Dr. John W. Parrish, Jr., Professor of Biology; B.S., Denison University, 1963; M.S., Bowling Green State University, 1970; Ph.D., Bowling Green State University, 1974; Postdoctoral Experience: The Reproductive Biology Institute, Department of Zoology, The University of Texas at Austin, 1974-1976; Department of Physiology, School of Veterinary Medicine, Cornell University, 1986.

CAMPUS PHONE (912)681-5487; PHONE MAIL (912)681-0053, FAX (912)681-0845   E-MAIL - jparrish@.georgiasouthern.edu: John Parrish

ADDRESS: Dr. John W. Parrish, Jr., Department of Biology, P.O. Box 8042, Georgia Southern University, Statesboro, GA 30460-8042

• Courses Taught
• Research Interests
• Graduate Students
• Ornithology
• E-Mail
• Links to other HTML sources
• Bird Field Guide
  
  
  
  
  
  
  
  

Courses Taught:

• Graduate Courses at Georgia Southern University:
  • Graduate Seminar (various topics)
  • Advanced Ornithology
  • Raptor Biology Projects
  • Comparative Animal Physiology
  • Endocrinology
  • Wildlife Management
• Undergraduate Courses at Georgia Southern:
  • Ornithology
  • Senior Seminar (various topics)
  • Environmental Biology
    

Research Interests:

Environmental Animal Physiological Ecology, Raptor Biology and Ecology, Avian Physiology and Ecology, and Wildlife Biology (Areas of current interest: Nesting Biology and Genetics of Southeastern American Kestrels, Ultraviolet Vision in Birds, Bioenergetics of Birds, and Pineal Gland Biology in Animals)

RESEARCH PUBLICATIONS:

1. Parrish, John W.  2006.  A Bullock’s Oriole and Baltimore Orioles in the winter 2005-2006,            in Bulloch County, Georgia.  Oriole (in press).
2. Heffron, Michael B. and John W. Parrish. 2005.  Apparent metabolizable energy of feeds in greater prairie-chickens.  Transactions, Kansas Academy of Science 108:93-98.
3. Parrish, John W.  2003.  Late wintering Baltimore Orioles in pecan orchards in Bulloch County, Georgia.  Oriole 68:14-15.
4. Snow, Frankie, and John W. Parrish.  2002.  Significant population of Southeastern American Kestrels in South-Central Georgia.  Oriole 67 (3,4):50-52.
5. Parrish, John W.  2002.  Early wintering Baltimore Orioles in pecan orchards in Bulloch County, Georgia.  Oriole 67(3,4):52-53.
6. Parrish, John W.  2000.  Possible prevention of European Starling nesting by Southeastern American Kestrels at a power substation in southern Georgia.  J. Raptor Research 34:152.  

7. Parrish, John W.  2000.  An effective one-handed band removal tool.  North American Bird Bander 23:81-82.

8. Breen, T. F. and J. W. Parrish. 1997. American Kestrel Distribution and use of nest boxes in the Coastal Plains of Georgia. Florida Field Naturalist 25:128-137.

9. Breen, T. F. and J. W. Parrish. 1996. First evidence for double-brooding in Southeastern American Kestrels in Georgia. Oriole 60:81-83.
10. Breen, T. F. and J. W. Parrish. 1996. Eastern Screech-Owl hatches an American Kestrel. J. Field Ornithol. 67:612-613.
11. Breen, T. F., J. W. Parrish, K. Boyd and B. Winn. 1995. Southeastern American Kestrel Nests in Bulloch, Evans and Columbia Counties. Oriole 60:33-36.
12. Parrish, J. W. 1995. Second Ross' Goose record for Georgia. Oriole 59:1-2.
13. Parrish, J. W. 1991. American Goldfinch and House Finch nesting in Bulloch County, Georgia. Oriole 56:50-51.
14. Parrish, J. W., P. S. Parrish, W. A. Parrish and C. D. Parrish. 1990. Wintering hawks along a Kansas roadside. Bulletin, Kansas Ornithol. Soc. 41:25-26.
15. Parrish, J. W. and D. K. Saunders. 1989. Simplified cage modification to reduce self-inflicted injury in confined birds. Wildlife Soc. Bulletin 17:80-81.
16. Saunders, David K. and John W. Parrish. 1987. Assimilated energy of seeds consumed by Scaled Quail. Journal of Wildlife Management 51 (4):787-790.
17. Parrish, John W. and Gina M. Wendt. 1987. Biliverdin and bilirubin in the serum of male Rock Doves. Bulletin, Kansas Ornithological Society 38(3):30-32.
18. Nash, Susan M., Brenda J. Anderson, Teresa L. Reed, John W. Parrish and Stephen F. Davis. 1986. Reward and nonreward odor cues: The role of the harderian gland. Bulletin of the Psychonomic Society 24(2):141-144.
19. Parrish, John W. and David Workman. 1984. Hematological characteristics of the Rock Dove. Bulletin, Kansas Ornithological Society 35(3):33-36.
20. Parrish, John W., James A. Ptacek and Kevin L. Will. 1984. Near-ultraviolet light detection in nonmigratory and migratory birds. Auk 101:53-58.
21. Parrish, John W. and Michelle L. Mote. l984. Chemical levels in the serum of captive female House Sparrows. Wilson Bulletin:96:138-143.
22. Parrish, John, Roger Benjamin and Ricky Smith. 1981. Near-ultraviolet light reception in the Mallard. Auk 98:627-628.
23. Andre, Joel S. and John Parrish. 1979. Inhibition of estrous cyclicity in golden hamsters by melatonin on the day of proestrus. J. Exp. Zool. 207:161-165.
24. Parrish, John W., Jr. and Elden W. Martin. 1977. The effect of dietary lysine level on the energy and nitrogen balance of the Dark-eyed Junco. Condor 79:24-30.

REARCH GRANTS

1. 2004-2006, Georgia Department of Natural Resources, Conservation of Southeastern American Kestrels, $104,980
2. 2003-2004, Georgia Power & Avian Powerline Interaction Committee, Reproduction of Southeastern American Kestrels in south-central Georgia, $14,000
3. 1999-2007, Arcadia Wildlife Preserve, Inc., Southeastern American Kestrel DNA Study, $5,000
4. 1998, Arcadia Wildlife Preserve, Inc., Southeastern American Kestrel Nestbox Study, $6,500
5. 1997, Arcadia Wildlife Preserve, Inc., Southeastern American Kestrel Nestbox Study, $6,500
6. 1996, Arcadia Wildlife Preserve, Inc., Southeastern American Kestrel Nestbox Study, $6,500
7. 1995, Arcadia Wildlife Preserve, Inc., American Kestrel Nestbox Study, $9,000
8. 1995, Georgia Department of Natural Resources, Southeastern American Kestrels in Georgia, $2,000
9. 1994, Georgia Department of Natural Resources, American Kestrels in Georgia, $2125
10. 1994, Arcadia Wildlife Preserve, Inc., American Kestrel Nestbox Project, $5,600
11. 1988, Quail Unlimited, Energetics of Northern Bobwhite and Ring- necked Pheasant chicks, $2000
12. 1988, Faculty Research Committee, ESU, Energetics of Northern Bobwhite Chicks, $4400.
13. 1987, Kansas Department of Wildlife & Parks, Energetics of the Greater Prairie-Chicken, $1500
14. 1987, Faculty Research Committee, ESU, Energetics of the Greater Prairie-Chicken, $1500.
15. 1985, Kansas Fish & Game Commission/Quail Unlimited, Bioenergetics of the Scaled Quail, $1010.
16. 1985, Faculty Research Committee, ESU, Bioenergetics of the Scaled Quail, $5,087.
17. 1984, National Science Foundation, Research equipment grant (co- principal investigator), $25,949.
18. 1984, Faculty Research Committee, ESU, The utility of near- ultraviolet vision to birds, $4,300.
19. 1983, Faculty Research Committee, ESU, The detection of near-ultraviolet light in nocturnal and aquatic mammals, $3,900.
20. 1981, Faculty Research Committee, ESU, The efficacy of near- ultraviolet light reception in birds, $4,196.
21. 1980, Faculty Research Committee, ESU, Near-ultraviolet light reception in birds, $3,000.
22. 1979, Faculty Research Committee, ESU, Endocrine role of the pineal gland, $2,970.
23. 1977, Faculty Research Committee, ESU, Effects of melatonin on reproductive function in golden hamsters, $1,485.
24. 1968, Josselyn Van Tyne Memorial Fund, American Ornithologists' Union, The effects of dietary lysine level on the energy and nitrogen balance of the Dark-eyed Junco., $200.
    
    

• Graduate Students
  
•  
  • Georgia Southern University:

1. 2007  M.S.  Beasley, Hope A.  -  Effects of High EMF on the Reproductive Success of Southeastern American Kestrels in South-Central Georgia (in progress) 
2. 2006  M.S.  Maney, Pamela L.  -  Southeastern American Kestrel Nesting in South-CentralGeorgia
3. 2003  M.S.  Lovell, Helen W. -  Population Increase and Range Expansion in Three Psittacids in Florida and California   
4. 2000  M.S.  Jolliffe, Alisha L. - Transfer Patterns among Free-ranging Infant Ring-tailed Lemurs
5. 1997 M.S. Shuford, Roy - Nesting biology of Southeastern American Kestrels below the Fall Line in Georgia
6. 1997 M.S. Halas, Cynthia - Ecology of American Kestrels in Southern Georgia (In Progress)
7. 1996 M.S. Turfler, Stacy - Effect of fire on neotropical migrants at Fort Stewart Military Base (Co-Advisor)
8. 1996 M.S. Goss, Jana - Neotropical migrants at Fort Stewart Military Base (Co-advisor)
9. 1995 M.S.Breen, Timothy - The use of nest boxes by American Kestrels (Falco sparverius paulus)below the Fall Line in Georgia
10. 1993 M.S. Cooler, Stephania - Morphology of near ultraviolet reflecting wings scales in southeast Asian butterflies
    • Emporia State University (Kansas):
11. 1989 M.S. Heffron, Michael B. - Bioenergetics of Wintering Greater Prairie-Chickens
12. 1986 M.S. Saunders, David K. - Bioenergetics of Wintering Scaled Quail.
13. 1986 M.S. Ptacek, James A. - Avian Community Structure in Kansas Riparian Forests.
14. 1986 M.S. Hamilton, Robert G. - Winter Avian Use of Fragmented Woodlands.
15. 1985 M.S. Lesher, Mark K. - Energy metabolism in doves and sparrows.
16. 1982 M.S. Pritchard, Donna M. - Genetic variance in the Prairie Deer Mouse.
17. 1978 M.S. Renner, Kenneth L. - The effects of melatonin and l-isoproterenol on estrous cyclicity in the Syrian hamster.
18. 1977 M.S. Ezeocha, Dickson O. - The role of temperature in determining the inhibitory activities of Ambrosia artemisiifolia, Ambrosia trifida, Bromus inermus and Helianthus annus on Spring and Winter wheat. (Co-advisor)
19. 1977 M.S. Villamil, Carmen R. - The effects of dietary protein level on plasma urea and transaminase activities in uremic rats.

• Ornithology
• Ornithology Lecture Notes (2002)
  
• I. Short History
  A. Birds are ubiquitous and conspicuous everywhere
  1. Snowy Owls - Arctic Circle
  2. Sand Grouse - Middle east deserts
  3. Finches - "Andean Mts
  4. Penguins- hundreds of meters below Antarctic seas
  5. Eagles & Parrots -rain forests
  6. Buzzards, plovers and sparrows - across arid plains
  B. Highly mobile creatures
  1. nicobar pigeons - move continuously among Indonesian islands
  2. Sooty shearwaters - migrate from coast of master
  Australia to CA & OR coasts
  3. Arctic Tern - migrate from New England to Arctic
  4, Rufous hummingbirds - from Alaska --> Mexico & often GA
  C. Highly attractive
  1. Bluebirds --> Painted Buntings
  D. Birds and Man
  1. Evidence that jungle fowl domesticated 3000 BC in India
  2. " Mallards & geese " 1000 BC in Far East
  3. Romans "raised" poultry/practice disappeared after "Fall of Roman Empire"
  a. Reappeared in 19th Century Europe
  4. Falconry- recent upsurge following pesticide-induced down in Penguins
  5. Feathers as ornaments spread worldwide (birds now protected)
  6. Birds in music
  a. Operas- Robin sings in Madam Butterfly Swallows sing in La Boheme
  b. America Pop- Mocking bird Hill, Rockin' Robin, Bye Bye Birdie
  7. Literature- Bible to Shakespeare to Audubon's Birds of America
  (1827-30) Ornithology of Shakespeare- 1871
  E. Early History of Ornithology
  1. 4th Century BC- Aristotle's History of Animals
  a. Land birds, water birds, etc.
  2. 1st Century AD- Alexander of Myndos
  a. 3 Volume work on animals- 2 volume for birds (Few fragments survive today)
  3. 1st Century AD- Pliny the elder- 37 Volume Natural History Encyclopedia
  4. Much early writing good, but some not believable
  a. Melian (170-235 AD) wrote that:
  1. Cranes only flew against wind and
  2. Cranes swallowed a stone for ballast
  3.Cranes sentinels held stone in beak at night stood on one foot
  a. If sentinel fell asleep, drop stone, wake up
  5. Many field observations began in 18th century
  F. Ornithology Today
  1. Nikko Tinberger (gulls) and Konrad Lorenz (ducks and geese)
  a.Studies on behavior earned them Nobel Prize and lead to
  beginning of modern Ethology
  2.Albert Szent-Gyorgyi won Nobel Prize for elucidation of Krebs
  Cycle in pigeon breast muscle
  3.Payton Rous- Noble Prize Winner for studies of avian sarcoma
  that linked viruses to cancer for 1st time
  4.Neotropical migrants forewarn us of global deforestration and effects
  ORIGINS
  I. Intro
  A. Millions of years ago a small bipedal reptile lived among dinosaurs
  1. Over time scales-> evolved into soft feathers
  a.Feathers provided insulation to permit control of high Tb which allowed greater activity & endurance1. Also permitted mobility- gliding & eventually flight
  2. A new class- Aves was Born
  II. Basic Characteristics
  A. Birds are bipedal vertebrates
  1. Possess uniquely--Feathers!
  a. Essential for both body temp. regulation & flight
  2. All birds have bills
  a. Modern birds- it is toothless & covered with Horny Sheath
  3. Flight also in INSECTS & BATS (Gliding in some Squirrels)
  4. Bones unique
  a. Reduced and fused
  b. Hollow and Strutted for strength
  c. Uncinate processes on ribs strengthen thorax
  d. Furcula (wishbone)- compresses and rebounds like spring during flight
  e. Arboreal birds most abundant
  1.Tendons cause automatic locking of claws to branch
  2. Passeriforms (Perching birds)
  a. Special ratcheting mechanism- permit perching birds to sleep on branches
  1.Large opposable rear toe (Hallux) further enhances perching- unique in vertebrates
  5. Avian Physiology
  a. Large flight muscles- extraordinary work capacity
  1. Also great for shivering thermogenesis
  b. 1st complete 4-chambered heart in birds
  c. Flow-through lungs &air sac arrangement unique in birds
  d. Reproduction
  1. Most elaborate external eggs of all animals
  a. No bird species bears live young
  2. Most birds form monogomous pairs
  a. Facilitates elaborate parental care system for raising young
  b. Some have extra-marital sexual relations
  e. Birds have large, well-developed brains & Senses
  1. Brain 6-11 times of like-sized reptiles
  2. Like primates- brains lateralized (left brain learning and vocal
  3.Highly developed neural systems & acute senses permit sensational feats of migration & communication
  a. Song birds (Oscines) greatest sound producers of all vertebrates
  b. Bird can navigate using polarized flight from Sun and Stars & possibly even Earth's magnetic fields
  4.Color Vision well-developed- even into near ultraviolet (near-UV)!!
  5.Hearing- incorporates infrasounds (long waves travel greater distances - hear thunderstorms long before humans)
  a. Single earbone (columella)=Stapes
  III. Adaptive Radiation
  A. ~ 300 x 10 9 birds of bout 10,000 species currently inhabit earth
  B. Earliest Known bird Archaeopteryx lithographica (155 x 10 6 yrs old)
  1. Had feathers- Flight? (Glide for sure)
  C. Vast array of world birds in all habitats & sizes
  1. Range in size from Hummingbirds(2 g) to Ostrich (100 kg)
  D. Vast array of bill sizes & Shapes
  1. Varied diets- herbivores, granivores, insectivores, nectivores & carnivores
  E. Vast arrange locomotion patters
  1. Swimmer, runners, soarers, agile acrobats, wing-propelled divers
  F. Many wing patterns
  1. Flight, flightless, soarers, agile acrobats, wing-propelled divers
  G. Feet many shapes and sizes
  
  IV. Bird Names and Classification
  A. ~10,000 species
  1. Standardized official common names (but still local names)
  a. American Goldfinch (yellow bird, finch-bird, wild canary, beet-bird)
  b. Based on system of nomenclature developed in 1758 by Carolus Linnaeus (Swedish Botanist)
  1. Binomial system
  a. American Goldfinch (Carduelis tista)
  Thistle Bold = Bold Thistle Bird
  
  Chapter 2 History
  I. Intro
  A. Diversity of life the result of three evolutionary processes:
  1. Phyletic evolution- gradual change in a lineage
  2. Speciation- splitting of one phyletic lineage
  3. Extinction- termination of a lineage
  a. Extinctions fragment historical record
  1. Erase connections b/n related lineage
  a. Systematists study and hypothesize evolutionary relationships
  II. Reptilian Features of Birds
  A. Reptilian affinities of birds
  1.Single occipital condyle (mammals evolved from reptiles with two condyles
  2. Single earbone (Columella-stapes
  3. Mandible in 5-6 parts (mammals only one part)
  4. Ankle site in tarsal bones (mammals between tibia and tarsi)
  5. Scales on legs~ reptiles
  6. Yolked, polar egg laid by both
  7. Female birds heterogametic (ZW) ~to some female reptiles male birds homogametic (ZZ)
  8. Nucleated RBC's
  9. Sclerotic ring supports eye
  III. Archaeopteryx: The Link
  A. Late Jurassic (Age of Dinosaur) 135-150 X 10 6 years ago
  1.Limestone deposits in central Europe contain excellent fossil records
  a. Central Europe tropical at that time with huge lagoons
  1.Bavarian quarry contained a complete skeleton of a
  small, crow-sized reptile with FEATHERS in 1861
  a. Named Achaeopteryx lithographica (ancient-winged print)
  1. Blunt snout with many small reptilian teeth
  2.Feathers on both wings and tail; probably
  also on body
  3. Probably semiarboreal, capable of gliding &
  weak flapping flight for short distances
  4. Had strong curved claws
  5. Had large furcula
  6. Vanes of feathers asymmetrical
  b. Archaeopteryx contributed to the acceptance of
  Darwin's Theory of Evolution - Origin of Species published in 1959
  IV. Evolution of Feathers
  A. Archaeopteryx was a feathered reptile
  1. Feathers likely originated for temperature regulation
  2. Modern lizards that live in hot, sunny environments tend to have
  large scales that reduce heat loads
  IV. Reptilian Ancestors of Birds
  A. Birds evolved from some line of Mesozoic reptiles
  1. Possibly from a group of reptiles called Thecodonts that gave
  rise to Dinosaurs & Crocodilians
  2. Other Possibility- from small Theropod dinosaurs
  B. Thecodont Theory
  1. Birds show 14 similarities between Thecodont crocodilians and
  modern birds
  a. Recently. Protoavis from late Triassic (75x 10 6 yrs before
  Archaeopteryx), was excavated from Western Texas (1961)
  1. May have had avian features, but specimens are not good
  C. Theropod Theory
  1. Supported because dinosaurs & birds co-existed
  2. Not only includes large carnivores (T. Rex) but many small dinosaurs (the coelurosaurs)
  a. Agile, bipedal reptiles such as Compsognathus (Jurassic Park)
  1. Share 23 of 42 special characteristics of hand,
  vertebrae, humerus, ulna, hindlimb or pelvis between
  they & Archaeopteryx
  V. Evolution of Avian Flight
  A. Arboreal theory
  1. Archaeopteryx had clawed wings- could climb trees & glide- modern
  a. Hoatzin young have clawed wings to climb trees
  
  B. Cursorial Theory
  1. Elongated forelimbs heightened ability of small theropod
  dinosaurs to jump and leap for insects they ate
  C. See figure 2-9 which suggests cursorial preceded arboreal
  VI. After Archaeopteryx
  A. Sinornis santensis
  1. Amazing fossil discovery in 1987 from China
  2. 140 x 10 6 years (late cretaceous)
  3. Toothed, sparrow-sized bird with many theropod features
  a.Many intermediate features between Archaeopteryx & modern birds
  1. Forelimb modified for true flight
  2. Possessed pygostyle to support tail feathers
  3. Feet with hallux for perching
  4. small keel on sternum
  B. Hesperornithiformes (toothed seabirds); also late cretaceous
  1. Hesperornis & Ichthyornis
  a. Hesperoinids were flightless, vestigial wings, powerful
  loon-like divers
  b. Ichthyornis- tern-like flying bird
  c. No extant species today
  1.. Only toothless birds survived the mass extensions
  C. Modern birds
  A. Tertiary period (60-1.5mya) -age of birds & mammals
  1. Many quite large
  a. Diatryma (2 m tall)
  b. Predatory birds (2-3 m tall in SA and Fla)
  c. Huge vulture-like TERATORNS present
  1. 4 m wing span in s. CA
  2. 5 m wing span in Nevada
  3. 8 m wing span in Argentina
  2. Recently some modern orders present by 60 mya
  a. Loons, ducks, auks, gulls, cranes, & petrels (37-54mya)
  b. Appearance of flower (26-7 mya) -> frugivores, nectivores
  VII. History of Avifauna
  A. Early earth unlike today
  1. Mesozoic- Pangea- single land mass before birds
  2. By Cretaceous
  a. N. Laurasia
  b. S. Gondwanaland
  VIII. Speciation
  A. 100,000 species existed on earth
  1. Population isolation
  
  
  #4 Feathers
  I. Intro
  A. Most distinctive feature of birds
  1. Provide insulation-probable main reason for 1st feathers -> homeothermy
  2. Provide aerodynamic power for flight
  3. Provide colors for communication and camouflage
  4. Provide secondary roles
  a. Some modified for swimming, sound production, sound
  collection, protection, cleanliness, water repellency,
  water transport, tactile sensation & support
  II. Feather Structure
  A. Consist mainly of keratin
  a. Also in nails, claws, and hair of other vertebrates
  b. Feather keratin (scales of legs) different than in reptilian scales
  B. Contour feather (body feather)
  a. Long central shaft
  1. Calamus (quill)- anchors feather in follicle in skin
  2. Rachis- rest of shaft; support vanes
  a. Barbs- lateral projecting form both sides of ramus
  1. Barbules projecting form both sides of ramus
  a. distal barbules have barbicels (hook-like)=hooklets
  b. Proximal barbules lack barbicels
  3. Vanes vary form downy nature at base to typical vane distally
  a. plumulaceous- Downy barbules lack barbicels (good insulation)
  b. pennaceous- out vane is stiff, tightly interlocking
  b. Body feathers with after feathers, or aftershaft
  1. Emerges from rachis where 1st barbs branch off
  a. primarily plumulaceous- excellent insulation
  1. Especially long in winter galliforms (Ptarmigan, PC)
  c. Rachis can be long, flexible or stiff = (woodpeckers rectrices)
  d. Curled barbicels in Sand Grouse
  1. Used to carry water (30km) by adults to young, flightless chicks
  a. carry 25-50 ml (chicks can suck up 10-20 ml H20)
  e. Vaned Feathers
  1. Include contour body feathers
  2. Flight feathers of rectrices (tail) and remiges(wing)'
  a. Almost entirely pennaceous -very aerodynamic
  1. Primary feathers attach to hand bones- forward thrust
  a. Asymmetrical- outer van narrower
  b. Most bird with 10
  1. Some passerines w/ 9
  2. Storks, flamingos, grebes, & rheas w/11
  3. Ostrich with 16
  4. flightless kiwi w/ 3-4
  c. Silent flight in owls
  a. Barbs on leading edge of wing long
  curved & fluffy
  b. long barbules minimize rubbing sounds
  c. male woodcock 1 feathers narrow-trilling
  noises during courtship
  2. Secondary feathers attached to ulna
  a. 6-humerus, 19-owls, 40-albatross
  b. Brightly colored for courtship display in
  many ducks
  1. Called speculums
  3. Rectrices attached to pygostyle
  a. usually 12 feathers
  1. Anis & Grouse=18
  2. Snipe=24
  a. modified for sound production in female snipes
  E. Down, Bristles and other feathers
  1. Down- soft and fluffy- plummulaceous
  a. Usually lack rachis- Barbules lack barbicels
  1. Water fowl exception
  2. All provide lightweight insulation
  2. Semiplumes
  a. Have large rachis-plummulaceous vanes
  1. Common of edge of feather tracts
  2. Provide insulation
  3. Fill out aerodynamics contours of body feathers
  4. Serve as courtship ornaments
  3. Filoplumes
  a. Hair-like feathers which monitor position and movement
  of adjacent vaned feathers- attached to sensory corpuscles
  1. Each flight feather may have 8-12 filoplumes
  a. aid in aerodynamic adjustments
  b. those in association with contour feathers
  may help to monitor air speed
  2. Absent in penguins and ratites
  4. Bristles- specialized for sensory & protective functions
  a. Rachis with few barbs at base
  1. Attached to sensory corpuscles on head of all
  except knee of Bristle-thighed curlew & Feet of some owls
  2. Form protective eye-lashes in ratites & cuckoos
  3. Form nostril coverings in woodpeckers & corvids
  4.Form insect-nets around mouths of aerial insectivores
  F. Feather Coats
  1. Swan- 25,000 feather (80% on head and neck)
  2. Passerines- 2,000-4,000 (30-40% on head and neck)
  3. Although light, feathers 2-3x heavier than bones
  a. Eagle: Bones- 300g, Feathers- 700g; total- 4 kg
  4. Attached in distinct tracts= pterylae
  a. Separated by featherless areas= apterylae
  1. Provides space for movements
  2. Also may facilitate heat loss
  3. Apterylae absent in Penguins, Screamer & Mousebirds
  b. Nine major tracts
  1. Capital tract (on head)
  2. Spinal tract (dorsal spine)
  3. Caudal tract (rectrices)
  4. Alan tract (remiges)
  5. Humeral tract (humerus)
  6. Femoral tract (femur)
  7. Crural tract (thigh-tibiotarsus
  8. Ventral tract (ventral- splits on abdomen)
  G. Feather care
  1. Dead structures which should become w/0 daily applications of
  waxy secretions from preen gland = uropygial gland
  a. Bilobed structures on rump of most birds
  1. secretes waxy oil rich in fat & fatty acids -cleans
  & waxes & waterproofs
  a. quite large in water birds
  b. secretions antibacterial
  c. But may promote some fungi that feather lice
  d. Antipredatory (Fowl smell- Hoopees & Woodhoopes
  2. Some birds have combs on middle toe to aid in feather grooming
  3. Secretion toxic in Shrike-thrushes of New Guinea
  a. Alkaloid (neurotoxin) ~ to Poison Arrow frogs of SA
  4. Dustlike talcum on contour feathers of many birds from powder
  down feathers which slough keratin continuously from barbs
  a. Effective waterproofing
  1. Especially evident in herons
  H. Feather growth
  1. Dead structures like hair- can only replace damaged feathers
  a. Epidermal in origin form feather follicle
  b. Some bird can lose feather when frightened (Nightjars)
  2.Although feathers can be regrown when pulled out- usually
  replaced seasonally in molt
  I. Feather molt
  1. Most bird molt annually
  a. Adults typically molt after breeding, replacing entire plumage
  1. May keep for 12 months
  a .Or may replace some feathers before nesting
  following year, converting camouflaged colors
  to bright reproductive feathers
  1. Those that keep them year-round
  may show changes with wear
  a. Starling is spotted in winter- spots wear off-> shiny bird by spring
  J. Feather molt patterns & terms
  1. Basic plumage- plumage renewed after breeding
  2. Alternate plumage- temporary breeding adornments
  a. Scarlet Tanager
  1. Basic- olive-green plumage
  2. Alternate- bright red breeding plumage
  b.Many birds (Robins, Starlings)- single annual molt in fall
  1.Others (Tanager, Goldfinch)- extra seasonal molts
  in Fall & Spring
  a. Some few have twice annual complete molts
  (Bobolinks, Sharp-tailed Sparrows- live in
  Abrasive, Course grass habitats; African
  Larks- Dessert sand wears feathers)
  b. Ruff- molts 3-4 times, but partial molt (reproductive)
  c. Most N.A. ducks drakes undergo rapid prebasic
  molt in Fall that renders them Flightless
  for few weeks- Basic Plumage (female-like)
  called "eclipse" plumage.
  1.Followed by early winter Pre-alternate
  molt- breeding Alternate Plumage by
  winter (Courtship & pairing)
  K. Sequence of feather loss
  1. Primaries: innermost outermost
  2. Secondaries: from both ends < -- > towards middle
  3. Rectrices: from middle >--< towards outer on both sides of middle
  4. Control mechanism poorly understood
  l. Feather colors
  1. Due to biochromes
  2. Due to light reflection
  m. Feather biochromes
  1. Three major types
  a. melanins- earth tones
  1. Eumelanins- dark brown, gray, & black
  2. Phaeomelanins- tans, reddish brown & some yellows
  3. Increased feather wear; absorb IR
  b. Carotenoids- red, orange and yellow colors
  1. From diet
  2. Most bird green due to yellow pigments & structural blue (iridescent)
  c. Porphyrins- reddish or brown feathers
  1. Derived from liver bile pigments (from Fe-containing Heme of hemoglobin)
  2. Some with copper instead of iron
  a. Turacin- magenta in turacos (crow-sized in
  African Forests)
  b. Turcoverdin- Bright green in a few birds
  n. Structural colors
  1.Many bird colors structural (parrot green, bluebird blues,
  Hummingbird iridescence)
  a. Blue & Greens from scattering of short wavelengths by
  tiny melanin pigments in feather barb
  1. Larger melanin particles scatter longer wavelengths-
  parrot green
  b. Carotenoids can convert structural blue green or violet
  c. Iridescence "eyes" of peacocks, Ruby-thr of hummingbirds
  1. Appear black at certain angles
  a. Interference filters reflect visible wavelengths at proper angle
  o. Bright red best male House Finches (Bright red to yellow- orange)
  1. Plumage due carotenoids in diet
  2. Bright red males plumage preferred by females
  a.Bring more food to females during incubation & to nestlings
  
  #5 Flight
  I. Skeleton
  1. Air-filled, hollow-struted long bones
  2. Light-weight toothless bill
  3. Sternum with heel (except most ratites)
  4. Furcula- Flexible, elastic spring
  II. Flight muscles
  A. Pectoralis (15% of body weight)
  1. On heel- pull wing downward power stroke
  B. Supracoracoides
  1. On keel- pulls wing upward recovery stroke
  a. Tendon through triosseal canal to dorsal humerus
  1. Pigeon cannot take off if cut
  a. However, can fly if launched
  b.5 x larger in hummer since upward stroke power
  necessary for hovering & flying backwards
  c. Also larger in penguins
  C. Slotting wings
  1. Prevents stalling at slow air speeds (birds landing)
  2. Also increase air speed over wing- up lift (soaring vultures, etc.)
  D. Flight in Formation
  1. Helps save energy (about 50%)
  E. Hummers
  1. Rate of wing beat constant (53bpm); up/down stroke duration equal
  a. Once oscillation set - neuromuscular impulse necessary only once every
  3-4 beats (like child in swing)
  F. Flightless birds
  
  
  Chapter 6 Physiology
  I. Physiology = internal metabolism & excretion
  A. Advanced physiology of birds permits both power & endurance
  II. High Tb
  A. Birds are endothermic- high internal metabolism
  1. Tb= 40-43 C
  a. Nerve conduct 2x ^ w/ 10C ^
  b. Muscle contraction 3 x " " ^
  2. High metabolism means birds need 20-30 x more energy than similar
  sized reptiles
  a. Warm reptiles are quick, but low endurance
  b. Warm birds are quick, But high endurance
  1. High oxygen & circulatory demands
  III. Avian Respiratory System
  A. Four main parts
  1. Nostrils
  2. Tracheal system
  3. Lungs
  4. Air sacs
  B. With each breath bird replace nearly all air inn lungs
  1.Mammal have residual air left in lungs with each breath (~25% air left)
  2. Birds get more oxygen per breath - more efficient
  C. Birds lack diaphragm
  1. ^ thoracic cavity by lower sternum during inhalation
  a. Exhale- contract muscles of sternum & ribs
  D. Rate of respiration down with ^ size
  a. Resting hummer- 140+ bpm
  b. Resting 10 kg turkey - 7bpm (resting 70 kg man- 15 bpm)
  1. Flight -10-20 x ^ in rate
  E. Most birds inhale thru nostrils (nares)
  1. Nasal cavity with highly coiled conchae
  a. Filter (mucus traps particulate) & warms inhaled air
  2. Olfactory tubercules also convoluted for enhanced smell
  3. Conchae have rete mirabile to control heat & water loss
  F. Lungs attached to air sacs
  1. Bird lung more dense- down volume
  2. Bird lung similar in weight to similar-sized mammals
  3. Primary bronchi ->secondary bronchi ->tertiary bronchi ->air capillaries
  (gas exchange with intertwined blood capillaries)
  G. Air Sac system
  1. Thin-walled (nonrespiratory)
  a. Attached to lungs and wing & leg bones
  2. No. 6 (weavers), 7 (turkeys), 9 (most), 12 (shorebirds)
  a.Paired cervical in neck (inflatable; colored for display in Frigate birds)
  b. Paired anterior thoracic
  c. Paired posterior thoracic
  d. Paired abdominal
  e. Single interclavicular
  H. Takes 2 Respiratory cycles to move air in and out of birds
  1.1st inhalation primary bronchus -> posterior air sacs (no gas exchange)
  2. 1st exhalation -> air capillaries of lungs (GAS exchange)
  3. 2nd inhalation -> anterior air sacs (no gas exchange)
  4. 2nd exhalation -> atmosphere
  IV. Avian Circulatory System
  A. Four chambered heart (50-100% ^ approx. sized mammal) "Birds all heart"
  B. Double circ.
  1 Pulmonary (from RT ventricle to lungs) Backwards in book
  2. Systemic (from left ventricle to lungs) " " "
  a. Right aortic arch (left in mammal)
  C. Resting HR
  1. Approx. 200 bpm
  a. Hummers 1200 bpm
  2. Avian heart more muscle fibers than mammals
  a. Also more mitochondria
  3. --high BP (300mmHg) (man approx. 120mmHg)
  a. Stress, fright can cause vessel rupture or death
  1. Ketterson (77) suggested female Prairie Warbler died of
  heart attack during courtship
  4. Typical diving Reflex in Waterfowl
  a. Bradycardia (down heart rate)
  b. Peripheral vasoconstruction
  1. Blood only to CNS, sense organs & endocrines
  2. muscles anaerobic
  a. High tolerance to ^ lactic acid
  b. High tolerance to blood acidosis
  V. Levels of metabolism
  A. BMR
  a.Metabolism at rest with minimal activity, postabsorptive at
  thermoneutral temperatures
  b. Passerines have highest BMR of all vertebrates
  1. 50-60% higher than similar-sized nonpasserines
  B. Acvitity Metabolism
  1. Birds can sustain 10-25X BMR in active flight for long periods
  of time a. ~-sized mammals: 5-6X for shorter periods
  VI. Insulation & Heat Loss
  A. Down feathers most effective insulators
  1. Arctic Finch- increased down
  2. Tropical " - no down
  B. Winter Sparrow 50% more feathers than in summer
  C. Increased insulation by "fluffing" feathers
  D. Increased pigmentation of feathers (solar collector)
  1. Roadrunner elevates scapular feathers so sun can fall on
  black-pigmented skin on dorsal apteria
  VII. Temperature Regulation
  A. w/i TNZ no change in O2 consumption in endotherms
  1. increased metabolism outside TNZ
  B. Responses to cold
  1. Below LCT --> shivering thermogenesis (esp. pectoral muscles)
  a. Acclimation-dependent
  1. Arctic birds shiver <10 C
  2. Temperate " " <20 C
  2. Hypothermia and Torpor
  a. Tb decreases several degrees at night (more for smaller
  birds) = Hypothermia
  1. TV --> decrease 6C
  2. B-c chickadee --> 8-12 C decrease in winter
  a. Can decrease Tb to profound hypothermia --> torpor
  1. Unresponsive to most stimuli
  b. Hummers show 75% decrease in O2 consumption if
  Tb decreases by 10C
  1. Tb = Ta but increase in metab if Tb<~20C!!!
  c. Warming up from torpor energetically expensive
  1. Hummers require ~1 hour to get Tb to normal
  (~41C) from 20C
  a. Would take kestrel 12 HOURS - so
  kestrels don't get torpid!
  d. Poorwills of SW US & Mexico
  1. "hibernate" for 2-3 MONTHS when Ta <5C
  a. decreases O2 consumption by 90%!
  2. Also show topor during inclement weather
  in summer months
  C. Responses to Heat (behavior)
  1. Above UCT, birds expend energy to cool (^resp-->^evap cooling)
  2. ^ resp (panting) - glular flutter (nonrespiratory); ^evap cooling
  3. Birds lack sweat glands - lose H20 directly thru thin skin
  4. Large water birds can lose most heat thru legs and feet while wading
  a. Storks and new world vultures enhance evap cooling from legs
  by directly excreting on own legs
  5. Ostrich (like camels) have large volume/low surface area and can
  therefore store heat (saves water - don't have to use evap cooling)
  D. Water Economy
  1. Food only source in most nectivores, frugivore & carnivores & many
  insectivores (swallows often drink surface water)
  a. Granivores- all drink (some arid bird better kidney
  function-don't drink)
  2.Metabolic water (sugar+ oxygen yields water + carbon dioxide + energy)
  3.Cool convoluted nasal cavity- condensation & water conservation
  especially in smaller birds
  E. Excretory Systems
  1. Urine mixed with feces
  2. Uricotelic (down water loss; 1/20th water need to excrete 1 g uric
  acid vs. 1 g urea)
  3. Short loops of Henle in Kidney
  a. Arid bird more nephrons
  F. Salt glands
  1. Sea birds drink 3 % NaCl water
  a. Would dehydrate & kill bird
  b. Gull can excrete 90% of salt load in 3 hours
  1. Can excrete salt at 5%
  c. Best developed in oceanic birds
  2. Mallard along coast- up size of salt glands
  3. Absent in all passerines
  a. Coastal marsh passerines use kidneys
  G. Daily Time & Energy Budgets
  1. Gambel's Quail
  a.Spend 28% of time foraging, which costs them 58% of daily
  energy budget
  b. Spend about 50% of time sleeping, which cost them only 25% of
  daily energy budget
  1. Thus, show greatly reduced night time metabolism
• #7 Feeding
  I. Bill Structure
  A. Key adaptation to feeding
  B. Form major features
  1. Upper mandible= maxilla
  a. Attached by nasofrontal binge- flexible
  2. Lower mandible articulates with quadrate (reptilian)
  3. Large jaw muscles
  4. Rhamphotheca- horn covering
  a. Sharp (bookies)
  b. Serrated (mergansers)
  c. Notched (falcons & toucans)
  5. Some bills flexible also at tips bill (scalopacids)
  C. Variously adapted
  1. Filter- feeding flamingos
  2. Seedcaching finches
  3. nectar-feeding hummers & sunbirds
  II. Avian Digestive System
  A.Lack of teeth mean digestive system specialized to process unmasticated
  food.
  B. Major parts
  1. Oral cavity
  2. esophagus
  3. Crop
  4. Two-chambered stomach
  5. Intestines
  6. Cloaca
  7. Also liver/pancreas
  C. Highly efficient energy extraction of rapidly processed, small volumes
  of food
  D. Saliva
  1. Bird's nest soup (swift's of Southeast Asia
  E. Tongue
  1. Taste buds (small number compared to mammals)
  2. Variously shaped
  F. Esophagus- muscular connection to stomach
  a. Expanded in fish-eater
  b. Pigeon milk in columbiformes
  c.Inflatable for display & sound production in ostriches, Prairie-
  chicken & male painted-snipe
  D. Crop
  1. An expanded section in many birds
  a. Stores and soften food
  1.Some digestion (hoatzin-eats only leaves)
  G. Stomach (2 chambered)
  1. Preventriculus (anterior glandular)
  a. Absent in reptiles; especially well- developed in fish-
  eating birds Raptors
  b. Low pH (0.2-1.2 pH)
  1. Quickly dissolves bones
  a. Shrike can digest mouse in 3 hours
  2. Gizzard- (posterior muscular grinding organ)
  a. Well developed in granivores
  1.Turkey can pulverize stainless steel lancets! b.Grebes swallow feathers (b/n gizzard & intestines)
  1. Filter scales & Fishbones- regurgitated
  c. Size can seasonal
  1. Parrotbill- granivores in winter (larger)
  - insectivoresinsummer (smaller)
  H. Food passage times
  1. Short in berry-eater (15-20 min)
  2. Longer others
  I. Assimilation varies
  1. 98% nectar in hummers
  2. Raptor 70-90%
  3. Herbivores 60-70%
  4. Spruce Grouse 30% spruce leaves
  5. Many passerines cannot digest sucrose (lack ???)
  a. Starling learn to avoid sucrose in lab tests
  J. Cecal- side sacs at posterior end of large intestines
  a. Best developed in ground-dwelling flightless birds
  IV. Feeding behavior & Energy Balance
  A. Tool use
  1. Woodpecker finch of Galapagos
  2. Egyptian Vulture (rock on ostrich eggs)
  3.Great Heron-seen using bird as fishing bait? What???
  4.50's in England- great tats learned to tear off paper bottle caps
  5. Sharp- beaked Ground Finch (Galapagos) drinks blood of nestling
  birds by pecking open growing pin feathers
  B. Increasing energy Profit
  1. Profit= energy gain- energy lost/ foraging time
  2. Birds tend to choose high energy foods
  C. Learning to feed
  D. Nutrition
  1. Little known
  2. Author encourage more research
  E. Energy Balance
  1. Fat reserve usually < 10%
  1. 10 yr warble cannot survive day w/out food
  2. 200g Kestrel can survive 5 days
  3. Female European Penguin fast 3-4 months during
  frigid incubation vigils ( down 45% BW)
  2. Food Caches
  a. Acorn woodpeckers
  b. RAPTORS- cache portion
  C. impales
  
  #8 Brain & Senses
  I. Intro
  A. Bird Senses beyond the ken of man
  1. Color vision, but also UV
  2. Hearing also includes infrasounds
  3.Use sun 7 stars for navigation, but also may use magnetic field
  4. Also can detect barometric pressure changes
  5. Large brain (6-11x approx. sized reptiles)
  a. Functional laterilization
  1. Left brain learning & vocalization
  II. Avian Intelligence
  A. Corvids do esp. well in laboratory learning experiments
  1. Raven and Parrots have been taught to count to 7
  2.Jays quickly learn to distinguish edible from inedible butterflies
  by watching other jay in adjacent cages
  3. Already mentioned tool use which is probable learned by observing
  others.
  a.Egyptian Vulture using rocks to open ostrich eggs
  b. English opening milk bottles
  III. Avian Brain
  A. Brain
  1. Forebrain ( olfactory bulbs & cerebral hemispheres)
  a. Complex behavioral instincts
  b. Sensory integration
  c. Learned intelligence
  2. Midbrain (optic lobes & cerebellum)
  a. Regulates vision
  b. Muscle coordination & balance
  c. Physiological controls
  d. Neurohormonal control of seasonal reproduction
  3. Hind brain (medulla)
  a. Autonomic control systems ( heart, resp, BP)
  4. Avian Brain with only a small cortex
  a.Lower brain parts for complex behaviors and learning
  b. Removal of cortex did not prevent pigeons from normal visual discrimination, nor prevent it form mating
  and rearing young successfully
  5. Hyperstructure (unique in birds)
  a. Center of avian learning & intelligence
  1. Well-developed in crows, parrots & passerines
  6. Optic Lobes (midbrain)
  a. Huge in birds
  1.Imply importance of visual balance input for flight
  IV. Control of Song
  A. Normally controlled by left hemisphere (as in humans)
  1.If left hemisphere damaged in young canary -right side
  2. Seasonal up testosterone- up nervous synapses
  a. These disintegrate in Fall when down testosterone
  V. Spatial Memory
  A. Hippocampus
  1. Destruction chickadees
  a. Still cache seeds, but can't find them!
  VI. Avian Vision
  A. Eyes are large
  1. Those of eagles & owls approx. to man's size
  2. Greater power (2.5-3X) that of man
  B. Binocular Vision
  1. Except owls, most birds eyes on sides of head- small binocular field
  a. Birds mostly view close objects with one eye (monocular)
  1. "Bob" head to view at more than one angle to Up depth
  perception
  VII. Eye Anatomy
  A. General 1. Anterior component: cornea & lens
  2. Large posterior component: main body & retina
  3. Sclerotic ring of 12-15 ossicles
  a.Muscles attached for focusing (Crampton's muscle, Burkes's
  muscle)
  4. Pecten (reptilian)
  a. Function still unclear
  B. Cornea & Lens
  1.In birds both cornea & lens change shape during focussing
  a. Crampton's muscle changes shape of cornea
  1.Not important in diving birds because RI of cornea = RI water
  b. Burke's muscle changes shape of lens
  2. Pupil round in all birds
  a.Vertical slit in bright light in skimmers (round in dim light
  3. Iris color varies
  a.Eyes "shine" from tapetum in most nocturnal birds
  C. Retina & Fovea
  1. Large # cones- good vision over all of retinas!
  a. Human =2 x 10 5 /mm 2
  b. House sparrow= 4 x 10 5 /mm 2
  c. Common Buzzard= 1 x 10 6 /mm 2
  d. Peripheral retina 10 x cones of human
  2. Fovea centralis- area of acute vision
  a. most bird w/ one fovea
  1. Deeper & more complex in most birds
  a. Less so in chicks & pigeons
  b.Bifoveal (temporal + central) in fast flying birds
  1. Hawks, eagles, terns, hummers, kingfishers, & swallows
  a. Give good binocular vision (temporal fovea)
  c.High cone densities also in ribbon-like stripes
  around retina in Albatrosses, grebes, & plovers
  1. Many up perception of horizon
  D. Pecten (unique in vertebrate)
  1. Large, pleated vascularized structure attached near optic nerves
  2. Most feel it provides nutrition to avascular retina
  a. REtina has blood vessels in mammals
  E. Color Vision
  1. Rods- dim light vision (predominant in nocturnal birds)
  2. Cones- bright light vision (predominant in diurnal birds)
  a. REd (670 mm)
  b. Green (540 mm)
  c. Blue (480 mm)
  d. UV (360 mm)
  1. Have associated oil droplets (lower verts)
  a. Act as red-yellow filters
  1. May enhance contrast
  b. Yellow droplets concentrated in central & lower
  retina
  c. Red droplets concentrated in peripheral & upper
  retina
  1. Function not really known
  3. Birds are sensitive to near UV wavelengths
  a.Bird lens transmits UV; human absorbs near- UV except
  children
  b. Hummer, Kingfisher, Mallards, and passerines have UV
  1. Chickens and probably owls- do not!!
  *c.Kestrels in Scandinavia- see UV reflective urine trails of
  voles to find best food territories
  VIII. Magnetic Field
  A. Birds posses crystals of magnetite
  1. Near olfactory nerves in pigeons
  2. In neck muscles of Bobolinks links
  B.Recently, photo pigment rhodopsin, which theoretically sensitive to
  magnetic fields has been suggested to be involved.
  IX. Hearing
  A. External ear, middle ear, inner ear
  1. Simpler structure than mammals, but equally sensitive
  a.Outer ear covered by auricular feathers- facial feather in
  owl collect sound
  b. Single? middle ear bone- columella (stapes)
  c. Cochlea with short basilar membrane with hair cells
  B. Auricular feathers protect ear form air turbulence noises
  1. Diving birds- protect ear from pressure changes
  C. Sound reception processed by nuclei in hind brain (medulla)
  1. Nocturnal owls have more nuclei (approx. 48,000)
  a. Duesenberg owl (approx. 11,000)
  b. Diurnal crow (approx. 14,000)
  D. Hearing ability
  1. Probably not extra ordinary
  a. Human can probably hear fainter sounds at most frequencies
  b. Range is narrower in birds
  1, Owls
  a. GHO better low
  b. Barn owl "hi
  c. NO bird ultra sonic
  2. Birds- Infrasounds<20 Hz
  a. Significance unknown
  1. Hear thunderstorms??
  2. Hear oceans for navigation??
  E. Echolocation Birds
  1. Navigation
  a. Cave Swiftlets (SE Asia) 2-10-KHz
  1. Approx. 1/10 efficient as ultrasonic echolocation
  b. Cave nesting Oilbird (SAmer); 1-15KHz
  1. 1. collide with objects smaller than <20MM
  2. Prey detection
  a. Barn owl hunts in DD
  1. Human ear horizontal plane discrimination, but 1/3 in
  vertical plane
  2. Assymetrical ear openings in owl and facial feather
  arrangements- up vertical plane detection
  a. Left facial ruff faces downward
  b. Right facial ruff faces upward
  *b. Canal b/n inner ears in songbirds
  1. Sound in one ear heard by other ear- may up directional
  sensitivity
  a. ??? since head small -ear separation small
  F. Mechanoreception 1. Herbst organs (touch)
  a. Abundant in tip of bill of mud probers (sandpipers, snipes)
  b. Abundant in tongue of woodpeckers
  c. Abundant in filoplume & bristle hair follicles
  2. Semicircular. canals
  a. Well-developed- enhance flight
  1. Bigger in better fliers (pigeons, owls, hawks)
  a. Smaller in ducks & galliforms
  G. Barometric responses
  1. Detect 5-10m difference in altitude (important for migration
  altitude)
  a. Receptors unknown
  H. Taste and Smell
  1. Few taste buds (24 chicken, 37 pigeon, 62 Quail)
  a. Human- 10,000
  2. OLfactory Bulb
  a. Large in Kiwis, vultures, & petrels
  b. small in passerines
  1. Detected orders approx. to rats & rabbits
  c. Evidence that drake mallard can smell sexual odor of hen
  d. Storm-petrels (seabirds) detect odor of krill (2.5-25km)
  e. Honey guides smell beehive odors
  I. Senses & Behavior
  
  
  #9. Visual Communication
  I. Intro
  A. Important in courtship displays, territory defense, flock maintenance
  1.Displays used in conjunction w/ plumage ornament, colors,
  postures, and vocalization, which have evolved together
  B. Sexual Dimorphism
  1. female usually more brightly colored
  II. Plumage Color Patterns
  A. Concealment 1 role
  1. Ptarmigan- white winter, brown-summer
  2. Breast bands of plovers- disruptive patterns- up concealment
  3. Countershading- up concealment
  a. Gray Plover (Black-billed Plover)- dark below, light above
  b. Breeding Bobolink- Black underparts, light above
  B. Visual Identity
  1. Many females with distinctive plumage patterns
  Usually feathers, but also bill, legs, feet
  2. Snow Goose (recessive-white, heterozygous- dark)
  a. Whites usually mate
  b. Darks usually ;mate
  1. REgardless of color- geese raised by white- mate white
  geese raised by dark- mate dark
  geese raised by mixed- mate mixed
  C. Evolution of displays
  1. Closely related birds tend to have similar behaviors
  a. Pelicanidae- all have "bowing" courtship displays
  D. Ritualized behavior
  1.Quail, pheasants, & peacocks- incorporate ritualized food
  behaviors into courtship
  a. Quail feed male before copulation
  b. Pheasants & peacocks- mock pecking- do not feed male
  E. Agonistic Behavior
  1. Generally little DIRECT contact
  a. Mostly use threats and appeasement displays
  2. Threat:
  a. Bill & Wing displays
  b. Submissive bird may lower head, turn away
  F. Sexual Selection
  1. Plumages highlight contests among males for mates
  2. Plumages highlight preferences of females for males
  3. RWB
  a. Female selects male with best territory
  1. Older males in INDIANA (males help feed young_
  2. Any males in WASHINGTON (male does not help)
  b. Blacken RED Epaulets
  1. males lose territory
  2. But can still mate
  ***** READ SUMMARY PAGE 231 *****
  
  #10 Vocal Communication
  I. Intro
  A. Syrinx- sound box in birds
  II. Physical Attributes
  A. Songs- Complex vocal displays of territorial male birds (short notes with
  broad frequency)
  1. Males- advertised sex
  attracts mates
  helps hold territory
  maintenance of pair bond
  B. Calls- short, simple vocalization given by either sex (usually faint,
  thin, highpitched, & hard to locate)
  1. Distress calls
  2. Flight calls
  3. Warning calls
  4. Feeding calls
  5. Nest calls
  6. Flock calls
  C. Low Freq. (grouse) -typical forest birds
  1. Long, distance, less alternation & interference
  D. Buzzy and Broad -band songs- typical open habitats
  1. Simple sustained notes would be disturbed by strong temperature
  gradients and air turbulence
  E. Sound production by Syrinx (larynx without vocal cords- open/closes
  glottis when swallowing)
  1. Swans, Turkey Vultures, Rheas- only hiss or grunt occasionally
  2. Contains tympaniform membranes on each side of syrinx
  a. Syrngeal muscle control
  1. Muscles absent in Ratites & New World Vulture (only
  grunt or hiss)
  2. Nonpasserines- 2 pair extrinsic
  3. Oscines- 2 pair + up to 6 pair intrinsic muscles
  3. Can be modified by length of trachea
  1. Swans & Cranes soiled in sternum
  F. Avian Vocabularies
  Chaffinch 1. many different calls (5-14 calls of varying functions)
  *2. Wood warblers- accented song- attract mates on territory
  unaccented song- edge of territory for defense
  (Chestnut-sided Warbler)
  G. Vocal Identity
  1. Song playback studies
  2. Female swamp & song Sparrows + E2 --> response
  H. Individual Recognition
  1. Recognize "voice"
  2. Duet- overlapping, cooridinated bouts of sounds
  a. 222 species (44 families)
  1. Mostly monogomous tropical birds which breed year round
  a. Territory defense & pair bond
  I. Song Repertoires
  A. female canaries build nest faster, lay eggs sooner & larger clutch
  in response to large -repertoire male songs
  B. May prevent declining interest by females
  J. Vocal Mimicry
  1. 15-20% of passerines of world
  a. N. Mockingbird &Starlings, Marsh Warbler, Australian
  Lyrebird
  1. Mocker- 150 songs (also imitate insects, frogs &
  mechanical sounds)
  a. Ton Tx - Bell's vireo, Great-tailed Grackles &
  Dickcissels
  2. Probably up size of song repertoire- up mating success
  3. Male mocker sings more when copulating,
  more when mate removed & more, especially at night if unmated!)
  2. Australian Lyrebird
  a. "Imitates": Gee up Bess, Piano, tractor, pigs, violin
  K. Learning to Sing (Inherited, learned, invented)
  1. Inherited
  a. Chickens, doves, and some Flycatchers
  b. Brood parasites: Cuckoos, honey guides, & cowbirds
  2. Learned
  a. E. Meadowlark
  1. Inherits call notes
  2. Learns songs from other meadowlarks
  b. Parrots, hummingbirds, & oscine song birds
  c. Four stages
  1. Critical period (usually < year)
  a. Isolation-- normal song
  2. Silent Period (up to 8 months) learned song stored
  3. Subsong period- plastic song -real song!!
  a. Auditory feedback essential
  1. Deafened birds- normal song
  2. Deafening after crystallization- normal song
  L. Dialects (coast to coast or hilltop to hilltop)
  1. New dialects can be learned as birds colonize new area
  2. Young males up reproduction by imitating older males
  3. Dialects may influence genetics of local population
  a. Fact that Iindigo Buntings take up song dominant males in new
  territories refutes #3.
  
  #11 Annual Cycles
  I. Annual Cycles A. In permanent residents, three main sequential tasks
  1. Breed
  2. Molt
  3. Survive until next breeding period
  B. Tropical bird in Borneo (Stable climate, unchanging daylength)
  1. Breed after heavy rains (Dec)
  2. Molt after young fledged (May)
  a. Continue thru 2 mo "Dry Season"
  1. Stay alive as food down
  3. Dec. rains- breed again
  C. Temperate Birds (similar sequence, but timing controlled)
  1. Breed spring (up gonad sized- up hormones)
  2. Molt after young leave in July (molt Aug-Sept)
  3. Survive Winter- breed again
  4. Migration complicates cycle in nonresident birds
  a. Up feeding after molt
  b. Nocturnal unrest (Zugenruhe)
  c. Migrate to winter grounds
  d. Many migrants molt twice
  1. After breeding
  2. Either late winter or early spring (usually not 1 or 2
  feathers)
  3. White crowned Sparrows (West Coast)
  a. Alaska breeders migrate to S. Calif (nest latest)
  b. S. Cal. breeders resident, nonmig (nest earliest)
  c. NW USA breeders migrate to S. Cal. too (nest
  intermed.)
  II. Circadian & Circannual Cycles
  A. Biological Clocks essentialforreproduction, migration, and
  navigation, and day-length measurement in birds.
  B. Circadian (circa= about, dia= day)
  1. All of us have innate daily rhythm approx. 24 hours in length
  a. Synchronized by "Zeitgeber"= time given
  b. w/o zeitgeber- freerun (24 hours +)
  ___________________________
  \------------\\\\\\\\\\\\\\\
  Light Dark
  ^^^^^^^^^^^^^^-------------- Bird entrained by photoperiod
  ^^^^^^^^^^^^^^-------------- (activity ends at lights out=Dark)
  ____________________________
  \\\\\\\\\\\\\\\\\\\\\\\\\\\\\
  Constant Darkness
  ^^^^^^^^^^^^^^-------------- Bird free-runs (activity onset delayed
  --^^^^^^^^^^^^^^------------ each day)(T=approx 24 hrs 15 min)
  ----^^^^^^^^^^^^^^----------
  ------^^^^^^^^^^^^^^--------
  Drawing above shows free-running rhythm in birds in DD
  
  C. Circannual Rhythms (circa=about, annu= year)
  1. Starlings & Warbler kept under 12L:12D for 4-8 hrs
  a. Still show molt & testes size changes (10-11 month intervals)
  2. In equatorial Sooty Terns
  a. Approx. 10 month reproductive cycle
  D. Role of Photoperiod
  1. Two components a. Environmental light & neural receptors
  b. Internal circadian clock measures day length
  1. Permits birds to reproduce at optimal season
  2. Synchronizes reproduction in mated pair
  3. Terminates reproductive function
  2. Internal clock
  a. Pineal gland- melatonin output
  E. Reception- Control System
  1. Hypothalamus- light sensitive (blind birds- up gonad in LD)
  a. Neurohormones pituitary release of LH & FSH (gonadal
  recrudescence & sperm production)
  F. Physiological control of molt
  1. Not well understood
  a. Probably down steroids permit molt
  1. Exog. steroids stop or slow molt
  2. Castrated birds molt on schedule
  G. Breeding Seasons
  1. Many proximate factors
  a. Desert species- rain
  b. temperature maybe most important (Although many temp. bird
  lay eggs in Jan-Feb. )
  c. Pinyon Jay -when seeds of Pinyon Pine available
  1. Sometimes breed in autumn in bumper pine seed crop!
  H. Energetic cost of Reproduction
  1. Energetic costs of molt & reproduction & migration above those of
  maintenance (staying alive)
  2. Peak reproductive activities increase daily energy expenditures by
  approx. 50%
  a. Day time activities may be 2-3 x up; overnight constant
  b. Female waterfowl (large eggs & large clutch)can 2x up total
  energy requirements
  1. Protein reserves down dramatically during egg laying
  I. Costs & Timing of molt
  1. Scrub Jays- 15-16% up energy requirements during peak periods of
  feather replacement
  2. Molt is so important to Sooty Terns (Ascension Is) that it might
  prevent continuous yr- round reproduction;
  3. Some female birds such as Eurasian Kestrel molt 1 while incubating
  & being fed by male
  4. Many northerly breeder (arctic) short breeding season- must molt
  before end of breeding to be ready for migration
  III. Annual cycle of the Waterfowl
  A. Intro
  1. Geese have simple annual molt
  2. Many northern latitude duck more unusual
  a. Prealternate molt starts before the fall, prebasic molt
  finishes
  b. Drake Mallards drop all flight feathers in a rapid
  postnuptial molt and become flightless
  1. The resulting "hen-colored", basic plumage is called an
  "Eclipse" plumage because of its short duration.
  a. Before the eclipse is complete, the drake begins
  the prealternate molt which complete by October when
  courtship activities begin
  2. Hen Mallards undergo prealternate molt just prior to
  spring migration
  B. Nonannual cycles
  1. Some birds follow other than 12 month cycles
  a. Rufous-collared Sparrow (Columbia), near equator
  1. Breeds and molts twice/yr- 2 "dry" seasons/yr
  2. In Costa Rica: Breed twice/yr- one molt and one partial
  molt/yr
  2. Many equatorial birds show 8-10 mo breeding cycles (Ascension
  Island, Galapagos Is.)
  3. Very Large birds
  a. Crowned eagles, Eurasian Griffons, Wandering Albatrosses
  1. Breed ever other year
  b. King penguins
  1. Incubate eggs 2 mo, raise young 10-13 months then molt
  a. Result: breed twice every 3 years
  4. many tropical and desert birds opportunistic- breed after rainfall
  maintain Partially developed gonads
  
  #12 Migration
  I. Intro
  A. Every fall approx. 5 x 10 9 birds (187 species) leave Europe for Asia & Africa
  B. Every fall approx. 5 x 10 9 birds (200+ species) leave NA for Central
  and South America
  1. Migration allows year-round activity
  2. Migration allows exploration of seasonal feeding
  3. Migration allows living in favorable climate
  a. Physiologically & energetically demanding
  C. The decline of neotropical migrants since 1950, a major conservation issue
  II. Patterns of migration
  A. Most NA birds migrate N-A(major rivers and mountains run N-S)
  B. Most European birds migrate E-W (Mediterranean, Alps, Coastline run E-W
  C. SA
  1. A goose & Ibis breed southern tip of SA, but migrate only to
  central Chile & Argentina
  2. Some swallows & flycatchers move to SA tropics- Exceptions
  D. Africa
  1. Only approx. 20 species winter to Equatorial Africa
  a. 183 Palearctic species migrate to sub-Sahara Africa for winter
  III. Why migrate?
  A. Vagrants
  1. Some follow specialized food supplies (Red-crossbills)
  2. In tropics- wandering common in nectar & fruit-eaters
  B. Some Altitudinal - grouse /hummers move up/down mts.
  C. Arctic Terns (70N. lat) Antarctica 12,000 km
  1. Also Buff-breasted sandpipers
  2. Some migrate shorter distances
  a. Wood Warblers NA- Central Am. & West Indies
  D. More than 1/2 small land birds never return next year in NA
  1. 100 X 10 6 waterfowl migrate south
  a. only 40 X 10 6 return
  E. HUge numbers of European migrants shot or trapped
  1. Only Israel had bird protection laws F. Migration continually evolving
  1. European Serrin- non migratory
  a. Expanded population to North in last 100 years
  1. Northern Population Migratory
  2. Blue Jay expanded northward into Canada- these population migrate
  3. Barn swallows migratory
  a. Recently began nesting in Argentina (winter home)
  G. In general
  1. Birds that migrate to tropics survive winter better than those
  that are resident in temperate zone
  2. Tropical residents have low productivity
  a. High survivorships
  1. Few nest succeed; several attempts per year
  2. Low clutch size
  3. Adults survive longer time
  H. In European Robin
  1. Some birds do not put on fat & show migratory unrest ( move 5 km)
  2. Some birds do put on fat & show migratory unrest ( 1000 km)
  I. In Juncos ( breed N. US & S. Canada)
  1. Adult females- southern most US
  2. Adult males & young females - middle states
  3. young males - northern most US( OH IN)
  J. In Sanderlings- appear to migrate to best available food resource from
  Washington to S. Chile (7500 km)
  III. Migratory feats
  A. Arctic terns- 25,000 km round-trip
  1. Lesser yellowlegs flew from MA- Martinique IS (3220 km)in 6 days
  B. Up to 12 X 10 6 birds migrated over Cape Cod in one night flying
  nonstop 80-90 hrs to SA
  1. Often exhausted birds land on SA island when flight conditions poor
  C. Northern Wheatear (Greenland- British Isles (2000 km of open ocean)
  D. Many European Migrants
  1. Fly 1100 km across Mediterranean , then
  2. Fly 1600 across Sahara Desert
  E. Many "Fall outs"
  1. Neotrops in Spring along gulf coast (TX, LA ) in April!
  IV. Fatty Fuels
  A. Premigratory hyperphagia
  1. up fat stores (2 X more energy & metabolic water /g)
  2. Black poll Warble - 2 X up body weight (11- 21 g)
  3. Ruby-throated hummer- 2 x up body weight (3-5+ g)
  B. Normally fat approx. 3-5% of Body weight
  1. Short range migrants- 13-25% BW
  2. Long range migrants- 30-50% BW
  C. Many birds refuel as migrate
  1. Song birds fly several hundred km
  a. Pause 1-3 day (rest & refatten)
  2. Arctic migrants often refuel 3-4 times
  3. Many shorebird congregate -Conservation needed
  V. Flight ranges
  A. Small birds w/ 40% fat reserve can fly 100 hrs (2500km)
  B. Migrant shorebirds can fly 3000-4000 km nonstop
  C. Hummingbird
  a. 2 g fat (consumed at 9.18 watts, flying at 40 km/hr) can fly
  1000km in 26 hrs- Nonstop across Gulf!
  D. Most birds use tailwinds!
  VI. Timing of migration
  A. Caged migrants
  1. Nightly unrest during migratory period (Zugenrube)
  a. Wakes shortly after dark- midnight
  2. Migratory birds- No zugenruhe
  3. Due to up adrenal hormones & prolactin
  B. Up winter day length (up photoperiod) stimulates spring restlessness,
  hyperphagia, fat deposition and up BW
  C> Some birds migrate w/ thaw line (Canada Goose; American Robin)
  D. In U.S., spring migration coincide w/ strong warm southern winds from
  Gulf to NE
  1. Fall migration follow strong NW winds
  F. Age & Sex
  1. Male generally migrate north before females
  2. In fall, many young migrate before parents
  3. Often females migrate farther south (Junco, Am Kestrels)
  G. Diurnal
  1. Hawks- soar on thermals
  2. Swifts, swallows feed
  Nighthawks, kingbirds
  Larger birds- Eagles, stork, Heron, Crows, Pelican
  Some Passerines- Robins, Bluebirds, Shrikes, granivorous passerines
  H. Nocturnal (starts 1-3 hours after sunset, peaks near midnight, end
  by 3 A.M.)
  1. Many passerines (Fly catchers, most thrushes, warblers, rails,
  woodcocks, orioles, tanagers, buntings, Harris' sparrow)
  I. Day or Night
  1. Waterfowl, shorebirds, cranes, auks
  J. Altitudes
  1. Nocturnal migrants < 700-800 m (0.5 mi)
  a. Many fly 3000-7000 m to escape air turbulence or for
  tailwinds
  2. Shorebirds typically fly at 2000-4000 m
  3. Migration over water up, then over land
  4. Migrant fly lower into opposing winds
  
  #13 Navigation % Orientation
  I. Intro
  A. Global navigation- extension of natural homing abilities (find
  nests, feeding grounds, and territories) Swallow return to
  Capistrano, Vultures to Hinkley
  B. Homing feats
  1. Pigeon- honed b Egyptian & Romans
  a. Can fly 800 km/day form unfamiliar places
  2. Many shearwater returned to nest borrow in Wales , 12.5 day after
  being transported to Boston
  C. How done?
  1. Use sun by day, stars by night; also olfactory cues, the Earth's
  magnetic field , and possibly infrasounds
  a. Inmate magnetic compass that includes both direction &
  duration form the base for development of advanced
  navigational abilities
  III. Visual Landmarks
  A. Used for both diurnal and nocturnal migrants
  1. Straits act as funneling points
  2. Coasts & rivers imp too
  3. Homing pigeons appear to use landmarks near home only
  a. Frosted Lens (<3m)
  1. Flew 170 km to loft (some lost, some crashed)
  IV. Solar Compass
  A. 1950,51- Gustav Kramer (Germany)
  1. Starling (Zugenruhe)
  a. As long as they could see sun, focussed attention of NE
  (Spring migration)
  B. On overcast days, no directional tendency
  B. 1951, 53, 68- Geoffrey Mathews (England)
  1. Pigeons homed well on sunny days, but poorly on overcast days
  2. From releasing pigeons at different times of day Mathews proposal
  that pigeons could tell time of day from sun's position
  a. Sun moves 15 degrees/hr (2.5degrees/min)
  1. Thus solar compass must be time-compensated
  3. Kramer trained starlings to feed from NW cup in circ. cages when
  they could see sun
  a. When all they could see was stationary bulb, they fed from
  cups increasingly farther left, as they compensated for
  hourly change in position of "Sun"
  4. Konrad Hoffman (54) trained starlings to feed from N tray in sunlight
  a. Placed experimentals in room 12L:12D (lights on midnight
  VS.0600)
  1. Shifted feed to east
  5. Clock-shift studies confirmed in pigeons
  C. Stellar compass
  1. Franz & Eleanor Sauer (57,58) put hand-reared Garden Warblers in
  circular cages in planetarium
  a. Migratory unrest observed through glass bottoms of cages
  1. Spring Hopped toward N
  2. Fall " " S
  a. When planetarium turned off- disorientation
  b. When axis of rotation moved 180- birds reversed
  compass headings
  2. Stephen Emlen (67,75) Duplicated studies w/ Indigo Buntings
  a. Used paper cone w/ink pad
  1. Buntings oriented NE w/ simulated spring sky
  2. " " SW " " Fall sky
  3. " " in reverse if axis of rotation reversed
  4. " disorientated when dim light & planetarium off
  3. Time compensated Stellar Navigation
  a. Warblers changed heading when planetarium was set ahead or
  behind correct times
  b. Buntings maintained NE orientation under phase shifted
  planetarium skies (birds use axis of rotation for N-polaris)
  1. Similar results w/ mallards & Wh-thr sparrows
  D. Olfaction Responses
  1. Leach's Storm-Petrels apparently locate nesting burrows on forest
  islands by using olfactory cues while flying downwind of nest
  burrows
  a. Those w/ bulbectomies did not return to nest burrows
  2. Italian Pigeons
  a. Earlier experiments (Papi 71,72)- bulbectomy down homing
  1. Not confirmed in American Pigeons
  2. Mixed results in German Pigeons
  E. Geomagnetic Responses
  1. Teagley (47) showed magnets disrupted pigeon homing
  2. Wiltsckko (65) showed European Robins oriented in steel cages with
  out sun
  a. Robins reversed orientation when magnetic field reversed
  3. Keeton (71,72) show free-flying pigeon with BAR magnets did not
  orient properly on cloudy days, wheras controls with BRASS barsdid
  4. Walcott Green (74) used Helmholtz coils with reversed coils flew
  opposite direction of home
  5. While magnetic in some birds- probabley photo receptors are
  sensitive too
  a. slight change (10%) in field's intensity disrupts
  orienttation for 3 days
  b. Sensitivity is low (10 -7 to 10 -9 gaus), which corresponds
  to sunspot & bills of iron ore
  1. These natural fluctations disrupt orientation of
  nocturnal migrants
  c. Inclination important, not true N-S
  F. Learning to Navigate
  1. Partly unnate & early experience
  a. Immatures get lost more often (rare bird sightings!)
  2. Duration of unrest to length of migration in a species
  a. Willow Warbler- 4 mo
  b. Clioffinch- 1-2 mo
  3. Hand-reared caged migrants exhibit well-defined orientation
  a. Banting can use Polaris or Betelgeuse as axis of orientation
  4. Suggestion that magnetism compass serves as initial basis for
  orientation for young birds; other compasses learned to form an
  integrated system
  a. Young pigeons do not home well if transported in a distorted
  magnetic field or if equipped with BAR magnets on 1st flight
  b. Homing pigeons reared with ut seeing sun & experienced on
  only overcast day , prob cannot use solar compass
  1. Do not respond to clock shifts
  G. maps & Bicoordinate Navigation
  1. Sun position can provide Lat & long info
  a. Clock-shift experiments however seemto imply that sun is only
  a simple campass
  2. Answers unknown!!
  
  
  #14 Social Behavior
  I. Territorial Behavior
  A. Birds establish, maintain & protect sp??? relationships
  1. Is a flexible and dynamic behavior
  a. Balance beneath costs & benefits (Great Tits forego
  territorality on coolest days)
  B. Types
  1. Feeding territories (simplest)
  2. Nesting territories (most complex)
  C. Average size
  1. body size, energy requirments & food habits
  a. benefits > cost
  D. Typically defend against some species
  1. Nectivorous often more general
  2. Wintery N. Mochers defend berries against other frugivores
  3. some birds also defend nesting territories against other closely
  related species
  4. Territories may be occupied and defended by a single bird, a mated
  pair, or extended family, or a group of unrelated individuals
  a. Wintering parids (tils, chichadees) defend woodlot
  territories containing both food & roosting holes
  II. Dominant Behvior
  A. Territorality imparts invincibility to territory owner
  1. Territory owners usually win encounters with intruders
  a. Usually have most to lose, fight harder & longer
  B. Territorality is one expression of dominancebehavior
  1. As birds encounter- subordinates (cease challenging dominants)
  2. Use larger > smaller; male > female; older > younger
  3. Dominant birds live longer (juncos & field sparrows)
  III. Flocking behavior
  A. Flocking often down predation and enables cooperative foraging
  1. Often enhanced when down food supplies
  IV. Feeding in Flocks
  A. Up feeding efficiency
  1. Flock s of white pelican encircle and trap school of fish in
  shallow water
  2. Group of cormorants and mergansers drive fish toward shallow
  shores
  3. Ravens steal more gull eggs when hunting in groups
  4. Flock benefits form "beater effect"- beat prey from bushes
  5. More prevalent in smaller birds with limited fasting abilities
  V. Safety in Flocks
  A. Joining flock decreases your chances of predation
  1. Predator confusion
  a. Starlings tighter flocks when hawk sighted
  1. Merlin success down as flock size increase in sandpiper
  flocks
  2. Up predator detection (more eyes)
  a. Means less time looking for predators, more time to feed in
  flock
  3. Can provide mobbing behavior to dissuade predator (crows mob
  hawks)
  
  
  
  #15 Reproduction
  I. Gonads
  A. males - mostly paired testes
  1. females usually only single ovary (left)
  a. Paired in many rapters, some pigeons, gulls, and passerines
  B. Endocrinology
  1. FSH - ganetes formation both male & female
  2. LH - steroid secretion both male & female
  a. testo - male secondary sex
  b. estrogen -important too
  II. 2 degree sex characteristics A. Influenced by T & E
  1. T causes starling bill to turn yellow
  2. E causes female quelins bill to turn red
  3. Wattles in male rooster- T
  B. Growth of coloful feathers by either sex- T
  1. Colorful female Phalerope > T than drubber male (polyandry)
  C. In weaver Finches (House Sparrows)
  1. Male plumage due to LH
  a. E in female inhibits LH effects
  D. Sperm production
  1. Seminal vesciles store sperm
  a. 4 degrees C cooler
  2. Most bird sperm approx. to mammalian sperm
  a. Oscine Passerines- sperm have spiral head and helical
  membrane or tail- corkscrew shaped(spin when swim)
  E. Copulation & Fertilization
  1. "Cloacal Kiss" few seconds
  2. Intromittent organ (Tinamous, Anseriforms, Currasow, Ostrich)
  a. Small penis in Galliforms & one passerine (White billed
  buffalo Weaver)
  E. Ovum maturation
  1. Yolk added in layers (count layers- yelloe day, white night=time)
  2. Mature ovum released for follicle- infurdibulin (24 hours) cloaca
  a. Spends 20 min infundib
  b. To magnum (3-4 hours)
  1. Albumen added (4 layers)
  c. To isth (1-2 hrs)
  1. Shell membranes
  d. to uteris (19-20 hrs)
  1. Shell added
  a. Pigments (porphyrinx mostly)
  3. Interval between eggs
  a. Usually 1-2 days
  1. One/day: passerines, hens, woodpeckers, grebes and small
  shorebirds
  2. Ratites & large rapters 3-5 days
  3. bookies and megapodes (7-8 dyas)
  4. Size
  a. Usually approx. to female body size
  1. Kuivi (500 g egg- 25% BW) Twoo eggs (4 week instead)
  F.
  1. Varies
  a. Many birds 3-5
  b. Larger birds usually fewer
  2. Determi??? vs. Indeterm
  1. Flicher (indeterm 71 eggs in 73 days0
  a. Domestic birds (one/day indefinitely)
  G. Egge Shapes
  1. Spherical
  a. Petrels, owls, kingfishers
  2. Ellipsoid (biconical)
  a. Grebes, pelican, bitterns
  3. Pyriform (roll in circles)
  a. Plovers, nurres
  4. Elongated
  a. Hummingbirds
  5. Conical
  a. Chickens, most passerines
  H. Egg shell color & texture
  1. Most hole, burrow nesters- white
  2 Most open nesters- colored & special patterns
  3. Some birds-blue eggs (Starling-cavity, Bluebird-cavity, Robin-cup)
  a. Ground-nesting thrushes- camouflaged, speckled eggs
  b. Tinamous- Brilliant blueish purple (very glossy- ceramic
  look)
  J. Eggshell structure
  1. Shell calcium and magnesium salts (carbonates & phosphates)
  embedded in a network of delicate collagenlike fibers
  a. Tonner based layer- adheres to shell membrane
  b. Palisade layer- makes up remainder of shell
  1. outer thin proteinaceous cullicle
  a. Fro of airbubbles that prevents microbial invasion
  c. Shelss have 10's of microscopic pores (gaseous exchange)
  1. Hen's egg (7500 pores)
  a. Most at blunt end
  2. Pores plugged or capped 9except columbiforms)
  3. Permit gaseous & water vapor exchange)
  d. Pestreeds thru egshells
  1. Esp. DDT & DDE (fat soluble)
  a. Affected brown Pelicans & Raptors
  
  #16 Nests & Incubation
  I. Intro
  A. All birds oviparous
  II. Nest architecture
  A. Most birds build isolated, hidden nest
  1. some colonial (2 -3 X 10 6 Quelas/ 100 hectares of thornbush savannah in Africa)
  B. Nest built to protect adults, their eggs,& hatchlings from predation
  and the elements
  C. Many types
  1. Single ground scrape (pebbles, stones)
  2. Cup nest most common (muds, sticks, grasses)
  3. Pendent nests most elaborate (orioles)
  D. materials
  1. Plant matter
  a. Twigs, grass, lichens, moss, and leaves
  b. Frsh green veg mostly by hole nesters
  1. Starlings select plant by orger
  a. Yarrow nettles and inhibits bacterial growth and
  the hatching of arthropod nest parasites
  1. Removal of plants - up blood sucking mites
  E. Nest safety
  1. Ground nesting most precarious
  a. In Oklahoma
  1. 70% of Mourning Dove ground nests destroyed (out of 130)
  2. 51% of Mourning Dove tree nests destroyed (out of 167)
  2. Cavity nesting safest
  a. Owls, Parrots, frodmouths, kestrels- natural cavities
  b. Kingfinches, petrels, tufted puffins, N. rough winged
  swallows- in ground
  c. trogons, tetmise, prothonotay, Warbler- soft, dead, rotten
  trees
  d. Woodpeckers- some even live tree (RCW -2 yrs to make) 80-100
  yr pines
  e. 2 degree cavity nesters compete intensivity for abandoned
  woodpecker holes
  3. some bird nest near large animal or insects for protection
  a. Starling, house Sparrows nest near Eagle aeries
  b. Eurasian Thick-knees nest near Crocidiles nests
  c. Banaquik (WEst Indies) nest near wasps
  d. In Central America, one parakeet nests only in termite mounds
  4. Some birds use displays to disuade predators
  a. Boken Wing display
  1. Killdeer & sandpipers
  b. Rodent-run display
  1. Low-running behavior (mouselike) to attract predator,
  then fly
  F. Colonial Nesting
  1. approx. 13% of birds, including most seabirds, nest in colonies
  III. Incubation
  A. Prl- incubation hormone
  B. Most birds delay incubation until multogy clutch is complete
  1. Owls & Hawks- incubate after 1st egg laid (asynchonous hatchlings)
  C. brood Patches
  1. Bare, Flacid skin areas on breast or abdomen- for heat transfer
  a. Single in most passerines & columbiforms
  b. Double in most shorebirds, larids and quail
  2. Columkids have natural apteria
  3. Others lose feathers to form brood patch
  4. Develop just before incubation (both sexes is both incubate)
  5. Not present in Brood Parasite (not produced by Prl & steroid
  injections)
  6. Also not present in Waterfowl, Penguins, & Pelicans
  a. Waterfowl line nest with breast feathers (acuter heat
  insualtion)
  1. G??, Boobies, Penguins and Murres use feet
  grasp eggs eggs on top
  D. Egg Temperature
  1. Optimum 37-38 C
  2. megapods (mount builder of Australia) 11 m mounds (composts)
  a. male tends mound alon 10-11 months
  1. Ta: 0-38 C
  2. Tm: 32-35 C
  E. Keeping eggs cool
  1. Eggs cooled by shorebirds (wet feathers)
  2. Incubating adults - part
  F. Turning eggs
  1. 15-25 % of daily activity
  a. Less than being active, However
  2. Birds abandon nest when foraging or body reserves dwindle
  a. kestrels & owls readily dessert eggs when hunting difficult
  3. Both sexes incubate in most birds
  a. Female alon approx. 25% families
  b. Male alone approx. 6% of families
  H. Incubation Period
  1. 10 days in woodpeckers, euchoos, small oscines to 80-90 days
  albatross & kiwis
  a. Short incubation- altricial young
  b. Longer incubations- preocial young
  
  
  #17 Mating
  I. Kinds of Pair bonds
  A. Monogomy (90%)
  1. main system
  B. Polygamy
  1. Multiple mates of opposite sex (approx.3%)
  C. Polygyny
  1. Male mates with 2 or more females (2%)
  D. Polyandry
  1. Female mates with 2 or more males (<1%)
  E. Polygynandry
  1. Both males and females with more than one mate
  a. Tinamous and Ratites (ostrich, rheas, Emu)
  b. Also Sith's Longspur
  c. Male Tatites & titamous incuabte mixed clutches of eggs form
  several females with different mates
  F. Promiscuity (approx 6%)
  1. Male hummingbirds, manakins, & grouse mat with any receptive
  female
  G. Sandpipers (Scolopacids) highly variable
  H. Sexual Selection
  1. Females select males
  a. Especially true on leks (communal display gound)
  2. Males often largers then females
  a. More vulnerable to short food supply
  3. Females bigger: shorebirds, raptors, & hummingbirds
  a. May be related male acrobatic aerial displays
  II. Monogamy
  A. Season or life
  1. Life: parrots, albatrosses, geese, eagles & pigeons (occass. divorce)
  a. Penguins - 16% separation of old pairs
  44% separation of young pairs
  1. increased divorce with increased breeding FAILURE
  2. Extra-Pair copulations
  a. Forced copulation (rape) - common among waterfowl!
  b. 30-40% Indigo Bunting chicks sired by extra-pair copulations
  1. All males in same breeding area sing exact same song!
  c. 50% extra-pair matings in Tree Swallows
  d. females of vasectomized male RWB's still show high fertility
  rates
  1. DNA fingerprinting shows that neighboring males fathered
  chicks
  B. Food abundance sometimes a factor
  1. Savannah Sparrows sometimes bigamous when food abundant
  
  
  III. Polygyny (~2%)
  A. 14 of 278 species of songbirds
  1. 11 of the species nest in marshes (female can feed on emirgent
  aquatic insects or grasses (also w/ many insects)
  B. Territory quality important
  1. Good territory - polygynous
  a. Bad " - often monogamous or unmated!
  2. Marsh Wrens
  a. % emergent marsh veg. important
  1. 50% - unmated
  2. 80% - monogamous
  3. 95% - polygynous
  3. Grass Lark Bunting (western US)
  a. Primary Female - aided by male
  b. Secondary " - raises young alone (but on good territory)
  
  IV. Polyandry
  A. females defend territory, compete for mates & lead courtship
  1. Females larger & brighter colored
  a. MALE incubates eggs and raises precocial young alone!
  
  
  V. Leks
  A. Promiscuous males display in courtship areana
  1. female visits to mate only
  a. build nest elsewhere and rasie precocial young alone
  2. Dominant males sire most offspring (80-90% of matings)
  a. Female Sage Grouse prefer males free of lice and malaria
  1. lice-infected grouse have RED spots on throat pouches
  B. Evolution of leks (why?)
  1. male flocking may reduce predation
  2. males gather at "hot spots" hwere most likely to encounter females
  a. Or females can best chose males if they can "see" all of them
  at the 'hot spots'
  1. mating in Ruff reflects both explainations.
  
  Skip #18
  
  #19 Parental Care
  I. Intro
  A. Peak breeding actively -50% up daily energy expenditures
  1. Most male raptors feed incubating females
  2. Most male passerines (211/250 species) help incubate or feed mates
  during incubations
  II. Intraspecific Brood Parasitism
  A. Most prevalent in waterfowl (Also in grebes, galligorms, larids,
  columbids, passerines and the ostrich)
  1. Most nests in some duck population contain eggs of other females
  a. When Wood Duck nest boxes close together (egg dumping) decreased
  success
  B. European Starlings often lay eggs in other's nests
  1. 25% of early nests have foreign eggs
  a. Female will discard if she has not begun to lay, but not so
  often she lays our first egg
  1. Roaming females often remove egg before laying her own
  a. Clutches with 6 egg - best success
  1. 7 egg clutches suffer losses- overcrowding
  hatchlings
  b. Since most females lay 5 eggs, can handle one extra
  egg
  1. Female might lay her 6th egg in another nest
  C. Black-billed & Yellow-billed Cuckoos will exhibit parasitism if
  abundant food supplies!
  III. Obligate Brood Parasitism (Cowbirds & European Cuckoos)
  A. Female Brown-headed Cowbirds lay 30-40 eggs/season (weekly set fo 2-5 eggs/week)
  1. Randomly laid; nost nests have one egg/nest (some up to 12 eggs)
  B. European Cuckoos lay 16-25 eggs/season (3-6/week)
  1. Only 1 egg/nest
  C. MImicry & other Adaptations of Brood Parasites
  1. Many are highly specialized for specific hosts
  2. Parasitic eggs usually thicker (can break host eggs when laid)
  a. usually ssme size as host (sometimes larger)
  3. Parasite chicks agressive
  a. European Cuckoos- "overboard reflex" (host eggs or young)
  b. African Honeyfuides- hooked bill to kill host's young
  c. Cowbirds- hatch earlier & bigger (get all food)
  4. Many have egg mimicry
  a. European Cuckoo eggs like hosts (differing egg colors/
  patterns in different parts of Europe) similar to regional hosts
  5. Some parasite hatchlings
  a. Same color as hosts
  b. Same mouth markings as host
  c. Some give begging calls of host
  6. How do you recognize own soecies? Genetics- inherited
  a. Isolated female B-hc raised in isolation respond to male
  songs and solicitcopulation with male
  b. Songs inherited in European Cuckoos too
  D. Effects of Brood Parasites on Hosts
  1. Rarely do hosts also fledge own young!
  2. Blackbird in Puerto Rico extinct on Island since Shiny Cowbirds
  expanded to island (only few Blackbirds left on remote island
  w/out Cowbirds
  3. B-hc's overwhelm populations of neotropical migrants
  a. Causes precipitious decline of Kirtlands Warbler (breeds only
  in MICHIGAN jack pine forests)
  1. Cowbirds now controlled
  a. Recent study with artificial bh cowbird eggs placed in RWB
  nests showed a 5-6 X decrease in cowbird parasitism!
  IV. Cooperative Breeding
  A. FLa Scrub Jays (altruism - placing good of species above own well-being)
  1. Social Units= breedign pair + up to 6 helpers
  2. "Helpers"- usually nonbreeding, young birds (usually help parents
  or step-parents)
  a. Help defend territory (most from snake predation)
  b. Feed & protect nestlings
  c. Helpers benefit too
  1. indirect benefits- kin selection
  B. Red-cockaded Woodpeckers too
  
  #20 Demography
  I. Intro A. Definition
  1. Study of Characteristics of population
  a. Age compostition
  b. Reproductive output
  c. Mortality
  II. Life History Patterns
  A. Patterns
  1. Mortality rates high in bird< one yr and decline to near constant
  lower rates amont\g adults
  2. Reproductive success and effort improve wtih age & experience
  3. Long-lived species have high survival rates(30% vs.15%), low
  fecundity (.2vs 3 young/yr), and low mortality rates (5% vs 50%
  year) than short-lived species
  4. Environments predictably impact life history pattern
  a. Clutch size large in north & arid, than south & wet
  environments
  B. Longevity
  1. Most small bird(2-5 years)
  2. Large birds(approx 20 years)
  3. Records
  a. 58 53 years albetross
  b. 36 years- oystercatcher
  c. 34 years- frigatebird
  4. Captive birds live longer (Parrot 80 years)
  
  

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• Last update: 6 June, 2006

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