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The Age of Dinosaurs in South America First Edition
Novas Digital Instant Download
Author(s): Novas, Fernando E.
ISBN(s): 9780253352897, 0253352894
Edition: First Edition
File Details: PDF, 36.08 MB
Year: 2009
Language: english
THE AGE OF DINOSAURS IN SOUTH AMERICA

ening
Life of the Past

James O. Farlow, editor

THE AGE OF DINOSAURS IN SOUTH AMERICA

Fernando E. Novas

ndiana University Press

3loomington & Indianapolis
Gray-scale illustrations facing front and back matter openers are details from color
plates 14, 15, 16 © by Maurilio Olveira and color plates 17, 18 © by Gabriel Lio.

This book is a publication of

Indiana University Press

601 North Morton Street
Bloomington, IN 47404-3797 USA

www.iupress.indiana.edu

Telephone orders 800-842-6796

Fax orders 812-855-7931
Orders by e-mail [email protected]

© 2009 by Fernando Novas

All rights reserved

No part of this book may be reproduced or utilized in any form or by any

means, electronic or mechanical, including photocopying and recording,
or by any information storage and retrieval system, without permission in
writing from the publisher. The Association of American University Presses’
Resolution on Permissions constitutes the only exception to this prohibition.

@The paper used in this publication meets the minimum requirements of

the American National Standard for Information Sciences—Permanence
of Paper for Printed Library Materials, ANS! Z39.48-1992.

Manufactured in the United States of America

Library of Congress Cataloging-in-Publication Data

Novas, Fernando E.
The age of dinosaurs in South America / Fernando E. Novas.
p. cm. — (Life of the past)
Includes bibliographical references and index.
ISBN 978-0-253-35289-7 (cloth : alk. paper)
1. Dinosaurs—South America. |. Title.
QE861.4.N69 2009
567.9098—dc22
2008042734

(234
S14 is 12 Wil IO Os
 This book is dedicated to Argentine paleontologists

Osvaldo A. Reig (1929-1992)

José Bonaparte (1928-)

Rodolfo Casamiquela (1932-2008 )

who envisioned the importance of exploring and studying the Mesozoic in South America
a
CONTENTS

ACKNOWLEDGMENTS X/ 3 Jurassic Dinosaurs 87

LIST OF ANATOMICAL ABBREVIATIONS Jurassic Dinosaur-Bearing Beds

USED IN THE FIGURES XiI/ in South America 88

INTRODUCTION X/xX The Jurassic Dinosaur Record

in South America 700

Sauropoda 700
An Overview of the Anatomy
Neosauropods 108
and Phylogenetic Relationships
of Dinosaurs 3 Theropoda 774

What Are Dinosaurs? 3

Ornithischia 723

Dinosaur Skeletal Anatomy 4

Jurassic Dinosaurs: An Overview 128

A Brief Historical Account of the

Systematics of Dinosauria 77
The Fossil Record of Cretaceous
Dinosaur Forerunners 73
Dinosaurs in South America 735
Characteristics That Distinguish Dinosauria The Cretaceous Dinosaur Record
from Other Archosaurs 77 in South America 138

Triassic Dinosaurs 25 Cretaceous Sauropods 767

Triassic Dinosaur-Bearing Beds Diplodocimorpha 169
in South America 26
Titanosauria 187
The Triassic Dinosaur Record
in South America 35 Cretaceous Sauropod Eggs and Nests 236

Dinosauria 47 Cretaceous Sauropod Footprints 238

Saurischia 47

Theropoda 54 Cretaceous Theropods 243

Ceratosauria 246
Sauropodomorpha 57
Abelisauroidea 249
Ornithischia 77
Abelisauridae 272
Ichnological Evidence 76
Tetanurae 286
Triassic Dinosaurs: An Overview 8&0

Cretaceous Theropod Footprints and Eggs Boo

7 Cretaceous Ornithischians 343 8 ASummary of the Cretaceous
Stegosauria 345 Dinosaurs 385
Dinosaur Diversification in the
Ankylosauria 346 Cretaceous of South America 386

Ornithopoda 352

Ceraptopsia 379 WORKS CITED 405

INDEX 439

Vill Contents
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i
ACKNOWLEDGMENTS

I would like to express my gratitude to Pat Rich, who suggested that I sub-
mit a book proposal on South American dinosaurs to Indiana University
Press, and to Jim Farlow and Bob Sloan for their kind encouragement.
Many thanks to all of them for their trust in the results of this proj-
ect. Jim made clever comments and suggestions about how to improve
my original manuscript, but I am particularly grateful to him for his
encouraging words and support.
Several colleagues and friends from Argentina, Brazil, Bolivia, Can-
ada, Chile, England, Italy, Uruguay, the United States, and Venezuela
contributed with information and corrections to the present volume.
Thanks are due to José Bonaparte, Jaime Powell, Jorge Calvo, Leonardo
Salgado, Diego Pol, Teresa Manera de Bianco, Rubén Martinez, Ri-
cardo Martinez, Olga Giménez, Bernardo Gonzalez Riga, Luis Chiappe,
Andrea Cambiaso, Silvina De Valais, Andrea Arcucci, Sergio Martin,
Alejandro Kramarz, Eduardo Bellosi, Rubén Juarez Valieri, Juan Porfiri,
Sebastian Apesteguia, Ismar de Souza Carvalho, César Schultz, Manuel
Medeiros, Alexander Kellner, Max Langer Cardoso, Luis Borges Ribeiro,
Roberto Candeiro, Marcelo Sanchez-Villagra, Matfas Soto, Mario Suarez
Riglos, Saswati Bandyopadhyay, Fabbio Dalla Vechia, Paul Barrett, Phil
Currie, Cathy Forster, Mike Brett-Surman, Martin Lockley, Dave Krause,
and my brilliant students Federico Agnolin, Juan Canale, Martin Ez-
curra, Augusto Haro, Ariel Méndez, Diego Pais, and Agustin Scanferla
for sharing their knowledge with me.
All my gratitude to Fernando Spinelli, Stella Alvarez, Gaston Lo
Coco, Federico Agnolin, Martin Ezcurra, Diego Pais, Agustin Martinelli,
Rodrigo Vega, and Marina Caporale for their valuable help in preparing
the illustrations and text. Kate Babbitt’s detailed edit improved the manu-
script. The book has benefited from the artwork done by Jorge Gonzalez,
Carlos Papolio, Maurilio Oliveira, Orlando Grillo, and Gabriel Lio.
Special thanks to the skillful technicians Marcelo Isasi, Pablo Puerta,
and Santiago Reuil for their discoveries in the field and for the prepara-
tion of fossil specimens. I would like to thank the Argentine institutions
Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET)
and Agencia Nacional de Promocién Cienttfica y Técnica (ANPCyT) for
the support that enabled me to conduct research in my country, and the
National Geographic Society (Washington) and the Jurassic Foundation
(Calgary) for support for field work.
Finally, my gratitude to my beloved family—my wife, Liliana, and
my sons, Mariano and Francisco—for their help and patience.

Xi
ANATOMICAL ABBREVIATIONS
USED IN FIGURES

acetabulum biceps tubercle of the coracoid

antorbital fenestra cannaliculii
articular facet of the fibula capitulum
alveolar ridge calcaneum
scar for M. ambiens caudal ascending process
angular of the astragalus

acromial process carpal bones

ascending process of the astragalus cranial articular surface

articular process of the lacrimal cancellous bone

ascending process of the maxilla caudal1

articular process of the squamosal caudosacral 1

articular centrodiapophyseal lamina

coracoid foramen
articular cavity of the quadrate
carpometacarpus
articular surface of the scapula
cnemial crest
astragalus
coracoid
articular surface of the astragalus
coronoid process
articular surface of the ilium
caudal process of the ilium
anterior trochanter
collateral ligament pit
antitrochanter
coronoid
atlas
conical scute
axis
calcaneal tuberosity
bulk
crista tibiofibularis
blind excavation
cuppedicus fossa
brevis fossa
cervical vertebrae 1-9
beak
cervical rib
blp bilobate process
dentary
bpt basipterygoid tuberosity
dorsal vertebra 1-15
bptr basipterygoid recess
distal bevel on the pubis
brt brachial tubercle
distal carpals
bs brevis shelf
diagonal crest on the dorsal
bsp basisphenoid
surface of the astragalus
bspw basisphenoidal wing
dgv distal groove
bst basisphenoid tubera

xiii
 distal ischiac foot humeral head
diapophysis core for the horny spike

deltopectoral crest hyposphene

distal pubic foot hypantrum

depression hypotarsus

dorsal rib interdental plate

dentary simphysis ilium
dorsosacral vertebra iliac pedicle
distal tarsals 3-4 internal malleolus
ectopterygoid infradiapophyseal lamina
elliptical depressions ischium
extensor ligament pit interspinal ligament scar
epipophysis ischiadic pedicle
epipophyseal-prezygapophyseal ridge internal tuberosity
“foot” for anchoring the neural spine jugal
fourth trochanter frontal knob
fossa knob-like nasal projection
facet for the ascending lacrimal
process of the astragalus
lateral bulge of proximal femur
fibular condyle
lateral condyle of proximal tibia
fibular crest
ligament pit
femur
left iliac blade
femoral head
lateral lamina of the neural spine
fib fibula
left nasal
flp flexor ligament pit
lateral pit
flexor tubercle
lateral primary ridge of the tooth
foramen magnum
lateral ridge of the pubis
foot
lateral temporal fenestra
frontal
medial acetabular wall
surface for fibuiar articulation
tuberosity for insertion of
fan-shaped expansion M. biceps femoris
ventral furrow mcfb scar of insertion of
furcula (fused clavicles) M. caudifemoralis brevis

collateral groove mdc mediodistal crest of the femur

gastralia mf mandibular fenestra

glenoid cavity mo mosaics

guilliotine-like (cutting) margin mps site of origin of M. pseudotemporalis

greater trochanter ms muscle scar on the distal femur

humerus mtcl-V metacarpals I-V

haemal arch mtti-V metatarsals I-V

XIV Anatomical Abbreviations

median vertical ridge on the distal tibia pneumatic cavity
maxilla pneumatic foramen
maxillary fenestra postorbital
notch on the distal tibia postacetabular wing
nasal paroccipital process
neural canal postspinal fossa
nasal openning postspinal lamina of the neural spine
neural spine postzygapophysis
orbit parapophysis
dermal ossification type 1-4 proximopalmar hook-like process
obturator process prepubic process
occipital condyle prearticular
odontoid process of the axis prefrontal
obturator foramen prespinal lamina of the neural spine
olecranon process prootic
outer malleolus of the distal tibia prominence for muscular
attachment on the scapula
opisthotic
prespinal fossa
ossified ligament
prezygapophysis
parietal
prezygapophyseal protuberance
process for articulation of the dentary
pubic shaft
palpebral bone
pterygoid
pubic apron
pterygoid ramus of the quadrate
parasphenoidal recess
pubis
proximal articular surface
of the phalanx pubic pedicle
preacetabular wing posteroventral margin of the ilium
predentary posteroventral heel of
the pedal phalanx
posterodorsal lip of the ungual
posteroventral quadrangular
posterodorsal process of the ischium
process of the pedal ungual
peg in a socket
quadrate
phalanx I- IV
quadrate foramen
phalanges
quadratojugal
pubic ischiac articulation
rib
palatine
radius
posterolateral ridge
raised eminence
(or crista lateralis plantaris)
rugosity
protuberance for muscle
attachment on the distal pubis ridge

premaxila ribl right iliac blade

promaxillary fenestra right nasal

Anatomical Abbreviations XV
 rostromedial process of the maxilla tooth
retroarticular process tarsal bones
sacrum tibial condyle
sacrals 1-8 tibia
surangular transverse process

supracetabular crest trochanteric fossa of the

proximal femur
supraacetabular process
trochanteric shelf
scapula
tibial articulation surface
scapular blade
tuberculum
sclerotic ring
tuberosity
subcondylar recess
tympanic recess
tooth serrations
ungual phalanx
ligament groove on the proximal femur
ulna
sms sulcus for M. supracoracoidei
vomer
soc supraoccipital
splenial
ventral keel
ventral notch
squamosal
sacral rib
ventral surface

spine table wrinkles

wear facet
sternal plates
supratemporal fenestra first to fifth digit of manus or pes

stapes first to fourth phalanges

strp supratrochanteric process

Xvi Anatomical Abbreviations

ae ape MOPUbA)
1 [sry my
qv ial a y
HBAs te!9 mi [ew
aoeogy'A Pits blin nig tom
7 Serib al. Of

Masri T=."

‘jee Od bd ( a

aot (e's af eel

INTRODUCTION

South America has been revealed as one of the most important continents
for dinosaur discoveries. Although numerically less impressive than that
of North America, the fossil record of South America is highly important
for understanding the evolution of dinosaurs: their origin in Triassic
times; the acquisition of gigantic size by some species; the effects of the
geographic fragmentation of Gondwana on the evolution of dinosaur
communities. The fossil record of South America includes many “unique
pieces” that offer a glimpse into the evolutionary history of dinosaur
clades not recorded in other parts of the world.
After dinosaurs were first documented in South America in 1883,
knowledge of their evolutionary history remained fragmentary and re-
stricted to a handful of poorly understood species. Although some dis-
coveries were published prior to 1930, little progress in understanding
dinosaur evolution in South America was made during the first half of
the twentieth century. A major change in fossil documentation occurred
at the end of the 1950s, when exploration of the ‘Triassic beds of the
Ischigualasto Formation, which crop out in northwest Argentina, resulted
in remarkable discoveries. This exploration represented a pivotal mo-
ment in the history of dinosaur paleontology in South America because
it involved the participation of the first Argentine specialists —Osvaldo
Reig, José Bonaparte, and Rodolfo Casamiquela—
who were committed
to studying Triassic fossils in different parts of the country. The work done
by these Argentine pioneers, in particular José Bonaparte, became the
starting point for a remarkable increase in our knowledge of Jurassic and
Cretaceous dinosaur faunas and inspired a new generation of research-
ers working mainly under Bonaparte’s direction. Among this group of
active younger paleontologists, Jaime Powell, Luis Chiappe, Guillermo
Rougier, Leonardo Salgado, Jorge Calvo, Rodolfo Coria, Rubén Mar-
tinez, Oscar Alcober, Andrea Arcucci, and Bernardo Gonzalez Riga
figure prominently.
In contrast to Argentina, the development ofvertebrate paleontology
in general and dinosaur studies in particular was slower in other South
American countries. However, Brazil has seen a sustained increase in
discoveries and research, including descriptions of several outstanding
discoveries of Cretaceous dinosaurs published by Llewellyn Ivor Price
between 1948 and 1969. Innovative studies of South American ichnology
were done by Giuseppe Leonardi and important advances in dinosaur
studies are currently being developed by Alexander Kellner, Ismar de
Souza Carvalho, and Max Langer.

XIX
 Activities carried out by foreign paleontologists also contributed to
better knowledge of dinosaur faunas, mainly in Argentina, including
pioneering studies by Richard Lydekker and Friedrich von Huene and
the sustained work by Alfred S. Romer in Triassic rocks of northwest
Argentina. In more recent years, Argentine paleontology has benefited
considerably from the activities conducted by Philip Currie in Creta-
ceous beds of northwest Patagonia, by Paul Sereno in outcrops of the
Ischigualasto Formation, by Patricia and Thomas Rich in Jurassic rocks
of Chubut, by Oliver Rauhut in Jurassic and Cretaceous formations in
the same Argentine province, and by Kenneth Lacovara and Matthew
Lamanna in Cretaceous fossil beds of central Patagonia.
Work carried out in Mesozoic outcrops of South America has resulted
in a remarkable string of discoveries, including more than 50 dinosaur
species (diagnosable on the basis ofderived osteological features), which
have revealed that a rich and complex evolutionary history took place in
this southern continent. The fossil record of South American dinosaurs
covers the Triassic, Jurassic, and Cretaceous periods, revealing remark-
able aspects of their origin and later diversification. We now know that
South America was once populated by dinosaurs of different pedigree,
some ofthem sharply different from their Laurasian counterparts. In com-
parison with dinosaurs from other southern continents, those from South
America are the best represented in terms of number of specimens, qual-
ity of preservation, and diversity of species. The fossil evidence amassed
in Argentina alone (consisting of skeletons, footprints, eggs, embryos,
nests, and skin impressions belonging to the main evolutionary streams
of Saurischia and Ornithischia) constitutes a comprehensive dataset that
is taxonomically more diverse, chronologically better documented, and
phylogenetically more informative than the fossil record obtained in
other continents that formed part of Gondwana (Africa, Madagascar,
India, Antarctica, and Australia).
Deficits of the South American fossil record mainly relate to basic
taxonomical problems (for example, questions about the validity of spe-
cies founded on partial evidence, in particular Cretaceous titanosaurs,
or clades such as stegosaurs and ankylosaurs that are known only from
incomplete specimens). There are geographic sampling problems as
well. Dinosaur discoveries are almost unknown from South American
regions other than Patagonia, northwest Argentina, and southern Brazil.
The stratigraphic representation is also patchy, and the most productive
sedimentary units of South America are still underrepresented in the
number of discovered taxa compared with formations of similar ages of
other parts of the world. For example, the intensively worked Campa-
nian—Maastrichtian Dinosaur Park Formation of western Canada has
yielded remains of nearly 30 different genera of dinosaurs (Weishampel
et al. 2004), double the number of taxa documented in the Bajo de la
Carpa and Anacleto formations, two of the most productive beds of South
America.

XX Introduction
 The purpose ofthis book is to offer an up-to-date and comprehensive
review of the anatomy, systematics, and evolution of South American
dinosaurs within paleogeographic and paleoecological contexts. To carry
out this task, I review information from a variety of sources, most of
them published in Spanish and Portuguese. Firsthand observations of
dinosaur specimens from South America form an important part of the
book, as well as comparisons with forms discovered in other Gondwanan
landmasses. Current understandings of phylogeny, paleoecology, and
paleobiogeography are considered in light of current controversies. Main
geological events (e.g., diastrophism, volcanism, continental breakup) as
well as faunal and floristic changes that occurred in South America are
also briefly examined.

Introduction XXi
THE AGE OF DINOSAURS IN SOUTH AME
RICA
Fig. 1.1. Cladogram
Va Ron (<2 @ @ &
depicting the phylogenetic 3 »e a OP FgeEY s = ES ses s sRed
RS we©
relationships of main e} Oe OM SRP RS OES RO rein ie)
Ss STROM
groups of Archosauria. SN Sm USSaR ae
yeNS a aS eess, asSe
Redrawn from Parrish
(1997).

ORNITHODIRA CROCODYLOTARSI
CRUROTARSI

ARCHOSAURIA
AN OVERVIEW OF THE
ANATOMY AND PHYLOGENETIC
RELATIONSHIPS OF DINOSAURS

This chapter provides the reader with some basic knowledge about how
dinosaurs are phylogenetically related to other reptiles, emphasizing
features of skeletal construction. It will also offer some generalities about
dinosaur anatomy to familiarize the reader with the main morphological
details this book will frequently use in its descriptions of South American
dinosaurs. Finally, the chapter presents a brief historical account of the
systematic interpretations of Dinosauria that highlights the impact of
discoveries in South America on the generation of currently accepted
phylogenetic hypotheses.

Dinosaurs are a diverse lineage of vertebrates that dominated terrestrial What Are
landscapes during most of the Mesozoic Era, a time span of approxi- Dinosaurs?
mately 135 million years. Because birds are a subgroup ofliving dinosaurs,
the group as a whole has lasted some 200 million years. Dinosaurs are
tetrapods, the group of vertebrates that are four-footed land animals. Di-
nosaurs are amniotes, a subgroup of four-footed vertebrates characterized
by internal fecundation and reproduction with a shelled egg. Amniota
includes two main evolutionary streams: Synapsida (the group leading
to the mammals) and Reptilia (the lineage that includes living turtles,
lizards, crocodiles, and birds plus a wide variety of extinct members). The
first members of Reptilia lived during the Carboniferous Period—320
million years ago—near the end of the Paleozoic Era.
Within Reptilia, dinosaurs belong to a subgroup known as Archosau-
ria, which is currently defined by modern systematics to include the last
common ancestor ofthe two extant groups of archosaurs, the crocodilians
and the birds, and all of the descendants of that common ancestor (e.g.,
Sereno 19914; Parrish 1997; Fig. 1.1). Archosauria originated during the
Triassic Period, and the group is subdivided into two main evolutionary
streams: Crurotarsi (a name that refers to a complex construction of ankle
bones involving a movable articulation between the astragalus and the
calcaneum) and Ornithodira (a word meaning “bird neck,” referring to
the S-shaped curvature of the neck). Crurotarsan archosaurs include liv-
ing crocodiles plus a wide spectrum of extinct crocodile-like creatures,
most of them quadrupedal, that prospered during the Triassic Period (e.g.,
ornithosuchians, parasuchians, aetosaurs, rauisuchids). On the other
side, Ornithodira is the archosaur lineage that joins pterosaurs (extinct
flying archosaurs that proliferated during ‘Iniassic, Jurassic, and Creta-
ceous times) and dinosauromorphs (the clade that gathers Mid-Triassic
 dinosaur forerunners such as Lagerpeton and Marasuchus and dinosaurs
themselves, including birds).
Dinosaurs and their kin underwent important transformations of
their locomotor apparatus when they acquired an upright posture of their
hindlimbs, resembling more the shape and posture of the hindlimbs of
living birds than the sprawling posture of living crocodiles or lizards. In
sharp distinction to these other reptile groups, the hindlimbs of a dino-
saur were vertically oriented below the body during both slow walking
and fast running. In other words, dinosaurs carried their bellies far from
the ground. In addition, while in the above-cited living reptiles the foot
contacts the ground with its entire sole, in dinosaurs only the toes rested
on the ground, as in living birds (footprints left by early dinosaurs that
lived around 200 million years ago are similar to the footprints that a
modern chicken leaves in the mud). Dinosaurs and their more immedi-
ate relatives (Lagerpeton, Marasuchus) were among the first reptiles that
abandoned crawling. Dinosaurs preceded by several tens of millions of
years other vertebrates (for example, jerboa rats, kangaroos, hominids)
that would evolve a bipedal carriage.
Bipedalism, which implies freedom of the forelimbs from their an-
cestral locomotory function, evolved in the immediate forerunners of
dinosaurs. The forelimbs of early dinosaurs presumably participated in
tasks other than supporting body weight (for example, handling food
items). Although later in their history many groups of plant-eating dino-
saurs regained a quadrupedal style of locomotion, other forms (in par-
ticular, the meat-eating dinosaurs) retained freed forelimbs, which had
enormous adaptive consequences for the evolution of wings and flying
capabilities in birds.
The word dinosaur means “fearfully great lizard,” and this image ap-
plies well to ferocious creatures like a Tyrannosaurus and Gigantosaurus
or even bigger beasts like Brachiosaurus and Argentinosaurus, which
weighed more than 50 tonnes. Nevertheless, many dinosaurs were tiny
animals, not fearsome giants. Scipionyx from Italy measured roughly 40
centimeters long, while Microraptor, from Cretaceous beds in China,
was just 25 centimeters from its nose to the tip of its tail.

Dinosaur Skeletal For better comprehension ofthe anatomical descriptions of South Ameri-
Anatomy can dinosaurs, I offer some general comments about the skeletal mor-
phology of these animals. The main sources of information for this part
of the book are Romer (1956) and Holtz and Brett-Surman (1997).
The skeleton of a tetrapod, dinosaurs included, can be divided into
the following sections from a topological point of view: the skull, the ver-
tebral column (including the ribs and haemal arches), the pectoral girdle
and forelimbs, and the pelvic girdle and hindlimbs. All the elements of
the skeleton, with the exception of the skull (“cranium’” in Latin), are col-
lectively called the postcranium (the part of the skeleton that is posterior
to the cranium).

The Age of Dinosaurs in South America

 Fig. 1.2. Sauropodomorph
dinosaur Plateosaurus.
Skull and jaw in (A) lateral,
(B) dorsal, (C) ventral, and
(D) occipital views. (E) Left
jaw in medial view. Re-
drawn from Galton and
Upchurch (2004a).

pra

Anatomy and Phylogenetic Relationships of Dinosaurs 5

 The skull is composed of paired bones repeated as mirror images of
each other on both sides of the axial plane (Fig. 1.2). There are also single
bones that lie along the midline of the skull. The skull houses the main
sense organs (nose, eyes, ears) and the brain. Two bones of the ventral
margin of the skull may have teeth: the premaxilla and the maxilla (to-
gether forming the upper jaw). In some dinosaurs (e.g., ornithischians,
titanosaurid sauropods, ornithomimid theropods, and living birds), teeth
are absent on premaxillae, on maxillae, or on both and a horny beak
covered the margins of the mouth.
The premaxilla is ventral to the nasal opening, and the maxilla is ven-
tral to the antorbital fenestra, an opening that is present in most dinosaurs
(and other archosaurian reptiles). Some dinosaurs (the ornithischians)
had reduced or lost the antorbital opening, while in others (particularly
in theropod dinosaurs) the antorbital fenestra is wide and was associated
with other pneumatic openings (the maxillary and the promaxillary open-
ings). The paired premaxillae and the maxillae articulate dorsally with
the paired nasals (sometimes fused in a single piece of bone) to surround
the nasal passage.
The eye socket, or orbit, is surrounded ventrally by the jugal, rostrally
(toward the snout) by the lacrimal, caudally (toward the tail) by the
postorbital, and dorsally by the frontal. The frontal is a paired bone that
meets on the midline of the skull, roofing part of the brain cavity. The
frontals articulate rostrally with the nasals, caudally with the parietals
(which also contribute to the walls of the braincase), and laterally with the
bones that encase the orbit (e.g., the lacrimal and postorbital). In the rear
end of the skull, a pair of openings, the temporal fenestrae, that housed
the jaw muscles: the lateral temporal fenestrae (located on both sides of
the skull) and the supratemporal fenestrae (lying on the dorsal surface
of the skull). These pairs of fenestrae evolved in the common ancestor
of diapsids, the group of reptiles that includes lepidosaurs (e.g., lizards
and snakes) and archosaurs (e.g., crocodiles, pterosaurs, and dinosaurs,
including birds).
In the posterior end of the skull there is a left and right quadrate,
a large bone that served for articulation with the lower jaw, which may
have teeth or a horny sheath. The tooth-bearing bone of the mandible
is the dentary, and the left and right dentaries form the symphysis that
unites the two sides of the jaws. Dinosaur teeth, when present, were
always growing and could always be replaced. They have different mor-
phologies and sizes depending on the dinosaur group. In theropods,
teeth are conical, transversely compressed, serrated, and caudally curved.
In sauropodomorphs, teeth are leaf-shaped with margins that might or
might not be serrated, although some derived sauropods had pencil-like
teeth. Ornithischians had a variety of types of teeth. Basal ornithischians
had a spatulate and serrated tooth but more-derived lineages evolved a
prismatic-shaped tooth that had no serrations.
The dentary articulates medially (toward the midline of the ani-
mal) with the splenial and caudally with three bones: dorsally with the

The Age of Dinosaurs in South America

 A cervicals
dorsals sacrais Fig. 1.3. Theropod
dinosaur Allosaurus: (A)
Silhouette of the body
indicating regions of the
vertebral column and tho-
rax. (B) Cervical 7 in lateral
view. (C-E) Dorsal 10 in (C)
left lateral, (D) cranial, and
(E) caudal views. (F) Dorsal
Neural rid in cranial view. (B—F)
Neural arch
Redrawn from Madsen
arch
(1976).

Centrum Centrum

surangular and ventrally with both the angular (on the outer side of the
mandible) and the prearticular (on the inner side of the jaw). These three
bones meet caudally with the articular, the bone that has a hinge-joint
articulation with the quadrate bone ofthe skull. The angular, surangular,
prearticular, and articular bound a wide fossa (open space) that housed
the adductor muscles that closed the jaw.
The braincase is made up of several tightly sutured bones. The
braincase proper lies beneath the roof bones ofthe outer skull. The spinal
cord exits from the braincase through a great opening called the foramen
magnum located on the posterior end ofthe skull. The foramen magnum
is dorsally and ventrally bounded by unpaired bones, the supraoccipital
and the basioccipital, respectively, and laterally by the paired exoccipitals.
The basioccipital forms a bulbous prominence, the occipital condyle, for
articulation with the vertebral column (i.e., the atlas and axis).
The vertebral column of dinosaurs can be divided into cervical
(neck), dorsal (trunk), sacral (hip), and caudal (tail) vertebrae (Fig. 1.3A).
Each of these sections is composed of vertebrae of different morpholo-
gies and the number of vertebrae in each module is variable, depending
on the group of dinosaur considered. In general, cervicals are low (with
a relatively small top-to-bottom dimension) and elongate (from front to
back). Dorsal vertebrae are more robustly constructed. The dinosaur
sacrum consists of two or more vertebrae, sometimes fused into a single
element. The sacrum attaches on either side to the left and right ilia,
which constitute the upper bones of the pelvic girdle (see below). The
caudal series is composed of several vertebrae that tend to diminish in
size toward the tailtip.
A single vertebra is formed by a large spool or cylindrical structure,
the centrum, which supports the neural arch dorsally (Fig. 1.3B-E). Both
the centrum and the neural arch bound the neural canal, which houses

Anatomy and Phylogenetic Relationships of Dinosaurs

 the spinal cord. The centrum has cranial and caudal surfaces for ar-
ticulation with contiguous centra. Articular surfaces may be flattened
(amphyplatyan) or slightly concave (amphicoelous), although in some
special cases the cranial surface is concave but the caudal end is convex
(procoelous, as in the tail vertebrae of titanosaurid sauropods) or the re-
verse condition occurs (the cranial surface is convex and the caudal end
is concave, a condition termed opisthocoelous, as in the neck vertebrae
of some derived theropod and sauropod dinosaurs). The lateral surface of
the centra may be perforated by pneumatic pores known as pleurocoels.
Such foramina connect with the interior of the vertebral centrum and may
have served as passage for air sacs. Such perforations occur in the cervical
vertebrae of most saurischian dinosaurs and are also present in the dorsal,
sacral, and caudal vertebrae of some derived theropods and sauropods.
Centra that are perforated by pneumatic pores are internally hollow.
From the neural arch projects a dorsal neural spine, which offers
surfaces of attachment for muscles and tendons of muscles running over
the vertebral column and ligamentous connections between successive
neural spines. Both cranially and caudally there are pairs of finger-like
projections called zygapophyses. The prezygapophyses (i.e., the ante-
rior zygapophyses) have articular surfaces oriented upward and slightly
inward and serve as support of the postzygapophyses (i.e., the posterior
zygapophyses), which have articular surfaces oriented downward and
slightly outward. The trunk vertebrae of saurischian dinosaurs evolved
additional surfaces for vertebral support: the hypantrum and the hypos-
phene. The hyposphene is a posteriorly directed median wedge project-
ing below the postzygapophyses that inserts into the hypantrum, a narrow
Fig. 1.4. Girdles and limb space between the prezygapophyses.
bones of several dinosau- The cervical vertebrae of most dinosaurs exhibit prominences on
rian taxa. (A) Allosaurus the dorsal surface of the postzygapophyses, called epipophyses, to which
fragilis, scapular girdle
and forelimbs in cranial
neck muscles attach.
view. (B) Brachiosaurus The cervical and dorsal vertebrae have conspicuous apophyses pro-
brancai, trunk and sacral jecting from both sides of the neural arch and centrum. Such projections
vertebrae articulated with are termed diapophyses (extending laterally from the neural arch) and
pectoral and pelvic girdles
parapophyses (present on the cranial margin of cervical and dorsal ver-
and fore- and hindlimbs
in left lateral view. (C, D) tebrae). These paired apophyses served for articulation with the double-
Pelvic girdles in left lateral headed cervical and dorsal ribs, which themselves proximally split into
view of (C) the basal a dorsal process, the tubercle, to attach to the diapophysis, and a ventral
saurischian Plateosaurus process, the capitulum, to attach to the parapophysis (Fig. 1.3F). In archo-
and (D) the basal ornithis-
saurian reptiles, including dinosaurs, the parapophysis migrates upward
chian Lesothosaurus. (E)
Allosaurus fragilis, pelvic and backward from the centrum to the neural arch from one vertebra to
girdle and hindlimbs in the next along the presacral series, finally lying close to the diapophysis
cranial view. (A) Redrawn so that diapophysis and parapophysis may fuse together into a single-
from Carpenter (2002). headed structure. Ribs articulated to the dorsal vertebrae form a ribcage
(B) Redrawn from
(thorax) around the vital organs. Along the belly of some dinosaurs are
Christiansen (2000). (E)
Composite reconstruction gastralia, or “belly ribs,” which strengthened the ventral side of the animal
based on Gilmore (1920) and acted as a girdle to hold in the viscera but probably also contributed
and Madsen (1976). dynamically in the breathing process.

The Age of Dinosaurs in South America

 mtc Ill
mtc Il

d
B
sc
cor

ry

car
manus | mtc
phs

The caudal vertebrae exhibit laterally projected transverse processes

from the neural arch that do not articulate distally with ribs (which
are absent in the caudal series). Ventral to the caudal vertebrae are the
chevrons, or haemal arches. These are Y-shaped in cranial aspect and
articulate below the connection between two contiguous caudal centra.

Anatomy and Phylogenetic Relationships of Dinosaurs 9

 Some dinosaurs have ossified tendons running along different por-
tions of the vertebral column. This is the case for most ornithischians,
in which a net of ossified tendons runs across the neural spines of the
dorsal, sacral, and caudal vertebrae as well as across the haemal arches of
the tail. Ossified tendons also evolved in the neck of certain titanosaurid
sauropods and in the tail of some clades of theropod dinosaurs (e.g.,
abelisauroids, dromaeosaurids).
The pectoral (or shoulder) girdle (Fig. 1.4A and B) attached through
ligament and muscle connections to the anterior region of the ribcage.
It is composed of asomewhat strap-like elongate scapula that articulates
with an elliptical or rectangular-shaped ventral element, the coracoid.
The junction of the scapula and coracoid (along their caudal margin)
forms the glenoid cavity, a deep concavity for articulation with the proxi-
mal element of the forelimb (the humerus). Running over the cranial
edges of both scapulae and coracoids were the paired clavicles, which in
most predatory dinosaurs fused into a single bone, the furcula (the avian
“wishbone”). The coracoids of both sides connect along the midline of
the body with a paired sternum, or “breastbone,” the shape and size of
which varies among groups of dinosaurs.
Dinosaur forelimbs (Fig. 1.4A and B) are made up of three main
bones: the humerus, the ulna, and the radius. The proximal humerus
joins the two forearm bones at the elbow. In most dinosaurs, the humerus
has a pronounced and cranially directed deltopectoral crest, which served
for the attachment of pectoral muscles that originated from the animal’s
chest. The ulna is larger and more posterior than the radius. It has a proxi-
mal prominence, the olecranon process, which is variably developed for
attachment of extensor muscles that straightened the limb. Both the ulna
and the radius articulate with the hand, or manus. The proximal compo-
nents of the manus are a series of small carpal (wrist) elements articulated
distally to elongate metacarpals, which are designated by Roman numerals
I to V. Lis the most medial (inner) and V is the most lateral (outer). Each
digit is numbered to match its corresponding metacarpal. The individual
bones of the digits are the phalanges. The distalmost (toward the finger-
tip) of the phalanges are the unguals, which supported horny claws or
hooves. Primitively, dinosaurs were bipedal reptiles with elongate and
slender forelimbs about half as long as the hindlimbs and hands suited
for manipulation rather than for progression. As a quadrupedal posture
was acquired, mainly by massively constructed dinosaurs (such as sauro-
pods, ankylosaurs, stegosaurs), the forelimb bones became robust but the
hands retained an almost unreduced digit count (Fig. 1.4B). The ungual
phalanges of quadrupedal dinosaurs were lost or were transformed into
hooves. In dinosaurs that retained a bipedal posture, rnainly the meat-
eating theropods, the number of outer digits of the manus was reduced
and the unguals became proportionally large and trenchant. Theropod
forelimbs manifested dissimilar trends; in tyrannosaurids and abelisaurids
they were strongly reduced, while in some derived avian-like theropods
(deinonychosaurs), particularly flying birds, the forelimb was elongated.

The Age of Dinosaurs in South America

 The pelvic girdle on each side of the body is composed of three
bones: a dorsal bone (the ilium); an anteroventral bone (the pubis); and
a posteroventral bone (the ischium) (Fig. 1.4C and D). At the junction
of these three pelvic bones is a perforated depression, the acetabulum,
which receives the femoral head. The left and right ilia are connected
to the sacral vertebrae. The ilium, the pubis, and the ischium served as
expanded surfaces for muscle attachment, mainly the origin of several
hindlimb protractors (to pull the limb forward) and retractors (to pull the
limb backward) and the muscles of the trunk and tail. The distal extremi-
ties of both pubes and ischia may have served as points of support of the
body while the animal was in a resting position.
Most of the saurischian dinosaurs retained from their archosaur an-
cestors a “triradiate,” or propubic, pelvis in which the pubis was projected
forward and downward (Fig. 1.4C). In contrast, from their early origins,
ornithischian dinosaurs manifested a backward-oriented (opisthopubic)
pelvis (Fig. 1.4D). However, in derived saurischians such as the coelu-
rosaurian therizinosaurids, mononykids, dromaeosaurids, and birds, the
pubis also became retroverted, independently acquiring a condition that
superficially resembles that of the omithischian dinosaurs.
The hindlimb bones are the femur, the tibia, and the fibula (Fig.
1.4B, E). The femur articulates proximally with the pelvic acetabulum
through a medially offset femoral head and joins distally at the knee to
the tibia and the fibula. The dinosaur femur has different prominences for
muscle attachment: the anterior trochanter proximally (to attach fibers
of protractor muscles originated on the ilium) and the fourth trochanter
at caudal midshaft (to attach retractor muscles that originated from the
ilium and the tail). The tibia constitutes the larger and more medial bone
ofthe lower leg and the fibula is the thinner and more lateral bone. Proxi-
mally, the tibia has a rectangular-shaped cranially projecting cnemial
crest. Distally, both the tibia and the fibula articulate with two tarsals (the
bones of the ankle), respectively the astragalus and calcaneum, which
contact distally with a small number of disc-shaped distal tarsals. The
tarsal bones articulate with the elongate metatarsals (numbered I to V
in medial-to-lateral fashion), and the metatarsals articulate distally with
their respective digits. The digits, metatarsals, and tarsals are collectively
called the foot, or pes.

English naturalist Richard Owen coined the name Dinosauria in 1841 for A Brief Historical
a group ofland-living reptiles oflarge size that exhibited features that were Account of the
more advanced than extant reptiles in the construction of the vertebral Systematics of
column and hindlimbs. After that time, dinosaur discoveries multiplied Dinosauria
considerably and it became clear that Dinosauria did not constitute a
homogeneous group but was a deeply diversified taxon with a wide vari-
ety of adaptations. Paleontologists became interested in bringing order
to this diversity and proposed numerous classifications. In 1887, British
paleontologist Harry Seeley made the case that dinosaurs could be sorted

Anatomy and Phylogenetic Relationships of Dinosaurs

into two different taxa: Saurischia for dinosaurs with a cranioventrally
projected pubis, and Ornithischia for dinosaurs with a caudoventrally
oriented pubis. These important distinctions in pelvic construction led
Seeley to argue that Saurischia and Omithischia were not closely related
to each other. Seeley reinterpreted the features that Owen recognized as
characterizing Dinosauria as superficial resemblances due to functional
similarities related to hindlimb posture. Over the course of the twentieth
century, Seeley’s interpretations gained wide acceptance and the name
Dinosauria began to be used as an informal term for a polyphyletic assem-
blage of archosaurian reptiles. Thus, the respective origins of saurischians
and ornithischians were sought in totally different ancestral groups.
The sharp morphological gap between dinosaurs and basal archosaurs
as well as the lack of a truly phylogenetic methodological approach cre-
ated confusion about dinosaur origins and interrelationships. Moreover,
during the 1970s the idea of amarked polyphyletism became predominant
among authors to the point of almost destroying not only Dinosauria but
also Saurischia and Theropoda as monophytetic groups (e.g., Thulborn
1975). However, this changed during that decade, mainly due to the avail-
ability of new fossil material from the Triassic of Argentina. Discoveries of
dinosaur-like archosaurs such as Lagerpeton chanarensis and Marasuchus
lilloensis (Romer 1971, 1972; Bonaparte 1975, 1984a) made it possible to
study, for the first time, advanced reptiles that were potential ancestors
of saurischians and ornithischians. Furthermore, more information on
early dinosaurs, such as Pisanosaurus from Argentina, Anchisaurus from
the United States, and Syntarsus from Zimbabwe, became available
(Raath 1969; Galton 1976; Bonaparte 1976). This new evidence provided
sufficient grounds for reevaluating the long-held belief that Dinosauria
was not a natural group (e.g., Seeley 1887, 1888; Baur 1891; von Huene
1914; Colbert 1964; Romer 1971).
Robert Bakker and Peter Galton (1974) provided the concept that
saurischians and ornithischians were closely related phylogenetically and
supported a hypothesis that both lineages descended from a common
archosaurian ancestor. In sharp contrast with Seeley, whose method-
ological procedure emphasized the anatomical differences between sau-
rischians and ornithischians, Bakker and Galton paid special attention to
the shared characteristics of these two groups. These authors (like Owen
before them) observed that early saurischians and ornithischians were
bipedal and that their hindlimbs adopted a vertical posture, constitut-
ing features that were absent in other archosaurian reptiles. Additional
characteristics Bakker and Galton listed in support of their hypothesis
included a completely open acetabulum, a femoral head that was distinct
from the rest of the bone, a reduced fibula, a mesotarsal type of tarsus,
and a foot with a digitigrade stance.
In 1975, Argentine paleontologist José Bonaparte published an exten-
sive study of Marasuchus (following Sereno and Arcucci [1994], the name
Lagosuchus talampayensis is regarded as a nomen dubium and the name
Marasuchus lilloensis is used here to refer the holotype of “Lagosuchus”

The Age of Dinosaurs in South America

lilloensis; Fig. 1.5) that demonstrated its resemblances to saurischians Fig. 1.5. Reconstructed
and ornithischians, thus filling an important morphological gap between skeleton of Marasuchus
early dinosaurs, which were mostly bipedal and digitigrade, and basal lilloensis. Illustration by
J. Gonzdlez.
archosaurs, which were mostly quadrupedal and plantigrade. Bonaparte’s
research offered strong support to Bakker and Galton’s hypothesis of the
common origin of Saurischia and Ornithischia.
The hypothesis that dinosaurs were monophyletic gained adherents
during the 1980s, and at present no alternative hypothesis (e.g., that dino-
saurs are nota clade) has been offered on the basis of derived characteris-
tics. In recent years cladistic analysis has been extensively applied to the
archosaur systematic. Gauthier (1986) offered an exhaustive analysis of
characteristics of saurischian dinosaurs, supporting the idea that Dinosau-
tia, Saurischia, and Theropoda were monophyletic. He argued that the
clade Ornithodira encompassed the common ancestor of Pterosauria and
Aves and all ofits descendants. Novas (1992a) found evidence in support
of amonophyletic group composed of Marasuchus plus Dinosauria, for
whom the name Dinosauriformes was coined.

Gauthier (1986) and Sereno (1ggia) studied in depth the monophyly of Dinosaur
Omithodira (a group including Pterosauria, Lagerpeton, and Dinosauri- Forerunners
formes and their most recent common ancestor). Sereno and Novas (1990),
Sereno (19914), and Novas (1992a) found evidence to support the notion
that Pterosauria were outgroups of a clade composed of Marasuchus plus
Dinosauria (see Fig. 1.1). Also, the phylogenetic placement of two taxa was
clarified: the bizarre archosaur Lagerpeton (Arcucci 1986, 1997; Sereno
and Arcucci 1993) is now considered the sister taxon of the Dinosauri-
formes (Sereno and Novas 1990; Sereno 1gg1a), and Pseudolagosuchus
(formerly placed within “Lagosuchidae”; Arcucci 1987) was more recently
interpreted as the immediate sister group of Dinosauria (Novas 1992a).
Pterosauria and Lagerpeton are now interpreted as successive ornithodiran
outgroups of Dinosauriformes (e.g., Sereno 19914).
The clade Dinosauriformes (Novas 1992a) is defined to include the
common ancestor of Marasuchus and Dinosauria and all the descendants
of that common ancestor (Fig. 1.1). Several synapomorphies concerned

Anatomy and Phylogenetic Relationships of Dinosaurs

Fig. 1.6. Cervical and
dorsal vertebrae of Mara-
suchus lilloensis. From
Bonaparte (1975). Re-
printed with permission of
the author and Fundacion
Miguel Lillo.

Fig. 1.7. The hindlimb

of dinosauriforms. (A-G)
Proximal end of femora
of several archosaur taxa:
(A, D) Right femur of La-
gerpeton. (B, E) Left femur
(reversed) of Marasuchus.
(C, F, G) Right femur of
Herrerasaurus, in (A-C)
proximal, (D-F) caudal, with neck anatomy and hindlimb morphology have been recognized to
and (G) cranial views. (H—J) support the hypothesis that Dinosauriformes are monophyletic. Mara-
Proximal end of left tibia suchus shows an incipient sigmoid curvature of the neck but it does not
in lateral view of several have the strongly S-shaped neck ofdinosaurs, including birds (Bonaparte
dinosauriform taxa: (H)
Marasuchus, (l) Herrera-
1975; Gauthier 1986; Fig. 1.6).
saurus, and (J) Dryosaurus. The hindlimbs of Marasuchus are notably similar to those of primi-
(K—M) Distal end of left tive dinosaurs, including a trochanteric fossa on the proximal articular
tibia in lateral view of surface of the femoral head (Fig. 1.7A-F), which served for sliding over
several dinosauriform taxa:
a prominent pelvic antitrochanter (located on the posterior portion of
(K) Pseudolagosuchus, (L)
Herrerasaurus, and (M) acetabulum; Fig. 1.8A—C). Marasuchus and Dinosauria also share an ante-
Dryosaurus. (NP) Distal rior trochanter on the femur, which conforms to a subvertical prominence
view of left tibia of several that is continuous distally with a subhorizontal ridge (the trochanteric
dinosauriform taxa: (N) shelf; Fig. 1.8F and G). The trochanteric shelf constitutes a prominent
Pseudolagosuchus, (O)
Herrerasaurus, and (P)
posterodistally oriented ridge on the lateroproximal surface of the fe-
Dryosaurus. (Q-T) Left mur that is uniquely documented among dinosauriform archosaurs (e.g.,
astragalus in proximal view Marasuchus, Pseudolagosuchus, Silesaurus, Herrerasaurus, Syntarsus).
of several dinosauriform Marasuchus and early dinosaurs also share important similarities in
taxa: (Q) Marasuchus, (R)
the shape of the tibia: on the proximal end of this bone, a cnemial crest
Pseudolagosuchus, (S) Her-
rerasaurus, and (T) Scutel-
developed that is square-shaped in side view (Fig. 1.7H). In addition, the
losaurus. Modified from distal end of the tibia is quadrangular and is laterally notched by a groove
Novas (1996a). © Copy- that runs longitudinally along the distal portion of the tibial shaft (Fig.
right 1996 the Society of 1.7K-O). This groove is expressed in distal aspect as a U-shaped notch
Vertebrate Paleontology.
that is bounded by the facet for the ascending process of the astragalus
Reprinted and distributed
with permission of the and the posterior descending process of the tibia. In dinosaurs this notch
Society of Vertebrate on the tibia correlates with the differentiation of amore complex dorsal
Paleontology. surface of the astragalus (Fig. 1.7L, O, and S).

14 The Age of Dinosaurs in South America

 Some dinosauriforms (e.g., the Mid-Triassic Pseudolagosuchus from
Argentina and the Late Triassic Silesaurus from Poland; Dzik 2003) ex-
hibit features that are closer to the dinosaurian condition than those pres-
ent in Marasuchus. For example, dinosaurs share with Pseudolagosuchus
and Silesaurus an elongate pubis that is equivalent to 70 percent or more
of femoral length, in contrast with more basal ornithodirans (e.g., Ptero-
sauria, Lagerpeton, Marasuchus), in which the pubis is shorter (nearly

Anatomy and Phylogenetic Relationships of Dinosaurs

Fig. 1.8. Pelvic girdles as long as the ischium) and is approximately 50 percent of the femoral
of dinosauromorph taxa: length (Fig. 1.8A—C).
(A) Lagerpeton. (B)
Pseudolagosuchus and Dinosauria also developed changes in astra-
Marasuchus. (C) Her-
rerasaurus, in right lateral galar anatomy, consisting of the acquisition of apyramid-shaped ascend-
view. (D-F) Right ilium of ing process and a caudal subvertical facet and an elliptical depression
Herrerasaurus in lateral (D), behind the process (Fig. 1.7S). In Archosauria ancestrally the ascending
ventral (E), and caudal (F) process of the astragalus constitutes a craniocaudally oriented ridge that
views. (G-l) Right ilium of
Torvosaurus in (G) caudal,
separates the tibial and fibular facets (Novas 198g9a). This condition was
(H) lateral, and (|) ventral retained in Lagerpeton and Marasuchus (Fig. 1.7Q). Pseudolagosuchus,
views. Modified from in contrast, has a pyramid-shaped ascending process resulting from de-
Novas (19966). © Copy- velopment of a posterior, subvertical, non-articular surface. Behind this
right 1996 the Society of
surface is an elliptical depression on the proximal face of the astrag-
Vertebrate Paleontology.
Reprinted and distributed alus. Dinosaurs are distinguished from other dinosauriforms in that the
with permission of the posterior surface of the ascending process articulates with the posterior
Society of Vertebrate process of the tibia, the latter being transversely broader than in basal
Paleontology. Dinosauriformes. In addition, the posterior process of the tibia con-
ceals the elliptical depression present on the dorsal astragalar surface. In

The Age of Dinosaurs in South America

Dinosauria, the ascending process ofthe astragalus shows important mor-
phological variation: in ornithischians (except Pisanosaurus; Bonaparte
1976; Novas 198ga) the ascending process is dorsoventrally reduced and
anteroposteriorly flattened (e.g., Scutellosaurus, Dryosaurus; Colbert 1981;
Galton 1981), while in tetanurine theropods the ascending process forms
an anteroposteriorly flattened, tall, and broad wedge (Gauthier 1986).
Nevertheless, all dinosaurs share a caudal facet on the ascending process
that articulates with the cranial surface (i.e., the astragalar surface) of the
distal tibia (Novas 1989a).

Dinosaurian diagnostic features (i.e., derived features that evolved in the Characteristics
common ancestor of Saurischia and Ornithischia but are absent in their That Distinguish
immediate sister groups) include several anatomical transformations in Dinosauria from
the head, the neck, and the pelvic girdles. Other Archosaurs
In the back of the skull, pseudotemporal muscles spread beyond the
boundaries of the supratemporal openings, partially invading the dorsal
surface of the frontal bones (Fig. 1.2B, mps), a modification that may
have increased the force of jaw closure. The neck vertebrae of dinosaurs
manifested a more pronounced inclination ofthe articular surfaces of the
cervical centra, conferring a distinctive sigmoid curve to the neck, thus
resembling the avian condition more closely. At least some ofthe cervical
vertebrae developed epipophyses above the postzygapophyses.
With the exception of Staurikosaurus, Guaibasaurus, and Saturna-
lia, in which the sacrum is apparently formed by two sacrals, most of
the early dinosaurs increased the sacral count to three by incorporating
a new vertebra from the presacral series (e.g., Welles 1984; Novas 1996a;
Fig. 1.9). The incorporation of a new sacral (from the dorsal vertebrae)
involved craniocaudal shortening of the last dorsal vertebrae and origi-
nal sacrals 1 and 2. This resulted in placement of the last dorsal vertebra
clearly behind the tip of the pubic pedicle of the ilium. The first sacral
ribs accompanied the craniocaudal shortening of its corresponding cen-
trum, articulating in a more caudal position than ancestrally. As a result
of this transformation, the last trunk vertebra was incorporated into the
sacrum (it became the first dorsosacral) and the perpendicular orientation
of the primordial sacral ribs allowed the articulation of the dorsosacral
ribs to the ilium. Axial shortening of the last dorsal and primordial sacral
vertebrae probably constituted the most important process that made
incorporation of more vertebrae to the sacrum possible, at least during
early dinosaur evolution. The sacral count increased independently in
Theropoda, Ornithischia, and Sauropodomorpha, in all of which the
sacrum has at least five vertebrae.
Dinosaurs have a deltopectoral crest (Fig. 1.4A and B) with the distal
corner positioned down the shaft away from the head of the humerus ata
distance greater than 30 percent of the maximum length of the humerus
(e.g., Bakker and Galton 1974; Sereno and Novas 1992; Sereno, Forster,
Rogers, and Monetta 1993; Sereno 1994).

Anatomy and Phylogenetic Relationships of Dinosaurs

 D
— pup

cd1 us ap di5 d14

d15 d14
F

aaa
$2 s1 dsai1
s2 s1 dsa1 d15
(s3) (s2) (s1)

cdsi S2 81 dsa1 dsa2 d15

cds1 s2 s1 dsa1 dsa2 d15
(s5) (s4) (s3) (s2) (s1)

Fig. 1.9. Ornithodiran Dinosaurs are distinguished from other dinosauriforms by a perfo-
sacra showing relative po- rated acetabulum (Fig. i.8C, shown in black), the result of the opening
sition of the dorsal, sacral,
of the medial acetabular walls of the ilium, pubis, and ischium. The
and caudal vertebrae
with respect to the ilium opening of the acetabulum is relatively small in basal dinosaurs (e.g.,
in several dinosauriform Guaibasaurus, Saturnalia, Staurikosaurus, Herrerasaurus, Lesothosau-
taxa. (A, B) Marasuchus. rus) but is larger in more-advanced dinosaurs (e.g., sauropodomorphs,
(C, D) Herrerasaurus.
theropods). The dinosaur pelvis is also characterized by a brevis shelf that
(E, F) Riojasaurus. (G, H)
Hypsilophodon. (A, C, E,
consists ofadistinct and prominent shelf on the posterolateral margin of
G) lateral view of sacral, the iliac blade that flares laterally above the posteroventral iliac margin
dorsal, and caudal verte- (Fig. 1.8D-I). Both the brevis shelf and the posteroventral iliac margin
brae (silhouette of ilium bound a ventral fossa that cradled the caudifemoralis brevis muscle, a
indicated by solid line).
femoral retractor. Most theropods, ornithischians, and early sauropodo-
(B, D, F H) dorsal view of
sacral, dorsal, and caudal morphs exhibit a well-developed brevis shelf and a transversely wide and
vertebrae (only the left dorsoventrally deep brevis fossa. Only Dinosauria among ornithodirans
ilium and left half of verte- have a brevis shelf and a fossa, since in basal forms (e.g., Pterosauria,
bral column is indicated). Lagerpeton, Marasuchus) the postacetabular portion of the iliac blade is

The Age of Dinosaurs in South America

transversely compressed, with the posteroventral margin running from Not to scale. The counting
the ischiadic peduncle toward the posterior end of the blade (Novas of dorsosacral vertebrae
1992a, 1996a). (dsa1, dsa2) starts with the
vertebrae attached to the
The dinosaur ischium has a slender shaft and a ventral keel (obtura-
ilium that articulates with
tor process) that is limited to the proximal third of the bone (Figs. 1.4C-D the original sacral 1 (s1).
and 1.8A—C). In basal archosaurs the ischium forms a plate-like structure Counting of caudosacral
that is distal to the acetabular region. This condition was retained in vertebra (cds1) starts with
Ornithodira ancestrally because it is present in Pterosauria, Lagerpeton, the vertebra attached to
the ilium that articulates
and Marasuchus, in which the ischium has a proximodistally expanded with the original sacral 2
and ventrally projected obturator process. In contrast, ornithischians, (s2). The count of sacrals
sauropodomorphs, and theropods share a distally elongate ischiadic shaft, based on vertebrae that
with the obturator process that is limited almost entirely to the proximal articulate with the ilium is
indicated parenthetically
third of the bone.
below the nomenclature
The hindlimb bones of early dinosaurs have several modifications (i.e., original sacrals 1 and
compared to more-basal dinosauromorphs. The proximal femur of cruro- 2, dorsosacrals 1 and 2,
tarsan archosaurs and basal ornithodirans (e.g., Lagerpeton, Marasuchus, and caudosacral 1). Light
stippling marks the cen-
and Pseudolagosuchus) shows a well-developed tuberosity on the caudal
trum of s2; dark stippling
surface of the head (Fig. 1.7A, B, D, and E). This tuberosity externally marks the transverse pro-
bounds the groove for a ligament attached to the femoral head and in cess and rib of s2 and its
basal archosaurs has a rounded and transversely wide prominence. The place of articulation with
tuberosity is placed nearly halfway between the medial and lateral mar- the ilium. Cross lines mark
$1 and its corresponding
gins of the proximal end ofthe femoral head. In Dinosauria, this tuberos-
transverse process, rib,
ity is significantly reduced and has a slight prominence that is medially and place of attachment
placed (Fig. 1.7C and F). Another modification in the proximal end to the ilium. Dashed lines
of the dinosaurian femur is related to the development of a prominent indicate the articulation
proximally projected anterior trochanter to attach femoral protractors. area of the first dorsosacral
rib. Modified from Novas
Interesting modifications are detected in the tibial-astragalar articula- (1996a). © Copyright 1996
tion of basal dinosaurs. In basal archosaurs the tibia articulates with the the Society of Vertebrate
proximal surface of the astragalus but does not extend laterally beyond Paleontology. Reprinted
the ascending process of the astragalus. Consequently, there is no clear and distributed with per-
mission of the Society of
distinction between the posterior process of the tibia and the remainder
Vertebrate Paleontology.
of the distal articular surface of the bone. This condition is retained in
Dinosauriformes ancestrally, since it is seen in Marasuchus and Pseudola-
gosuchus, in which the distal end of the tibia is transversely convex and
posterodistally inclined (Fig. 1.10). In contrast, the tibia in dinosaurs,
although quadrangular in basal dinosaurs (e.g., Herrerasaurus, Pisano-
saurus; Novas 1989a, 1994), covers the ascending process posteriorly,
resulting in the articulation of the latter beneath the craniolateral corner
of distal articular surface of the tibia (Fig. 1.7L, O, and S). The distal
tibial articular surface becomes topographically complex and it is pos-
sible to distinguish an anterolateral facet for the ascending process of the
astragalus and a ventrally projected posterior process. The latter covers
posteriorly the ascending process of the astragalus and its dorsal surface
behind the ascending process. The dorsal astragalar surface that is poste-
rior to the ascending process becomes enlarged toward the rear, and the
posterior margin of the bone becomes straight in dorsal view (Fig. 1.75
and T). By contrast, in Pseudolagosuchus this portion of the astragalus

Anatomy and Phylogenetic Relationships of Dinosaurs

 is deeply notched in dorsal view and is a non-articular surface. Also, in
the dinosaur astragalus, a deep elliptical basin is present immediately
posterior to the ascending process (F'ig. 1.7S).
Two more modifications occurred in the tarsal bones of basal dino-
saurs. The proximal surface of the calcaneum became concave, creat-
ing a tighter articulation with the distal end of fibula (Fig. 1.D). In
contrast, in Pseudolagosuchus and Marasuchus the calcaneum has a
hemicylindrical proximal condyle for articulation with the fibula (Fig.
1.nA and C). Also, distal tarsal 4 of dinosaurs acquired a proximodistally
flattened condition and a triangular-shaped contour in proximal view
(Fig. 1.11K—M), differing from the more block-like distal tarsal 4 of more
Fig. 1.10. Right tibia, basal ornithodirans (Fig. 1.11E-J).
fibula, and tarsus of From the available evidence, the early evolution of the Ornithodira
Marasuchus in caudal as- manifested a sustained improvement in locomotor capabilities. Bipedal
pect. From Novas (1989a). and digitigrade postures were acquired by the ancestral ornithodiran
Reprinted with permission
species (Gauthier 1986) and were inherited by dinosauromorphs, includ-
of The Paleontological
Society and the Society for ing Dinosauria ancestrally. Many of the synapomorphies of Ornithodira
Economic Paleontology (Sereno 1991a) pertain to the hindlimb. Dinosauromorphs inherited
and Mineralogy. such adaptations, evolving transformations in the pelvic and hindlimb
skeleton that are also related to locomotion. On the basis of studies
made by Sereno (1gg1a) and Sereno and Arcucci (1994), features of the
tarsus and pes constitute 70 percent of the apomorphies diagnostic of
Dinosauromorpha.
Fig. 1.11. Fibulae, tarsals, Moreover, although the synapomorphies that diagnose Dinosauri-
and metatarsals of several formes include a strong sigmoid neck curvature, most of the evolutionary
ornithodiran taxa: (A)
Right fibula and calcaneum
novelties pertain to the locomotor apparatus. In early dinosauriforms,
of Marasuchus in lateral some outstanding skeletal modifications include the remodeling of the
view. (B) Right fibula of femoral head as a trochanteric fossa developed, which probably restricted
Marasuchus in lateral femoral movement in the acetabular socket more than in Archosauria
view. (C) Right calcaneum
ancestrally. The presence of the anterior trochanter and trochanteric shelf
of Pseudolagosuchus in
lateral view. (D) Right
suggests greater development of or differentiation of both the femoral
calcaneum of Herreras- protractors and the retractors. Also, the proximal and distal ends of the
aurus in lateral view. (E) tibia exhibit interesting modifications: a cnemial crest on the proximal
Left distal tarsals 3 and tibia, which suggests differentiation and enlargement of the protractive
4 of the basal pterosaur
(extensor) muscles of the lower leg, and flexor muscles of the ankle.
Dimorphodon in proximal
view. (F) Left distal tarsals In contrast to the transformations noted above, modifications of
and metatarsals of Lager- pelvic elements were delayed for the hindlimb bones during the early
peton in proximal view. evolution of dinosauriforms. Since pelvic bone morphology retained the
(G) Left distal tarsal 4 of ancestral archosaurian condition (e.g., a craniocaudally short ilium and
Lagerpeton in caudal view.
(H-J) Left distal tarsals and
a pubis and ischium that were not ventrally elongated), it is assumed
metatarsals of Lagosuchus that no major modifications occurred in the placement of origin of the
in (H) proximal, (|) lateral, pelvic musculature. This implies that bone morphology, mainly the
and (J) caudal views. (K—-M) acetabular-femoral and tibiofibular-tarsal articulations (but not muscular
Right (reversed) distal tarsal rearrangement at the area of origin), were major constraints of hindlimb
4 and metatarsal IV of Her-
rerasaurus, in (K) proximal,
movements in a parasagittal plane in the early evolution of ornithodirans.
(L) lateral, and (M) caudal Modifications occurred at the insertion areas of the protractive and retrac-
tive muscles of the femur, probably allowing an increase in the power

20 The Age of Dinosaurs in South America

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Title: Follow the Ball

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 FOLLOW THE BALL

By RALPH HENRY BARBOUR

Yardley Hall Series

FOURTH DOWN
FORWARD PASS
DOUBLE PLAY
WINNING HIS Y
GUARDING THE GOAL
FOR YARDLEY
AROUND THE END
CHANGE SIGNALS

Purple Pennant Series

THE LUCKY SEVENTH

THE SECRET PLAY
THE PURPLE PENNANT

Hilton Series

THE HALF-BACK
FOR THE HONOR OF THE SCHOOL
CAPTAIN OF THE CREW
 Erskine Series

BEHIND THE LINE

WEATHERBY’S INNING
ON YOUR MARK

The “Big Four” Series

FOUR IN CAMP
FOUR AFOOT
FOUR AFLOAT

The Grafton Series

RIVALS FOR THE TEAM

HITTING THE LINE
WINNING HIS GAME

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COXSWAIN OF THE EIGHT

Books Not In Series

THE LOST DIRIGIBLE

FOR THE FREEDOM OF THE SEAS
KEEPING HIS COURSE
THE BROTHER OF A HERO
FINKLER’S FIELD
DANFORTH PLAYS THE GAME
THE ARRIVAL OF JIMPSON
UNDER THE YANKEE ENSIGN
 BENTON’S VENTURE
THE JUNIOR TROPHY
THE NEW BOY AT HILLTOP
THE SPIRIT OF THE SCHOOL
THE PLAY THAT WON
OVER TWO SEAS (With H. P. HOLT)
FOR THE GOOD OF THE TEAM
INFIELD RIVALS

D. APPLETON AND COMPANY, Publishers, New York

THEN HAP FERRIS MADE A PASS TO SAWYER

FOLLOW THE BALL

BY

RALPH HENRY BARBOUR

 AUTHOR OF “INFIELD RIVALS,” “FOR THE GOOD OF THE TEAM,”
“COXSWAIN OF THE EIGHT,” ETC.

D. APPLETON AND COMPANY

NEW YORK :: 1924 :: LONDON

copyright,
1924, by
D. APPLETON AND COMPANY

Copyright, 1920, 1921, 1922, 1923, by The Sprague Publishing Company

Copyright, 1920, 1922, 1923, by The Century Company

PRINTED IN THE UNITED STATES OF AMERICA

 CONTENTS
CHAPTER PAGE
I. For the Team 1
II. A Sacrifice for Kenton 14
III. Friends at Outs 23
IV. Ghosts 37
V. The Vigilantes 48
VI. Joe Finds a Clue 59
VII. The Lone Chase 70
VIII. Joe Resigns 80
IX. Gus Billings Narrates 93
X. Gus Billings Concludes 104
XI. Camp Resthere 116
XII. Uninvited Guests 127
XIII. Down the Brook 139
XIV. Alonzo Jones Speaks 149
XV. Alonzo Goes On 163
XVI. Ginger Burke 176
XVII. One All 187
XVIII. The Deciding Game 199
XIX. Ginger Signs Up Again 212
XX. Called to the Colors 223
XXI. Joe Follows the Ball 237
FOLLOW THE BALL
 CHAPTER I
FOR THE TEAM

J oe Kenton, tilted back in his swivel chair, was thinking.

The school year was nearly over and there were many things
that he had meant to do and hadn’t done. There was that extra
course in the spring term, there was that reading that was to have
made next year easier, there was—well, several other things. Such
as getting on better terms with his roommate. That, too, had got by
him, in spite of all his good intentions. There was some excuse for
abandoning the extra course and the reading; playing on the school
nine hadn’t left much time for additional work; but attaining the
reputation of being the cleverest second baseman in the history of
the school needn’t have kept him from making up with Hal Norwin.
The silly part of it was that there was no apparent reason for the
estrangement. They had entered Holman’s together last fall, and,
although they had never chummed much at home, it had seemed
natural that they should room together. But it hadn’t worked out
well. They had managed to get along without a real quarrel, but that
was the best that could be said. And now, although no word had
been spoken of it, it was mutually understood that next year they
should separate. There were moments when Joe regretted it. It did
seem that they should have hit it off better. Why hadn’t they? He
had nothing against Hal; or nothing much. He did think him a bit
snobbish, inclined to make too much of the fact that his school
friends were of the “smart crowd.” And sometimes he acted “stuck-
up” about his playing. Perhaps, though, he had a right to, for he was
easily the best man on the team, not even excepting Captain Bob
Stearns. As for his trying to get Wilder on second instead of Joe,
why, he had a right to his judgment. Still, that rankled.
 Perhaps, thought Joe, if he had made the effort when he had
meant to, away last autumn, they might have got together, and life
in 14 Routledge would have been fairly jolly. Fourteen was a dandy
study. They had been lucky to get it. He wished he could be certain
of having as good a one next fall; for, of course, he would get out
and let Hal fill his place with a more congenial roommate. In case
the trouble had been more his fault than Hal’s, that would sort of
make up. And speaking of Hal, where the dickens was he?
The clock on his dresser said twenty-two past eleven. At Holman’s
you were required to be in hall at ten unless you had secured leave,
and even then eleven was the limit of absence. And here it was
twenty-two minutes after! Well, Hal must have obtained permission,
for he couldn’t get in now without ringing, and he surely wouldn’t be
idiot enough to risk a row with faculty! And yet, he reflected as he
began to undress, it wouldn’t be unlike Hal to take a chance just at
the wrong time. He was forever doing it—and forever getting by with
it! The crowd he trained with thought it clever to show contempt for
rules and had, as Joe well knew, a long list of unpublished
escapades to their credit; or discredit. Oh, well, he should worry!
What happened to Hal was none of his business. He had plenty of
troubles of his own; one of which was to get the light out before
“Granny” Maynard, second floor proctor, began his nightly snooping
expedition. However, there were still full three minutes—
There was a sound at the open window. A hand slid over the sill
and then the upper part of a body appeared against the outer
darkness. “Give me a hand, Joe! That’s some climb. Thanks.” Hal
Norwin swung over the ledge, breathing hard but grinning in
triumph. Then the grin changed to a frown. “Rotten luck,” he
continued. “I thought maybe they’d forget to lock the door for once,
but of course they didn’t. And ‘Granny’ stuck his silly old bean out
and saw me. I beat it around back, but I’ll bet he recognized me.
Got the door locked?”
Joe nodded. “Yes, but we’ll have to let him in if he comes. Funny
he hasn’t been around if he saw you.”
 “Well,” panted Hal, “if he stays away another ten seconds I’ll beat
him.” He struggled out of his clothes rapidly. “But if he did recognize
me and reports me—well, you know the answer; probation for yours
truly! And pro doesn’t suit me just now; not with the Munson game
the day after to-morrow. There, now let him come! I—listen!”
There were footsteps in the corridor. Joe leaped toward the
switch. In the sudden darkness he heard Hal’s bed creak. The
footfalls came nearer. Joe, standing silent in the darkness, listened
and hoped. Perhaps Maynard was only making his rounds, after all.
Perhaps he hadn’t seen— The steps stopped outside. There was a
moment of suspense. Then three brisk raps sounded.
“Pretend you’re asleep!” whispered Hal.
But Joe, remembering that he was still attired in his underclothes
and that he had but the moment before put the light out, saw the
uselessness of that. Instead, he fumbled his way to the door and
opened it. The proctor stood revealed in the dim light of the corridor.
“Norwin,” he began.
“I’m Kenton,” said Joe placidly. “What’s up?”
“Turn your light on, please.” Maynard pushed past Joe into the
room. The radiance showed the apparently sleeping form of Hal, a
litter of hurriedly discarded garments about his bed and Joe but
partly undressed. Maynard viewed the motionless form beneath the
covers perplexedly. Then:
“Which of you came in by the window just now?” he demanded.
“By the window!” echoed Joe incredulously. “What is it, a joke?”
“Now stop, Kenton!” Maynard raised a hand. He was tall and thin
and bespectacled, and had a way of holding his head slightly
forward from his shoulders as he talked, perhaps because the
glasses did not quite overcome his nearsightedness. “Don’t trouble
to lie. I know what I’m talking about, for I watched from the lavatory
window and saw one of you climb in there. And I’m pretty certain
which one it was.” He turned toward the form huddled under the
covers. “I’m sorry,” he went on, “but I’ll have to report you. I can’t
understand your doing a crazy thing like this, though.” His tone was
indignant. “You must have known what it meant to be caught. If you
didn’t care on your own account you ought to have realized what it
would mean to the team, to the school. Hang it, it isn’t fair to risk
defeat just for the sake of some piffling escapade in the village!”
The form under the bed-clothes stirred, an arm was thrust forth
and Hal groaned sleepily. Then, as though disturbed by the sound or
the light, he thrust the clothes down and blinked protestingly. It was
a good piece of acting. Joe wondered whether Maynard was
deceived by it. It was hard to tell.
“Put out that light, Joe,” muttered Hal. Then, wakefully: “Hello,
what’s the row?”
Maynard viewed him doubtfully. “I think you heard what I said,” he
observed.
“He says he saw some one climb in our window a while ago.” Joe
nodded smilingly at the proctor.
Hal turned and looked at the window, blinking and rubbing his
eyes. Then: “Wh-what for?” he asked stupidly.
“I don’t think he said,” replied Joe gravely. “You didn’t say, did
you, Maynard?”
“I’ve had my say.” The proctor turned toward the door. “I’m sorry,
fellows.”
“Just a minute!” said Joe. “Do you still think you saw—what you
said, Maynard?”
“Naturally.”
“And you feel that it’s—it’s up to you to spoil Saturday’s game?”
“It’s up to me to report to faculty. You should have thought of the
game before.”
 “It seems sort of tough,” muttered Joe. Maynard flashed a puzzled
look at him. Hal sat up impulsively.
“Oh, well,” he began, “I suppose—”
“Never mind,” interrupted Joe, shrugging. “I can stand it, I guess.”
“You mean—it was you?” demanded Maynard, staring hard.
Joe shrugged again. “I thought you said you knew,” he scoffed.
“I think I do,” replied Maynard meaningly, with a quick side glance
at Hal’s troubled face. “But I can’t prove I’m right, I suppose. Seems
to me it would be the decent thing for one of you to own up,
though.”
Again Hal started to speak and again Joe interrupted. “Oh, piffle,
Maynard! A fellow’s innocent until he’s proved guilty. Anyway, I
guess the—the circumstantial evidence is all you need.”
“All right, have it your way, Kenton. You know where the evidence
points. I’m sorry to have—I’m sorry it happened. Good night.”
“I’m sorry, too,” answered Joe soberly. “Good night, Maynard.”
The door closed behind the proctor and Joe snapped off the light.
After a long moment of silence: “What did you do that for?”
demanded Hal, truculently.
“Well, he was sure it was one of us. If I don’t play Saturday it
won’t much matter. If you don’t, it’ll matter a lot. You’re the only one
of us who can hit Cross, and unless some one hits him we’re going
to get licked. Besides, I didn’t lie to him.”
When Joe had struggled into his pajamas and crawled into bed
Hal spoke again. “Mighty decent of you,” he said. “Don’t know that
I’d have done it for you.”
“Wouldn’t expect you to. I didn’t do it for you, so that needn’t
worry you. I did it for the team; or the school; or maybe just
because I want to see Munson beaten.”
 “Oh,” replied Hal in relieved tones. “That’s different!” A minute
later he added: “Sorry you’re in a mess, though.”
“That doesn’t matter. G’night!”
Doctor Whitlock seemed the next day much more grieved than
Joe. Of course, the doctor explained gently, it meant probation for
the balance of the term, and probation meant that he wouldn’t be
allowed to take part in athletics, but in view of the fact that Kenton
had maintained good standing for the school year and was well up
near the head of his class there would be no further—ah—penalties
inflicted. Joe thanked him gravely. Outside again, he laughed
mirthlessly. Just what other penalty, he wondered, did the principal
think mattered now?
He and Hal had not mentioned last evening’s incident again. For
that matter, there had not been many opportunities, for they had
seen each other but a few minutes before breakfast. While dressing
Hal had seemed morose and out of sorts. After the interview in the
office Joe returned to Number 14. He might have gone over to the
field and watched practice, and would have done so if he hadn’t
funked the explanations that would have been required of him.
There was a bad ten minutes just at dusk when Bob Stearns came
in. The captain was hurt rather than angry and said one or two
things that made Joe want to crawl under a bed—or weep. But he
went away finally, leaving Joe feeling very small and mean, and
liking Bob more than ever for the things he might have said and
hadn’t. Then there was another knock and Joe’s silence didn’t
protect him, for “Granny” Maynard opened the door and descried the
lone occupant of the study in the twilight.
“Mind if I come in a minute, Kenton?” he asked. “You know the
fact is I feel particularly rotten about what’s happened and I do wish
it had been some one else besides me. How bad did they treat you?”
“Not very, thanks. Pro, of course. You needn’t feel badly, though.
You only did what you had to.”
 “I know, but—being proctor is fairly rotten sometimes. If it wasn’t
for the difference it makes in my term bill I’d quit it. But I really can’t
afford to. I suppose you’re out of the game to-morrow?”
“Oh, yes. But my being out of it won’t matter much.”
“Not so much as Norwin,” said Maynard significantly.
“Norwin? Oh, no! Hal’s the best player we’ve got. Don’t you think
so?”
“I’m not much of an authority, but I’ve heard it said that he is.”
There was a moment of silence. “It’s none of my business, Kenton,
but I must say I think it was very decent of you.”
“Thanks,” replied the other dryly. “What?”
“I guess you know what I mean. I’d rather not put it in words
because—well, I’m not supposed to know anything about it.”
Maynard laughed as he arose. “As I said before, Kenton, I’m beastly
sorry.” He held out his hand and Joe, a trifle surprised, took it. “Hope
we win to-morrow, eh?”
“Rather!” agreed Joe. After Maynard had gone he frowned into the
darkness beyond the open window. “He knows. Or he thinks he
knows. Well, it doesn’t matter. Nothing does—much. I wonder if I
told Hal the truth last night, though. Did I do it for the school or
didn’t I? Of course I want Holman’s to win, but—I don’t know! But
I’d hate to have him suspect that—that—oh, shucks, that’s
tommyrot! Why should I do it on his account? Of course I didn’t!
Surly brute!”
Hal came in a few minutes later. He didn’t see Joe until he had
turned the light on. Then: “Hello!” he said awkwardly.
“Hello. How did practice go?”
“All right, I guess. Wilder played second.”
Joe nodded. “I supposed he would. That ought to please you.”
“Me? Why?”
 “You wanted him there, didn’t you?”
“Sure! With you out of it—”
“I mean before. Last month.”
“I don’t know what you’re talking about.”
“Oh, rot! You tried your best to get Wilder on second in place of
me, didn’t you?”
“Who told you that?” demanded Hal sternly.
“Why, I don’t know that any one exactly told me. Anyhow, it didn’t
matter much. He’s got the place finally.”
“So you’ve been holding that in for me?” sneered Hal. “Let me tell
you, then, that I did not try to get Wilder on second. I didn’t even
want him there. Why would I? You’re the better player.”
“Oh!” murmured Joe, somewhat blankly.
“Yes, ‘oh!’” retorted the other. “I don’t say I wouldn’t have tried for
Wilder if I’d wanted him. But I just didn’t. Now chew that over.”
“All right. But I thought—”
“You’re always thinking something that isn’t so,” grumbled Hal. “I’ll
bet you’re doing it right now, too!”
“What do you mean?”
“You’re thinking that I—that I let you take the blame for last night
because I want to play to-morrow,” flared Hall. “I do, but, if that was
all I wouldn’t have let you. I’m standing for it because I know
plaguey well that if I don’t play we’ll get beaten. Oh, I dare say that
sounds cocky, but it’s so. I can hit Cross’s curves and not another
one of you fellows can come anywhere near ’em.”
“I know, and I’m not kicking, am I? I said it was me because I
knew we’d get ‘Finis’ written all over us if you were out of the game.
So what’s the use of chewing the rag about it now?”
 “Because I won’t have you think I’m a—a sneak and a coward!
And you do think so—inside.”
“I don’t!”
Hal had come close and now he stood staring down at Joe
menacingly. “You don’t?” he demanded suspiciously.
“No, I don’t.”
“All right. See that you don’t. If I thought you were lying I’d—I’d
knock your head off! Mind you, I appreciate what you’ve done for
me—”
“You!” shouted Joe, jumping up. “For you? Don’t you dare say I
did it for you! I did it because I wanted to.” He waved a finger under
the other’s nose. “Just one more crack like that and I’ll punch your
ugly face in!”
“I didn’t mean me personally,” growled Hal. “Anyhow, we
understand each other, I guess.”
 CHAPTER II
A SACRIFICE FOR KENTON

H olman’s School had won the first contest with Munson, and she
wanted very much to win the second and do away with the
necessity of playing a third on neutral territory. This warm, blue-and-
gold June afternoon found them well matched and eager, how well
matched is shown by the fact that until the sixth inning neither side
scored. Then Prentiss got Holman’s first hit, a rather scratchy affair
at that, and although Cummins was thrown out at first Prentiss
reached second. Cross, Munson’s really remarkable twirler, let down
long enough to pass Wilder and, with one down, Holman’s cheered
hopefully. “Babe” Linder flied out to shortstop, however, and it
remained for Cochran, Holman’s left-hand pitcher, to do the trick, or,
rather, to bring it about. Cochran was no batsman, and he knew it,
just as every one else did, but he had a wonderful faculty for getting
in the way of the ball. I’m not prepared to say that it was
intentional, but Cochran’s average was just about one base per
game owing to being struck by a pitched ball. This time he got it on
the thigh, started right off for first and, it may be, decided the
matter for an umpire who was inclined for an instant to be doubtful.
That filled the bases and there was a good deal of noise from
coaches and spectators, and Cross, disgruntled, sought revenge by
trying to catch Stearns off second, or by pretending to. At all events
the ball went over the shortstop’s head, Prentiss scored and Stearns
raced for third but was caught when the center fielder pegged a
swift one to the third sack.
But Munson evened things up in the eighth, just when the home
team had visions of a one-to-nothing victory, by getting two clean
hits off Cochran and combining them with a clever steal. And at 1—1
the game dragged—no, it never dragged for an instant. But at 1—1
it stayed until the last of the eleventh. Holman’s had no hope of
doing anything in that particular inning, for the tail end of her
batting list was up: Wilder, Linder, Cochran. But you never can tell
when the break will come. Wilder was passed, Babe Linder laid down
a sacrifice bunt and Cochran, in spite of almost Herculean efforts,
took the fourth ball pitched squarely on his shoulder! Cross
complained bitterly when the rival pitcher was waved to first, and I
think the incident affected his delivery. At all events, Torrey, left
fielder and head of the batting list, rolled one toward third and after
baseman and pitcher had each politely left it to the other during a
tragic moment the latter threw late to first. With bases filled, but
one out and Hal Norwin swinging his two bats as he stepped to the
plate, there could have been but one outcome. Cross had to pitch
’em and he knew it. Perhaps Cross already read the writing on the
wall, for Hal said afterwards that that third delivery came to him with
nothing on it but a sunbeam. He said that it looked so good he was
almost afraid of it. Possibly Cross intended he should be. But Hal
didn’t scare quite so easily as that, and so he took a fine healthy
swing at it and it traveled. It went straight and far and came safe to
earth yards out of reach of right fielder and to Cummins went the
honor of scoring the winning tally!
Joe didn’t march back to the campus with the triumphant horde
but cut across back of the gymnasium and made his way to Number
14 in a somewhat depressed frame of mind. He had watched the
game from start to finish and was well satisfied at the outcome, but
he hadn’t been happy. When you have worked hard from February
on to win your position and have set your heart on playing in the Big
Game, why, you just can’t help feeling a bit glum when the Big
Game finds you perched among the noncombatants of the
grandstand. I don’t think Joe really regretted what he had done. One
can be sad without being sorry. But there were moments when he
was rather self-contemptuous, when he told himself that he had
done a silly, quixotic thing for which no one thanked him.
 They were still cheering and singing over in front of School Hall
when he reached his room, and the sounds came to him around the
corner of the building and floated in at the open window. Although it
was nearly five o’clock the golden sunlight still streamed across the
meadows beyond the little river and save for the disturbing and
discordant sounds from the campus the world was dreamily silent. It
was beautiful, too, with the fresh, new green of grass and leaves
and the peaceful sky and the mellow sunlight, but he was glad that
in a few more days he would see the last of it for a while. In fact, he
wasn’t sure that he ever wanted to return to Holman’s. He felt so
horribly like a failure.
The shadows lengthened and the sunlight became tinged with
flame. The dormitory echoed to laughter and the tramp of feet and
the slamming of doors. Then, presently, his own door opened and
Hal came in, bustlingly, radiating triumph and high spirits. “Some
game, Joe!” he cried. “By jiminy, though, I thought they had us for a
while! Didn’t you?”
“Yes,” replied Joe listlessly. “Cross was in great form.”
“Wasn’t he? I couldn’t get near him—until the last inning. Well, we
won, thank goodness!”
Joe made no answer and Hal busied himself at the washstand.
After a while: “You’re coming to the dinner, aren’t you?” asked the
latter.
Joe hesitated. He had forgotten that the team would dine in state
to-night in the visitors’ hall, with speeches and songs and at the end
of the modest banquet, the election of a new captain. “I don’t
know,” he said finally. “I suppose I have a right to, but—”
“Of course you have. Any fellow who has played on the team
during the season has. I asked because—” Hal hesitated, and Joe,
looking across, saw him as near embarrassment as he ever got. “The
fact is,” he began again, and again stopped.
“Don’t worry,” said Joe. “I intend to, anyway.”
 “Intend to what?” asked Hal, looking puzzledly over the towel with
which he was drying his face.
“Vote for you for captain.”
“Oh, that! Thanks, but you needn’t if you’d rather not. I sha’n’t
mind if you don’t. That isn’t what I was going to say, though.” He
tossed the towel aside and, hands in pockets, came over to the
window. “Look here, Joe. I haven’t been feeling any too easy
yesterday and to-day. I thought it was all right to let you take the
blame for—for my foolishness because it might mean winning the
game to-day. And I guess it did mean that, as it’s turned out. But
I’ve sort of hated myself, just the same, and I guess what I ought to
have done was stand the racket myself and let the game look after
itself. But I didn’t and post mortems don’t get you anything. But
there’s no reason for carrying the thing any further. What we’ve got
to do now is get you squared up with faculty and the school and—
and every one. So I’m going to tell ’em the truth at dinner to-night.”
“That’s a brilliant idea!” scoffed Joe.
“Why not?”
“Why not? Because there’ll be at least two faculty there, and if
you think they’ll let you accept the captaincy after ’fessing up to that
stunt you’re all wrong.”
“I don’t. They’ll have me in probation to-morrow, of course. That
isn’t the question.”
“Of course it’s the question,” said Joe impatiently. “You’re
practically sure of the captaincy. I know it and so do you. If faculty
gets this on you you’re a goner. Besides, what good’s it going to do
any one? School’s over in three days, and just as long as they’re
going to let me pass with my class I don’t mind three days in
bounds.”
“That’s all right,” replied Hal stubbornly, “but right is right. I let
you suffer because I wanted to win the game. The game’s won. Now
it’s my turn to stand the gaff.”
 “And lose the captaincy!”
Hal shrugged. “I know. I thought of that, though. It can’t be
helped. Besides—”
“It can be helped!” said Joe angrily. “All you need to do is get this
fool idea out of your head. You talk like a—a sick fish!”
“Just the same—”
“No, sir! I won’t stand for it! What sort of a silly fool do you think
I’d feel like with you getting up before all that bunch and—and
spouting all that rot? If you tell that yarn I’ll deny it!”
Hal smiled. “I can prove it, though. I can produce five fellows who
will testify that I was in Gus Billing’s room at eleven o’clock that
night.”
“Is that where you were?” asked Joe eagerly.
“Yes.”
“Oh! Why, that isn’t—there’s no harm—”
“Of course there’s no harm, but I stayed too late. Gus’s clock was
about an hour slow and I never thought to look at my watch.
Anyhow, it won’t do you any good to deny it, Joe.”
“Well, then—” Joe spoke slowly, frowning intently across the
shadowy room. “Maybe you sort of feel that you—you owe me
something. Of course I didn’t do it just for—just to oblige you, but
you wanted to win, and I guess I helped—”
“Of course I owe you something. I’m trying to make you
understand it. And I’m going to pay what I owe.”
“Not that way,” replied Joe firmly. “If you do want to—to square
things there’s just one way you can do it.”
“How’s that?” asked Hal suspiciously.
“Forget it!”
“No, sir!”
 “Yes, I mean it, Hal.” Their eyes challenged. After a moment Hal
shrugged.
“All right,” he said, “but I don’t get your idea. It isn’t as if you’d
done it for me—” He stopped and there was a long moment of
silence. Then he asked brusquely: “You didn’t, did you?”
“No!” answered the other. Hal walked over, picked up his jacket
and began to put it on. “And what if I did?” added Joe defiantly.
Hal stopped with one sleeve on. “I knew mighty well you did,” he
growled.
“You know a lot, don’t you?” grumbled Joe sarcastically.
“I know that if you don’t wash up and get ready we’ll be late,”
laughed Hal. “Get a move on, Grumpy!”
“Well—but no speeches, Hal!”
“Nary a spooch!”
Joe splashed and gurgled and Hal watched, grinning broadly.
Presently he observed carelessly: “I say, Joe, we’ve only got two
more days to get our application in if we want this room next year.”
Joe dried his face with unusual care. “That’s right,” he said at last.
“Guess we’d better get busy, eh?”

Maynard fell in with Naylor, assistant manager, on his way out.

Naylor was still figuring his totals in the official score book and
Maynard peered over his shoulder.
“What did you give Kenton on that last play?” he asked.
“Kenton? Kenton wasn’t in it, you idiot! Wilder played—”
“Still,” said “Granny” soberly, “I think you should have credited him
with a sacrifice.”
And he went on, leaving Naylor looking after him commiseratingly.
 CHAPTER III
FRIENDS AT OUTS

H al won the captaincy, and two days later he and Joe and Bert
Madden started for home. About three hundred other youths
also started for home, but none of them lived in Central City, and so,
beyond the Junction, Joe and Hal and Bert went on westward alone.
Bert was well over seventeen and would be a senior next year, as
would Hal, a year younger. Joe, who was Hal’s age within a few
months, was returning to Holman’s in the fall as a junior. He and Hal
had been friendly at high school, and when Hal had decided to go to
Holman’s for the last two years Joe had decided to go also. It wasn’t
so easy for Joe, however, for Joe’s folks weren’t wealthy by any
means, while Hal’s were. But he had found employment last summer
and worked hard, and, when September had arrived, his earnings,
with what his father had been able to provide, had been sufficient to
put him through the first year.
It wasn’t going to be nearly so hard next fall, for Mr. Kenton’s
business had improved. Nevertheless, Joe meant to find some sort
of employment for the summer months, and on the journey home
this matter occupied his thoughts a good deal of the way. He
couldn’t go back to Murray and Bankhead’s, for his place there was
occupied permanently by another, but he was certain that he could
find a job of some sort. While Joe considered ways and means, Hal
was telling Bert about the good time he was going to have at his
father’s camp up north and Bert was picturing the delights of
summer life at one of the nearby summer resorts. Hal had invited
Joe to visit the camp some time toward the last of the summer and
Joe had half accepted the invitation. He didn’t really expect to get
there though.
 Hal left town about a week after their return home, and Joe
missed him a good deal at first, even though they didn’t get
together very often in Central City. Hal moved in a different circle
than Joe. Looking for work, however, occupied much of Joe’s time
during that week and the next, for he had been home more than a
fortnight before he secured the job with Donaldson and Burns, who
operated the Central City Market. His principal duty was to deliver by
bicycle, orders that could not await the trucks or that had been
forgotten by them. When not occupied in that way he sometimes
helped to put up orders. His hours were from eight to five, save on
Saturdays, when the store kept open until nine. Thursday afternoons
he had off, for in Central City Thursday was the weekly half holiday
from July to September.
It was on the first Thursday afternoon after starting to work that
he sat on an empty soap box by the window of the stable loft and
listlessly distributed type from a “stick” in his left hand to the case
before him. The July day was hot, and from the printing press that
stood on a stout packing case came a strong though not unpleasant
odor of fresh ink. Joe wasn’t very happy this afternoon. On a shelf
under the type case lay the results of his recent labor, twelve printed
invitations still sticky from the press. Now, having distributed the last
of the type, he lifted one of the invitations, held it at arm’s length
and read it. Beginning in script, it ran the gamut of Old English,
italics and small Roman, and it read as follows:
You are Cordially Invited
to Attend a House Warming at
Camp Peejay, Squirrel Lake,
Thursday, July 6.
Philip Levering Joe Kenton
R. S. V. P.
It really looked awfully well, but he couldn’t get much of a thrill
from that fact since, as sightly as they were, those invitations would
probably never be used.