Mammalian Diversity and Evolution
Mammalian Diversity and Evolution
1627-BS-Z-23
BS (HONS) ZOOLOGY
SEMESTER: 2nd
SUBJECT: zoology
Diversity of mammals with evolutionary
perspectives
Table of Contents
Introduction:..........................................................................................................................................3
Overview of mammalian diversity:........................................................................................................4
Terrestrial mammals..........................................................................................................................5
Aquatic mammals..............................................................................................................................5
Flying mammals.................................................................................................................................5
Evolutionary Origins of Mammals.........................................................................................................6
The ancestry of mammals.....................................................................................................................8
Amniotes...........................................................................................................................................8
Synapsids...........................................................................................................................................9
Therapsids.......................................................................................................................................10
Biarmosuchia...................................................................................................................................10
Dinocephalians................................................................................................................................11
Anomodonts................................................................................................................................11
Theriodonts......................................................................................................................................11
Cynodonts........................................................................................................................................12
Fossil evidence and transitional forms................................................................................................13
Triassic takeover:.............................................................................................................................13
From cynodonts to crown mammals................................................................................................14
Fossil record....................................................................................................................................14
Morganucodontidae........................................................................................................................15
Docodonts.......................................................................................................................................15
Kuehneotheriidae.........................................................................................................................16
Monotremes.....................................................................................................................................17
Theria..........................................................................................................................................17
Metatheria.......................................................................................................................................18
Eutheria...........................................................................................................................................19
Adaptive Radiation of Mammals:....................................................................................................20
Key features of the adaptive radiation of mammals include:..........................................................21
Overview of extinction events in mammalian history:........................................................................21
Anthropogenic threats to mammalian diversity:.............................................................................21
Future directions for research and conservation efforts:....................................................................22
Conservation efforts and mitigating human impact on mammals:......................................................22
References:..........................................................................................................................................23
Topic:
Diversity of mammals with evolutionary perspectives
Introduction:
Mammals use two bones for hearing that all other amniotes use for eating. The earliest
amniotes had a jaw joint composed of the articular (a small bone at the back of the lower jaw)
and the quadrate (a small bone at the back of the upper jaw). All non-mammalian this system
including amphibians, turtles, lizards, snakes, crocodilians, dinosaurs (including
the birds), ichthyosaurs, pterosaurs and therapsids. But mammals have a different jaw joint,
composed only of the dentary (the lower jaw bone, which carries the teeth) and
the squamosal (another small skull bone). In the Jurassic, their quadrate and articular bones
evolved into the incus and malleus bones in the middle ear. Mammals also have a
double occipital condyle; they have two knobs at the base of the skull that fit into the topmost
neck vertebra, while other tetrapods have a single occipital condyle.
In a 1981 article, Kenneth A. Kermack and his co-authors argued for drawing
the line between mammals and earlier synapsids at the point where the mammalian pattern
of molar occlusion was being acquired and the dentary-squamosal joint had appeared. The
criterion chosen, they noted, is merely a matter of convenience; their choice was based on the
fact that "the lower jaw is the most likely skeletal element of a Mesozoic mammal to be
preserved."
Terrestrial mammals:
Aquatic mammals:
These mammals also have evolved specialized adaptations for life in water, including
streamlined bodies, fins or flippers, and blubber for insulation. This group includes cetaceans
(whales, dolphins, and porpoises), pinnipeds (seals, sea lions, and walruses), and sirenians
(manatees and dugongs). Aquatic mammals inhabit oceans, seas, rivers, and freshwater
habitats, where they play important ecological roles as top predators, filter feeders, and
ecosystem engineers.
Flying mammals
They are represented primarily by bats, are the only mammals capable of sustained
powered flight. Bats have evolved unique wing structures and echolocation abilities, enabling
them to navigate and hunt prey in the air. With over 1,400 species, bats are one of the most
diverse groups of mammals and inhabit diverse ecosystems worldwide, from tropical
rainforests to deserts and even urban areas.
Permian period: During its late, approximately 260 million years ago, a lineage of
synapsids known as therapsids emerged. Therapsids exhibited several important evolutionary
innovations that foreshadowed the characteristics of mammals, including modifications to the
skull, teeth, and limbs. These adaptations likely conferred advantages in terms of locomotion,
feeding, and [Link] of the key innovations of therapsids was the development
of a secondary palate, a bony structure that separated the nasal passages from the mouth. This
allowed for more efficient breathing while eating and is considered a precursor to the
mammalian condition. Therapsids also evolved differentiated teeth, with specialized incisors,
canines, and molars adapted for various feeding strategies.
Triassic period: During this, approximately 230 million years ago, a group of therapsids
known as cynodonts underwent further evolutionary changes that brought them closer to the
mammalian condition. Cynodonts possessed more mammal-like jaw joints, which allowed for
a more efficient chewing motion. Additionally, they had enlarged temporal fenestrae and
began to show evidence of mammalian ear structures, with modifications to the jaw bones
that eventually gave rise to the middle ear bones of [Link] the late Triassic and early
Jurassic periods, approximately 200 million years ago, true mammals had emerged. These
early mammals, known as the mammaliaforms, exhibited several key mammalian features,
including mammary glands, hair, and a specialized middle ear structure.
The ancestry of mammals
Amniotes
The first fully terrestrial vertebrates were reptilian amniotes — their eggs had
internal membranes that allowed the developing embryo to breathe but kept water in. This
allowed amniotes to lay eggs on dry land, while amphibians generally need to lay their eggs
in water (a few amphibians, such as the common Suriname toad, have evolved other ways of
getting around this limitation). The first amniotes apparently arose in the
middle Carboniferous from the ancestral reptiliomorphs
Within a few million years, two important amniote lineages became distinct: synapsids, from
which mammals are descended, and sauropsids, from which lizards, snakes, turtles/tortoises,
crocodilians, dinosaurs, and birds are [Link] earliest known fossils of synapsids and
sauropsids (such as Archaeothyris and Hylonomus, respectively) date from about 320 to 315
million years ago. The times of origin are difficult to know, because vertebrate fossils from
the late Carboniferous are very rare, and therefore the actual first occurrences of each of these
types of animal might have been considerably earlier than the first fossil
Synapsids
Synapsid skulls are identified by the distinctive pattern of the holes behind each eye, which
served the following purposes:
A number of creatures often – and incorrectly – believed to be dinosaurs, hence part of the
reptile lineage and sauropsids, were in fact synapsids. This includes the well-
known Dimetrodon.
When referring to the ancestors and close relatives of mammals, paleontologists also use the
following terms of convenience:
Pelycosaurs — all synapsids, and all of their descendants, except for therapsids – the
eventual ancestor of mammals. The pelycosaurs included the largest land vertebrates of
the Early Permian, such as the 6 metre (20 foot)-long Cotylorhynchus hancocki. Among
the other large pelycosaurs were Dimetrodon grandis and Edaphosaurus cruciger.
Stem mammals (sometimes called protomammals or paramammals, and previously
called mammal-like reptiles) all synapsids, and all of their descendants, except
for mammals [Link] mammals therefore include all pelycosaurs, and also all
non-mammalian therapsids. Traditionally these were known as "mammal-like reptiles",
but this is incorrect; terms such as "stem mammal" are preferred instead, because these
synapsids were neither reptiles nor even part of reptile lineage.
Pelycosaur and "mammal-like reptile" are both paraphyletic terms. The modern reptiles,
all being sauropsids, evolved in parallel to the synapsids, thus under the crown group use of
the term "reptile", mammals did not evolve from them. For that reason are disfavored and
outdated terms rarely used in modern literature.
Therapsids
In the middle Permian, therapsids supplanted sphenacodonts, a primitive
synapsid, as the main land vertebrates. Several characteristics of the skull and jaws, such as
greater temporal fenestrae and equal-sized incisors, set them apart from previous synapsids.
Following a series of stages in the therapsid lineage, cynodonts evolved in the late Permian,
with some of them starting to resemble early mammals. The slow emergence of a secondary
palate made of bone. Due to their ability to breathe and eat simultaneously, mammals' high
metabolic rate is interpreted by most books and articles as having evolved as a result of this.
However, some scientists point out that a bony palate provides a surface on which the tongue
may manipulate food, facilitating chewing rather than breathing, and that some current
ectotherms use a fleshy secondary palate to separate the mouth from the airway
Biarmosuchia:
The Biarmosuchia were the most primitive and pelycosaur-like of the therapsids
Dinocephalians:
Dinocephalians ("terrible heads") included both carnivores and herbivores. They
were large; Anteosaurus was up to 20 ft (6.1 m) long. Some of the carnivores had semi-erect
hindlimbs, but all dinocephalians had sprawling forelimbs. In many ways they were very
primitive therapsids; for example, they had no secondary palate and their jaws were rather
"reptilian
Lystrosaurus, one of the few genera of dicynodonts that survived the Permian–Triassic
extinction event
Anomodonts
The anomodonts ("anomalous teeth") were among the most successful of the
herbivorous therapsids — one sub-group, the dicynodonts, survived to the end of the Triassic.
But dicynodonts were very different from modern herbivorous mammals, as their only teeth
were a pair of fangs in the upper jaw (lost in some derived kannemeyeriiformes) and it is
generally agreed that they had beaks like those of birds or ceratopsians.
Theriodonts
The theriodonts ("beast teeth") and their descendants had jaw joints in which
the articular bone of the lower jaw tightly gripped the very small quadrate bone of the skull.
This allowed a much wider gape and allowed one group, the
carnivorous gorgonopsians ("gorgon faces"), to develop "sabre teeth". However, the jaw
hinge of the theriodont had a longer term significance — the much reduced size of the
quadrate bone was an important step in the development of the mammalian jaw joint and
middle ear.
The gorgonopsians still had some primitive features: no bony secondary palate (other bones
in the right places perform the same functions); sprawling forelimbs; hindlimbs that could
operate in both sprawling and erect postures. The therocephalians ("beast heads"), which
appear to have arisen at about the same time as the gorgonopsians, had additional mammal-
like features, e.g. their finger and toe bones had the same number of phalanges (segments) as
in early mammals (and the same number that primates have, including humans.
Cynodonts
The cynodonts, a theriodont group that also arose in the late Permian, include
the ancestors of all mammals. Cynodonts' mammal-like features include further reduction in
the number of bones in the lower jaw, a secondary bony palate, cheek teeth with a complex
pattern in the crowns, and a brain which filled the endocranial cavity.
Multi-chambered burrows have been found, containing as many as 20 skeletons of the Early
Triassic cynodont Trirachodon; the animals are thought to have been drowned by a flash
flood. The extensive shared burrows indicate that these animals were capable of complex
social behaviors.
Their primitive synapsid and therapsid ancestors were very large (between 5–8 ft (1.5–
2.4 m)) but cynodonts gradually decreased in size (to 1.5–5 ft (0.46–1.52 m)) even before
the Permian-Triassic extinction event, probably due to competition with other therapsids.
After the extinction event, the probainognathian cynodont group rapidly decreased in size (to
4–18 in (100–460 mm)) due to new competition with archosaurs and transitioned
to nocturnality, evolving nocturnal features, pulmonary alveoli, bronchioles and a
developed diaphragm for a larger surface area for breathing, enucleated erythrocytes, a large
intestine which bears a true colon after the cecum, endothermy, a hairy, glandular and
thermoregulatory skin (which releases sebum and sweat), and a 4-chambered heart to
maintain their high metabolism, larger brains, and fully upright hindlimb (forelimbs remained
semi sprawling, and became like that only later, in therians). Some skin glands may have
evolved into mammary glands in females for fulfilling the metabolic demands of their
offspring (which increased 10 times).
Fossil evidence and transitional forms
Triassic takeover:
The catastrophic mass extinction at the end of the Permian, around 252 million
years ago, killed off about 70% of terrestrial vertebrate species and the majority of land
plants.
As a result, ecosystems and food chains collapsed, and the establishment of new stable
ecosystems took about 30 million years. With the disappearance of the gorgonopsians, which
were dominant predators in the late Permian, the cynodonts' principal competitors for
dominance of the carnivorous niches were a previously obscure sauropsid group,
the archosaurs, which includes the ancestors of crocodilians and dinosaurs.
The archosaurs quickly became the dominant carnivores, a development often called the
"Triassic takeover". Their success may have been due to the fact that the early Triassic was
predominantly arid and therefore archosaurs' superior water conservation gave them a
decisive advantage. All known archosaurs have glandless skins and eliminate nitrogenous
waste in a uric acid paste containing little water, while the cynodonts probably excreted most
such waste in a solution of urea, as mammals do today; considerable water is required to keep
urea dissolved.
The therapsid trend towards differentiated teeth with precise occlusion accelerated,
because of the need to hold captured arthropods and crush their exoskeletons.
As the body length of the mammals' ancestors fell below 4 in (100 mm), advances
in thermal insulation and temperature regulation would have become necessary for
nocturnal life.
Acute senses of hearing and smell became vital.
o This accelerated the development of the mammalian middle ear (though the complete
detachment of the middle ear bones from the jaw happened independently
in monotremes and therians).
o The increase in the size of the olfactory lobes of the brain increased brain weight as a
percentage of total body weight Brain tissue requires a disproportionate amount of
energy. The need for more food to support the enlarged brains increased the pressures
for improvements in insulation, temperature regulation and feeding.
Probably as a side-effect of the nocturnal life, mammals lost two of the four cone opsins,
photoreceptors in the retina, present in the eyes of the earliest amniotes. Paradoxically,
this might have improved their ability to discriminate colors in dim light.
This retreat to a nocturnal role is called a nocturnal bottleneck, and is thought to explain
many of the features of mammals.s
From cynodonts to crown mammals
Fossil record
Mesozoic synapsids that had evolved to the point of having a jaw joint composed of the
dentary and squamosal bones are preserved in few good fossils, mainly because they were
mostly smaller than rats:
They were largely restricted to environments that are less likely to provide
good fossils. Floodplains as the best terrestrial environments for fossilization provide few
mammal fossils, because they are dominated by medium to large animals, and the
mammals could not compete with archosaurs in the medium to large size range.
Their delicate bones were vulnerable to being destroyed before they could be fossilized
— by scavengers (including fungi and bacteria) and by being trodden on.
Small fossils are harder to spot and more vulnerable to being destroyed by weathering
and other natural stresses before they are discovered.
In the past 50 years, however, the number of Mesozoic fossil mammals has increased
decisively; only 116 genera were known in 1979, for example, but about 310 in 2007, with an
increase in quality such that "at least 18 Mesozoic mammals are represented by nearly
complete skeletons".
Cynodonts to crown group mammals
Morganucodontidae
The Morganucodontidae first appeared in the late Triassic, about 205
million years ago. They are an excellent example of transitional fossils, since they have both
the dentary-squamosal and articular-quadrate jaw joints. [42] They were also one of the first
discovered and most thoroughly studied of the mammaliaforms outside of the crown-
group mammals, since an unusually large number of morganucodont fossils have been found.
Docodonts
Docodonts, among the most common Jurassic mammaliaforms, are noted for the
sophistication of their molars. They are thought to have had general semi-aquatic tendencies,
with the fish-eating Castorocauda ("beaver tail"), which lived in the mid-Jurassic about
164M years ago and was first discovered in 2004 and described in 2006, being the most well-
understood example. Castorocauda was not a crown group mammal, but it is extremely
important in the study of the evolution of mammals because the first find was an almost
complete skeleton (a real luxury in paleontology) and it breaks the "small nocturnal
insectivore" stereotype
Hadrocodium skull.
Kuehneotheriidae
The family Kuehneotheriidae, known from the Late Triassic and Early Jurassic,
was originally classified as part of either 'Symmetrodonta' or 'Pantotheria' based on their
tooth structure, with Kuehneotherium once being considered the oldest known representative
of Theria. They have since been recovered as among the closest relatives of crown-group
mammals. As only tooth fossils have been discovered, they nevertheless remain poorly
known, and have rarely been included in phylogenetic studies.
Monotremes
Teinolophos, from Australia, is the earliest known monotreme. A 2007 study
(published 2008) suggests that it was not a basal (primitive, ancestral) monotreme but a full-
fledged platypus, and therefore that the platypus and echidna lineages diverged considerably
earlier. A more recent study (2009), however, has suggested that, while Teinolophos was a
type of platypus, it was also a basal monotreme and predated the radiation of modern
monotremes. The semi-aquatic lifestyle of platypuses prevented them from being
outcompeted by the marsupials that migrated to Australia millions of years ago, since joeys
need to remain attached to their mothers and would drown if their mothers ventured into
water (though there are exceptions like the water opossum and the lutrine opossum; however,
they both live in South America and thus do not come into contact with monotremes).
Genetic evidence has determined that echidnas diverged from the platypus lineage as recently
as 19-48M, when they made their transition from semi-aquatic to terrestrial lifestyle
Monotremes have some features that may be inherited from the cynodont ancestors:
like lizards and birds, they use the same orifice to urinate, defecate and reproduce
("monotreme" means "one hole").
they lay eggs that are leathery and uncalcified, like those of lizards, turtles and
crocodilians.
Unlike other mammals, female monotremes do not have nipples and feed their young by
"sweating" milk from patches on their bellies.
Theria
Theria ("beasts") is the clade originating with the last common ancestor of
the Eutheria (including placentals) and Metatheria (including marsupials). Common features
include
no interclavicle.
coracoid bones non-existent or fused with the shoulder blades to form coracoid processes.
A type of crurotarsal ankle joint in which: the main joint is between
the tibia and astragalus; the calcaneum has no contact with the tibia but forms a heel to
which muscles can attach. (The other well-known type of crurotarsal ankle is seen in
crocodilians and works differently — most of the bending at the ankle is between the
calcaneum and astragalus).
tribosphenic molars.
Metatheria
The living Metatheria are all marsupials (animals with pouches). A few fossil genera,
such as the Mongolian late Cretaceous Asiatherium, may be marsupials or members of some
other metatherian groupsThe oldest known metatherian is Sinodelphys, found in 125M-year-
old early Cretaceous shale in China's northeastern Liaoning Province. The fossil is nearly
complete and includes tufts of fur and imprints of soft tissues. Didelphimorphia (common
opossums of the Western Hemisphere) first appeared in the late Cretaceous and still have
living representatives, probably because they are mostly semi-
arboreal unspecialized omnivores. Tracks from the Early Cretaceous of Angola show the
existence of raccoon-size mammals 118 million years ago.
The mother develops a kind of yolk sack in her womb that delivers nutrients to
the embryo. Embryos of bandicoots, koalas and wombats additionally form placenta-like
organs that connect them to the uterine wall, although the placenta-like organs are smaller
than in placental mammals and it is not certain that they transfer nutrients from the
mother to the embryo.
Pregnancy is very short, typically four to five weeks. The embryo is born at a very early
stage of development, and is usually less than 2 in (51 mm) long at birth. It has been
suggested that the short pregnancy is necessary to reduce the risk that the
mother's immune system will attack the embryo.
The newborn marsupial uses its forelimbs (with relatively strong hands) to climb to
a nipple, which is usually in a pouch on the mother's belly. The mother feeds the baby by
contracting muscles over her mammary glands, as the baby is too weak to suck. The
newborn marsupial's need to use its forelimbs in climbing to the nipple was historically
thought to have restricted metatherian evolution, as it was assumed that the forelimb
could not become specialised intro structures like wings, hooves or flippers. However,
several bandicoots, most notably the pig-footed bandicoot, have true hooves similar to
those of placental ungulates, and several marsupial gliders have evolved.
Eutheria
The time of appearance of the earliest eutherians has been a matter of
controversy. On one hand, recently discovered fossils of Juramaia have been dated to 160
million years ago and classified as eutherian. Fossils of Eomaia from 125 million years ago in
the Early Cretaceous have also been classified as eutherian.A recent analysis of phenomic
characters, however, classified Eomaia as pre-eutherian and reported that the earliest clearly
eutherian specimens came from Maelestes, dated to 91 million years ago. That study also
reported that eutherians did not significantly diversify until after the catastrophic extinction at
the Cretaceous–Paleogene boundary, about 66 million years ago.
Eomaia was found to have some features that are more like those of marsupials and earlier
metatherians:
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