Chapter IV: Part 4

The largest representative of the family is the Tasmanian wolf, or
thylacine, alone representing the genus _Thylacinus_, in which the
dentition numbers i. 4/3, c. 1/1, p. 4/4, m. 3/3 = 46; with the
incisors small and vertical, the outer one in the upper jaw being
larger than the others. Summits of the lower incisors, before they are
worn, with a deep transverse groove, dividing it into an anterior and
a posterior cusp. Canines long, strong and conical. Premolars with
compressed crowns, increasing in size from before backwards. Molars in
general characters resembling those of _Sarcophilus_, but of more
simple form, the cusps being less distinct and not so sharply pointed.
Deciduous molar very small, and shed before the animal leaves the
mother's pouch. General form dog-like, with the head elongated, the
muzzle pointed, and the ears moderate, erect and triangular. Fur short
and closely applied to the skin. Tail of moderate length, thick at the
base and tapering towards the apex, clothed with short hair. First
hind toe (including the metacarpal bone) absent. Vertebrae: C. 7, D.
13, L. 6, S. 2, Ca. 23. Marsupial bones unossified. The gradual
passage of the thick root of the tail into the body is a character
common to the Tasmanian wolf and the aard-vark, and may be directly
inherited from reptilian ancestors (see THYLACINE).

The next genus is represented solely by the Tasmanian devil,
_Sarcophilus_ (or _Diabolus_) _ursinus_, a medium-sized animal with a
dental formula similar to that of the dasyures, but with teeth (fig.
2) approximating to those of the thylacine, though markedly different
in details. The first hind toe is absent.

In the "native cats," or dasyures, constituting the genus _Dasyurus_,
the dental formula is i. 4/3, c. 1/1, p. 3/3, m. 3/3: total 42. The
upper incisors are nearly equal and vertical, with the first slightly
longer, narrower, and separated from the rest. Lower incisors sloping
forward and upward. Canines large and sharply pointed. First two
premolars with compressed and sharp-pointed crowns, and slightly
developed anterior and posterior accessory basal cusps. Molars with
numerous sharp-pointed cusps. In the upper jaw the first two with
crowns having a triangular free surface; the last small, simple,
narrow and placed transversely. In the lower jaw the molars more
compressed, with longer cusps; the last not notably smaller than the
others. Ears of moderate size, prominent and obtusely pointed. First
hind toe rudimentary, clawless or absent; its metatarsal bone always
present. Tail generally long and well clothed with hair. Vertebrae: C.
7, D. 13, L. 6, S. 2, Ca. 18-20 (see DASYURE).

The genus _Phascologale_ comprises a number of small marsupials, none
exceeding a rat in size, differing from the dasyures in possessing an
additional premolar--the dentition being i. 4/3, c. 1/1, p. 4/4, m.
3/3: total 46--and in having the teeth generally developed upon an
insectivorous rather than a carnivorous pattern, the upper middle
incisors being larger and inclined forward, the canines relatively
smaller, and the molars with broad crowns, armed with prickly
tubercles. The muzzle is pointed. Ears moderately rounded, and nearly
naked. Fore feet with five sub-equal toes, with compressed, slightly
curved pointed claws. Hind feet with the four outer toes sub-equal,
with claws similar to those in the fore feet; the first toe almost
always distinct and partially opposable, though small and nailless,
sometimes absent.

In some respects intermediate between the preceding and the next genus
is _Dasyuroides byrnei_, of Central Australia, an animal of the size
of a rat, with one lower premolar less than in _Phascologale_, without
the first hind toe, and with a somewhat thickened tail. The pouch is
incomplete, with two lateral folds, and the number of teats six.

_Sminthopsis_ includes several very small species, with the same
dental formula as _Phascologale_, but distinguished from that genus by
the narrowness of the hind foot, in which the first toe is present,
and the granulated or hairy (in place of broad, smooth and naked)
soles. A pouch is present, and there are eight or ten teats. Nearly
allied is the jumping _Antechinomys laniger_, of East Central
Australia, an elegant mouse-like creature, with large oval ears,
elongated limbs, a long and tufted tail and no first hind toe. In
connexion with the large size of the ears is the excessive inflation
of the auditory bulla of the skull.

From all other members of the family the marsupial, or banded,
ant-eater (_Myrmecobius fasciatus_) differs by the presence of more
than seven pairs of cheek-teeth in each jaw, as well as by the
exceedingly long and protrusile tongue. Hence it is made the type of a
distinct sub-family, the _Myrmecobiinae_, as distinct from the
_Dasyurinae_, which includes all the other members of the family. From
the number of its cheek-teeth, the banded ant-eater has been regarded
as related to some of the primitive Jurassic mammals; but this view is
disputed by Mr Bensley, who regards this multiplicity of teeth as a
degenerate feature. On the other hand, it is noteworthy that this
marsupial retains in its lower jaw the so-called mylo-hyoid groove,
which is found in the aforesaid Jurassic mammals. _Myrmecobius_ has a
total of 52 or 54 teeth, which may be classed as i. 4/3, c. 1/1, p. +
m. (8 or 9)/(8 or 9). The teeth are all small and (except the four
posterior inferior molars) separated from each other by an interval.
Head elongated, but broad behind; muzzle long and pointed; ears of
moderate size, ovate and rather pointed. Fore-feet with five toes, all
having strong pointed, compressed claws, the second, third and fourth
nearly equal, the fifth somewhat and the first considerably shorter.
Hind-feet with no trace of first toe externally, but the metatarsal
bone is present. Tail long, clothed with long hairs. Fur rather harsh
and bristly. Female without pouch, the young when attached to the
nipples being concealed by the long hair of the abdomen. Vertebrae: C.
7, D. 13, L. 6, S. 3, Ca. 23. The single species, which is a native of
western and southern Australia, is about the size of an English
squirrel, to which its long bushy tail gives it some resemblance; but
it lives entirely on the ground, especially in sterile sandy
districts, feeding on ants. Its prevailing colour is chestnut-red, but
the hinder part of the back is marked with broad, white, transverse
bands on a dark ground.

FIG. 4.--The Marsupial or Banded Ant-eater (_Myrmecobius fasciatus_).]

With the bandicoots, or _Peramelidae_, we come to a family of
polyprotodonts which resemble the diprotodonts in the peculiarly
specialized structure of their hind limbs; an adaptation which we must
apparently regard as having been independently acquired in the two
groups. The dentition is i. 5/3, c. 1/1, p. 4/4, m. 3/3; total, 48;
the upper incisors being small, with short, broad crowns; the lower
incisors moderate, narrow, proclivous; canines well developed.
Premolars compressed, pointed; and the molars with quadrate
tuberculated crowns. Deciduous premolar preceded by a minute
molariform tooth, which remains in place until the animal is nearly
full grown. Fore feet with two or three of the middle toes of nearly
equal size, and provided with strong, sharp, slightly curved claws,
the other toes rudimentary. Hind feet long and narrow; the first toe
rudimentary or absent; the second and third very slender and united in
a common integument; the fourth very large, with a stout elongated
conical claw; the fifth smaller than the fourth (see fig. 6). The
terminal phalanges of the large toes of both feet cleft at their
extremities. Head elongated, with the muzzle long, narrow and pointed.
Stomach simple. Caecum of moderate size. Pouch complete, generally
opening backwards. Alone among marsupials bandicoots have no
clavicles. More remarkable still is the development of a small
allantoic placenta.

FIG. 5.--Gunn's Bandicoot (_Perameles gunni_).]

In the true bandicoots of the genus _Perameles_ (fig. 5) the fore-feet
have the three middle toes well developed, the third slightly larger
than the second, the fourth somewhat shorter, provided with long,
strong, slightly curved, pointed claws. First and fifth toes very
short and without claws. Hind feet with one or two phalanges, in the
first toe forming a distinct tubercle visible externally; the second
and third toes very slender, of equal length, joined as far as the
terminal phalange, but with distinct claws; the fifth intermediate in
length between these and the largely developed fourth toe. Ears of
moderate or small size, ovate, pointed. Tail rather short, clothed
with short depressed hairs. Fur short and harsh. Pouch opening
backwards. Vertebrae: C. 7, D. 13, L. 6, S. 1, Ca. 17. (see
BANDICOOT.)

c, calcanium; a, astralagus; cb, cuboid; n. navicular; c³,
ectocuneiform; II. and III. the conjoined second and third digits; IV.
the large and only functional digit; V. the rudimentary fifth digit.]

The rabbit-bandicoot, _Peragale_ (or _Thylacomys_) represents a genus
in which the cheek-teeth are curved, with longer crowns and shorter
roots than in the last. Hind extremities proportionally longer with
inner toe represented only by a small metatarsal bone. Muzzle much
elongated and narrow. Fur soft and silky. Ears very large, long and
pointed. Tail long, its apical half-clothed on the dorsal surface with
long hairs. Pouch opening forwards. Vertebrae: C. 7, D. 13, L. 6, S.
2, Ca. 23.

The one species, from Western Australia, is the largest member of the
family, being about the size of a rabbit, to which it bears sufficient
superficial resemblance to have acquired the name of "native rabbit"
from the colonists. It burrows in the ground, but in other respects
resembles bandicoots in habits.

In the pig-footed bandicoot (_Choeropus castanotis_) the dentition
generally resembles that of _Perameles_, but the canines are less
developed, and in the upper jaw two-rooted. Limbs very slender;
posterior nearly twice the length of the anterior. Fore feet with the
functional toes reduced to two, the second and third, of equal length,
with closely united metacarpals and short, sharp, slightly curved,
compressed claws. First toe represented by a minute rudiment of a
metacarpal bone; the fourth by a metacarpal and two small phalanges
without a claw, and not reaching the middle of the metacarpal of the
third; fifth entirely absent. Hind foot long and narrow, mainly
composed of the strongly developed fourth toe, terminating in a
conical pointed nail, with a strong pad behind it; the first toe
represented by a rudimentary metatarsal; the remaining toes completely
developed, with claws, but exceedingly slender; the united second and
third reaching a little way beyond the metatarso-phalangeal
articulation of the fourth; the fifth somewhat shorter. Tail not quite
so long as the body, and covered with short hairs. Ears large and
pointed, and folded down when the animal is at rest. Fur soft and
loose. Pouch opening backwards. Vertebrae: C. 7, D. 13, L. 6, S. 1,
Ca. 20.

The only species of this genus is about the size of a small rat, found
in the interior of Australia. Its general habits and food appear to
resemble those of other bandicoots. A separate family, _Notoryctidae_,
is represented by the marsupial mole (_Notoryctes typhlops_), of the
deserts of south Central Australia, a silky, golden-haired, burrowing
creature, with a curious leathery muzzle, and a short, naked stumpy
tail. The limbs are five-toed, with the third and fourth toes of the
front pair armed with enormous digging claws; there are no external
ear-conchs; and the dentition includes four pairs of upper, and three
of lower, incisors, and distinctly tritubercular cheek-teeth. The
small pouch, supported by the usual epipubic bones, opens backwards.
In correlation with its burrowing habits, some of the vertebrae of the
neck and of the loins are respectively welded together. The eyes have
degenerated to a greater extent than those of any other burrowing
mammal, the retina being reduced to a mass of simple cells, and the
cornea and sclerotic ("white") to a pear-shaped fibrous capsule
enclosing a ball of pigment. The reason for this extreme degeneration
is probably to be found in the sandy nature of the soil in which the
creature burrows, a substance which would evidently irritate and
inflame any functional remnant of an eye. The portion of the lachrymal
duct communicating with the cavity of the nose has, on the other hand,
been abnormally developed, apparently for the purpose of cleansing
that chamber from particles of sand which may obtain an entrance while
the animal is burrowing. (See MARSUPIAL MOLE.)

FIG. 7.--The Pig-footed Bandicoot (_Choeropus castanotis_).]

FIG. 8.--Skull of _Caenolestes obscurus_.]

2. _Paucituberculates._--The second sub-order of marsupials, the
Paucituberculata, is exclusively South American, and typically
represented by the family _Epanorthidae_, the majority of the members
of which are extinct, their remains being found in the probably
Miocene Santa Cruz beds of Patagonia, although one existing genus
(_Caenolestes_) survives in Ecuador and Colombia. One of the two
living species was, indeed, described so long ago as the year 1863,
under the preoccupied name of _Hyracodon_, but attracted little or no
attention, as its affinities were not fully recognized. Externally
_Caenolestes_ has a shrew-like appearance. The elongated skull (fig.
8) has four pairs of upper incisors and long upper canines, while in
the lower jaw there is a single pair of procumbent incisors, followed
by several small teeth representing the canine and earlier premolars.
The three pairs of molars in each jaw are, like the last premolar,
quadritubercular oblong teeth. The five-toed feet are of normal
structure, and the rat-like tail is prehensile towards the tip. The
female has a small pouch. The extinct members of the family are
represented by the genera _Epanorthus_, _Acdestis_, _Garzonia_, &c. In
a second family--_Abderitidae_--also from the Patagonian Miocene, the
penultimate premolar is developed into an enormous tooth, with a tall,
secant and grooved crown, somewhat after the fashion of the enlarged
premolar of _Plagiaulax_. From the structure of the skull, it is
thought probable that _Abderites_ had an elongated snout, like that of
many Insectivora. As a sub-order, the Paucituberculata are
characterized by the presence of four pairs of upper and three of
lower incisor teeth; the enlargement and forward inclination of the
first pair of lower incisors, and the presence of four or five sharp
cusps on the cheek-teeth, coupled with the absence of "syndactylism"
in the hind limbs.

FIG. 9.--Front view of Skull of the Koala (Phascolarctus cinereus) to
exhibit Diprotodont type of dentition.]

3. _Diprotodonts._--The third and last sub-order of marsupials is the
Diprotodontia, which is exclusively Australasian and includes the
wombats, koala, cuscuses, kangaroos and their relatives. There are
never more than three pairs of upper and one of lower incisors, of
which the middle upper and the single lower pair are large and
chisel-like (fig. 9); the canines are small or absent; the cheek-teeth
have bluntly tuberculate or transversely-ridged crowns in most cases;
and the hind-feet are syndactylous. With one exception, the intestine
has a caecum, and the pouch is large and opens forwards. It should be
added that Professor Elliot Smith has pointed out a certain
peculiarity in its commissures whereby the brain of the diprotodonts
differs markedly from that of the polyprotodonts and approximates to
the placental type. Dr Einar Lönnberg has also recorded certain
adaptive peculiarities in the stomach. Most of the species,
particularly the specialized types, are more or less completely
herbivorous.

The first family, _Phascolomyidae_, is typified by the wombats; but
according to the view adopted by Mr H. Winge, and endorsed by
Professor Max Weber, is also taken to include the koala. In this wider
sense the family may be characterized as follows. The tympanic process
of the alisphenoid bone of the skull is short, not covering the cavity
of the tympanum, nor reaching the paroccipital process. The tail is
rudimentary, the first hind-toe opposable, the first pair of upper
incisors very large, but the second and third either absent or small
and placed partially behind the larger pair; and only five pairs of
cheek-teeth in each jaw. The stomach has a cardiac gland, and the
number of teats is two.

In the wombats (_Phascolomys_) the dentition is i. 1/1, c. 0/0, p. +
m. 5/5, total 24; all the teeth growing from persistent pulps, and the
incisors large and chisel-like, with enamel only on the front surface.
The cheek-teeth strongly curved, forming from the base to the summit
about a quarter of a circle, the concavity being directed outwards in
the upper and inwards in the lower teeth. The first of the series
(which appears to have no predecessor) single-lobed; the other four
composed of two lobes, each subtriangular in section. Limbs equal,
stout and short. Fore-feet with five distinct toes, each furnished
with a long, strong and slightly curved nail, the first and fifth
considerably shorter than the other three. Hind-feet with a very short
nailless first toe, the second, third and fourth toes partially united
by integument, of nearly equal length, the fifth distinct and rather
shorter; all four with long and curved nails. In the skeleton the
second and third toes are distinctly more slender than the fourth,
showing a tendency towards the character so marked in the following
families. Tail rudimentary. Caecum very short and wide, with a
vermiform appendage (see WOMBAT).

In addition to remains referable to the existing genus, the
Pleistocene deposits of Australia have yielded evidence of an extinct
giant wombat constituting the genus _Phascolonus_ (_Sceparnodon_).

The koala, or "native bear" (_Phascolarctus cinereus_), which differs
widely from the wombats in its arboreal habits, is less specialized as
regards its dentition, of which the formula is i. 3/1, c. 1/0, p. + m.
5/5, total 30. Upper incisors crowded together, cylindroidal, the
first much larger than the others, with a bevelled cutting edge (fig.
9). Canine very small; a considerable interval between it and the
first premolar, which is as long from before backwards but not so
broad as the molars, and has a cutting edge, with a smaller parallel
inner ridge. The molar-like teeth slightly diminishing in size from
the first to the fourth, with square crowns, each bearing four
pyramidal cusps. The lower incisors are partially inclined forwards,
compressed and tapering, bevelled at the ends. Cheek-teeth in
continuous series, as in the upper jaw. Fore-feet with the two inner
toes slightly separated from and opposable to the remaining three, all
with strong curved and much compressed claws. Hind-foot (fig. 10) with
the first toe placed far back, large and broad, the second and third
(united) toes considerably smaller than the other two; the fourth the
largest. No external tail. Fur dense and woolly. Ears of moderate
size, thickly clothed with long hair. Caecum very long and dilated,
with numerous folds. Vertebrae: C. 7, D. 11, L. 8, S. 2, Ca. 8. Ribs
eleven pairs (see KOALA).

Here may be noticed three genera of large extinct marsupials from the
Pleistocene of Australia whose affinities appear to ally them to the
wombat-group on the one hand and to the phalangers on the other. The
longest known is _Diprotodon_, an animal of the size of a rhinoceros,
with a dental formula of i. 3/1, c. 0/0, p. 1/1, m. 4/4, total 28. The
first upper incisor very large and chisel-like, molars with prominent
transverse ridges, as in _Macropus_, but without the longitudinal
connecting ridge. Complete skeletons disinterred by Dr E. C. Stirling
indicate that in the structure of the feet this creature presents
resemblances both to the wombats and the phalangers, but is nearer to
the former than to the latter. On the other hand, the considerably
smaller _Nototherium_, characterized by its sharp and broad skull and
smaller incisors, seems to have been much more wombat-like, and may
perhaps have possessed similar burrowing habits.

FIG. 11.--Front view of Skull of _Thylacoleo carnifex_, restored.]

The last of the three is _Thylacoleo carnifex_, so named on account of
its supposed carnivorous habits. In the adult the dentition (fig. 11)
is i. 3/1, c. 1/0, p. + m. 4/3, total 24. The first upper incisor is
much larger than the others; canine and first two premolars
rudimentary. In the lower jaw there are also one or two small and
early deciduous premolars; third premolars of both jaws formed on the
same type as that of the rat-kangaroos, but relatively much larger;
molars rudimentary, tubercular. The functional teeth are reduced to
one pair of large cutting incisors situated close to the middle line,
and one great, cutting, compressed premolar, on each side above and
below. As already mentioned, _Thylacoleo_ was originally regarded as a
carnivorous creature, but this view was subsequently disputed, and its
diet supposed to consist of soft roots, bulbs and fruits, with an
occasional small bird or mammal. Recently, however, the pendulum of
opinion has swung back towards the original view: and Dr R. Broom
believes _Thylacoleo_ to have been "a purely carnivorous animal, and
one which would be quite able to, and probably did, kill animals as
large or larger than itself." The affinities of the creature are
clearly with the phalangers.

By means of the little musk-kangaroo, the cuscuses and phalangers
constituting the family _Phalangeridae_, are so closely connected with
the kangaroos, or _Macropodidae_, that in the opinion of some
naturalists they ought all to be included in a single family, with
three sub-families. Theoretically, no doubt, this is correct, but the
typical members of the two groups are so different from one another
that, as a matter of convenience, the retention of the two families
seems advisable. From the _Phascolomyidae_, the two families, which
may be collectively designated Phalangeroidea, differ by the
circumstance that in the skull the tympanic process of the alisphenoid
covers the tympanic cavity and reaches the paroccipital process. The
tail is long and in some cases prehensile; the first hind-toe may be
either large, small or absent; the dentition usually includes three
pairs of upper and one of lower incisors, and six or seven pairs of
cheek-teeth in each jaw; the stomach is either simple or sacculated,
without a cardiac gland; and there are four teats.

With the exception of the aberrant long-snouted phalanger, the members
of the family _Phalangeridae_ have the normal number of functional
incisors, in addition to which there may be one or two rudimentary
pairs in the lower jaw. The first in the upper jaw is strong, curved
and cutting, the other two generally somewhat smaller; the single
lower functional incisor large, more or less inclined forwards;
canines 1/(1 or 0) upper small or moderate, conical and sharp-pointed;
lower absent or rudimentary; premolars variable; molars 3/3, or 2/2,
with four obtuse tubercles, sometimes forming crescents. Limbs
subequal. Fore-feet with five distinct subequal toes with claws.
Hind-feet short and broad, with five well-developed toes; the first
large, nailless and opposable; the second and third slender and united
by a common integument as far as the claws. Caecum present (except in
_Tarsipes_), and usually large. The lower jaw has no pocket on the
outer side. All are animals of small or moderate size and arboreal
habits, feeding on a vegetable or mixed diet, and inhabiting
Australia, Papua and the Moluccan Islands.

FIG. 12.--The Long-snouted Phalanger (_Tarsipes rostratus_).]

As the first example of the group may be taken the elegant little
long-snouted phalanger (_Tarsipes rostratus_, fig. 12), a west
Australian creature of the size of a mouse, which may be regarded as
representing by itself a sub-family (_Tarsipediinae_), characterized
by the rudimentary teeth, the long and extensile tongue, and absence
of a caecum. The head is elongated, with a slender muzzle and the
mouth-opening small. The two lower incisors are long, very slender,
sharp-pointed and horizontally placed. All the other teeth are simple,
conical, minute and placed at considerable and irregular intervals
apart in the jaws, the number appearing to vary in different
individuals and even on different sides of the jaw of the same
individuals. The formula in one specimen was i.(2 - 2)/(1 - 1), c.(1 -
1)/(0 - 0), p. + m.(3 - 4)/(2 - 3); total 20. The lower jaw is
slender, nearly straight, and without a coronoid process or inflected
angle. Fore-feet with five well-developed toes, carrying small, flat,
scale-like nails, not reaching the extremity of the digits. Hind-feet
rather long and slender, with a well-developed opposable and nailless
first toe; second and third digits united, with sharp, compressed
curved claws; the fourth and fifth free, with small flat nails. Ears
of moderate size and rounded. Tail longer than the body and head,
scantily clothed with short hairs, prehensile. Vertebrae: C. 7, D. 13,
L. 5, S. 3. Ca. 24.

As indicated in the accompanying illustration, the long-snouted
phalanger is arboreal in habits, extracting honey and probably small
insects from long-tubed flowers by means of its extensile tongue.

The remaining members of the family may be included in the sub family
_Phalangerinae_, characterized by the normal nature of the dentition
(which shows rudimentary lower canines) and tongue. Cuscuses and
phalangers form a numerous group, all the members of which are
arboreal, and some of which are provided with lateral expansions of
skin enabling them to glide from tree to tree like flying-squirrels.
The typical members of the group are the cuscuses (_Phalanger_),
ranging from the Moluccas and Celebes to New Guinea, in which the
males are often different in colour from the females. The true
phalangers, or opossums of the colonists, constitute the genus
_Trichosurus_, while the ring-tailed species are known as
_Pseudochirus_; the latter ranging to New Guinea. _Dactylopsila_ is
easily recognized by its attenuated fourth finger and parti-coloured
fur; the flying species are classed as _Petauroides_, _Petaurus_,
_Gymnobelideus_ and _Acrobates_, the last no larger than a mouse;
while Dromicia, _Distaechurus_ and _Acrobates_ are allied types
without parachutes (see PHALANGER).

An equally brief notice must suffice of the kangaroo tribe or
_Macropodidae_, since these receive a special notice elsewhere. The
dentition is i.(3/1) c.(0 or 1)/0 p.(3/3) m.(3/3); the incisors being
sharp and cutting, and those of the lower jaw frequently having a
scissor-like action against one another. The broad molars are either
bluntly tuberculated or transversely ridged; the outer side of the
hind part of the lower jaw has a deep pocket; and the hind-limbs are
generally very long, with the structure of the foot similar to that of
the bandicoots. The family is connected with the _Phalangeridae_ by
means of the musk-kangaroo (_Hypsiprymnodon moschatus_); forming the
sub-family _Hypsiprymnodontinae_. Then come the rat-kangaroos, or
kangaroo-rats, constituting the sub-family _Potoroinae_; while the
tree-kangaroos (_Dendrolagus_), rock-wallabies (_Petrogale_), and
wallabies and kangaroos (_Macropus_) form the _Macropodinae_ (see
KANGAROO).

_Extinct Marsupials_

Reference has been made to the Australasian Pleistocene genera
_Phascolonus_, _Diprotodon_, _Nototherium_ and _Thylacoleo_, whose
affinities are with the wombats and phalangers. The same deposits have
also yielded remains of extinct types of kangaroo, some of gigantic
size, constituting the genera _Sthenurus_, _Procoptodon_ and
_Palorchestes_. Numerous types more or less nearly allied to the
phalangers, such as _Burramys_ and _Triclis_ have also been described,
as well as a flying form, _Polaeopetaurus_. It is also interesting to
note that fossil remains indicate the former occurrence of thylacines
and Tasmanian devils on the Australian mainland. Of more interest is
the imperfectly known _Wynyardia_, from older Tertiary beds in
Tasmania, which apparently presents points of affinity both to
phalangers and dasyures. From the Oligocene deposits of France and
southern England have been obtained numerous remains of opossums
referable to the American family _Didelphyidae_. These ancient
opossums have been separated generically from _Didelphys_ (in its
widest sense) on account of certain differences in the relative sizes
of the lower premolars, but as nearly the whole of the species have
been formed on lower jaws, of which some hundreds have been found, it
is impossible to judge how far these differences are correlated with
other dental or osteological characters. In the opinion of Dr H.
Filhol, the fossils themselves represent two genera, _Peratherium_,
containing the greater part of the species, about twenty in number,
and _Amphiperatherium_, with three species only. All are comparatively
small animals, few of them exceeding the size of a rat.

Besides these interesting European fossils, a certain number of
didelphian bones have been found in the caves of Brazil, but these are
either closely allied to or identical with the species now living in
the same region.

The occurrence in the Santa Cruz beds of Patagonia of fossil
marsupials allied to the living _Caenolestes_ has been mentioned
above. The alleged occurrence in the same beds of marsupials allied to
the thylacine is based on remains now more generally regarded as
referable to the creodont carnivores (see CREODONTA).

_Mesozoic Mammals._--Under the heading of MULTITUBERCULATA will be
found a brief account of certain extinct mammals from the Mesozoic
formations of Europe and North America which have been regarded as
more or less nearly related to the monotremes. The same deposits have
yielded remains of small mammals whose dentition approximates more
nearly to that of either polyprotodont marsupials or insectivores; and
these may be conveniently noticed here without prejudice to their true
affinities. Before proceeding further it may be mentioned that the
remains of many of these mammals are very scarce, even in formations
apparently in every way suitable to the preservation of such fossils,
and it hence seems probable that these creatures are stragglers from
a country where primitive small mammals were abundant. Not improbably
this country was either "Gondwana-land," connecting Mesozoic India
with Africa, or perhaps Africa itself. At any rate, there seems little
doubt that it was the region where creodonts and other primitive
mammals were first differentiated from their reptilian ancestors.

FIG. 13.--Lower Jaw of _Triconodon mordax_ (nat. size).]

FIG. 14.--Lower Jaw and Teeth of _Phascolotherium bucklandi_ (nat.
size in outline).]

FIG. 15.--Spalacotherium tricuspidens (twice nat. size), Purbeck
beds.]

Of the Old World forms, the family _Triconodontidae_ is typified by
the genus _Triconodon_, from the English Purbeck, in which the
cheek-teeth carry three cutting cusps arranged longitudinally. There
seems to have been a replacement of some of these teeth; and it has
been suggested that this was of the marsupial type. To the same family
are referred _Phascolotherium_ (fig. 14), of the Lower Jurassic
Stonesfield slate of England, and _Spalacotherium_ (fig. 15), of the
Dorsetshire Purbeck; the latter having the three cusps of the
cheek-teeth rotated so as to assume a tritubercular type. Other genera
are _Menacodon_ and _Priacodon_, the former American, and the latter
common to Europe and North America. By one authority _Amphilestes_
(fig. 16), of the Stonesfield Slate, is included in the same group,
while by a second it is regarded as representing a family by itself.
_Amphitherium_, of the Stonesfield Slate, typifies the family
_Amphitheriidae_, which includes the American _Dryolestes_, and in
which some would class the European Purbeck genus _Amblotherium_,
although Professor H. F. Osborn has made the last the type of a
distinct family. Yet another family, according to the palaeontologist
last named, is typified by the genus _Stylacodon_, of the English
Purbeck. To mention the other forms which have received names will be
unnecessary on this occasion.

FIG. 16.--Lower Jaw and Teeth of _Amphilestes broderipi_ (twice nat.
size).]

It will be observed from the figures of the lower jaws, which are in
most cases the only parts known, that in many instances the number of
cheek-teeth exceeds that found in modern marsupials except
_Myrmecobius_. The latter has indeed been regarded as the direct
descendant of these Mesozoic forms; but as already stated, in the
opinion of Mr B. A. Bensley, this is incorrect. It may be added that
the division of these teeth into premolars and molars in figs. 14 and
16 is based upon the view of Sir R. Owen, and is not altogether
trustworthy, while the restoration of some of the missing teeth is
more or less conjectural. As regards the affinities of the creatures
to which these jaws belonged, Professor Osborn has referred the
_Triconodontidae_ and _Amphitheriidae_, together with the
Curtodontidae (as represented by the English Purbeck _Curtodon_), to a
primitive group of marsupials, while he has assigned the
_Amblotheriidae_ and _Stylacodontidae_ to an ancestral assemblage of
Insectivora. On the other hand, in the opinion of Professor H. Winge,
a large number of these creatures are primitive monotremes. Besides
the above, in the Trias of North America we have _Dromotherium_ and
_Microconodon_, extremely primitive forms, representing the family
_Dromotheriidae_, and apparently showing decided traces of reptilian
affinity. It may be added that a few traces of mammals have been
obtained from the English Wealden, among which an incisor tooth
foreshadows the rodent type.

AUTHORITIES.--The above article is partly based on that by Sir W. H.
Flower in the 9th edition of this work. See also O. Thomas, Catalogue
of Monotremata and Marsupialia in the British Museum (1888); "On
_Caenolestes_, a Survivor of the _Epanorthidae," Proc. Zool. Soc.
London_ (1895); J. D. Ogilby, Catalogue of Australian Mammals (Sydney,
1895); B. A. Bensley, "A Theory of the Origin and Evolution of the
Australian Marsupialia," _American Naturalist_ (1901); "On the
Evolution of the Australian Marsupialia, &c.," _Trans. Linn. Soc._
(vol. ix., 1903); L. Dollo, "Arboreal Ancestry of Marsupials,"
_Miscell. Biologiques_ (Paris, 1899); B. Spencer, "Mammalia of the
Horn Expedition" (1896); "Wynyardia, a Fossil Marsupial from
Tasmania," _Proc. Zool. Soc. London_ (1900); J. P. Hill,
"Contributions to the Morphology of the Female Urino-genital Organs in
Marsupialia," _Proc. Linn. Soc. N. S. Wales_, vols. xxiv. and xxv.;
"Contributions to the Embryology of the Marsupialia," _Quart. Journ.
Micr. Science_, vol. xliii.; E. C. Stirling, "On _Notoryctes
typhlops_," _Proc. Zool. Soc. London_ (1891); "Fossil Remains of Lake
Cadibona," Part I. _Diprotodon, Mem. R. Soc. S. Australia_ (vol. i.,
1889); R. Broom, "On the Affinities of _Thylacoleo," Proc. Linn. Soc.
N. S. Wales_ (1898); H. F. Osborn, "Mesozoic Mammalia," _Journ. Acad.
Nat. Sci. Philadelphia_ (vol. ix., 1888); E. S. Goodrich, "On the
Fossil Mammalia from the Stonesfield Slate," _Quart. Journ. Micr.
Science_ (vol. xxxv., 1894). (R. L.*)

FOOTNOTE:

[1] The presence or absence of the corpus callosum has been much
disputed; the latest researches, however, indicate its absence.

MARSUPIAL MOLE (_Noloryctes typhlops_), the "Ur-quamata" of the natives, an aberrant polyprotodont from central South Australia, constituting a family (_Noloryctidae_). This is a small burrowing animal, of a pale golden-yellow colour, with long silky hair, a horny shield on the nose, and a stumpy leathery tail. The feet are five-toed, and the third and fourth toes of the front pair armed with enormous claws adapted for digging. Neither ear-conches nor eyes are visible externally. There are but three pairs of incisor teeth in each jaw, and the upper molars are tricuspid. This animal spends most of its time burrowing in the sand in search of insects and their larvae, but occasionally makes its appearance on the surface.

MARSUS, DOMITIUS, Latin poet, the friend of Virgil and Tibullus, and contemporary of Horace. He survived Tibullus (d. 19 B.C.), but was no longer alive when Ovid wrote (c. A.D. 12) the epistle from Pontus (_Ex Ponto_, iv. 16) containing a list of poets. He was the author of a collection of epigrams called _Cicuta_ ("hemlock")[1] from their bitter sarcasm, and of a beautiful epitaph on the death of Tibullus; of elegiac poems, probably of an erotic character; of an epic poem _Amazonis_; and of a prose work on wit (_De urbanitate_). Martial often alludes to Marsus as one of his predecessors, but he is never mentioned by Horace, although a passage in the _Odes_ (iv. 4, 19) is supposed to be an indirect allusion to the _Amazonis_ (M. Haupt, _Opuscula_, iii. 332).

See J. A. Weichert, _Poetarum latinorum vitae et reliquiae_ (1830); R.
Unger, _De Dom. Marsi cicuta_ (Friedland, 1861).

FOOTNOTE:

[1] According to others, a reed-pipe made of the stalks of hemlock;
the reading _scutica_ ("whip") has also been proposed.

MARSYAS, in Greek mythology, a Phrygian god or Silenus, son of Hyagnis. He was originally the god of the small river of the same name near Celaenae, an old Phrygian town. He represents the art of playing the flute as opposed to the lyre--the one the accompaniment of the worship of Cybele, the other that of the worship of Apollo. According to the legend, Athena, who had invented the flute, threw it away in disgust, because it distorted the features. Marsyas found it, and having acquired great skill in playing it, challenged Apollo to a contest with his lyre. Midas, king of Phrygia, who had been appointed judge, declared in favour of Marsyas, and Apollo punished Midas by changing his ears into ass's ears. In another version, the Muses were judges and awarded the victory to Apollo, who tied Marsyas to a tree and flayed him alive. Marsyas, as well as Midas and Silenus, are associated in legend with Dionysus and belong to the cycle of legends of Cybele. A statue of Marsyas was set up in the Roman forum and colonies as a symbol of liberty. The contest and punishment of Marsyas were favourite subjects in Greek art, both painting and sculpture. In Florence there are several statues of Marsyas hanging on the tree as he is going to be flayed (see GREEK ART, fig. 54, Pl. II.); Apollo and the executioner complete the group. In the Lateran museum at Rome there is a statue representing Marsyas in the act of picking up the flute, a copy of a masterpiece by Myron (Hyginus, _Fab._ 167, 191; Apollodorus i. 4, 2; Ovid, _Metam._ vi. 382-400, xi. 145-193), for which see GREEK ART, fig. 64 (Pl. III.).

MARTABAN, a town in the Thaton district of Lower Burma, on the right bank of the Salween, opposite Moulmein. It is said to have been founded in A.D. 573, by the first king of Pegu, and was once the capital of a powerful Talaing kingdom; but it is now little more than a village. Martaban is frequently mentioned by European voyagers of the 16th century; and it has given the name of "Martavans" to a class of large vessels of glazed pottery, also known in India as "Pegu jars." It was twice captured by the British, in 1824 and 1852. The Bay of Martaban receives the rivers Irrawaddy and Salween.

MARTELLO TOWER, a kind of tower formerly used in English coast defence. The name is a corruption of Mortella. The Martello tower was introduced in consequence of an incident of the French revolutionary wars. In September 1793 a British squadron of three ships of the line and two frigates was ordered to support the Corsican insurgents. It was determined in the first place to take a tower on Cape Mortella which commanded the only secure anchorage in the Gulf of San Fiorenzo. This tower, according to James, was named "after its inventor"; but the real derivation appears to be the name of a wild myrtle which grew thickly around. The tower, which mounted one 24-pounder and two 18-pounders on its top, was bombarded for a short time by the frigates, was then deserted by its little garrison, and occupied by a landing party. The tower was afterwards retaken by the French from the Corsicans. So far it had done nothing to justify its subsequent reputation. In 1794, however, a fresh attempt was made to support the insurgents. On the 7th of February 1400 troops were landed, and the tower was attacked by land and sea on the 8th. The "Fortitude" and "Juno" kept up a cannonade for 2½ hours and then hauled off, the former being on fire and having sixty-two men killed and wounded. The fire from the batteries on shore produced no impression until a hot shot set fire to the "bass junk with which, to the depth of 5 ft., the immensely thick parapet was lined." The garrison of thirty-three men then surrendered. The armament was found to consist only of two 18-pounders and one 6-pounder. The strong resistance offered by these three guns seems to have led to the conclusion that towers of this description were specially formidable, and Martello towers were built in large numbers, and at heavy expense, along the shores of England, especially on the southern and eastern coasts, which in certain parts are lined with these towers at short intervals. They are structures of solid masonry, containing vaulted rooms for the garrison, and providing a platform at the top for two or three guns, which fire over a low masonry parapet. Access is provided by a ladder, communicating with a door about 20 ft. above the ground. In some cases a deep ditch is provided around the base. The chief defect of the tower was its weakness against vertical fire; its masonry was further liable to be cut through by breaching batteries. The French _tours modèles_ were somewhat similar to the Martello towers; their chief use was to serve as keeps to unrevetted works. While the Martello tower owes its reputation and its widespread adoption in Great Britain to a single incident of modern warfare, the round masonry structure entered by a door raised high above the base is to be found in many lands, and is one of the earliest types of masonry fortification.

MARTEN, HENRY (1602-1680), English regicide, was the elder son of Sir Henry Marten, and was educated at University College, Oxford. As a public man he first became prominent in 1639 when he refused to contribute to a general loan, and in 1640 he entered parliament as one of the members for Berkshire. In the House of Commons he joined the popular party, spoke in favour of the proposed bill of attainder against Strafford, and in 1642 was a member of the committee of safety. Some of his language about the king was so frank that Charles demanded his arrest and his trial for high treason. When the Great Rebellion broke out Marten did not take the field, although he was appointed governor of Reading, but in parliament he was very active. On one occasion his zeal in the parliamentary cause led him to open a letter from the earl of Northumberland to his countess, an impertinence for which, says Clarendon, he was "cudgelled" by the earl; and in 1643, on account of some remark about extirpating the royal family, he was expelled from parliament and was imprisoned for a few days. In the following year, however, he was made governor of Aylesbury, and about this time took some small part in the war. Allowed to return to parliament in January 1646, Marten again advocated extreme views. He spoke of his desire to prepare the king for heaven; he attacked the Presbyterians, and, supporting the army against the parliament, he signed the agreement of August 1647. He was closely associated with John Lilburne and the Levellers, and was one of those who suspected the sincerity of Cromwell, whose murder he is said personally to have contemplated. However, he acted with Cromwell in bringing Charles I. to trial; he was one of the most prominent of the king's judges and signed the death warrant. He was then energetic in establishing the republic and in destroying the remaining vestiges of the monarchical system. He was chosen a member of the council of state in 1649, and as compensation for his losses and reward for his services during the war, lands valued at £1000 a year were settled upon him. In parliament he spoke often and with effect, but he took no part in public life during the Protectorate, passing part of this time in prison, where he was placed on account of his debts. Having sat among the restored members of the Long Parliament in 1659, Marten surrendered himself to the authorities as a regicide in June 1660, and with some others he was excepted from the act of indemnity, but with a saving clause. He behaved courageously at his trial, which took place in October 1660, but he was found guilty of taking part in the king's death. Through the action, or rather the inaction of the House of Lords, he was spared the death penalty, but he remained a captive, and was in prison at Chepstow Castle when he died on the 9th of September 1680. Although a leading Puritan, Marten was a man of loose morals. He wrote and published several pamphlets, and in 1662 there appeared _Henry Marten's Familiar Letters to his Lady of Delight_, which contained letters to his mistress, Mary Ward.

Marten's father, Sir Henry Marten (c. 1562-1641), was born in London and was educated at Winchester school and at New College, Oxford, becoming a fellow of the college in 1582. Having become a barrister, he secured a large practice and soon came to the front in public life. He was sent abroad on some royal business, was made chancellor of the diocese of London, was knighted, and in 1617 became a judge of the admiralty court. Later he was appointed a member of the court of high commission and dean of the arches. He became a member of parliament in 1625, and in 1628 represented the university of Oxford, taking part in the debates on the petition of right.

See J. Forster, _Statesmen of the Commonwealth_ (1840); M. Noble,
_Lives of the English Regicides_ (1798); the article by C. H. Firth in
_Dict. Nat. Biog._ (1893); and S. R. Gardiner, _History of the Great
Civil War_ and _History of the Commonwealth and Protectorate_.

MARTEN,[1] a name originally belonging to the pine-marten (_Mustela martes_), but now applied to all members of the same genus of carnivorous mammals (see CARNIVORA). Martens are limited to the northern hemisphere, ranging throughout the greater part of the northern temperate regions of both Old and New Worlds, and southwards in America to 35° N. lat., while in Asia one species is met with in Java.

The species appear to be similar in their habits. They live in woods and rocky places, and spend most of their time in trees, although descending to the ground in quest of prey. They climb with great facility, and are agile and graceful in their movements. Some are said occasionally to resort to berries and other fruit for food, but as a rule they are carnivorous, feeding chiefly on birds and their eggs, small mammals, as squirrels, hares, rabbits and moles, but chiefly mice of various kinds, and occasionally snakes, lizards and frogs. In proportion to their size they are among the most bloodthirsty of animals, though less so than the weasels. The female makes her nest of moss, dried leaves and grass in the hollow of a tree, but sometimes in a hole among rocks or ruined buildings, and produces several young at a birth, usually from four to six. Though wild and untameable to a great degree if captured when fully grown, if taken young they are docile, and have frequently been made pets, not having the strong unpleasant odour of the smaller _Mustelidae_. The pine-marten appears to have been partially domesticated by the Greeks and Romans, and used to keep houses clear from rats and mice. In the same way, according to Brian Hodgson, the yellow-bellied weasel (_Putorius kathia_) "is exceedingly prized by the Nepalese for its service in ridding houses of rats. It is easily tamed; and such is the dread of it common to all murine animals that not one will approach a house where it is domiciled." It is, however, to the great value attached to the pelts of these animals that their importance to man is chiefly due. Though all yield fur of serviceable quality, the commercial value varies immensely, not only according to the species from which it is obtained, but according to individual variation, depending upon age, sex, season, and other circumstances. The skins from northern regions are more full and of a finer colour and gloss than those from more temperate climates, as are those of animals killed in winter compared to the same individuals in summer. Fashion has, moreover, set fictitious values upon slight shades of colour. Enormous numbers of animals are caught, chiefly in traps, to supply the demand of the fur trade, Siberia and North America being the principal localities from which they are obtained.

With the exception of the pekan (_M. pennanti_), the martens are much
alike in size, general colouring and cranial and dental characters.
The following description by Dr Elliott Coues of the American marten
(_M. americana_) will apply almost equally well to most of the others.
"It is almost impossible to describe the colour of the marten, except
in general terms, without going into the details of the endless
diversities occasioned by age, sex, season, or other incidents. The
animal is 'brown,' of a shade from orange or tawny to quite blackish;
the tail and feet are ordinarily the darkest, the head lightest, often
quite whitish; the ears usually have a whitish rim, while on the
throat there is usually a large tawny-yellowish or orange-brown patch,
from the chin to the fore legs; sometimes entire, sometimes broken
into a number of smaller, irregular blotches, sometimes wanting,
sometimes prolonged on the whole under surface, when the animal is
bicolor like a stoat in summer. The general 'brown' has a greyish
cast, as far as the under fur is concerned, and is overlaid with rich
lustrous blackish-brown in places where the long bristly hairs
prevail. The claws are whitish; the naked nose pad and whiskers are
black. The tail occasionally shows interspersed white hairs, or a
white tip."

The following are the best-known species:--

_Mustela foina_: the beech-marten, stone-marten or white-breasted
marten.--Distinguished from the following by the greater breadth of
the skull, and some minute but constant dental characters, by the dull
greyish-brown colour of the fur of the upper parts and the pure white
of the throat and breast. It inhabits the greater part of the
continent of Europe, but is more southern than the next in its
distribution, not being found in Sweden or Norway.

_M. martes_, the pine-marten (see figure).--Fur rich dark brown; under
fur reddish-grey, with clear yellow tips; breast spot usually yellow,
varying from bright orange to pale cream-colour or yellowish-white.
Length of head and body 16 to 18 in., of tail (including the hair) 9
to 12 in. This species is extensively distributed throughout northern
Europe and Asia, and was formerly common in most parts of Great
Britain and Ireland. It is still found in the northern counties of
England and North Wales, but in decreasing numbers. In Scotland it is
rare, but in Ireland may be found in almost every county occasionally.
Though commonly called "pine-marten," it does not appear to have any
special preference for coniferous trees.

Next comes _M. zibellina_, the sable (German, _Zobel_ and _Zebel_;
Swedish, _sabel_; Russian, _sobel_, a word probably of Turanian
origin), which closely resembles the last, if indeed it differs except
in the quality of the fur--the most highly valued of that of all the
group. The sable is found chiefly in eastern Siberia.

Very distinct is the brilliantly coloured orange-and-black Indian
marten (_M. flavigula_), found from the Himalaya and Ceylon to Java.

The North American _M. americana_ is closely allied to the pine-marten
and Asiatic sable. The importance of the fur of this animal as an
article of commerce may be judged of from the fact that 15,000 skins
were sold in one year by the Hudson's Bay Company as long ago as 1743.
It is ordinarily caught in wooden traps of simple construction, being
little enclosures of stakes or brush in which the bait is placed upon
a trigger, with a short upright stick supporting a log of wood, which
falls upon its victim on the slightest disturbance. A line of such
traps, several to a mile, often extends many miles. The bait is any
kind of meat, a mouse, squirrel, piece of fish or bird's head. It is
principally trapped during the colder months, from October to April,
when the fur is in good condition, as it is nearly valueless during
the shedding in summer. It maintains its numbers partly in consequence
of its shyness, which keeps it away from the abodes of men, and partly
because it is so prolific, bringing forth six to eight young at a
litter. Its home is sometimes a den under ground or beneath rocks, but
oftener the hollow of a tree, and it is said to take possession of a
squirrel's nest, driving off or devouring the rightful proprietor.

The pekan or Pennant's marten, also called fisher marten, though there
appears to be nothing in its habits to justify the appellation, is the
largest of the group, the head and body measuring from 24 to 30 in.,
and the tail 14 to 18 in. It is also more robust in form than the
others, its general aspect being more that of a fox than a weasel; in
fact its usual name among the American hunters is "black fox." Its
general colour is blackish, lighter by mixture of brown or grey on the
head and upper fore part of the body, with no light patch on the
throat, and unlike other martens generally darker below than above. It
was generally distributed in wooded districts throughout the greater
part of North America, as far north as Great Slave Lake, lat. 63° N.,
and Alaska, and extending south to the parallel of 35°; but at the
present time is almost exterminated in the settled parts of the United
States east of the Mississippi. (W. H. F.)

FOOTNOTE: