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Biology · Phylogeny, Classification & the Tree of Life

Homology, convergent similarity and character evidence

A bird and a bat both fly. A bird and a human both have forelimbs with corresponding bones. Which resemblance should guide a tree? First specify the feature and the historical claim: the answer need not be the same for the whole limb and its use in flight.

Heredity, Variation & Evolution introduces inherited differences and common ancestry; Evolutionary Processes & Adaptation distinguishes observations from evolutionary mechanisms. Species, Speciation & Macroevolution provides useful context for species concepts and reticulate histories, though its detailed cases are optional here. You need to read small tables, add branch values and compare fractions. Each tree and probability rule is explained before it is used; no programming or previous taxonomic vocabulary is assumed.

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By the end, you can

Define the character being compared and distinguish shared ancestry from independently acquired similarity.

01

A resemblance needs a precise comparison

A character is a feature selected for comparison; character states are its scored versions. “A limb element is present” is a different character from “powered flight is present.” Anatomy, behaviour, development and sequence positions can all provide characters, but each comparison needs an operational description.

Homology describes correspondence through common ancestry. Vertebrate forelimbs can retain corresponding skeletal elements despite differing shapes and functions. Our schematic marks a humerus connected with a radius and ulna in two modified limbs. It shows those relationships rather than actual bone proportions or every joint.

Bird and bat forelimbs have this deeper shared history, while their modifications for powered flight arose independently in the two lineages. The flight resemblance is therefore convergent at that level of comparison. Calling the limbs homologous does not make every detail of the wings a single ancestral invention. Conversely, convergence in flight does not erase the inherited forelimb relationships.

Researchers evaluate a proposed correspondence using anatomical position and connections, development, fossils and molecular evidence. Similar performance alone is insufficient. A character’s historical interpretation is a hypothesis that can change when better evidence shows a mistaken correspondence or a different origin.

Inherited limb organisation and independently modified flight surfaces

Inherited limb organisation and independently modified flight surfacesOriginal schematic limb-element connections are paired with distinct feather and membrane surface symbols to identify what is being compared.Bird forelimbHumerusRadiusUlnaFeather-based flight surface; simplifiedBat forelimbHumerusRadiusUlnaMembrane flight surface; hand supports omittedCorresponding skeletal elementsInherited humeruscorrespondenceBird forelimb: Humerus ↔ Batforelimb: HumerusInherited radius correspondenceBird forelimb: Radius ↔ Batforelimb: RadiusInherited ulna correspondenceBird forelimb: Ulna ↔ Batforelimb: UlnaCorresponding skeletal elementsPowered-flight modifications evolved independentlyElement map, not an anatomical scale drawingConceptual bird/bat forelimb correspondence only. No actualproportions, complete digit counts, joint mechanics or ancestral wingreconstruction.
Matching labels identify corresponding limb elements. The membrane polygon and five feather symbols illustrate flight surfaces; they do not give real dimensions, feather counts or digit counts.
Read the diagram in words

Humerus, radius and ulna correspond in inherited forelimb organisation; simplified feather versus membrane symbols identify distinct wing modifications.

Bird forelimb

  • Humerus
  • Radius
  • Ulna
Bird forelimb
DescriptionRelationship
Humerus → RadiusCorresponding skeletal elements
Humerus → UlnaCorresponding skeletal elements

Feather-based flight surface; simplified

Count: 5

Bat forelimb

  • Humerus
  • Radius
  • Ulna
Bat forelimb
DescriptionRelationship
Humerus → RadiusCorresponding skeletal elements
Humerus → UlnaCorresponding skeletal elements

Membrane flight surface; hand supports omitted

Corresponding skeletal elements
Bird forelimbBat forelimbRelationship
HumerusHumerusInherited humerus correspondence
RadiusRadiusInherited radius correspondence
UlnaUlnaInherited ulna correspondence

Conceptual bird/bat forelimb correspondence only. No actual proportions, complete digit counts, joint mechanics or ancestral wing reconstruction.

02

A shared state can have more than one history

A state inherited from a distant ancestor may occur in several branches without identifying one recent subgroup. A shared derived state is a change inferred to have arisen in the relevant common ancestor and then inherited by its descendants. “Derived” is relative to an ancestral comparison, not a word for better or more complex.

Convergence, parallel changes and reversals can produce homoplasy: similar scored states that do not reflect one origin in the ancestor being used to group those taxa. A trait can also be lost. Whether two presences support a particular clade depends on the proposed tree, ancestral-state reasoning and plausible alternatives.

The invented table has five sampled taxa, O, A, B, C and D. O is a stipulated outgroup for the example. Character X has state 0 in O/A/B and 1 in C/D. On the assigned tree, a single 0-to-1 change on the C/D stem explains X. Character Y has state 1 in A/C but 0 in O/B/D. On that same tree, two independent gains explain Y, and alternative histories may include gain and reversal.

This model makes the difference visible; it does not establish that every binary character changes independently, at equal cost, or only once. The observed table contains tip states. The internal changes are candidate reconstructions attached to an assumed history.

Observed states and candidate origins

Observed states and candidate originsA scored character table sits beside a stipulated rooted tree with separate X and Y change overlays.Observed tip statesCharacter XCharacter YO00A01B00C11D10One X gain on C/D stemFocalancestorA/B ancestorC/D ancestorAssignedoutgroup OEarlierancestorTaxon ATaxon BTaxon CTaxon DOne possible two-gain Y historyFocalancestorA/B ancestorC/D ancestorAssignedoutgroup OEarlierancestorTaxon ATaxon BTaxon CTaxon DUnobservable: unknown, not absenceC / YUnknown (?)Observed tip statesAssigned-tree change mappingUnknown (?)Invented binary states and supplied outgroup/root. Equal cost foreither transition; overlays are possible histories, not measured originsor unique reconstructions.
Table cells are observations, while edge marks represent one inferred history. An unknown score is missing information, not observed absence. The rooted ancestry is assumed for this example, rather than established by the table alone.
Read the diagram in words

X=1 in C/D fits one stem gain; Y=1 in A/C fits two separate gains on this supplied tree. The alternate unobservable C/Y cell is unknown, not 0.

Observed tip states
DescriptionCharacter XCharacter Y
O00
A01
B00
C11
D10
Assigned-tree change mapping
DescriptionRelationship
Earlier ancestor → Assigned outgroup ODescent
Earlier ancestor → Focal ancestorDescent
A/B ancestor → Taxon ADescent
A/B ancestor → Taxon BDescent
C/D ancestor → Taxon CDescent
C/D ancestor → Taxon DDescent
Focal ancestor → A/B ancestorDescent
Focal ancestor → C/D ancestorDescent
One X gain on C/D stem
DescriptionAssigned state transition
Focal ancestor → C/D ancestor0 → 1; X: 0 to 1
One possible two-gain Y history
DescriptionAssigned state transition
A/B ancestor → Taxon A0 → 1; Y: 0 to 1
C/D ancestor → Taxon C0 → 1; Y: 0 to 1
Unobservable: unknown, not absence
DescriptionUnknown (?)
C / Y?

Invented binary states and supplied outgroup/root. Equal cost for either transition; overlays are possible histories, not measured origins or unique reconstructions.

03

Count evidence rather than repeated descriptions

Several measurements of one tightly linked structure may record the same underlying change. Treating them as fully independent votes can overstate the evidence. Likewise, adjacent sites, duplicated copies and selected gene regions can have relationships that a simple independent-character model ignores.

Fossils can provide anatomical combinations and time constraints even when recoverable DNA is absent. Missing information should remain missing rather than being scored as absence. An incomplete specimen cannot be said to lack a bone simply because the relevant region was not preserved.

Useful comparisons make their sampling and scoring choices visible. Researchers can examine whether conclusions survive alternative character definitions, added taxa and different models. A tree supported by a broad, well-characterised comparison is more informative than a grouping chosen because one familiar resemblance fits it.

Worked example

Locate evidence before claiming an origin

On the assigned rooted tree, O branches before the focal A/B and C/D lineages. X = 1 appears in C and D, whose shared stem can carry one gain. If you group all Y = 1 taxa instead, you select A and C. Their common ancestor in this tree also has B and D as descendants, so the selected pair is not a clade.

Two Y gains, one on A’s terminal branch and one on C’s, fit the scored observations. This demonstrates a possible convergent history under the assigned tree and equal-cost binary rule. It does not prove that the real history of an unknown trait contained exactly those two events.

Now suppose C’s Y state could not be observed. Record it as unknown, not 0. The available observations would no longer justify the same complete mapping. Missing data and a measured absence are different evidence, even when a blank table cell makes them look similar.

Check your understanding

  1. Can bird and bat wings be homologous in one comparison and convergent in another?

    Show reasoning

    Yes. Their underlying vertebrate forelimbs include inherited corresponding structures, while modifications for powered flight arose independently. State whether you mean the forelimb organisation, a specific component or the flight adaptation.

  2. Does state X = 1 in C/D prove that 1 is the ancestral state for all five taxa?

    Show reasoning

    No. In the assigned example the outgroup and A/B have 0, and a gain on the C/D stem explains 1. Ancestral-state inference depends on the tree, comparison and change model; “present in two taxa” does not settle it.

  3. A fossil preserves no wrist region. Why is scoring “wrist element absent” unsafe?

    Show reasoning

    The element might simply be unobservable because that region is missing. Score unknown unless preservation or other evidence permits a reliable absence claim. Converting unknowns to absences invents character evidence.

What to carry forward

Specify the character, test its correspondence and keep observed states separate from reconstructed changes.

References

Resources supporting this lesson’s explanations and examples.

  1. Developmental homoplasy: convergence in cellular differentiation (Journal of Anatomy, 2010)Anthony GrahamAuthor research review — selected textpmc.ncbi.nlm.nih.govRead source
  2. The Tree Room — Reading trees and comparing evidence (Live teaching text; inspected October 2026)Understanding Evolution team · UC Museum of Paleontology, University of California, BerkeleyInstitutional scientific educationThe Tree Room — Reading trees and comparing evidence (Live teaching text; inspected October 2026)Common ancestry and relationshipsTip order, rotation and progress misconceptionsComparing charactersParsimony and its limitsResolution and branch supportevolution.berkeley.edu
  3. Foundational likelihood and bootstrap methods (Journal of Molecular Evolution 17:368–376, 1981; Evolution 39:783–791, 1985)Joseph FelsensteinResearch abstractEvolutionary trees from DNA sequences: a maximum likelihood approachConfidence limits on phylogenies: an approach using the bootstrappubmed.ncbi.nlm.nih.gov