1983 Biological Significance of Mammalian Milk Teeth c2

CHAPTER 1
INTRODUCTION
Dentition has always been a major tool in the study of mammals,
largely because mammal teeth exhibit a rich array of fine variation
superimposed on a basically conservative, persistent structure. Of
the lifetime complement of two sets of teeth that mammals are
conventionally said to possess, the second or permanent set has been
analyzed in the contexts of age determination, chemical and physical
structure, comparative morphology, development, dietary
specialization, mechanical engineering, paleontology, pathology,
population genetics, predatory ecology, social behavior, and
systematics (some recent examples: Butler 1982, Grue and Jensen 1979,
Kay 1978, Leutenegger and Kelly 1977, Lucas 1979, Lumsden 1979,
McKenna 1975, Radinsky and Emerson 1980, Townsend 1980). Such studies
now form the core of what we know about mammalian phylogeny and have
added considerably to knowledge of adaptative processes. By contrast,
the first dentition, the deciduous or "milk tooth" set, has been
largely ignored. One reason for this neglect may be that milk teeth
have relatively short tenure in an individual. Yet the juvenile
period in mammals is always a time of high growth rate, high
mortality and consequent high potential to affect fitness. An
assessment of the importance of milk teeth in the lives of juvenile
mammals would be of clear interest, the more so because the adaptive
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2
significance of permanent teeth to adult mammals has already been so
abundantly demonstrated.
In this study I make such an assessment. The first necessary
task was to grasp the scope of what is and can be known about milk
dentition in mammals; these patterns are documented in Chapter 2. I
then selected one group, the family Mustelidae, for a more detailed
analysis of milk tooth morphology and timing in relation to important
features of development (Chapter 3). This led to investigation of
some problems of tooth use that young carnivores face, in a
laboratory study of acquisition of predatory efficiency in
long-tailed weasels (Chapter 4). Ultimately, the clarified patterns
of milk tooth presence and use in extant mammals called into question
some current interpretations of what the two-tooth-set system
reflects about parental care in the first mammals; Chapter 5
reevaluates the paleontological evidence and proposes a different
hypothesis for the origin of mammalian milk teeth.
Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.
CHAPTER 2
MAMMALIAN MILK TEETH: PRESENCE AND PATTERNS
INTRODUCTION
Mammals are allegedly diphyodont. Every general textbook on
mammalogy or vertebrate biology claims so; attempts to characterize
the Class Mammalia invariably include the possession of two
dentitions (the milk teeth and a replacement set) as a diagnostic
feature (e.g. DeBlase and Martin 1982, McFarland et al 1979). The
diphyodont condition is one member of that suite of conditions
conventionally signalling the attainment of mammalian grade from
polyphyodont reptilian ancestors. Recently other mammalian inventions
have received review leading to theoretical frameworks and the posing
of investigative questions (e.g. Case 1978a and McNab 1978 on
endothermy, Crompton and Parker 1978 on the masticatory apparatus,
Case 1978b and Gubernick and Klopfer 1981 on parental care, Oftedal
1980 and Pond 1977 on lactation); but the latest comprehensive review
of diphyodonty in the extant Mammalia appeared in 1871, when William
Henry Flower addressed the Odontological Society of Great Britain on
this topic.
Despite the lack of recent synthesis, there have been some
generalizations on the significance of diphyodonty and of the milk
4
dentition (e.g. see Pond 1977). These carry a certain degree of
logical appeal. First, since humans and an array of domestic species
have two definite tooth sets, this familiar pattern seems likely to
characterize the Class as a whole. Replacement of teeth in humans
proceeds in a roughly consecutive order from front to back, so this
might seem a general mammalian development schedule. When baby
mammals begin to eat solid food, they appear to have an adult diet:
thus deciduous teeth might be expected to be morphologically like
adult teeth. All mammals first suckle, then wean, their young; the
replacement of "milk teeth" might thus be expected to show a
consistent relationship to the weaning process and to acquisition of
nutritional independence.
Evidence for these ideas, when supplied at all, is usually in
the form of individual cases. Cumulative data have not previously
been assembled. However, on closer examination it becomes apparent
that a great many species have been examined in some context.
Developmental biologists have reported tooth eruption as an
ontogenetic milestone. Wildlife population ecologists have drawn up
schedules for aging specimens from tooth eruption. Dental researchers
have described tooth development in laboratory species as human
analogs. Paleontologists have sought evidence of diphyodonty in
ancestral mammals. Systematists have made forays into use of milk
tooth characters in tracing phylogenies. Much descriptive information
thus exists: it would be important to exploit these data from
disparate contexts and disparate terminological conventions for a
5
realistic assessment of what mammal milk teeth really do.
Beginning with the two most relevant recent monographs, Peyer’s
(1968) on comparative odontology of the vertebrates and Ziegler’s
(1971) on changes in the dental formula through therian history, I
have located over 800 works which describe milk dentition or its
replacement in some way. The information from these is compiled in
the following review with two general aims. The first part tabulates
milk tooth presence for every species for which there is published
information and summarizes, for each order of mammals, what is known
about timing and function of the milk dentition, along with notes on
issues peculiar to that group. With these data in hand, the second
section examines four prevalent notions about mammalian milk teeth:
1) that mammals are typically diphyodont, 2) that mammal tooth
development is typically consecutive and front-to-back, 3) that
juvenile teeth typically have little significance other than as
temporary stand-ins for adult replacements, and 4) that milk teeth
have some typical relationship to milk feeding. The cumulative data
show that, for these characters, exceptions are often as "typical" as
the supposed mammalian rule.
TOOTH DEVELOPMENT AND REPLACEMENT IN EXTANT ORDERS OF MAMMALS
Taxonomic treatment here follows Honacki et al (1982). Tooth
terminology from various disciplines is so unstandardized and
confusing that I have avoided it wherever possible; Peyer
6
(1968:197-200) gives a lucid explication of several of the most
common conventions in use. "Milk" dentition as used here in
unmodified form refers to a functional dentition present in juveniles
that is replaced (by a "replacement" or "permanent" dentition) later
in life. Where symbolized in the text, milk tooth abbreviations are
conventionally preceded by a "d" (from "deciduous"), as in "di2;" for
brevity, the "d" is omitted in the tables.
Monotremata
Three modern species of monotremes are recognized. For the two
echidnas (Tachyglossidae) no dentition is known in juvenile or adult
(Grasse 1955). The platypus (Ornithorhynchidae) forms a complete
dental laminar tissue with incisor, canine, premolar and molar
regions evident XGreen 1937, Wilson and Hill 1907). In the upper jaw
a canine, two premolars and three molars on each side, and in the
lower jaw five incisors, a canine, two premolars and three molars, or
a total of 22 teeth, begin to develop. Most of these are resorbed,
however, and eventually only twelve teeth (an upper premolar, two
upper molars, and three lower molars on each side) come to function.
Even these are lost as the animal matures and forms cornified palatal
plates. The molars have no predecessors (Simpson 1929), but the
premolar may undergo replacement.
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Since at most a single tooth is replaced in monotrernes and the
"functional" dentition deteriorates early in life, this order may not
be considered functionally diphyodont.
Marsupialia
Marsupial neonates are extremely altricial. Accordingly, they
are all toothless at birth. The earliest teeth to erupt in
macropodids (kangaroos and wallabies) are the adult incisors, but in
the Virginia opossum (Didelphis vlrginianus) the deciduous premolar
comes in first (Table 1); by the end of pouch life the young of both
species have at least a portion of the adult dentition.
For a long time, marsupial dentition has caused debate in terms
of homology. Shortly after his original proposal that milk and
permanent dentitions in placental mammals are homologous with each
other, Owen (1849) sought to extend this scheme to marsupials as
well. He noted that the great gray kangaroo Macropus glganteus
replaces two functional premolars with a single tooth per quadrant,
and claimed in addition that small deciduous milk incisors (3 upper,
I lower) and canines (upper only) are shed when the young leaves the
pouch. The small deciduous teeth and the two replaced premolars Owen
interpreted as homologous with the milk dentition of placentals,
while the functional adult dentition correpsonds to the placental
permanent set.
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permission of the copyright owner. Further
Table 1. Dental ontogeny, length of pouch life, and diet in Marsupialia.
Species Reference First tooth (age) Size of functional Age at premolar Length of Diet
replaced premolar replacement pouch life
Didelphidae:
Didelphis virginiana 5,6 Last premolar large 28 wk 8 wk 0
(9 wk.)
Desyuridae:
Dasyurus maculatus 1,11 none 18 wk C
1,11 "long persistent" 9 wk C
Myrmecobiidae:
Myrmecobius fasciatus 10 small never shed I
TRy
hylacinidae
Tbylacinus c 1 very small "infancy" C
Peramelidae:
1,11 large "functional till (P. nasuta: I
CD half grown" 8 wk.)
■o
3 Macrotis sp. 1 large "long persistent" C
Q. Phalangeridae:
C Trichosurus vulpecula 1 small "not long H oo
o
I—H
o' persistent"
3 Petauridae:
T3 1,8 small "shed early" 16 wk H
S Macropodidae:
Macropus eugenii 3 Incisors & last large 150 wk 36 wk li
O 2 premolars
Q. (18 wk)
Macropus giganteus 7,8 large 78 wk 43 wk H
Macropus parma 4,8 Lower incisor large 200 wk 30 wk H
(19 wk)
Macropus robustus 2 Incisors (15 wk) large 156 wk 31 wk H
T3 Setonix brachyurus 9 Lower incisors large 101 wk 25 wk H
CD
(18 wk)
Phascolarctidae:
(/) Phascolarctos cinereus 1 none 26 wk
in H
* References: 1) Brazenor 1950, 2) Ealey 1967, 3) Inns 1982, 4) Maynes 1972, 5) McCrady 1938, 6) Petrides 1949,
+ 7) Poole 1982, 8) Sharman et al 1966, 9) Shield 1968, 10) Tate 1951, 11) Walker 1975.
Diet: C = Carnivorous, H = Herbivorous, I « Insectivorous, 0 = Qnnivorous.
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Leche (1892, 1895), from histological investigations of timing
and position of tooth bud development, disagreed with Owen and deemed
the persistent dentition homologous with milk teeth of placentals. An
exception was the replacement premolar which he interpreted as the
sole functional remnant of a permanent set. Woodward (1896a)
recognized "traces of" four mammalian tooth sets (premilk, milk,
permanent, postpermanent). He assigned most of the functional
dentition of marsupials to the milk set as did Leche. Uncalcified or
weakly calcified tooth vestiges observable in several genera he
called "premilk" teeth, while the permanent set he said was
represented only by "a series of lingually placed swellings"
(1896a:287).
Bensley (1903) returned to a version of Owen's interpretation,
recognizing only the posteriormost premolar as the remnant of a milk
dentition and all other functional teeth as part of the "permanent"
set. More recent work by Berkovitz (1966, 1968a, 1968b, 1972) follows
this interpretation and includes vestigial incisors and canines as
milk tooth remnants as did Owen.
Finally, Fosse (1969a, 1969b) and Fosse and Risnes (1972a,
1972b) tentatively suggested that marsupial tooth replacement may not
correspond to that in placentals at all. This position was argued
more forcefully by Archer (1978), who proposed that marsupial cheek
teeth basically consist of a premolar series (three teeth in
ancestral forms) and a molar series (five teeth, the first of which
10
is commonly shed) with no placental-type milk dentition nor
replacement of postcanines.
However homology is interpreted, the situation is quite
uncomplicated when only the functional dentition is considered (Table
2). In no known marsupial is there more than one functional tooth
replaced per quadrant. This is always a cheek tooth and is commonly
designated the last premolar. There is considerable variation,
however, in the degree of devlopment of this tooth and the timing of
its replacement (Table 1). Small premolars may be replaced during
pouch life or may never function, while large ones may be retained
for years.
Size of the premolar and its time of replacement are relatively
consistent within a family. Ride (1970) proposed that the marsupials
be split into four orders partly on the basis of dentition. It might
be expected that tooth replacement would show a stable pattern within
each of these orders, but such is not the case for Marsupicarnivora
(Didelphidae, Dasyuridae, Thylacinidae, and Myrmecobiidae) and
Diprotodonta (Macropodidae, Phalangeridae, Petauridae), each of which
has both early-replacing and late-replacing forms (Table 1). Neither
is there a clear correspondence of tooth replacement with diet. For
both carnivorous and herbivorous habits there are early- and
late-replacing species; however, in both groups, replaced premolars
are decidedly molariform. The functional significance of molariform
’’premolars" is clear, since these are the teeth farthest back in the
Table 2. Functional and non-functional dentition in Marsupialia.
Species Reference Non-functional Functional Adult dentition

teeth resorbed teeth shed Up (I C P H) Low (I C P M)
Didelphidae:
Didelphis virginiana McCrady 1938 Last premolar 5 1 3 4 4 13 4
Dasyuridae:
Dasyurus maculatus Brazenor 1950 None 4 1 2 4 3 12 4
Dasyucus sp. Woodward 1896 Up 1 I, 1 C, 1 P 4 1 74 3 17 4
Low 1 I, 1 C, 1 P
Ehascogale tapoatafa Brazenor 1950 Last premolar 4 1 3 4 3 13 4
fhascogale sp. Woodward 1896 Up 4 I, 1 C 4 1 ? 4 3 17 4
Low 4 I, 1 C
Sminthopsis Fosse 1969a & b Last premoiar 4 1 3 4 3 1 3 4
crassicaudatus
Antechinus stuartii Fosse 1969a Last premolar 4 1 3 4 3 13 4
Myrroecobiidae:
Myrmecobius fasciatus Woodward 1896, Up 4 I, 1 C 4 1 4 4 4 14 4
Tate 1951 Low 2 I, 1 C (Last premolar never shed)
Thylacinidae:
Ihylacinus cynocephalus Brazenor 1950 Last P shed in 4 1 3 4 3 13 4
infancy
Peramelidae:
Chaeropus sp. Bensley 1903 Last premolar '? 1 3 4 3 1 3 4
Isoodon obesulus Fosse & Risnes Last premolar 5 1 3 3 3 13 3
1972a & b
Perameles bougainville Brazenor 1950 Last premolar 5 1 3 4 3 13 4
Perameles gunnii Fosse & Risnes Last premolar 5 1 2 4 3 12 4
1972a
Thylacomyidae:
Macrotis sp. Bensley 1903 Last premolar ? 1 3 4 3 13 4
Phalangeridae:
Fhalanger orientalis Berkovitz 1968a Up 2 1 ? 1 ? ? 70 7 7
Low 3 I
Phalanger sp. Woodward 1896 Up 3 1
Low 1 I
Trichosurus vulpecula Berkovitz 1968b, Up 4 I, 0 C, 1 P Last premolar 3 1 2 4 2 0 14
Brazenor 1950 Low 2 I, 1 C, 2 P
Petauridae:
Pseudocheirus peregrinus Brazenor 1950 Last premoiar 3 1 34 20 3 4
Table 2 (contd).
Species Reference Non-functional Functional Adult dentition

teeth resorbed teeth shed Up (I C P M) Low (I C
Macropodidae:
Aepyprymnus rufescens uerkovitz 1968a Up 2-3 :L 7 7 7 7 7 7 7 7
Low 2-4 I
Macropus eugenii Berkovitz 1972 Up 2 I, 1 c One P replaces 3 0 1 4 1 0 1 4
Low 2 I , 0 c last 2 premolars
Macropus giganteus Owen 1849 Up 3 I, 1 C, 1 P One P replaces 3 0 1 4 1 0 1 4
Low 1 I,, 0 C, 1 P last 2 premolars
Macropus parma Maynes 1972 Upper C, 3 0 1 4 1 0 1 4
last 2 premolars
Macropus robustus Ealey 1967 One P replaces 3 0 1 4 1 0 1 4
last 2 premolars
Patrogale penicillata Berkovitz 1968a Up 3 I 7 7 7 7 7 7 7 7
Low 2 I
Setonix brachyurus Berkovitz 1966, Up 2 I One P replaces 3 0 1 4 1 0 1 4
1968a Low 2 I last 2 premolars
Thylogale billardieri Berkovitz 1968a Up 3 I 7 7 7 7 7 7 7 7
Low 3 I
Phascolarctidae:
Ehascolarctos cinereus Brazenor 1950 Up 1 P, Low 1 P None 3 1 1 4 1 0 1 4
Vombatidae:
Vombatus ursinus Rose 1893 Up 4 I, 1 C Last premolar 1 0 1 4 1 0 1 4
Low 4 I, 1 C
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jaw and are performing the job of grinding or crushing before the
molars erupt. Although this morphology is consistent with Archer’s
(1978) suggestion that these last "premolars" are indeed molars, it
is often true in placental mammals as well that last milk premolars
are particularly molariform (Butler 1939).
For the most part, then, marsupials function through life with
only the adult set of teeth. Two schemes for extending the usefulness
of the functional dentition have arisen in marsupials: evergrowing
cheek teeth (in wombats, Vombatidae) and ’’molar progression’’ (in many
macropodids). In "molar progression," posterior molars migrate
forward in the jaw as the anterior ones wear down and are eventually
shed (Sanson 1980), but the teeth are not replaced from underneath.
With a maximum of one tooth per jaw being replaced, it is clear that
the Marsupialia cannot be considered functionally diphyodont.
Edentata
The anteaters (Myrmecophagidae) completely lack functional teeth
after birth. Rose (1892a) described tooth buds in Myrmecophaga
embryos which he interpreted as representing two tooth sets, but in
an embryo of tridactlya and another of Tamandua tetradactyla,
Leche (1892, 1895) found no evidence whatever of tooth development.
Sloths (Bradypodidae, Choloepidae) are born with a set of
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erupted enamelless teeth which they retain through life (Leche 1895,
Veselovsky 1966). The five upper teeth of Choloepus are usually
considered to be a canine and four unspecified cheek teeth, while the
four lowers are cheek teeth (Grasse 1955); there is no evidence to
indicate whether these functional teeth are homologous with the
permanent or the milk set of other eutherians. The situation in the
Bradypodidae is slightly more complicated. Bradypus erupts five upper
and five lower teeth; Grasse (1955) considered the upper row as
similar to those in Choloepus; the lowers he interpreted as a canine
and four cheek teeth, but the putative canine is shed early and not
replaced. In an embryo of Bradypus, Leche (1895) found yet another
tooth developing anterior to the functional upper row. This tooth was
calcified but apparently would not have developed further. From its
position and state of development, Leche deemed it a milk tooth, the
last remnant of a former diphyodonty. The functional set, he
concluded, is homologous with the permanent dentition of eutherians.
Why sloths should have teeth at birth is not clear, and the
available information on feeding ontogeny is not very helpful.
Goffart (1971) described Bradypus variegatus as eating solids at four
days, but Beebe (1926) found B. cuculliger does not begin to eat
leaves until five weeks of age (though it takes regurgitate from the
mother in the third week). Choloepus hoffmanni eats solids at four to
five weeks (Goffart 1971) and C. didactylus at two and a half months
(Veselovsky 1966). Reported age at weaning varies from nine weeks for
B. cuculliger (Beebe 1926) to five months for CL_ didactylus
15
(Veselovsky 1966) and up to one to two years for B. variegatus
(Goffart 1971). It appears that sloths are probably dependent on milk
for at least a month. Perhaps the early eruption of teeth is simply a
manifestation of selection for general precocity at birth; newborn
sloths are fully furred and have their eyes open (Goffart 1971).
However, if Leche*s (1895) conclusion is correct and sloth teeth are
indeed homologous with other eutherians* permanent teeth, their rate
of development has been accelerated beyond what would be expected in
a generally precocious young.
Armadillos (Dasypodidae) were considered monophyodont until
Gervais (1855) described and figured functional tooth replacement in
a specimen of Dasypus novemcinctus. Although the permanent teeth are
simple cylinders, the milk teeth in this species are bilobed and
double-rooted. The permanent teeth erupt in wedge fashion between the
lobes of the milk teeth, eventually splitting them in two (Gervais
1855). In this way function of the teeth is apparently maintained
throughout the replacement process. Gervais* figure shows seven milk
teeth being replaced on each side of upper and lower jaws, with a
single additional tooth at the back of the jaw erupted but showing no
evidence of replacement. From examination of more specimens, Flower
(1868) confirmed Gervais* observations, and noted that tooth
replacement must take place when the animals are nearly full size. In
addition to the functional teeth, Leche (1892,1895) found five
calcified toothlets and three less developed buds farther forward in
the lower jaw of D. novemcinctus and D. hybridus; Weber (1928)
16
interpreted these vestigial teeth as part of the milk dentition.
The Edentata are generally considered a natural group on the
basis of features other than dentition. In tooth structure and
replacement phenomena, however, they are more diverse than any other
order. The sloths are functionally monophyodont, the anteaters are
genuinely edentulous, and only the armadillos (if in fact all follow
the pattern of Dasypus) are functionally diphyodont (Table 3).
Insectivora
Recent taxonomic arrangements have reduced this formerly diverse
order to a more strictly defined set (Butler 1972), but the
Insectivora sensu strictu are still not uniform with respect to tooth
replacement (Table 3). All the species that have been examined
histologically develop two sets of tooth buds; but the shrews
(Soricidae) resorb all or nearly all rudiments of milk dentition
before birth and so are functionally monophyodont. Friant (1965)
reported two erupted milk premolars in a Pleistocene shrew and
possibly in a modern Sorex, but these are tiny teeth and of doubtful
functional significance even if they pierced the gums. True moles
(Talpidae) fall into two groups. Some have only a rudimentary milk
dentition which is either resorbed without erupting or consists of
tiny spicule-like teeth of short duration. Other moles develop a
robust and apparently functional set of milk teeth. (Ziegler (1971b)
Table 3. Functional milk and replacement dentition in diphyodont eutherian mammals. Taxonomy
follows Honacki et al (1982). Numbers of functional teeth (incisors, canines, premolars, molars)
on each side of jaw are given for upper and lower jaws of milk and permanent sets. Although "PI"
and molars are conventionally considered late-erupting members of the milk dentition (see text),
they are, in this functional classification, included as members of the permanent dentition. For
species in which additional nonfunctional teeth are known to develop, these are listed in parentheses
after the functional formula.
Species Reference Milk upper Milk lower Perm, upper Perm, lower
(I C P ) (I C P) (I C P M) (I C P M)
EDENTATA
Dasypodidae:
Dasyjxjs novemcinctus Flower 1868 0 0 7 0 0 7 0 0 7 1 0 0 7 1
INSECTIVORA
Solenodontidae:
Solenodon sp. Thomas 1882 3 1 2 3 1 2 3 1 3 3 3 1 3 3
Tenrecidae:
Echinops telfairi Thomas 1882 2 1 3 21 3 2 1 3 2 2 1 3 2
Hemicentetes semi'spin s Leche 1897 3 1 1? ? 7 ? 2 ? ? ? 2 ? ? ?
Oryzorictes hova Forsyth Major 1897 3 1 3 3 1 3 3 1 3 3? 3 1 3 37
Oryzorictes Forsyth Major 1897 3 1 3 3 1 3 3 1 3 3 3 1 3 3
tetradactylus
Microgale cowani Forsyth Major 1897 3 1 3 3 1 3 3 1 3 3 3 1 3 3
Microgale dobsoni Forsyth Major 1897 3 1 3 3 1 3 3 1 3 3 3 1 3 3
PotamogaTe sp. Leche 1907 3 1 3 3 1 3 3 1 3 3 3 1 3 3
Setifer setosus Woodward 1896 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Tenrec ecaudatus Woodward 1896 2 1 3 3 1 3 3 1 3 3-4 3 1 3 3-4
Chrysochloridae:
Chrysochloris sp. Leche 1907 3 1 3 3 1 3 3 1 3 2-3 3 1 3 2-3
Eremitalpa granti Kindahl 1963 3 1 2 3 1 2 3 1 2 3 3 1 2 3
Erinaceidae:
Echinosoirex gymnurus Woodward 1896 2 1 3(1 0 0) 3 1 2(0 0 1) 3 1 3 3 3 1 3 3
Erinaceus europaeus Woodward 1896, 2 0 2(1 1 1) 1 0 1(2 1 1) 3 1 3 3 2 1 2 3
Kindahl 1959
Hylomys sp. Leche 1902 2 1 3(1 0 0) 3 1 3 3 1 4 3 3 1 4 3
Table 3 (contd).
(I C P ) (I C P) (I C P M) (I C P M)
Soricidae:
Cryptotis sp. Choate 1970 None None 7 ? 7 7 7 7 7 7
Sorex araneus Kindahl' 1959 (2 1 4) (2 1 1) 3 1 3 3 2 0 1 3(0 1 0 0)
Sorex sp. Woodward 1896 (2 1 2) (1 o 1) 3 0 1 3 3 1 7 7.
Suncus varilla Kindahl 1959 & 1967 (2 1 2) (2 1 1) 3 1 2 3 2 0 1 3(0 1 0 0)
Talpidae:
Condylura cristata Leche 1895 (2 0 3) (3 0 3) 3 1 4 3 3 1 4 3 00
Desmana sp. Ziegler 1971b 3 1 3 ■ 3 1 3 3 1 4 3 3 1 4 3

Galemys sp. Ziegler 1971b 3 1 3 3 1 3 3 1 4 3 3 1 4 3
Neurotrichus sp. Ziegler 1971b 3 1 3 3 1 3 3 1 4 3 3 1 4 3
Parascalops breweri Eadie 1944, Ziegler (3 1 3-4) (3 1 3-4) 3 1 4 3 3 1 4 3
1971b
Scalopus aquaticus Leche 1895; see also (3 1 3)’ (3 1 2) 3 1 3 3 2 0 3 3(1 1 0 0)
Conaway & Landry 1958
Scapanus latimanus Ziegler 1972a & b 0 0 3(3 1 0) 0 0 3(3 1 0) 3 1 4 3 3 1 4 3
Scaptonyx sp. Ziegler 1971b 3 1 3 2 1 3 3 1 4 7 2 1 4 7
Talpa europaea Sicher 1917, Ziegler 2 1 2(0 0 1) 2 1 1(1 0 2) 3 1 4 3 2 1 2 3(1 0 2 0)
1971b
Uropsilus sp. Ziegler 1971b 2 1 2-3 1-2 1 2-3 2 1 3-4 7 1-2 1 3-4 7
Urotrichus pilirostris Ihomas 1908, Ziegler 3 1 21 2 1 2 3 1 3 3 2 1 3 3
1971b
Urotrichus talpoides Leche 1897 3 1 3 2 0 3 3 1 3 3 2 O' 3 3
SCANDENTIA
Tupaiidae:
Tupaia Rlis Shigehara 1980 2 1 3 3 1 3 2 1 3 3 3 1 3 3
'lupaia iavanica Butler 1980, Kindahl 2 1 3 3 1 3 2 1 3 3 3 1 3 3
1957
Table 3 (contd).
(I C P) (I C P) (I C P M) (I C P M)
DERMOPTERA
Cynocephalidae:
Cynocephalus variegatus Dependorf 1896 2 0 3 2 1 3 2 0 3 3 2 1 3 3
Cynocephalus volans Dependorf 1896 2 0 3 2 1 3 2 0 3 3 2 1 3 3
CHIROFTERA
Pteropodidae:
Dobsonia peroni Temminck 1827 2 ? ? 2 1 7 7 2 2
2 7 2 3 3
Dobsonia praedatrix Andersen 1912 2 1 2 2 1 2 1 1 2
2 1 1 3 3
Cynopterus brachyotis Miller 1907 2 1 2 2 1 2 7 7 7 7 7 7 7 7
Cynopterus sphinx Leche 1878 2 1 2 2 1 2 7 7 3 1 7 7 3 2
Epomophorus gambianus Leche 1878 2 1 2 2 1 2 7 7 2 1 7 7 3 2
Epomophorus wahlbergi Sowler 1980 2 1 2 2 1 2 2 1 2 1 2 1 3 2
Eporoops franqueti Friant 1965 2 1 2 2 1 2 7 7 ? 7 7 7 7 7
Macroglossus minimus Leche 1878 2 1 2 2 1 2 7 7 3 2 7 7 3 3
Nyctimene cephalotes Leche 1878 1 1 2 1 1 2 1 0 3 1 0 0 3 2
Pteropus alecto Leche 1878 2 1 2 2 1 2 7 7 3 2 7 7 3 3
Pteropus poliocephalus Leche 1878 2 1 2 2 1 2 7 7 3 2 7 7 3 3
Pteropus samoensis Leche 1878 2 1 2 2 1 2 7 7 3 2 7 7 3 3
Rousettus aegyptiacus Friant 1951 2 1 r 2 1 1 2 1 U 2 2 1 3 3
Rousettus amplexicaudatus Leche 1878 2 1 2 2 1 2 7 7 3 2 7 7 3 3
Emballonuridae:
Peropteryx macrotis Leche 1878 2 1 2 2 1 2 1 1 2 3 3 1 2 3
Megadermatidae:
Lavia frons Dorst 1953 None None 0 1 1 3 2 1 2 3
Rhinolophidae:
Hipposideros caffer Gaunt 1967 (1 1 3) (2 1 1) 1 1 2 3 2 1 2 3
Rhinolophus hipposideros Leche 1877 (1 ? 2) (? ? 2) 1 1 2 7 2 1 3 7
Table 3 (contd).
(I C p) (I C P) (I C P M) (I C P M)
Phyllostomidae:
Ametrida centurio Leche 1878 2 1 2 0 1 2 2 1 2 3 2 1 2 3
Artibeus jamaicetTsis Leche 1878 2 1 2 1 1 1 2 1 2 2 2 1 2 3
Carollia brevicauda Leche 1878 2 1 3 2 1 2 2 1 2 3 2 1 2 3
Carollia perspicirTata Kleiman & Davis 1979 2 1 3 0 1 2(2 0 0) ? ? ? 7 7 7 7 7
Choeronyctens mexicana Phillips 1971, Stains 2 1 3 2 1 2 2 1 2 3 0 1 3 3
& Baker 1954
Chrotopterus auritus Leche 1878 2 1 1 ? ? 2 2 1 2 3 2 1 3 3
Desmodus rotundus Miller 1896 2 0 0(0 1 1) 2 0 0(0 1 2) 2 1 3 0 2 1 3 0
Dlphylla ecaudata Birney & Timm 1975 2 1 2 2 1 2 2 1 1 2 2 1 2 2
Glossophaga soricina Leche 1878 2 1 3 2 1 2 2 1 2 3 2 3 3
1
Leptonycteris sanborni Phillips 1971 2 1 3 2 1 2 2 1 2 2 2 3 2
1
Nelson 1966 2 1 2 2 1 2 ? ? 7 7 7 7 7 7
Macrotus waterhousii
rtiyllostomus sp. Miller 1907 2 1 2 2 1 2 7 7 ? 7 7 7 7 7
Sturnira lilium Leche 1877 2 1 2 2 1 2 2 0 2 3 2 0 2 3
Tonatia silvicola Dorst 1957 2 1 3 ? ? ? 2 1 2 7 7 7 7 7
Vespertilionidae:
Antrozous pallidus Orr 1954 2 1 2 3 1 2 1 1 1 3 2 1 2 3
Eptesicus fuscus Miller 1907, Baker 2 1 2 3 1 2 2 1 1 3 3 1 2 3
1956
Eptesicus serotinus Kleiman 1969 2 1 2 3 1 2 2 1 1 3 3 1 3
2
3 1 2 1 1 2 7 3 1 7
2
Lasiurus sp. Miller 1907 2 1 2
Mlniqpterus schreibersi Friant 1963 2 1 2 3 1 2 2 1 2 3 3 1 3 3
Myotls albescens Webster 1981 2 1 2 3 1 2 2 3
1 3 3 1 3 3
Myotls blythil Friant 1963 2 1 2 3 1 2 2 3
1 3 3 1 3 3
Myotis daubentoni Leche 1877 2 1 2 *3 1 2 ? ? 3 3 7 7 3 3
Myotis luv~.ifufiui~ Fenton 1970 2 1 2 3 1 2 2 1 3 3 3 1 3 3
Myotis myotis Kratky 1970 2 1 2 2 1 2 2 1 3 3 3 1 3 3
Table 3 (contd).
Myotis mvstacinus Leche 1878 2 1 2 2 1 2 7 7 3 3 7 7 3 3

Myotis vivesi Reeder 1963 2 1 2 3 1 2 2 1 3 3 3 1 3 3
Nyctalus leisleri Matthews 1950 2 1 2 3 1 2 7 7 7 7 7 7 7 7
Nyctalus noctula Kleiman 1969 2 1 2 3 1 2 2 1 2 3 3 1 2 3
Nvcticeius humeralis Jones 1967 2 1 2 3 1 2 1 1 1 3 3 1 2 3
PipistreTIus kuhlii Dorst 1949 2 1 2 3 1 2 2 1 2 3 3 1 2 3
Pipistrellus nanus Dorst 1949 2 1 2 3 1 2 7 7 7 7 7 7 7 7
Pipistrellus natfiusii Weber 1928 2 1 2 2 1 2 7 7 2 3 7 7 2 3
Pipistrellus pipistrellus Kleiman 1969 2 1 2 3 1 2 2 1 2 3 3 1 2 3
Plecotus auritus Leche 1877 2 1 2 3 1 2 7 7 2 3 7 7 3 3
Plecotus rafinesquii Jones & Suttkus 1975 2 1 2 3 1 2 2 1 2 3 3- 1 3 3
Scotophilus heathi Peterson 1968 2 7 7 7 7 7 1 1 7 7 3 1 7 7
Vespertiilo murinus Leche 1878 2 1 2 3 1 2 7 7 3 3 7 7 3 3
Vespertiflo superans Yoshiyuki 1968 2 1 1 3 1 2 7 7 7 7 7 7 7 7
Molossidae:
Molossus sp. Temminck 1827 7 7 7 2-3 7 7 7 7 7 7 7 7 7 7
Promops nasutus Spillman 1927 2 1 1 7 7 7 7 7 7 7 7 7 7 7
Tadarida brasiliensis Miller 1907 2 1 2 3 1 2 2 1 1 7 3 1 2 7
Tadarida condylura Dorst 1957 2 1 2 3 1 1 7 7 7 7 7 7 7 7
Archaeonycteridae+:
Archaeonyc teris Revilliod 1917 7 7 7 7 7 2 2 1 7 7 7 1 2 3
PRIMATES
Adapidae+:
Notharctus sp. Hill 1953 2 1 4 1 2 4 2 1 4 3 2 1 4 3
Leptadapis magnus Leche 1897 2 1 3 2 1 3 2 1 4 3 2 1 4 3
Table 3 (contd).
Species Reference Milk upper Milk lower Perm, upper Perm. lower
Lemuridae:
Lemur sp. Leche 1897 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Lepilemur mustelinus Leche 1897j see also 1-2 1 3 2 1 3 0 1 3 3 2 1 3 3
Schwartz 1975a
Hapalemur sp. Gingerich 1979, 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Swindler 1976
Megaladapidae+:
Megaladapis sp. Hill 1953 ? 1 3 3 ? 3 0 1 3 3 2 1 3 3
Daubentoniidae:
Daubentonia Peters 1866, Remane 2 1 2 2 0 2 1 0 1 3 1 0 0 3
madagascariensis 1962
Indriiaae:
Indri indri Friant 1935, Swindler 2 1 3 2 1 3 2 1 2 3 1 1 2 3
1976
Propithecus verreauxi Milne-Edwards & 2 1 2 2 1 3 2 1 2 3 2 0 2 3
Grandidier 1875-97
Propithecus
opi diadema
_____ Leche 1897 2 1 3 2 1 3 2 1 2 3 1 1 2 3
Licnanotus laniger Milne-Edwards & 2 1 2 2 1 3 2 1 2 3 2 0 2 3
Grandidier 1875-97
Archaeolemuridae+:
Archaeolemur sp. Hill 1953 2 1 3 2 1 3 2 1 3- 3 2 0 3 3
Hadroptthecus sp. Lamberton 1938 2 1 3 2 0 3 2 1 3 3 1 1 3 3
Cheirogaleidae:
Cheirogaleus major Schwartz 1975b, 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Swindler 1976
Microcebus murinus Schwartz 1975a 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Ftianer furcifer Schwartz 1975b 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Table 3 (contd).
^ (I C P) (I C P) (I C P M) (I C P M)
Lorisidae:
Schwartz 1975b, 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Loris tardigradus
Swindler 1976
Schwartz 1975b, 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Nycticebus coucang
Swindler 1976
James 1960 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Perodicticus potto
Galagxdae:
Leche 1896 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Galago demidovii
Tarsiidae:
Luckett & Maier 1982, 0 1 2(2 1) 1 1 2(0 0 1) 2 1 3 3 2 1 3 3
Tarsius bancanus
Greiner 1929
Tarsius spectrum Leche 1897 2 1 2(0 1) 1 1 2(0 0 1) 2 1 3 3 1 1 3 3
Tarsius syrichta Dahlberg 1948 0 1 2 0 1 2 2 1 3 3 1 1 3 3
Callithricidae:
Cnllithrix argentata Hershkovitz 1977 2 1 3? 2 1 3 2 1 3 2 2 1 3 2
Callithrfx jacchus Johnston et al 1970 2 1 3 2 1 3 2 1 3 2 2 1 3 2
Saguinus nigricollis Chase & Cooper 1969, 2 1 3 2 1 3 2 1 3 2 2 1 3 2
Tapper & Severson 1971
Cebidae:
Alouatta sp. Schultz 1960 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Ateles sp. Schultz 1960 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Cebus sp. Della Serra 1952, 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Schultz 1960
Long & Cooper 1968 2 1 3 2 1 3 2 1 3 3 2 1 3 3
Saimiri sciureus
Cercopi thecidae: o 3
Cercopithecus aethiops Ockerse 1959 2 1 2 2 1 2 2 1 £ 2 1 2 3
Cercopithecus mona Selenka 1899 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Macaca fascicularis Spiegel 1934 & 1952 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Macaca mulatta Hurme & Van Wagenen 2 1 2 1 2 2 1 2 3 2 1 2 3
1953 & 1961
Table 3 (contd).
Nasalis larvatus Schultz 1942 2 1 2 2 1 2 2 1 2 3 2 1 2 3

Papio hamadryas Freedman 1962 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Hylobatidaes
Hylobates sp. Schultz 1944, 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Selenka 1898
Pongidae:
Gorilla gorilla Randall 1943, 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Selenka 1899
Pan paniscus Johanson 1974 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Pan troglodytes Nissen & Riesen 2 1 2 2 1 2 2 1 2 3 2 1 2 3
1945 & 1964
Pongo pygmaeus Schultz 1941 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Hominidae:
Homo sapiens Kronfeld 1935, 2 1 2 2 1 2 2 1 2 3 2 1 2 3
Miller et al 1965
to
CARNIVORA
Canidae;
Canis familiaris Evans 1960 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Canis mesomelas~ Lombaard 1971 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Otoevon megalotis Guilday 1962 3 1 3 3 1 3 3 1 4 3 3 1 4 4
Vulpes macrotis Efeoscue 1962 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Vulpes vulpes Linhart 1968 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Ursidae:
Tremarctos omatus Dittrich 1960 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Ursus americanus Dittrich 1960, 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Marks & Erickson 1966
Table 3 (contd).
Specie.1; Reference Milk upper Milk lower Perm, upper Perm, lower
Ursus arctos Couturier 1954 3 1 3 3 1 3 3 1 4 2 3 1 4 3

Ursus maritimus Dittrich 1960 2 1 3(1 0 3 1 3 3 1 4 2 3 1 4 3
Ursus thibetanus Dittrich 1960 3 1 3 3 1 3 3 1 4 2 3 1 4 3
Procyonidae:
Bassariscus astutus Toweill & Toweili 1978 3 1 2? 3 1 2? 3 1 4 2 3 1 4 2
Procyon lotor Montgomery 1964 3 1 4 3 1 4 3 1 4 2 3 1 4 2
Mustelidae:
Enhydra lutris Fisher 1941, Jacobi 3 1 3 3 1 3 3 1 3 1 2 1 3 2
1938, Kenyon 1969.
Scheffer 1951
Gulo r u Io Krott 1959 2 1 3 ? 1 3(3? 0 0) 3 1 4 1 3 1 4 2
Lutreola lutreola Moshonkin 1979 1 1 3 0 1 3 3 1 3 1 3 1 3 2
Martes Fuina Habermehl & Rottcher 3 1 3 3 1 3 3 1 4 1 3 1 4 2
1967
tteles meles Corbet & Southern 1977 (3 1 3)7 (3 1 3)7 3 1 3-4 1 3 1 3-■4 2
Mellivora capensis Lydekker 1912 3 1 3 ? 7 ? 3 1 3 1 7 7 7 7
Mephltis~roepnitis Verts 1967 None None 3 1 3 1 3 1 3 2
Mustela frenata Hall 1951 7 1 3 7 1 3 3 1 3 1 3 1 3 2
Mustela nivalis Heidt et al 1968 2 1 1? 2 1 2? 3 1 3 1 3 1 3 2
Mustela putorius Berkovitz 1973, 3 1 3 0 1 3(3 0 0) 3 1 3 1 3 1 3 2
Mazak 1963
Mustela vison Aulerich & Swindler 3 1 3 1 1 3(2 0 0) 3 1 3 1 3 1 3 2
1968
Taxidea taxus Long 1965 1-2 1 3 0 1 3 3 1 3 1 3 1 3 2
Viverridae:
Chrotoeale owstoni Thomas 1912 3 1 4? 3 1 4? ? 7 7 ? 7 7 7
Cynogale jp. Mivart 1882 3 1 3 3 1 3 3 i 4 2 3 1 4 2
Genetta sp. Mivart 1882 7 7 3 7 7 3 3 l 4 2 3 1 4 2
Table 3(contd)
(I C P ) (I C P) (I C P M) (I C P M)
Nandinia binotata Charles-Dominique ‘ 3 1 4 3 1 4 3 1 4 2 3 1 4 2

1978
Paradoxurus sp. Mivart 1882 ? ? 3 ? 7 3 3 1 4 2 3 1 4 2
Viverra sp. Mivart 1882 3 1 3 3 1 3 3 1 4 2 3 1 4 2
Herpestidae:
Suricata suricatta Ducker 1962 3 1 3 3 1 3 3 1 3 2 3 1 3 2
Protelidae:
Proteles cristatus Gingerich 1974 3 1 3 3 1 3 3 1 4 1 3 1 3 1
Hyaenidae:
Crocuta crocuta Pournelle 1965 3 1 ? 3 1 7 3 1 4 1 3 1 3 1
Felidae:
Felis concolor Schneider 1959 3 1 3 3 1 2 3 1 3 1 3 1 2 1
Felis silvestris Meyer-Holzapfel 1968 3 1 3 3 1 3 3 1 3 1 3 1 2 1
Lynx canadensis Saunders 1961 3 1 2 3 1 2 3 1 2 1 3 1 2 1
Panthera leo Smuts et al 1978 3 1 3 3 1 2 3 1 3 1 3 1 2 1
Panthera onca Schneider 1959 3 1 ? 3 1 7 7 7 7 7 7 7 7 7
Panthera pardus Schneider 1959 3 1 2 3 1 2 7 7 7 7 7 7 7 7
Panthera tigris Volf 1972 3 1 3 3 1 2 7 7 7 7 7 7 7 7
Panthera uncia Pocock 1916 3 1 3 3 1 7 3 1 3 1 3 1 2 1
Otariidae:
Arctocephalus pusillus Rand 1950 (3 1 3) (2 1 3) 3 1 6 2 1 5 *
Callorhlnus ursinus Chiasson 1957, Kubota 0 1 0(3 0 3) 0 1 0(2 0 3) 3 1 4 2 2 1 4 1
1963, Scheffer &
Kraus 1964
Eumetopias jubatus Spalding 1966, (3 1 3) (2 1 3) 3 1 5* 2 1 5 *
Scheffer 1967
Odobenidae:
Odobenus rosnarus Cobb 1933, Allen (3 1 3-4) (3 1 3-4) 0 -•2 1 8 1 0 1 3 1
1880, Kellogg 1922
Table 3 (contd).
Phocidae:
Cystophora cristata Reinhardt 1865 (2 1 3)7 (1? 1 3)7 2 14-6 * 1 1 4-6*
Lobodon sp. Scheffer 1958 (2 1 3) (2 3)1 7 7 7 7 7 7 7 7
Mirounga angustrostris Briggs 1974 (? 1 3) (7 3)1 2 1 4 1-2 1 1 4 1-2
Mlrounga leonina Flower 1881, Laws 1953 (2 1 3) (I 3)1 2 1 4 1 1 1 4 1
Monachus monachus King 1956 (2 1 3) (2 3)1 2 1 5* 2 1 5*
Fhoca groenlandica Steenstrup 1861 (3 1 3) (2 3)1 3 1 5* 2 1 5*
Fhoca hispida Steenstrup 1861 (3 1 3) (2 3)1 3 1 5* 2 1 5.*
Fhoca vitulina Van Beneden 1871 (3 1 3) (3? 0? 3) 3 1 5-6 * 2 1 5*
to
•o
CETACEA Not functionally diphyodont; see text
SIRENIA Not functionally diphyodont; see text
PROBOSCIDEA Not functionally diphyodont; see text
PERISSODACIYLA
Equidae:
Equus burchelli Erz 1964 3 1 3 3 1 3 3 1 4 3 3 1 3 3
Equus caballus~ Sisson 1914 3 0 3 3 0 3 3 1 3-4 3 3 1 3 3
Equus zebra Penzhorn 1982 3 1 3 3 1 3 3 1 4 3 3 1 47 3
Tapiridae:
Tapirhs indicus Parker 1882 3 1 4 3 1 3 3 1 4 3 3 1 1 3
Table 3 (contd).
Tapirus pinchaque Blainville 1839-64 3 1 4 ? 7 7 3 1 4 3 3 1 4 3

Tapirus terrestris Blainville 1839-64 3 1 4 3 1 3 3 1 4 3 3 1 4 3
Rhinocerotidae:
Owen 1840 2 0 4 2 0 4 7 7 4 3 7 7 4 3
Diceros bicornis Anderson 1966; see 0 0 4 0 0 4 0 0 4 3 0 0 4 3
also Foster 1965
HYRACOIDEA
Procaviidae:
Procavia capensis Adloff 1903 1 1 4(1 0 0) 2 0 4(1 1 0) 1 1? 4 3 2 0 4 3 00
TUBULIDENTATA Not functionally diphyodont; see text
ARTIODACIYLA
Suidae:
Fhacochoerus aethiopicus Child et al 1965 1 1 2-4 2-3 1 1-3 1 1 1-3 3 2-3 1 1-2 3
Sus scrola Matschke 1967 3 1 3 3 1 3 3 1 4 3 3 1 4 3
Tdysssuidae•
Tayassu taiacu Kirkpatrick & Sowls 2 1 3 3 1 3 2 1 3 3 3 1 3 3
1962
Table 3 (contd).
Camelidae:
Camelus dromedarius Droandi 1936 3 1 3 3 1 3 1 1 3 3 3 1 2 3
Tragulidae:
Tragulus napu Milne-Edwards 1864 0 1 3 7 7 7 0 1 3 3 3 1 3 3
Cervidae:
Alces aIces Bromee-Skuncke 1952 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Capreolus capreolus Baumann 1949 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Cervus dama Bromee-Skuncke 1952 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Cervus elaphus Baunann 1949, Qjimby 0 1 3 3 1 3 0 1 3 3 3 1 3 3
& Gaab 1957
Cervus nippon Kuwano 1929 ? 1 3 3 1 3 0 1 3 3 3 1 3 3
Odocoileus hemionus Rees et al 1966 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Odocoileus virginianus Ryel et al 1961 ? ? ? 3 1 3 0 0 3 3 3 1 3 3
Pangifer Taranaus Miller 1972 7 ? 7 3 1 3 0 1 3 3 3 1 3 3
Giraffidae:
Giraffa Camelopardalis Dagg & Foster 1976 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Bovidae:
Aepyceros melampus Child 1964 ? ? 7 3 1 3 0 0 3 3 3 1 3 3
Ammotragus lervia~ Qgren 1965 ? ? 7 3 1 3 7 7 7 7 3 1 3 3
Antidorcas marsupialis Rautenbach 1971 ? ? 7 3 1 3 7 7 7 7 3 1 3 3
Antllocapra amencana Dow 1952 ? ? 7 3 1 3 7 7 7 7 3 1 3 3
Antilope cervicapra Mungall 1976 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Bison bison Fuller 1959 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Bubalus bubalis Rollinson 1974 0 0 3 3 1 3 7 ? 7 7 3 1 7 3
Capra hircus Nickel et al 1967 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Damaliscus dorcas Ludbrook & Ludbrook 0 o- 3 3 1 3 0 0 3 3 3 1 3 3
1981
Gazella granti Spinage 1976 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Hemitragus lemlahicus Caughley 1965 ? ? 7 3 1 3 7 7 7 7 3 1 3 3
Kobus einpsiprvmnus Spinage 1967 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Oreamnos americanus Brandborg 1955 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Table 3 (contd).
Oreotragus oreotragus Wilson & Child 1965 0 0 3 3 1 3 0 0 3 3 3 1 3 3

Ovibos inoschatus Henrichsen & Grue 0 0 3 3 1 3 0 0 3 3 3 1 3 3
1980
Ovis annion Turcke & Schmincke 9 7 9 3 1 3 ? ? 7 7 3 1 3 3
1965
Ovis aries Koch 1%3 0 3
0 3 1 3 0 3
0 3 3 1 3 3
Ovis canadensis Deming 1952 0 3
0 3 1 3 0 3
0 3 3 1 3 3
3 1 3 9 7 7 7 3 1 3 3
Ovis dalli Henning 1969 0 3
0
Rupricapra rupricapra Baumann 1949 0 3
0 3 1 3 6 0 3 7 3 1 3 7
Sigmoceros lichtensteini Mitchell 1965 0 3
0 3 1 3 0 0 3 3 3 1 3 3
Saiga tatarica Bannikov et al 1961 0 3
0 3 1 3 0 0 3 3 3 1 2 3
Sylvicapra erlmmia Riney & Child 1960 0 3
0 3 1 3 0 0 3 3 3 1 3 3
Syncerus caffer Grimsdell 1973 0 3
0 3 1 3 0 0 3 3 3 1 3 3
Tragelaphus oryx Jeffery & Hanks 1981 ? ? 3 3 1 3 ? 7 3 3 3 1 3 3
Tragelaphus scriptus Wilson & Child 1964 0 0 3 3 1 3 0 0 3 3 3 1 3 3
Tragelaphus strepsiceros Simpson 1966 0 0 3 3 1 3 0 0 3 3 3 1 3 3
PHOLIDOTA Not functionally diphyodont; see text
RODEOTIA
Aplodontidae:
Aplodontia sp. Cederblom 1900 0 0 2 0 0 1 1 0 2 3 1 0 1 3
Sciuridae:
Sciurus lis Miyao 1971 0 0 2 0 0 1 1 0 2 3 1 0 1 3
Spermophilus beecheyi Bryant 1945 0 0 2 0 0 1 1 0 2 3 1 0 1 3
Table 3 (contd).
Spermophilus franklinii Bryant 1945 0 0 2 0 0 1 1 0 2 3 1 0 1 3

Spermophilus townsendii Bryant 1945 0 0 2 0 0 1 1 0 2 3 1 0 1 3
Spermophilus undulatus~ Mitchell & Carsen 1967 0 0 2 0 0 1 1 0 2 3 1 0 1 3
Tamias minimus Bryant 1945 0 0 1 0 0 1 1 0 2 3 1 0 1 3
tamlas striatus Bryant 1945, Allen 0 0 1 0 0 1 1 0 1 3 1 0 1 3
1938
Geomyidae:
Geomys sp. Cederblom 1900 (0 0 1)7 (0 0 1 )? 1 0 1 3 1 0 1 3
Heteromyidae:
Dipodomys sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Perognathus sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Castoridae:
Castor canadensis Cook & Maunton 1954 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Anomaluridae:
Anomalurus sp. Wood 1962 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Pedetidae:
Pedetes sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Cricetidae:
Mesocricetus auratus Keyes & Dale 1944 None None 1 0 0 3 1 0 0 3
Neotoma floridana Hamilton 1953 None None 1 0 0 3 1 0 0 3
Muridae:
Mus musculus Bhaskar et al 1948, (1 0 0) (1 o °> 1 0 0 3 1 0 0 3
Gaunt 1966, Moss-
Salentijn 1978
Rattus norvegicus Hoffman & Schour 1940, (1 0 0) (1 o 0) 1 0 0 3 1 0 0 3
Moss-Salentijn 1978
Gliridae:
Graphiurus sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Myoxus sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Table 3 (contd).
Dipodidae:
Dipus sp. Cederblom 1900 None? None? 1 0 1 3 1 0 0 3
Hystricidae:
Atherurus sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Hystrix sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Eretfuzbntidae:
Erethizon dorsatum Earle & Kranrn 1980 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Sphigeurus sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Caviiaae:
Cavia porcellus Berkovitz 1972 (0 0 1) (0 0 1) 1 0 1 3 1 0 1 3
Hydrocnaeridae:
Hydrochaeris sp. Cederblom 1900 (0 0 1 ) (0 0 1) 1 0 1 3 1 0 1 3
Dasyproctidae: •
Dasyprocta sp. Cederblom 1900 0 0 1 0 0 1 1 0 1 3 1 0 1 3
Chinchillidae:
Chinchilla sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
LaRidium sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Lagostomus sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Capromyidae:
Gapromys sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Myocastoridae:
Hyocastor sp. Aliev 1965 None? None? 1 0 1 3 1 0 1 3
Octodontidae:
Octodon sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Abrocanidae:
Abrocotna sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
Echimyidae:
Dactylomys sp. Cederblom 1900 None? None? 10 13 10 13
Table 3 (contd).
Thryonomyidae:
'Ihryonomys sp. Thomas 1894, Wood (0 0 1) (0 0 1) 1 0 0 ? 1 0 0 ?
1962. Woods 1976
Petromyidae:
Petromus sp. Wood 1962 0 0 1 0 0 1 ? ? ? ? ? ? ? ?
Bathyergidae:
(composite of Bathvergus. Cederblom 1900 None7 None? 1 0 2--5 * 1 0 2--5 *
Georhychus, Heliophobius
and Heterocephalus)
Ctenodactyliaae:
Ctenodactylus sp. Cederblom 1900 None? None? 1 0 0 3 1 0 0 3 OJ
OJ
Pectinator sp. Cederblom 1900 None? None? 1 0 1 3 1 0 1 3
LAG0M0RPHA
Leporidae:
Oryctolagus cuniculus Moss-Salentijn 1978 1 0 3(2 0 0) 0 0 2(2 0 0) 2 0 3 3 1 0 2 3
Sylvilagus floridanus Dice & Dice 1941 1 0 3 0 0 2 2 0 3 3 1 0 2 3
MACROSCELIDEA
Macroscelididae:
Elephantulus Hill 1938 2 1 3 3 1 3 3 1 4 2 3 1 4 3
brachyrhynchus
Elephantulus myurus Kindahl 1958 2 1 3 2 1 3 ? ? 4 ? ? ? 4 ?
Leche 1897, Evans 1942 0 1 3(1 0 0) 3 1 3 1 1 4 2 3 1 4 2
* Premolars and molars reported jointly as cheek teeth or postcanines.

34
describes these patterns and correlates robust milk dentition with
greater degree of fossorial life). The golden moles (Chrysochloridae)
and most hedgehogs are unequivocally diphyodont, but the European
hedgehog Erinaceus has a feeble set of milk teeth with dubious
function.
Information on amount of use of milk teeth in the diphyodont
Insectivores is scanty and inconsistent. No Insectivores are reported
to bear functional teeth at birth (Table 4). The golden moles and
some tenrecs retain their well-formed milk dentitions until their
skulls are nearly adult size (Table 4). The milk teeth of the spiny
tenrec Echinops (Thomas 1892) are almost indistinguishable from their
permanent successors in size and form, as are those of its relatives
Hemicentetes (Butler 1941) and Setifer (Butler 1937). Of the known
Insectivora, nine species are described as having little or no
functional milk dentition while 23 have milk teeth which actively
function (Table 3): thus Woodward's (1896b:591) statement that "the
living Insectivora are specialized froms tending toward a
Monophyodont condition" is not a good overall characterizaton of

4
tooth replacement in this order.
The Insectivores have recently been examined as "primitive"
examples for potential insight into the otherwise inaccessible
physiology of early mammals (Schmidt-Nielsen 1980). It would be
encouraging if one could likewise deduce something about the milk
dentition of the early mammals from the extant Insectivora.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission
Table 4. Dental ontogeny, growth, and nutritional development in Insectivora.
Species Reference Functional Age at first Skull size Age at

teeth at solid food at tooth weaning
birth replacement
Tenrecidae:
Echinops telfairi Eisenberg & Gould 14 da. Full grown 28 da.
1970, Leche 1907
Uemicentetes semispinosus Eisenberg & Gould 18 da. Full grown 25 da.
1970, Leche 1897
Microgale dobsoni Eisenberg & Gould <22 da. Full grown
1970, Leche 1907
Setifer setosus Eisenberg & Gould 14 da. Full grown 14-21 da.
1970, Leche 1907
Tenrec ecaudatus Eisenberg & Gould 25 da. Less than >28 da.
1970, Leche 1907 full grown
Chrysochloridae: CJ
Ln
Chrysochloris sp. Leche 1907 Full grown
Eremitalpa sp. Kindahl 1963 Nearly full
grown
Erinaceidae:
Erinaceus europaeus Kindahl 1957a None Less than
full grown
Soricidae:
Cryptotis parva Choate 1970 None Less than
full grown
Talpidae:
Parascalops breweri Eadie 1944 None Before tooth
developmt complete
Scapanus latimanus Ziegler 1972a & b None 35 da.
(Nonfunctional milk antemolars erupt 10-35 da.)
Urotrichus talpoides Usuki 1967 <3-4 mo.
(before tooth
replacement)
36
Erabryological and morphological evidence (e.g. Friant 1965, Sicher
1917) indicates that the monophyodonty of shrews and some moles is
surely a derived condition. Evidence from the remaining Insectivora
would lead one to think that early mammals were born toothless, then
acquired a set of milk teeth which persisted to adulthood and were
replaced by permanenent teeth similar in morphology and perhaps size.
Unfortunately this scenario conflicts with the best
paleontological evidence for earliest mammals, that for the Triassic
triconodont Eozostrodon, which apparently replaced its tiny milk
teeth early in life (Parrington 1971, Crompton and Jenkins 1981).
Even though the Insectivora as an order evolved early in mammalian
history and apparently gave rise to many other groups (see McKenna
1975), the original pattern of tooth replacement may not be well
represented in the modern representatives of the order.
Alternatively, by the time the early Insectivora appeared, the tooth
replacement pattern seen in Eozostrodon may have been modified in the
direction of longer retention of milk teeth. The appears to have been
the condition in the Cretaceous eutherian Kennalestes, in which the
basicranial skull length of a juvenile still replacing teeth is 81
percent of the corresponding length of an adult skull (data from
Kielan-Jaworowska 1975, 1981).
Scandentia
37
Tree shrews (Tupaiidae) are functionally diphyodont, and each
milk tooth is replaced (Table 3). The milk teeth resemble their
successors in form and are so nearly the same size that it is
sometimes difficult to tell them apart. The last milk premolar,
however, is molariform and is morphologically more like the first
molar than it is like its replacement (Butler 1980).
Developmental data come largely from laboratory colonies of
Tupaia glis. In this species, the first milk teeth erupt at 11 days
(Gensch 1962). Parental care is rather perfunctory. The mother
returns to nurse her young every two days, and weaning is
accomplished by about five weeks (Martin 1966). Leche (1897)
concluded from skull examination of T^ glis, T. tana and Dendrogale
melanura that the milk teeth persist until the animals are nearly
full size. Presumably, then, tooth replacement occcurs after weaning,
though in 1\_ glis the permanent teeth are replaced by the time the
young reach physiological sexual maturity at 3 to 4 months (Shigehara
1980).
The developmental sequence of milk teeth in Tupaia javanica was
examined histologically by Kindahl (1957), and eruption sequences of
permanent teeth were described for Tupaia glis by Slaughter et al.
(1974) and Shigehara (1975) and for a composite of species by Lyon
(1913) and Butler (1980). These sequences are complex and
individually variable. In general the molars erupt from front to
back, upper incisors are replaced late, and of the three premolars it
38
is the middle one that is first to develop as a milk tooth yet the
last to be replaced. The middle upper premolar is the "main
puncturing tooth" in the milk and permanent dentition of the
Tupaiinae (Butler 1980), so its early development and late
replacement may be related to this function. In the pen-tailed tree
shrew Ptilocercus it is the posteriormost milk and permanent premolar
that is most modified for puncturing, and it is this premolar that is
the last to be replaced. It may also be of functional significance
that the milk incisors are shed well before the permanent ones erupt,
so that for a period of time the young animals have no functional
anterior dentition (Shigehara 1980). This is a situation familiar in
humans but seldom seen in other mammals. Since a major function of
the incisors is to act as a tooth comb for grooming (Butler 1980), it
would be interesting to know if during this period young tree shrews
are groomed by parents or others.
The possesion of a late-retained, functional milk dentition does
little to resolve the controversy surrounding tupaiid affinities,
since this pattern is seen in both of the proposed related groups,
Insectivora and Primates. On this and other grounds, Butler (1980)
prefers to assign tree shrews to their own separate order.
Dermoptera
39
The literature on tooth replacement in extant Dermoptera
(colugos or gliding lemurs) is based on ten specimens: five of
Cynocephalus variegatus (Owen 1840, Leche 1886, Dependorf 1896) three
of volans (Blainville 1839-64, Dependorf 1896) and two unspecified
embryos (Dependorf 1896). Despite, or perhaps because of, this small
sample size, the general picture of dental development is relatively
unambiguous (Table 3). No known-age material is available, but
’'newborn'1 specimens (Dependorf 1896, Leche 1886) are toothless, while
a fully haired young (Leche 1886) had erupted only the tips of four
milk teeth (incisors and premolars). The milk dentition may erupt
rather suddenly, since a specimen of variegatus with milk
dentition entirely erupted is only 6.3 cm longer (14 percent of adult
length longer) than a toothless neonate (data from Dependorf 1896),
Once the milk teeth are in place they remain functional until the
animal is nearly full size, and the molars (except upper M3) have
already erupted before tooth replacement occurs (Leche 1886). The one
exception to this pattern is the third tooth in the lower jaw (di3 of
Leche 1886, dc of Dependorf 1896) which is shed and replaced quite
early. This tooth is small and peglike in contrast to its elaborately
lobed successor, but the rest of the milk dentition of colugos is
morphologically almost identical to the permanent dentition (Leche
1886). The comb-shaped lower milk incisors even have the same number
of "teeth" as do the permanent ones (Blainville 1839-64).
How weaning is timed in relation to dentition is not known for
colugos. Presumably the adult-like milk dentition allows the young to
40
process the same plant parts eaten by adults, but there are no
relevant observations.
Although the comblike lower incisors of modern Dermoptera are
particularly spectacular, the whole antemolar dentition is so unusual
as to have confounded expression in terms of a conventional dental
formula. Dependorf (1896) cites eight different dental formulas from
the literature before adding his own, and Findley (1967) lists yet
another from Cabrera (1925). Observed tooth replacement patterns
(Table 3) invalidate over half these formulas, since only the three
posteriormost cheek teeth are unreplaced and can therefore properly
be designated "molars.” Dependorf (1896) claimed to find, in newborn
and fetal specimens, vestigial germs of a lower milk and permanent
incisor anterior to the first functional tooth; his formula for the
functional dentition (upper I2P3M3., lower I2CP3M3) is the only one
consistent with that evidence.
The systematic position of colugos has frequently been debated.
Before being elevated to their own order, they had been classed with
both Insectivora and Primates (see Findley 1967). Leche (1886)
considered them incipient bats. The morphological and chronological
relationships of milk to permanent dentition in extant Dermoptera are
more reminiscent of several "primitive" insectivores (e.g. Tenrec,
Erenri talpa) which also have permanent-like milk teeth that are
retained late, than of other groups. This evidence is consistent with
modern taxonomic arrangements giving Dermoptera closest affinities
41
with broadly-defined Insectivora (Findley 1967), but less so with
those treating tupaiids, bats, dermopterans and primates as a branch
separate from either macrosceledids or lipotyphlans (McKenna 1975).
The only relevant fossil evidence, a jaw interpreted as that of a
juvenile dermopteran from the Eocene (Rose 1973, Rose & Simons 1977),
has adult-like premolars and indicates that large, persistent milk
teeth have been in this lineage for a long time.
Chiroptera
Of over 60 bat species that have been described from seven
families, all appear to form a milk dentition (Table 3). For the most
part these teeth complete development and erupt; exceptions have been
found in three genera (Megadermatidae: Lavia; Rhinolophidae:
Rhinolophus and Hipposideros) where they are resorbed before birth.
The most complete bat milk dentition is 22 teeth (several
vespertilionids, some phyllostomids). Below this maximum, milk and
replacement tooth number is almost whimsically variable from genus to
genus (Table 3). Such variation is most evident in the premolars.
Some milk premolars fail to be replaced while some permanent
premolars erupt without predecessors, sometimes with both phenomena
occurring in the same species. Only Nyctalus and Pipistrellus
(Vespertilionidae) have a milk dentition that is matched
tooth-for-tooth with the replacement series (if indeed these teeth
42
are homologous; it is not clear that they are). Evidently whatever
selection pressures have reduced the numbers of teeth in bats have
acted differently upon the milk and permanent dentition.
Five species of Megachiroptera and 17 of Microchiroptera have
all or part of the milk dentition erupted at birth (Table 5).
Hipposideros caffer, which resorbs its milk dentition, is born with
several permanent lower teeth erupted. The several apparent cases of
toothlessness at birth are based on single late-fetal or neonatal
specimens and need confirmation. In known-age cases, eruption of the
permanent dentition occurs rapidly, the last tooth having erupted
well before weaning and usually by the time any solid food is taken
(Table 5). Thus a young bat launching into independent feeding
already has a full set of dental equipment, so that the exact
sequence of acquisition of these teeth may be less functionally
significant than in some other groups. The only quantitative data on
intraspecific variation in timing of tooth eruption are Sowler’s
(1980) for Epomophorus wahlbergi. She reports a wide range of ages
for eruption of each permanent tooth; yet while variations in
sequence do occur, the first to erupt are always upper C and lower
P3, and the last are always lower M2 and upper 12.
Although adult dentitions in Chiroptera are a classic example of
morphological adaptive radiation, bat milk teeth are remarkably
uniform. Much smaller than the permanent teeth, they take the form of
spikelets or recurved hooks (e.g. Weber 1928:143, Figure 88). These
Table 5. Dental and nutritional development in Chiroptera.
Species Reference Functional Age at first Age span of Age at

teeth at solid food tooth weaning
birth replacement
Pteropodidae:
Epomophorus wahlbergi Sowler 1980 Milk teeth 30-90 da. >90 da.
Pteropus samoensis Leche 1878 Milk teeth
Pteropus poliocepHalus Leche 1878 Milk teeth
Rousettus aegyptiacus Friant 1951 Milk teeth
Rousettus Leche 1878 Milk teeth
olexTcaudatus
Emballonuridae:
Peropteryx macrotis Leche 1878 None
Megadermatidae: ■p'
Lavia frons Dorst 1953 None? OJ
Rhinolophidae:
Hipposideros caffer Gaunt 1967 Some perm. teeth
Rhinolophus hipposideros Leche 1877 Some perm. teeth
Mormoopidae:
Pteronotus gyimonotus Nelson 1966 None
Phyllostomiaae:
CaroIlia perspicillata Kleiman & Davis Milk teeth 22-31 da. 56 da.
1979
Desmodus rotundus Greenhall et al None to 35 da. 300 da.
1983, Leche 1878
Glossophaga soricina Leche 1878 Milk teeth
Macrotus waterhousii Nelson 1966 None
Vespertilionidae:
Ametrida centurio Leche 1878 Milk teeth
Antrozous pallidus Orr 1954 Milk teeth 42 da. 28-35 da. >42 da.
Eptesicus serotinus Kleiman 1969 Milk teeth 11-19 da. 24 da.
Table 5 (contd).
Species Reference Functional Age at first Age span of Age at

teeth at solid food tooth weaning
birth replacement
Miniopterus schreibersi Van der Merwe 1981 Milk teeth

Myotis albescens Webster 1981 After tooth
replacement
Myotis leibii Leche 1878 Milk teeth
Myotis lucifugus Fenton 1960, After tooth
Stegeman 1956 replacement
Myotis myotis Kratky 1970 Milk teeth 10-35 da.
Myotis velifer Kunz 1973 Milk teeth 21-28 da. 21-28 da. 42 da.
Nyctalus leisleri Matthews 1950 Milk teeth
Nyctalus noctula Kleiman 1969 Milk teeth 31 da. 18-39 da. 49 da. -O'
to 35 da. 40 da.
■C'
Nycticeius hunneralis Jones 1967 Milk teeth
Pipistrellus Kleiman 1969 Milk teeth 15-23 da. 24 da.
pipistrellus
Plecotus rafinesquii Jones & Suttkus Milk teeth 15-18+ da.
1975
Molossidae:
Tadarida brasiliensis Herreid 1959, Milk teeth 28-42 da.
Short 1961
Tadarida condylura Dorst 1957 Milk teeth
45
teeth resemble nothing in the early Mammalia and apparently
differentiated to their reduced form well after the full achievement
of Chiropteran grade: a specimen of the Eocene bat Archaeonycteris
has milk premolars quite like its permanent ones (Revilliod 1917).
For many years it has been stated that modern bat milk teeth are
specialized to allow the young to grasp its mother (Kunz 1973:26;
Phillips 1971:125; Phillips et al 1977:125; Stegeman 1956:60; Tomes
1880:402). More specifically, the young are said to use the milk
teeth to grip the mother's fur (Dorst 1957:133) or her teat (Birney &
Timm 1975:204; Dorst 1949:47; Fenton 1970:817; Peyer 1968:226;
Spillman 1927:252; van der Merwe 1981:180; Greenhall et al 1983:3),
particularly while she is carrying the young in flight (Allen
1940:168; Friant 1963:98; Grasse 1955:1758; Miller 1896:114 and
1907:21; Orr 1954:230; Reeder 1953:13; Walker 1975:184; Weber
1928:143). It is not easy to separate myth from reality here.
Curiously, not one of these authors actually documents any such
function. Many of them seem to assume that it is a common practice
for mother bats to carry their young in flight, but this itself is a
dubious assumption. Davis and Cockrum's (1964) wing-loading
experiments on Microchiroptera demonstrate that weights comparable to
those of small young could be kept aloft by adults, but they make no
claims about the extent of such behavior in the field. Mother bats of
several species have been observed to carry young short distances
when disturbed by humans at nursery sites, yet obervations under
undisturbed conditions are generally lacking (Jones and Suttkus 1971,
46
Davis and Cockrum 1964, and references therein). Of the authors that
comment on both tooth development and maternal behavior in a species,
none finds evidence of young bats being carried on foraging flights
(Short 1961, Orr 1954, Kunz 1973, Jones and Suttkus 1971 and 1975;
see also Davis et al 1968). Pine (1972:67) speculates that such
behavior may be restricted to bat species that change roost
frequently. Bradbury (1977) notes several species which may carry
young far enough to hide them while the mothers forage, but he also
lists many others whose habit is to leave their young in the roost.
If there are many bat species that under normal conditions do not
carry their young, the idea that milk teeth are peculiarly
specialized "to hold tightly to the mother's nipple when she is in
flight" (Orr 1954:230) loses credence. Further, Kleiman (1979) notes
that milk teeth are more reduced in the phyllostomids than in the
vespertilionids; phyllostomids sometimes carry young but
vespertilionids seldom if ever do (Bradbury 1977).
On the other hand, young bats have been observed to cling to
their roosting mother's teat so tenaciously that attempts to remove
them artificially cause injury (Kleiman 1969, Orr 1954), so these
teeth do at least have a potential for increasing the effectiveness
of attachment. For those bats that do not routinely transport young,
what other advantages might close physical attachment of baby to
mother provide? Some obvious possibilities are quick emergency
removal from nursery-site predators, enhancement of social bonds, and
thermoregulatory aid to the young. If it is the social bond function
47
that is important, a testable hypothesis is that colonial roosting
species should have greater development of hook-like dentition than
solitary-roosting ones; if thermoregulatory, temperate bats ought to
have more developed hooks than tropical ones. There is enough
variation in degree of development of bat milk dentition to make a
comparative assessment of certain of these potential functions, but
more behavioral observation is needed. Whatever proposal is made for
selective value of milk tooth morphology in bats, it must allow for
the apparent supression of this dentition in at least three genera
(see above). Intriguingly, these are the same genera in which
auxiliary "false mammae" (Verschuren 1940) have been described.
"False mammae" are also present in Nycteris and Rhinopoma (K.
Koopman, pers. comm.), but the milk dentition of these taxa has
apparently not been investigated. It seems likely that the functional
significance of bat milk teeth is more precise and possibly more
diverse than is currently appreciated. Until some of these
possibilities are explored, it seems wise not to perpetuate what
amounts to a folklore surrounding these highly modified structures.
Primates
Tooth replacement in Primates has been more thoroughly studied
than in any other group. Swindler (1976) provides a comprehensive
review of most aspects of primate dentition. Deciduous and permanent
48
dental formulas in anthropoid primates are quite straightforward:
apes, monkeys, and humans are fully diphyodont and replace all milk
teeth over a long time span that begins after weaning (Table 3, Table
6). Among other groups (lemurs, indriids, tarsiers, lorises,
aye-aye), tooth homologies and the status of replacement have always
been confusing and have recently been made even more so by proposals
for new homologies (Luckett and Maier 1982, McKenna 1975, Schwartz
1979; see Szalay and Delson 1979 for a critical review). In at least
two groups (Daubentoniidae, Indriidae), some milk teeth are present
which are shed without replacement; but in non-anthropoid taxa there
is no clear morphological distinction among incisors, canines, and
premolars, and therefore little agreement on dental formulas for
either the milk or replacement sets. The formulas in Table 3 for
these groups are thus somewhat arbitrary.
Some comparative observations can be made. In most anthropoids,
the last milk premolars closely resemble molars in morphology. In
toothcombed strepsirhines (lemurs and lorises) both adults and
juveniles have anterior lower teeth modified into a comblike
apparatus, but the toothcomb may be formed from different numbers of
teeth and/or different tooth positions in the two sets (Schwartz
1974, 1978, 1979; Gingerich 1979). The simplest functional
explanation for these two patterns is that they represent
conservation of similar function from juvenile to adult. By contrast,
many primates have enlarged canines as adults, but their milk canines
are generally unmodified relative to neighboring teeth (Ashton 1956,
Table 6. Dental and nutritional development in Primates.
Species Reference First milk teeth Age at first First perm. Age at
erupted (age) solid food teeth (age) weaning
Callithricidae:
Callithrix iacchus Johnston et al Incisors Incisors 12 wk.
1970 (0-1 wk.) (7 mo.)
Saguinus nigricollis Chase & Cooper 1969 Incisors & canine Incisors 10-13 wk,
(birth) (19-20 wk.)
Cebidae:
Saimiri sciureus Long & Cooper 1968 Incisors 1 mo. Incisors 4 mo.
(birth) (9-10 mo.)
Cercopi theeidae:
Macaca mulatta Hurme & Van Wagenen Incisors Incisors 1 yr.
1953, 1961 (2 wk.) (2.4 yr.) >£>
Walker 1975
Papio hamadryas Freednan 1962 Incisors Incisors 7 mo.
(1 wk.) (33 mo.)
Pongidae:
Gorilla gorilla Schaller 1963 Incisors 2.5 mo. 1.5 yr.
(2.5 mo.)
Pan troglodytes Nissen & Riesen Incisors Incisors 2 yr.
1945, 1964 (2.5 mo.) (67 mo.)
Hominidae:
Homo sapiens Kronfeld 1935, Incisors Incisors 3-4 yr.
Pond 1977 (6 mo.) (6.5 yr.)
50
Johanson 1974). Canine size in adults is frequently sexually
dimorphic and apparently important in aggressive displays, and canine
replacement is often retarded until late in the replacement sequence
(Freedman 1962, Hurme and van Wagenen 1961, Sirianni 1978, Spiegel
1952). These facts suggest that canine enlargement in primate
permanent teeth has a primarily social significance and may even have
been selected against in the milk dentition.
Despite the fact that numerous studies have shown primate tooth
form and size to be related to diet and body size in remarkably
precise ways (Hylander 1975, Kay 1975 and 1978, Kay et al 1978, Pirie
1978, Seligsohn and Szalay 1978), and despite the fact that
intraspecific variation in diet (on the basis of sex) is known for
primates (Gauthier-Hion 1980), there have been no investigations to
see if the morphology of primate deciduous teeth is predictably
related to juvenile diets. Age-differentiated field observations
should be possible, since juveniles of most primate species are
identifiable in the field for several years before tooth replacement
is complete.
Carnivora
Nearly all Carnivora are true diphyodonts (Table 3). The only
apparent exceptions are a few mustelids (skunks and badgers; see
chapter 3) and the termite-eating aardwolf Proteles cristatus, which
51
has rudimentary teeth in both permanent and milk sets.
Few Carnivora have teeth at birth (Table 7): the sea otter
Enhydra and spotted hyena Crocuta are the only well-documented
exceptions. Milk teeth erupt o^er a short time span but are typically
retained until after weaning, though a few species begin replacement
before weaning is complete (Table 7). The Viverridae have especially
retarded tooth replacement (Charles-Dominique 1978, Pocock 1933a,
1933b). Although two species of bears are alleged to have their
permanent teth before weaning ends a nearly two years, another
species with similar tooth replacement is reported to be weaned much
earlier. Weaning dates for bears need to be confirmed before these
species can be considered genuinely exceptional in timing of tooth
replacement.
Carnivore milk dentition is morphologically similar to the
permanent set, but with remarkable disparity between milk and
permanent homologues. In most Carnivores, sectorial function in the
permanent dentition is concentrated in the "carnassial apparatus"
composed of the opposing shearing surfaces of the posteriormost upper
premolar and the anteriormost lower molar. Although there are no
molars in the functional juvenile dentition, analogous shearing
surfaces are found in the penultimate upper and last lower milk
premolar, so that juveniles have their own "carnassial apparatus"
developed at different tooth positions. Frequently the juvenile's
last upper milk premolar also resembles the adult's first upper
Table 7. Dental and nutritional development in Carnivora.
Species Reference First milk teeth Age at fir^t Age span of Age at +
erupted (age)* solid food tooth replacemt. weaning
Canidae:
Canis familiaris Kremenak 1967 & 69 I & P (23 da.) 21 da. 117-154 da. 35-42 da.
Canis latrans Bekoff & Jamieson I (14 da.) 28 da. 5 wk.
1975
Canis lupus Mech 1970 I (15 da.) 16-26 wk. 5 wk.
Canis mesomelas Lombaard 1971 1 & C (1 wk.) 17.5-23 wk. 9 wk.
Nyctereutes procyonides Novikov 1962 C (14-16 da.) 18-20 da. 1.5-2 mo.
Vulpes vulpes Linhart 1968 I & C (18-22 da.) 30 da. 15-21 wk. 8 wk.
Ursidae: Ln
Ursus americanus Dittrich 1960, C (6.5 wk) 5.5-9 mo. 5 mo. ro
Rausch 1961
Ursus arctos Couturier 1954 I (2.5 mo.) 3.5-18 mo. nearly 2 yr.
Ursus maritimus Dittrich 1960, I & P (7.5 wk.) 5 mo. 5.5-11 mo. nearly 2 yr.
Pohle 1923
Procyonidae:
Bassariscus astutus Toweill & Toweill I (26 da.) 36-40 da. 75-120 da. 80 da.
1978, Richardson
1942
Nasua narica Kaufmann 1962 I (15 da.) 40 da. 4 mo.
Procyon lotor Montgomery 1964, I (25 da.) 30-40 da. 9-18 wk. 16 wk.
Hamilton 1936,
Stuewer 1943
Mustelidae:
Eira barbara Poglayen-Neuwal1 I (37 da.) 19 da. 115-225 da. 72 da.
1978
Enhydra lutris Loughlin et al 1981, I, C, P Soon after 2.5-5.5 mo. 8-9 mo.
Schneider 1973, (birth) birth
Kenyon 1969
Gulo gulo Krott 1959, Shilo & I & C (5-6 wk.) 12 wk. 12-21 wk. 10 wk.
Tamarovskaya 1981
Table 7 (contd).
Species Reference First milk teeth Age at first Age span of Age at
erupted (age) solid food tooth replacemt. weaning
Ictonyx striatus Bail 1978, Rowe- C (32 da.) 6 wk. ca. 13.5 wk. 8 wk.
Rowe 1978
Martes aroericana Brassard & Bernard I (17 da.) 5 wk. 9-16 wk. 6 wk.
1939, Markley &
Bassett 1942
Martes foina Habermehl & Rottcher C & P (2-3 wk.) 6 wk. 7-16 wk. 6 wk.
1967, Schmidt 1943,
Camivenc et al 1981
Martes martes Corbet & Southern I (40 da.) >36 da. 2-4 mo. 1.5 mo.
1977, Ryabov 1962, <_n
u>
Novikov 1962
Martes pennanti Powell 1982 C (7 da.) 8-10 wk. 4 mo.
Meles roeles Cox 1962, Gucwinska I (17-30 da.) 75-80 da. ca. 4 mo. 3 mo.
& Gucwinski 1968, (prob. non
Howard & Bradbury functional)
1979, Leche 1910
Mephitis mephitis Verts 1967, None 38 da. 36-49 da. 2-4 mo.?
Stegeman 1937
Mustela erminea East & Lockie 1965, I (5-6 da.) 3-5 wk. 5-8.5 wk. 5-7 wk.
Muller 1970, limanov
et al 1970
Mustela frenata Hamilton 1933, C & P (21 da.) 21 da. 55-75 da. 36 da.
Sanderson 1949
Mustela nivalis Heidt 1970, East & C (2-3 wk.) 22-23 da. 4.5-6.5 wk. 6 wk.
Lockie 1964, Corbet i
Southern 1977
Mustela putorius Goethe 1940, Hahn & C & P (2-3 wk.) 23 da. 7.5-12 wk. b wk.
Wester 1969
Table 7 (contd).
Mustela vison Aulerich & Swindler I (19 da.) 2 wk. 45-68 da. 42 da.
1968, Travis & Schaible
1961, Kainer 1954
Poecilogale albinucha Rowe-Rowe 1978 C (35 da.) 35 da. ca. 13 wk. 11 wk.
Taxidea taxus Long 1974, Long I,C, P (2 wk.) 1-4 mo. 3 mo.
& Killingley 1983,
Wright 1966
Viverridae
Arctitis binturong Aquilina & Beyer I & C (34 da.) 36 da. >172 da. cn
1979
Genetta maculata Rowe-Rowe 1971 C (3-4 wk.) 5 wk. ca. 51 wk.
Herpestidae:
Suricata suricatta Ducker 1962 I (5 da.) 22 da. After weaning 58 da.
Hyaenidae:
Crocuta crocuta Kruuk 1972, I & C (birth) 5 mo. ca. 9.5 mo. 19 mo.
Poumelle 1965
Felidae:
Acinonyx jubatus Florio & Spinelli (20 da.) 1 mo. 5 mo.
1968
Felis caracal Grobler 1982 (all by 4-6 wk.) 6 wk. ca. 25 wk. 10-25 wk.
Felis concolor Currier 1983, Volf I (10-20 da.) 5.5-8+ mo. 1-2 mo.
1972, Schneider 1959
Felis nigripes Armstrong 1975, I & C (18 da.) 30-35 da. ca. 146 da.
Leyhausen & Tonkin*
1966
Felis silvestris Haltenorth 1957, I (11 da.) 1-1.5 mo. 4 mo.
Meyer-Holzapfel 1968,
Tonkin & Kohler 1981
Felis wiedii Petersen & Petersen C? (20 da.) 52-57 da. ca. 132 da.
1978
Table 7 (contd).
Crowe 1975 11 da. 130-240 da. 60 da.

David 1962, de I (12-21 da.) 5 wk. 8.5-17 mo. 7-8 mo.
Carvalho 1968, Schaller
1972, Smuts et al 1978
Panthera onca Schneider 1959 I (<17 da.) 70 da. 22 wk.
Panthera pardus de Carvalho 1968, I (20 da.) 42 da. ca. 280 da. 120-125 da.
Schneider 1959
Panthera uncia Freuh 1968, Marma & I (17 da.) 30 da. 8-10 wk.
Yunchis 1968
____________________________________________________________________________________ cn
* ^
"Tooth designations: I = incisor, C = canine, P = premolar.
Age of first solid food and weaning: some supplementary data are from Ewer 1973.
56
molar.
The positional shift from milk to permanent carnassial has been
thoroughly described for a number of species (Broom 1949, Gaunt 1969,
Hall 1951, Leche 1915, Mivart 1882, 1885, Pococlc 1916, Tims 1896,)
and appears to occur in every species where a specialized carnassial
can be identified. A natural corollary of milk and permanent
carnassial being positionally different is that carnassial function
can be maintained even during tooth replacement, if the permanent
carnassials erupt before the milk carnassials are shed. Just such an
overlap has been documented in several cases (e.g. Aulerich and
Swindler 1968, Berkovitz 1973, Long 1974). Ewer (1973) proposes that
this eruption pattern may be general for carnassial-dependent
carnivores, and Slaughter et al (1974) confirm it for a number of
species throughout the order. Butler (1946) even suggests that the
lack of a positional shift from milk to permanent carnassials may
have been fatal to the extinct carnivorous order Creodonta.
In addition to the highly specialized carnassials, Carnivora
also possess greatly developed canine teeth, large even in species
with no predatory habits. If canines are functionally vital it would
seem important for them to be maintained through the period of tooth
change as is the carnassial, but the milk canines occupy the same
position in the jaw as do the permanent ones. Unlike other groups
with developed canines, however, Carnivore milk canines do not drop
out before the permanent ones erupt; instead, the permanent canine
57
typically erupts just anterior to the milk canine and closely
appressed to it, so that until the permanent canine is approximately
as long as the milk one, the two coexist in the jaw (Couturier 1954,
Ewer 1973, Smuts et al 1978). Thus there is continuous function of
the two functionally specialized portions of the Carnivoran dentition
throughout the period of change from milk to permanent teeth.
In contrast to the terrestrial carnivores, the pinnipeds
(Otariidae, Odobenidae, Phocidae) produce only a rudimentary milk
dentition which is resorbed or shed in utero (Kubota and Matsumoto
1963) or shortly after birth (Briggs 1974). Usually the permanent
teeth have alreaady begun to erupt in newborns. The pinniped
carnivores are therefore functionally monophyodont (Table 3).
Cetacea
The mysticete or baleen whales (Balaenopteridae, Balaenidae,
Eschrichtiidae) produce no erupted teeth, ingestive function being
performed instead by the baleen plates which originate from oral
epithelial membrane. It has been known for many years, however, that
tooth buds are formed in the embryo. Eschricht (1845) described such
buds in the upper jaws of the right whale Balaena mysticetus. He also
(1845) reported 51 upper and 42 lower tooth buds on each side in the
fin whale Balaenoptera physalus and 44 upper and 40 lower buds per
side in the Sei whale Balaenoptera borealis. Pouchet and Chabry
58
(1882) found a dental lamina in upper and lower jaws of the blue
whale Balaenoptera musculus. These reports were based on single
fetuses. The most extensive data on dentition in baleen whales are
from Karlsen (1962). Examining 22 embryos of Balaenoptera physalus,
he showed that the number of tooth buds present is subject to
considerable variation. A dental lamina is present even in tiny
embryos (2 cm crown-tail length) of this species. Dentin is formed in
the little teeth of both jaws and an enamel organ is present, though
no enamel is actually laid on. Resorption of the vestigial teeth
begins at the front of the jaw, and all tooth germs had disappeared
in a fetus 330 cm long (Karlsen 1962).
Since only a single set of teeth is formed, there is little
evidence to determine its homology with the milk or permanent set of
other eutherians. From the high degree of intimacy that the tooth
buds show to the dental lamina, Karlsen (1962) concluded that the
vestigial teeth of baleen whales are milk teeth, while the permanent
set has been altogether lost.
All species in the other major division of Cetacea, the
odontocete or toothed whales, produce functional teeth; but the
number of such teeth is extremely varied, from a single ever-growing
tusk on the upper left side in the male narwhal Monodon monoceros
(Ness 1960) to a total of 260 homodont teeth in some delphinids
(Walker 1975). Eschricht (1855) found 12 upper and 11 lower teeth per
side in a fetus of the bottlenose whale Hyperoodon ampullus, a
59
species that at maturity has no upper and only one or two lower teeth
functional (Walker 1975). Since only one of these, the seventh upper,
had erupted, Eschricht (1855) proposed that the erupted tooth
represented a remnant milk dentition while the rest corresponded to a
permanent set. This appears to be the only evidence for presence of
two tooth sets in whales. Later, Leche (1892) described tooth buds in
the common porpoise Phocoena phocoena. Apparently arguing from the
positional relationship of these buds to the dental lamina, he
concluded that the persistent teeth in odontocetes are the milk
dentition.
Little is known about timing of tooth development in
odontocetes. The bottle-nosed dolphin Tursiops truncatus is toothless
at birth, erupts its functional teeth at six weeks, ingests solids at
six months and is weaned at 16 to 18 months (Slijper 1978, Walker
1975).
Baleen and toothed whales have had a very long separate history,
and common ancestors are unknown (Rice 1967). A third group of
whales, the extinct Archaeocetes, were relatively heterodont and may
have had two sets of teeth (Grasse 1955:359). The eutherian ancestors
of whales presumably had two functional tooth sets, and at some point
in the mysticete lineage there must have been intermediate forms with
only one tooth set, since vestiges of it remain in modern embryos. It
would be interesting to know if it is indeed the permanent dentition
that has been totally suppressed in both mysticete and odontocete
60
whales, or if instead in either it was the milk dentition that was
reduced first, as is the case with the other major group of marine
mammals, the pinniped carnivores.
Sirenia
Dental morphology and tooth replacement is quite different in
the two families of Sirenia. In the Dugongidae (Dugong dugon), the
cheek teeth are homodont, cylindrical, enamelless columns. As early
as 1825, Cuvier observed that lower incisors occur only in the very
young and are probably nonfunctional, and that the number of
functional cheek teeth actually decreases with age from five to two
per jaw quadrant. Mitchell (1973) substantiated Cuvier’s observations
and demonstrated how the reduction in cheek tooth number occurs. Very
young skulls have five cheek teeth erupted. A sixth is later added at
the back of the jaw, but meanwhile the first has been shed without
replacement. As the dugong ages, the second, third, and fourth cheek
teeth are also shed without being replaced. Mitchell (1973) also
observed a single upper incisor on each side that is resorbed rather
than erupting, and a later-developing upper incisor that calcifies
but may or may not erupt. Dugongs begin to feed on aquatic vegetation
very shortly after birth (Kenchington 1972), but their teeth may be
relatively unimportant in this process since the palate and part of
the lower jaw are covered with horny plates used in chewing or
61
Ingestion. The recently extinct Steller’s sea cow Hydroamalis
apparently had no functional dentition at all (Walker 1975).
Manatees, family Trichechidae, also have rough oral plates. Two
incisors develop beneath the plates on each side of upper and lower
jaws, but these teeth never become functional (Howes and Harrison
1892, Walker 1975). Cheek teeth, in contrast to those of dugongs,
appear to be highly modified for long-lasting function (Domning
1983). Enameled crests extend across the tooth crowns to form a
washboard-like surface. Although the teeth are low-crowned and
eventually wear down anteriorly, each worn tooth is shed and a new
tooth erupts at the back of the jaw. The alveolus forms a continuous
channel in the jaw, so that as each tooth is shed the row shifts
forward and function is preserved. Trichechus senegalensis produces
in its lifetime up to 20 cheek teeth in each quadrant, with no
cessation of production evident (Thomas 1897), and T. inunguis may
produce 27 or more (Domning 1983).
The homologies of sirenian teeth with those of other eutherians
have never been very satisfactorily resolved. Kukenthal (1896)
described and figured tooth buds in an embryo of Trichechus manatus,
comprising three upper and three lower incisors, a lower canine and
three lower premolars, as well as three upper and lower true molars.
From their relationship to the dental lamina, Kukenthal designated
all these as milk teeth. He had no older specimens, however, so the
correspondence of these buds to the functional teeth is unclear. The
62
small nonfunctional teeth that are present in young animals are
sometimes designated "milk" or "deciduous" teeth (e.g. Abel 1906,
Heuvelmans 1941, Jones and Johnson 1967); but again there is no known
correspondence between these and a permanent dentition. Whatever the
embryonic condition, however, in functional terms, even where tooth
replacement occurs in this order, it does not follow the diphyodont
pattern.
Proboscidea
There is some debate over whether or not elephants produce milk
tusks; although Dittrich (1974) reported them in Indian elephants
(Elephas maximus, Lang (1965) did not observe any milk tusks in six
captive African elephants (Loxodonta africana). Permanent tusks in
this species erupt at about 17 months in males and at two to two and
a half years in females (Lang 1980, Laws 1966), and grow throughout
life (Ness 1960).
Modern elephants are apparently born with a single cheek tooth
in each jaw quadrant erupted; at least this has been reported in
Loxodonta (Morrison-Scott 1947). As the surface of each molariform
tooth wears down, its roots deteriorate also, and eventually the
tooth is shed. The tooth that has been behind this one moves forward
as it does in manatees, thus making space at the back of the jaw for
a new tooth to erupt. Young Elephas are weaned at about three years
63
(Peters et al 1972), presumably after the tusks and first two cheek
teeth are already in use, if timing of tooth succession is like that
in Loxodonta (Lang 1980). Although the jaw grows considerably over
the years, there are never more than two functional cheek teeth in
use at once in each quadrant, because each succeeding molariform
tooth is larger than the one before it. In Loxodonta there is a
maximum of six (Lang 1980) or seven (Laws 1966) such cheek teeth over
a lifespan. Only the last three are conventionally designated molars
(Peyer 1968). The final tooth erupts at about 23 years (Krumrey and
Buss 1968), and the last two molars may be in use on the order of
thirty years each (data from Lang 1980, Laws 1966, Krumrey and Buss
1968). Tooth replacement in' Elephas is similar (Peyer 1968). Thus,
although the living Proboscidea do have a functional tooth
replacement resembling that in manatees and macropodid marsupials,
they cannot be considered truly diphyodont.
Perissodactyla
All modern perissodactyls (horses, rhinos, tapirs) produce a
functional milk dentition and replace it over a period of several
years (Table 3, Table 8). As young adults all have a row of four
premolars in at least the upper jaw. The replacement status of the
first of these teeth is of interest, since in other mammal groups
that have four premolars, the first is almost never replaced (Ziegler
Table 8. Dental and nutritional development in Perissodactyla.
erupted(age) solid food tooth replacemt. weaning
Gquidae:
Equus burchelli Erz 1964, King 1965, Incisors & Few days 2.5-4 yr. 7 mo.
Klingel 1965 premolars
(1 wk)
Equus caballus Nickel et al 1967 Incisors & 2.5-5 yr.
premolars
(birth)
Equus grevyi King 1965 Incisors 2.5+ yr. 9 mo.
(<1 wk.)
Equus zebra Penzhorn 1982 Incisors & 2-3 yr.
premolars ON
4N
(<3 mo.)
Rhinocerotidae:
Diceros bicornis Anderson 1966, Premolars (Ceratotherium: 1.5-2 yr. (Rhinoceros:
Walker 1975, (birth) 4 mo.") TyrTJ----
Wallach 1969
65
1971a). In horses (Equidae), the first premolar remains unreplaced;
it erupts later than other milk premolars but earlier than the
replacement premolars behind it, so is often considered a milk tooth
without successor (Ziegler 1971a). At any rate in equids this tooth
is quite small, variable in form, and often lost early (Ellenberger
and Baum 1932, Erz 1964, Penzhorn 1982).
In rhinoceroses (Rhinocerotidae), the first premolar apparently
is replaced, contra Ziegler (1971a). Anderson (1966) described its
change in Diceros bicornis and Owen (1840) documented it in
Dicerorhinus sumatrensis. Anderson's (1966) size measurements for the
putative milk and permanent first premolar show no difference between
the two, however, so confirmation of the change would be useful. In
the tapirs (Tapiridae), replacement of the upper first premolar has
been shown for Tapirus indicus (Parker 1882), T. pinchaque
(Blainville 1839-1864, Ziegler 1971a), and an unspecified American
tapir (Cuvier 1822).
There is little specific information available about dental
ontogeny in relation to feeding development in perissodactyls. Equids
have functional teeth at birth or soon thereafter and begin to eat
vegetation within a few days; weaning occurs months before any teeth
are replaced (Table 8). A Malayan tapir 18 months old had not yet
begun to replace its milk teeth (Parker 1882), but weaning age is not
known for this species. The black rhino Diceros bicornis has
premolars erupted at birth; its relative the white rhino is consuming
66
a substantial amount of solid vegetation at four months (Table 8). If
Diceros nurses for two years as is reported for Rhinoceros, it
replaces its teeth quite near the time of weaning, a situation that
would be unusual among mammals (see later).
Phylogenetic trends among fossil Equidae toward increasing
hypsodonty of the cheek teeth especially in relation to increased
grazing habit are well known (Janis 1976, Simpson 1951). Butler
(1952) examined the relationships among milk premolars, true molars,
and replacement premolars in this regard. He found that milk
premolars of one species tend to resemble those of other species, but
that replacement premolars differ considerably from species to
species. Milk premolars moreover tend to be more molariform than are
permanent premolars, and in several parallel lineages they became
increasingly more molariform with time. Butler pointed to this trend
as evidence that these teeth are part of a continuous primary series
with the true molars while the replacement teeth are of a separate
secondary set. In functional terms, however, this progressive
"molarisation of the milk-molars" is what might be expected if the
milk premolars perform the same function in early years that true
molars later assume. Butler's interpretation assumes that morphology
of the milk teeth is heavily influenced by developmental canalization
(specifically in this case by influence of the molar developmental
field; Butler 1978a), while the functional interpretation assumes
that milk tooth morphology is responsive to the selective forces it
encounters. Unfortunately, not enough is yet known about what
67
determines tooth morphology to choose between these alternatives
(Butler 1982). It is interesting to note that in Merychippus, while
the permanent cheek teeth were hypsodont and had the cementum found
in modern forms, the milk cheek teeth were short-crowned and lacked
cementum, thus were less modified from the ancestral condition (Lull
1907).
Hyracoidea
Historically, hyrax tooth replacement was one of the first
explicitly described and illustrated (Cuvier 1812). These animals
unquestionably have milk incisors and at least three milk premolars
per jaw quadrant that erupt, experience wear, and are replaced by
permanent successors with similar morphology, as shown in Table 3
(Adloff 1903; Blainville 1839-1864; Brauer 1913; Cuvier 1812, 1822-;
Fairall 1980; Lataste 1886a, 1886b; Roche 1978; Thomas 1892b;
Woodward 1892). In addition, vestiges of additional milk incisors and
possibly a lower milk canine have been observed to form and calcify
but not to become functional (Adloff 1903, Brauer 1913, Thomas 1892b,
Woodward 1892).
What has generated controversy in the hyrax dentition is the
status of three remaining teeth, namely the anteriormost two upper
and single lower cheek teeth. Beca'use they are located directly in
front of the premolar row, these teeth are usually recognized as a
68
canine and first premolar in the upper jaw and a first premolar only
in the lower. Lataste (1886a) claimed that all three undergo
replacement; he described the morphology of milk and permanent
versions of all of them, though admitting that the permanent canine
may be lost in old individuals. Nobody else seems to have seen this
permanent canine, however, and the canine is usually described as a
milk tooth which is shed early without replacement (Brauer 1913,
Fairall 1980, Roche 1978, Thomas 1892b). On the other hand, the upper
and lower premolars have been repeatedly described and even figured
as being replaced (e.g. Blainville 1839-64; Brauer 1913). If they
are, this would be an unusual situation, since when four premolars
are present in modern mammals the first virtually always erupts late
and remains unreplaced (Ziegler 1971a). Ziegler (1971a) suggested
that perhaps the replacement of the upper first premolar is an
illusion, and that what actually occurs is the replacement of the
milk canine by a preraolariform permanent canine which is large enough
to displace both the milk canine and the first premolar behind it.
This leaves the replacement of the lower first premolar still
explained, however, and it seems most parsimonious to assume that the
upper and lower first premolars of the hyrax really are replaced. It
would be interesting to know whether the supposed canine replacement
described by Lataste (1886a) is real as well, or whether perhaps he
was misled by morphological variation among individuals.
Hyraxes are precocious at birth; Procavia capensis at least has
incisors and three premolars per quadrant already erupted (Roche
69
1978). Young begin to eat solid vegetation within three or four days,
and weaning occurs at about three months (Mendelssohn 1965). Growth
is remarkably slow for such small mammals, adult weight being
approached only after about six years (Fairall 1980). Thus even
though skulls showing tooth replacement are still quite small (Roche
1978), the milk teeth are in use a long time after weaning. In
Procavia capensis incisors are replaced at nine months, premolars at
about a year (Fairall 1980, Roche 1978). Such a persistent milk
dentition might well be associated with differences in diet between
juveniles and adults, particularly perhaps in a species such as
Procavia capensis, a habitat specialist with elaborate social
organization (Walker 1975). Recent scanning electron microscope
analyses have shown it possible to deduce differences in diets among
hyrax species by microwear on teeth (Walker et al 1978). It would be
interesting to extend this technique to compare diets of juveniles
with those of adults within species.
Tubulidentata
The one living species assigned to this order, the aardvark
Orycteropus afer, has approximately five evergrowing cheek teeth per
quadrant as an adult (Peyer 1968) and lacks functional permanent
incisors. Although the aardvark was initially thought to be totally
monophyodont, Thomas (1890) reported a nonfunctional milk dentition
70
consisting of up to eight small calcified pegs in the upper cheek
tooth row and four in the lower. Broom (1906-09) described a cleared
head of a newborn aardvark as containing ten such milk teeth in each
quadrant. He interpreted these as three incisors, a canine, and six
premolars. From histological study, Heuser (1913) expanded this
number to 12 upper and 14 lower cheek tooth buds on each side in the
milk dentition. A near-full-term fetus examined by Anthony (1934) had
no teeth erupted, but contained 13 teeth on a side in both upper and
lower jaws. Anthony designated the last three of these as true molars
that would erupt in the adult. The rest he felt were milk teeth that
would be shed very early, only the last two being replaced by
permanent premolars. There appears to be no evidence that the milk
teeth of the aardvark ever erupt, and from Broom's (1906-09)
description of partially resorbed vestiges it seems likely that these
small teeth degenerate rather than being exfoliated.
The permanent tooth structure of the aardvark is unique. Each
cheek tooth is composed of hundreds of hexagonal prisms of dentine
arranged around tubules of pulp. There is no enamel; instead, the
teeth are coated with a layer of cementum (Grasse 1955:700). Since
aardvarks subsist almost exclusively on ants and termites (Melton
1976), the bizarre tooth architecture is presumably an adaptation for
this specialized diet. The earliest fossil orycteropodids had such
teeth, and no intermediate forms are known (Dawson 1967). It would be
interesting to know whether the milk teeth resemble the permanent
ones in structure or are instead more conventional. As figured and
71
described by Heuser (1913), the buds for these teeth contain an
enamel organ and overall are quite similar to those of other mammals.
However, Heuser also claimed to find enamel organs on the budding
permanent teeth. Grasse (1955:700) stated that the milk teeth are not
prismatic like the permanent teeth, but his source for this statement
is unclear. Thomas (1890:247), on the other hand, described the
vestigial milk teeth as showing the beginnings of "the remarkable
histological structure characteristic of the adult teeth." Since the
phylogenetic position of Orycteropus has always been somewhat
puzzling, it seems odd that the nature of the aardvark milk dentition
has not been reexamined in nearly fifty years.
Artiodactyla
Apparently, all artiodactyls are functionally diphyodont (Table
3). Ruminant artiodactyls (deer, bovids, camels, giraffes) routinely
produce a functional milk dentition morphologically analogous to the
permanent one, and (except in the dromedary) all milk teeth are
replaced. Throughout the order milk teeth (incisors and premolars)
are present at birth or within days thereafter, and they are used
until quite worn before being replaced, often after five years or
more (Table 9). Milk premolars in many artiodactyls resemble molars
morphologically, so that the grinding surface undergoes a positional
shift from last premolars to molars during development.
Table 9. Dental and nutritional development in Artiodacryla.
Species Reference First milk tegth Age at first Age span of Age at
erupted (age)" solid food tooth replacemt. waning
Suidae:
R iacoch oeru s a e th io p ic u s Quid et al 1%5 C or more (birth) 7-17 mo. 11 mo.?
Sus s c r o fa Matschke 1967 I & C (birth) 7-19 mo. 4 mo.
T a y a s s u id a e :
l'ayassu t a ja c u Kirkpatrick & I & C (birth) 29-83 wk. b -8 wk.
Sovls 1%2
Camelidae:
Caroelus dromedarius Droandi 1936 I (10 da.) 4.5-8 yr. 3 yr.
Cervidae:
Alces aIces Ifeterson 1955 I 1 C (birth) 10-lb mo. 2-12 mo.
Cervus~dama Chaplin & Mute 1959 I, C, P (birth) 12-24 mo. 3-4 mo.
Cervus elaphus Murie 1951, bowe I & C (birth) 18-39 mo. 9 mo. NS
1967
Cervus nippon Kuwano 1929 I, C, P (birth) 20-32 mo. 9 mo.
Odocoileus hanionus Jones 1953 I, C, P (birth) 11-22 mo.
Odocoileus vTi r a rmn ia n u s Brokx 1972 I, C (birth) 8-20 mo.
Rangifer~taraandus Bergerud 1970 I, C, P (birth) 1-2 wk. 9-27.5 mo. 2 mo.
Giraffidae:
Giraffa Camelopardalis Dagg & Foster 1976 I, C, P (birth) 1-12 wk. 3-6+ yr. 8-12 mo.
Bovidae:
Antidorcas narsupialis Rautenbach 1971 I, C, P (birth) <1 mo. 12-22 mo.
Antilocapra americana Dow 1952, Hoover et I, last P (birth) 14-35 mo. 4 mo.
al 1959
Bison bison Fuller 1959 I, C, P (few days) 2-4 yr. 7 mo.
Bos taurus Ellenberger & Baun I, C, P (birth) 2-5 yr.
1932
Capra hircus Koch 1963 I, P (birth) 15-36 mo.
Table 9 (contd).
Connochaetes taurinus Talbot & Talbot P (birth) 10 da. 26-40 mo. 12-16 mo.
1963
Hemitragus jemlahicus Caughley 1965 I, P (birth) 17-42.5 mo.
Oreamnos americanus Brandborg 1955 I (birth) Few days 1-4 yr. 2.5-3.5 mo.
Ovibos moschatus Henrichsen & Grue I, P (birth) 3-5 yr. 4-5 mo.
1980
Ovis aries Nickel et al 1967 I, C, P (birth) 12-48 mo.
Ovis canadensis Cowan 1940, Deming I (2 da.) 2 wk. 24-48 mo. 4-6 mo.
1952, Geist 1971
Ovis dalli Cowan 1940, Geist I, C, P (birth) 13-48 mo. 4-6 mo.
1971, Hemming 1969
Saiga tatarica Bannikov et al 1961 I, C, P (birth) 3-4 da. 13-24 mo. 2-2.5 mo.
Syncerus caffer Grimsdell 1973 I, P (birth) 3.5-4.5 yr.
^Tooth designations: I = incisor, C = canine, P = premolar.

Weaning age: some supplementary data are from V. Hayssen (unpub.)
74
Many artiodactyls (Bovidae, Cervidae) lack upper incisors
throughout life. In sheep, there is an early embryological
invagination of oral ectoderm in the upper jaw, similar to the one in
the mandible that later becomes the dental lamina; but in the upper
jaw this deteriorates at nine weeks fetal age (Hatt 1967). Whether
this is a remnant of a milk or a permanent dentition is unclear.
Pigs and peccaries (Suidae, Tayassuidae) produce complete milk
tooth sets, but at least in pigs the milk incisors and canines differ
substantially from their permanent successors, being tiny sharp
spikes. Observing that littermates often inflict slash wounds on each
other during struggles at the mother’s teats, Fraser (1975) proposed
that the canines of piglets are specially adapted implements for
effecting swift establishment of "teat order." He compared wounding
rates, teat consistency, and weight gain in 37 untreated litters with
a carefully matched set in which the "eye teeth" had been clipped.
For weight gain the mean and variance both were higher in "unclipped"
litters. This suggests that teat dominance, as determined by eye
tooth fighting, may be important in piglet growth. Fraser’s is the
only controlled manipulative study of functional significance of milk
dentition; it points out the theoretical.feasibility of conducting
such studies but also the difficulty of using this approach with any
but domestic, high-production species, since even with carefully
paired, large sample sizes, comparative results were subtle.
75
Because artiodactyls retain their milk dentition for so long,
there is, as in the case of primates, hyraxes, and perissodactyls,
considerable potential for specialization to juvenile diet. Natural
diets for some ungulates do differ from juvenile to adult (e.g.
saiga, see Bannikov et al 1961) but no work has yet been done on
dietary adaptations of artiodactyl milk teeth.
Pholidota
The pangolins are specialist ant- and termite-eaters and are
functionally toothless throughout life. Rose (1892b) reported a
dental lamina in the upper jaw and a swollen tooth bud in the lower
jaw of an embryo of Manis. In M. tricuspis Leche (1892, 1895) found
no traces of tooth development, however, and Rose's findings remain
unconfirmed.
Rodentia
With the exception of one meticulous monograph (Cederblom 1900),
little has been written on tooth replacement in the largest order of
mammals. This is probably because the majority of rodents are
muroids, all of which apparently are functionally monophyodont (Gaunt
1966). Vestigial incisors have repeatedly been observed in laboratory
Table 10. Dental and nutritional development in Rodentia.
Species Reference First teeth Age at first Age at premolar Age at

erupted (age) solid food* replacement weaning*
Sciuridae:
Sciurus carolinensis Uhlig 1955 Perm, incisor 31 wk. 7-10 wk.
(3-4 wk.)
Tamias striatus Allen 1938, Yerger Perm, incisor 10 wk. 2 mo.
1955 (1 wk.)
Tamias minimus Forbes 1966 Perm, incisor 37 da. 90-100 da.
(27 da.)
Castoridae:
Castor canadensis Cook & Maunton 1954, Perm, incisor 11 mo. 10-12 wk.
Van Nostrand & (birth)
Stephenson 1964
Erethizontidae:
Erethizon dorsatum Earle & Kramm 1980, Perm, incisor & 9 da. <2 yr. 3 mo.
Sutton 1972 milk premolar
(birth)
Caviidae:
Cavia porcellus Harman & Smith 1936 Perm, incisor & 1 hr. Before birth 21 da.
perm, premolar
(birth)
"Solid food, weaning age: some supplementary data are from Bee et al. 1981 and Hazard 1982.
77
rats and mice and are usually interpreted as deciduous teeth
(Fitzgerald 1973, Moss-Salentijn 1975 and 1978, Peters and Strassburg
1969, Woodward 1894), but these never erupt nor fully calcify. The
functional incisors, thought to be of the permanent set, are
evergrowing and never replaced in any rodent, and canines are totally
absent even as rudiments throughout the order. The last three cheek
teeth are virtually always unreplaced and are considered molars; so
only those rodents that have more than three cheek teeth have
potential tooth replacement. Still, there are a number of at least
partially diphyodont rodents (Table 3), since cheek teeth do undergo
replacement in several families. Tooth change never involves more
than two teeth per jaw quadrant. Surprisingly, there are common
rodents which have more than three cheek teeth and therefore "should"
have replacing premolars, but whose tooth replacement has never been
documented (e.g. pedetids, geomyids and heteromyids). Some of these
may shed the milk premolars very early, but in at least one case
(Thryonomys: Wood 1962, Woods 1976) the replacement teeth appear to
have been suppressed.
Few rodents are reported to have teeth at birth (Table 10), but
many erupt incisors within a few days thereafter. On first eruption
in a suckling young, the incisors may have tips modified or splayed,
apparently allowing the entry of the mother's teat between the teeth
(Hamilton 1953, Lawrence 1941). Data on replacement of the milk
premolar in relation to feeding development appears in Table 10: with
one case of replacement before weaning, three considerably after, and
78
one at about the time of weaning, there is clearly no general rodent
pattern evident.
Rodent milk premolars may be smaller and simpler than their
replacements (Cavia, Berkovitz 1972; Myoxus, Sciurus, Cederblom 1900)
or complex and of the same size (Castor, Erethizon; Cederblom 1900).
Those that are as large as the permanent premolars tend to be
retained late and show wear (Earle and Kramm 1980, Ray 1964), so they
are clearly functional. Function for the small ones is undocumented.
One puzzling feature of rodent premolar replacement is that in
several groups (Caviidae, Harman and Smith 1936; Dasyproctidae,
Castoridae, Aplodontidae, Cederblom 1900) the milk premolar forms
typical roots while the replacement one is "rootless," i.e. never
forms root apices and therefore remains evergrowing throughout life.
Since there is almost surely some impairment of function during the
period of tooth change, it is odd that the production of milk teeth
has not been suppressed with the acquisition of a perpetually active
tooth growing apparatus. Perhaps the guinea pig, which sheds a rooted
milk tooth in utero before forming an evergrowing premolar (Berkovitz
1972, Tims 1901), represents such an intermediate stage, but this
seems an inelegant solution for representatives of an order that has
managed to suppress completely the formation of many teeth.
Lagomorpha
79
Most of what is known about tooth replacment in lagomorphs
(Table 3) is based on the domestic rabbit Oryctolagus cuniculus. Like
rodents, the rabbit has a single large evergrowing incisor on each
side of upper and lower jaws. These do not undergo replacement. Just
behind the upper one is a second, peglike incisor. The original tooth
in this position is shed and replaced at 35 days (Moss-Salentijn
1978, Owen 1840, Woodward 1894). In addition, a rudimentary incisor
forms in the upper jaw and the lower, but it is apparently exfoliated
in the uterus before birth (Hirschfeld et al 1973, Moss-Salentijn
1978, Ooe 1980, Peters and Strassburg 1969, Woodward 1894).
Moss-Salentijn (1978) reported another maxillary incisor bud as well.
There is no consensus on homology of the rabbit incisors.
Hirschfeld et al (1973) considered the functional adult upper
incisors to be the permanent II and 12 of the ancestral placental
formula; Moss-Salentijn (1978) and Ooe (1980) called them a milk di2
and a permanent 13. Even more confusing, the single functional lower
incisor has been considered to be permanent II (Hirschfeld et al
1973), milk di3 (Moss-Salentijn 1978), and milk di2 (Ooe 1980). The
formula in Table 3 is based on Hirschfeld et al's interpretation only
because it best expresses functional relationships.
Oryctolagus1 milk premolars erupt before birth and are replaced
at 30 to 35 days (Horowitz et al 1973). Since young domestic rabbits
suckle for about 3 weeks (V. Hayssen, unpub.), the teeth are not
replaced until after weaning. In the eastern cottontail Sylvilagus
80
floridanus, apparently the only other lagomorph whose tooth
development has been described, the tips of the large functional
incisor are erupted in upper and lower jaws at birth (Dice and Dice
1941). The same teeth are replaced in this species as in
Oryctolagus; the deciduous premolars are considerably worn by the
time they are shed. Using other developmental criteria, Dice and Dice
(1941) estimated that young cottontails still have their milk
dentition when they leave the nest at about a month of age. Weaning
has been reported as occurring at 17 days (V. Hayssen, unpub.), also
well before the age of any tooth replacement.
All members of the adult dentition of lagomorphs are
evergrowing; so it is remarkable that the milk predecessor of the
peglike upper incisor, and all the milk premolars, are rooted. These
milk teeth are virtually the same size as their replacements. It
would be interesting to know why, in a lineage that has apparently
suppressed development of canines and some incisors, these rooted
milk teeth are produced at all.
Macroscelidea
Elephant shrews are decidedly diphyodont (Table 3). Their milk
teeth are similar to, but often more elaborate than, the permanent
successors, and sometimes are larger as well (Hill 1938, Kindahl
1958, Leche 1897a).
81
A neonatal Elephantulus brachyurus had no teeth fully erupted,
but several incisors and premolars of the milk dentition had started
to pierce the gums (Hill 1938). Although known-age specimens are not
available, it is apparent from the size of the skull at tooth
replacement that the milk dentition is functional for a long time
(Hill 1938, Kindahl 1958). Since elephant shrews are precocial at
birth and are weaned early (15 days for Rhynchocyon chrysopygus, 22
for Elephantulus rufescens; V. Hayssen, unpub.), tooth replacement in
these animals probably occurs after weaning.
THE "TYPICAL" MAMMALIAN MILK DENTITION
The order-by-order review above shows considerable variety among
groups with respect to presence, function and timing of the milk and
replacement dentitions. One way to summarize this diverse information
is to measure it against some prevalent ideas of how tooth
replacement "typically" operates in mammals.
Extent of mammalian diphyodonty
The comparative anatomist Owen (1840) originated the term
"diphyodont" to refer to the condition of bearing one set of teeth
82
replaced by a second. The term is now used to indicate a general
mammalian trait, in contrast to tooth replacement in reptiles, which
produce one successor after another in each tooth position. But what
Owen actually intended was to divide the Mammalia into diphyodonts,
which replace teeth, and "monophyodonts," which do not.
Flower (1869) questioned such a division, since close
examination of embryos and young of presumed monophyodonts (seals,
for example) gave some evidence of another set of teeth. The search
for homologous structures was at its peak of interest, and the
ensuing emphasis on tracing both dental sets through the various
groups of the Mammalia obscured Owen's valid point that in
functional terms there are indeed many monophyodont mammals.
To assess the relative proportions of functionally diphyodont
and monophyodont mammals, I have compiled the set of described milk
dentitions and extrapolated these representative cases to related
species whose tooth replacement has never been described (Table 11).
Replacement characteristics of relatively few species (about 10
percent of the total) have actually been described, so unknown
species are classed with closest relatives.
A species is here considered diphyodont if it replaces more than
one tooth in any jaw quadrant and if the teeth of both the milk and
replacement sets function. The "non-diphyodont" category of Table 11
contains a variety of types. First, 24 species of mammmals are
functionally toothless throughout life. These include the pangolins,
83
Table 11. Estimated numbers of functionally diphyodont and

non-diphyodont mammal species.
DIPHYODONT NON-DIPHYODONT
(Monophyodont, toothless, etc.)
Taxon Number of Taxon Number of

species species
Edentata: Monotremata 3
Dasypodidae 20 Marsupialia 262
Insectivora (most) 83 Edentata (most groups) 9
Scandentia 16 Insectivora:
Dermoptera 2 Soricidae 277
Chiroptera (most) 789 Talpidae (part) 17
Primates 181 Chiroptera:
Carnivora (most) 227 Rhinolophidae 128
Perissodactyla 18 Carnivora:
Hyracoidea 7 Otariidae 14
Artiodactyla 186 Odobenidae 1
Rodentia: Fhocidae 19
Aplodontidae 1 Mustelidae
Sciuridae 261 (Mephitinae,
Lagomorpha 60 .. Meles) 10
Macroscelidea 15 Cetacea 76
Sirenia 5
Proboscidea 2
Orycteropodidae 1
Eholidota 7
Rodentia (most) 1433
TOTAL 1866 2264

(45%) (55%)
84
anteaters, monotremes, and baleen whales. Another group of
approximately 1500 species bear but one set of teeth with no fully
formed predecessors or replacements: these include the toothed
whales, sloths, skunks, shrews, many rodents, and a few bats. A third
group produces a rudimentary set of weakly calcified milk teeth for
which no function has been demonstrated or even proposed; often these
teeth never erupt. Pinnipeds and some moles and rodents belong in
this category. A fourth type, including elephants and manatees,
exhibits tooth succession but of a nonstandard type in which teeth
are shed from the front of the tooth row and replaced from the back;
they are thus not functionally diphyodont. Finally, most marsupials
and many rodents show what appears as a rather standard tooth
replacement, but of only one tooth per quadrant (last premolar); they
too are not strictly diphyodont. Tooth replacement in rodents is
poorly understood. The mountain beaver (Aplodontidae) and some
sciurids do replace more than one premolar in some jaw quadrants, so
as a conservative estimate all members of these groups are counted as
diphyodont. The actual number of diphyodont may be somewhat lower.
The non-diphyodont cases above comprise 2264 species. The true
diphyodonts, on the other hand, number approximately 1866. Thus the
term "diphyodont" as intended by Owen applies to only 45 percent of
the extant mammalian species. Over half the mammals produce fewer
than the "typical" two sets of teeth, and this fraction includes some
very speciose groups, notably the shrews and muroid rodents. Clearly
diphyodonty is not a requisite for success as a mammal; even if
85
(ifphyodonty way the ancestral, mammalian condition, it: has boon
subjected to a tremendous amount of modification.
MammaJ. tooth eruption sequence
Mamma J. teeth are generally sup posed to erupt from front to back;
indeed, f:ront-to-back tooth eruption is often cited as a defining
character of the Ciasy (McFarland et al 1979, Romer 8 Parsons 1.977,
Weller 1968). The significance of this pattern Is that it sets
mammals apart from reptiles, whose tooth eruption sequences are well
studied and often involve replacement of alternate teeth in an
apparent back-to-front pattern (Udmurt d I960, Osborn 1970). A number
of theoretical models have been proposed to account for the reptilian
and mammalian eruption-sequence patterns and to suggest how the
transition from alternate "back-to-front" to consecutive
f:ront-to-back sequences might have occurred (lidmund i960, Osborn
1970). The most elaborate recent version (Osborn 197'J) claims that
part of the mammalian dentition, the premolars, actually emerges from
back to front, and cites several, supporting examples. Osborn's
challenge to the traditional model begs the question; in what
sequence do mammal teeth actually erupt?
To investigate this issue I assembled eruption sequences for 215
mammal species from the literature (Appendix I); taxonomic
distribution of these is summarized in Table 12. When eruption
Table 12. Correspondence of known tooth eruption or development sequences

to three models. Columns three through five are the percentage of tooth
eruption sequences from the literature that are consistent with the model
designated at head of column. Sequences that have tied ranks and thus do
not discriminate between two models are listed with both; thus these are
maximum possible percentages. In parentheses below the total maximum per
centages are the minimum percentages of sequences consistent with the model
if those with tied ranks are excluded.
Order No. of No. of Percent agreement with model sequence

species sequences
MODEL: Front-to-back: Delayed-canine Osborn
Marsupialia 6 8 13% 13% 100%

Insectivora 8 24 17 17 63
Scandentia 2 7 29 29 43
Dermoptera 1 2 0 0 50
Chiroptera 16 44 59 59 64
Primates 59 209 35 56 57
Carnivora 55 161 25 27 57
Perissodactyla 4 12 8 8 25
Hyracoidea 1 4 25 25 25
Artiodactyla 48 139 51 51 68
Rodentia 11 20 95 95 95
Lagomorpha 2 4 50 50 100
Macroscelidea 2 4 0 0 50
Total maximum 215 638 38% 45% 61%

Total minimun (15%) (22%) (23%)
if tie ranks
excluded
87
sequences for upper and lower teeth and for milk and permanent series
are considered separately, the number of known sequences is 638.
305 of these cases show a generally front-to-back eruption order
(i.e. positive correlation between tooth position and eruption order,
Spearman rank correlation p<.05), but 333 do not. When a strict
front-to-back model (Figure la) is considered, only 38 percent of the
sequences are consistent with it (Table 12). Moreover, most of these
compatible sequences contain tie ranks (groups of teeth in which true
eruption sequence is not known), so that the actual fraction of cases
which fit the strict front-to-back model may be as low as 15 percent
(Table 12). As a variant on this model, Romer and Parsons (1977)
suggest that even though progression is usually front-to-back,
canines may erupt late (Figure lb). This pattern accounts for a
number of additional sequences, but the total consistent with the
model is still only 22 to 45 percent (Table 12). Osborn (1973), by
contrast, proposes a fundamental mammalian (or at least eutherian)
pattern wherein incisors and molars develop front to back but
premolars develop in the opposite direction (Figure 1c). This version
is the one best supported by the known sequences, since between 23
and 61 percent of them are consistent with it (Table 12). However,
even the best model still leaves at least 39 percent, and perhaps as
much as 77 percent, of the observed mammal tooth eruption sequences
unaccounted for. Thus a large fraction, and perhaps a majority, of
mammal tooth eruption sequences are "exceptions” that fit none of the
proposed "rules."
88
Figure 1. Models of "typical" tooth eruption sequences in mammals. On
the TOOTH axis, each block represents one tooth position of the
supposed "primitive" eutherian dentition; the left end of the axis
represents the front of the jaw, the right end the back. Positions
for three incisors, a canine, four premolars and three molars are
indicated. Along the ERUPTION ORDER axis, highest value represents
the earliest erupting tooth, and successively lower values later
erupting ones. la. Front-to-back model. Stair-step eruption sequence
to be expected in a mammal that erupts all teeth consecutively from
front to back. Jjb. Delayed-canine model. Eruption sequence expected
from Romer and Parsons (1977) model of delayed canine eruption in an
otherwise front-to-back progression, lc. Osborn model. Eruption
sequence according to Osborn's (1973) concept of dental development
in primitive Eutheria.
I
89
10
oc 9
LU
Q 8
oc
O 7
z 6
o 5
I— 4
CL.
3 3
oc
2
1
II 12 13 C PI P2 P3 P4 Ml M2 M3
TOOTH
10
oc 9
LU
Q 8
cm
O 7
Z 6
o 5
H- 4
a.
3 3
oc
2
I
II 12 13 C PI P2 P3 P4 Ml M2 M3
TOOTH
10
oc 9
LU
Q 8
oc
o 7
Z 6
o 5
I— 4
a.
3 3
04
2
I
II 12 13 C PI P2 P3 P4 Ml M2 M3
TOOTH
90
Milk teeth as stand-ins for permanent teeth
Mammalian milk teeth are seldom credited with any specialized
function; they are commonly supposed to be miniature place-holders
scaled down for a juvenile-size jaw (e.g. Pond 1977); if they
function at all it is presumed to be in the same fashion as their
permanent successors. Human deciduous teeth appear to fit this model,
but those of many other mammals do not. The exceptions fall into
several categories as follows:
a) Cases in which milk tooth is larger than successor. It is not
uncommon for a cheek tooth of the milk dentition to be larger than
its homologous replacement. In the giraffe, the upper posterior
premolar of the milk set is 18 percent, and the lower 34 percent,
longer (anterior-posterior length) then its replacement (Singer and
Bone 1960). This relationship holds for the pygmy chimpanzee (milk
upper 18 percent, lower 14 percent longer than permanent; Johanson
1974) and even humans (milk upper 33 percent, lower 36 percent longer
than permanent; Moorees et al 1957). The last milk premolar is
reported to be larger than its successor in springbok (Rautenbach
1971), duiker (Riney and Child 1960), elephant shrew (Hill 1938),
tree shrew (Leche 1897) European otter (Pohle 1919), and a variety of
hominoids (Ashton and Zuckerman 1950a and 1950b, Remane 1962) as
well, and the pattern is no doubt widespread. While many milk teeth
91
are small, the common-sense notion that "small jaws need small teeth"
clearly does not explain all milk dentitions.
b) Cases in which milk tooth is morphologically similar to a
non-homologous permanent tooth. Most of the examples of large milk
teeth cited above, and a suite of additional examples, are milk
premolars which are strikingly molariform yet are replaced by much
less molariform successors. In taxa where molar function is very
important (e.g. herbivores, omnivorous primates) there is obvious
functional advantage to having a milk tooth which will act as a
grinder until true molars erupt. Similarly, among the terrestrial
Carnivora, where the upper milk dp3 and lower milk dp4 form the
carnassial apparatus, there teeth morphologically resemble the true
carnassials rather than their own homologues. Surprisingly, no
current theory of tooth morphogenesis addresses how such changes of
form might develop ontogenetically, much less how they might have
evolved; yet a distinct morphological shift or switch between milk
and permanent homologues is very common.
c) Cases in which milk tooth is morphologically and/or
functionally unique. As discussed above, for the entire order
Chiroptera the milk teeth, where they exist, differ markedly in
morphology and presumably in function from the permanent set, and
show none of the dietary modifications known in adult bats. The
spikelet milk teeth of young pigs are also clearly functionally
different from those of adults. And in species such as baboons where
92
enlarged adult canines play a role in social interaction, the canines
of juveniles are not enlarged.
Overall, though there are many mammal species whose milk teeth
are morphologically similar to their permanent teeth, there are also
many exceptions. Even where milk and permanent dentitions function in
similar ways, it is often non-homologous teeth that are performing
analogous functions. There is no adequate evolutionary model to
account for these patterns, perhaps because the patterns themselves
have not previously been recognized.
"Milk teeth" and milk
A fourth prevalent assumption is that "milk teeth" (German
"Milchgebiss" , French "dents de lait") have some necessary connection
to lactation. Most commonly, milk teeth are believed to be present
during the suckling period, but are then "replaced at weaning" (Pond
1977). Weaning is sometimes even said to be instigated by tooth
replacement, as the juveniles' new teeth make nursing uncomfortable
for the mother.
The text and tables above summarize what is known about tooth
replacinent in relation to weaning among mammals. Of the 76 truly
diphyodont species for which there are sufficient data, 12 replace
all their teeth well before weaning and 56 do not begin tooth
93
replacement until weeks or months after weaning is complete. Only
Crocuta, two procyonids, and four mustelids appear to replace teeth
near the time of weaning, and in the case of the mustelids the
functionally most important teeth (canines and carnassials) are
replaced a considerable time after weaning. In summary, 89 percent of
known cases present "exceptions" to the "rule" of milk teeth being
replaced at the time of weaning.
CONCLUSIONS
This review of what is known of mammalian milk dentitions shows,
contrary to conventional assumptions, that most extant mammals are
not truly diphyodont; that few of them erupt their teeth from front
to back; that milk teeth often differ strikingly from permanent
teeth; and that there is no universal nor necessary relationship
between the timing of tooth replacement and the timing of weaning.
One occasionally encounters additional related generalizations,
explicit or more often implicit. Milk teeth have been alleged to be
more "primitive" than are permanent teeth (Bardenfleth 1913,
Dependorf 1896, Jorgenson 1956, Leche 1915, Lull 1907, Mivart 1882),
though the criteria for primitiveness are often vague. If this idea
could be substantiated, the milk dentition could provide valuable
phylogenetic clues; but if milk dentition is as evolutionarily
with permission of the copyright owner. Further reproduction prohibited without permission.
94
plastic as the permanent dentition, systematic inferences should be
viewed with caution. Recently tooth development and eruption
sequences have been promoted as phylogenetic characters (Broom and
Robinson 1951, Byrd 1981, Clements and Zuckerman 1953, Schwartz 1975a
and 1975b, Slaughter et al 1974). Again, the validity of the approach
depends upon the relative adaptive inflexibility of tooth eruption
sequences, but this has not been addressed. Further, attempts to
trace generalized mammalian-type "consecutive tooth replacement"
(Crompton and Jenkins 1973) from reptilian-type "alternate tooth
replacement" are misdirected if "consecutive tooth replacment" is a
myth. It is important before proceeding further with such
generalizations to evaluate them critically in reference to the whole
range of extant Mammalia. It is particularly important to try to
assess the role of adaptive variation and convergence in morphology
and timing, if milk tooth characters are to be used as systematic
tools.
Little consideration has been given in the past to the
functional significance of milk dentition nor to the potential for
evolutionary divergence between milk and permanent dentitions and the
timing of tooth replacement. From this review it appears that mammal
milk teeth are complex, variable, often functional, sometimes
specialized, and no more amenable to simplistic generalizations than
permanent teeth are.

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CHAPTER 1

Dentition has always been a major tool in the study of mammals,

largely because mammal teeth exhibit a rich array of fine variation

superimposed on a basically conservative, persistent structure. Of

the lifetime complement of two sets of teeth that mammals are

conventionally said to possess, the second or permanent set has been

analyzed in the contexts of age determination, chemical and physical

structure, comparative morphology, development, dietary

specialization, mechanical engineering, paleontology, pathology,

population genetics, predatory ecology, social behavior, and

added considerably to knowledge of adaptative processes. By contrast,

have relatively short tenure in an individual. Yet the juvenile

period in mammals is always a time of high growth rate, high

mortality and consequent high potential to affect fitness. An

assessment of the importance of milk teeth in the lives of juvenile

mammals would be of clear interest, the more so because the adaptive

significance of permanent teeth to adult mammals has already been so

In this study I make such an assessment. The first necessary

dentition in mammals; these patterns are documented in Chapter 2. I

analysis of milk tooth morphology and timing in relation to important

features of development (Chapter 3). This led to investigation of

some problems of tooth use that young carnivores face, in a

laboratory study of acquisition of predatory efficiency in

long-tailed weasels (Chapter 4). Ultimately, the clarified patterns

some current interpretations of what the two-tooth-set system

reflects about parental care in the first mammals; Chapter 5

reevaluates the paleontological evidence and proposes a different

hypothesis for the origin of mammalian milk teeth.

MAMMALIAN MILK TEETH: PRESENCE AND PATTERNS

Mammals are allegedly diphyodont. Every general textbook on

mammalogy or vertebrate biology claims so; attempts to characterize

the Class Mammalia invariably include the possession of two

dentitions (the milk teeth and a replacement set) as a diagnostic

feature (e.g. DeBlase and Martin 1982, McFarland et al 1979). The

diphyodont condition is one member of that suite of conditions

conventionally signalling the attainment of mammalian grade from

polyphyodont reptilian ancestors. Recently other mammalian inventions

have received review leading to theoretical frameworks and the posing

of investigative questions (e.g. Case 1978a and McNab 1978 on

endothermy, Crompton and Parker 1978 on the masticatory apparatus,

of diphyodonty in the extant Mammalia appeared in 1871, when William

Henry Flower addressed the Odontological Society of Great Britain on

Despite the lack of recent synthesis, there have been some

generalizations on the significance of diphyodonty and of the milk

dentition (e.g. see Pond 1977). These carry a certain degree of

logical appeal. First, since humans and an array of domestic species

characterize the Class as a whole. Replacement of teeth in humans

proceeds in a roughly consecutive order from front to back, so this

might seem a general mammalian development schedule. When baby

thus deciduous teeth might be expected to be morphologically like

replacement of "milk teeth" might thus be expected to show a

consistent relationship to the weaning process and to acquisition of

Evidence for these ideas, when supplied at all, is usually in

the form of individual cases. Cumulative data have not previously

been assembled. However, on closer examination it becomes apparent

that a great many species have been examined in some context.

Developmental biologists have reported tooth eruption as an

ontogenetic milestone. Wildlife population ecologists have drawn up

schedules for aging specimens from tooth eruption. Dental researchers

have described tooth development in laboratory species as human

analogs. Paleontologists have sought evidence of diphyodonty in

ancestral mammals. Systematists have made forays into use of milk

tooth characters in tracing phylogenies. Much descriptive information

thus exists: it would be important to exploit these data from