich in eighteenth-century England, and the English/ Arabian thoroughbred became Stubbs’ most frequent model. He had an ability to capture the distinctive demeanors and expressions of individual horses—and the same animal can be spotted appearing again and again in many of his paintings. Stubbs soon diversified into other genres of painting. For example, his 1772 oil paintings of a kangaroo and a dingo—Kongouro from New Holland and Portrait of a Large Dog—were the first Western paintings of Australian mammals and, indeed, the first time most Europeans had ever seen these creatures. However, it is Stubbs’ surge of anatomical-artistic creativity in the early 1760s for which he will be most remembered. george stu bbs: the a natomy of the horse

George Stubbs (1724–1806), Anatomy of the Horse... c.1823, Horse from the side (above) and 1766, Muscles, bones and blood-vessels of a horse: écorché figure (detail) (left).


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George Stubbs (1724–1806), A Lion Attacking a Horse (1762). Having spent many months immersed in the smell and filth of dissection (see page 72), Stubbs was determined to make use of his newfound anatomical knowledge. He made several paintings of horses attacked by lions, presumably because they provided him with an equine subject whose every muscle and sinew was tensed in terror. These images are never gory, and the lion is no more than a dramatic foil for the main actor-victim in these scenes—in some, the predator is drawn noticeably more crudely than the prey, and sometimes almost disappears into its own demonic darkness. The horse is always in the same pose, caught in the moment of numb fear when it first feels its assailant’s teeth cut the skin.

George Stubbs (1724–1806), Whistlejacket (1762). Stubbs produced his best work in the early 1760s, just before The Anatomy of the Horse was published. In my opinion, Whistlejacket is the most striking animal portrait in the entire history of animal art. This is no wild, ferocious beast, but a washed and brushed status symbol with shiny horseshoes as his jewelery—the sports car of an earlier century. The dull olive background, which is entirely disconnected save for the shadows of the horse’s two hindfeet, creates a stark composition and has led to rumors that the animal was originally intended to bear a painted rider and prance in a Palladian landscape. Near-life-sized, Whistlejacket now hangs at the end of a series of rooms in the National Gallery, in London, where all lines of sight seem to converge on him, and whence the piercing glint in his eye can draw the viewer in.



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Philippe-Étienne Lafosse (1738–1820), Cours d’hippiatrique, ou Traité complet de la médecine des chevaux: orné de soixante & cinq planches gravées avec soin; The distribution of ten pairs of nerves leaving the base of the skull. Lafosse was a prominent French veterinarian and published widely on horse management and medicine. His representations of the anatomist’s job can be shockingly realistic. Here an animal is suspended from the ceiling, flayed and eviscerated, the long, hairpin loop of its enormous ascending colon draped on the floor in a pool of blood. Today, during operations to treat colic, the colon is often exteriorized in this way and laid on a sterile surface, so that other, smaller abdominal organs may be accessed.

Philippe-Étienne Lafosse (1738–1820), Cours d’hippiatrique, ou Traité complet de la médecine des chevaux: orné de soixante & cinq planches gravées avec soin; Natural skeleton of a foal. The engravings in Lafosse’s book do not have a unified artistic tone, but instead represent a selection of the anatomical imagery available at the time. In this depiction, the skeleton is suspended from something that resembles a gallows.


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Philippe-Étienne Lafosse (1738–1820), Cours d’hippiatrique, ou Traité complet de la médecine des chevaux: orné de soixante & cinq planches gravées avec soin; Miological horse, side view (suspended). This is another of Lafosse’s images of what he called “hippotomy,” which may be unpalatable to many. The decision to include the cloth shroud used to cover the cadaver is, perhaps, a strange one.

Jeremiah Bridges after Charles Grignion (1721–1810) and Francis Sartorius (1734–1804), Muscles of the Horse, 1772. At first sight, Bridges’ engraving appears to be a meticulous representation of the musculature of the horse, but closer inspection provides less useful information than many diagrams that had come before it. The large muscles converging from the head, neck, and back onto the forelimb are vaguely described, and the tendons of the lower limbs are incomplete and confusing.


Like a glance at the chopping board of un boucher chevalin (“a horse butcher”), this image takes the viewer on a tour of the less appetizing parts of a horse. Clockwise from top left: skull and brain, two brains, liver, right kidney, stomach, eyeballs, bladder, left kidney, spleen.


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Philippe-Étienne Lafosse, plate Philippe-Étienne Lafosse (1738–1820), Cours d’hippiatrique, ou Traité complet de la médecine des chevaux: orné de soixante & cinq planches gravées avec soin; Assorted visceral organs.

the horse stripped bare

[William Youatt (1776–1847) and Isambard Kingdom Brunel (1906–1859)], The Horse, with a Treatise on Draught, 1831; Section of the head. In the nineteenth century, cross sections of body parts became a common way to represent anatomical structure, a trend that faded in the twentieth century, only to be resurrected in the twenty-first when modern diagnostic imaging techniques started to yield what are, effectively, slices through patients. This diagram shows how a horse’s muzzle is dominated by the nasal cavities (“s” and “r”) and the tongue (“y”). Once its voicebox (“3”), throat (“9”), and neck muscles (“l”) are included, little space is left for the brain (“m” and “n”).


Charles Landseer was the older brother of the more famous painter (especially of animals), Edwin Landseer, but he has left us preliminary sketches for his own paintings. The musculature of the horse’s head has long fascinated artists, as it is the cause of the facial expressions they strive to depict. Emphasized here are the elements of the facial musculature that are the most extravagantly developed in the horse: the muscles that flare the nostrils and the muscles that close the eye—even a strong man cannot open a horse’s eye if it does not wish him to.

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Unknown, The Ages of the Horse, 1848.

Although many practitioners debate the accuracy of estimating horses’ ages according to the wear evident on their teeth, learning this skill remains a rite of passage for anyone working with horses. As the central image suggests, half the battle is pulling the wriggling tongue out of the way to expose the occlusal surfaces of the lower incisor teeth. These erupt from the gums, with a set of identifiable internal structures running throughout their length, and wear away in a predictable order.


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Charles Landseer (1799–1879), The muscles of the head of a horse, c.1815.

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Benjamin Waterhouse Hawkins (1807–1894), A Comparative View of the Human and Animal Frame, 1860. Hawkins was a sculptor and artist who specialized in zoological and paleontological subjects—he is best known for creating the dinosaur statues still on display in Crystal Palace Park, in London, for the Great Exhibition of 1851. In this image he compares the bones of man and horse, in a tradition dating back to Leonardo da Vinci, but which in this composition has something of the apocalyptic about it.

J. A. McBride and T. Walton Mayer, Anatomical Outlines of the Horse, 1878; Digestive and vascular systems of the horse. The images in Mayer’s book start to take on a modern appearance, their schematic nature reflecting an attempt to clarify the complexity of nature rather than to hide gaps in the author’s knowledge. The lower diagram of the cardiovascular system, for example, shares the clarity and abstraction of twentieth-century wiring diagrams of London Underground maps.


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John T. Share-Jones, The surgical anatomy of the horse, 1906; Section of the foot and caudo-lateral view of the pes. Improvements in anatomy and anesthesia paved the way for veterinarians to undertake invasive, complex, and often successful surgeries. In the twentieth century, equine anatomy began to be presented in styles that suggested that the viewer might one day encounter the structures in live animals as well as dead ones. Thus, equine anatomy became a practical roadmap, rather than a fanciful medieval atlas.

Henry Fairfield Osborn, Equidae of the Oligocene, Miocene, and Pliocene of North America, 1918; Upper teeth of Mesohippus. Horses have become biologists’ favorite explanatory paradigm for evolution because they have the most complete fossil record of any animal. Multiple species and subspecies appear in the strata, as if to demonstrate the incremental gradations of dental and locomotor change that brought this branch of the animal kingdom to its modern state. Here, for example, are presented the increasingly convoluted teeth of Mesohippus, the “middle horse.”


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W. Ellenberger and Hermann Baum, Handbuch der Vergleichenden Anatomie der Haustiere, 1912; Vertical view through the center of the front toe of the horse. Some pieces of anatomical knowledge are more directly applicable than others. To a clinician, almost all the structures in the foot are a needle’s prick or a scalpel’s slice away, whereas any earlytwentieth-century vet seeing the arteries wriggling through a horse’s head had probably reached a surgical point of no return.


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O. Charnock Bradley, The Topographical Anatomy of the Thorax and Abdomen of the Horse, 1922; The heart and great blood vessels seen from the right (dissected) and left side (intact). These twentieth-century diagrams return us to a place not too far from Carlo Ruini’s sixteenth-century heart on page 48, albeit with the addition of a few more centuries of scientific confidence. Arteries taking blood from the heart are shown in red, and veins returning blood to the heart are in blue.


chapter 3

bewildering variety A pictorial menagerie: 16th–19th centuries

Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich... 1918; Anatomy of a domestic dove (1762). Colored etching.

The sheer variety of animal life has proved irresistible to the artist.

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efore the sixteenth century, the most common pictorial depictions of animals were in bestiaries—densely illustrated catalogs of animals familiar, exotic, and fabricated. Rather than a figure legend containing a scientific anecdote, most were accompanied by a short moral parable, explaining how God’s creations—every beast of the earth, and every fowl of the air, and every thing that creepeth upon the earth, as Genesis would have it—existed to make clear His design for us. Because of this spiritual slant, it was not important for the creatures in bestiaries to actually exist, and there was also no reason to display their insides. One might think that the Hand that guided the creation of beasts would be best evidenced by demonstrating their internal complexity, but the medieval Christian mindset did not agree. However, attitudes changed in the late Renaissance, with the production of a veritable menagerie of images depicting the skeletons and internal organs of vertebrate animals—a scientific and artistic urge that persisted as late as the nineteenth century. Relatively quickly, the sage and instructive stories of the bestiaries were replaced by images created by people who believed creation had a deeper story to tell. Some of this movement’s earliest works show pregnant sharks and inquisitive apes, or juxtapose the differing placentas or bony architectures of diverse species. In one particularly striking picture (see page 106) from Pierre Belon’s 1555 L’Histoire de la Nature des Oyseaux, the skeletons of human and bird appear suspended by their skulls as if from a necklace, and are explicitly compared. The anatomists and artists of the time were fascinated by the creeping realization that each creature is not, it turns out, created anew, an entirely novel design formed without reference to any other. Instead, it was revealed that animal life is not a cataloger’s inventory of discrete unrelated forms, but rather consists of variations on a basic theme. The bird and the man are shown as essentially the same, human separated from fowl by a few minor skeletal tweaks and distortions. Move beyond the backboned animals, however, and this anatomical commonality breaks down. Among the invertebrates one encounters worms, molluscs, jellyfish, and insects with body plans 94

so fundamentally different to our own that there seems little point in comparing them. In contrast, the internal similarity of backboned animals, which we now know reflects their common evolutionary heritage, was already so obvious in the sixteenth century that it seems to have obsessed anatomical artists. The idea that the same structures could be modified and adapted to serve the purposes of a human, a bat, a dove, a serpent, and a halibut spoke of far more profound evidence of the Creator’s plan than the inventoried animal sermons of the Middle Ages. The vertebrate animals fall into seven groups, although even this basic classification remains contentious. The group most distantly related to man is the jawless fish, comprising the hagfishes (which were little known to early scientists because they lived in the ocean depths) and the lampreys, more familiar (and edible) creatures that are harvested from rivers and seas across the world. At close quarters lampreys are alarming-looking, eel-like creatures, mainly because of their varied specializations for an unpleasant parasitic lifestyle—they will rear their ugly head only once in this chapter (see page 149). More closely related to us are the cartilaginous fish—the sharks, rays, and chimaerae. To us they appear archaic, relics of a past age, but, 95

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Thomas Bartholin (1660–1680), De Unicornu Observationes Novae, 1645.

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W. Schimkewitsch, Lehrbuch der Vergleichenden Anatomie der Wirbeltiere, 1921; Shark’s head, after Steno.

of course, this sense of evolutionary time is a relatively modern idea. Before the nineteenth century they must simply have seemed strange, ferocious dwellers of an unknown world beneath the waves. However, natural philosophers had for some time realized there was something disconcertingly familiar about sharks: apart from mammals, sharks are the animals that most often gestate internally and give birth to their precious offspring live. As a result, early anatomical depictions of cartilaginous fish focused not only on the savagery of their jaws, as in Steno’s Lamia Piscis or “Great White Shark” (see page 116), but also the strange internal life of the cosseted shark embryo. The third group of vertebrates, the bony fishes, while more closely related to us than their gristly cousins, are in some ways a more exotic group. This more diverse and distorted category includes eels, flatfish, lungfish, flying fish, and a huge range of anatomical weirdness trawled from the abyss. Although we now know they are just one evolutionary step away from land vertebrates, they still seem very different, and centuries of their anatomical depiction represent a protracted attempt to determine the hidden equivalences between fish and quadrupeds. In amphibians, artists were surrounded by the miraculous and bizarre in their everyday lives. Frogs, toads, newts, and salamanders are everywhere, but their metamorphosis from tadpole to adult must have seemed as remarkable to late-Renaissance and Enlightenment 96

Gerardus Leonardus Blasius (1626–1692), Anatome animalium, terrestrium variorum, volatilium, aquatilium, serpentum, insectorum, ovorumque, structuram naturalem exveterum, recentiorum, propriisque observationibus proponens, 1681; Porpoise & sea fox [thrasher shark].


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thinkers as it does today. Also, as we shall see, the humble and ubiquitous frog represents one of the most dramatically distorted and specialized versions of the vertebrate body plan. And, once this anatomical format evolved, it seems to have been so successful that it has hardly changed since, radiating out to fill varied ecological niches around the world. Recent studies have suggested that the fifth group of vertebrates, the reptiles—which were once thought to be an incoherent rag-bag of unrelated species—is, in fact, a neat classificatory group of creatures with a common ancestry and shared anatomical features. Turtles, crocodiles, lizards, and snakes share many physiological adaptations for living life away from water, and we now believe that, in spite of their starkly differing anatomies, they are, in fact, all related. Today, we consider them alongside dinosaurs and huge, extinct flying and aquatic forms, but for most of history those species were unknown or unrecognized. Reptiles, like the gristly fish, have something otherworldly and ancient about them, which seems to fascinate anatomical artists, and, of course, the biblical representation of

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Bernhardus Siegfried Albinus (1697–1770), Tabulae Sceleti et Musculorum Corporis Humani, 1747; Male skeleton with a rhinoceros.

the serpent added greatly to their artistic allure. Birds, like frogs, are a group of vertebrates that hit upon a successful anatomical design and then stuck to it. Birds are numerous, varied, and ecologically diverse, yet, to be honest, they all look very similar on the inside. Students who dissect a chicken are well prepared should they ever have to deal with an emu, eagle, or penguin. For the same reason, birds represent good value for the artist: a little knowledge goes a long way. We now know that birds are actually the diminutive survivors of a group of feathered bipedal dinosaurs, a story to which I will return 98

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866–68; Dermal muscles of a hedgehog.


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in later chapters. However, until recently, they must have seemed like separate, spontaneous, godly inventions—created, as we are told in the biblical story, on the fifth day, discrete from most other creatures. Finally, come the mammals. Philosophers have long realized that the ability to nourish offspring with milk unifies this otherwise exceptionally varied assemblage of beasts. Most are furry, livebearing, four-footed creatures with expressive faces, but whales, bats, armadillos, seals, platypuses, and people stray significantly from anything that might be called a “standard” mammalian body plan. The twenty-or-so classificatory subgroups of mammals offer a wonderful resource for artists, and the fact that they constitute the majority of domestic animals means some of them are always at hand to pose as artists’ models. Also, the suspicious similarity of some mammals to humans must have raised worrying questions in the Christian mind about man’s special place in the scheme of things. And so the artistic comparisons of animal anatomy began. From the mid-sixteenth century onward, animal anatomical art might be regarded as the forerunner of today’s television nature documentaries—a leisure pastime justified by a veneer of nonfrivolous intellectual respectability. These images are fascinating, even if one has no interest in learning more about their spiritual or scientific meaning. The internal structure of a toucan’s beak (see page 153) is not important for most of us, but it is beautiful and surprising nonetheless.

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Giovanni Maria Lancisi (1654–1720) after Bartolomeo Eustachi (1500/14–1574), from Tabulae anatomicae clarissimi viri... 1722; Dog placenta (fig. VII) and sheep placenta (fig. X). Early comparative biologists developed something of an obsession with reproductive biology, and hidden behind the variety of animal forms lies an even greater diversity of reproductive strategies. For example, anatomists as far back as Eustachi, a contemporary of Vesalius, noticed that the placenta is the most variable of all organs. Here, the puppy’s placenta wraps around it like a girdle, while the lamb’s placenta is divided up into multiple “mini-placentas,” just like those erroneously inserted by Leonardo da Vinci into his diagram of a human fetus (see page 28).

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Anatomists have long been intrigued by the fact that, other than mammals, the backboned animals that most often give birth to live young are sharks. That some species of these otherwise frightening creatures gestate their young in this way seems surprising, although we now know that it simply reflects the relative advantages and disadvantages of live birth and egg-laying. The upper image shows a viviparous female dissected to reveal an unrealistically oversized pup still attached by its umbilical cord. The animal in the lower image lies alongside an enlarged depiction of its egg, the “mermaid’s purse” that is typical of oviparous sharks and rays.


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Guillaume Rondelet (1507–1566), Libri de piscibus marinis, in quibus verae piscium effigies expressae sunt... 1554–55; Viviparous and oviparous sharks.

Volcher Coiter (1534–1576):

De Partibus Similaribus Humani Corporis, 1575 Born in Groeningen, in the Low Countries, Volcher Coiter was a physiciananatomist who studied and practiced extensively in northern Italy during its sixteenth-century flowering of anatomy. Coiter’s interests were extremely wide: in human anatomy he wrote detailed descriptions of the structure of the inner ear, bone development, and the nervous system, and he also studied the development of chicks within the egg. He was fascinated by comparative anatomy, and many of his most famous images compare the skeletal structures of different

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vertebrates—indeed, it was he who started the science of “comparative osteology” in earnest. Coiter was unusual in that he drew his own diagrams and made his own engravings, and was cited as artist in both his own works and the works of others. His images are detailed and accurate, and they also

Volcher Coiter (1534–1576), Externarum et Internarum Principalium Humani Corporis Partium Tabulae... 1573; Primate skeleton.


volcher coiter: de pa rt i bus si m i l a r i bus h u m a n i cor por i s

Volcher Coiter (1534–1576), from Gabriele Falloppio (1523–1562), Lectiones... de partibus similaribus humani corporis, 1575; Bird skeletons. Birds are excellent candidates for comparative anatomy. Indeed, Coiter was among the first biologists to develop a taxonomic “key”—a decision tree to allow others to identify species of birds.


possess compositional features unique to their creator. Coiter was unusually parsimonious with the space available in his engravings, and often the skeletons of many animals jostle for space in a single composite image— sometimes sideways, sometimes upside down, and occasionally crammed into the irregular gaps between other creatures’ bones. Coiter had an active but short life: on his return to the north he became the city physician of Nuremberg, before becoming involved in the French Wars of Religion, as surgeon to a palatine prince. It was during those wars that he died. His main legacy is the clear, fine lines of his beautiful images, and the idea that animal species can be classified according to their structure. He was also among the first scientists to explicitly discuss the

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remarkable anatomical similarities between humans and other primates.

Volcher Coiter (1534–1576), from Gabriele Falloppio (1523–1562), Lectiones ... de partibus similaribus humani corporis, 1575; Mixed skeletons (a marten, rabbit, piglet, and parrot). Four creatures compete for space in this maelstrom of bones, but the detail in this image goes beyond anything drawn before. A perfect tongue and windpipe are appended to the parrot; the piglet has its unique snout-bone in place; the marten displays his os penis (bony penis) dangling below his spine; and the hunched-up rabbit’s bones are perfectly oriented.


volcher coiter: de pa rt i bus si m i l a r i bus h u m a n i cor por i s

Volcher Coiter (1534–1576), from Gabriele Falloppio (1523–1562), Lectiones ... de partibus similaribus humani corporis, 1575; Mixed skeletons (tortoise and a deformed chick). The anatomy of turtles and tortoises has always fascinated anatomists, and continues to do so. Above, Coiter correctly illustrates the ribs of the turtle as being part of the shell itself, closely applied to the horny upper carapace, whereas the underside of the shell, the boneless plastron, is shown separately from the skeleton. On the right is the skeleton of a pulli monstrosi (“deformed chick”), with two super-numerary legs protruding from its rear.


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Pierre Belon (1517–1564), L’Histoire de la Nature des oyseaux, 1555; The skeleton of a man compared with that of a bird. Belon’s bird and man compared is one of the most charming curios in all of anatomical art. Apart from a few vertebrae and digits here and there, there is an almost direct correspondence between the bones of the two vertebrates. In fact, the only error in these diagrams is the result of a rare inequivalence between the two: Belon suggests that the human clavicle or collar bone (“L”) is equivalent to the coracoid bone of birds, when, in fact, the coracoid does not exist as a separate entity in humans. Instead, it is the wishbone (denoted by an irregular squiggle in the image) that forms from two fused clavicles. These two images appeared on facing pages in Belon’s book, as an artistic expression of just how much humans share with the beasts.

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Jean Germain (fl. 1625–37), Breve e Sustantiale Trattato Intorno alle Figure Anathomiche delli Piü Principali Animali Terestri, Aquatili, et Volatili, 1625; Spoonbill. La paletta in Italian (Platalea in Latin), this is almost certainly a spoonbill, a wading bird that is widespread in Europe and around the rest of the world. Spoonbills spend much of their time wading through shallow water, waiting to snap their sensitive, spatulate bills onto any tiny prey that brush past. This individual has been transplanted to a drier, upland region that is entirely unsuited to its way of life.


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Hieronymus Fabricius, Dissection of a live-bearing shark, De Formato Foetu, Padua: L. Pasquatus (1604). Fabricius, or Girolamo Fabrizio d’Acquapendente, was a pioneering Italian anatomist who was professor of surgery at Padua. This closely observed study continues, with greater clarity, his predecessors’ fascination with viviparous sharks—showing bulbous, lobulated ovaries as well as pups developing in the uterus. In contrast, bony fish often have diminutive, transient, or even absent reproductive tracts, and merely squirt their eggs and sperm directly from the body cavity into the surrounding water.

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This image demonstrates the distinctive features of the fetal membranes and placenta of ruminants. Whereas a human child is bathed in a single cavity of amniotic fluid, the lamb lies within an inner amniotic cavity (the oval region around the fetus), which is in turn partially surrounded by an elongated allantoic cavity (the large, C-shaped sac around the periphery in this image). Between the two is the cotyledonary placenta, which, unlike the single disc of the human placenta, is divided into multiple regions of materno-fetal attachment.

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Hieronymus Fabricius (Girolamo Fabrizio, 1537–1619), De Formato Foetu. [De brutorum loquela. De venarum ostiolis. De locutione et eius instrumentis liber], 1604; Sheep fetus and placenta.

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Giulio Cesare Casseri (Julius Casserius; c.1552–1616), De Vocis Auditusque Organis Historia Anatomica, 1601; Pike ears. Yet another anatomist of the Paduan school, Casseri wrote extensively on many areas of anatomy, but his volume on the organs of vocalization and hearing is of the most interest to comparative anatomists. Here the brains and sensory organs of some pike are being progressively investigated. Fish lack the outer ear tube, middle ear cavity, and the cochlear region of the inner ear that are present in land vertebrates, but Casseri was still able to identify the nervous links between the brain and the organ of hearing.

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Giulio Cesare Casseri (Julius Casserius; c.1552–1616), De Vocis Auditusque Organis Historia Anatomica, 1601; Mammal ears. Unlike fish, land vertebrates must funnel sound into an outer ear canal before it can be transmitted to the biological microphone of the inner ear. Mammals, in particular, have developed a wide variety of external ear flaps to collect sound efficiently, and here Casseri attempts to find equivalences between the funnel-like ears of pigs, deer, sheep, cats, and mice, and the irregular flattened spiral of the human ear.


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Giulio Cesare Casseri (Julius Casserius; c.1552–1616), De Vocis Auditusque Organis Historia Anatomica, 1601; Ox throat. The underside of the mammalian head can seem a confusing tangle of muscles, but this reflects the several complex activities that take place there. When alive, this ox grazed, chewed, breathed, swallowed, coughed, and lowed, and perhaps half of all the muscles involved in those processes are visible in this diagram. Finally, the tableau is garnished with decorative pendants of fragrant herbs and fruit.

Giulio Cesare Casseri (Julius Casserius; c.1552–1616), De Vocis Auditusque Organis Historia Anatomica, 1601; Cat, and rabbit larynx. Casseri was just as interested in the organs that generate sound as in those that detect it, and this composition details the anatomy of the voicebox (larynx) and windpipe (trachea), and the cartilages, bones, and muscles that move them.

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Marcello Malpighi (1628–94), De pulmonibus observationes anatomicae, 1661; Lungs of a frog. Most frogs’ lungs have a relatively simple, balloon-like structure, and this simplicity allowed Malpighi, professor at Bologna and then Pisa, to discover the capillaries that convey blood from the arteries to the veins. In this image, the capillaries are depicted as a mesh of tiny lines that connect the arteries and veins once they have become too small to be observed with the naked eye. Thus, in discerning the capillaries, Malpighi effectively completed the theory of the circulation of the blood, first proposed by the Englishman William Harvey.

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Willis wrote two major works on the brain, one on anatomy and one on pathology, and each was more comprehensive and detailed than anything published before. Much of his work focused on the human brain, but he was also interested in comparative neurology, as in this image based on a drawing by Sir Christopher Wren, the architect of St. Paul’s Cathedral in London. The brain is the most structurally diverse organ in the body, with a wide variety of fleshy geography for anatomists to map, yet its configuration differs little between species. This sheep’s brain is notable, however, for its olfactory bulbs (“D”), which are far larger than would be seen in a human due to our species’ poor sense of smell.


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Thomas Willis (1621–1675), Cerebri Anatome: Cui Accessit Nervorum Descriptio et Usus, 1664; Sheep’s brain (right) with the circle of Willis (the anastomotic arterial circle).

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Nicolaus Steno (1638–1686), Elementorum Myologiae Specimen, Seu Musculi Descriptio Geometrica, 1667; Head of a sea fish [shark]. Often copied, this is Steno’s original engraving of a shark’s head, probably a great white shark—the original scientific name for the great white was “Lamia,” from the flesh- and blood-eating demons of Greek mythology. Here decapitated, the shark looks like a ferocious gargoyle, with its endless supply of teeth cascading over its lips, although the tongue is strangely fleshy and mammalian-looking.

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Samuel Collins (1617–1685), A Systeme of Anatomy, treating of the body of man, beasts, birds, fish, insects, and plants, 1685; Head of a spaniel bitch opened, and the head of a cat opened. It is the choice of a spaniel that may bring this image too close to home for some observers—the use of dogs and cats in anatomy and scientific research has often been viewed as less acceptable than the use of hoofed animals and rodents. Rather than merely showing a superficial view, these images depict dissections beyond the convoluted cerebral hemispheres, down to the brainstem, the internal fluid-filled ventricles, and the hippocampus (named after its resemblance to a “seahorse”), which is now known to play an important role in memory.


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Edward Tyson (1650–1708), Phocaena, or, The Anatomy of a Porpess, dissected at Gresham Colledge: with a praeliminary discourse concerning anatomy, and a natural history of animals, 1680. Tyson researched widely in animal anatomy, studying opossums, lions, ostriches, and snakes, but this image recounts his studies of cetacean anatomy. He noted how the internal organs resemble those of land mammals, especially the convoluted nature of the brain. He is credited with “discovering” that cetaceans are mammals, though Conrad Gessner’s image (see pages 42–43) suggests that this had been known for some time.

Edward Tyson (1650–1708), Orang-outang, Sive Homo Sylvestris: or, The anatomy of a pygmie compared with that of a monkey, an ape, and a man... 1699; Chimpanzee skeleton. The title of Tyson’s work is self-explanatory, except that he actually dissected a young chimpanzee rather than an orang-utan. Despite noting a long list of similarities between the chimp and humans, he still decided that apes were animals and humans were not. However, he did propose that the apes— chimpanzees, gorillas, and orang-utan—represented a distinct group, separate from the monkeys. In this image, the ape is drawn in an unnaturally humanlike, erect stance, with his palms turned forward, probably to simplify the anatomy of the forearm rather than as a gesture of supplication. Despite the creature’s forest origins, he is drawn on the arid, rubble-strewn ground that was so popular with Renaissance anatomical artists.

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Nehemiah Grew, Musaeum Regalis Societatis. Or a catalogue & description of the natural and artificial rarities belonging to the Royal Society... Whereunto is subjoyned the comparative anatomy of stomachs and guts, 1694; Human urethra stone, armadillo, baby wild pig, tusk of a wild boar, head of a hippopotamus. Grew was primarily a plant biologist, but he did publish one volume of striking images of animal anatomy, which included this unusual assortment of objects—why a human bladder stone is included is not clear. The hippopotamus (or “behemoth”) has an especially grotesque majesty about it—its baroquely sculpted jaws and crushing teeth being perfectly adapted for a life of chewing through tons of water plants.


Grew developed a strangely stylized way of depicting the gastrointestinal tract of different vertebrates, which bears an uncanny resemblance to the pipelike, tubular forms in the works of the twentieth-century Cubist painter Fernand Léger. Fish guts are often relatively short and simple, but those of the salmon (top left) are more ornate than most, due to a cluster of blind-ending tubes that bud off the lower end of the stomach— the “pyloric caeca,” depicted here as a forest of sideways-pointing spikes.


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Nehemiah Grew, Musaeum Regalis Societatis. Or a catalogue & description of the natural and artificial rarities belonging to the Royal Society... Whereunto is subjoyned the comparative anatomy of stomachs and guts, 1694; Stomach 7 guts of a salmon, and a plaice.

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Gerardus Leonardus Blasius (1626–1692), Anatome animalium, terrestrium variorum, volatilium, aquatilium, serpentum, insectorum, ovorumque, structuram naturalem exveterum, recentiorum, propriisque observationibus proponens, 1681; A beaver. In his Anatome Animalium, the Amsterdam physician-anatomist Blasius published a series of tableaux, illustrating various creatures, along with inset “manuscripts” illustrating the structures he considered to be the most interesting. In the case of this beaver, these included its genitalia and accessory sex glands, and its forepaws.


Perhaps the most unusual and specialized of all reptiles, the chameleon is a favorite of artists. These images focus on its arboreal habits—showing its prehensile tail and opposable toes.


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Gerardus Leonardus Blasius (1626–1692), Anatome Animalium... 1681; Chameleons.

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Gerardus Leonardus Blasius (1626–1692), Anatome Animalium... 1681; A dromedary. Among the body parts illustrated alongside this dromedary are its abrasive tongue, eyeball with attached muscles, udder, and complex stomach.


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Gerardus Leonardus Blasius (1626–1692), Anatome Animalium... 1681; A monkey. The ape skeleton in this composition is an almost perfect mirror image of that drawn by Volcher Coiter more than a century earlier (see page 102).



Georges-Louis Leclerc, Comte de Buffon (1707–1788), Skeleton of a stag, Histoire Naturelle, Générale et Particulière, avec la déscription du Cabinet du Roi, 1749–1766; Skeleton of a stag. Leclerc was perhaps the most influential, and certainly one of the most productive, naturalists of the eighteenth century. In the 36 volumes of his Histoire Naturelle, he not only illustrated an enormous variety of animals, but also hinted at many of the concepts that future evolutionary biologists would tackle—heredity, animal change, the nature of the species. Here, a skeleton of a stag stands, statue-like, on a plinth.

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Born in Leicestershire, Cheselden was an immensely influential London surgeon who pioneered several ground-breaking clinical procedures. He was also an anatomist, whose Osteographia was the most detailed survey of the human skeleton to date, but which also touched on comparative osteology. Here, the turtle makes another appearance. Unlike other animals, turtles’ limbs take their origins from inside the rib cage, and this image captures the large, bony levers and pivots that are required to wield

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the flippers which drive these powerful reptiles through the water.

Frederick Ruysch (1638–1731), Thesaurus animalium primus... Het eerste cabinet der Dieren, 1710; Zoological preparations (Surinam toads). Ruysch was a Dutch plant and animal anatomist who is mainly remembered for the dioramas he created using dried and preserved specimens. He chose this species of toad because of its strange habit of gestating its eggs within the skin of its own back. In fact, his zoological compositions, such as these potted Surinam toads, topped with shells, snakes, coral, and leaping froglets, were among his most orthodox. Infamously, many of his dioramas include skeletons of miscarried human fetuses arranged in a variety of theatrical poses.


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William Cheselden (1688–1752), Osteographia or The anatomy of the bones, 1733; Water tortoise.

Andrew Snape (1644–1708), The Anatomy of an Horse : containing an exact and full description of the frame, situation and connexion of all his parts, (with their actions and uses) exprest in forty nine copper-plates... 1683; Chick development. The developing chick was such an inexpensive trope of the anatomist that it somehow made its way into volumes to which it had no apparent nourished by the large, yellow yolk sac, which gradually ejects its contents into the embryo’s intestines. The yolk sac dangles beneath the chick’s belly, leaving a large window through which the internal organs may be glimpsed.

William Cheselden (1688–1752), Osteographia or The anatomy of the bones, 1733; Sparrow and bat. This image draws attention to the fundamental skeletal differences between the two extant groups of flying vertebrate.

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relevance, including Snape’s Anatomy of an Horse. In the egg, chicks are

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August Johann RÖsel von Rosenhof, Skeleton of a frog, Historia Naturalis Ranarum Nostratium in qua Omnes Earum Proprietates, Nuremberg: J.J. Fleischmann (1758). The body plan of the frogs and toads is one of the most specialized of all vertebrates, yet once it evolved, it seems to have changed little. Eight front toes, ten back toes, greatly lengthened hind limbs and pelvis, reduced vertebrae and ribs, and a delicate, simplified skull—all are here, brought into warm relief by the gentle shadows they cast.

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Mettel’s Meer Drachen (“Sea Dragon”) seems to have been drawn from a real dried specimen, rather than copied from Steno’s picture (see page 116), like so many others. The jaws and teeth are, if anything, more ferocious, and a cold, dead eye leers ominously at the viewer. The desiccated skin has contracted into a shriveled mask, making the animal’s outline more angular and aggressive.

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Nicolaus Mettel (1739–1772 fl.), Head of a shark, with dried flesh and an eye adhering to the upper part, 1770s.

Georges Cuvier (1769–1832): Le Règne Animal, 1817

Georges Cuvier, often credited with being the initiator of modern paleontology, was also remarkable for his ability to survive the political upheavals that tore France apart during his lifetime. Born in the Jura Mountains, and educated in Stuttgart, Cuvier spent most of the rest of his life in France, always in the right place at the right time, to avoid the repercussions of both the French Revolution and the fall of Napoleon Bonaparte. Cuvier’s knowledge of animal structure was second to none, and he

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was appointed to the Chair of Animal Anatomy at the Musée National d’Histoire Naturelle in Paris. For centuries, many scientists had suspected that fossils were petrified remains of long-dead creatures, but none had had Cuvier’s ability to reconstruct fossilized animals’ bodies from the fragmentary and damaged vestiges that remained—the essential skill of the modern paleontologist. Cuvier propounded his theory that many fossilized species are actually now extinct, and that today’s animals represent just a fraction of those with which God populated the Earth. Many could not countenance the idea that the Lord would create perfect creatures only to destroy them, and this

Georges Cuvier, Baron (1769–1832), Le règne animal distribué d'après son organisation: pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparée / par M. le cher. Cuvier; avec figures, dessinées d'après nature, 1817; Head of a cod.

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Georges Cuvier, Baron (1769–1832), Le règne animal... 1817; Anomalous mammals. The upper three images are of the skull of an aye-aye, an unusual lemur with protruding teeth and long, spindly fingers for extracting insects from tree trunks. The lower images are of a wombat, a stocky and endearing, yet anatomically unremarkable, marsupial.

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led to the idea that the dark hearts of the continents were still inhabited by the strange animals dug from the ground in Europe. Cuvier was adamant, however, that the Earth is immensely old, and that repeated cycles of catastrophe have shaped its inhabitants—an idea that has come back into vogue in recent decades. Although he was often cited by those old-earthers who were later to espouse the idea of evolution, Cuvier himself did not believe that animal species change over time. Rather than any religious conviction, his belief was fueled by the idea that all the parts of an individual animal are each perfectly adapted to their role, and that if one of them changes, then all those parts will cease to work together harmoniously. Thus, he contended, if the anatomy of a fossil elephant and a modern elephant differ, then they

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are separate species, and have been so ever since they were created.

Georges Cuvier, Baron (1769–1832), Le règne animal... 1817; Head of a great Javanese python, and head of a rattlesnake. The skulls of snakes are anatomically bizarre, as snakes appear to have gone through a phase in their evolutionary history when many components of the head were “lost.” The upper three diagrams are of a nonvenomous, constricting species, and the first image shows the double arcades of teeth in the upper jaw. The lower row of images demonstrates how venomous species’ skulls are reduced to little more than a set of levers and pushrods to bare the fangs and widen the gape.

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Georges Cuvier, Baron (1769–1832), Le règne animal... 1817; Anomalous snakes. Cuvier correctly identified the species illustrated in these three rows of images as not being snakes at all. The upper row is three views of the skull of a caecilian, a limbless scaly amphibian, and below are the skulls of an amphisbaenid and a glass lizard, both limbless lizards.

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Ludwig Heinrich Bojanus (1776–1827), Anatome Testudinis Europaeae, 1819–21; Internal organs of a European tortoise. This image was one of the first to make clear the strange arrangement of the internal organs of turtles and tortoises. The upper half of the shell is filled with lungs, whereas the lower part is depicted as a writhing mass of intestinal coils and bubbling ovarian follicles—tortoises lay large, hard-shelled, yolky eggs, and can store an entire clutch of them inside the body. Throughout the book, Bojanus’ images are exceptional in their clarity, detail, and lightness.

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H. [Henri] Milne-Edwards (1800– 1885), Cours élémentaire d'histoire naturelle, Zoologie, 1852; Skeleton of a cetacean (Dugong) (above) and Circulatory system in a lizard (right). For some time, the dugong, or sea-cow, was thought to be a close relative of whales and dolphins, although modern anatomical and molecular studies have indicated a close, if unexpected, affiliation with elephants and hyraxes.

Roger Fenton (1819–1869), Dinornis elephantopus, 1860; Heavy-footed moa. Although photography does not provide the same scope for creative interpretation as drawing, its advent permitted new ways of seeing the world. In its hastily-arranged studio setting, it is tempting to see this heavy-footed moa as looking slightly embarrassed, caught in a state of ultimate undress.


In this composition, the human skeleton looks as if it is begging or praying, perhaps because of the attentions of the predator immediately behind it. Hawkins’ montages certainly draw in the eye of the viewer, but it is less clear to what extent they actually advanced the science of comparative anatomy.

James Erxleben (1830–40), Transactions of the Zoological Society of London; Little Bush Moa (Dinornis parvus). The other half of this odd couple (the first shown on page 141) is a little bush moa. Up to 12 feet in height, the moas of New Zealand were among the largest birds ever to exist. Unlike modern ostriches and emus, they lacked wings entirely, and their skeletons were dominated by their robust legs. No moa species survived the arrival of man in New Zealand.


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Benjamin Waterhouse Hawkins (1807–1894), A Comparative View of the Human and Animal Frame, 1860; Skeleton of a man, with those of a male and female lion.

Alfred Brehm (1829–1884): Tierleben, 1864–1869

Son of a Thuringian ornithologist, Brehm was raised in a house filled with literally thousands of stuffed birds. Perhaps this formative environment explains why he was eventually drawn away from his early training as an architect to study zoology. The epitome of the nineteenth-century traveler-savant, Brehm voyaged to Spain, Scandinavia, Egypt, Sudan, and Abyssinia, and was soon in demand by European publishers for his floridly descriptive travelogs. Brehm was commissioned to write his own multi-volume journey

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through the animal kingdom, the Tierleben (“Life of Animals”), and a cadre

Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich, 1918; Anatomy of a domestic dove. Untimely plucked from the dovecote, this dramatic single image captures many elements of avian anatomy. The gray, grain-filled crop bulges from the throat; the shiny red heart hangs like a jewel at the animal’s core; and the finely stippled, pink lungs on either side nestle against the glassy-looking respiratory air sacs (which would actually have been destroyed during the dissection process). An artistic device used elsewhere in the Tierleben, the tip of the beak protrudes across the boundary of the image to give the impression the bird is escaping from the page.


Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich, 1918; Skeleton of a domestic pigeon.

of professional artists was assembled to illustrate the new work. It is probably for this reason that the huge, multi-volume series lacks the unity of artistic purpose seen in shorter works. However, the series was certainly appealing to a wide audience, and some of the images are among the most vibrant anatomical depictions ever produced. Although the science of the Tierleben now sounds outdated, the art remains as vibrant as ever. Many of the figures depict live animals in the wild, including the troglodytic but tellingly titled “man-shaped apes.” Yet the dissections and skeletons have been more influential. Indeed, Charles Darwin commented that the book’s illustrations were the finest he had seen. The Tierleben made Brehm famous, and he became director of the Hamburg Zoological Gardens and founded the Berlin Aquarium. Although he died in his fifties, he is remembered for casting an educated

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traveler’s eye over a new world—newly mapped, accessible, and perhaps even understandable. From a cultural standpoint, he was the forerunner of today’s celebrity wildlife documentary makers.

Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich, 1918; Anatomy of a rattlesnake. Packaging a vertebrate’s internal organs into a narrow cylindrical body is a challenge, but it can be done. Working from head to tail, this admirably clear image shows the trachea (windpipe), the tiny pink thyroid gland, the heart, the pale lungs, the dark liver, the veiny stomach, the bean-shaped, green gall bladder, the tortuous intestines, and what may represent the pallid gonads.


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Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich, 1918; Anatomy of a female grass frog. Inspiration for a thousand school dissection classes, the use of color in this image clearly defines internal organs that might otherwise appear an unintelligible tangle. The front end of the body cavity is dominated by ruby-red liver lobes and a pink, almost tonguelike heart (the actual tongue is shown protruding from the mouth). Behind the liver and the mauve, lobulated lungs, a branching red blood vessel meanders lazily across the pink stomach as it traverses from the animal’s left to its right. Behind this, the pink tangles of intestine mingle with the unrealistically green follicles of the ovary—spawn-inwaiting. Even the muscles of the hind limb are delineated with care, and almost every one has a direct evolutionary equivalent in the human leg.

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St George Mivart (1827–1900), On the Genesis of Species, 1871; Skeleton of a Flying-Dragon. The title of Mivart’s tome may not sound particularly original, but it provided an excuse for yet another nineteenth-century comparative biologist to commission images of his favorite anatomical oddities. Much of the skeleton of the flying dragon is of little interest, except for the enormously extended middle group of ribs, which has given up its original ventilatory function and taken on a new role in supporting large, aerodynamic gliding surfaces. Several other groups of vertebrates can glide, but most of them make less profound alterations to their anatomy, instead using overgrown skin flaps, as in the mammalian sugar glider, or fins in the case of flying fish.


The jawless fish are among the most strange and distorted of all vertebrates. They are classified into two groups: the hagfish, which are specialized for a life of deep-sea carrion-eating, and the lampreys, an array of freshwater and marine forms, many with unpleasant parasitic lifestyles. Instead of jaws, lampreys have oval mouth-plates studded with denticles, with which they burrow into their hosts before extracting blood and dead tissue.


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Karl Gegenbaur (1826–1903), Principles of Comparative Anatomy, 1870; Mouth opening of Petromyzon marinus [Sea lamprey] with the “horn teeth.”

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Auguste Chauveau (1827–1917) and Saturnin Arloing (1846–1911), Traité d'anatomie comparée des animaux Domestiques, 1891; Skull of a cat and camel skeleton. From the nineteenth century into the twentieth, veterinary medicine became a more important discipline in its own right, and broadened its focus from horses and cattle to other species, including pets, as well as species domesticated outside Europe and North America. Some of the anatomical knowledge gathered in ruminants such as cattle and sheep was transferrable to their relatives, the camels, and their South American relatives, the alpaca, vicuña, and llama. Also, veterinary focus has gradually turned toward pet carnivores—cats and dogs.

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Richard Owen (1804–1892):

The Anatomy of Vertebrates, 1866 Few scientists have generated as much debate as Richard Owen—his contributions to anatomy and paleontology were enormous, but it is hard to find anyone who speaks kindly of him. He wrote 600 books, coined the term “dinosaur,” and founded London’s Natural History Museum, yet much of his life seems to have been spent in conflict with others. Originally from Lancashire, Owen became fascinated by anatomy even before he trained as a doctor, first in Edinburgh and then in London. His anatomical skills allowed him to identify the true nature of many

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fossils for the first time, including those of the giant flightless moas of New Zealand (see pages 141 and 142). Although Owen accepted the process of evolution, he disagreed with Charles Darwin over the mechanism underlying it, leading to considerable acrimony between Owen and Darwin’s more acerbic supporters. Owen was also known for failing to credit the contributions of others, writing unduly harsh reviews, and in some cases even attempting character assassination of his competitors. It is also suspected that he wrote anonymous articles that praised his own work and disparaged those with whom he did not agree. Owen has undergone something of a rehabilitation in recent years, with historians now calling him

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866– 68; Skeleton of the cobra (Naija tripudians).

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richard owen: the a natomy of v ertebr ates

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866–68; Skeleton of flamingo (top); Section of head showing vertical nostrils, Toucan (Ramphastos).

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“stubborn” and “uncompromising” rather than the less flattering epithets used in the past. Certainly, his contributions to comparative biology were wider than any other scientist—he reappraised the relationships between many strange groups of vertebrates, including lungfish, giant ancient amphibians, the proto-bird Archaeopteryx, and, of course, dinosaurs. He also wrote extensively on South American mammals, as well as the egglaying and pouched mammals of Australia. Yet the name Richard Owen still often has negative associations. There is even a permanent monument to one of his few scientific mistakes: the concrete Iguanodon and Megalosaurus still on display at Crystal Palace in London, following the Great Exhibition of 1851, stand frozen in the

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erroneously quadrupedal position insisted upon by Owen.

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866–68; Skeleton of the haddock. As this image suggests, bony fish have more bones than any other backboned animal, and much of the story of the evolution of land vertebrates has been the progressive discarding of many of those bones. Despite their various specializations, bony fish have sometimes been used as a “model” basal body plan for studies of comparative morphology. Certainly, the profusion of bony spars seen in this image made their way into Owen’s theoretical “vertebrate archetype.”

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richard owen: the a natomy of v ertebr ates

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866– 68; Base of skull of a male narwhal, with a section of the tusk (above, left); Section of the cranium and tusk of Elephant. Many mammalian species possess tusks, either modified canine teeth, as in the narwhal, or incisors, as in the elephant. Narwhals’ tusks are particularly unusual in that they are generally present on only one side, and have a helical “thread” running along their length—in those rare individuals with two tusks, the pitch of the helix is in the same direction, rather than opposite, in both. Apart from the tusk, the most noticeable feature of the elephant skull is the extent to which it is aerated and lightened by sinuses, air-filled outpouchings of the nasal cavity.

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Ernst Heinrich Philipp Haeckel (1834–1919), Anthropogenie; oder, Entwickelungsgeschichte des menschen. Keimes-und Stammesgeschichte (Anthropogeny or the developmental history of man), 1891; The Water Dachshund [duck-billed platypus]. Published in English as The Evolution of Man, the work of Ernst Haeckel (see pages 196–203) took a circuitous route toward the origins of our own species, with apes only appearing in its second half. Looking at these images, it is easy to understand why many in Europe did not believe early reports of the platypus—with its duck bill, beaver tail, otter trunk, birdlike eggs, and venomous claws. Even once accepted, it remained a focus of much scientific study: as one of the few egg-laying mammals, it was seen as an archaic missing link between reptiles and mammals. We now realize that it is, in fact, a remarkably successful animal, specialized for hunting aquatic invertebrates by detecting the electric fields created by their tiny muscles.

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Horace Jayne (1859–1913), Mammalian Anatomy; a preparation for human and comparative anatomy, 1898; The cat skull: the front and left side views. A quest for ever-increasing detail and accuracy has sometimes led to images that could be argued to prefigure the hyper-realism of some late twentieth-century painting and sculpture.

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Frank Evers Beddard (1858–1925), A Book of Whales. With forty illustrations by W. Sidney Berridge, 1900; Skeleton of Right Whale (top); Foetus of Beluga. Berridge’s Book of Whales is a rare book: dedicated to the largest and strangest of all animals, it is affectionate and scientifically incisive at the same time. The upper image demonstrates the extreme specialization of the whale skeleton, with enormous, filter-feeding jaws, fused neck vertebrae, specialization of the forelimb into a flipper, almost complete loss of the pelvis and hind limb, and adaptation of the tail skeleton to support horizontal flukes. The lower image is a baleen whale, whereas the beluga is a toothed whale, albeit an unusual one.

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[Thomas George Browne], Atlas of the Anatomy and Physiology of the Ox... With original plates by George Dupuy, 1927; Internal parts of a bull. As much at home in a butcher’s store as in a veterinarian’s surgery, the Atlas is a practical person’s guide to the internal parts of a bull. It consists of a series of pages that are opened to reveal ever deeper layers of bovine tissue, and some also include pop-up sections or organs that may be moved by pulling on attached tabs.

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Frank Evers Beddard (1858–1925), Mammalia, 1902; Three figures showing the cranial evolution of Titanotherium... (After Osborn). Now termed Megacerops, this gnarled, protuberance-laden creature, the size of an elephant, was actually a close relative of today’s rhinoceroses and horses, and lived in North America 35 million years ago.

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Beddard was Prosector of the Zoological Society of London, and had access to a large resource of exotic post-mortem material. Particularly arresting is his prey’s-eye-view of the Tasmanian devil, a famously predatory and bellicose marsupial. This species is currently under threat from the spread of an unusual contagious tumor, which is transmitted by devil-to-devil bites, but recent data suggests that the species’ reproductive biology is adapting to ensure more individuals produce offspring before succumbing to the tumor.


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Frank Evers Beddard (1858–1925), Mammalia, 1902; Front view of the skull of a Tasmanian Devil (Sarcophilas ursinas), showing Polyprotodont and carnivorous dentition.

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J. E. V (Johan Erik Vesti) Boas (1855–1935) and Simon Paulli (1865–), The Elephant’s Head: studies in the comparative anatomy of the organs of the head of the Indian elephant and other mammals, 1908; Superficial musculature of an elephant, a boar, a wolf, and a tapir.


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Among vertebrates, the muscles of facial expression are a characteristic feature of mammals—only our furry, warm-blooded relatives can snarl, suckle, and smile. Boas and Paulli’s landmark study illustrated the many ways that subcutaneous muscles animate the mammalian visage.


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Wilhelm Ellenberger, Hermann Baum, and Hermann Dittrich, Handbuch der Anatomie der Tiere für Künstler, [Manual of the Anatomy of the Animals for artists], 1911–25; Lion; Dog. Aside from the obvious technical proficiency and anatomical accuracy of these images, they are also perfectly posed to capture the personalities of the two species—the dog stands erect and alert, while the lion’s head has drifted lazily to the side, searching for opportunities across the savannah.



chapter 4

EMBRYOS & ANCESTORS Evolution and development in the 19th century

Ernst Haeckel (1834–1919), The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny, 1879; The cleavage of the forgs ovum. Engraving.

In the nineteenth century, science needed art and art assimilated science.

embryos & ancestors


iology changed utterly in the nineteenth century. Before 1800 there was description and classification, whereas by 1900 there were mechanisms and processes. Between those two dates, a cascade of fundamental changes in natural philosophy created our modern view of the living world. The art of animal anatomy remained as vibrant and diverse as before, but its focus changed. Instead of merely illustrating the creatures around us, the aim was now more often to make a point about where those creatures came from. The main forerunner to this shift dates back to the invention of the microscope in the seventeenth century, and a drawing by one of its pioneers, Antonie van Leeuwenhoek, appears on page 174. The microscope was to become part of the routine apparatus of anatomical

Carl Vogt (1817–95), Lectures on Man; His place in creation, and in the history of the earth, 1864; Side view of a skull, with Baer-Busk measurements.

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research, but it is easy to forget how its appearance led to a step-change in our understanding of animal structure and function. Rather than being an amorphous bag of protoplasm, the animal body was for the first time revealed to comprise millions of individual compartments, or cells, each of which has a little life of its own. And the idea that every animal is a community of these individual functional units raised the possibility that their interrelationships might one day be understood. Suddenly, animals had a novel microscopic inner life, and the scene was set for a grand era of discovery. Yet the first new realization to come in the eighteenth century was a radical change in our view of the large, not the small—a change in how we view our planet. For centuries, humans had been told that the Earth was several thousand years old, and that its creatures had been installed by God during an initial week of frenzied creation. Now, advances in the study of rocks, fossils, and radioactivity showed that our world is at least millions of years old, and also that for much of that time it has contained unfamiliar and sometimes monstrous animals that have not survived to the present day. This is why several of this chapter’s images depict early attempts to understand fossils for what they truly were—not the drowned vestiges of antediluvian monsters, 16 9

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Richard Owen (1804-1892), On the Anatomy of Vertebrates, 1866–68; Embryo Snake (Natrix).

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Ernst Heinrich Philipp Haeckel (1834-1919), The Evolution of man: a popular exposition of the principal points of human ontogeny and phylogeny, 1879; The great chain of being.

but real animals related to those we see today. There was suddenly a great deal of time, geological time no less, for interesting developments to have happened to animals. Indeed, natural philosophers had long suspected that the apparent similarities between animals reflected something deeper than mere coincidence or divine design. Thinkers such as Jean-Baptiste Lamarck developed the idea that animal species might change over time, and even that they can split into multiple new descendant species. The term “evolution” was appropriated from the linguisticians who, in the previous century, had deduced that the world’s languages had undergone a similar process. Unfairly, Lamarck is now mainly remembered for suggesting an erroneous mechanism for how this evolutionary change occurs, but his idea that animal types slowly change, diverge, and die out was incredibly powerful—it entirely reset the way we think about animals. The next crucial discovery was the force that drives this evolutionary change. Although animal change had been accepted by many scientists, the mechanism by which it occurs was still mysterious. The story of how Alfred Russel Wallace and Charles Darwin, both separately and in collaboration, formulated their theory of natural selection is one of the most famous in biology—and that theory is itself biology’s 17 0

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most important. Natural selection seems simple in retrospect, and its power lies in that simplicity: individual animals’ characteristics can be inherited by their offspring, and those characteristics predominate in future generations if they increase those descendants’ chances of themselves producing successful offspring. Beneficial traits spread, and unhelpful ones fade away by the uncaring arithmetic of animals’ own success or failure. Natural selection was a theory initially argued in words rather than pictures. Darwin’s On the Origin of Species (1859) only contains one figure: a workmanlike attempt at an early evolutionary tree. Both Darwin and Wallace made drawings while they were conducting their researches abroad, but it was in the aftermath of their breakthrough that the pictorial legacy of natural selection really started to appear. The British paleontologist Richard Owen, for example, became obsessed with evolutionary homologies—structural similarities shared by species because they inherited them from some unknown ancient ancestor— producing striking diagrams that draw direct parallels between crocodiles, ostriches, and human babies. Another biologist, Ernst Haeckel (see pages 196–203), was a major force behind the other great theme of nineteenth-century biological science—embryology. Continual improvements in microscopy now permitted biologists to elucidate the story of animal development. The mammalian egg and sperm had already been discovered, and now the embryonic program that creates the complexity of every individual animal could be dissected. The idea that something as apparently amorphous as a fertilized egg could proliferate, differentiate, fold, and distort to produce a perfect little animal still seems amazing today, and many of the most striking anatomical images of Darwin’s century relate not to evolution, but to development. Bats, cats, lizards, and sharks—all are recruited to the artist-embryologist’s palette. Yet the boundary between evolution and embryonic development was blurred almost as soon as it was recognized. Early in the twentieth century, scientists realized that the two processes must be linked. For a species’ shape to change over millennia, it is not the adult-creatures that must evolve, but the developmental program that forms each animal is slowly rewritten. Long before computing or genetics, it was understood that progressive modification of the “program” and data used to create each animal is the fount of the amazing natural diversity around us. The images in this chapter show how the links between the evolution of the species and the development of the individual came to obsess Haeckel and others—to wonderful artistic effect.

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A dark side to this scientific quest emerged, however, when these ideas were carelessly applied to humans. People evolve just like any other animal, but many biologists applied their understanding of evolution incautiously, and in thrall to their own racial preconceptions. In this chapter, there are examples of misguided anatomical comparisons between different human populations and apes, and pseudosciences such as “craniometry” dominated many scientists’ understanding of race. Worst of all was the misuse of links between human evolution and development, and the concept that some races are “more advanced.” Indeed, the powerful images created to promote this idea were to inspire the racial policies of early twentieth-century Europe. The dark clouds were surely gathering alarmingly when Haeckel wrote that non-Europeans are “physiologically nearer to the mammals—apes and dogs—than to the civilized European. We must, therefore, assign a totally different value to their lives.” The century closed with the flourishing of a more benign science in southern Europe. Working almost alone, the Spanish scientist Santiago Ramón y Cajal was mapping the pathways that together tangle that densest of all anatomical thickets—the brain. There will be more on Cajal later (see pages 216–219), but for now it is enough to say that his delicate pen-and-ink traceries are so beguiling that we still use them today to explain the workings of the central nervous system. Suddenly, the computer in our heads was no longer such a daunting challenge: Cajal’s images made it seem possible for the first time that scientists would one day understand the brain. In retrospect, the nineteenth century was the most important phase of animal anatomy. Classification and cataloging were suddenly redundant, and biological science was now fluid and dynamic in a way it had never been before—causes and mechanisms were now the focus. This century’s anatomical art illustrates the pace of change and the profundity of the new understandings. The boundary between animal and human art had become necessarily fuzzy because, after all, we now knew that the same biological forces craft both man and beast. In the process, creatures had been stripped down to their essence, thrown back in developmental time, hurled back in geological time, and dissected into the processes that had formed them—and form us, too.

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Santiago Ramón y Cajal (1852–1934), Textura del Sistema Nervioso... 1899; Stages in the growth of granule cell dendrites.

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Antonie van Leeuwenhoek (1622–1733), “Observationes de natis e semine genitali animalculis,” from Philosophical Transactions, 1665–1775; Spermatozoa of rabbit (figs 1–4) and dogs (figs 5–8). The discovery of spermatozoa, the smallest cells in the body, had to wait until the invention of the microscope in the seventeenth century. Here we see varying morphologies of different species’ normal and abnormal sperm, with the imprecision of early microscopes leaving space for a certain amount of artistic license.

Hendrik Bary (c.1602–1707) after Reinier de Graaf (1641–73), Anatomical image of the cervix, uterus, fallopian tubes and ovaries, 1672. Hendrik’s drawing of a ruminant animal’s reproductive tract is remarkably accurate, starting from the vagina (“N”), and depicting the cervix, uterine body (“A”) and horns (“B”), oviducts, and ovaries (“H”). Most striking are the tortuous blood vessels supplying the ovary—these are, in fact, only found in ruminant mammals, in which they allow a particular reproductive hormone to take a “short cut” from the uterine vein to the ovarian artery.


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Georges Cuvier, Baron (1769–1832), Essay on the Theory of the Earth 1815; Skeleton of the Megatherium, dug out of the Alluvial Strata near Buenos Aires. With the nineteenth century came the realization, or at least the suspicion, that the Earth is older than previously thought, and that many of the outsized and bizarre bones unearthed from the ground represent ancient animal species that are no longer living. The name Megatherium simply means “great beast.” This was identified as an extinct giant ground sloth by Cuvier at the start of the century from drawings sent back following its discovery in South America.

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Contemporaries of the dinosaurs, one of the most exotic groups of vertebrates ever to exist was the pterosaurs, or “wing-lizards.” This specimen, first described by Cuvier, was given the name Ornithocephalus, or “bird-head,” but is now classified as Pterodactylus, or “wing-finger.” It is hard to imagine the impact that the discovery of this third group of flying vertebrates must have had: they possessed long jaws full of teeth, unlike birds and bats, and their wings were suspended on hugely enlarged fourth digits—they flew, effectively, with their ring fingers.

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Georges Cuvier (1769–1832), Essay on the Theory of the Earth 1817; Ornithocephalus.

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Hugh O’Neill (1784–1824), Ichthyosaurus chiroligostinus, drawn from nature on stone, 19th century. Found in stone and depicted in stone, evidence of huge ancient animals came also from the sea. Clearly a modified quadruped, this early ichthyosaur was evidence that, just like turtles, several extinct reptile lineages also reverted to an aquatic environment. The ichthyosaurs were to become the most specialized, losing their hind limbs, giving birth to live young, and eventually bearing a striking resemblance to dolphins.

Mary Anning (1799–1847), Autograph letter to Sir Henry Banbury with a drawing of a Plesiosaurus dolichodeirus, found at Lyme Regis, 1823. Anning is one of the key figures in paleontology, largely due to her studies of the anatomy of fossil marine reptiles, including this plesiosaur. Unable, as a woman, to join any learned societies, Anning’s scientific contributions were frequently overlooked or wrongly attributed to men. The often-longnecked plesiosaurs were a distinct group from the ichthyosaurs (which Anning also studied) and the two may not be closely related.


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Karl Ernst von Baer (1792–1876), Über Entwickelungsgeschichte der Thiere (“Developmental history of animals”), 1828–37; Lateral view of embryo. As well as describing the processes of embryonic development more precisely than any before him, the Estonian von Baer was also one of the first scientists to investigate the processes and principles that underlie embryogenesis. For example, his observation that distantly related species share aspects of early embryonic development, whereas closely related species share late phases of development, was to form the basis of Ernst Haeckel’s later theories linking evolution and embryology. In this diagram the embryo is at a contorted stage, with the bulbous, gray, brain-filled head on the left tapering to the spindly tail on the right. The heart and blood vessels are depicted in bright red. The fetal membranes surround the embryo, connected at the body stalk (“pq”), later to become the umbilical cord.

Karl Ernst von Baer (1792–1876), Über Entwickelungsgeschichte der Thiere (“Developmental history of animals”), 1828–37; Embryos. Some of von Baer’s diagrams are remarkably schematic, illustrating how he attempted to look beyond the anatomical details to discern the twists, folds, flows, separations, and swellings that conspire to create a new animal. Today’s developmental biologists still study these very basic formative processes.


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Gideon Mantell (1790–1852), Reconstruction of an Iguanodon, 1834. Iguanodon was the second dinosaur to be discovered, and it was, in fact, initially spotted by Mantell’s wife, Mary Ann, in rubble unearthed during the building of a road. Mantell himself was already credited with having identified the first fossil teeth, and this new herbivorous creature was eventually named “iguana-tooth” due to its dental similarities to modern iguanas. Mantell had few bones to work with and his reconstruction is posed like a giant quadrupedal forerunner of the modern iguana, with a bony horn on its nose. Subsequent discoveries indicated that Iguanodon was, in fact, bipedal, and that its nasal ornament had actually been one of its defensive thumb spikes.


Although the discovery of Megalosaurus preceded that of Iguanodon (see opposite page), this reconstruction dates from 20 years after Mantell’s. It demonstrates how little information was available to early paleontologists: just like Iguanodon, Megalosaurus was later understood to be a biped, although a carnivorous one in this case. Owen had made the same mistake as Mantell, although he was quick to dismiss his own errors and focus on others’, and he spent much of the rest of his life attempting to discredit Mantell and belittle his undoubted contributions to paleontology.


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Richard Owen (1804–1892), Geology and Inhabitants of the Ancient World, 1854; Megalosaurus.

The most famous “missing link” of all, Archaeopteryx (“ancient-wing”) had been named from a single feather just five years before this reconstruction was published—ten fossil specimens have now been discovered. Birds are a specialized subgroup of bipedal, carnivorous, theropod dinosaurs, and Archaeopteryx is an obviously transitional form between what would colloquially be called “dinosaurs” and “birds.” The body plan is birdlike, with forelimbs specialized as wings and asymmetrical feathers indicating that the animal could actively fly as well as glide. However, the jaws are toothed and the pelvis retains ancestral characteristics. The style and composition of this image not only shows the skeletal evidence on which paleontologists based their reconstructions, but also strongly implies that this creature was an agile, modern avian, not a lumbering ancient lizard.

Richard Owen (1804–1892), The Life of Richard Owen, 1894; Cyamodus laticeps (cynodont skull). Placodonts were marine reptiles that were contemporary with the dinosaurs, and possibly related to plesiosaurs (see page 180). Their skulls had a distinctive morphology adapted to exerting large, crushing forces through the blunt, rounded teeth visible in the lower image.


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Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866– 68; Restoration of Archaeopteryx, a Mezoic bird.

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Richard Owen (1804–1892), On the archetype and homologies of the vertebrate skeleton, 1848; Skeleton of a baby; Crocodile. Paleontologists and morphologists have become obsessed with the evolutionary equivalences, or “homologies,” between the disparate vertebrate groups, not least because their depiction often carries considerable visual impact. Frequently, it is slightly horrific creatures that are being studied, and unsettling juxtapositions are often invoked, as in this comparison of a baby and a crocodile.


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Benjamin Waterhouse Hawkins (1807–1894), A Comparative View of the Human and Animal Frame, 1860; Ostrich, camelid, and prosmian. One of Hawkins’ most striking compositions sees a skeletal comparison between an ostrich, a deer, and a carnivore turn into an unnatural and frightening six-legged chimera. However, there is some good comparative anatomy around the periphery of this disturbing scene: on the left, the uniquely tortuous human spine is illustrated for comparison, and elsewhere contrasts are drawn between the hands and feet of humans and apes.

George Murray Humphry (1820–1896), Observations in myology: including the myology of Cryptobranch, Lepidosiren, dog-fish, Ceratodus and Pseudopus pallasii, with the nerves of Cryptobranch and Lepidosiren and the disposition of muscles in vertebrate animals, 1872; Cryptobranch. The muscles and nerves of the limbs are remarkably similar throughout the limbed vertebrates, and these images illustrate their configuration in an entire amphibian forelimb viewed from the side (above) and the underside of a forepaw (below). Almost all the structures shown have direct equivalents in mammals, including humans, and such homologies have been an immensely powerful tool for understanding animal structure and evolution. The artistic value of these images in undoubted: muscles fold like drapery, and tendons and nerves writhe like ivy among tree branches.

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Extinct giant sloths were favorites of eighteenth-century anatomical artists. The increased size of ancient land sloths renders their bones more attractively lumpen and tuberous than today’s already plodding and gnarled tree-dwellers. Thus they are presented not just as giants, but as giants weighed down, almost crushed, by their own gigantism. At least one of this slightly asymmetrical individual’s forepaws bears the grasping claws still seen in modern sloths.

W. Lens Aldous (1792–1878), Brain of a horse: cross-section, 1853. Aldous was a medical illustrator from London who specialized in microscopy, yet here he turns to the gross anatomy of a horse’s brain for his subject. The composition is unusual, and unexplained, giving the image the appearance of a bisected mushroom. The “stalk” is the brainstem and the “cap” is the cerebral cortex. The mushroom’s “gills” are the fine striations of the corona radiata, the distinctive “radiating crown” of fibers that connects the nerve cells of the cortex to those of lower regions.

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Joseph Dinkel (1806–1891), Skeleton of a mammal (giant sloth).

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John Gould (1804–81), in Charles Darwin (1809–1882), Journal of Researches into the Natural History and Geology of the Countries Visited During the Voyage of H.M.S. Beagle... from 1832–6, 1845; Beaks of the different species of Geospiza, or Galapagos finches. The story of Darwin’s youthful circumnavigation of the world, and how the animals he encountered sowed the seed of his later theory of natural selection, is probably the most famous in science. However, it was not until he had returned to Britain and his avian specimens had been cataloged and described by the ornithologist John Gould that Darwin realized that the profusion of subtly different forms across the islands of the Galapagos archipelago, and the tiny gradations of anatomical difference between them, suggested that all might have originated from a single founder species. He soon realized that this species may have diverged into multiple descendant forms in response to the environmental conditions on different islands. Darwin’s On the Origin of Species was published 14 years after this image, in 1859.

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The evolution of man, and our species’ relationship to other primates, were major focuses of late nineteenth-century biology, yet much of what was written is unacceptable by today’s standards. It was not until a more mature understanding of evolution arose and racist assumptions were discounted that real progress could be made. The human species is one of the most varied in existence, but this reflects the enormous variety of environments to which our ancestors adapted, rather than the inherent superiority of one human population over another.

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Carl Vogt (1817–95), Lectures on Man; His place in creation, and in the history of the earth, 1864; Cranium of an Australian (aborigine) in profile after Lucae; Skull of the Weeper monkey, Cebus apella, in profile.

Ernst Haeckel (1834–1919): Development of the embryo, development of the race

Ernst Haeckel, Professor of Zoology at Jena, was an unusual person—a traveler-savant, embryologist, artist, ultra-Darwinist, and self-promoter who even expounded his own pseudo-religion premised upon his ideas of development, evolution, race, and religion. Haeckel’s scientific and artistic output was profuse, from the watercolors of his youthful wanderings around the globe to his later sharp engravings extolling the patterns and geometry present in the animal kingdom. As well as being a working scientist, he was also

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a hugely successful popularizer of science, his many books selling worldwide, and he traveled around Europe with his roadshow of skeletons, embryos, and diagrams. Few have so successfully used their art to impose scientific opinions on future generations.

Ernst Heinrich Philipp Haeckel (1834–1919), Basic plan of a gilled, segmented ancestral vertebrate, Anthropogenie; oder, Entwickelungsgeschichte des Menschen (“The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”), 1879; Basic vertebrates.

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Most vertebrate embryos are nutrified not by a placenta, but by an egg yolk. This image illustrates the myriad ramifications of a developing vertebrate’s yolk arteries and veins, although there has been some artistic license in its creation. These “vitelline” vessels are relevant to both human and veterinary medicine, as they are modified during embryonic development to form the vasculature of the gut.

Haeckel’s life changed when he read Darwin’s On the Origin of Species, and there followed an uneasy scientific correspondence between the cautious, cerebral Darwin and his argumentative, bombastic protégé. Haeckel pushed the theory of natural selection further than anyone else by firmly binding it to his own main field of study, embryology. In the nineteenth century, many biologists already realized that evolution and development must be linked in some way, but it was Haeckel who claimed they were directly equivalent processes. 19 7

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Ernst Heinrich Philipp Haeckel (1834–1919), “The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”, 1879; Vitelline vessels in the germinative area of the chick embryo.

According to his Law of Recapitulation, every animal passes through all the stages of its evolutionary history as it develops as an embryo—thus humans would be said to go through single-celled, wormlike, fishlike, reptilelike, shrewlike, and monkey-like phases before becoming truly human. This idea may seem strange now, but it has been repeated to biology students ever since, even though it was largely refuted in the 1920s.

Ernst Heinrich Philipp Haeckel (1834–1919), Kunstformen in Natur (“Art Forms in Nature”), 1904; Chiroptera. As well as being a scientist, Haeckel was also an explorer who wrote extensively about his travels throughout the world as he absorbed and cataloged the animal diversity around him. Later in his life, he published “Art Forms in Nature”—a beautiful, multipart portfolio of the patterns he had observed, with flourishes that prefigure the

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nascent Art Deco style. There are few vertebrates in the book, but it seems Haeckel could not resist the Rococo ornamentation of bats’ snouts, which we now know modify the acoustic properties of the face in these echolocating creatures.

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A confusing feature of vertebrates is that they are constructed from at least two apparently separate systems of segmentation. Both are evident in this picture: the dark “somitic” segments running down the embryo’s back to the bottom of the image, and the bulbous, gill-like “pharyngeal” swellings in the throat. Still, today, no embryologist has succeeded in reconciling these two systems, nor explained why both exist. Indeed, a third system of segmentation is now known to be important in the developing brain.

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Ernst Heinrich Philipp Haeckel (1834–1919), “The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”, 1874; Segmental arrangement of a mammalian embryo.

So potent was the idea that it imbued a spurious sense of “progress” into evolutionary biology that was scientifically misleading when used to claim, for example, that monkeys are more “advanced” than sharks. But when Haeckel used his theories to claim that Europeans are superior to other humans, or that Judaism is an inferior, primitive forerunner to Christianity, one can sense the seeds of Nazi racial policy being sown.

Ernst Heinrich Philipp Haeckel (1834–1919), “The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”, 1874; Human embryo with its membranes, six weeks old. There is little to distinguish a primate embryo from any other mammal, and Haeckel used this similarity to support his theories linking evolution and development. Although schematic, this image is largely correct, except for the forest of fingerlike placental

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projections encircling the conceptus; in most primates, this shaggy region of placental exchange is focused into a single region, such as the discoid placenta of humans.


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Ernst Heinrich Philipp Haeckel (1834–1919), “The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”, 1874; Development of the Face: M, Man; B, Bat; S, Sheep; C, Cat. 201

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Ernst Heinrich Philipp Haeckel (1834–1919), “The Evolution of Man: a popular exposition of the principal points of human ontogeny and phylogeny”, 1879; Embryos of two lower and two higher vertebrates. This image is the most controversial in all of animal anatomy. To comply with his theory that animals re-enact their evolutionary history as they develop as embryos, Haeckel needed the early stages of all species’ development to look as similar as possible, and to diverge only in the later stages.


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twentieth century were Haeckel’s observations repeated, showing that many of the similarities he reported were fictitious. Some, especially those who find evolution unpalatable for religious reasons, have claimed that this represents deliberate fraud on Haeckel’s part, whereas others argue that it shows how indistinct was the boundary between scientific representation and interpretation before the twentieth century.


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Reproduced in textbooks for a hundred years, only at the end of the

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Francis Balfour, A Monograph on the Development of Elasmobranch Fishes, 1898; Pristiurus embryo hardened in chromic acid (above); Aired fin development (opposite). The dogfish is a popular animal with embryologists and evolutionary biologists. Not only is it readily available, but it also looks superficially as if it might be close to the ancestral body plan of vertebrates. Certainly, animals very like this little shark have been swimming in the oceans for hundreds of millions of years, so it may indeed have retained many ancestral characteristics. These delicate, precise images date from the early days of developmental biology at Cambridge University, the start of a strong tradition that continues to the present day.


Edweard Muybridge (1830–1904): Animal Locomotion: An electro-photographic investigation of consecutive phases of animal movements, 1887 Edweard Muybridge had an unusual life. Born in Kingston-upon Thames, in England, he changed his name from Edward Muggeridge to what he felt was a more archaically distinguished form, emigrated to the United States (only to nearly die in a stagecoach accident), shot dead his wife’s lover (only to be acquitted), and returned to England in his later years having changed the art and science of photography and, indeed, the way we see the world. Muybridge was initially a landscape photographer, specializing in

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images of the grandeur of the newly accessible American West. In the 1860s and 1870s, the Western states were still relatively untouched by the hand of man and Muybridge captured a world of untamed rivers, uninhabited coasts, and vast stands of gargantuan trees that had disappeared just 50 years later. Throughout his life, Muybridge was a pioneer in photographic technology, experimenting with time-lapse, stereoscopic images, panoramic photography, and double exposures, and this inventiveness was eventually to yield the images for which he is best known. At the time, there was considerable debate among horse-lovers as to the nature of the four equine gaits—and particularly whether horses lose contact with the ground during the trot and the gallop. Perhaps unsurprisingly, there were considerable sums wagered on the matter, so the hunt was on for definitive, scientific evidence.

Edweard Muybridge (1830–1904), Animal Locomotion, 1887; Bucking mule.


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The second and third frames of this series are proof that, at one point during the stride, a galloping horse loses contact with the ground. This occurs at a point when all its feet are gathered together beneath it—not when they are stretched far apart, as many artists had previously depicted. It is remarkable that this is now considered common knowledge, yet it was entirely unknown before Muybridge captured this photographic series. It is perhaps appropriate that a .GIF file of this image series was, in 2017, the first moving image to be encoded into the genome of a living creature, a bacterium.


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Edweard Muybridge (1830–1904), Animal Locomotion, 1887; 16 frames of racehorse “Annie G,” galloping.

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Edweard Muybridge (1830–1904), Animal Locomotion, 1887; Buffalo, running.

Over a period of years, Muybridge developed a photographic system for examining gaits, by arranging a series of cameras in a row to take sequential pictures of an animal as it ran past, hitting trip-wires as it went. Along with faster shutter speeds and more sensitive photographic emulsions, this contraption allowed Muybridge to produce images, unmatched artistically or scientifically‚ of the movements of people and an enormous range of different creatures. A previously hidden world of human and animal movement was suddenly revealed—a revelation satisfying in its own right, but which was also to lead directly to the unique visual art form of the twentieth century: cinematography.

Edweard Muybridge (1830–1904), Animal Locomotion, 1887; Baboon climbing a pole.


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Edweard Muybridge (1830–1904), Animal Locomotion, 1887; Cockatoo in flight. Muybridge adapted his original trip-wire-activated system so that a clockwork mechanism could trigger the exposures. As long as the velocity of the subject could be predicted, this allowed the capture of more fragile creatures that would have been ensnared by trip-wires.


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Although Muybridge’s name is not well known, his images permeate how we see the modern world. The graphic arts had always tried to freeze movement at a certain point in time, but photography now gave artists the idea of depicting movement by the superimposition of multiple Muybridge-like frames in a single image. This found its first expression in Marcel Duchamp’s Cubist/Futurist 1912 Nude Descending a Staircase No. 2 and also in this Futurist painting from the same year, in which dog, leash, and owner are propelled forward as a linked, resonating, bio-mechanical system. Particularly charming is how this painting conveys one of life’s truisms: a dachshund’s legs must move faster than everyone else’s.

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Giacomo Balla (1871–1958), Dynamism of a Dog on a Leash, 1912.

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I. Vaughan, Strangeways’ Veterinary Anatomy, 1904; Caecum and Great Colon of a horse. Perhaps the most unusual feature of mammalian physiology is that so many species are able to survive by eating grass, or other similarly unpromising herbage. In the late nineteenth century, it was already suspected that this is made possible by the presence of vast numbers of microbes living symbiotically in the gut, breaking down cellulose to produce digestible products. Here, the upper image shows the four-chambered stomach of an ox, distorted into a giant fermenting vat. The lower image is the tortuous, sacculated large intestine of a horse. Both structures take up at least half of their respective owner’s abdomen.


This diagram represents paleontologists’ attempts to classify extant and extinct reptiles, birds, and mammals according to the holes in their skulls. These skull fenestrae (”windows”) usually allow for the attachment of jaw muscles and are thought to be retained stably over the course of evolutionary time. The only forms recognizable to the untrained eye are the lower three in the rightmost row—from top to bottom: bird, mammal, and snake.

Hans Gadow (1855–1928), Amphibia and Reptiles, 1901; Head of Lachesis lanceolatis after removal of the skin. The snake skull is perhaps the most outlandish of any vertebrate, and is reduced in many species to a delicate framework of vestigial bony struts. An array of complex muscles allows these struts to pivot and slide on one another, widening the serpent’s gape and everting its venomous fangs.

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Hans Gadow (1855–1928), Amphibia and Reptiles, 1901; Diagrams of skulls, showing especially the composition of the bony arches of the orbito-temporal region.

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(Above) W.N. Parker (1857–1923), “On some points in the structure of the young of Echidna aculeata,” Proceedings of the Zoological Society of London, 1894; Structure of young Echidna. (Left) Frank Evers Beddard (1858–1925), Mammalia, 1902; Ovarian egg of Echidna. The monotremes—the egg-laying echidna (above) and platypus (shown on the opposite page)—have long been viewed as strange evolutionary relics. Because of this, they have been a mainstay of comparative mammalogy, on the assumption that they retain many ancestral traits that can help us understand the evolution and biology of mammals as a whole. In fact, both are very specialized creatures that are well adapted for distinctive ecological niches, and thus have evolved an array of specializations that have largely obliterated any evidence of the archaic traits they were once assumed to possess.

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(Above right) J.T. Wilson, “Description of a young specimen of Ornithorhynchus anatinus from the collection of the Australian Museum, Sydney,” Proceedings of the Linnean Society of New South Wales, 1895; Young echidna. (Above left) J.T. Wilson, “On the skeleton of the snout of the mammary foetus of monotremes,” Proceedings of the Linnean Society of New South Wales, 1901; Ornithorhyncus.

Santiago Ramón y Cajal (1852–1934):

Textura del Sistema Nervioso del Hombre y de los Vertebrados, 1899–1904 Santiago Ramón y Cajal was exceptional for three main reasons. First, he is one of the few scientists who single-handedly transformed his chosen discipline, neuroscience. Second, more than any other scientist, he became a national hero in his native country, Spain, which he was instrumental in bringing into the world’s intellectual mainstream. And finally, he wrote one of the most endearing scientific autobiographies ever produced,

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Recuerdos de mi Vida (1917). Born in Aragon, young Santiago was something of a tearaway, almost dying in an attempt to scale a cliff to visit an eagle’s nest, destroying part of his neighbor’s house with a homemade cannon, and eventually being thrown into the cells of the local police station. His father, a doctor, decided his son needed more intellectual focus for his energy, and perhaps strangely decided to take him on nocturnal grave-robbing sorties to acquire bones for the artistic child to draw. Santiago was transfixed with wonder by the bony landscapes he discovered, and so started a lifelong quest to understand human and animal structure. Many biologists had tried to use microscopy to understand the structure of the brain and nervous system, but most tissue-staining methods revealed no more than a bewildering mass of overlapping cells and

Santiago Ramón y Cajal (1852–1934), Textura del Sistema Nervioso... 1899; Glial cell of the spinal cord of a cow.


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The light-sensitive retina at the back of the eye is one of the most regularly ordered neural structures in the body. Typically there are three layers of cells—the densely packed photoreceptor cells at the top of this image, a less dense intermediate layer of processing cells, and, at the bottom, the ganglion cells which collate visual information from the other cell types and send it to the brain via the optic nerve.

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Santiago Ramón y Cajal (1852–1934), Textura del Sistema Nervioso... 1899; Cross-section of the retina of a mammal (green lizard).

densely tangled fibers. Cajal applied a silver-staining method developed by the Italian Camillo Golgi, who had discarded it largely because it was so inefficient. Only a tiny fraction of the nerve cells in a specimen took up the stain, and while this made others despair of its value, Cajal realized that this selectivity was the key to producing simpler, more comprehensible images. Also crucial was Cajal’s artistic skill, crafting delicate pen-andink diagrams so uncluttered and beautiful that they remain the clearest

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images of the cellular machinery of the brain ever made.

Santiago Ramón y Cajal (1852–1934), Textura del Sistema Nervioso... 1899; Schema of the philosophical and ontogenetic evolution of the pyramid cells. Harking back to Ernst Haeckel’s obsession with linking evolution and embryonic development, Cajal here compares these processes for the “pyramidal” cells of the cerebral cortex. The upper series depicts, from left to right, the pyramidal cells of a frog, a lizard, a rat, and a human. The lower series illustrates, from right to left, the development of a single embryonic pyramidal cell.


Before Cajal, many scientists thought the cells of the brain were fused into a huge continuous mass of thinking protoplasm, but Cajal showed this of individual nerve cells (or the Cajal-coined “neurons”) clearly separated from each other by tiny communication gaps, or “synapses.” Once Cajal had worked out which end of each neuron was the “input” end and which was the “output” end, he was able to map neural pathways throughout the entire nervous system. And so, with a flourish of artistic brilliance, modern neuroscience began.

Cajal discovered the pyramidal cells—billions of tiny units that form the basis of thought and movement in animals. The image on the left depicts a veritable forest of them in the cerebral cortex, while the image on the right is a rose-tinted portrait of a single cell and its communicating tendrils.

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Santiago Ramón y Cajal (1852–1934), Textura del Sistema Nervioso... 1899; Granular layer and cat neuron.

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was not the case—instead, the nervous system comprises tens of billions

chapter 5

onward, inward, outward The wonders of life since 1900

Thomas Russell and David Linstead, Mouse nose, transverse section. Polarized light micrograph (composite of 18 images) from a slide mount made by Russell between 1868 and 1901.

The last century has seen more biological discovery and artistic innovations than any other.

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he art critic Robert Hughes once wrote that centuries rarely begin on time, and this is certainly true of the anatomical art of the twentieth century: it was late. The distinctive visual flavors of the twentieth century did not infuse the art of animal anatomy until the Great War of 1914–18 was over. That conflict is often credited with changing attitudes more than any other historical event, and its industrialized, futile extermination shocked art into repeated cycles of re-evaluation. Since 1914, the pace of social, technological, and scientific change has been so rapid that art has had trouble keeping up. There have been more artistic movements, “-isms,” in the ensuing hundred years than in the rest of our history. Cubism, abstract expressionism, pop art—these new ways of seeing the world could not help but change future forms of expression. Perhaps this is why everything before 1914 simply looks “old,” while everything since looks somehow “modern.” This shift to a rapid succession of artistic styles is one reason why the images in this final chapter are the most diverse, but it is not the only reason. In addition to this -ism overload, twentieth-century scientists were accumulating new information at a ferocious and exponentially increasing rate. There was so much novelty to depict and interpret, and most of it demanded new modes of depiction—fossil reconstructions, cellular mechanisms, the genetic code, biomechanics, the processes of thought. New things to draw and new ways to draw them. Paralleling these innovations was another change in the entire philosophy of science. Just as flux for flux’s sake was abroad in art, scientists too became more and more obsessed with questioning rather than answering. Each new discovery seemed to raise more problems than solutions, and the sum total of what humans did not know was increasing at an alarming rate. Yet scientists relished this new uncertainty, and their messy, fluid ignorance of the universe inspired rather than intimidated them. Instead of respecting the knowledge of the ancients, scientists now deliberately challenged everything they had been told. Nothing was unchallengeable, and thinking the unthinkable became de rigueur. Some of the most radical and distorted depictions of animal form came from a man­—D’Arcy Wentworth Thompson (see page 230)— 222

working in the incongruously genteel environment of the Universities of Dundee and St Andrews. Indeed, distortion was his thing. It had been known for some time that mathematics crops up time and again in nature, in the spiraling seeds of a sunflower and the expanding coils of the nautilus’ shell, for example. Thompson took these observations and combined them with the obvious fact that biological systems have physical forces acting on them—forces that can be calculated with simple formulae—and developed his own world of animal geometry. In Thompson’s images, chromosomes divide with arithmetic precision, bones appear as wondrous works of engineering, and, most striking of all, animals evolve by distortion. His view of things was strictly analog, with fish, pterosaurs, and horses pinned to a grid, and then stretched and transformed by mysterious extrinsic forces to generate new life-forms. Every animal becomes a data set, a matrix, waiting to be forged into new shapes by the stresses and strains of the physical world. Thompson’s pictures are convincing and compelling, but they now seem outdated. They were published not long before the 223

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Frank Evers Beddard (1858–1925), Mammalia, 1902; Cerebrum of a female chimpanzee at two years old (left); Skull of chimpanzee, Anthropopithecus troglodytes (After de Blainville).

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Sir D’Arcy Wentworth Thompson (1860–1948), On Growth and Form, 1917; Chromosomes, undergoing splitting and separation.

discovery of the structure of DNA, a discovery that implied that life is controlled not only by complex analog numerical equations, but also by a crudely simple digital genetic code. The early twentieth century also saw, for the first time, artists using animal anatomy purely as a prop—using it not because of what it is, but simply because of how it looks. One of these artists was Georgia O’Keeffe, who moved from New York City to New Mexico in the 1930s and was entranced by the light, colors, contours, and sharp lines of the American Southwest. O’Keeffe drew landscapes, and famously tumescent flowers, but she also drew bones. Flesh rots quickly in the desert, and is then desiccated away by the wind to reveal bones that last for years in the dry, clear air, bleaching in the sun. Obviously, bones were a symbol of O’Keeffe’s new home, and acted as potent memento mori too, but she also used them as visual tools divorced from their actual identity. In her paintings, an ox skull forms the white in a spacious but ominous version of the homely American red, white, and blue; a disarticulated pelvis becomes part of the majestic desert landscape around it, a vast, white rock-arch looming over the red grit beneath. Ever since, bleached bones have become a symbol of the region, from tacky, themed ox-skull restaurants (see pages 246–247) to the 1970s album covers of certain later eastern migrants to the great open West. Back in the laboratory, new preparation techniques were changing the timeframe of anatomy from “draw-it-before-it-rots” haste to an altogether more relaxed pace. The preservation of specimens with formaldehyde, phenol, and their less toxic successors allowed 224

dissections to be slow, painstaking, and almost permanent. Preservation often causes the garish colors of fresh flesh to fade to subtle browns, but the longevity of pickled specimens—lasting for decades in some cases— meant they could be displayed in museums as works of the anatomist’s art. When Damien Hirst revealed his famous pickled sheep, Away from the Flock, and shark, The Physical Impossibility of Death in the Mind of Someone Living, he was, as he so often does, simply mimicking the techniques and processes of others, but on his new and distinctively overblown scale. A more recent technique, plastination, produces anatomical artifacts of an even more remarkable nature. In this technique, fats and cell contents are removed with acetone and suction, and replaced by silicone plastic that is pumped in under pressure. The result is a dry, odorless, perfect replica of the original tissues, retaining much of their original color. Plastinated specimens can be tiny or huge, depending on how big one’s acetone and silicone tanks are, and entire horses have been immortalized in this way. Combined with processes in which plastics are injected into blood vessels, windpipes, and any other available biological tube, plastination has been used to produce a veritable Noah’s ark of dissected plastic creatures, often displayed in exhibitions in the great galleries of the world. 225

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Eugène Nicolas (1867–1928), Ophtalmologie Vétérinaire et Comparée (“Veterinary and Comparative Ophthalmology”), 1917; Dog—normal eye; Cat—normal eye.

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Edwin Stephen Goodrich (1868–1946), Studies on the Structure and Development of Vertebrates, 1930; Hesperornis regalis, Marsh, Upper Cretaceous, Kansas. Restoration of a skeleton 1/8.

Another source of dramatic new anatomical art has been recent advances in diagnostic imaging. Radiographs have been with us for over a century, with their ghostlike simulacra hovering in a darkened virtual space, but they have now been joined by a variety of new imaging techniques. Computed Axial Tomography (CAT) scans are effectively three-dimensional radiographs that can be computer processed to create color-enhanced, rotatable images, or video footage traveling through a series of cut sections—a view of the body no anatomist from a previous era could have imagined. Magnetic Resonance Imaging (MRI) scans use magnetic fields and radio waves to map the spin of the protons inside animal and human bodies, while Tensor-Diffusion Imaging generates multicolor renderings of nerve fibers in the living brain and muscle bundles in the beating heart. 226

anatomical street art, architecture, and animal body-painting is inspiring, and anatomy as art is as potent a force now as it has always been. The more we know, the more we are drawn to depict, it seems. Film, pixels, oil on canvas: there is no end to our obsession with the interior life of beasts.

Rachel Murray, Magnetic resonance image of the distal radius, carpal bones, and proximal third metacarpal bone of the horse, 2017.


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The vast amounts of data these techniques generate must be simplified and color coded for the human mind to comprehend them, and the resulting images are made beautiful by the artistic instincts of the computer programmer. Despite the scientific insights that the cool, dispassionate process of modern biology has given us about the inner working of animals, depictions of animals’ anatomy still strike a deep emotional chord. Appropriately, they still strike us, one might say, as visceral. For example, it is no surprise that one of the longest-running modern film franchises is built around the Swiss artist H.R. Giger’s alien, and the oozing, fleshy caverns she inhabits. Giger’s unremitting horrors were not created from nothing—instead, he drew on the anatomy of eels, crabs, and insects, and combined them with the visual shock of phalluses, ribs, and offal straight from the butcher’s store. He used familiar organic tropes to create a callously violent, violating world that is, well, alien. And so the art of animal anatomy continues onward into the future, along with its connotations of beauty, death, awe, fear, and enlightenment. The level of sheer craft evident in contemporary

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Edwin Stephen Goodrich (1868–1946), Studies on the Structure and Development of Vertebrates, 1930; Diagram of segmentation of the head in a Selachian [shark], above; Diagrams of components of dorsal root cranial nerves of Gnathostomes (below). The quest to discover the plan underlying the segmented structure of the vertebrate head continued throughout the twentieth century, and still continues today. It is possible that modern molecular techniques will soon resolve the question of how intrinsically regular, or irregular, this most complex of anatomical constructs actually is.


The embryonic development of the eye is centered around the lens (speckled yellow). This forms from a thickened plate of cells on the surface of the head (“A”), which folds inward to take on an oblate spheroidal shape (“F”). The cell nuclei (black specks) then arrange themselves across the “equator” of this spheroid and the constituent cells secrete crystalline proteins and become transparent. The image at left demonstrates how Cajal’s drawings (see pages 216–219) continued to dominate microscopic anatomy well into the twentieth century.


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W. Schimkewitsch (1858–1923), Lehrbuch der Vergleichenden Anatomie der Wirbeltiere (“Textbook of the Comparative anatomy of the vertebrates”), 1921; Development of the eye (above); Structure of the retina (left).

D’Arcy Wentworth Thompson (1860–1948): On Growth and Form, 1917

By the start of the twentieth century, it was known that the universe could be explained, to any attainable degree of accuracy, by physics— and thus that mathematics is the language by which it may be described and predicted. Living organisms may seem too complex to explain in these terms, but early in the twentieth century one man from Scotland decided to do just that.

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Thompson was a key figure in the development of the science of morphology—a step beyond descriptive anatomy toward an understanding of why animals are the shapes they are, and what processes guide the formation of that structure. Thompson was working at a time before it was understood how genes control cellular processes and embryonic development, and in a way, this ignorance liberated him. Without DNA or proteins to distract him, he was able to examine living beings as an arena for physics, mathematics, and geometry. Sometimes he was obviously examining physical phenomena, such as the forces acting on spicules of bone (as in the figures on the opposite page), but often his enquiries veered into the purely numerological or geometric. Nature may seem organic and imprecise, but look carefully and it is full of spirals and helices, arithmetic and geometrical series, areas and volumes, trigonometry and calculus. Indeed, it is difficult to look at the graceful helices of a ram’s horns without wondering what rules and principles underlie their formation. Despite its short title, On Growth and Form is a long book in which Thompson’s observations and imagination are given free rein to examine many different aspects of mathematical biology. However, it is his attempts to explain evolving animal forms as the result of simple mathematical transformations in a two-dimensional plane that are most eye-catching to the student of anatomical art. Humdrum beasts are stretched, compressed, and twisted until they become pterosaurs, sunfish, and a wide variety of other exotica. Today it is easy to assume that animal evolution is all about the digital code of genes, but long before the discovery of the structure of DNA, Thompson taught us that we must never lose sight of the fact that animals are also entities existing in an analog, physico-mathematical universe. 230

d ’a r c y w e n t w o r t h t h o m p s o n :


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D’Arcy Wentworth Thompson (1860–1948), On Growth and Form, 1917; Crocodile porosus, b. C. americanus, c. Notosuchus terrestris; Argyropelecus Olfersi; Sternoptyx diaphana; Diodon; Orthagoriscus.

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Sir D’Arcy Wentworth Thompson (1860–1948), On Growth and Form, 1917; Diagram of ram’s horns, A. frontal; B. orbital; C. nuchal surface. One of the most commonplace examples of natural geometry is the horns of the sheep. Horns are concretions of keratin protein produced by microscopic, fingerlike projections covering approximately triangular regions on the surface of the ovine head. The shape of the triangle and subtle deviations of the germinal projections determine the direction of horn growth, and the direction and pitch of the tapered helical shape of the horn. The triangular base dictates that the horn has three surfaces, and sporadic growth creates “growth rings” called St Venants’s curves (“A–B”).

Sir D’Arcy Wentworth Thompson (1860–1948), On Growth and Form, 1917; Geometric transformation during evolution of the horse. In this montage, Thompson uses the near-complete fossil record of the horse to suggest how the evolution of the ancestral Hyracotherium (lower right) to larger modern forms might be explained as the result of a series of geometric transformations.



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Oil on canvas. Gift of Anne Marmon Greenleaf in memory of Caroline Marmon Fesler, Indianapolis Museum of Art.

Georgia O’Keeffe, Cow’s Skull: Red, White, and Blue, 1931. O’Keeffe’s imagery is strongly influenced by the light, contrasts, colors, and signature objects of her adopted New Mexico home. Horizons are hazily hued, but sharply edged, and bones are dry-sun-blasted white as only desert air can. A skull becomes a morbidly patriotic stripe on the red, white, and blue; a pelvis becomes a giant looming rock arch—a frame, a lens for the landscape of the endless West.


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Georgia O’Keeffe, Pelvis with the Distance, 1943.

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Eugène Nicolas (1867–1928), Ophtalmologie Vétérinaire et Comparée (“Veterinary and Comparative Ophthalmology”), 1928; Composite of four plates showing ocular diseases of the horse, top left and right, and bottom right detail showing papillary stasis with edema in the cat. Clinical imaging techniques reveal new visual worlds, and one of the first was ophthalmoscopy. The inside of the human eye is an unexciting homogenous orange-red, but domestic animals’ eyes are a dazzling display of iridescence and sparkle. Before these images could be photographed, veterinary ophthalmologists painted them.


Donald Steven was Professor of Veterinary Anatomy at Cambridge and produced a series of pen-and-ink drawings to clarify some particularly tortuous anatomical concepts. This image is a left view of the first two chambers of a sheep’s stomach, which develop as twisted, distorted outpouchings of the rest of the stomach. Because of this, its three overlapping muscular coatings (white, gray, and hatched) run confusingly writhing courses over its surface. As an artwork in its own right, this image has echoes of some of the more sinuous works of 1960s Op artists such as Bridget Riley.


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Donald Steven, Muscle layers in the rumen, 1967.

Robert T. Bakker, from John H. Ostrom, “Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana,” 30 Peabody Museum of Natural History Bulletin 1–165, 1969; Deinconychus in full sprint (below); Reconstruction of the skeleton of Deinconychus antirrhopus, based on the hypodigm (right). Rarely can scientific images have made a point so clearly. Described in the late 1960s, Deinonychus (“terrible-claw”) was eventually to find fame as the model for the “raptors” in the film Jurassic Park, but here it is reconstructed and fleshed out for the first time. At a time when dinosaurs were often assumed to be lumbering, cold-blooded behemoths, Deinonychus simply did not fit that preconception. Its limbs were agile and clearly those of a biped, and its vertebrae were fused into an unyielding rigid fuselage like those of fast-running birds. Each hindfoot bore an outsized sickle-shaped claw for slashing its prey, presumably while the raptor hopped deftly on its other foot. Although Ostrom’s scientific paper suggested that Deinonychus was adapted for “moderately, but not

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unusually, fast running,” this is not the impression given by Bakker’s images. This is a hot-blooded, speeding monster, ready for the silver screen.

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Vasile Ghetie et al., Anatomical Atlas of Domestic Birds, 1976; Superficial head vessels of the turkey. One does not usually think of turkeys as possessing artistic impact, but this anatomical image from the Soviet Bloc is certainly striking. Birds evolved from dinosaurs closely related to Deinonychus (see previous page) and a hint of these affinities has crept into this depiction. Even the usually comical snood has morphed into a menacing Triceratops-like horn.

Ludovic Collin, Metaphase in a rat embryonic fibroblast cell, 2006. A modern counterpart to the D’Arcy Wentworth Thompson image of chromosomes (see page 224), this micrograph of a dividing cell shows blue-stained chromosomes lining up, ready to be torn apart by the green and red mitotic spindle and partitioned into two daughter cells. Normally, all these structures would be difficult to see, but here they are labeled with fluorescent stains. The stains can be used alone, but also in multicolored combinations for scientific or, it must be suspected, artistic reasons.


Von Hagens is best known for developing the technique of plastination (see page 225), and touring his human and animal zoo of plastinated cadavers around the world. However, this specimen has been prepared instead by infusing plastics into its blood vessels and then removing the surrounding tissues. Rarely has the humble duck been transformed into biological art.

Professor R. Bellairs, Embryonic chick (13 days old) stained to highlight the skeleton, 2007. Most of the vertebrate skeleton forms as an initial cartilaginous precursor or “model,” which is then replaced by bone. This chick has been captured mid-transition with its cartilage stained pale blue and its mineralized bone stained dark red. Considering the late stage of development, a surprisingly large fraction of the skeleton is still to be mineralized.


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Gunther von Hagens (b. 1945), Blood vessel configuration of a duck.

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Raffael Latorre and David Bainbridge, Plastinated specimen of a bisected sheep’s head, 2015. Plastination is a technique that allows anatomical specimens to be preserved in a robust, dry, odorless, plastic-like form that retains their original structure and color. The relative permanence of these preparations means that detailed dissections may be kept for decades. In this specimen, the head has been divided almost perfectly in two—the yellowish brain and spinal cord are visible on the upper right, and to the left of them lie the complex, folded bony scrolls of the nasal cavities. The mouth cavity is visible as a long, black space immediately above the fleshy tongue, while in the lower right the tubular windpipe disappears off toward the lungs.

Gillian Higgins, Horses Inside Out, 2016; The biomechanics of jumping: the high-jump. Higgins’ is an organization dedicated to education in equine management and training using anatomical principles. Others have painted anatomical structures on living animals, but only rarely have the results been accurate enough to be usable as teaching aids. Sitting somewhere between animation and still anatomical images, this technique allows the observer to visualize and even manipulate anatomical structures in real time.


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The Longhorn Bar & Grill, Amado, Arizona, in the shape of a steer skull, built 1970s, 2016. Ever since O’Keeffe, bleached skulls have been a symbol of the arid American Southwest, appearing everywhere from Eagles’ album covers to this kitschy bar in Amado, Arizona.

Rachel Murray and David Bainbridge, Horse foot polyptych, 2017. Magnetic Resonance Imaging (MRI) is a technique that involves the detection of protons in living tissues by disrupting and then measuring their spin inside a strong magnetic field. Within a fraction of a second,

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a radio signal emanating from the patient is processed to create a threedimensional image of its internal structure. These are MRI cross-sections of a horse’s foot taken by Rachel Murray of the Animal Health Trust in Newmarket, and further processed and arranged by the author. Read left to right and then from the top to the bottom row, they start at the level of the first phalanx bone of the digit (first to third images), pass through the second phalanx (third to tenth images) to the crescent-shaped third phalanx (eleventh to last images), before the tip of the toe disappears.


Street art is one of the oldest forms of artistic expression, but rarely has it touched on animal anatomy. One notable exception is SHOK-1, an artist working mainly in the south of England who specializes in spectacular, ghostly artworks based on X-rays, sometimes distorted or modified into novel and unexpected forms. This image of a dog’s skull is remarkable for its precision and accuracy, however, especially as it was created freehand without the use of stencils. As with most street art, the viewer is left to draw their own conclusions, but the sheer incongruity of a beautifully tinted, anatomically correct, outsized dog skull on a dull hoarding in East London is surely enough to snap passerbys out of their everyday routine.


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SHOK-1, X-ray of a dog’s skull, graffiti on a wall in Walthamstow, London, 2016.

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Martin Krzywinski, Comparison between human, chimp, mouse, and zebrafish genomes, 2013. Modern biological techniques produce enormous amounts of data, so much in fact that it can be difficult for the human brain to assess them without becoming lost in detail. Krzywinski attempts to render large data sets into a visually comprehensible form, and this frequently involves making them artistically attractive, too. Often he decides that the best format is circular, as in this example. Each quadrant in this image represents the 20–25 chromosomes of one vertebrate species (clockwise from upper right: human, chimpanzee, mouse, and zebrafish), and the arcuate lines connecting them across the central circle illustrate genetic similarities between the different species. The amount of information contained in this diagram is vast, but the image itself is both accessible and appealing.


Diffusion Tensor Tractography is a specialized type of Magnetic Resonance Imaging that detects nerve fiber bundles as they traverse the brain of live patients—something not possible by any other imaging technique. This image is a “tractogram” of the fiber tracts in a marmoset brain, with a range of colors used to allow different bundles of living nerve cells to be distinguished, creating an artistically inspiring image in the process.


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Hideyuki Okano and Junichi Hata, Tractography of a marmoset brain, 2016.



Illustrations are in italics. A A bear’s foot (Da Vinci) 17 A greyhound (Dürer) 38 A horse in profile divided by lines (da Vinci) 30 A Lion Attacking a Horse (Stubbs) 44–45, 76 Albinus, Bernhardus Siegfried 98 Aldous, W. Lens 192 amphibians 96, 213 Anatomia del Cavallo (Ruini) 6, 10, 50, 54 Anatomy of the horse... (Stubbs) 72 Anning, Mary 181 anomalous animals 135, 137 Archaeopteryx 187 Aristotle 14 armadillo 120 art, and science 168–195, 222–227 Away from the Flock (Hirst) 225 B baboons 209 Baer, Karl Ernst von 182–183 Bainbridge, David 244, 248 Bakker, Robert T. 238–239 Balfour, Francis 204 Balla, Giacomo 211 Banbury, Sir Henry 181 Bartholin, Thomas 95 Bary, Hendrik 175 Bat (Dürer) 35 bats 24, 35, 131, 198, 201 Baum, Hermann 51, 90 beavers 122 Beddard, Frank Evers 158, 160–161, 223 Belon, Pierre 94, 106 birds 98, 103, 106, 131, 153, 194, 226 Blasius, Gerardus Leonardus 97, 122–125 boars 120, 162 Boas, J. E. V. 11, 162–163 Bojanus, Ludwig Heinrich 138–139 Bradley, O. Charnock 91

Brehm, Alfred Edmund 92–93, 144, 144–147, 146 Bridges, Jeremiah 81 British Museum, London 15 Browne, Thomas George 159 Brunel, Isambard Kingdom 82 buffalos 208 C Cajal, Santiago Ramón y 172, 173, 216, 216–219, 218–219 camels 124, 150–151, 190 Casseri, Giulio Cesare 110–113 cataloging and cutting 46–53 cats 113, 117, 150, 157, 201, 225, 236 cells 173, 218–219, 218–219, 241 chameleons 123 Chauveau, Auguste 150–151 Cheselden, William 128, 131 chicks 105, 130, 197, 242 chimaerae 95 chimpanzee 119, 250 chimpanzees 223 chromosomes 224 circulatory system 140 cockatoos 210 cod 134 Coiter, Volcher 102, 102–105, 104 Collin, Ludovic 241 Collins, Samuel 117 Computed Axial Tomography (CAT) scans 226 Cours élémentaire d’histoire naturelle, Zoologie (MilneEdwards) 140 cows 216, 246–247 Cow’s Skull: Red, White, and Blue (O’Keeffe) 234 cranium 18, 195 crocodiles 188–189, 231 cutting and cataloging 46–53 Cuvier, Georges 134, 134–138, 136, 176–177 cynodonts 186 D Da Vinci, Leonardo 17–18, 17, 18, 26, 27–31 Darwin, Charles 146, 152, 170–171, 197


De humani corporis fabrica... (Vesalius) 50 deer 24 Deinconychus 238–239 digestive system 87 Dinkel, Joseph 193 dinosaurs 152, 176–177, 176–181, 180–181, 184, 184–186, 186–187, 238–239 dissection 15 divination 20–21 dogs anatomy 165 eyes 225 Greyhound 38 head 117 movement 211 placenta 100 skull 249 Spaniel 117 spermatozoa 174 doves 92–93, 144 ducks 243 dugongs 140 Dürer, Albrecht 18, 19, 35–39, 36–39 E echidnas 214–215 elephants 16, 24, 136, 162 Ellenberger, Wilhelm 51, 90 embryos 171, 182–183, 197, 199, 200, 202–203, 242 Equus (Schaffer) 48 Erxleben, James 142 evolution 170 F Fabricius, Hieronymus 108–109 face development 201 Falloppio, Gabriele 103–105 Fenton, Roger 141 Ferrari, Giovanni Battista 52 finches 194 fish 8–9, 95–96, 96, 110, 121, 149, 204–205, 231 flamingos 153 flying-dragon 148 foals 57, 69, 79 fowl, domestic 7 frogs 97, 114, 132, 147

G Gadow, Hans 213 Galen, Claudius 15, 18 Gegenbaur, Karl 149 genomes 250 Germain, Jean 107 Gessner, Conrad 19, 41–43 Ghetie, Vasile 240 Giger, H. R. 227 Gnathostomes 228 Golgi, Camillo 218 Goodrich, Edwin Stephen 226, 228 Gould, John 194 Great War, 1914–1918 222 Greeks, Ancient 14–15 Grew, Nehemiah 120–121 greyhounds 38

K Kitâb al-baytara (“Treatise on Hippiatry”) 10 Krzywinski, Martin 250 L Lafosse, Philippe-Étienne 78–81 Lamarck, Jean-Baptiste 170 lampreys 95, 149 Lancisi, Giovanni Maria 100 Landseer, Charles 83 Latorre, Raffael 244 Law of Recapitulation (Haeckel) 198 Leclerc, Georges-Louis 126 Leeuwenhoek, Antonie van 174 L’Histoire de la Nature des oyseaux (Belon) 94 lions 16, 44–45, 76, 143, 164 lizards 140, 217

I Ichthyosaurus chiroligostinus 180 Iguanodon 154, 184 Ilyas, Mansurf ibn Muhammed 71

M McBride, J. A. 87 Maerlant, Jacob van 16, 24 Magnetic Resonance Imaging (MRI) scans 226 Malpighi, Marcello 114 Mantell, Gideon 184 Markham, Gervase 53 marmoset 251 martens 104 Mayer, T. Walton 87 Megalosaurus 154, 185 Megatherium 176 Mettel, Nicolaus 133 mice 250 microscopes 168, 171 Middle East 16 Milne-Edwards, Henri 140 Miological horse, side view (suspended) (Lafosse) 80 miology 80, 191 Mivart, St George 148 moas 141–142, 152 monkeys 125, 195 morphology 230 mules 206 Murray, Rachel 227, 248 Muybridge, Edweard 206, 206–210, 208 Muzzle of an Ox (Dürer) 39

J Jayne, Horace 157

N narwhals 155



H haddock 154 Haeckel, Ernst Heinrich Philipp 156, 170, 171, 196–200, 196–203 Hagens, Gunther von 243 hagfish 95 Hata, Junichi 251 Hawkins, Benjamin Waterhouse 86, 143, 190 Head of a stag, pierced by an arrow (Dürer) 18–19 hedgehogs 99 Héroard, Jean 62–63 Hesperornis regalis 226 Heyden, Pieter van der 8–9 Higgins, Gillian 245 hippopotamus 120 Hirst, Damien 225 Hoffmann, Hans 12–13 horses anatomy 32–33, 64 in art 44–45, 76–77 arteries 90 biomechanics of 245 bladder 81 blood vessels 71, 74, 91 bones 227 brain 59, 65, 81, 192 circulatory system 53 colon 212 curing of 25 cutting and cataloging 46–51 diaphragm 66 digestive system 87 evolution 233 eyeballs 81

eyes 65, 236 face veins 51 feet 88, 90 foals 57, 69, 79 gait 206, 206–207 head 72, 74, 82–83, 90 heart 48, 67, 91 and humans 47–48 internal organs 50, 56 intestines 50, 55, 71, 212 kidneys 81 limbs 27, 46–47, 58, 62, 68 liver 81 lungs 67 membranes 69 meninges 59 muscles 46, 60, 74, 80, 81, 83 nerves 65, 78 in profile 30 racing 72, 75 side view 80 skeletons 10, 52–53, 63, 70, 73, 86 skull 81 spleen 81 stomach 81 sublumbar 66 teeth 84–85, 89 vascular system 87 visceral organs 81 windpipe 47 Horses Inside Out (Higgins) 245 Horse’s leg (Da Vinci) 27 Hughes, Robert 222 humans evolution 170 faces 201 genomes 250 and horses 47–48 skeleton 86, 98, 106, 143, 190 skulls 168, 195 uretha stone 120 Humphry, George Murray 191

Natural History Museum, London 152 natural selection 170–171 Nicolas, Eugène 225, 236


O Okano, Hideyuki 251 O’Keeffe, Georgia 224, 234–235 On Growth and Form (Thompson) 230 On the Anatomy of Vertebrates (Owen) 169 On the Origin of Species (Darwin) 171, 197 O’Neill, Hugh 180 Ornithocephalus 178–179 Osborn, Henry Fairfield 89 ostriches 190 Owen, Richard 99, 152, 152–155, 154, 169, 185 oxen 28, 29, 39, 112, 159 P Parker, W. N. 214 Parrocel, Charles 70 parrots 104 Paulli, Simon 11, 162–163 Pelvis with the Distance (O’Keeffe) 235 petroglyphs 16, 23 photography 206–210 The Physical Impossibility of Death in the Mind of Someone Living (Hirst) 225 Piacenza Liver 14, 15 pigeons 145 pigs 104, 120 pike 110 plaice 121 platypus, duck-billed 156 Plesiosaurus dolichodeirus 181 porpoises 97 preservation, specimen 224–225 primates 102, 190 pythons 136

R rabbits 41, 104, 113, 174 rams 22, 232 rattlesnakes 36, 146 rays 95 Recuerdos de mi Vida (Cajal) 216 reproductive system 175 reptiles 97, 136, 186, 187, 213 rhinoceros 36, 98 Rondelet, Guillaume 101 Rosenhof, August Johann Rösel von 132 Ruini, Carlo 6, 10, 48, 50–51, 54, 54–61 Ruysch, Frederick 129 S salmon 121 Schaffer, Peter 48 Schimkewitsch, W. 96, 229 science and art 168–195, 222–227 sea fox 97 sea lamprey 149 Share-Jones, John T. 88 sharks 95, 96, 96, 101, 116, 133, 225, 228 sheep in art 225 brain 115 faces 201 fetus 109 head 232, 244 intestines 21 liver 15, 20 placenta 100, 109 slain 23 stomach 237 skeletons animal 104–105 chimpanzee 119 fish 154 human 86, 98, 106 Megatherium 176 sloths 193 snakes 137, 146, 152, 169, 213 Snape, Andrew 47, 64–69, 130

sparrows 131 spermatozoa 174 Spoonbill 107 stags 19, 19, 126 Steno, Nicolaus 96, 116 Steven, Donald 237 Stubbs, George 44–45, 51, 72, 72–77, 75 The surgical anatomy of the horse (Share-Jones) 88 T tapirs 163 Tasmanian Devil 161 Tensor-Diffusion Imaging 226 Thompson, D’Arcy Wentworth 222–224, 224, 230, 230–233 Tierleben (Brehm) 144, 146 Titanotherium 160 toads 129 tortoises 105, 128, 138–139 toucans 153 Tractography of a marmoset brain (Okano and Hata) 251 turkeys 240 Tyson, Edward 118–119 U Uccello, Paolo 30 V vascular system 87 Vaughan, I. 212 vertebrates 95–99, 152–165, 154, 196 Vertical view through the center of the front toe of the horse (Ellenberger and Baum) 90 Vesalius, Andreas 40, 48, 49, 50 Vogt, Carl 168, 195 W Wallace, Alfred Russel 170–171 water tortoise 128 whales 19, 42–43, 158 Whistlejacket (Stubbs) 76 Willis, Thomas 115 Wilson, J. T. 215 wolves 163 Y Youatt, William 82 Z zebrafish 250


Credits Every attempt has been made to trace the copyright holders of works reproduced. Many illustrations are kindly provided under open access or Creative Commons licenses by their holding institutions or licensors. For these we’d like to thank:

Heidelberg University Library, Creative Commons license (CC BY-SA 3.0): p.149 J. Paul Getty Museum, Los Angeles: p.141 Library of Congress, Washington D.C : pp. 185, 207, 209, 210 © Luc Viatour / (CC 3.0 Unported license) (Biblioteca Ambrosiana, Milan): p.30 The Metropolitan Museum of Arts, New York: pp. 8–9 National Central Library of Rome: pp. 97, 122–125 National Gallery of Art, Washington, Rosenwald Collection, 1964.8.697: p.36 Rijksmuseum, Amsterdam: p.175 Saint Mary's College of California: p.148 University of Maryland, Baltimore, Digital Archive: p.116 University of Toronto Library: half title, 155 University of Virginia Library, Charlottesville, VA (Health Science Library): p.100 U.S. National Library of Medicine, Bethesda: pp. 48, 64, 66–67, 69, 75, 128, 131

Wellcome Library, London: pp. 20, 49, 53, 70–71, 74, 81 (bottom), 83–86, 92–93, 95, 98, 106, 114– 115, 117–118, 120–121, 130, 132, 143–144, 153, 170, 174, 180–181, 187–188, 190–193, 201–203, 220–221, 241–242 Yale Center for British Art, New Haven; Paul Mellon Collection: pp. Frontis, 44–45, 76, 78–81 (top)

Specific further acknowledgements for illustrations on the following pages are: Alamy Stock Photo, London/Science History Images: p.184 Art Digital Studio © Sotheby’s, Paris: p.102 Augsburg, Staats- und Stadtbibliothek: pp.110–113 Robert Bakker: pp.238–239 Bayerische Staatsbibliothek München, Res/2 Anat. 13 Beibd.2, Tab. I, II, III, IIII: pp. 103–5 © BNF/Bibliothèque nationale de France, Paris: pp. 10, 19, 34, 62–63, 108, 109, 126 © Bridgeman Images, London: pp. 211 (Albright–Knox Art Gallery, Buffalo, New York); 15 (De Agostini Picture Library / M. Carrieri); 40 (Glasgow University Library); 12–13, 37 (Graphische Sammlung Albertina, Vienna); 235 (Gift of Anne Marmon Greenleaf in memory of Caroline Marmon Fesler, Indianapolis Museum of Art, acc. 77.229); 35 (Musée des Beaux-arts et d'archeologie de Besancon); 77 (National Gallery, London); 73 (© Royal Academy of Arts); 17, 18, 22, 28–30, 38 (Royal Collection Trust © Her Majesty Queen Elizabeth II, 2018) Göttingen State and University Library: p.107 Gunther von Hagens’ Body Worlds: ANIMAL INSIDE OUT & Institute for Plastination, Heidelberg, Germany, www.animalinsideout. com: p.243 Hideyuki Okano, Dean, Dept of Physiology, Keio University School of Medicine, Tokyo, Junichi Hatai, Laboratory for Marmoset Neural Architecture, Riken Brain Science Institute, Saitama, and Keio University School of Medicine: p.251



Biodiversity Heritage Library (BHL), http:// pp. 156, 208, 224, 231–233 (Boston Public Library); 138–139, 238–239 (Harvard University, Museum of Comparative Zoology, Ernst Mayr Library); 189 (King's College London, Foyle Special Collections Library); 156–158, 169, 194, 215 (Marine Biological Laboratory, Woods Hole Oceanographic Institution (MBLWHOI) Library); 142, 214 top (Natural History Museum, London); 42–43, 186 (Naturalis Biodiversity Center); cover, title-page, 41, 101, 119, 129, 134–137, 145–147, 160–161, 214 bottom, 223 (Smithsonian Libraries); 140 (University Library, University of Illinois Urbana Champaign); 212–213 (University of California Library); half-title, 152, 154–55 (University of Toronto – Gerstein Science Information Centre)

Utrecht University Library: pp. 47, 65, 68


Gillian Higgins,, by kind permission: p.245

Giacomo Balla © DACS, 2018: p.211

Martin Krzywinski, Canada's Michael Smith Genome Sciences Centre: p.250

Georgia O’Keeffe © The Metropolitan Museum of Art: p.234

Mesrop Mashtots Institute of Ancient Manuscripts, Yerevan: p.25

Georgia O’Keeffe © Georgia O’Keeffe Museum / DACS 2018: p.235

National Library of the Netherlands, The Hague: p.24

Cover image

Photo © RMN-Grand Palais, Paris: p.19 (© BNF/Bibliothèque nationale de France, Département des estampes et de la photographie); 21 (Musée du Louvre/ René-Gabriel Ojéda); 27 (Bibliothèque de l’Institut de France / René-Gabriel Ojéda)

Alfred Edmund Brehm (1829–1884), Brehms Tierleben. Allgemeine kunde des Tierreich... 1918; Anatomy of a domestic dove. Half title (ii)


© 2018 Photo SCALA/ Art Resource, Florence; The Metropolitan Museum of Art, Alfred Stieglitz Collection, 1952 (52.203): p.234

Richard Owen (1804–1892), On the Anatomy of Vertebrates, 1866–1868; Bronchial cartilages of the Dugong. Frontispiece (iii)

Rachel Murray, Animal Health Trust, by kind permission: p.227, 248

Philippe-Étienne Lafosse (1738–1820), Cours d'hippiatrique, 1772; Horse, vessels.

Raffael Lattore, Veterinary Anatomy, University of Murcia, Spain, by kind permission: p.244

Title page (iv) Frank Evers Beddard (1858–1925), Mammalia, 1902; Front view of the skull of a Tasmanian Devil (Sarcophilas ursinas), showing Polyprotodont and carnivorous dentition.

Royal Collection Trust, London, on loan to the British Museum © Her Majesty Queen Elizabeth II, 2018: p.22 © 2016 SHOK-1: p.249 Susan Steven, by kind permission: p.237 Kurt Stüber, under GNU Free Documentation License, (detail): p.198 © Trustees of the British Museum, London: pp. 39, 133 Università degli Studi di Firenze-Ateneo, Florence: p.52 University of Wisconsin Digital Collections, Madison: p.164–651 Unıversity Rare Works Library, Istanbul: pp. 32–33 Vasile Ghetie, by permission: p.240 Wellcome Images, under Creative Commons licence (CC BY 4.0): pp.242 (Prof. R. Bellairs); 220–221 (David Linstead); 241 (Ludovic Collin. CC BY-NC) All other illustrations appear by kind permission of the author.