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    Robert T Bakker

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      foot brontosaur head, with only a handful of pencil-size front teeth,

      had to feed twenty or thirty tons of body. Obviously, the standard

      orthodoxy has it, the brontosaur's extreme microcephaly imposed

      severe dietary restrictions. Only the most nutritious and softest of

      water vegetation would have met the stringent requirements. And

      even with a superabundant supply of such green mush, the bron-

      tosaur's metabolism would still had to have been incredibly low—

      somewhere between the level of a tortoise's and a cactus's—for

      the great beast to survive at all.

      This argument has been repeated hundreds of times by

      schoolteachers and Ivy League professors alike. A recent issue of

      National Geographic featured a long piece by a respected curator

      at a university museum. Typically, this author scoffed at the idea

      of any brontosaur's having a high metabolism. He dismissed any

      such notion with a single fact: its head was too small. In a 1984

      article in a technical journal, a young paleontologist presented a

      mathematically reasoned argument that proved beyond the least

      GIZZARD STONES AND BRONTOSAUR MENUS I 125

      Yard-wide gizzard of a

      Brontosaurus. With its

      thick muscular walls and

      lining of hard rocks, the

      brontosaur gizzard could

      grind enough tough

      leafage to fuel a warm-

      blooded body.

      doubt that the big brontosaur's meager cranial apparatus was

      hopelessly undersized to provide for any sort of high metabolism.

      Several years before, a graduate student from Yale lecturing be-

      fore an enthralled audience at Harvard used the rate at which moose

      chew water lilies to prove irrefutably that a twenty-ton brontosaur

      simply could not support anything but the most subdued and slug-

      gish life style. Documentary proof. Irrefutable logic. The giant

      brontosaurs could only have spent all their lives in a somnolent

      state of semi-torpor, just barely moving their long necks to reach

      into the lukewarm water, poking slowly about for the softest part

      of the Jurassic swamp salads.

      But all these arguments, both popular and professional, leave

      out important pieces of the brontosaur puzzle: gizzards, stones, and

      moas.

      A white mouse sacrificed to a hungering alligator posthu-

      mously provides a most important clue. The bones of the mouse

      show up quite clearly in the alligator's stomach on the laboratory's

      television X-ray monitor. But the mouse's bones are not alone.

      The alligator's after-stomach is lined with hard, dense objects—

      gizzard stones. The gizzard stones are convulsed by sudden mus-

      cular contractions of the gizzard's walls. The monitor clearly shows

      the mouse is being chewed, not by teeth in the mouth but by stones

      in the gizzard.

      Naturalists who study big 'gators and crocs in the wild find

      huge masses of gizzard stones when they cut open the animals to

      126 | THE HABITAT OF THE DINOSAURS

      study their feeding habits. The stones are found only in one cham-

      ber of the stomach—the gizzard—and this one chamber has walls

      with grooves and folds to permit expansion and contraction. Even

      without X-ray monitoring, it is obvious that this stomach chamber

      is a churning compartment designed to crush and pulp the prey's

      body after the gastric juices begin their preliminary chemical

      treatment. Crocs usually select very hard stones—quartz and gran-

      ite pebbles, for example—to line their gizzards. If such materials

      are lacking in their native streams, they may use angular bits of

      hard wood, pieces of glass bottles, or whatever else is available. I

      have also seen one or two near-perfect fossil alligator skeletons

      containing a neat bundle of hard pebbles clustered between the

      ribs precisely where the gizzard was in life. These fossilized gastric

      mills demonstrate plainly that gizzard stones have been an essen-

      tial functional component of crocodilian food processing for many

      millions of years. And the study of crocodilian gizzards leads to

      some intriguing conclusions about evolution both in birds and in

      the Dinosauria.

      Zoos mislead their visitors by the way the species are housed.

      Birds are in the Bird House, of course, and crocodiles are always

      segregated to the Reptile House with the other naked-skinned,

      scale-covered brutes. So the average visitor leaves the zoo firmly

      persuaded that crocodilians are reptiles while birds are an entirely

      different group defined by "unreptilian" characteristics—feathers

      and flight. But a turkey's body and a croc's body laid out on a lab

      bench would present startling evidence of how wrong the zoos are

      once the two stomachs were cut into. The anatomy of their giz-

      zards is strong evidence that crocodilians and birds are closely re-

      lated and should be housed together in zoological classification, if

      not in zoo buildings.

      Both birds and crocs have the identical plan to their special-

      ized gizzard apparatus, and this type of internal food processor is

      absent in the other "reptiles"—lizards, snakes, and turtles. In both

      birds and crocs, the gizzard is a thick-walled, muscular, crushing

      compartment with two great tendons reinforcing the walls of mus-

      cle (these are the shiny sheets of tough tissue you cut off the tur-

      key gizzard before cooking it). In both birds and crocs, the muscular

      gizzard is just aft of the thin-walled glandular stomach where food

      is softened by gastric juices.

      This croc—bird digestive system makes a lot of mechanical

      GIZZARD STONES AND BRONTOSAUR MENUS | 127

      sense. We humans chew our food first, then pass it to the glan-

      dular stomach, where it is softened by stomach juices. Our system

      makes our teeth do the heavy work; they must crunch up the food

      as it comes directly through the lips. If the human diet is a civi-

      lized one, full of soft TV dinners and tender cuts of meat, our teeth

      don't wear much. But in primitive human societies the natural foods

      are often tough and gritty—the Anasazi Indians of ancient New

      Mexico wore their teeth down to the gums because tiny bits of

      sand got mixed into their cornmeal when it was ground on stone

      matates. Even horses wear out their huge molars if they have to

      feed on grass growing in gritty soil. But consider the advantages

      of the croc—bird system. They swallow without chewing and pass

      their food directly into the glandular stomach, where the food rests,

      softened by the gastric biochemistry. Then sphincter muscles act

      as gastric gatekeepers, letting the food pass on to the gizzard where

      it is chewed. The "teeth" of this system (the gizzard stones) don't

      begin their crunching work until the food has been rinsed, soaked,

      and softened.

      Crocs have powerful digestive processes. However, no croc

      species eats vegetation purposely; sometimes weeds are swallowed

      accidentally when the croc swallows turtles or fish. So crocs don't

      provide a complete picture of how a gizzard might work in an her-

      bivorous dinosaur lik
    e Brontosaurus. Fortunately many species of

      bird are plant-eaters, and vegetarian birds perform some truly

      spectacular gastric feats with their rock-lined gizzards. Ducks and

      geese shovel up hard nuts and grains and even live clams, chug

      them down to the gizzard, and crunch them up with the gizzard's

      lining. Clamshells, acorns, and corn kernels are all equally cracked

      into small pieces by this formidable gastric mill. Fruit pigeons do

      even better; their gizzard is especially tough and contains horn-

      covered "teeth" growing from the inside lining. Even the hardest

      of tropical nuts are swallowed hole, passed into the gizzard, and

      cracked with an audible thunk. Ostrichs shot in the wild have giz-

      zards lined with the hardest rocks—usually those rich in quartz—

      available in the countryside. And a large bird can carry around as

      much as a double handful of these stong gastric tools.

      Now the problem of tooth wear in nature is not a minor one.

      When wild species wear out their adult teeth and can't replace them,

      they die. Elephants possess huge adult teeth, the largest ever

      128 I THE HABITAT OF THE DINOSAURS

      evolved. But every elephant eventually wears out its last molar and

      wastes away along some swampy shore, attempting to gum soft

      water plants for nourishment. Having a continuous supply of teeth

      in each socket, as was the case for dinosaurs, eases the tooth-wear

      problem but doesn't remove it entirely. The basic adaptive diffi-

      culty is that the hardest material in a tooth—the enamel—is still

      much softer than the grit that covers most foods in nature. Wind-

      blown dust generally contains tiny specks of silica. Silica is natural

      glass, a very common material in rocks and soil. Plants growing in

      natural soils become coated with windblown grit and with dirt

      containing silica particles.

      Not only do soil and wind tend to make plant food gritty, but

      plants themselves sometimes evolve silica armor to discourage the

      plant-eaters. Horsetails are one such armored type of plant, an an-

      cient group dating back to long before the dinosaur. Modern

      horsetail species are sometimes called "scouring rushes" because

      peasant housewives used to scrub pots with horsetail stems. They

      scour well because evolution has provided them with special cells

      that concentrate silica from the soil. The silica cells armor the en-

      tire stem with row after row of glass-hard microlumps. A plant-

      eater learns quickly that a diet of horsetails will erode its teeth

      down to the gumline.

      Gizzards not only give plant-eaters an edge in their evolu-

      tionary struggle with plants. They also confer the freedom to do

      other things besides constant chewing. Pity the poor plant-eater

      with neither gizzard nor ruminating stomach—a zebra, for exam-

      ple. The zebra must chew each lump of grass directly, without

      soaking or softening. Zebra heads are large for their bodies and

      are provided with huge molars—twelve on each side of the mouth

      (twice the number humans have). Even with this dental armory,

      when grass is tough and sparse, zebras are nonetheless forced to

      spend nearly all their working hours plucking and chewing. All this

      chewing demands that the zebras remain out on the plains, ex-

      posed to rain, wind, and constant danger from lions and hyenas.

      What would happen if a zebra were supplied with a hypo-

      thetical gizzard? Such a zebra could pluck up grass quickly, with-

      out masticating, fill its forestomach chamber, and retreat to the

      shade and safety of a bush-covered hill to let its gizzard do all the

      work of mastication. Gizzards also free the animal's mouth for other

      GIZZARD STONES AND BRONTOSAUR MENUS 129

      activities—such as sex. With its gizzard doing all the work of

      chewing, the zebra could use its mouth to snort and whinny and

      make all sorts of elaborate noise display to attract mates and frighten

      sexual rivals. Ever wonder how tiny songbirds can afford to spend

      so much of their time singing? Little birds are notorious for their

      high metabolism, but when do they find the time to chew? They

      don't. As the warbler sings, its gizzard and forestomach are doing

      the food processing without interfering with the music.

      Cud-chewing mammals have evolved a soak-and-soften mech-

      anism almost as good as the gizzard. A cow or deer plucks a

      mouthful of gritty grass, swallows it without chewing, and passes

      the lump of grass to a series of special stomach chambers. These

      chambers are fermentation vats where gastric juices and yeastlike

      microorganisms clean the wad of food and break down the tough

      plant fiber. Only after the lump of grass has soaked and softened

      is it passed back up to the mouth to be chewed by the molars. The

      technical name for this stomach vat system is "rumen," and such

      cud-chewing mammals are called ruminants.

      The ruminant system must be reckoned as one of the best de-

      vices mammals have evolved for coping with tough plant food. Most

      of today's successful big plant-eating mammals are in fact rumi-

      nants—all the cattle, sheep, goats, antelope, deer, giraffes, and

      others. But the gizzard system must be considered superior.

      Imagine a twenty-ton Brontosaurus equipped with an ad-

      vanced, avian-style, rock-lined gizzard. A two-hundred-pound os-

      trich may possess a gizzard four inches across and a pound in weight.

      A roughly proportionate gastric grinder would provide a twenty-

      ton brontosaur with a gizzard of approximately one hundred pounds.

      One hundred pounds of tough muscle contracting a lining of big

      quartz pebbles could crush up Jurassic vegetation at a rate more

      than adequate to supply any level of metabolism. A hundred pounds

      of gizzard muscle weighs more than four times the jaw muscles of

      a five-ton African elephant. So four elephants, totaling twenty tons,

      possess less chewing power than the single hypothetical brontosaur.

      But what about that tiny head—would a brontosaur be able

      to engorge enough food to keep a giant gizzard apparatus going at

      full capacity? That question can be answered by turning to New

      Zealand, where up until a few centuries ago a giant, long-necked,

      pinheaded herbivore waddled about the landscape plucking leaves

      130 | THE HABITAT OF THE DINOSAURS

      from trees and crushing them with its gizzard. This native New

      Zealand plant-grinder was the moa—or more precisely, the moa

      family, a group of flightless species of bird that achieved a weight

      of half a ton. New Zealand's ecosystems evolved without any na-

      tive land mammals, so the role of large plant-eater was filled by

      the evolution of these big ground birds. Unfortunately for mod-

      ern science, the Polynesian colonists, the Maoris, who arrived in

      New Zealand about A.D. 1300, found the moas tasty and easy to

      kill, so moas were extinct before Western civilization could meet

      them alive.

      But moas created a sensation when they first turned up as

      fossils in New Zealand bogs and stream gravels. European zoolo-

      gists already knew ostriches well, because
    they had been circus fa-

      vorites from the time of the Caesars. But no one suspected that a

      plant-eating bird as large as a small buffalo could have existed. In

      1838, Sir Richard Owen, Queen Victoria's favorite anatomist, re-

      ceived a packet from New Zealand containing a curious bone

      fragment the size of an ox's femur. Owen was such an accom-

      plished comparative anatomist that he instantly recognized the

      fragment as from a bird—a bird possessing a body five times heav-

      ier than any previously known. With a courage few other young

      scientists might display, Owen publicly announced his discovery.

      Six-foot Maori hunter and

      the great moa, Dinornis

      GIZZARD STONES AND BRONTOSAUR MENUS

      131

      Based on this single fragment, he deduced the existence of huge

      birds rivaling the mammals in size. Owen's name for the extinct

      bird was an emphatic superlative: Dinornis maximus "enormous

      terror bird."

      Owen's announcement met with skepticism, but his judgment

      was vindicated by more and better discoveries in New Zealand:

      partial hind limbs, vertebrae, and then astonishingly complete

      skeletons found standing upright, buried in quicksandlike depos-

      its. Owen's deductions, based on one thigh fragment, were on tar-

      get—the greatest species of moa were twelve feet tall and must

      have weighed a half a ton. Other species varied down to pony size.

      Moa anatomy was full of surprises for biomechanical anatomists.

      The wings were nearly totally absent—unlike ostriches, moas re-

      tained not even a tiny feathered remnant. And the moa's head was

      tiny—a twelve-foot-high moa carried a skull no bigger than a

      poodle's.

      Moas are delightful objects of study in their own right, but

      their importance in the present discussion lies in the pinheaded

      configuration of their head and neck. At a distance, moas would

      have appeared as microcephalic as any brontosaur, with the tiny

      moa skull perched atop a very long, gracefully tapering neck. And

      moas were without doubt herbivores; their beaks were con-

      structed like those of living leaf-eating species. Unassailable evi-

      dence for their food preferences subsequently came from skeletons

      found in bogs and caves, where the stomach contents from the giant

      birds' last meal were mummified with the bones. These fossil meals

     


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