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    Mammalian Specializations

    Chapter 21

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    Mammalian groups Monotremes

    Marsupials

    Eutherians

    Have varying reproductive modes Egg laying in monotremes

    Eutherians have long gestations Marsupials have very short gestation

    lengths

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    Common reproductive aspects Blastocyts

    An embryonic ball of cells that forms the embryo

    All mammals grow from this blastocyst Trophoblast

    An embryonic tissue of mammals specialized forimplanting the the embryo onto the uterine wall

    (in Therians), obtaining nutrients from the mother,and secreting hormones to signal the state ofpregnancy to the mother

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    Common reproductive aspects Endometrium

    Glandular uterine epithelium of the

    mammals that secrete materials thatnourish the embryo in uterus

    Presence of corpus luteum Formed by the ruptures follicles after

    releasing egg

    Secretes hormones that sustain earlystages of pregnancy

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    Monotreme Reproduction Primitive reproductive tract

    2 oviducts remain separate, do not fuse

    during development except at the basewhere they join with urethra to formurogenital sinus (Fig 21.4 a)

    Oviducts swell to form uterus that retains

    the fertilized egg Fertilization occurs in the anterior portion

    of the oviduct (fallopian tube)

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    Monotreme Reproduction Ovaries larger in compares to Therians

    Monotremes provide embryo with moreyolk

    Produce smaller eggs at ovulation

    Eggs retained in uterus & nourished bymaternal secretions, increase in sizeafter which the shell is secreted.

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    Monotreme Reproduction Egg shell is leathery

    1-2 eggs laid at hatching

    Hatching is rapid (7-10 days) In platypus only left oviduct is functional and

    hatching is ~ 12 days

    Lay eggs in burrows, but echidnas lay eggs in

    a ventral pouch Young hatch as embryos, and brooding has

    to continue for about 16 weeks (fig. 21.1)

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    Reproduction in Therians All have placentation: 2 types

    Choriovitelline placenta

    Placentas developed from the yolk sac seen in allTherian animals during early development

    Chorioallatontoic placenta Developed from the chorionic & allantoic extra-

    embryonic membranes Grows out & takes over from the CV placenta

    Typical trait of all eutherians

    Most marsupials have only one CV placenta, butsome show a transitory CA placenta at end ofgestation

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    Reproduction in Therians Embryonic diapause

    Maintaining eggs in a state of arrested

    development before implantation as in Kangaroos; Carnivores; Rodents; Bats

    Enables mating and birth of young to occur

    at optimal times of the year

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    Reproduction in Therians Male reproductive anatomy

    Monotremes retain testes in abdomen

    In therians, testes descend into scrotum

    Descent is genetically controlled inmarsupials and hormonally controlled in

    eutherians Scrotum in front of penis in marsupials and

    behind in most eutherians.

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    Reproduction of Eutherians Ureters enter into the bladder rather than the

    cloaca

    Oviducts fuse anterior to the urogenital sinusto form a uterus

    All have a single midline vaginam but only afew have a single midline uterus as seen in

    humans Some have a bipartite uterus for some or all

    of its length. Bipartite uterus is abnormal inhumans

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    Reproduction of Eutherians Urogenital sinus and alimentary canal

    have separate openings

    Space between them is perineum(space between anus and vagina)

    In primates the urogenital sinus

    separates into distinct vaginal andurethral openings

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    Reproduction of Eutherians Corpus luteum is maintained for a larger period than

    one estrus cycle

    Allows for larger gestation lengths Some young are altricial (rodents & insectivorous)

    Other young are precocial (most ungulates)

    All young require lactation for transfer of essential

    antibodies Almost all ungulates bear one precocial young

    Parturition and lactation are hormonal

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    Reproduction of Marsupials Females

    Female oviducts do not join on midline becauseureters pass medial to reproductive ducts to enterbladder

    2 separate uteri

    2 vaginae, Lateral one for sperm passage only

    Pseduovaginal canal for parturition Corpus luteum is not maintained

    Young ejected at end of estrus cycle

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    Reproduction of Marsupials Young ones are neonates

    Well developed limbs, jaws, secondary palate,

    large lungs, tongue and facial muscles Climb up the pouch and attach to the nipples

    Some ejected directly into the pouch or mammaryarea of pouchless animals

    Pouch absent in some: mice & Opposums Lactation continues after young ones detach from

    the pouch

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    Feeding specializations:

    Dentition Incisors to seize food

    Canines to stab prey

    Premolars to pierce and crash food

    Molars: to break down food into fineparticles

    Therians have tribosphenic molars

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    Feeding specializations:

    Dentition Canines

    Lost in herbivores or modified

    Tusks of pigs and walruses: modifiedcanines

    Upper canines larger in male primates

    Male horses have small functionlesscanines

    Maybe used in male fighting and display

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    Feeding specializations:

    Dentition Incisors

    Tusks of elephants= modified incisors

    Enlarged in gnawing mammals and growcontinuously throughout life (rabbits,rodents)

    Rodent incisors have only enamel in theanterior

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    Feeding specializations:

    Dentition Premolars

    Single cusped for slicing food

    Molars:

    3- cusps for thorough food processing

    In many herbivores both molars and

    premolars are the same as in horses

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    Feeding specializations:

    Dentition Molars of Omnivorous & Fruit eating

    mammals

    Cusps are rounded, flattened structuresideal for crushing

    Upper molars: 4th cusp

    Molars called bunodonts (fig 21.6 e) since they appear like a square rather than

    triangular and also the rounded nature

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    Feeding specializations:

    Dentition Molars of herbivores

    Teeth have ridges called lophs that help to phone

    some kind of shearing blades Lophodont teeth

    Straight lophs (kangaroos, rabbits)

    Selenodont teeth

    Molars have crescent lophs as in of artiodactyls (deer) Multilophed teeth: lamellar

    Wombarts, warthogs, rodents, elephants

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    Feeding specializations:

    Dentition Dental durability

    Diphyodont condition: adult dentition must

    last a life time Problem for herbivores who have to deal

    with more abrasive vegetation

    Grazers also have to deal with high toothwear due to silica in grasses

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    Feeding specializations:

    Dentition Solutions

    Hyposodont teeth

    Highly crowned teeth. Crown extends deep intothe jaw bone

    Deep lower jaws & deep cheek regions Brachyodont: low crowned teeth- primitive mode

    Larger hyposodont mammals (ungulates) Layer of cementum to cover whole tooth

    Usually covers only root & base of crown

    Cementum is a bone-like material, fills the highlophs of teeth

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    Feeding specializations:

    Dentition Still teeth get worn out

    Animals cant eat anymore

    Horses (20-30) should be fed soft food No molars left

    Hypselodont mammals

    Molar teeth with evergrowing crowns Roots do not close

    Unique in small mammals: rodents and rabbits

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    Carnivorous mammals Have large canines to subdue prey

    Specialized post-canine teeth for

    shearing

    E.g Carnassials

    A pair of teeth specialized as tearing blades

    Formed by last premolar in upper jaw andist molar in lower jaw

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    Craniodental Specializations Generalized mammals: Primitive mode

    Molars triangular

    Pointed individual cusps

    E.g in insectivorous & opossum

    Anteaters Most elongated jaws

    Progressively reduced teeth Highly elongated tongue

    Enlarged salivary glands

    Teeth reduction in nectar sucking mammals

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    Craniodental SpecializationsAquatic feeders

    Highly elongated jaws

    Anterior-most teeth lost (dolphins,porpoises)

    Teeth single cusped, pointed and increased

    in number

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    Craniodental SpecializationsAquatic Feeders

    Baleen Whales

    Teeth replaced by baleen Sheets of fibrous hornlike epidermal tissue that

    extend from downward from the upper jaw

    Used for filter feeding

    Walruses Flat postcanine teeth

    For crushing shells of sea food

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    Craniodental Specializations:

    Carnivores vs Herbivores Jaw closing muscles

    Masseter

    Temporalis

    Pterygoideus

    Jaw opening muscles in therians

    Digastric

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    Craniodental Specializations:

    Carnivores vs Herbivores Carnivores

    Large temporalis

    muscles to allow aforceful bite tosubdue prey

    Herbivores

    Reduced size oftemporalis muscles

    Large size of masseterto create force requiredto grind large amountsof fibrous materials withback teeth and to allowside to side movementof jaws. Skull & teethmodified to grind toughresistant food in large

    quantities

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    Craniodental Specializations:

    Carnivores vs Herbivores Large coronoid process

    of the jaw for insertionof temporalis muscles

    Temporal fossa is large.(area from which thetemporalis originates)

    Presence of a

    postglenoid process toprevent dislocation ofjaw muscles

    Reduced size ofcoronoid process

    and temporal fossafor insertion oftemporalis

    Absence of

    postglenoid process

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    Craniodental Specializations:

    Carnivores vs Herbivores Large occipital

    region to reflect

    extensivemusculature linkinghead to neck. Idealfor resisting

    struggling prey

    Small occipitalregion except for

    pigs that root withtheir snouts

    Elongated snouts

    Diastema: gapbetween cheekteeth and incisors

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    Digestion in Herbivores Plant cell walls: cellulose, require cellulase

    enzymes which cannot be produced by any

    mammal Thus mammals unable to digest cellulose

    Microbes in gut: symbiotic microorganisms,produce enzymes that degrade cellulose andlignin into digestible nutrients

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    Digestion Two types of fermentative digestion

    Hindgut fermentation &

    Foregut fermentation

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    Monogastric Animals Hindgut fermentors

    Horses, elephants, wombarts, koalas, rabbits, rodents, otherperissodactyls

    Simple stomach Enlarged colon and cecum

    Chew food thoroughly to release cell contents

    Cell contents digested & absorbed in stomach and smallintestine

    Cellulose digested in the cecum and colon by microorganisms

    Products of fermentation are volatile fatty acids

    Most eat large quantities to get enough nutrients

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    Hindgut (Monogastric)

    Digestion Coprophagy

    Eating the first set of feces that are produced

    thereby recycling nutrients that would beotherwise be lost

    Characteristic of small monogastric animals

    such as rabbits and rodents Ferment food in cecum, but do not absorb

    much, thus eat the feces

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    Foregut (Ruminant)

    Fermentors E.g. cows and other ruminant artiodactyls

    Camels lack an omasum

    Forestomach: 3 chambers store & processfood Rumen :1st chamber

    Reticulum: 2nd chamber

    Omasum: 3rd

    Chamber Fourth chamber:

    abomasum: for digestion

    Figure 21.9

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    Foregut (Ruminant)

    Fermentors Food initially retained in the rumen and reticulum.

    Degraded by microorganisms

    Microorganisms breakdown cellulose

    Food regurgitated and re-chewed (cud)

    Food in small particles then passes to omasum andthen abomasum (true stomach)

    Digestion in abomasum similar to monogastricanimals

    Note: all cellulose is broken down before reachingsmall intestines

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    Advantages in foregut

    fermentation Absorption occurs in small intestine, thus

    absorb most of the energy from plantmaterials. Hindgut fermentors rely on cecum& large intestine for breakdown of celluloseand lignin. But absorption is not as efficientas in the small intestines, thus loose energy isfecal matter

    Microorganisms attack plant material beforereaching small intestines-which is anadvantage vs hindgut fermentors

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    Advantages in foregut

    fermentation Microorganisms are themselves a source of nutrients

    to the ruminant animals

    Microorganisms play a role in nitrogen cycling, since

    they can convert urea into microbial protein that canbe used by the animals. Thus, microbes make allessential amino acids required by the animal

    A ruminant animal can be more limited in its selectionfor plant species than a monogastric animal whichhas to eat a wide variety of plant spp to get its aminoacids

    Detoxify chemical compounds No such benefit for monogastrics

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    Disadvantages in foregut

    fermentation Foregut system is slow

    Movnt thru a cows gut takes 70-100

    hrs whereas thru a horse its 30-45 hrs

    Do not thrive well on fibrous diets sinceits takes time to finish the processing in

    rumen and reticulum (slows passagerate)

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    Specializations for Locomotion Scansorial

    Generalized form as seen in shrews and squirrels

    Limbs and back are flexed during locomotion See figure 21-10 a & b

    Larger animals move with a stiffer back andstraighter legs and gallop rather than bound

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    Cursorial Limb Morphology Cursorial means specialized for running

    Specializations include

    Elongated legs

    to maximize strides

    Long legs provide a long outlever arm for forthe major locomotor muscles such as triceps inforelimbs and gastrocnemius in hindlimb

    Enhance speed of motion

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    Cursorial Limb Morphology Only certain portions of the limb are

    elongated primarily the lower limb

    portions Radius & ulna in the forelimb

    Tibia and fibula in hind limb

    Humerus and femur and phalanges arenot elongated

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    Cursorial Limb Morphology Muscles concentrated to the proximal

    portions of the limb to reduce the mass in thelower limb No muscles below horses knee (wrist) joint or

    ankle (hock) joint

    Foot is light

    Long elastic tendon transmit force of musclecontraction from upper limb to the lowerlimb. Tendons are long to increase stretch &recoil

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    Cursorial Limb Morphology Number of digits reduced to decrease

    weight of foot, some lost completely

    while others are compressed See slides below on Artiodactyls and

    Perissodactyls

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    Terminology related to

    locomotion Plantigrade

    Type of locomotion in which the entire sole

    of the foot contacts the groundAs in humans and primates who have

    retained all the 5 digits

    These mammals called pentadactyls

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    Terminology related to

    locomotion Digitigrade

    Condition in which an animal walks on the ends ofits metacarpals and metatarsals; only the toescontact the ground in walking

    Their wrists and ankles are elevated and thethumb has been reduced or lost

    Run or walk faster than plantigrade animals, walkmore silently and more agile

    Common in rabbits, rodents and many carnivores

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    Terminology related to

    locomotion: Ungulates Unguligrade

    Type of locomotion in which only the tips of thedigits contact the ground

    These animals have reduced number of digits

    Possess either 4, 3, 2 or 1 and thus walk on tips ofremaining fingers & toes

    Weight of body is borne on hoofed whichrepresent modified claws that have becomehardened and thickened.

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    Terminology related to

    locomotion: Ungulates The metacarpals corresponding to the

    missing digits have been either reduced in

    size or lost and those that are remainingare elongated and often united, amodification that greatly strengthens thelower leg and foot

    Limbs of unguligrades are only capable offorward and backward motion, no twistingor rotation is capable

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    Ungulates Muscles activating the lower portion of the

    limbs are located closer to the body to lessen

    the weight of the limb each time it is raised. The appendicular muscles attach to the limb

    bones by long lightweight tendons

    Thus the limbs and feet of hoofed mammals

    which are long and light and only capable ofonly aft movements are highly specialized forrunning and/ maneuvering on rocky terrain

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    Two groups of ungulates1. Artiodactyls (even toed)

    Retained digits 3 & 4 as functional digits

    Digits 2 & 5 reduced or lost in others Digit 1 is lost in all

    Pigs & hippopotamus: 4 digits (3, 4, 2, 5)

    Camels, deer, elk, giraffes, antelopes,bisons, buffalo, cattle, gazelles, goats,sheep: 2 digits (3& 4) (digits 1, 2 & 5 lost)

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    Two groups of ungulates2.Perissodactyls (odd # of digits)

    Digit 3 retained as primary functional

    digit Bears all of the weight

    Digits 2 & 4 are reduced

    Digits 1 & 5 usually lost

    Horses, zebras, rhinoceros.

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    Fossorial Limb Morphology Limbs specialized for burrowing underground or

    digging

    Digging limbs maximize power at the expense of

    speed Short forearm with a long olecranon process (elbow)

    Retain all five digits, tipped with stout claws

    Large bone projections on limbs for attachment of

    strong muscles E.g large acromion on scapula for attachment of deltoid

    muscles

    Examples are: African golden mole; Australianmarsupial mole; ferrets

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    Semiaquatic mammals

    (Amphibious) Have paddle-like limbs, use limbs to swim (paraxial

    swimming) as we do ourselves.

    Denser fur; webbing between their toes

    Examples are: Platypus (monotreme)

    Marsupials (water opossum, yapok)

    Water shrews, desmans, river otter, beavers, muskrat andmink

    Hippopotamus

    Inhabit a variety of waterways and associatedwetlands

    Require both aquatic and shoreline habitats for

    feeding

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    Aquatic mammals Use undulations of the body for swimming

    (axial swimming) via dorso-ventral flexion

    Do not use lateral undulations

    Swimming enabled by flexion of the vertebralcolumn

    Have short paddle-like limbs

    Limbs have short proximal ends Have elongated phalanges

    Limbs used for breaking & steering

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    Aquatic mammals

    In summary, the front limbs of aquaticmammals are modified for life in the

    sea and superficially resemble themodified appendages of of sea turtlesand penguins. Appendages become

    flattened, short and stout and may havea greatly increased number ofphalanges.

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    Aquatic mammals: Examples

    Order: Cetacea Whales and dolphins

    Have lost hind limbs Short necks

    Forelimbs modified into paddles

    Order: Sirenia Dugongs & manatees:

    have lost hind limbs

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    Aquatic mammals: Examples

    Order Carnivora: Seals, sea lions, andwalruses

    Have large naked front flippers andreversible hind flippers that can be broughtunder the body for locomotion on land

    In hair seals (earless) front flippers are smaller

    than hind flippers, which are not reversible.Thus in these seals, hind flippers are notreversible.

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