On the Prowl, by Mark Hallett and John M. Harris. Illustrated by Mark Hall (preface)

PREFACE

A young woman treads the narrow path rising from Silpuri village near the edge of Kanha National Park.

Her feet pound softly, syncopating with the click of her ankle bracelets and producing clouds of dust that rise to mingle with the dawn air, barely cool from the stifling heat of the previous day. Shafts of light cut through the shadows of the nearby forest to illuminate tall grass along the path in crisp detail. Suddenly she halts, stops, her gaze locked on two golden, staring eyes inside a concentric maze of coppery-yellow, stark white, and black. The wide brown eyes of the woman are apprehensive; the golden ones of the female tiger, secure and mildly curious. Both are mothers. They are separated physically by a distance of several meters, but the sources of their reactions were crafted millions of years apart, and the passing seconds contain flashbacks of raw flesh, flint blade points, and bullets; of thrusting mountain strata and glacial cold; of gnawing animal and human needs; of royal grandeur, lifeless trophy heads, and now vanished forests. The tiger finally yawns, exposing a long, crenellated pink tongue and yellow, pointed canines, before calmly padding across the clearing and downhill to join her cubs at the streambed. Breathing out, the woman continues along the path to check on her family’s rice paddy.

. . . . .

T

he great or big cats, known scientifically as the pantherins (terms in boldface type are explained in the glossary), are perhaps the most iconic of carnivores. Our instinctive fear of their lethal teeth, claws, and muscular power is balanced by an equally inborn fascination with their exquisite fur patterns, supple grace, and compelling presence. They have been represented for thousands of years on the walls of caves and palaces, on weapons, and in countless other images, ranging from loaded political symbols to innocent cartoons. As children we grow up with them as legends and myths, but only within the last sixty years or so, in the twilight of the pantherins’ existence on our planet, have we started to begin to understand their behavior in the wild. We are now in the process of uncovering their origin, and this book will explore what is currently understood on the basis of new discoveries and current scientific interpretations. Although much is still unknown, enough clues have now been uncovered so as to create a tentative, sketchy trail of where the great cats came from, sometimes crossing paths with our own emergence as modern humans. The story of the great cats’ evolution is based firmly in the fossil record, new interpretations of comparative anatomy, and the decades of patient field studies by wildlife biologists. Where the trail becomes faint, however, we must occasionally resort to intelligent, informed speculation, admitting what we do not know. This new summation in turn provides the basis for how we can work to ensure that pantherins and other wild cats will continue into the future.

xvi Preface The late British paleontologist Alan Turner (Liverpool John Moores University) and the Spanish artist-paleontologist Mauricio Antón (Museo Nacional de Ciencias Naturales) in 1997 jointly produced a superbly presented, highly readable, and beautifully illustrated book, The Big Cats and Their Fossil Relatives (Columbia University Press), which recounts what was known at that time about cat evolution, focusing in particular on both the true felid and nimravid “sabertoothed” cats. This was followed by the excellent Sabertooth (Indiana University Press, 2013), by Antón, and Biology and Conservation of Wild Felids, by David MacDonald and Andrew Loveridge (Oxford University Press, 2010). These two titles are classic works of paleontology and zoology, and because of the thoroughness with which the nimravid and machairodontine sabertoothed cats have now been assessed, their subject matter is therefore generally outside the scope of our book. Instead, we focus on the evolution and paleobiology of tigers, jaguars, leopards, lions, and other pantherin cats within the broad context of ancient changing ecosystems and on their possible relationship with other animals and humans, including the first documented contacts during more recent prehistory.

Chronologically, our story of big cat evolution focuses on the last 23 million years of the Earth’s history, an interval of time that contains the Neogene and Quaternary periods. The Neogene comprises two epochs.

The Miocene epoch started at 23.03 million years ago or, as abbreviated by earth scientists, at 23.03 Ma. The subsequent Pliocene epoch started at 5.33 Ma. The Quaternary period contains the Pleistocene epoch, which began at 2.59 Ma, and the Holocene epoch that began about 10,000 years ago (~10 ka) and has lasted until the present day. The epochs are in turn subdivided into stages or ages (appendix 1). The stages of the epochs are, for historical reasons, defined by different sequences of marine sedimentary rocks located in Europe; an age is the interval of time that it took to accumulate that rock sequence or stage.

Most fossil mammals occur in terrestrial rocks, and it is not always easy to precisely match or correlate European sequences of terrestrial rocks with their marine chronological equivalents; correlating the terrestrial rocks of other continents with the European marine stages is even more difficult. Mammal paleontologists have, therefore, subdivided the terrestrial fossil-bearing sequences of the different continents into their own unique sequences of land mammal stages (LMS) and land mammal ages (LMA).

Each stage has been assigned a type locality and is defined by its unique assemblage of mammals. For example, the type locality of the North American Rancholabrean stage is the Rancho La Brea Tar Pits of Los Angeles, and the Rancholabrean LMA begins with the arrival of bison into North America, recently established by Froese (University of Alberta, Edmonton) and colleagues as about 195,000 years ago. The largest fossil cat known

Preface

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from complete skeletal material, the American lion Panthera atrox, occurs only in the Rancholabrean LMA. In North America, the Neogene LMAs most important to our story are the Hemphillian (10.3–4.9 Ma) and the Rancholabrean (240 ka–11ka; see appendix 2).

Much of our story follows big cat history in Europe and Asia. The first mention of pantherin animals (species, families, etc.) that are of particular importance will also appear in bold and are listed in appendix 4. When describing the different events, we will—where possible and for clarity— use the epoch names and dates in millions of years. Where we need to be even more precise, we’ll provide the European or Asian stage names.

For Europe, the most important of these for our story are the Villanyian (3.6–2.58 Ma), Gelasian (2.58–1.81 Ma), and Calabrian (1.81–0.78 Ma).

The Asian LMAs most important to our story are the Mazegouan (3.6–2.59 Ma) and the Nihewanian (2.59 Ma–700 ka). Our emphasis on Asia, a crucial region for the origin of big cats, reflects the extensive but generally lesswell-known discoveries and academic studies over the last fifteen to twenty years in Asian paleontology by Chinese and other workers.

In addition, the timeless Middle and Late Paleolithic paintings and portable sculptures of pantherins in caves such as Chauvet-Pont-d’Arc and others in Europe offer a unique insight into the impact these cats had on the human mind in these places and times, and these are examined in chapter 5, “Testimony of the Caves.” We also seek to reconstruct the past ecologies of the pantherins’ endangered and, in some cases, now lost subspecies, such as the Caspian tiger and Barbary lion, beginning with the first records of the animals and ending with a survey of present-day surviving populations. The historic relationship between ethnic cultures and great cats, combined with the most recent field studies of pantherins, offers the possibility that new conservation attitudes and practices could bring these animals back from their currently endangered status to once again become wild, free carnivores. It is the authors’ fond hope that this book will in some way contribute to this outcome.

Praise for On the Prowl “On the Prowl is a superb book, both enjoyable and instructive. I’m greatly impressed with the huge amount of data that Mark Hallett and John M. Harris have collected and presented in a most readable manner. The complex evolutionary history and relationships of the cats will keep researchers arguing for years.”

—GEORGE SCHALLER, senior conservationist, Wildlife Conservation Society, and author of The Serengeti Lion: A Study of Predator-Prey Relations

“On the Prowl brings together biological and paleontological research related to the origins and history of the big cats, with an original emphasis on Asiatic data, paying homage to their majesty, dignity, and beauty. Hallett’s artwork has succeeded brilliantly in capturing their inner spirit and character, their vulnerability and vitality. This book will serve the noble goals of making people aware of the fragility of ecosystems and revealing the moral obligation to preserve and protect extant species.”

—CHRISTINE ARGOT, research professor and curator of collections, Muséum National d’Histoire Naturelle

“Hallett and Harris guide the reader on a 20-million-year tour of predator and prey evolution, using environmental change as an engine for adaptation and extinction. The narrative is engaging, spiked with a touch of controversy, and supported by superb illustrations. For those interested in the evolutionary origins of big cats, On the Prowl would be a valuable addition to your library.”

—CHRISTOPHER SHAW, Idaho Museum of Natural History

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