Journal of Vertebrate Paleontology 27(3):778–779, September 2007
© 2007 by the Society of Vertebrate Paleontology
MAMMALS FROM THE AGE OF DINOSAURS—ORIGINS, EVOLUTION, AND STRUCTURE, by Zofia Kielan-
Jaworowska, Richard L. Cifelli, and Zhe-Xi Luo, 2004. Columbia University Press, New York, 630 + xv pp., 239 figures, US $195,
ISBN 0-231-11918-6.
For a book about Mesozoic mammals, there are far fewer pictures of
teeth than might be expected in Mammals from the Age of Dinosaurs—
Origins, Evolution and Structure. This reflects well on the state of the
subject, and especially so on the three authors of an extremely compre-
hensive book. All known genera at the time of publication—283—are
illustrated and discussed in a phylogenetic context, with the phylogeny
presented in the last chapter shaping the structure of the rest of the
volume. The format consists of (in addition to a foreword, preface, ap-
pendix, references, etc.) the following 15 chapters; 1: Introduction; 2:
Distribution of Mesozoic Mammals in Space and Time; 3: Origin of
Mammals; 4: The Earliest-Known Stem Mammals; 5 to 14: ten chapters
discussing particular clades or grades of Mesozoic mammals; 15: Inter-
relationships of Mesozoic Mammals. The introduction starts by pointing
out that in comparison with the Cenozoic ‘Age of mammals,’ Mesozoic
mammalian history is longer (155 Ma compared with 65 Ma), and that in
terms of diversity of clades, the three major extant lineages are a small
proportion of the 25 known from the Mesozoic. The authors’ definition
of Mammalia is introduced (all descendents of the common ancestor of
Sinocodon and extant monotremes, marsupials, and placentals). Having
defined the size of the subject and the scope of the book, a brief look is
taken at the premammalian synapsids the Mammalia are nested within,
the changing perception of the shape of Mesozoic mammal phylogeny,
and the timing of major episodes of diversification. Mention is made of
the small size of the vast majority of Mesozoic mammals, possible evo-
lutionary causes for this are examined, and the apparent domination of
Mesozoic terrestrial ecosystems by non-avian dinosaurs noted.
The second half of the introduction deals with the state of research
into Mesozoic mammals, starting by listing major advances over the past
25 years, since the publication of this volumes predecessor, Mesozoic
Mammals: The First Two-thirds of Mammalian History (Lillegraven et
al., 1979). A graph of total described Mesozoic mammal genera, and the
number described per decade, against time, shows both to be on an
exponentially increasing curve, indicating the majority of fossilised taxa
remain undiscovered. Key morphological studies are highlighted, show-
ing how an increase in ontogenetic and morphogenetic understanding
have been coupled with new studies, such as those on enamel micro-
structure, and advances in technology (i.e., computed tomography scan-
ning) to greatly aid the interpretation of mammalian evolution. An im-
provement in the quality of morphological data is noted, with informa-
tive characters other than those of the molar teeth available across a
variety of taxa. The authors view integrating this with the vast amount of
molecular data available on extant mammals as a positive opportunity to
reciprocally illuminate both subjects, especially in areas such as deep
mammalian relationships, calibration of molecular clocks, and the origin
and relationships of modern mammalian orders. A section on paradigm
shifts in mammalian evolutionary studies reiterates recent comprehen-
sion of the tree as bushy, requiring many stem lineages to encompass the
great diversity of Mesozoic mammals, and the reinterpretation of a num-
ber of groups as paraphyletic grades. The discovery of Southern tribo-
sphenic mammals from the Middle Jurassic-Early Cretaceous and the
possibility of non-monophyly of tribosphenic mammals is also noted as a
paradigm shift. The chapter is rounded out with notes on systematic
conventions followed throughout, and two alternative taxonomic classi-
fications are presented at the suborder/family level, one in the traditional
Linnean form and one cladistics based.
Chapter 2 is a comprehensive account of worldwide localities known to
have produced Mesozoic mammalian remains. Faunal lists for the sites
are presented by age and region, with maps indicating localities (where
necessary amalgamating a number of nearby coeval localities). Diagrams
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BOOK REVIEW
summarise the distribution of localities through time by geographic re-
gion, and the final figure displays the huge disparity between the number
of recorded occurrences of fossil mammals in the late Cretaceous of
North America and all other regions and time periods. This is necessarily
reflected in the content of the chapter, with over a quarter dedicated to
the most heavily sampled epoch and continent. The geological and pa-
leoenvironmental context of each locality is discussed along with brief
morphological and taxonomic comments on the fossils present, and plate
tectonics, biogeography and relative abundance of clades noted for each
epoch.
There follows an excellent chapter on the origin of mammals, discuss-
ing the stepwise acquisition of diagnostic mammalian features across the
cynodont-mammalian transition. Six character complexes of dental and
osteological synapomorphies are identified, and a table presents 34 in-
dividual characters with the condition for the mammalian crown group,
Mammalia, Mammaliamorpha and the eucynodont precursor/
pleisomorphic condition. Eight categories of character complex are then
examined, each broken into subsections (i.e. ‘Evolution of the snout’
encompasses palatal structures; nasal structures; and orbitosphenoidal
and ethmoidial structures). The condition in clades and taxa displaying
the transition of a feature is discussed, and clearly illustrated either with
the cynodont, Mammaliamorpha and Mammalia conditions, or mapped
onto a more inclusive cladogram. Alternative hypotheses for character
transformations and homologies across taxa are presented, and the most
parsimonious explanation for the changes with respect to the phylogeny
adopted in the book noted. The chapter represents a thorough summary
of knowledge on the origin of mammals, including detailed characteri-
sation of the transformations that occurred in the non-mammalian syn-
apsid—mammal transition.
Eleven chapters containing anatomical and taxonomic detail for 283
Mesozoic mammalian genera constitute the bulk of the book. Each chap-
ter deals with one of seven clades or three grades of Mesozoic mammals,
apart from the first, chapter four, which describes ‘The earliest-known
stem mammals’. The latter is so that each of the numerous (but generally
taxon poor) monophyletic groups branching off early from the main
lineage of mammalian evolution does not have to be assigned a separate
chapter. A Linnean classification is presented for the genera discussed in
each chapter, to either the genus or the species level, and where available
a cladistic classification is also included. A brief characterization of the
taxa is provided, either at the start for the seven chapters on clades, or for
each monophyletic clade for the four chapters on other groupings. This
presents basic information on the age and distribution of the group along
with a non-technical summary of the anatomy, the size of individuals,
mode of life etc. Sections on more detailed anatomy and the systematics
of the group are followed by an examination of each genus in turn, listing
a diagnosis, age and distribution, species in the genus, synonymy, and
often a comments section looking in further detail at one or two aspects
of the taxon (e.g., systematics, differing interpretations of anatomy etc.).
Each and every genus is illustrated, either by full skeleton, skull, par-
tial skull (often braincase), dentition or individual teeth. It is worth men-
tioning that, as noted at the start of the review, the vast majority of the
283 genera are illustrated by more complete remains than individual
teeth. In addition to clearly presented illustrations of taxon specific
anatomy, there are; reconstructions of taxa, limb posture, musculature,
nerves and veins/arteries, and range of movement of the pelvis, tail and
hind limbs; molar cusp homology, occlusion, tooth wear and shearing
pattern diagrams; transverse sections through skulls showing details of
the braincase; and scanning electron microscopy micrographs of tooth
enamel microstructure. A number of diagrams show comparative differ-
 BOOK REVIEWS
ences in characters or character complexes within a clade, and across
Mammaliamorpha.
The organisation of these chapters, and the classifications shown in
chapter 1, is based on the phylogeny presented in chapter 15. The chapter
aims “to demonstrate that the cladistic relationship for most (if not all)
Mesozoic mammals can be resolved by parsimony methods on the basis
of currently available morphological data (p. 520).” Issues of taxon and
character sampling are discussed, as are missing data and conflicts of
characters. Mention is also made of the differences and similarities be-
tween morphological and molecular phylogenies of extant mammalian
clades where the evolutionary history of these groups stretches into the
Mesozoic—it is noted that different gene sequences (e.g., those of mito-
chondrial versus nuclear origin) appear to support different hypotheses,
and that some are more consistent with morphological evidence than
others. A plea is made for field paleontologists to search for ever-earlier
members of modern lineages (especially placentals) in order to resolve
the conflicts that do exist, but the authors are encouraged by the number
of areas where morphological and molecular phylogenies have con-
verged over time.
The analysis presented is based upon a previous paper (Luo et al.
2002), and extended to include 48 taxa (including outgroup) and 275
morphological characters. The results of a parsimony analysis on this
data were 50 most parsimonious trees (MPTs), of which the strict con-
sensus is used to summarise the results (Figure 15.1), provide the cladistic
classification in chapter 1, and inform the systematics sections of chapters
4–11. An alternative phylogeny is presented in which a parsimony analy-
sis was constrained to place Allotheria (Haramiyavia plus multitubercu-
lates), a clade not found in the first analysis but supported by dental
characters, outside of the mammalian crown group. A strict consensus of
seven MPTs, each eight steps longer than the unconstrained trees, shows
the placement of a number of taxa is affected (Figure 15.2); Kuehneo-
therium moves from being considered the sister taxon to crown group
Mammalia to within the order Morganucodonta; and a monophyletic
clade within crown group Mammalia consisting of Tinodon plus Eutri-
conodontans is found to be paraphyletic and placed outside the crown
group (Luo et al. 2002 mention this in the text but include a paraphyletic
Eutriconodonta and Tinodon within crown group Mammalia in their
constrained consensus tree, Figure 2).
In combination with the data set presented in the appendix, and rea-
soning on characters and taxa from Luo and colleagues (2002, chapter
15) is an extremely thorough analysis of Mesozoic mammal phylogeny. A
large dataset has been assembled, both in character and taxa numbers, in
order to include virtually all Mesozoic mammal lineages and to use every
aspect of their anatomy to infer phylogeny. The authors acknowledge
that the “definition of highly transformed characters and their character
states can be controversial,” often in part due to the quality of the per-
tinent fossils, but express hope that the dataset “will be critically re-
evaluated, corrected, and added to”—a rare and commendable stance.
However, given that the phylogeny presented is an extremely important
and integral aspect of the volume, there is one aspect that I feel could be
improved upon—the use of ‘stable’ and ‘well-supported’ in respect to
nodes representing monophyletic clades. Ten of the 39 nodes within
Mammalia in Figure 15.1 are numbered, and referred to as “the nodes of
the stable monophyletic clades, discussed in the text” (p. 521). Trivially,
not all of the numbered clades are discussed in the text, and a number of
unnumbered clades important to the author’s arguments (e.g. Australo-
sphenida) are discussed. More importantly, it is uncertain how these
clades have been designated as stable. In systematic terms, stability of a
node on a cladogram is usually related to a measure of its resistance to
dissolution with additional steps (as measured by decay indices/Bremer
support—Bremer, 1994), the degree to which it is retained in randomly
permuted pseudo-replicate datasets (bootstrap support—Felsenstein,
1985), or other such measures. No stability analyses or values are re-
ported either in the book or in Luo and colleagues (2002). It may be that
stable in this case refers to nodes retained between the unconstrained
(Figure 15.1) and constrained (Figure 15.2) analyses; however, the num-
bered nodes are not the only ones shared between analyses, and not all
numbered nodes are identical in both phylogenies. Similarly, under a
heading of ‘Major Mammalian Clades,’ “well supported monophyletic
groups of Mesozoic mammals”(p. 526) are discussed, but no justification
for how a well (or poorly) supported clade is defined is provided. It is
unfortunate that the authors are so explicit about the rest of the analysis
(coding, choice of taxa and characters, analyses run etc.) but don’t pro-
779
vide support values or justification for their choice of well-supported or
stable clades.
To investigate this, the character codings presented in the appendix for
the two taxa Nanolestes and Asfaltomylos were added to the data matrix
of Luo and colleagues (2002). Running the resulting matrix in PAUP
with all settings as in the present volume results in seven MPTs instead
of the 50 in figure 15.1; step length (943), CI (0.495) and RI (0.764) values
are however identical to those reported. Bootstrap analysis (1000 rep-
etitions) gives nodes 1,3,4,5 and 9 (as labelled in figure 15.1 p 521) boot-
strap partition values of less than 50%, indicative of poorly supported
clades. Only node 10 (Boreosphenida) has a bootstrap value of over 90%
(92%) indicating strong support. Similarly, using TreeRot (Sorenson
1999) to calculate decay or Bremer support indices (Bremer, 1988) shows
that nodes 1–5 have a decay index of one—these clades can be consid-
ered poorly supported. Nodes 6–10 have decay indices of 3, 4, 4, 2 and 4
respectively and so can be considered more strongly supported, but nine
other clades in the analysis have equally high decay indices, and one
(Monetremata plus Ausktribosphenida, 5) has higher. Looking at the
broader pattern as identified by bootstrap and decay analyses, support
outside Crown Mammalia is almost universally weak (decay indices of 1,
bootstrap support of 70% or lower). All clades within Australosphenida
plus Shuotherium are moderately to well supported (bootstrap 70% to
98%, decay indices 3–5). Clades outside Trechnotheria (the common
ancestor of Zhangeotherium plus Crown Theria and all its descendants)
but within Crown Mammalia are poorly supported with the exception of
two well-supported terminal sister group relationships (Priacodon plus
Trioracodon, 95% bootstrap/decay index 4; Plagiaulacidans plus Cimolo-
dontans, 96% bootstrap/decay index 4). Within Trechnotheria, support
for clades ranges from low to high (decay index 1–4; bootstrap 38% to
94%). In summary, the clades singled out as being stable or well sup-
ported are no more so than others in the phylogeny, and six (1–5, 9) are
considered poorly supported (decay index 2 or less, bootstrap support
52% or less).
Returning to the positive, a theme that runs throughout the book is the
acknowledgment of alternative hypotheses. Whether the homology of
molar cusps, the validity of a species or genus, the systematic position of
a taxon or group, or the multiple origin of an important feature (i.e.,
tribosphenic molars), opposing theories are presented in a favorable
light, and the tentativeness of some of the conclusions reached well
expressed. Another pleasing aspect is the explicitness and pragmatism
with which the authors describe and use stem and crown groups, clades
and grades, Linnean taxa etc, while in each case again presenting alter-
native viewpoints (i.e., the definition of Mammalia, p. xiii).
Overall, this is a superbly well-written and comprehensive compilation
of our knowledge on Mesozoic mammals, and as such is recommended to
all, especially students and researchers, with so much as a passing interest
in the field. The provision of information on all known genera, and the
almost exhaustive literature section, allow its use as the foremost refer-
ence on the subject. Additionally, data like the rate of discovery of
Mesozoic mammal genera, graphs of discoveries by geographical area
and geological epoch, and the most comprehensive data set and phylo-
genetic analysis for the group at date of publication, make it much more
than simply a review volume. Like its predecessor, this book will inspire
and guide Mesozoic mammal research for at least the next quarter of a
century.
IAN J. CORFE
University of Bristol
Department of Earth Sciences
Wills Memorial Building
Queens Street
Bristol BS8 1RJ
LITERATURE CITED
Bremer, K. 1994. Branch support and tree stability. Cladistics 10:
295–304.
Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using
the bootstrap. Evolution 39:783–791.
Lillegraven, J. A., Z. Kielan-Jaworowska, and W. A. Clemens (eds.).
1979. Mesozoic Mammals: The First Two-thirds of Mammalian His-
tory. University of California Press, Berkeley, 311 pp.
Luo, Z.-X., Kielan-Jaworowska, Z., and Cifelli, R. L. 2002. In quest for a
phylogeny of Mesozoic mammals. Acta Palaeontologica Polonica
47:1–78.