Invertebrate species evolution, taxonomy and

phylogeny
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• Introduction of evolutionary theory into animal

taxonomy
– Changed taxonomist’s role from one of classification to
systematization

• Classification
– Denotes the construction of classes
• Grouping or organisms that possess a common feature
called an essence
– Used to define the class
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• Systematization
– Places groups of species into units of common evolutionary
descent
• Character variation
– Used to diagnose systems of common descent
– No requirement that an essential character be
maintained throughout the system for its
recognition as a taxon
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• In classification
– Taxonomist asks whether a species being classified
contains the defining feature of a particular
taxonomic class
• In systematization
– Taxonomist asks whether the characteristics of a
species confirm or reject the hypothesis that it
descends from the most recent common ancestor
of a particular taxon
 Species We lead

• Biologists have repeatedly used certain criteria for

identifying species

– Common Descent
• Central to nearly all modern concepts of
species
• Smallest distinct groupings of organisms sharing
patterns of ancestry and descent
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– Reproductive Community
• Member of a species must form a reproductive
community that excludes members of other
species
– Sexually reproducing populations
» Interbreeding is critical for maintaining a
reproductive community
– Asexually reproducing populations
» Entails occupation of a particular ecological
habitat in a particular place so that a
reproducing population responds as a unit to
evolutionary forces.
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• Any species has a distribution through space and

distribution through time
– Geographic range
• Species having very large geographic ranges or
worldwide distributions
– Cosmopolitan
• Species with very restricted geographic distributions
– Endemic
– Evolutionary duration
• Distribution through time
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• Typological Species Concept

– Before Darwin, a species was considered a distinct
and immutable entity
• A type specimen was labeled and deposited in a
museum
– Represented the ideal form of a specimen
• Small differences from the type
– Considered accidental imperfections
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– Today, organismal morphology still important in

recognizing species
– However, species no longer viewed as classes of
organisms defined by possession of certain
morphological features
– Type specimens serves only as a guide to general
morphological features that one may expect to
find in a particular species
– Variation is not viewed as an imperfect
manifestation of an eternal “type”
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• Biological Species Concept

– Proposed by Theodosius Dobzhansky and Ernst

Mayr
– Been refined and reworded several times
• A species is a reproductive community of
populations (reproductively isolated from
others) that occupies a specific niche in nature.
– Ability to successfully interbreed is central to the
concept
– Criteria of “niche” tie in ecological properties
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– Sometimes species status can be evaluated

directly by conducting breeding experiments
• However, controlled breeding experiments can be
difficult to conduct
– Molecular and other studies may detect sibling
species
• Species too similar in morphology to be diagnosed as
separate species by morphological characters alone
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– Biological species concept

• Lacks an explicit temporal dimension
• Gives little guidance regarding the species
status of ancestral populations relative to their
evolutionary descendants
– Proponents of the biological species concept
• Disagree on the degree of reproductive
isolation necessary for considering two
populations separate species
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• The biological species concept has received

strong criticism
– A species has limits in space and time
• Boundaries between species may be difficult to locate.
• Interbreeding is not an operational definition in asexual
organisms
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• Evolutionary Species Concept

– Simpson proposed the evolutionary species

concept in the 1940s
• Concept persists but with modification
– An evolutionary species
• A single lineage of ancestor-descendant populations
that maintains its identity from other such lineages and
that has its own evolutionary tendencies and historical
fate.
– Definition accommodates both sexual and asexual
forms as well as fossils
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• Phylogenetic Species Concept

– Phylogenetic species
• An irreducible (basal) grouping of organisms
diagnosably distinct from other such groupings and
within which there is a parental pattern of ancestry and
descent.
– Both asexual and sexual groups are covered
– Phylogenetic species is a single population lineage
with no detectable branching
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– Main difference in practice between the

evolutionary and phylogenetic species concepts
• The latter emphasizes recognizing as separate
species the smallest groupings of organisms
that have undergone independent evolutionary
change
• Discerns the greatest number of species but
may be impractical.
• Disregards details of evolutionary process
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• Dynamism of Species Concepts

– Disagreement is a sign of dynamic research
• Should not be considered discouraging
– Cannot predict which species concepts will remain
useful in the future
– Possible that disagreements regarding species
boundaries may identify interesting cases of
evolution in action
– No one concept is comprehensive or final;
• All need to be understood to understand future
concepts
Taxonomic Characters and Phylogenetic Reconstruction
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• Major goal of systematics

– Infer the evolutionary tree or phylogeny that
relates all extant and extinct species
– Accomplished by identifying organismal features
called characters that vary among species
– Characters are any feature used to study variation
within and among species
• Identified by observing patterns of similarity of
morphological, chromosomal, and molecular features
• Less frequently, behavioral and ecological features are
used
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• Homology
– Character similarity resulting from common
ancestry
• Similarity does not always reflect common
ancestry
• Homoplasy
– Character similarity that misrepresents common
descent
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Using Character Variation to Reconstruct

Phylogeny
• First step is to determine which variant form of each
character was present in the common ancestor of
the entire group
• The character state present in the common ancestor
– Ancestral character
• All other variant forms of the character arose later
within the group
– Evolutionarily derived character
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• Polarity of a character
– Refers to identifying which one of its contrasting states is
ancestral and which one(s) is derived
• Outgroup comparison
– Method used to examine the polarity of a variable
character
– Outgroup
• Group that is phylogenetically close but not within the
group being studied
• Infer that any character state found both within the
group being studied and the outgroup is ancestral for
the study group
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• Organisms or species that share derived character

states
– Form subsets within the study group called a clade
• Derived character shared by members of a clade
– Synapomorphy of that clade
• Synapomorphies are used as evidence of homology
– Infers that a particular group of organisms forms a clade
• A clade corresponds to a unit of evolutionary
common descent
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• Includes all descendants of a particular ancestral

linage
• Pattern of derived states of characters within study
group
– Forms a nested hierarchy of clades within clades
• By identifying the nested hierarchy of clades or
branches
– Patterns of common descent can be formed
• Character states ancestral for a taxon are
plesiomorphic for that taxon
– Sharing of ancestral states is termed
symplesiomorphy
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• Identifying the level at which a character state is a

synapomorphy may identify a clade
• Cladogram
– Nested hierarchy of clades
• To construct a phylogenetic tree
– Additional information concerning ancestors, duration of
lineages, and amount of evolutionary change must be
included
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• Sources of Phylogenetic Information

– Comparative Morphology
• Examines shapes, sizes and development of organisms
• Skull bones, limb bones, scales, hair and feathers
• Living specimens and fossils are used
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– Comparative Biochemistry
• Analyzes sequences of amino acids in proteins and
nucleotides sequences in nucleic acids
• Recent studies show comparative biochemistry can be
applied to fossils

– Comparative Cytology
• Examines variation in number, shape and size of
chromosomes
• Used almost exclusively on living specimens
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 Theories of Taxonomy We lead

• Two currently popular theories of taxonomy

– Traditional Evolutionary Taxonomy
– Phylogenetic Systematics (cladistics)
• A relationship between a taxonomic group and a
phylogenetic tree or cladogram important in both
theories Relationship can take on one of three forms
– Monophyly
• A monophyletic taxon includes the most recent
common ancestor and all descendants of that ancestor
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– Paraphyly
• A taxon is paraphyletic if it includes the most recent
common ancestor of all members of a group and some
but not all descendants of that ancestor
– Polyphyly
• A taxon is polyphyletic if it does not include the most
recent common ancestor of all members of a group
• The group has at least two separate evolutionary
origins
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 Theories of Taxonomy We lead

• Both evolutionary and cladistic taxonomy

– Accepts monophyletic
– Rejects polyphyletic groups
– Differ on accepting paraphyletic groups.
– Difference has important evolutionary implications
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• Traditional Evolutionary Taxonomy

• The two main principles
– Common descent
– Amount of adaptive evolutionary change
• A particular branch on an evolutionary tree
– Considered a higher taxon if it represents a distinct
adaptive zone
• Distinct “way of life”
– A taxon that represents an adaptive zone is a grade
– Evolutionary taxa may be either monophyletic or
paraphyletic
– By accommodating adaptive zones, nomenclature
reflecting common descent is not as clear
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• Phylogenetic Systematics/Cladistics
– Challenge to evolutionary taxonomy
– Willi Hennig (1950)
• First proposed cladistics or phylogenetic
systematics
• Emphasizes common descent and cladograms
– Cladists avoid paraphyletic groups by defining a
long list of sister groups to each more inclusive
taxon.
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• Current State of Animal Taxonomy

– Modern animal taxonomy
• Established using evolutionary systematics and recent
cladistic revisions
– Phylocode
• New taxonomic system
• Being developed as an alternative to Linnean taxonomy
• Replaces Linnean ranks with codes that denote the
nested hierarchy of monophyletic groups conveyed by
cladograms
– The terms “primitive,” “advanced,” “specialized”
and “generalized”
• Used for specific
 Major Division of Life We lead

• Aristotle’s two kingdom system

– Included plants and animals
– One-celled organisms became a problem
• Haeckel (1866)
– Proposed Protista for single-celled organisms
• R.H. Whittaker (1969)
– Proposed a five-kingdom system to distinguish prokaryotes
and fungi
• Woese, Kandler and Wheelis
– Proposed three monophyletic domains above kingdom
level—Eucarya, Bacteria and Archaea—based on
ribosomal RNA sequences.
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• More revisions are necessary to clarify taxonomic

kingdoms based on monophyly
• “Protozoa”
– Neither animals nor a valid monophyletic taxon
• “Protista”
– Not a monophyletic kingdom
– Most likely composed of seven or more phyla
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Major Subdivisions of the Animal Kingdom
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• Traditional groupings based on embryological and

anatomical characters

• Branch A (Mesozoa): phylum Mesozoa,

the mesozoa
• Branch B (Parazoa): phylum Porifera,
the sponges and
phylum Placozoa
• Branch C (Eumetazoa): all other phyla
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– Grade I (Radiata): phyla Cnidaria, Ctenophora

– Grade II (Bilateria): all other phyla
• Division A (Protostomia):
– Acoelomates: phyla Platyhelminthes,
Gnathostomulida, Nemertea
– Pseudocoelomates: phyla Rotifera, Gastrotricha,
Kinorhyncha, Nematoda, Nematomorpha,
Acanthocephala, Entoprocta, Priapulida, Loricifera
– Eucoelomates: phyla Mollusca, Annelida,
Arthropoda, Echiurida, Sipunculida, Tardigrada,
Onychophora
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• Division B (Deuterostomia):
phyla Phoronida,
Ectoprocta,Chaetognatha, Brachiopoda,
Echinodermata, Hemichordata, Chordata
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• Recent molecular phylogenetic studies have challenged

traditional classification of Bilateria
– Grade II: Bilateria
• Division A: (Protostomia):
– Lophotrochozoa: phyla platyhelminthes, Nemertea, Rotifera,
Gastrotricha, Acanthocephala, Mollusca, Annelida, Echiurida,
Sipunculida, Phoronida, Ectoprocta, Entoprocta,
Gnathostomulida, Chaetognatha, Brachiopoda
– Ecdysozoa: phyla Kinorhyncha, Nematoda, Nematomorpha,
Priapulida, Arthropoda, Tardigrada, Onychophora, Loricifera
• Division B (Deuterostomia):
– phyla Chordata, Hemichordata, Echinodermata
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Thank you