INTRODUCTION
Why Study Birds
Scientific Inquiry
There are 10,700 species of birds
Very diverse forms (huge ostrich to tiny hummingbird)
Very diverse ecosystems (penguins swim underwater in cold, desert birds)
Fascinating behaviors (brood parasitism, migrations, breeding habits)
Economic Importance
Food (chickens, eggs/meat)
Hawk mountain
Models for research
Physiology - krebs cycle discovered studying pigeons
Easily observed and counted
Conservation
Used to be billions, now none of great awk
Health
Highly pathogenic avian influenza = bird flu
Snow geeses, canada geese
Lecture
Bird biology
Characters, evolutionary history, foraging
Flight, life history and reproduction, conservation
Evaluation
3 exams
3 assignments
Research article summary
Research paper
eBird training
Reading Assignment
pp 1-7, 11-18 of textbook
Lecture #1
What is a bird?
Feathers, Beak (no teeth), Claws, hollow bones, Keeled sternum, Wings, fly, 4 chambered heart, double circulation, Gizzard (Processes their food), Air sacs (in addition to lungs)
Flight
Feathers
Unique to (living) birds
Flight (gives them a stiff surface to allow them to fly)
Insulation
Mate attraction (males usually more colorful)
Bill or Beak
Turtles and platypus’ also have
Rhamphotheca - keratin based covering over bone
Stringing out food, stabbing fish, cracking seeds, tearing flesh
Why do they lack teeth?
They are heavy (gizzard is prolly heavier than teeth, turtles dont need to save weight and also dont have teeth)
Beak is proficient and adaptable for capturing prey
Teeth take too long to develop
chicks need to develop really fast in nest to avoid predators
Fused or Reduced Bones
Benefit
Increases strength, reduces the mass
“Hand” bones
Only 3 digits
Carpometacarpus = fused metacarpals (palm)
Hand supports primary feathers of the wing
Reduced fibula (tiny wishbone when eating drumstick)
Digitigrade - walk on their toes
“Foot” bones
Only 4 digits
Tarsometatarsus = fused metatarsals
All you see on bird is foot and toes
Synsacrum (pelvic bone)
Fusion of pelvic bones and vertebrae
Strong pelvis for bipedal stance and flight
Pygostyle
Fusion of caudal vertebrae
Support tail feathers and muscles operating it (short tails)
Pectoral Girdle
Keeled sternum
Very deep (deep/low projection, helps boats stay upright)
Furcula
Two clavicle fused together (our collarbone)
Wishbone
Flight muscle attachment
⅓ of body weight
OUTLINE
Bird Characters and Importance
Skeletal
Perching foot
Internal anatomy
Physiology
Origin and Evolution of Birds
Darwin
Fossil Intermediate
Which Group of Reptiles
Reading
pp. 4-7, 24-28
Assignment
pick article for slide, email underwoo with pdf
Review notes with flashacrds
Lecture #2
Features of Birds
Uncinate Processes*
Rearward projections on ribs
Strengthens thorax (keeps body rigid for flight, sleek line to be efficient when moving through the air)
Pneumatic Bones*
Hollow bones
Air sacs penetrate some
Internal struts called trabeculae (for support, m maintain strength)
Like internal anatomy of airplane structure
Perching Foot
3 toes forward, 1 toe back
Rear opposable toe called hallux, digit #1 (grips like thumb, allows them to perch)
Tendons posterior to ankle
Body mass drops down to stretch tendons
Maintains grip
Digestive Modifications
Gizzard
Muscular stomach
Grinding (they lack teeth, this processes their food)
Largest in seed eaters
Reproduction Modifications
Oviparous - they lay eggs, never give birth to live young
Don't have to carry around heavy developing fetus
Only left gonad fully developed (more dramatic in females than males)
Gonads regress after breeding
All save mass for flight
Hickman et al. 2007
Heart
Four chambers
> 40% larger than mammals of equal mass
Pumps oxygen and nutrients (glucose) efficiently
Maintains their high metabolism and activity levels
Body Temp and Metabolism
Endothermic - can regulate their body temperature through the production of metabolic heat (produce their own heat, don't need it from the sun)
Higher body temp for higher activity
Birds run much hotter than mammals, higher body temp >40*C (ours is ~36*C)
Increases their metabolism (when you add heat to a reaction it goes quicker), proteins break down at higher temps
Figure 6.1, p 142
Respiratory System (highly efficient)
Air sacs - most species have 9
Expand volume of system
Allow 1-way air flow through lung (unidirectional air flow)
Maximizes contact with fresh air in lungs
Replace almost all air in lung with each breath
Hickman t al 2015
Fig. 6.2, p 143
Lungs = faveolar
Bronchi branch into fine network of air spaces called air capillaries
Air capillaries surrounded by blood capillaries
Cross current flow for gas exchange
Blood flows in opposite direction to air in lungs
Fish have this too
Fig 6-4, p 144
Respiratory cycle
2 cycles of inspiration/expiration to move air through
Gas exchange happening on both inspiration and expiration
Most efficient of all terrestrial vertebrates
Why fly at high altitudes?
Bar headed goose has to fly over the himalayas at 30,000 feet (not enough oxygen for a human to survive up there), but they good
American birds fly at 5,000-10,000 feet
Origin and evolution of birds
1859 - darwins/s publication “On the Origin of Species”, highly controversial, lots of debate, critics really focused on birds and how they look nothing like other verebrates
Fossilized feather in germany 1 year later
1861 - Archaeopteryx lithographica (archaeo = ancient, pteryx = wing, type of slate they were extracting from that quarry), to date 12 species found, originated 150 billion years ago
Archaeopteryx traits
Bird Traits
Beak
Feathers (asymmetrical primaries, main feathers on hand of bird, like birds capable of flight)
3 digits on their hand
Furcula
Digitigrade (walk on their toes) and bipedal
3 toes forwards, one back
Some fused foot bones
Reptile Traits
Teeth
Lack a keeled sternum
Lack synsacrum
No uncinate processes
Long tail
Claws on the wing
No fused bones in the hand
Intermediate between birds and reptiles, what darwins skeptics were lacking
OUTLINE
Origin and Evolution of Birds
Which Group of Reptiles
Theropods vs Thecodonts
Evidence for each
Evolution of Feathers
Hypotheses
Evidence
Reading
pp. 24-45
Assignment
Quiz at beginning of lab (all areas of lab)
summarize article for slide
identify ducks
review notes with flashcards
Lecture #3
Archeopteryx Traits
Which group of reptiles did birds evolve from
Theropod dinosaurs (velociraptors)
Small, bipedal, predators
~180 million years ago
Thecodonts
Very small (lizard sized), arboreal (lived in trees), quadrupeds
Elongated scales on back (gliding?)
~230 million years ago
Huxley - 1867
Believed birds evolved from theropod dinosaurs
Shared similar characteristics in pelvis and hind limb
35 characters in common
Critics said due to convergent evolution (both bipedal)
unrelated species evolve the same structures because they do the same thing with it, common function (wings in birds, bats, butterflies, doesn't mean they are closely related)
Heilman - 1926
Believed birds evolved from theropod dinosaurs, share many features
But thought therapods lacked a furcula therefore must have evolved from thecodonts
Shortly after this, theropods with a clear furcula were found
Ostrom - 1970
Discovered deinonychus, a theropod
Found they share a lot of features with birds that no other reptile groups had
If Archaeopteryx wasn’t found with feathers, people would have thought it were a reptile
Gauthier - 1980s to present
Used cladistics (systematic technique)
More objective and robust than traditional systematics
Confirmed theropod orogin
~70 characteristics unite with birds
9 important we will learn
Bird Evolved From Theropod Dinosaurs
Shared traits as evidence
Hind foot
Loss of 5th toe
1st digit reversed (one points to the back), hallux
Pubic bones
Pubic boot (pubis expands)
Eventually disappears in modern birds
Faces backwards
Furcula
Pneumatic bones
Air sacs
Feathers
All major bird traits ^^^^^^^^^^^
Caudipteryx - covered in feathers (veined and DOWNY)
Trait Also from Theropod
Semi-lunate carpal bones (our wrists)
Allows wing to flex properly and move to allow folding of the wing
Flexion and side to side movement
Wing folding
Flight stroke
Hand bones
Loss of digits
5 and 4
Only 3 digits like modern living birds
Fused bones
Foot
Tarsus
Hand
Carpometacarpus
Tail
Birds evolved from theropods!!!!!!!!!!!
Thecodont Evidence
Feduccia - 1980s to 1990s
Birds and crocodilian ancestors share a few characters
Longisquama had feathers not scales
Argued flight evolved from arboreal ancestor
Issues
No definitive fossil link to thecodonts
80 million year gap between thecodonts and birds
No hard evidence really to support
Conclusion: birds evolved from therapods
Dinosaurs are not extinct, they evolved feathers and are birds
Who do we consider a bird
Aves - modern, living birds (when teeth go)
Avialae - theropod dinosaurs, tings not fully evolved with all traits, not really adopted yet, but suggested by Gill and Prum 2019
Evolution of Feathers
Functions
Flight
Insulation
Carry water (pretty unusual)
Waterproofing
Sensory
Stability / Support (woodpeckers use it to prop themselves up)
Capture sound (owls)
Aid in prey capture
Mate attraction
Hypotheses
Epidermal scales
Both made of beta keratin
Elongation, split,
Evidence
Some lizards (in hot desert environments) have elongated scales
Feathers = B keratin
Feathers likely first evolved for thermoregulation (either for reflecting or conserving heat)
ONLINE LECTURE
OUTLINE
Origin of Flight
Incline Running
Conclusions
Vision
Resolution
Binocular vision
Eye structure
Color vision
Reading
pp. 172-181
Assignment
Quiz at beginning of lab (all areas of lab)
identify beaks / feet
review notes with flashcards
Lecture #4
Newer Hypothesis (origin of flight)
Incline Running
Flapping protowings assisted climbing inclines
Flapping motion similar to flight stroke (early function of wings must be useful for something before fully developing)
Proto wings still functional
Experiment, trimmed 50% of wing feathers and they could still climb incline, if they could do that, they could climb trees and climb down (could lead to arboreal or cursorial flight)
Problem with idea
Therapods did not yet have some flapping ability (shoulder structure needed to flap wings like this)
Conclusion
Origin of flight not yet resolved
Avian Senses
Vision
Live in a world of color
Can see a rabbit from a mile away
Largest eyes for their body size
Visual acuity / resolution (separate out fine details)
Ability to distinguish between two points from a distance
Some birds 2-3x better than humans
Especially raptors
Binocular vision
3 dimensional with depth perception
Requires
Eyes must be able to focus on same object (have overlapping field of vision)
Examples
Evans and heiser 2004
Woodcocks have 360 field of mostly monocular vision, they have better vision behind them (to see predators approaching because they are always in the dirt
Pigeons bobble their head so they can look at the same object from many different dimensions
American bittern throw their head up when they get startled, their binocular vision is downward cuz they catch fish, so they do that to aim binocular vision at predator, they also do it to add to their camouflage (improves vertical camouflage)
Eye structures - external
Most transparent covering = nictitating membrane
Moistens their eyes without losing their vision
Underwater mask, important for diving birds who are underwater
Eye anatomy - internal
Light passes through cornea and lens to retina
Retina contains photoreceptor cells (rods and cones)
Both cornea and lens change shape to focus image on retina (in mammals only the lens changes shape)
Sclerotic rung - ring of bones around their eyes to support all the internal eye muscles (lacking in mammals)
Pecten
Unique to birds
Highly vascularized (lots of blood vessels)
Supplies oxygen and nutrients to retina (retina lacks a direct blood supply)
Iris
Function - act like a diaphragm on microscope to adjust the opening of how much light comes in to adjust the size of the pupil
Pupil contracts in bright light and dilates in dim lights
Color function - mate attraction and species recognition
Retina cross section
Photoreceptors
Rods detect black and white
Found in nocturnal birds
Cones detect color
Diurnal birds
Most diurnal birds have tremendous amounts of cones (up to 1 million/mm2), the more cones the better the resolution
Fovea - depression in retina with high densities of cones, high resolution, usually 1 per eye = central fovea, allows birds to detect really fast movements (flying through environment, capturing prey, need to be able to really pick up on these movements)
2nd Fovea in diurnal raptors
Temporal fovea, gives them really good forward binocular vision
Pecten benefit
Allows for a high density of cones (not possible in mammals)
No need for blood vessels in retina that would interfere with vision
Tapetim lucidum
Reflective layer at the back of eye
In low light conditions it bounces some of that light back through the eye to help capture more light (light back → retina), found in nocturnal birds (owls)
Eyes light up when you shine light on them
OUTLINE
Origin of Flight
Lecture #5
MISSED - get notes from jake
SIght
Humans - trichromatic vision
Birds - tetrachromatic vision
Birds are capable of seeing some of the UV light (another dimension that we camt)
Birds see 300-400nm (near UV)
Functions of UV vision
Orientation (perceive environment, navigate themselves)
Foraging (some berries reflect UV light differently than foliage around them, makes them stand out, voles pee reflects UV light, kestrel can pick up on where they are most active)
Mate selection
Mate selection in Zebra finch
Measured UV reflectance in males
Compared female choice between males with and without UV, used glass filters between males and females to prevent them from seeing
Results - strong evidence that females significantly preferred the males with UV cues
Monomorphic Species
European starling (got here because people love shakespeare)
Monomorphic plumage = one plumage type, males and female look the same
Many of these birds are actually dimorphic when UV is measured
Avian Hearing
Some birds have great ability (owls, under 2 feet of snow) but most birds hear worse than us
External ear
Just a hole in the head
Feathers cover ear = auriculars
Keep wind / bugs out of the ear
Lets sound through
Owls - have an operculum
Allows them to adjust the opening
Middle and inner ear
Middle
Tympanic membrane
Columella (1 bone) - transmits the vibrations
Humans have 3 bones
Inner
Cochlea - large fluid filled chamber, lined with hair cells, hair cells convert vibrations to nerve impulses
Semicircular canals - balance and equilibrium (important for animals with high activity levels), fluid with calcium carbonate crystals, hair cells at base, stimulated by movement
On roller coasters you get wobbly after
Hearing ability
Frequency and sensitivity, vast majority of birds can't hear faint noises (except for strigiformes / owls who can hear better than us)
Directional hearing in horizontal plane
Sound hits right or left ear first, we turn head to hear where its coming from, birds do too
Birds also hear vertically too
In barn owls
Localize sound in vertical plane too
Ears and facial disk asymmetrical (one set is higher than the other)
Facial disk oriented to detect
Sounds above - right
Sounds below - left
Ear tufts in owls
Communication and camouflage, but not for hearing
Echolocation
2 families of cave nesting birds
Oilbirds and swiftlets
Feed on the nuts of oil palm trees, natives use them as a source of oil
15-20 millisecond clicks (not ultrasound, not the same as bats)
1-5khz, ~10% as effective as bat echolocation, allows them to move around in the dark, but not find tiny insects
Mechanoreception
Herbst corpuscle - tactile / touch receptors, layers of cells that surround a nerve or sensory cell, detect physical pressure changes put on that cell
Numerous in certain areas (bills of ducks and shorebirds, probe around in mud and water to find food, base of sensory feathers)
LAB Lecture - Feathers
Quill = calamus
Modifications to Shapes
Club-winged Manakin - creates sound with wings (vibrates wings really fast, twice as fast as hummingbirds), thick rachis allows them to vibrate
Birds-of-paradise - have biofluorescent feathers (reflects light back at a higher wavelength)
Narrow modification = makes sounds
No such thig as sea gulls, shorebirds tho
Shorebirds
Sandpipers
Scolopacidae
Stand on one leg
Long, thin beaks
More modeled with browns/reds, variations
Quite hard to differentiate
Between
Semipalmated sandpiper - semipalmated webbed feet (hard to see), look at leg color, yellow blackish greenish, go off shape and thickness of beak, stouter
Least sandpiper - little tony curve, more of a point, narrows to the point
Between
Lesser Yellowlegs - smaller, fairly proportionate beak, beak points up
Greater Yellowlegs - bigger, very long beak compared to the width of its head, more barring on side of body
Between
Wilson’s Snipe - vertical stripes on top of head, more streak colored breast
American Woodcock - horizontal bars of dark color on top of head, buffy colored more solid colored breast
Plovers
Charadriidae
More uniform color patterns
OUTLINE
Baroreceptor
Smell and Taste
Structures and examples
Bird Brains
Basic structure and variation
Spatial memory
Sleep
Assignment
Page 183-203
Lecture #6
Baroreception
Detect changes in atmospheric pressure
Paratympanic organ, middle ear
10m difference in altitude
Important for
Detect approaching storms
Find right altitude for migratory flights
Sense of Smell
Olfactory bulb - process scent in brain
Birds have poor sense of smell (great horned owls eat skunks)
Relative size of olfactory lobe - highest is 33% of cerebrum to 5% of cerebrum
Nocturnal predators have best sense of smell
Nasal cavities - olfactory receptors on mucosal membranes covering their nasal turbinal bones
Smell and Foraging
Black vultures detect carrion (rotting flesh) releasing ethyl mercaptan
Kiwi are nocturnal bird preying on soil invertebrates, has a great sense of smell to find food, rely entirely on scent to find their food in the dark in the soil
ONLY BIRD to have nostrils at the tip of their bill
Mallard courtship, scent required for mating, odors from uropygial gland, secretes oil for conditioning feathers and waterproofing, some species also produce pheromones (like in mallard)
Crested auklets sniff each others necks, smell good like oranges, pheromones smell like organs, in courtship they sniff each other
Taste
May not be well developed (don't have as many taste buds s mammals), taste buds located on rear of tongue and walls of oral cavity
Chilli peppers have capsaicins that repel (most) mammals, but have no effect on birds
Why do plants deter mammals with capsacians and not birds = birds are much better at dispersing seeds over a large distance
Brain and Intelligence
Display insight learning aka problem solving (crows and parrots are great at this)
Crows in japan crack walnuts by dropping them on roads to have cars crack them open, and that when the light is red cars wont drive by
Social learning aka learning by observing others
Chickadees (or tits in europe) learned how to break into milk bottle and drink cream on top, then they all did ot
Tool use
Cactus finch from galapagos, uses cactus spine to probe into trees and get bugs to come out
Avian Brain
Forebrain
Olfactory bulb - processes scent
Cerebrum - center for learning, sensory processing, and integration of information, problem solving
Pallial neurons - nerve cells for advanced learning (ONLY in birds and mammals)
Midbrain
Optic and auditory lobes - vision processing and hearing
Cerebellum - balance and equilibrium (works with semicircular canals of inner ears), high activity level = big cerebellum
Hindbrain
Medulla oblongata - controls autonomic functions (heart rate/breathing)
Variants in bird brains
Modified to reflect something about their lifestyle / foraging abilities
Flamingoes gave relatively small cerebellum and optic lobe - filter feeders, dont fly much, pump water through their beak
OUTLINE
Brain and Intelligence
Structural variation
Spatial memory
Sleep
Foraging behavior
Optimal foraging theory
Where to forage
How long to stay
Reading
pp. 190-202
Assignment
Quiz at beginning of lab (all areas of lab)
identify taxonomic
Finish lab packet
review notes with flashcards
Spinal feather tract
Apterya feather tracts
Lecture #7
Variations in Brains
Shearwater has hygge olfactory bulb = very good at scent, use it to find plankton that they feed on and their nest burrows in the dark
Crow has largest cerebral hemisphere = great at problem solving, intelligence well developed
Avian Brain vs Primate Brain
Birds have higher number of pallial neurons (indicator of intelligence)
Birds have smaller brain in mass, but more of their brain devoted to intelligence
Birds have more forebrain and pallial neurons than similar sized primates
Spatial Memory
Seed catchers
Store food caches for winter (in trees or ground), can remember for up to 8 months later, 2000-3000 seed caches
Titmice, chickadees, nuthatches, corvids, jays in pa do this
More common in places that get cold over winter
Test
Remember where things are in relation to things in the environment
Use spatial reference to large objects
Move objects around, birds would search in that direction of the big rock they moved
Controlled by hippocampus of cerebrum
Hippocampus is very large in seed caching birds (larger than those who do not rely on memory to find food)
Shows neurogenesis (production of new neurons)
Add new brain cells
Chickadees hippocampus expands ~30% in fall, shrinks back in spring
Same thing for male birds who sing more versions of songs, grows in summer and shrinks outside breeding season
Sleep in Birds
Short bouts (minutes, seconds)
Same basic patterns of sleep as humans
Rapid Eye Movement (REM)
Dreaming and restore brain function
Slow Wave Sleep (SWS)
Deep sleep
Restores body and brain function (physical and mental health)
Birds can sleep while flying!!
Unihemispheric short wave sleep
Sleep with one eye open
Half of brain is functioning at once
Predator vigilance hypothesis
Studied flocking birds (mallards), birds at edges of the flock use unihemispheric sleep more than birds at the center, strong support
At the edge of the flock 90% were looking away from the flock
Foraging Behavior
When it gets colder birds are feeding much more intensely
Work outside in the winter make you more hungry than working inside
Foraging Decisions
Where to forage
Patch of habitat to go to
How long to stay
What to eat
How to search
Optimality Theory
Our profit from eating something, making a certain decision = (our energy gained - energy cost to obtain that) / foraging time
Costs and benefits to any behavior
Maximize benefits and minimize costs
Natural selection favors individuals (for reproduction) that “behave” optimally (make the smartest decision)
Bird with calculator counting calories\
WHere to forage
Prediction - most profitable feeding areas
Evidence
Great tits in captivity, made 6 artificial patches of habitat that differed in prey density, looked at percent of time spent searching in each of those patches
Over time, the birds to forage in the most profitable areas, to start there isn't much difference but as they learn their surroundings, its obvious, but still visited low density patches
Prey abundance varies in time and space in the nature, not believing high density patches ALWAYS have food and low density NEEVER had food, making sure it didn't change
Do forage in most profitable areas, but keep their options open
How long to stay
Capture rate declines over time because prey supply is depleted (eat them all) or prey take evasive action (try to hide)
Exponential like growth at start, then decreases
Leave when capture rate drops below average for other available patches
LAB Lecture
Field Trips
Look for movement, listen for sounds, scan for irregular shapes
Describe where birds are in reference to landmarks, clock to describe position
Outline
Foraging Behavior
How long to stay
What to eat
How to search
Learning
Foraging Niches
Bill Structure
Variation
Cranial kinesis
Reading Assignment
452-453, 538-540, 12-15
Lecture #9
Diminishing Returns
Pied wagtails moved on when the capture rate really drops
What to Eat (Prey Selection)
Prediction - max energy gain and minimum foraging time (optimal foraging)
Handling time - deal with prey, break into a seed, crack open the shell, subdue and swallow an animal
Most profitable is a fly 7mm long, black bars = flies available, white bars = flies captured
8mm flies are most abundant, but ate the 7mm flies the most
Exception to the rule
Not always optimal
8 finches and sparrows - seed preference
Did not eat optimal seeds
Chose those with less handling time
Trade off to mitigate predation risk
How to Search for Prey
Much like we do, someone loses something, we ask what it looks like, blue color, try to pick out where it is in the environment
Set of prey characteristics that act as filter to remove unimportant stimuli and focus on the key item = search image
Search image testing on blue jays
Prediction - if they're using a search image, then their foraging success will improve over time
With 1 species of moth, improve over time, supports prediction
2 species moth photos mixed - no improvement, cant form a search image because prey keeps changing
What influences foraging success
Weather
Stealth ability / camouflage of birds
Compietioton
Bird size
Age / experience
Influence of time / age on learning
American oystercatcher
Specialize on worms or molluscs
Worm feeders - 7 weeks
Worms are easier to catch
Mollusc feeders - up to 1 year training
Sneak up on clam and jab beak in and snip muscles
Foraging niches
Niche - sum total of an organism's resource use (abiotic and biotic factors) and general ecological requirements
Principle of competitive exclusion - if two species share the same ecological niche, one or both species will go extinct or one or both will develop a new niche
Boreal forest warblers suggested to be an exception to this principle (all seemed to forage in confierous tree)
They all forage in different PARTS of the tree, slight overlap but for the most part foraged in different parts of tree and had slightly different foraging styles
And nesting periods were slightly offset as well, peak food needs were not at the same time
Evolved niche segregation
Feeding - Bill Structure
Natural selection doesnt just work to improve feeding, still has to be functional for other things as well
Other uses of bill other than capturing prey
Defense
Preen
Build nests
White winged cross bills - use the cross to lever open pinecone seeds