Ornithology Lecture Notes
Evolution & Ornithology
Intro to Phylogenies
Phylogenies are trees that show hypothesized evolutionary relationships.
They illustrate ancestor-descendant relationships at various taxonomic levels: populations, species, and larger taxa.
Phylogenies are constructed using molecular, morphological, and/or behavioral data.
Alternative names: phylogenetic trees, cladograms, or simply "trees."
Key Phylogeny Terms:
Root: The common ancestor (CA) at the base of the tree.
Nodes: Represent ancestors at speciation events in the past.
Tips/Terminals: Represent the most recent taxa in the present.
Transitions:
Derived Characters: Newly evolved traits.
Evolutionary Reversals: Loss of traits or reduction in complexity.
Shared Transitions & Conserved Traits: Help determine relationships between taxa.
Outgroup: The earliest divergent taxon in the tree, sharing a common ancestor with the ingroup.
Ingroup: The taxa of focus, sharing a more recent common ancestor than the outgroup-ingroup ancestor.
Sister Taxa: Tips/terminals sharing an immediate common ancestor not shared by other tips/terminals.
Polytomy: A split of three or more lineages, indicating uncertainty in relationships.
Phylogeny Representation:
Phylogenies can be shown in multiple formats without altering the relationships.
Terminals can rotate on branches without changing relationships.
Removing taxa changes the relationships depicted.
Assembling Phylogenies via Parsimony:
Parsimony assumes that the tree with the fewest transitions is the most logical and reflects reality.
Example: Three species A, B, and C with ancestral characters (0,0,0) and derived characters (c, h, w).
Why phylogenies are hypotheses/estimates:
We are never 100% sure if we have all the data, and cannot observe the past.
Confounding factors:
Missing species (extinct and/or undiscovered).
Lack of characters.
Unique characters.
The common ancestor is hypothetical.
Homoplasies (similarities not due to shared ancestry).
Homoplasies:
Convergence: Similarities due to similar selection pressures, not shared ancestry.
Example: Endothermy in birds and mammals.
Reversal: Loss of a derived trait, resembling ancestral taxa.
Example: Flightlessness in ostriches and penguins.
Atavism: Reappearance of an ancestral trait after loss, usually due to mutations.
Example: Hoatzin's two-clawed wing vs. Archaeopteryx's three-clawed wing.
Phylogeny Terms:
Monophyly: A group with an ancestor and all its descendants.
Paraphyly: A group with an ancestor and some, but not all, of its descendants.
Polyphyly: A group with some, but not all, of its descendants, and no common ancestor.
Birds as Reptiles
Birds are vertebrates.
Birds are amniotes.
Birds are avian reptiles.
Reptilian Characteristics Shared by Birds:
Life Cycle: Direct development (no larval stage).
Amniotic Eggs:
Amniotic membranes for gas exchange, waste storage, and feeding.
Larger at hatching with longer incubation due to more yolk.
Calcareous-shelled eggs resist desiccation.
Integument:
Epidermal scales or scutes.
True claws.
Few integumentary glands.
Resistant to fluid, solute, and gas exchange.
Ecto/Poikilothermic (many are "Heliothermic").
12 pairs of cranial nerves.
Ribs encircle the body cavity.
Internal fertilization.
Reptiles vs. Other Reptiles (Birds and Mammals):
Reptiles are transitional and ancestral to birds and mammals.
Identifying unique reptilian characters can be difficult.
Birds: endothermic, feathered, bipedal with forelimb wings and pneumatic bones.
Mammals: endothermic with mammary glands and hair; more glandular skin; mostly give birth to live young.
Reptiliomorphs (ancestral to amniotes) diverged from amphibians ≈355 mya.
Sauropsid reptiles and synapsid reptiles (ancestral to mammals) diverged ≈320 mya.
Amniotes = Synapsids + Sauropsids
Early Evolution History of Reptiles
Archosauromorphs and Lepidosauromorphs derived from early reptiles and diverged ≈280 mya.
Extant Lepidosaurs include lizards, snakes, amphisbaenians, and tuataras.
Archosaurs and turtle ancestors derived from early Archosauromorphs and diverged ≈254 mya.
Archosaurs = The “Ruling Reptiles”
Birds as Dinosaurs
Hypotheses for Bird Divergence:
A: Birds diverged from Theropods ≈150 mya (late Jurassic).
B: Birds diverged from stem Thecodonts ≈230 mya (mid Triassic).
Stem Thecodonts = Ancestral Archosaurs (ancestors of dinosaurs, crocodilians, pterosaurs, etc.)
Archaeopteryx:
Discovered in 1861 (Jurassic period ≈150 mya).
First feathered dinosaur fossil discovered.
Shows clear link between Theropod Dinosaurs and Birds.
Provided major support for Darwin’s Descent with Modification.
Considered the first known bird, but not a “true bird” or a direct ancestor of modern birds.
Baminornis zhenghensis: Discovered 2023; Described 2025!
Dinosaur Phylogeny:
Historically, 3 major clades: Ornithischians sister to Saurischians, and Sauropods sister to Theropods.
Based mostly on hip structure.
Recent phylogenetic analysis finds all three arrangements statistically indistinguishable.
Despite the debate, birds are very much “settled in” Theropoda.
Derived Traits Inherited from Non-Avian Dinosaurs:
Lower body: Upright posture, hip structure, foot structure.
Reproduction: Hard-shelled amniotic eggs, egg brooding.
Flight Facilitation: Feathers, forelimb wing structure, furcula, hollow pneumatic bones, and possibly air sacs.
Traits Derived in Birds:
Axial Skeleton Modifications:
Toothless beaks
Loss of gastralia
Keeled sternum
Sternal ribs
Uncinate processes
Synsacrum
Reduced caudal vertebrae and pygostyle
Important Muscles of Respiration Associated with Vertebral Ribs:
Intercostal Muscles
Inhalation: contract to expand rib cage (pull rib cage upward and outward)
Exhalation: Relax to contract rib cage
Appendicocostal Muscles
Inhalation: contract to rotate vertebral ribs cranially via uncinate processes and sternum ventrally via sternal ribs Exhalation: relax
External Obliques
Inhalation: relax Exhalation: use uncinate process as brace while contracting to help reset action of appendicocostals; moves sternum dorsally
Appendicular Skeleton Modifications:
Reduced digits 1-3
Alula
Pubis angled parallel to ischium
Tibiotarsus
Reduced fibula
Tarsometatarsus
Adaptive Radiation of Birds
Adaptive Radiation of Dinosaurs
Adaptive Radiation of Bird-Like Dinosaurs and Birds
Dromaeosauria:
Non-Avian Theropod Outgroup, commonly called "raptors" (≈168-65 mya).
"Long-Tailed” Bird-Like Theropods
First known "birds", but not "true birds" nor direct ancestors of modern birds.
Lineages may represent the first and second outgroups to “true birds."
Derived Traits: unserrated teeth; fully reversed hallux, reduced caudal vertebrae
Examples: Archaeopteryx (≈ 155-150 mya) and Jeholornis(≈ 130-120 mya)
"True Bird” Lineages
Baminornis (?-150-? Mya)
Confuciusornis
Lineage may represent first outgroup to Ornithurines (more modern birds)
DT: no teeth; carpometacarpus; sternal ribs; posteroventral pubis; short tail w/ pygostyle
However, 3 wing digits/claws and gastralia still present; no keeled sternum
Enantiornithines
e.g., Sinornis
One of two major, highly-diverse (species & ecologies) bird sister groups during Cretaceous
Derived Traits: weakly keeled sternum, alula, and 2 wing claws (not 3)
Very similar appearance to sister Ornithurines, but still had teeth, gastralia, and wing claws
Died out 65 mya; survived by sister Ornithurines
Hesperornithines & Icthyornithines
Two aquatic, cormorant-like lineages that existed during Cretaceous
Derived Traits: uncinate processes and no wing claws; Hesperornithines had reduced traits
However, still had teeth.
Died out 65 mya (possibly earlier); survived by Neornithines(AKA modern Avians/Birds)
Adaptive Radiation of Birds:
Birds may have diverged as early as ≈ 172-164 mya (mid Jurassic) and radiated throughout Jurassic.
Birds diversified even more during Cretaceous, way before Cretaceous extinction!
Extant Birds:
Two extant infraclasses possibly diverged as early as ≈ 120-110 mya (≈ early Cretaceous) or as late as ≈ 95-90 mya (≈ mid-late Cretaceous).
Palaeognathae (Ratites: ≈ 0.5% of extant species)
Neognathae (All other extant birds: ≈ 95.5% of extant species)
Galloanserae and Neoaves possibly diverged as early as ≈ 105-100 mya (mid-late Cretaceous) or as late as 85 mya (late Creataceous).
Galloanserae (All land and water fowl: ≈ 4.5% of extant species)
Neoaves (All other extant birds: ≈ 95% of extant species)
Key Points:
Modern birds were contemporaries of non-avian dinosaurs!
No “March of Progress”, just adaptive radiation of dinosaur groups!
Birds are dinosaurs!
Modern birds survived Cretaceous Extinction!
Diversified rapidly after Cretaceous Extinction and even more throughout Cenozoic!
Dinosaurs are not extinct; they’re still perhaps the most successful group of tetrapods ever! We just call them birds now…