Extant Sauropsida birds (finally) lilac breasted roller, Coracias caudatus Photo: AMMurray Learning objective 2: to be able to recognize and describe different types of feathers, and understand how contour feathers and other anatomical adaptations affect bird flight. Amniota Tetrapoda (land vertebrates) Sarcopterygii (lobe-finned “fish”) Osteichthyes (bony “fish”) Gnathostomata (jawed vertebrates) Vertebrata Mammalia Reptilia Lissamphibia (modern amphibians) Dipnoi (lungfish) Actinistia (coelacanths) Actinopterygii (ray-finned fish) Chondrichthyes (cartilaginous fish) Cyclostomata (jawless fish) Reptile Phylogeny Archosauria Aves (birds) Crocodylia Reptilia Testudinata (turtles) Sphenodon (tuatara) Lepidosauria NB: The avian crown group is sometimes called Neornithes, but we’ll use Aves in ZOOL 224 (as in text book). Squamata (lizards) Aves • birds and their extinct relatives • extant are immediately recognizable BUT: – most ‘bird’ features evolved prior to bird lineage • feathers, furcula, pneumatic bones, unique wrist joint, uncinated processes photos: AMMurray feathers and flight • evolved in non-avian dinosaurs • inherited by their descendants (birds) • (videos on bird origins and origin of flight) Acheroraptor Emily Willoughby, Own work, CC BY 3.0, https://commons.wikimedia.org/w/index.php?curid=30194091 Artist's concept of Caihong, a crow-sized iridescent dinosaur from the Jurassic Period. smithsonianmag.com; Velizar Simeonovski/The Field Museum origin of feathers • (from the video with Dr. Julia Clark) – evolved first in non-avian dinosaurs – after feathers evolved, were ‘co-opted’ for flight = exaptation • why did they evolve? – insulation – gliding – thermoregulation (insulation and shading) anatomy of a feather • develop from feather follicles = infoldings of surface layer of skin • each has its own blood supply, muscles and nerves • predominantly formed of keratin • calamus anchors feather to body within the sheath follicle calamus veterinkey.com/feather-follicle-extirpation anatomy of a feather • rachis = central axis • from largest to smallest branching: rachis, barb, barbules • barbs branch off from the rachis, and the barbules are what hold the barbs of the feather together to provide a smooth outline https://academy.allaboutbirds.org/feathers-article/ pterylae • feathers are laid out on the body in distinct tracts = pterylae – some birds lack pterylae and feathers are uniformly distributed over body www.zoo.ufl.edu/courses/vertzoo/Images/Birdlab/FeatherTrack.jpg Nestling Common Grackle showing pterylae photo: Caleb G. Putnam types of feathers classified based on morphology: • plumulaceous – no vane, rudimentary rachis, elongate barbules on tuft of barbs • pennaceous – centra rachis with hooked barbs to form vane with smooth edge Prum and Brush 2003 types of feathers • classify based on function (5 types) – contour – down – semiplumes – filoplumes – bristles Wing and tail feathers are pennaceous, and down feathers are plumulaceous. Contour and semiplume feathers are plumulaceous at their bases, and pennaceous at their tips. The filoplume has barbs at its tip, and the bristle has barbs at its base https://academy.allaboutbirds.org/feathers-article feather functions • contour: cover surface of bird in overlapping layers – on the body = coverts or deck feathers – specifically for flight = flight feathers – on the wing = remex (pl. remiges) https://www.fws.gov/lab/feat heratlas/idtool.php • asymmetrical to prevent twisting • remiges on outer tip of wing = primaries; provide thrust • remiges on trailing edge of wing = secondairies; form airfoil – on the tail = retrix (pl. retrices) • generally arranged in a fan shape; steering • normally 6 pairs, more asymmetrical towards outer edges of tail feather functions • down: thermal insulation – small feathers with short rachis – lie closest to skin – no barbules, entirely plumulaceous in form By Wouter Hagens - Own work, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?cur id=103988002 feather functions • semiplume: thermoregulation and streamlining – – – – between down and contour in morphology lie underneath contour feathers long rachis but no vane no barbules (entirely plumulaceous) study.com feather functions • filoplume: sensory – – – – short, simple with a few barbs at the tip base is associated with nervous system underlie contour feathers, mostly on the wings nerves sense movement of filoplume and provide information to brain about position and movement of overlying contour feathers http://people.eku.edu/ritchisong/feathers.html feather functions • bristle: sensory (tactile) and protection – simplest feathers – central rachis with no or few barbs at base – concentrated at the base of the beak and around eyes Eastern Whip-poor-will at Magee Marsh Wildlife Area, Ohio, May 3, 2015, by Joan Tisdale; https://www.birdwatchingdaily.com/news/science/bird-basics-six-different-feather-types-explained/ feather colour • from pigments Plum-headed Parakeet ebird.org – melanins in skin cells – blacks, greys, browns – carotenoids from diet – reds, oranges, yellows – porphyrins containing metals – e.g., iron red • from structure – scattering of light by cells of barbs – blues, greens, purples • mix of two – e.g., parakeet green from yellow carotenoid mixed with structural blue bird aerial locomotion • • • • very complex! gliding soaring wing flapping But well explained in section 22.2 Wings and Flight (pp.404–407) of the text book. www.liveanimalslist.com principles for flight • thrust must equal drag, or a flier will slow down • lift must equal weight, or a flier will sink • Bernoulli’s principle: for a wing with a convex upper surface, air must move faster above the wing than below, leading to greater pressure below the wing. https://www.flight-mechanic.com/aircraft-theory-of-flight/ Gliding – relatively simple • requires an initial movement (e.g., jump) to establish a forward trajectory • wing then produces lift by a combination of downwash (if tilted) and Bernoulli’s principle (if the upper surface is convex) – The wing can never produce quite enough lift to equal weight, so the animal slowly sinks downwash = force that creates lift Gliding and Soaring • gliders can maintain constant speed while slowly sinking – wing shape and susceptibility to drag determine ratio of lift to drag forces, which dictates the angle of descent (glide angle) – low glide angle = can travel a long distance before contacting the ground – albatross released at an altitude of 1 km could glide ~20 km before contact! • soaring is when gliders regain altitude by exploiting air movement, e.g., updrafts caused by air rising in the heat of the morning sun – used by many large vultures Wandering albatross (Diomedea) www.dw.com Turkey vulture (Cathartes) nature.mdc.mo.gov Avian Flapping • must produce thrust and lift with muscular activity • thrust produced by movement of the outer part of the wing, with the longest feathers (primaries) • lift produced by the inner part of the wing (secondaries) acting as an airfoil (as in gliding, but aided by the speed resulting from the extra thrust) • flapping movements are complex and adjustable, and cause the flight feathers of the outer part of the wing to twist along their length and act analogously to propellers Bushtit (Psaltriparus) Avian flapping flight - vortices • major role in flight – for the individual – create lift, but also drag – for the group – leading bird’s vortices provide lift to following birds snow geese; birdwatching.com theguardian.com flight muscles • downstroke – pectoral major • upstroke – supracoracoideus muscle • both insert on the sternum – these two muscles work in opposite directions, so why do they insert ventrally on the same bone? – ventral mass stabilizes centre of gravity https://www.youtube.com/watch?v=aFdvkopOmw0 flight specializations • larger/heavier birds need longer runs to takeoff • landing is tricky! must reduce velocity but only ‘stall’ at landing – increase pitch of wing, spread tail to increase drag – alula – feather-covered second digit is extended to create lift https://www.featheredphotography.com/blog/ Other Factors in Flight Performance • wing loading is mass divided by wing area (area of wings and body between) – If too large, flight is impossible • wing aspect ratio is ratio of wing length to wing width; a high aspect ratio (long, narrow wing) has a high lift : drag ratio • camber is wing curvature, and contributes to production of lift • slots between outer wing feathers allow each to act as an independent airfoil • different combinations of these factors are suitable for different types of flight wing types in lab Chiappe et al. 2014, PeerJ 2:e234