CHAPTER 7

Animal Taxonomy, Phylogeny, and Organization

TAXONOMY AND PHYLOGENY

 Zoologists' first question upon encountering a new species:

 “What is it?”
 This question reflects curiosity about both the species' identity and its
evolutionary history.

 Understanding a species involves:

 Not just naming it, but understanding its evolutionary history.

 Evolutionary processes have created order that is reflected in phylogenetic
relationships.

 Systematics and Taxonomy:

 The study of organisms' kinds, diversity, and evolutionary relationships is called

systematics or taxonomy.
 Systematics comes from Greek systema (system) and ikos (body of facts).
 Taxonomy comes from Greek taxis (arrangement) and Latin nominalis
(belonging to a name).

 Role of taxonomists:

 Describing new species and organizing animals into groups (taxa) based on
evolutionary relatedness.
 Identifying and describing inherited characteristics that animals share.
 Ranking these characteristics based on their evolutionary order of
appearance.

 Phylogeny:

 The evolutionary history of a group of organisms.

 Phylogenies are usually depicted using tree diagrams, which will be explored
later in the chapter.

A Taxonomic Hierarchy

 Modern Taxonomy and Karl von Linné (Carolus Linnaeus):

 Rooted in the work of Karl von Linné (1707–1778).

 Linnaeus' binomial system (two-part naming system) is still in use today.

 Grouping based on shared characteristics:

 Species can be grouped into broader categories based on common traits.

 Example: Housefly (Musca domestica) shares characteristics with other flies
(e.g., single pair of wings) and forms a taxon with other flies.
  Flies, bees, butterflies, and beetles share traits and form a broader taxon:
insects.

 Taxonomic Ranks:

 Animals are classified using eight taxonomic ranks in a hierarchical order

(from broad to specific):
1. Domain
2. Kingdom
3. Phylum
4. Class
5. Order
6. Family
7. Genus
8. Species
 As you move down the hierarchy from domain to species, the groups become
smaller and more closely related.

 Subdividing taxonomic ranks:

 Taxonomists may subdivide ranks (e.g., subphylum, superclass, infraclass) to

express relationships between ranks.

 Limitations of the Linnaean Taxonomic Hierarchy:

 The Linnaean hierarchy has some limitations for modern biology:

o No clear definition for ranks above species (e.g., what exactly defines a
family).
o Example: The cat family (Felidae) has 40 species, while the ground
beetle family (Carabidae) has over 40,000 species.
 There's no clear way to compare the level of divergence between such vastly
different families.

 Phylogenies and Modern Methods:

 The Linnaean ranking system doesn't always accurately reflect the

phylogenetic relationships revealed by modern methods.
 Phylogenetic relationships are sometimes depicted without traditional Linnaean
ranks.
 Phylogenies are considered hypotheses that are continually tested and
revised.
 The precision and accuracy of phylogenetic hypotheses are more important than
adhering to traditional ranking systems.

Nomenclature

 Common Names and Their Issues:

 Common names vary from country to country and region to region within a
country.
 A species can have many different common names (e.g., crawdads, crayfish,
crawfish for freshwater crustaceans).
  Common names can refer to taxonomic categories higher than the species level,
making it hard to distinguish between different species based on just the name.
 Example: Pillbugs (class Crustacea, order Isopoda) and crayfish (class
Crustacea, order Decapoda) might look similar under superficial examination but
refer to different species.

 Nomenclature and the Binomial System:

 Nomenclature is the process of assigning a distinctive name to each species.

 The binomial system of nomenclature helps avoid confusion caused by
common names and is universal, ensuring clarity in biology.
 Each animal has only one correct binomial name, as required by the
International Code of Zoological Nomenclature.
 The binomial name consists of two parts:
1. Genus (capitalized)
2. Species epithet (lowercase)
 The scientific name is italicized or underlined because it is derived from Latin
or is latinized.
 Example: Humans are named Homo sapiens, and when the genus is
understood, it can be abbreviated as H. sapiens.

Taxonomic Methods

 Traditional Taxonomy:

 Taxonomists use observable phenotypic traits to establish evolutionary

relationships (phylogenies).
 Observable traits are usually based on genetic makeup but can be influenced by
epigenetic factors.
 Taxonomists use computational tools and databases to create and evaluate
phylogenetic trees.
 Taxonomic databases contain species descriptions, distribution, ecology, keys,
and literature citations.
o Example: FishBase is a global species database for fishes.

 Molecular Biological Techniques:

 Molecular biology provides important data for taxonomic studies by analyzing

genes and proteins.
 Related animals have more similar DNA and proteins than distantly related
ones.
 Gene sequencing technologies use the polymerase chain reaction (PCR) to
amplify DNA sequences and automated sequencers to read them.
 Sequencing nuclear DNA and mitochondrial DNA is now common in
taxonomic studies.
o Mitochondrial DNA is inherited maternally and is useful for tracing
maternal lineages.

 Ribosomal RNA (rRNA) in Taxonomy:

  rRNA genes are valuable for studying ancient evolutionary relationships
(millions to billions of years ago).
 rRNA changes slowly due to evolutionary conservation, meaning closely
related organisms have similar rRNA, and distantly related ones have more
differences.
 rRNA plays a key role in protein production in ribosomes.

 Extragenic DNA:

 Animals have extragenic DNA (DNA that doesn’t code for proteins) that can
change without harming the organism.
 Changes in extragenic DNA accumulate over time, making it useful for studying
recent phylogenetic changes (e.g., over 50,000 to a few million years).
 Techniques like microsatellites and AFLP analyze extragenic DNA to track
evolutionary changes.

 Molecular Databases and Tools:

 Tools analyze DNA base sequences and compare them to molecular databases
(e.g., NCBI).
 Molecular systematists use these tools to compare organisms and determine
evolutionary relationships.

 Combining Traditional and Molecular Methods:

 Molecular techniques complement traditional taxonomy but will not replace it.
 The most reliable phylogenetic hypotheses come from combining both traditional
and molecular methods to reach similar conclusions.

Animal Systematics

 Goal of Animal Systematics:

o Arrange animals into groups that reflect evolutionary relationships.
o Ideally, groups should include the most recent ancestral species and
all of its descendants, forming a monophyletic group.
o Monophyletic Group: Includes a common ancestor and all its
descendants (the ideal group).
o Polyphyletic Group: Does not include the most recent common ancestor
and has at least two phylogenetic origins. Reflects insufficient knowledge.
o Paraphyletic Group: Includes some, but not all, descendants of the most
recent common ancestor. May indicate insufficient knowledge or need for
clarification.
 Characters Indicating Relatedness:
o Taxonomists look for characters that indicate relatedness. A character is
anything with a genetic basis that can be measured.
o Types of characters:
 Morphological: e.g., structure of insect genitalia.
 Embryological: e.g., early cleavage patterns of a fertilized egg.
 Biogeographical: e.g., distribution across mountain ranges.
 Physiological: e.g., hormone functions.
 Ecological: e.g., habitat requirements.
 Behavioral: e.g., courtship behaviors.
  Molecular: e.g., DNA base sequences.
 Types of Characters:
o Homologous Characters:
 Related through common descent.
 Example: Vertebrate forelimbs (arms, forelegs, wings) are
homologous.
o Analogous Characters:
 Result from similar evolutionary pressures (convergent evolution),
not common ancestry.
 Example: The wings of birds and insects are analogous
(homoplasy).
 Use of Homologies vs. Homoplasies:
o Homologies (similarities due to common ancestry) are useful in
classification.
o Homoplasies (analogous resemblances) are not useful for classification.
 Approaches to Animal Systematics:
o Two popular approaches include:
 Evolutionary Systematics
 Phylogenetic Systematics (Cladistics)

In animal systematics, taxonomists use genetic and observable traits to classify

animals and understand their evolutionary relationships. Groups must reflect
evolutionary history, with homologous characters being the key to identifying
relatedness.

Phylogenetic Systematics or Cladistics

Cladistics (Phylogenetic Systematics):

 Goal: Generate hypotheses of genealogical relationships among monophyletic

groups of organisms.
 Focus: Cladists focus on homologies of recent origin for phylogenetic
studies.

Character States:

 Ancestral Character States (Plesiomorphies):

o Old traits retained from a common ancestor.
o Shared by all members of a group but cannot describe relationships within
the group.
o Help establish a valid starting point for study.
 Derived Character States (Apomorphies):
o Traits that arose after common ancestry with an outgroup.
o Used to describe relationships within the group.
o More recent traits, shared by closely related members.
 Synapomorphies:
o Derived characters shared by members of a group, indicating a common
origin.

Outgroup Comparison:
  A related group (outgroup) is used to determine ancestral and derived
character states.
 The outgroup shares ancestral characters with the study group but doesn’t
include them in the group.

Cladogram (Tree Diagram):

 A cladogram depicts evolutionary relationships by showing the sequence of

derived characters.
 Branches represent evolutionary pathways.
 Nodes represent points of divergence (common ancestors).

Application of Cladistic Principles:

 Example with Vertebrates:

o The outgroup Cephalochordata shares characters like pharyngeal slits
and notochord with the study group (vertebrates), making them
symplesiomorphies.
o Synapomorphies (derived characters) like vertebrae and paired
appendages are used to determine relationships within the vertebrate
groups.

Clades:

 A clade is a related subset within a cladogram, defined by shared derived

characters (synapomorphies).
 For example, Tetrapoda and Lobe-finned fishes form a clade based on shared
derived characters like appendages with muscular lobes.

Autapomorphies:

 Autapomorphies are unique derived characters that define a taxon (e.g., the
sucker-like mouth of lampreys).
 For a specific taxon, autapomorphies are shared within that taxon but not useful
for describing relatedness with other groups.

Sister Groups:

 Sister groups are two taxa that share a most recent common ancestor.
 Two sister groups form a monophyletic clade.
 Perch and related fishes are not sister groups to lobe-finned fishes because
their most recent common ancestor is shared with tetrapods.

Cladogram of Vertebrates:

 Amniota (reptiles, birds, mammals) are defined by the presence of

extraembryonic membranes.
 Derived characters like skull characteristics distinguish reptiles, birds, and
mammals.
 Within a taxonomic group, synapomorphies define larger groups, while
symplesiomorphies are shared by members of the clade but cannot distinguish
them.
Taxonomic Hierarchy:

 Cladograms show a hierarchy of relatedness:

o Groups within a nested group are more closely related.
o For example, Amniota (reptiles, birds, mammals) are a nested group
within Gnathostomata (vertebrates with jaws).

In summary, cladistics focuses on determining evolutionary relationships by identifying

derived characters (synapomorphies) to group organisms. It uses cladograms to
show the evolutionary paths and relationships between organisms based on shared
ancestral and derived traits.

Evolutionary Systematics

 Criteria Used:
o Recency of common descent: How recently two groups share a
common ancestor.
o Amount and nature of evolutionary change: The evolutionary
changes that have occurred since divergence from a common ancestor.
 Character States:
o Evolutionary systematists recognize plesiomorphic (ancestral) and
apomorphic (derived) character states, similar to phylogenetic
systematists.
o Derived character states are used to evaluate branching patterns
within phylogenies.
 Monophyletic Groups:
o Evolutionary systematists define a monophyletic group as one that has
descended from a common ancestor.
o Unlike cladists, evolutionary systematists do not insist that a monophyletic
group must include all descendants of the common ancestor.

Differences Between Evolutionary and Phylogenetic Systematics:

 Example with Amphibians:

o Evolutionary systematists consider Amphibia (the class of amphibians) as
a valid taxonomic group with a single temnospondyl ancestor.
o Cladists, however, place amphibians as part of the larger monophyletic
group Sarcopterygii (which includes lobed-finned fishes, reptiles, birds,
and mammals), and Amphibia would be considered paraphyletic in
cladistic analyses.
 Example with Reptiles and Birds:
o Evolutionary systematists recognize Aves (birds) as a monophyletic
group with a common single ancestor.
o Cladists, on the other hand, point out that crocodylian reptiles
(crocodiles) share a more recent common ancestor with birds than
with other reptiles. Therefore, in cladistic analysis, Reptilia and Aves are
considered paraphyletic groups and should be combined.
 Importance of Derived Characters:
o Evolutionary systematists value derived characters like wings, feathers,
and flight adaptations in defining birds and consider them as defining
an adaptive zone—a unique set of characteristics that distinguish the
group.
Tree Diagrams:

 Both evolutionary systematists and cladists use tree diagrams (phylogenetic

trees) to depict relationships.
 However, evolutionary systematists often integrate fossil record information into
their trees to show:
o Time periods of evolution.
o The relative abundance of taxa within a lineage.

Key Differences Between Evolutionary and Cladistic Approaches:

 Evolutionary systematics allows for groups that are not strictly monophyletic
(i.e., groups that don’t include all descendants of a common ancestor), while
cladistics requires monophyletic groups to include all descendants.
 Evolutionary systematists place more emphasis on derived characteristics,
especially those that define adaptive zones, such as adaptations for flight in
birds, while cladists focus on the shared derived characters within a
monophyletic group.

In summary, evolutionary systematics uses a more flexible approach to grouping

organisms, considering both the amount of evolutionary change and the recency of
common descent. While it shares some methods with cladistics, it differs in its
definition of monophyletic groups and the inclusion of adaptive characters in the
classification of organisms.

Two Approaches, One Goal

Cladistics vs. Evolutionary Systematics:

 Different Approaches, Same Goal:

o While cladistics and evolutionary systematics may lead to different
interpretations of phylogeny, most zoologists agree that both methods
offer valuable insights and provide different perspectives on the same
evolutionary relationships.

When Evolutionary Systematics is Useful:

 Evolutionary systematics is particularly valuable when the fossil record is well-

known, as it allows for a chronological sequencing of ancestral lineages.
 Example: The evolution of the horse and primate evolution are often
studied using evolutionary systematics due to the richness of their fossil records.

When Cladistics is Useful:

 Cladistic studies do not focus on ancestral-descendant relationships but instead

test hypotheses regarding evolutionary relationships (branching patterns)
among organisms.
 Cladistics offers a method for investigating the branching sequences that link
different organisms.

Complementary Nature of Both Approaches:

  Differing interpretations from the two methods lead to a more complete
understanding of evolutionary relationships. These differences prompt
scientists to re-examine old hypotheses, ensuring that the field of animal
systematics remains dynamic and evolving.

Animal Systematics in Zoology:

 Chapters 9 through 22 in zoology books typically offer a survey of the animal

kingdom, following traditional taxonomy, which most zoologists find familiar and
comfortable.
 Cladograms are often included in the “Evolutionary Connections” section at
the end of chapters, with any alternative interpretations of animal phylogeny
explained and discussed.

Conclusion:

 The different approaches used in systematics help refine our understanding of

animal evolution. These differences force researchers to challenge and test
ideas, keeping the field vibrant and continually evolving.

ANIMAL ORGANIZATION

Key Concepts of Animal Organization:

 Multicellularity & Heterotrophy: Animals are multicellular organisms that

consume organic material (heterotrophic) and are classified as eukaryotes (with
complex cells).
 Patterns of Organization:
o Animals can be characterized by how their cells are organized into
tissues, and how those tissues form organs and organ systems.
o The patterns of organization in animals are often similar in closely
related species due to common ancestry and similar environmental
adaptations.
o However, animals with common ancestry may have different body
forms due to adaptations to different environments.
o Conversely, distantly related animals may appear to have similar body
forms because they evolved to fit similar environments (this is known
as convergent evolution).

Complexity and Adaptation:

 The use of terms like “simple” or “primitive” can be misleading. These terms
should be used carefully:
o For instance, animals that lack organs or organ systems, or that resemble
early ancestors, may be seen as "primitive," but this doesn’t mean they
are less evolved or inferior.
o Some simpler structures, such as parasites that have lost digestive
organs, are highly adaptive to their environment and have survived for
millions of years.
o Complexity in animals (e.g., the nervous system of octopuses and
squid) reflects advanced specialization, but this doesn’t necessarily
imply superiority.
Understanding Evolution:

 "Advanced" animals with detailed organ systems are often very different from
the earliest animal ancestors. However, it’s important to recognize that no body
form is inherently "better" or "worse".
 The survival of any group of animals for hundreds of millions of years reflects its
success in adapting to its environment, not the level of complexity or the use of
"simpler" or "primitive" descriptors.

Conclusion:

 When discussing animal evolution and organization, it's important to avoid using
terms like "simple" or "primitive" in a judgmental way. These terms should be
used to describe the adaptive success of a group rather than implying
inferiority or inadequacy. Each animal's organizational structure is the result of
successful adaptation to its unique environment.

Symmetry

Types of Symmetry:

1. Asymmetry:
o Definition: The absence of a central point or axis, meaning the body
parts are not symmetrically arranged around any reference point or axis.
o Example: Sponges are asymmetrical and do not develop complex
systems like communication, sensory, or locomotor functions. Their lack of
symmetry allows them to grow and adapt to any available substrate,
making them highly successful over millions of years.
2. Radial Symmetry:
o Definition: The arrangement of body parts such that any plane passing
through the central oral-aboral axis divides the animal into two mirror
images. The body parts are symmetrically arranged around a central axis.
o Example: Sea anemones exhibit radial symmetry, where sensory and
feeding structures are distributed uniformly around their body. Radial
symmetry allows for a relatively simple organization but limits the
complexity of sensory and motor systems. Radial symmetry is sometimes
modified in animals, leading to biradial symmetry (where structures are
arranged in pairs) or pentaradial symmetry (arrangement in fives).
o Limitation: Radial symmetry makes it difficult to evolve elaborate
structures like complex sensory or locomotor systems, as these would
require specialized structures distributed around the animal.
3. Bilateral Symmetry:
o Definition: The body can be divided into two mirror-image halves by a
single plane passing through the longitudinal axis (from head to tail).
o Example: Active animals, like crawling or swimming animals, often
exhibit bilateral symmetry because they move primarily in one direction.
This symmetry allows for the development of specialized structures at the
anterior (head) end, which encounters the environment first.
o Cephalization: This is the evolutionary development of a distinct head at
the anterior end of the animal, where sensory, nervous, and feeding
structures are concentrated. This organization allows for better
coordination and responsiveness to environmental stimuli.
 o Other Terminology: Bilateral symmetry also involves additional
directional terms (e.g., anterior for head, posterior for tail) and body
planes for anatomical reference (as shown in tables and figures).

Summary:

 Asymmetry is seen in animals like sponges, which have simpler body forms and
are not focused on complex functions.
 Radial symmetry allows animals like sea anemones to have uniform structures
around their central axis, useful for slow-moving, sedentary lifestyles but limits
complex sensory and locomotor development.
 Bilateral symmetry is found in more active animals, providing the ability to
develop specialized sensory, nervous, and feeding structures in the head,
promoting a more complex nervous system and behavioral responses.

Symmetry plays an important role in the evolutionary success and adaptation of

animals to their environment.

Cellular and Tissue Organization

In addition to body symmetry, animals are classified based on several other structural
features, including cellular organization, tissue layers, and the presence or
absence of a body cavity. Here’s a breakdown of each criterion:

The Unicellular (Cytoplasmic) Level of Organization

 Unicellular organisms are composed of a single cell that performs all the
functions necessary for life, including locomotion, food acquisition,
digestion, water and ion regulation, sensory perception, and
reproduction.
 This level of organization is not "simple." Although unicellular organisms may
seem basic, each cell is highly specialized and complex, capable of carrying out
multiple tasks simultaneously within a single plasma membrane.
 Protists, such as amoebas and paramecia, are classic examples of unicellular
organisms.
 Some zoologists use the term "cytoplasmic organization" to emphasize the
fact that these organisms operate within the confines of a single cell, where all
metabolic processes take place.
 Despite the complexity, unicellular organisms are still highly efficient at surviving
and thriving in a variety of environments, often performing complex activities
like sensory perception and movement.

Cellular Aggregates (Colonies):

 A cellular aggregate is a loose association of cells that are not highly

interdependent, and there is little cooperation or coordination of functions
compared to true tissues. These aggregates are sometimes called colonies.
 Although these aggregates do not form tissues (as cells do not work as a single
coordinated unit), they can still have cells or groups of cells that are supported
by an extracellular matrix (ECM) that gives structure and support.
 Cells within a colony may become specialized for specific functions, such as:
o Reproductive functions (producing offspring or spores),
 o Nutritive functions (absorbing or processing food),
o Structural functions (providing physical support to the colony).
 This specialization in function is sometimes referred to as a division of labor,
though it is not as advanced or integrated as tissue-level organization.
 An example of cellular aggregates in the animal kingdom could be seen in
certain colonial protozoans or sponges, where individual cells can carry out
specific roles for the group but do not form a fully integrated tissue system.

Summary:

 Unicellular organisms operate as a single cell performing all necessary

functions, which is quite complex despite being "single-celled."
 Cellular aggregates are groups of loosely associated cells with minimal
cooperation, which may show some specialization of cells for different functions
but do not form true tissues.

These organizational levels represent the simplest forms of life but still showcase the
remarkable adaptability and efficiency of living organisms.

Diploblastic Organization:

 Diploblastic animals have two embryonic tissue layers:

o Ectoderm: Forms the outer body layer (epidermis).
o Endoderm: Forms the inner layer (gastrodermis), which lines the gut
cavity.
o Mesoglea: A non-cellular layer found between the ectoderm and
endoderm. It may contain cells derived from the ectoderm or mesoderm in
some species.

These animals display tissue-level organization, where cells in each tissue

layer are functionally interdependent. Examples include Hydra and jellyfish,
where cooperation among cells allows for movements like feeding and
swimming. The organization in these animals is simpler compared to triploblastic
animals.

Triploblastic Organization:

 Triploblastic animals have three embryonic tissue layers:

o Ectoderm: Forms the outer body layer.
o Endoderm: Lines the gut.
o Mesoderm: A new layer between the ectoderm and endoderm that gives
rise to muscles, connective tissue, and blood cells.

Triploblastic animals typically have organ-system level organization, where

tissues form organs, and organs work together to form systems such as
digestive, circulatory, and nervous systems. These animals are typically
bilaterally symmetrical and are more complex and active than diploblastic
animals.

Body Cavities:
Body cavities provide various benefits to triploblastic animals, including:

1. More space for organ development.

2. Increased surface area for diffusion of gases, nutrients, and waste.
3. Storage space for various substances.
4. Hydrostatic support (acting as a hydrostatic skeleton).
5. Enhanced body size and flexibility.

Triploblastic Acoelomates:

 Acoelomates have a solid mass of cells between the ectoderm and endoderm,
without a body cavity.
 The parenchyma (a mass of loose, unspecialized cells) fills the space between
tissues.
 Acoelomates are typically small and flat, limiting nutrient and gas exchange due
to the absence of a body cavity.

Triploblastic Pseudocoelomates:

 Pseudocoelomates possess a body cavity, but it is not entirely lined by

mesoderm.
 The cavity, called a pseudocoelom, develops from the blastocoele (the initial
cavity in the early embryo).
 Organs are suspended freely in the pseudocoelom without mesodermal support.
The body cavity allows for greater movement and size compared to
acoelomates.

Triploblastic Coelomates:

 Coelomates have a true coelom, a body cavity completely surrounded by

mesoderm.
 The peritoneum (mesodermal sheet) lines the inner body wall, and serosa lines
the visceral organs.
 Mesenteries are structures that suspend the organs within the coelom, allowing
for more complex organ systems and greater flexibility.
 The coelom forms differently in two major groups:
o Protostomes: The coelom forms from the splitting of mesodermal
masses.
o Deuterostomes: The coelom forms from the outpocketing of the
embryonic gut.

Summary of Benefits of Body Cavities:

1. More space for organs to develop.

2. Better diffusion of gases, nutrients, and wastes.
3. Hydrostatic skeletons for support and flexibility.
4. Increased body size.

This classification system helps zoologists understand the evolutionary relationships

and complexity of different animal groups based on their body organization and
internal structures.