CRANIATA: Animals with Skulls
The earliest ancestors of craniates were plate-skinned fishes known as
OSTRACODERMS. (from the Greek ostraco meaning "shell" and
derm meaning "skin"). This was a diverse array of armored fishes
that lacked a lower jaw (mandible) or paired fins (remember the
lancelets! Do they have paired fins? What is the advantage of paired
appendages?).
One group of these fish (OSTEOSTRACANS) evolved paired pectoral fins just
behind the head. A separate group (ANASPIDS) closely resembled modern-day lampreys.
The Devonian period saw the rise of the first jawed fishes, the PLACODERMS, all of which are now extinct. Another
group, the ACANTHODIANS which lived at the
same time as the placoderms, may have included the ancestors of all modern
bony fishes.
Craniates can be divided thusly:
- Hyperotreti (hagfishes)
- Vertebrata
- Hyperoartia (lampreys)
- Gnathostomata (jawed vertebrates)
Jawed vertebrates all show
similar ontogeny with respect to jaw formation, and so are believed to be monophyletic. The jaw bones develop
from portions of the cartilaginous arches that once formed the support of
the anterior gill slits.
The condition of being "jawless" is primitive, and is not a good diagnostic character to use for constructing phylogenies.
Let's Meet the Vertebrates
I. Hyperotreti, the Hagfishes
Let's meet the Gnathostomes
The extant Jawed Vertebrates are presently divided thusly...
I. Chondrichthyes (sharks, skates and rays)
II. Actinopterigii (ray-finned fishes)
III. Sarcopterigii (lobe-finned fishes and tetrapods)
Chondrichthyes - The Cartilaginous Fishes
Sharks...

Skates...

...and Rays
Chondrichthyan Features and Facts
The chondrichthyan skeleton is composed entirely of cartilage.
The cartilaginous skeleton of sharks and their relatives is believed to have
been secondarily derived, with a bony skeleton representing the primitive
vertebrate condition.
In most vertebrates, the skeleton is first cartilaginous during embryonic development, and then is
ossified (made bony) later. In cartilaginous
fishes, some major genetic event apparently interfered with that process,
resulting in a permanently cartilaginous skeleton.
all chondrichthyans are carnivorous/predatory (though skates, rays and some sharks may prey primarily on molluscs, crustaceans, and other invertebrates, and some
sharks feed on plankton)
many pelagic (free-swimming) sharks must remain in constant motion to
keep fresh water moving over their gills, or they will die from lack of
oxygen.
skin is covered with placoid scales (derived from the dermis, just
underneath the epidermis), which form the famous replaceable
teeth at the edges of the mouth.
Sharks hunt primarily via their keen sense of smell, and localize prey
with the lateral line system, which can detect movement in the water.
Chondrichthyans may be
- oviparous (laying eggs that hatch outside the
mother's body)
- ovoviviparous (brooding eggs that hatch within the
mother's body, and then releasing the young)
- viviparous (young develop
within a uterus inside the mother's body, and are nourished prior to birth
via a connection with the mother's bloodstream (placenta).
Actinopterygii - The Ray-finned Fishes
These are the most numerous of vertebrates, both in species (about 30,000) and
individuals.
Vast diversity in form, function and natural history
Fun Fish Facts
Skeleton composed primarily of bone
ectothermal poikilotherms
- poikilotherm - body temperature controlled by the environment
- homeotherm - body temperature metabolically controlled
- ectotherm - body heat obtained from the environment
- endotherm - body heat produced metabolically
Skin covered with dermal scales that protrude through the epidermis and shed readily.
Skin copiously supplied with mucous glands (science terminology note: mucous is an adjective. mucus is a noun.)
Fins may be either medial and unpaired, or lateral and paired

Primitively, fins have rays made of cartilage or bone
Toothy, terminal (i.e., at the end of the body) mouth
Gill arches covered by a bony operculum
Swim bladder, derived from the digestive tract, may or may not be open
to the pharynx, and provides buoyancy.
Heart is two chambered (one ventricle (caudal) and one atrium (cranial)
Brain differentiated, with small olfactory lobes and cerebrum; large
optic lobes and cerebellum (what does this tell you about what fish do
best, and what they might not be quite as good at?)
Most species dioecious, a few hermaphroditic; some change sex during
the lifespan, depending on environmental circumstances
Sarcopterygii - Lobe-Finned Fishes and Tetrapods
The lobe-finned fishes include such "living fossils" as the Coelacanths and the lungfishes. (More cool lungfish information here.)
These are the closest living relatives to the four-legged vertebrates, or tetrapods.
Tetrapods
These are the four-legged, primarily terrestrial vertebrates, derived from a specialized group of
shallow-water fishes that probably resembled extant lobe-finned fishes.
Tetrapods include the
- Amphibia (salamanders, frogs, caecilians)
- Reptiliomorpha (reptiles, birds, dinosaurs, mammals, etc.)
Amphibia
The amphibians made first landfall during the Devonian (400 mya). The
first tetrapod had come ashore, with these defining characteristics:
- four limbs connected to the trunk
- each limb ending in five digits
There are three main Orders:
Modern Caudates (salamanders and newts) most closely resemble the
ancestral amphibians, in that they retain a tail as well as the five-digits on each limb. But all three extant
orders of amphbians are highly derived in their own ways, with each
species showing amazing specializations in both morphology and behavior.
Fun Amphibian Facts
ectothermal poikilotherms
Primitively, larvae are aquatic, with gills and lateral lines
Skin is scaleless, and serves a major respiratory function: must remain moist
Skin is well supplied with glands, including poison glands. Some species are far more poisonous than others!
large mouth with very small teeth
Mostly insectivorous/carnivorous (they are sit-and wait/ambush predators)
Heart is three-chambered
dioecious
- internal fertilization in caecilians and salamanders
- external fertilization in most frog species
Amniota: An Egg to Beat them All
The main homology linking the amniotes together is the Amazing Amniotic
Egg
All amniotes share this character, though it is highly derived
in some groups (e.g., mammals, in which it has given rise to the placenta and other structures).
There is some argument as
to the proper classification of amniotes, but they are generally
divided into three main groups on the basis of skull morphology:
- Anapsida - No temporal opening behind the eye orbits
- Synapsida - Single temporal opening behind the eye orbits
- Diapsida - Two temporal openings behind the eye orbits
Today's living remnants of these three lineages:
- Anapsids - turtles (maybe)
- Synapsids - mammals
- Diapsids - tuataras, lizards, snakes, crocodilians, and birds
Of greatest note here is the reorganization of what we used to call "Class
Reptilia." This has traditionally included the animals that are now known
to be Anapsids (turtles) and Diapsids (dinosaurs, birds, crocodilians,
snakes, lizards, and tuataras), so "Reptilia" as it usually has been constructed is polyphyletic, and is now in the process of being reorganized.
Anapsida - Turtles and their relatives
- Anapsids are characterized by an external bony shell
consisting of a dorsal carapace and a
ventral plastron. The shell is
fused to the vertebrae and ribs, and is an integral part of the
skeleton.
- Turtles lack teeth, but have sharp cutting edges to the maxilla (upper
jaw) and mandible (lower jaw).
- Turtles may be marine, fresh water, or terrestrial.
- Here's a nice
gallery of a few of the many turtle species for you to enjoy at your leisure.
Diapsida -
Dinosaurs, Birds, Crocodilians, Tuataras, Snakes, and Lizards
Reptile Features and Fun
- derived from the epidermis (not the dermis, as in fish), diapsid skin is primitively tough, scaley and resistant to desiccation
- dermal chromatophores (pigment-bearing cells) give reptiles skin its many distinctive colors and patterns
- jaws designed for gripping and crushing prey with large, powerful
jaw muscles
- internal fertilization (sperm and egg must fuse before the shell is
formed!)
- three- (all reptiles except crocs) or four-chambered (crocodilians and birds)
heart is more efficient than the three-chambered amphibian version. Higher blood
pressure means higher levels of activity.
- well-developed lungs allow complete independence from wet habitat
- many reptiles have evolved water conservation mechanisms, primarily
metabolic. (Recall the differences between the three types of nitrogenous waste. Which ones do reptiles make?)
- skeleton and limbs designed for terrestrial locomotion
- complex, highly developed nervous system with complex sense organs.
- visual system is at least trichromatic, and some reptiles equipped
with colored oil droplets that filter the color-sensing retinal cells may
perceive more colors than we primates do!
- Jacobson's organ - specialized olfactory organ on the roof of the mouth
(our vomeronasal organ is a vestige of this reptile
characteristic).
Birds are Reptiles!
Birds are perhaps the most derived of all Diapsids
Birds began as feathered reptiles derived from a group of small,
carnivorous dinosaurs known as therapods
The most famous of these is Archaeopteryx.
Feathers are homologous to scales/scutes of crocodilians.
Diagnostic Characteristics of Birds
For you to get all the details we discussed in class, it is ESSENTIAL that
you do your text readings and read at least the first page of any link above. Additional links you must read and know were mentioned in class, so be sure to note and read those, too.
VERTEBRATES: Evolution and Diversity
Our Friends, the Mammals
Synapsida:
The Mammals
- In mammals, the ancestral vertebrate jaw bones have become modified to form the bones of
the inner ear.
- Mammalian synapomorphies
- mammary glands
- fleshy lips for suckling (marsupials and placentals only)
- hair (grows between the analog of the reptilian scute/scale)
- There are about 4500 extant species in more than 14 orders.
Monotremes - Egg-Laying Mammals (Platypus and Echidna)
These are the only living mammals that lay eggs, which are reptilian in
structure and development.
Marsupials - Pouched Mammals
Eutherians - Placental Mammals
These are the mammals that give birth to live young that develop in the
mother's uterus, and are nourished during development via the
connection to the mother's bloodstream, the placenta. The placental
membranes are homologous to the amniotic egg membranes. YOUR
ASSIGNMENT: Find out which membrane (amnion, chorion and allantois)
has developed into WHAT in the mammalian system. (HINT: Review the
Amniotic Egg link at the UC Berkeley Museum of Paleontology)
The Synapsid vertebrates are probably the most familiar to all of us, as we happen to
be one of the many species of synapsids. What's so special about the synapsids?
They were the first vertebrates to radiate into a wide diversity of terrestrial
habitats
They diversified into herbivorous, carnivorous forms known as PELYCOSAURS
(they looked somewhat like lizards, but were NOT! Lizards are diapsids.)
You've probably seen pictures of one of the most famous pelycosaurs, Dimetrodon.
One group of carnivorous pelycosaurs gave rise to the THERAPSIDS, the
only synapsids to survive beyond the Paleozoic great extinction (about
245 mya).
Therapsids had limbs positioned directly beneath the body, allowing
for a more energy-efficient gait and movement (Think of the sideways limb
attachment of crocodilians and other reptiles for comparison.)
Most of the therapsids vanished during the great extinction at the
end of the Permian (245 mya)
The last surviving therapsids, the CYNODONTS, were the dominant
mammal ancestors in the Mesozoic
- high rate of metabolism
- jaw musculature increased in size and complexity
- bony plate (now seen as the hard and soft palates) separates the mouth and nasal cavities, allowing the
animal to hold food in its mouth and still breathe
- This inovation would also allow young to breathe while suckling,
a major mammalian inovation
The Importance of Teeth
The earliest true mammals showed up in the late Triassic, and were
mouse-sized carnivores.
They had diphyodont dentition, meaning they
were unable to constantly replace lost teeth (as their amniote ancestors
could), but rather replaced them only once over the span of a lifetime.
- deciduous ("baby") teeth
- permanent teeth
Humans and all other mammals show this pattern today.
Another dental inovation: differentiated
teeth (we are HETERODONTS, as
opposed to HOMODONTS as all other vertebrates
are.)
- incisors
- canines
- premolars
- molars
The number of each of these types of teeth varies with
species, and the
"dental formula" is one diagnostic character of mammal species.
In humans, the adult dental formula is 2:1:2:3/2:1:2:3 - This is the
number of teeth in half of the upper jaw (2 incisors, one canine, 2
premolars, and 3 molars) and in half of the lower jaw (two incisors,
1 canine, 2 premolars and 3 molars).
Note that a few mammals have secondarily reverted to the homodont condition, but this is derived with respect to other mammals.
Some mammals are hypsodonts. This means that their teeth grow continually throughout their lives, and must grind against each other to remain a normal, manageable shape. Some examples: rodents, lagomorphs (hares, rabbits, and pikas), horses, elephants and many other herbivores. What do you suppose is the advantage of having continually growing teeth? What could be some disadvantages?
What makes us mammals?
Hair (a derivative of the integument)
- undercoat - provides most insulation
- guard hairs - coarser and longer, this provides protection against
wear, and is usually the part of the pelage that has color patterns
- hair length is genetically determined, as we already have discussed
- hair usually sheds annually or twice a year
- specialized hairs called vibrissae ("whiskers") are used for touch
sensation
- Horns and antlers (another derivative of the integument)
- horns are sheaths of keratinized epidermis (keratin is the
protein that makes up hair) wrapped around a bone core rising from
the skull. They are permanent, and never shed. (Note: Rhino horn
isn't a true horn: it'sa swirl of hair-like filaments rising from the
dermis and firmly "cemented" together with glycoproteins to form the
structure, which is not attached to the skull.
- antlers are composed of solid bone at maturity, but are shed
annually.
Integumentary glands
- sweat glands
- eccrine - secrete water and salts, some organic compounds, trace amounts of urea and other waste products. Function: cooling
- apocrine - secrete a milky substance associated with reproduction
and emotion. The sweat from apocrine glands contains organic molecules (lipids, proteins, pheromones) that are broken down by resident bacteria, and produce a stong odor. Function: intraspecific (and possibly interspecific) communication
- scent glands - vary greatly in number, function, and location
across species
- sebaceous (oil) glands - function to keep skin and hair soft and
pliable with its secretion, sebum.
- mammary glands - the gland that gives our class its name,
evolved for feeding the young. Our fleshy, muscular lips have evolved
for the function of suckling.
MAMMAL NUTRITION
Mammals may be
- herbivores (including frugivores)
Such mammals often have a complex intestinal flora of specialized
microorganisms that can digest cellose (a complex plant starch), which is
not digestible by mammals.
Many herbivores (horses, rabbits, elephants, some primates), have a
cecum, a side pocket that branches off the junction between small and
large intestine. This contains a complex ecosystem of microorganisms
that provides essential nutrients via fermentation of the herbivore's
food.
Lagomorphs (hares and rabbits) and some rodents re-ingest their
cecotropes, strong-smelling pellets delivered from the cecum via the
anus. Your text calls this "coprophagy", which means "ingestion of
feces. This is not truly correct, as cecotropes are NOT feces. They are
an essential part of the lagomorph diet. Hence, your bunny engages, not
in coprophagy, but in cecotrophy.
Other herbivores (bovids (cow family) and ovids (sheep family)) are ruminants which have a large,
four-chambered stomach. Ruminants swallow food, which passes to the rumen,
where it is partially digested and regurgitated as cud. This is re-chewed
to break down fiber, and then sent back to the rumen for further
digestion.
insectivores - feed on insects
carnivores -feed primarily on herbivores
Let's talk about the "carnivore intelligence" myth....
omnivores - feed on a variety of plant and animal matter
MAMMAL REPRODUCTION
Egg-laying: Monotremes
Live birth of altricial young which are then raised in a pouch:
Marsupials
Internal brooding of young via a placenta - Eutherians (placental
mammals)
Most mammals have defined mating seasons, with females accepting male
advances only during her estrous cycle. When not in "heat"
(estrus), the female is unreceptive to male sexual advances.
- One estrus per breeding season: monoestrous
- Multiple estrus per breeding season: polyestrous
- Induced ovulators: ovulate upon the act of mating
Let's meet some interesting mammalian orders.
Family Ties: The Primates
The first primate was
- nocturnal
- small (about the size of a rat)
This lineage gave rise to the
- prosimians - lemurs,
tarsiers, lorises
- simians - monkeys and apes (including us)
(And some very derived
primates.)
And of course, of great interest to most of us are our own primate relationships.
The primate ancestors that gave rise to the simians were
- diurnal, with good trichromatic color vision
- arboreal
- social
How has our evolutionary history made us who we are today? The
earliest hominids (human-like apes) were not only social, but
cooperative, and evolved into the most primitive human social group,
the hunter-gatherer:
male - hunter of meat
female - gatherer of vegetable matter
What do these two types of foraging strategies require, and what
genetically-based behaviors might confer a selective advantage to
each sex?
Our Favorite Ape: Homo sapiens
Paleoanthropology is the study of human origins and evolution. Because
humans and chimpanzees have existed as separate species for only a few
million years, this branch of science examines only a very small, recent
portion of the fossil record.
Terms to know:
- anthropoid - of or pertaining to monkeys and apes
- hominoid - of or pertaining to the great apes (including humans)
- hominid - member of family Hominidae
Humans, apes, and monkeys all diverged from
a common anthropoid ancestor that shared all the characteristics common to all primates. Natural selection has driven the specialization of each primate species to be what it is today, and that includes our own species.

The earliest fossil members of our genus (Homo) range in age from
about 2.5 to 1.6 million years, and are currently classified as Homo
habilis:

Homo erectus shared common ancestry with H. habilis, and
shows up in the fossil record from about 1.8 million to 0.5 million years
ago:

Homo sapiens neanderthalensis , named for the Neander Valley in
Germany where its fossils were first found, may have arisen from an H.
erectus-like ancestor, as H. erectus is known to have migrated into both Europe
and Asia (timing uncertain).

There's a good deal of argument about how anatomically modern Homo sapiens
sapiens came onto the scene, but it's generally agreed that we
originated in Africa, where all the oldest hominid fossils are found.
- Multiregional Hypothesis:
Homo sapiens evolved in each of the
regions where its fossils are now found from ancestral Homo
erectus that migrated out of Africa about 1.5 million years ago.
Advocates of this hypothesis consider H. erectus to be an early
version of H. sapiens, and not a different species.
Constant interbreeding between neighboring populations of this "archaic"
Homo sapiens
may have prevented reproductive isolation, resulting in our
present-day races of Homo
sapiens,
rather than multiple species of Homo.
- "Out of Africa" Hypothesis:
All H. sapiens now living evolved from a second major migration out of
Africa that occurred about 100,000 years ago, and not from wandering Homo
erectus. These later migrants replaced the descendants of the
earlier H. erectus migrants.
So far, DNA analyses have supported the "Out of Africa" ("Replacement")
hypothesis. But as any hypothesis, this one is subject to further
testing as new analytic methods become available.

Here's a nice
overview of our hominid relatives, for those who would like to delve a
little deeper.
What makes humans different from the other great apes?
- Brain size
Hominoids of 6 million years ago had brains of about 400 -
450cm2. This is about the same as modern chimpanzees. Modern
humans' average brain volume is 1300cm2. This threefold
increase in volume is associated with cultural trends such as development
of complex language
- Jaw shape
Anthropoids and ancient hominoids have prognathic jaws: the
upper and lower jaws protrude beyond the nose. Recall that the human
face is paedomorphic with respect to that of other modern apes. The
face is flatter, the prognathous jaws lost. Changes in dentition
accompanied this change in jaw shape.
- Bipedal posture
Ancestral anthropoids and some of the earliest hominoids walked on all
fours, though--like modern apes--they could probably walk on their hind
legs with some degree of balance and skill. Humans are different from
all other apes in that the body posture is fully upright and
locomotion is entirely bipedal. A number of skeletal and muscular
modifications make this possible, but there is still a great deal of
academic argument about why humans became bipedal.
- Reduced sexually dimorphic size differences
In orangutans and gorillas, the male weighs about twice as much as the
female. In chimpanzees and Bonobos, males weigh about 1.4 times as much
as females. In humans, the difference is still less, with males
averaging 1.2 the body weight of females. (Why do you suppose this is the
case? HINT: social structure is a major selective factor here.)
- Key changes in family and other social structures
- Gibbons are social, with a dominant male defending a group of
females from other male rivals.
- Orangutans are solitary and do not form permanent social groups
- Gorillas are social, with a single "silverback" dominant male
getting all the mating opportunities with the females in his band.
Immature and subordinate males are allowed to stay in the group, but
do not get many mating opportunities.
- Chimpanzees are social and promiscuous. When a female comes into
estrus, all males will attempt to mate with her, and she will mate
with multiple males.
- Bonobos will have sex with anyone who holds still long enough.
- Hunter-gatherer human societies may be polygamous or
polyandrous. But in most human societies, monogamy is the norm.
(But is it biologically programmed?)
- Young are even more altricial than other great apes' young.
Newborn humans are exceptionally dependent on their mothers, and parental
care lasts longer after birth than in other ape species. This extended
period, coupled with the enlarged brain, enhances learning and is one
factor that contributes to the behavioral complexity of the human
animal.