Lecture

1
Inheritance of contemporary systematics
Carolus Linneaus and Charles Darwin
o Linnaeus Scala Naturae (all organisms have a fixed place on the scale with humans at the
summit)
Haeckel introduced phylogeny
Willi Hennig cladistics
o Only method that produces only monophyletic groups
Groups that include an ancestor and oof its descendants
o Completely changed biological classifications
o Permitted a better comprehension of the structure of biodiversirsit

A brief history of biotic classification
Taxonomy
o Taxis order or arrangement
o Nomos law
o First proposed to designate the science of biotic classification
o Need for rules and rigorous principles started to be appreciated
o Two broad types of logic were employed
Aristotle
Dividing groups of organisms into subgroups according to predefined criteria and
repeating this process until the species level is reached
Divisive results in a dichotomous choice with one term opposing the other
Agglomeration
Assembling species into groups based on similarity criteria
Process is reiterated using the newly formed groups as the primary units, until all
organisms are united within a single category
o Goal: to find a logic that enabled access to the natural classification
o Different characters should not be considered as individual units, but each following its relative
value, in such a way that a single constant character might be of equal or greater value than
several inconsistent characters taken together
Lamarck inheritance of acquired traits
o Did not gave priority to the variation of species
o Consider factors from external environment as the motors of organic change
Charles Darwin
o Before seeing the spcies, he saw that variation among individuals
o Nominalism enabled him to discover the hidden value of the variation that had previously been
considered negligible
o Natural selection
Under the particular environmental conditions of a given moment in time, certain
variants are favored and become more numerous because they leave behind more
descendants than do competing variants
The key was to study the variability not of acquired character but only that of inherited
characters, those transmitted to future generations
Resemblance between species is thus due to characters inherited from an ancestral
species
Homologues
o Characters inherited form a common ancestor
 Paraphyletic
o Possess members that are more closely related to organisms outside the grade than to other
members within the grade
Phylogenetic systematics / cladistics
o Attempts to recognize evolutionary relationships among species by assembling them into
monophyletic group
o Best understood by focusing on taxonomic sampling
Choosing a number of representative individuals about whom we ask who is related to
whom
Symplesiomorphy
o Shared ancestral trait
o Should never be taken into account in cladistics because it does not provide any information
about the relationships among species within the group
Synapomorphy
o Shared derived character state
Convergent
o Result of independent evolution
Parsimonious tree
o The one with the fewest number of evolutionary steps
o Tree with the fewest number of transformation assumptions

Definition and importance of systematics
Systematics
o Science of classification
o First task is the identification, description nad inventory of natures living organisms, past and
present
o Classification
o Objective of all science: to understand the complexity of nature in terms of categories that can
give rise to the most coherent language and reasoning possible and that may serve as basis for
other studies
o Pertinent logic for classifying species evolutionary relatedness
o Classifications should reflect the succession of branches of the unique tree of life
Importance
o Many applications medical, pharmaceutical, agronomical, ecological, geological

Systematics and biodiversity
Biodiversity can be viewed in two ways
o Habit-based (ecosystem) approach
Ecological biodiversity
Species are classified in relation to the habitat they occupy, their position in the trophic
chain, or their reproductive strategy
o History
Species are classified on the basis of comparative, structural data and a re considered on
paleontological time scale
Where does species come from?



Character and homology
Character
o Observable attribute of an organism
o Designation of that which we observe
o In a comparative approach, it is tightly associated with the notion of similarity and homology
o Complete definition: any set of observable organismal attributes for which we can formulate at
least one hypothesis of homology
State
o The way we discriminate it within a sample of organisms
Homology
o Structures are homologues when they are inherited from a common ancestor
o Structure, compared among organisms with the same body plan, has the same arrangement
with neighboring structures regardless of its form of function.
Homoplasy
o Similarity not inherited from a common ancestor

What is a tree?
Noncyclic, connected graph
o Noncylic because only a single pathway links two tips (tree is not a network)
Comprises links that join up the tips
Kinds of trees according to the method used for construction
o Dendogram
Neutral and refers to a tree
o Cladogram
Dendogram constructed by cladistics analysis
Expresses the phylogenetic relationships among taxa and nodes are defined by
synapomorphies
o Phenogram
Dendogram constructed on the basis of phonetic (observable) characteristics of the taxa
Expresses the degree of global similarity among taxa
o Phylogram
Cladogram in which the length of branches is proportional to the divergence of taxa

What is a taxon?
Group of organisms that is recognized as a formal unit
Certain descendants are missing form paraphyletic group
Common ancestor is missing from polyphyletic groups

What is evolutionary relatedness?
Look for relatedness in sister groups
Searching for a group that shares a unique common feature with another group, a feature not shared
with another group
Presence of the unique feature attests to an inheritance from an exclusive hypothetical common
ancestor

How do we determine evolutionary relatedness?
First step in the construction of a phylogeny is to polarize character changes, distinguish the primitive
state (plesiomorphy) from the derived states (apomorphy) deduce the relationships
Steps for construction a phylogenetic classification
Post an initial question
o Define the phylogenetic problem
Choose the organisms to be samples to answer the question: the taxonomic sampling
o Choosing one or several external reference points or out-groups
Choose or identify the characters that will be informative for the question posed: the character sampling
Code the data and the character matrices
o Establishing primary homologies and polarizing connections
Explore all possible trees
Evaluate each tree with regard to the character matrix
Choose the tree-selection criterion: parsimony
Choose a tree
o Shortest tree

Phenetics
Analyze data in another manner
In contrast to cladistics, phonetics attempts to quantify the general similarity between organisms
o Distance is calculated for each pair of taxa

When is a tree a phylogeny?
Phylogeny the historical pattern of descent of organized beings
Tree is not necessarily a phylogeny
o Because it can be used to symbolize a hierarchy
To classify is to create logical links among objects according to a predefined purpose
Purpose of classification describe the degree of evolutionary relatedness among organisms

Lecture 2
Fishes have occupied nearly all major aquatic habitats
Evolved obvious and striking anatomical, physiological, behavioral, and ecological adaptations
Aided by an extensive fossil record dating back more than 500 years
Fishes are excellent showcases of the evolutionary process, exemplifying the intimate relationship
between form and function, between habitat and adaptation

What is a fish?
Poikothermic, aquatic chordate with appendages (when present) develops as fins, whose chief
respiratory organs are gills and whose body is usually covered with scales
A fish is an aquatic vertebrate with gills and with limbs in the shape of fins

The diversity of fishes
Fish is singular and plural for a single species
Fishes is singular and plural for more than one species







Superlative fishes
Lungfishes can live in a state of dry suspended animation for up to 4 years, becoming dormant when
their ponds dry up and reviving quickly when immersed in water
Antarctic fishes live in water that is colder than the freezing point of their blood. They keep from
freezing by avoiding free ice because of the presence of antifreeze proteins that depress their bloods
freezing point
Deep sea fishes include many forms that can swallow prey larger than themselves
Gender change is common
Fishes engage in parental care
Fishes are unique among organisms with respect to the use of bioelectricity
Fishes are unique among vertebrates in their ability to produce light
Although though of as cold-blooded, some pelagic sharks and tunas maintain body temperature warmer
than their surroundings
Predatory tactics include attracting prey with modified body parts
Fishes can significantly change the depth of their bodies by erecting their fins or by filling themselves
with water, an effective technique for deterring many predators

A brief history of ichthyology
Georges Cuvier and Achille Valenciennes produce the first complete list of the fishes of the world

Summary
Fishes account for more than half of all living vertebrates and are the most successful vertebrates in the
aquatic habitats worldwide
Most of living fishes are primarily marine and the remainder live in freshwater

Lecture 3
Why do we need a system of classification?
o Things must be named and divided into categories before we can talk about and campare them
o We cannot deal with all the members of a class individually, so we must put them into some sort
of classification
The most general classification is considered for special purposes the most general classification is
considered to be the most natural classification
o Classification that best represents the phylogenetic (evolutionary) history of an organism and its
relatives
o A phylogenetic classification of taxa holds extra information

Species
Species are the fundamental unit of classification schemes
A community, or a number of related communities whose distinctive morphological characters are, in
the opinion of a competent systematist, sufficiently definite to entitle it, or them to a specific name
Biological species concept
o Groups of actually or potentially interbreeding populations which are reproductively isolated
from other such groups
Evolutionary species
o Single lineage of ancestor-descendant populations which maintined its identity from other such
lineages and which has its own evolutionary tendencies and historical fate


Taxonomy versus systematics
Taxonomy
o Deals with the theory and practice of describing biodiversity (including naming undescribed
species), arranging this diversity into a system of classiciation and devising identification keys
o Includes rules of nomenclature that govern use of taxonomic names
Systematics
o Emphasizes the study of relationships postulated to exist among species or higher taxa, such as
families and orders

Approaches to classification
Cladistics
o Apomorphies
More recently evolved, derived, or advanced characters
o Pleisiomorphies
More ancestral, primitive or generalized characters
o Goal is to find synapomorphies (shared derived characteristics) that diagnose monophyletic
groups
o Symplesiomorphics shared primitive characters
Do not provide data useful for constructing phylogenetic classifications because primitive
characters may be retained in a wide variety of distantly related taxa
o Autapomorphies, specialized characters that are present in only a single taxon, are important in
defining that taxon but are also not useful in constructing a phylogenetic tree
o Taxa are arranged in a cladogram
o Monophyletic groups are defined by at least one synapomorphy at a node, or branching point
o Primary goal: definition of monophyletic groups
o
Phenetics
o Starts with species or other taxa as operational taxonomic units
o Clusters the OTUs on the basis of overall similarity, using an array of numerical techniques
Evolutionary systematics
o Holds that anagenesis, the amount of time and differentiation that have taken place since groups
divided, must also be taken into consideration along with cladogenesis, the process of branch or
lineage splitting between sister groups
o Tree- phylogram

Taxonomic characters
Characters are needed to differentiate taxa and assess their interrelationships
Variations of a homologous structure and to be useful, they must show some variation in the taxon
under study
Categories
o Meristic
Characters that correspond to body segments, such as numbers of vertebrae and fin rays
Used for almost any countable structure
Useful because they are clearly definable, and usually other investigators will produce the
same counts
Easier to treat statistically
o Morphometric characters
Measurable structure such as fin lengths, head length, eye diameter, or ratios
 Can be measured to different levels of precision and so are less easily repeated
Problem of allometry
Lengths of different parts change at different rates with growth
o Anatomical characters
Include characters of the skeleton and soft anatomy such as position of the viscera,
divisions of muscles, and branches of blood vessels
o Molecular characters
Nuclear DNA and mitochondrial DNA
Closely related organisms show a high degree of similarity in molecular structure
Barcoding
Relies on differences between species in a relatively short segment of mtDNA

Units of classification
Order iformes
Suborder oidei
Family idea
Subfamily inae
Tribe ini

International code of zoological nomenclature
System of rules designed to foster stability of scientific names for animals
Rules deal with such topics as the definition of publication, authorship of new scientific names and types
of taxa
Species and subspecies are based on type specimens
o Primary types
Holotype
Single specimen upon which the description of a new species is based
Lectotype
A specimen subsequently selected to be the primary type from a number of
syntypes (a series of specimens upon which the description of a new species was
based before the code was changed to disallow this practice)
Neotype
A replacement primary type specimen that is permitted only when there is strong
evidence that the original primary type specimen was lost or destroyed and when
a complex nomenclatorial problem exists that can only be solved by the selection
of neotype
o Secondary types
Paratypes
Additional specimens used in the description of new species
Paralectotypes
The remainder of a series of syntypes when a lectotype has been selected from
the syntypes
Topotype
A specimen taken from the same locality as the primary type, and therefore
useful in understanding variation of the population that included the specimen
upon which the description was based
Allotype
 A paratype of opposite sex to the holotype and useful in cases of sexual
dimorphism

Name changes
Why do scientific names of fishes sometimes change?
There are four primary reasons that systematists change names of organisms
o Splitting
Considered to be a single species into two (or more)
o Lumping
Two species that were considered distinct into one
o Changes in classification
o An earlier name is discovered and becomes the valid name by the principile of priority

Summary
The best classification is the most natural one, that which best represents the phylogenetic
(evolutionary) history of an organism and its relatives
Species are the fundamental unit of classification and can be defined as a single lineage of ancestor
dependent populations that maintains its identity from other such lineages. Species are usually
reproductively isolated form other species
Taxonomy deals with describing biodiversity (including naming undescribed species), arranging
biodiversity into a system of classification, and devising identification keys. Rules of nomenclature
govern the use of taxonomic names. Systematics focuses on relationships among species or higher taxa
Cladistics is a widely used system of classification in which characters are divided into apomorphies
(derived or advanced traits and plesiomorphies (primitive or generalized traits). The goal is to find
synapomorphies (shared derived characters_ that define monophyletic groups, or clades (groups
containing an ancestor and all its descendant taxa)
Taxonomic characters can be meristic (countable), morphometric (measurable), morphological
(including color), cytological, behavioral, electrophoretic, or molecular
Ray finned: Kingdom: Animalia; Phylum: Chordata; Subphylum: Vertebrata; Superclass: Gnathostomata;
Grade: Teleostomi; and Class: Actinopterygii
Holotype the single specimen upon which the description of a new species is based
Paratype additional specimens used in the description of a new species

Lecture 4
Fishes were the first vertebrates
Understanding the evolutionary history of fishes is therefore important not only for what it tells us
about fish groups, but for what it tells us about evolution of the vertebrates and ultimately our own
species

Jawless fishes
The very first fishlike vertebrates undoubtedly evolved from invertebrates, perhaps a cephalochordate

Subphylum Craniata, Infraphylum Vertebrata (Dermal skeleton, and neural crest)
Conodonta (extinct)
o Consist of tooth like structure made from calcium phosphate
o Protocondodonts V-shaped muscle blocks, a bilobate head and cartilaginous head skeleton,
eyes contained in otic capsules, extrinsic eye muscles, a compressed body, axial lines suggestive
of a notochord, and unequal tail fins supported by ray like elements
 o Conodont elecments were dentition homologous to the rasping, book-closing action of modern
hagfishes
o They are more derived than hagfishes and lampreys and may even be basal to the jawed fishes
that arose later
Ostracoderms (extinct)
o First species, conodonts aside, were historically named ostracoderms (shell skinned) in reference
to a bony shield that covered the head and thorax
o Considered an artificial designation that includes perhaps four distinct superclass of jawless
craniate fishes
Pteraspidomorphi (extinct)
Or Diplorhina two nares
Inside of the head plates indicates 2 separate olfactory bulbs in the brain
Jawless filter feeders in both marine and freshwater environments
Three subclass
o Astraspida (extinct)
Symmetrical tails
Full body armor
Multiple branchial openings
o Arandaspida (extinct)
Same with astraspida
o Heterostraci (extinct)
those with a different shell
Had dermal armor extended from the head almost to the tail,
necessitating swimming by lashing the tail back and forth, like
tadpole
Hypocercal the notochord extended into the enlarged lower lobe
of the tail
Plowed at the bottom, pumping sediments into the ventral mouth,
and filtering digestive material through the pharyngeal pouches
Armor is generally sutured and shows growth rings, indicating
incremental growth
Anaspida (extinct)
More fusiform, compressed anspidiforms
Seldom larger and had pronounced hypocercal tails and terminal mouths
Originated in near shore marine habitats and gradually entered fresh water
Covered largely with overlapping, tuberculate scaled
One advance was the development of flexible, lateral, finlike projection that has
muscles and an internal skeleton, thus giving these small fishes considerable
maneuverability
Thelodonti (extinct)
nipple teeth
Diminutive, fusiform, jawless fishes that were covered with denticles rather than
bony plates
Most were depressed with a horizontal mouth, asymmetrical hypocercal tails,
and a detectable lateral line that ran the length of the body
mode of life probably similar to pteraspidiforms skimming and filtering small
organisms from bottom sediments while swimming
Osteostracomorphi (extinct)
 One class and 3 orders of jawless fishes
Had two semicircular canals and evidence of truve bone cells

Gnathostomes: early jawed fishes
Jaws are present, derived from gill arches
Paired limbs with skeletal support are usually present
Endochondral bone, three semicircular canals, and dentine-based rather than horny teeth
Perhaps the greatest of all advances in vertebrate history was the development of jaws and the
consequent revolution in the mode of life of early fishes
This revolution included a diversification of the food types that early fishes could eat
Large animal prey could be captured and dismemeberd, and hard-bodied prey could be crushed
Agnathous fishes were probably limited to planktivory, detrivory, parasitism and microcarnivory
Stomachs for storage of food evolved, probably as a consequence of jaws that could bite off pieces of
food
With the advent of jaws, both carnivory and herbivory on a grand scale became possible
Jaws also allowed for active defense against predators, leading to de-emphasis on armor
This increase in agility was greatly enhanced by the development of paired, internally supported
pectoral and pelvic appendages the most outstanding shared derived character of the gnathostomes
besides tha jaw

Class Placodermi
Plate skinned
Peculiar bony, often ornamentaed plated that covered the anterior of the body
Most placoderms had depressed, even flattened bodies, suggesting benthic existence
As in ostracoderms, they have occurred first in marine habitats but later moved into fresh water. Many
placoderm groups show an evolutionary trend toward reduction in external armor, leading to a mobile
existence in the water column
Ossified haemal and neural arches along the unconstrictred notochord and three semicircular canals
Acanthothoracidofrms basal group and are the oldest known jawed vertebrates
Arthrodirifoms larges order
o Posses a unique hinge at the back top of the head between the braincase and the cervical
vertebrae termed craniovertebral joint
o Joint allowed opening of the mouth by both dropping the lower jaw and raising the skull roof,
thus increasing gape size
o As the group evolved, this joint became larger and more elaborate and dentition diversifiend into
slashing, stabbing, and crushing structures
o Largest among placoderms
Rhenaniforms
o Exteremely dorsoventrally depressed and bore a striking resemblance to modern skates, rays
and angel sharks,
Antiarchiforms
o Predominantly freshwater, heavily armored, benthic fishes with spiral valve intesting and jointed,
arthropod-like pectoral appendages that had internal musculzarization
Ptyctodontiforms
o Resembled modern holocephalans in body form
o First fishes known to possess apparent male intromittent organs in the form of claspers
associated with the pelvic fin, an indication of internal fertilization

Advanced Jawed fishes 1: teleostomes (Osteichthyes)
Teleostomes are groups together because they share cranial, scale and fin similarities, but specially
because both acanthodians and actinopterygians possess three otoliths
Acanthodians
Sarcopterygians
o Euteleostome
o fleshy or lobe fins
o diversified into extinct and modern coelacanths, lungfishes, and tetrapodomorphs
(rhizodontimorphs, osteolepidomorphs, and the elpistostegalians that gave rise to tetrapods)
Actinopterygians
o Euteleostome
o ray fins
o Underwent tremendous multiple radiations, producing the cladistian bichirs, the chondrosteans
and neopterygians including gars and related fossil groups, bowfin and related fossil groups and
ancient and modern teleosts
Eutelostomes share numerous characteristics, including the bone series in the opercular and pectoral
girdles, the pattern of their lateral line canals, fins supported by dermal bony rates, a heterocercal tail
with an epichordal (upper) lobe, replaceable dentition, and a swim bladder that developed as an
outpocket of the esophagus

Class Acanthodi (extinct)
Oldest fossils of relatively advanced, jawed fishes
spiny sharks
Stout median and paired spins evident in most fossils
Generally small
Occurred in both salt and fresh water
Had cartilaginous skeletons
Body covered with small, non-overlapping scales; large heads; large eyes.
Streamlined, round bodies reduced armor when compared to ostracoderms

Class Sarcopterygii
Remain one of the most actively studied fossil groups of fishes

Subclass Coelacanthimorpha order Coelacanthiformes
Occurred worldwide in both marine and freshwater
More specialized than other sarcopterygians
Possess a unique spiny rather than a lobate first dorsal fin
Tree-lobed caudal fin with a middle fleshy, fringed lobe
Rostral organ

Subclass Dipnoi (Dipnotetrapodomorpha)
Besides coelecanths, sarcopterygians consist of two other subclass
o Dipnoans
Variety of extinct fishes with stout bodies and paddlelike paired fins
Major groups are dipnomorphs and tetrapodomorphs
o Tetrapoda
That emerged onto land to become amphibains, reptiles, birds, and mammals

Dipnomorphs
Unranked taxon made up of extinct fishes in the superorder Porolepimorphe, two superorders of
lungfishes, the extinct Dipterimorpha and extant Ceratodontimorpha
Lungfishes as a group have been generally referred to as Dipnoi or dipnoans
Dipterimophs arose in the Devonian in marine environments, expanded into freshwater habitats, and
died out by the end of the Triassic
Primitive lungfishes: two dorsal fins; fleshy, scale-covered, paired, leaflike archipterygial fins with a bony
central axis and with fin rays coming off the central axis; a lack of teeth on the marginal jaw bones, but
with tooth plates inside the mouth, and with premaxilla, maxilla, and dentary missing; a solid braincase;
and a pore-filled cosmine xoating on the dermal bones that covered the skull and scales that may have
been associated with eletroreception
Ceratodontimorphs
o Appear in the lower Triassic
o Represented today by the freshwater order Ceratodontiformes
Modern lungfishes take the anatomical trends to the extreme, having eel-like, largely cartilaginous
bodies, lacking any cosmine bony layers, and possessing diphycercal tails
Lungfishes underwent extensive diversification during the Devonian, evolving more than 60 genera and
100 species
Many lungfish species are known only from fossilized toothplates, with toothplates and other structures
found in fossilized lungfish burrows

Tetrapodamorohs: tetrapod ancestors
Large, predatory fishes characterized by sarcopterygian traits such as two dorsal fins, cosmine covering
of the bones and scaled, kinetic skulls, lobbed fins, and replacement teeth on the jaw margins
Remained common throughout the latter half of the Paleozoic
Evolutionary trends include reduction in dermal bon thickness, a change from diamond to round-
shaped scales, and and increasingly symmetrical tail
Possessed any homologies with later tetrapods, these fishes were unlikely to have been transitional
forms to living on land, even temporarily
Epistostegalians
o Considered the most likely sister group of modern tetrapods
o Many apparent homologies, including eye position, skull roof bones, paired fins, dentition, and
vertebral accessories
o Both have eyes set close together on the top of the skull facing upwards, with eyebrowlike ridges

Class Actinopterygii
Just as ancient as most of the other groups, primitive ray-finned fishes are similar in size and shape to
many extant fishes, and many of their fossils are very well preserved

Subclass Cladista
Also referred to as brachiopterygians

Subclass Chondrostei
Early fishes, collectively known as palaconiscoids were relatively small and were distinguished from
saropterygians by the presence of a single triangular dorsal fin, a forked heterocercal tail with no upper
lobe above the unconstructed notochord, paired fins with narrow rather than fleshy bases, dermal
bones lacking a cosmine layer, scaled joined by a peg-and-socket arrangement and covered with
ganoine (ganoid scales), relatively large eyes, and a blunt head
 ray fin refers to the parallel endoskeletal fin rays that were derived from scales
o Rays supported the median and paired fins, which were moved by adjacent body musculature
In contrast, the fins of the Sarcopterygii had a thick, bony central axis and muscles contained in the fin
itself

Paleoniscoid trends
Flourished throughout the latter Paleozoic
Ostracoderms, acanthodians, and placoderms disappeared and sarcopterygians diminished in
abundance
This correlation suggests ecological interaction among groups, and possible replacement of primitive
jawed and jawless fishes with more advanced actinopterygian and chondrichtyan lineages
What innovations did the ray-finned fishes possess that might have given them ecological superiority?
o Changes in jaw and fin structure leading to diversified feeding habits and increased mobility
were critical to actinopterygian success and dominance
The highly ossified braincase of early actinopterygians makes it possible to determine the origins,
insertions, and approximate sizes of the different muscle masses involved in jaw function, from which
we can estimate the forces in operation
During actinopterygian evolution, culminating in advanced teleosts, changes in the angles and
connections between the skull case, dermal bones, muscles, and ligaments of the head and jaws have
been most influential
o Hyomandibula has been reoriented from oblique to vertical, the posterior end of the maxilla has
been freed from the cheek bones, and the jaw musculature has increased in size and complexity
o These changes increased the speed and strength of the gait
o Allowed for enlarging of the mouth both vertically and laterally
o When the mouth was opened, its volume increased and it assumed a more tubular shape
Paleoniscoid scaled changed from heavy, interlocking, diamond-shaped units to thinner, lighter, circular,
cycloid structure
o Reduction was accomplished by elimination of the dentine, vascular, and ganoine layers
Because paleoniscoid fins consisted of jointed scaled, reduction in scale thickness meant increased
flexibility in fins; fins became mobile structures composed of dermal fin rays that could be erected or
lowered and also moved laterally
Associaited with scale reduction was increased ossification of the vertebral column, leading to
recognizable centra with dorsal and ventral accessory structures
These accessory structures are closely related to modifications in the caudal region, where a major
trend has been toward an increasingly symmetrical, homocercal tail
Caudal fin rays became supported by a series of ventral accessory, hypural bones
All these changes during palaeoniscoid phylogeny imply increased reliance on locomotion, integrated in
both escape and prey capture

Subclass Neopterygii
new fins
First appear in the fossil record during the Upper Permian
Underwent an initial radiation in the Triassic and Jurassic and then expanded more extensively in the
late Cretaceous
Pycnodontiforms
Aspidorhynchiforms
o Needlefishlike body from
Pachycormiforms
 o Sister group to early teleosts
Lepisosteifrom gars and amiiform Bowfin extant pre-teleostean
o Intermediate between paleoniscoids and teleosts in a number of structures:
Gars retain the ganoidlike scales of primitive neopterygians
Bowfin have a primitive gular plate under the head
Both groups have identifiably heterocercal tail elements
Bowfin, and their fossil relatives justify their placement together in a separate group,
sometimes referred to as the division Holostei
Neither gars nor Bowfin are considered to be on a direct line to the teleosts

Division Teleostei
perfect bone
Outnumber all other living fish groups
Form a definable group with a recognizable ancestry
Constitute a monophyletic group
o Chief among these are ural neural arches elongated to from the uroneurals of the tail support
o Unpaired basibranchial toothplates
o Distinctive urohyal
o Prevalence of a mobile premaxilla
Arose in the Middle or Late Triassiac
Evolution apparently involved four major radiations
o Three that each gave rise to distinct, primitive subdivisions
o Fourth that produced the major advanced groups alive today
Multiple radiations imply that modern teleosts as a group could be polyphyletic, more a developmental
grade than a single clade
Radiations
o Osteoglossomorphs
Bony tongues
o Elopomorphs
Tarpons and true eels
o Otecephalans or osttarioclupeomorphs
Herrings and minnow relatives
o Euteloestei

Trend during teleostan phylogeny
Reduction in bony elements
o Fusion or actual loss of bones
Reposition and elaboration of the dorsal fin
o Dorsal fin in primitive telesots is a simple, spineless, fixed, single, midbody keel that prevents
rolling and serves as a pivot point for fishes that typically swim in open water situations
o In higher teleosts, the trend is for the dorsal fin to become elongate and diversified
Change in placement and function of paired fins
o In basal teleosts, pectoral fins are oriented horizontally and located in the thoracic position,
below the edge of the gill cover
o Pelvic fins occur at mid-body in an abdominal location
o Both fins act primarily as planes that help stabilize movement up and down or from side to side
o Teleostean phylogeny: pectoral fins move up onto the sides of the body and their base assumes
a vertical orientation; pelvic fins move into thoracic or even jugular position
 Structural modifications to and interaction between the caudal fin and gas bladder
o Progressive increase in symmetry of the tail fin
o Tail fins became externally and functionally homocercal fairly early in the groups history
o Apparent conjuction with tail and paired fin modifications, an additional teleostean trend is
added control over gas bladder function
Physostomous
Gas bladder in which a pneumatic duct connects the gas bladder with the gut and
ultimately the mouth
Filled with gas by gulping air
Gas is expelled largely via the same route
Physoclistous
Lost the pneumatic duct
o Gas bladder makes an otherwise dense fish neutrally buoyant
Jaw improvements
o Teleosts developed a protrusible pippet mouth capable of generating powerful, directed,
negative pressures
Results from enlargement of the muscles and modifications to the bones in the jaw
apparatus, most notably the maxilla and hyomandibula
o In teleosts, it has been the dentition and musculature of the pharyngeal jaws that have
diversified to serve this chewing, crushing, and grinding function
o Pharyngeal pads lie posterior to the marginal jaws, just anterior to the esophagus, and are
derived from dermal tooth plates in the pharynx

Advanced jawed fishes II: Chondrichthyes
Cartilaginous fishes
Rapidly specialized as marine predators
Many of the characters of modern Chondrichthyes are secondarily derived and represent specializations
for a very different, parallel mode of life in water

Subclass Elasmobranchii
plate or strap gills
Divided into three infraclasses
o Cladoselachimorphs
Cladoselachidae
5 gill slits
Terminal mouth
Dentition cladodont: multicuspid teeth in which the central cusp was usually
largely
Teeth were made of enamel-covered dentine and were homologous with scales
Often large, pelagic, marine predators with an unconstructed notochord
protected by calcified cartilaginous neural arches and small precaudal, lateral
keels analogous to those found in modern pelagic sharks
o Xenacanthimorphs
Common in tropical water
Pleuracanth
o Euselachians
Earliest: Ctenacanthiformes
2 dorsal fins with prominent spines
 anal fin set far back on the body
slightly overhanging snout along with a terminal mouth

Subclass Holocephalii
Calcified cartilaginous skeleton and internal fertilization, among other traits link the Holocephali with the
elasmobranchs
whole heads
Position and structure of the gill chamber, which is located further forward than in sharks and has a
single opercular opening covering four gill openings
Holocephalans have non-protrusible jaws because the palatoquadrate is fused to the brain case

Summary
First fishes to fossilized lacked jaws but possessed bony armor and had a muscular feeding pump. 5
superclasses of diverse jawless vertebrate craniates:
o Conodont
o Pteraspidomorphs
o Anaspids
o Thelodonts
o Osteostracomorphs
Bony shield that covered the head and thorax
Conodont were well known from tooth like structures hat fossilized abundantly during Precambrain
Development of jaws was a critical step in the advancement of fishes
o Placoderms were early jawed fishes that arose in the Silurian
Had bony, ornamented, platelike skin, many were predators and achieved monstrous size
Hinge at the back top of the head that allowed for greater opening of the mouth
Teeth consisted of dermal bony plates attached to jaw cartilage and could not be repaired
or replaced
First advanced jawed fishes Acanthodians / spiny sharks
o Unrelated to modern sharks
o Water column swimmers
o Share common ancestry with modern bi=ony fishes
o Often placed with sarcopterygians and actinopterygians in the grade Teleostomi
Eustelestomes Sarcopterygii and Actinopterygii
o Gave rise to modern bony fishes
o Sarcopterygians Coelacanthimorphs (coelacants), dipnoans (lungfishes, osteolepidomorphs,
and elpistostegalians), tetrpods
Elpistogalians most likely ancestors of tetrapods
Shakre skull and neck characteristics and fin patterns with stem tetrapods
o Actinopterygians cladistians (bichers), chondrosteans (palaeoniscoids, sturgeons and
paddlefishes) and neopterygians (semionotoids, gars, bowfin and teleosts)
Actinopterygians arose during the Silurian
o Successful group palaeoniscoids
Triangular dorsal fin, heterocercal tail, paired ray-supported fins with narrow bases, and
ganoid scales
Structural changes occurred in the jaw apparatus that strengthened the bit, increased the
gape, and created suction forces
Neopterygian modern ray-finned
o Most successful
 o Gars and bowfin extant pre-teleostean
Five major trends characterize teleostean evolutnio
o Reduction of bony elements
o Shifts in position and function of the dorsal fin
o Placement and function of paired fins
o Caudal fin and gas bladder modification
o Improvements to the feeding apparatus
Chondricthyans
o Elasmobranchs
Late Ordovician
Improvements in jaws, dentition, vertebrae and fins that paralleled locomotry and feeding
changes in bony fishes
o Holocephalans
Devonian
Share with elasmobranchs a calcified skeleton and pelvic fin claspers
Differ by having non-protrusible jaws (upper jaw fused to braincase) and by a single
opercular opening

Lecture 5
Jawless fishes: lancelets, hagfishes, and lampreys
Amphioxiforms
Not fishes because they lack many diagnostic characters
Cephalochordates invertebrate chordates along with urochordate and hemichordates
o Evolutionary and anatomical affinities are much closer to the vertebrates
Lancelets are small, slender organisms that as adults occupy sandy, usually shallow bottoms in all major
oceans
Commonly bury themselves in the sediments with just the anterior portion of the body protruding from
the bottom
Filter diatoms and other small food items from the water via the cilia that transport water through the
mucus-laden mouth and pharynx and out through the atriopore
Food-trapping mucus is produced by the endostyle
o Pharyngeal organ that also functions in iodine uptake and may therefore be homologous to the
thyroid of higher vertebrates
Intriguing in their lack of many typical chordate structures
Lacking most parts of a head (there is no cranium, brain, complex eyes, external nostrils, or ears)
Lack vertebrate, scales, genital ducts, a heart, red blood cells, hemoglobin, and specialized respiratory
structures
Only have 1 layer of epidermis
Number of internal gill clefts increases throughout life in lancelets
Notochord extends beyond the anterior end of the dorsal nerve tube to the anterior end of the body
Commonly used in biology laboratories as a study animal
o Example of primitive chordate features, it is morphologically convergent with the ammocoete
larva of the lampreys

Hagfishes and lampreys: Evolutionary relationships
Cyclstomata round mouths
Similarities in the body morphology of modern hagfishes and lampreys are though to reflect convergent
evolution
 Agnatha
o Polyphyletic
Hagfishes are considered a more primitive, separate, non-vertebrate group in their own superclass
Lamreys and hagfishes share a host of anatomical, physiological, and biochemical traits but have an
even greater number of differences
Both groups are scaleless, lampreys lack the mucus-producing capability of hagfishes
Lampreys have one or two dorsal fins supported by radial muscles and cartilage, whereas hagfishes have
a single continuous caudal fin
Lampreys have a terminal mouth, hagfishes a subterminal mouth
Lampreys have a larval stage, hagfishes have direct development
In adult lampreys, the external opening of the nasohypophysis is dorsal and the tract ends internally in a
blind sac above the branchial region
In hagfishes, the external opening is terminal and the internal opening is into the pharynx
Lampreys have two semicircular canals, hagfishes only one
Lampreys have a pineal organ and functional eyes, hagfishes possess neither
A major similarity between the two groups involves their immune responses
All gnathostomes, including fishes, have immune systems that involve immunoglobulin-type antigen
receptors that produce pathogen-specific antibodies in response to infectious agents such as microbes
Lampreys and hagfishes also produce pathogen-specific defensive substances, but instead of antibody
proteins, jawless fishes produce different kinds of proteins called variable lymphocyte receptors
Lampreys possess lateral line neuromasts that are touch sensitive; these are lacking in hagfishes
All lampreys have seven gill openings, hagfishes vary between one and 16
Although the tongue possesses keratinous, replaceable teeth in both groups, it is anatomically and
functionally different
Myxinoids use the tongue for biting and tearing, whereas lampreys use it for rasping and suction

Myxiniforms
Hagfishes
Slime ells or slime hags
Mucus production is the combined result of these holocrine slime glands as well as merocrine exudate
from the epidermis itself
Produce slime in response to being disturbed or handled
Mucus undoubtedly serves some antipredator function perhaps by making the fish too slippery to
handle or by clogging the gills of a potential predator and threatening it with suffocation
Hagfishes are highly specialized animals belying their typification as primitive fishes
Possess four rudimentary hearts: a primary, three-chambered branchial or systemic heart posterior to
the gills, and three auxiliary, single-chambered hearts located just behind the mouth, at midbody, and at
the end of the tail
Other hagfish peculiarities characterize the respiratory, digestive, immune, and sensory systems
Oxygen uptake in hagfishes occurs both at the gills and at capillary beds in the sin
Hagfishes lack a true stomach, having instead an intestine that begins at the pharynx and ends at the
anus, with an anterior muscular subdivision that prevents water inflow
Hagfish taxonomy is based on the arrangement of efferent gill ducts, number of slime pores, finfolds,
and tentacle and dentition patterns



Petrmyzontiforms
Hagfish scavenging habits, many lampreys are parasitic on other vertebrates
Lampreys superficially resemble hagfishes in general body form
As with hagfishes, lampreys lack constricted vertebrae, the body being supported by a notochord
Lack paired fins, jaws, a sympathetic nervous system, and a spleen
They too are scaleless, have a single nostril, and have horny teeth on the tongue
Adult lampreys possess functional eyes, dorsal fins, an additional semicircular canal, a cerebellum,
separate dorsal and ventral roots of the spinal nerves, and a spiral-like rather than a straight intestine
Differences between the two jawless fish - mode of reproduction
o Hagfish spawn repeatedly during their lives, and produce a few large eggs each time
o Lampreys produce many small eggs and the adults die after spawning

Primitive bony fishes
Subclass coelacanthimorpha: the living coelacanths
Extinct fishes that represented a side branch of the lineage that presumably gave rise to the tetrapods
Conservative sarcopterygian fishes that had gone unchanged in many respects since the Devonian
Latemeria, and coelacanths, are not ancestral to the tetrapods but represent an offshoot lineage within
the sarcopterygians
o Drifts in the water column with the currents, sculling with its paired fins in an alternating
diagonal pattern
o Highly electro sensitive as are most primitive fishes
o Detects weak electric currents via a unique series of pits and tubes in the snout called the rostral
organ similar to the enlarged ampullae of Lorenzini in sharks

The living coelacanths, at least for now
Coelacanths occupy a unique place in the consciousness of man: they represent a level of tenacity and
immortality which man will never achieve during his short stay on earth

Subclass Dipnoi, Order Ceratodontiforms: the lungfishes
Lungfishes dipnoans because of their two methods of breathing
Though to be a reptile because of the structure of its lung and the placement of the nostrils near the lip
Protopterus proclaimed to be an amphibian based on its heart structure
Recent cladistics analyses indicate that tetrapods are another subclass within the Sarcopterygii.
o Makes lungfishes in fact phylogenetically closer to tetrpaods and hence to amphibians than to
most other bony fishes
A general and distinctive characteristic existence and location of massive toothplates
o Teeth are not attached to the jaw margins as in most other living fishes, but instead occur only
on interior bones
o Often quite large and apparently function in crushing aquatic insects, crustaceans, and mollusks
African lungfishes are best known for their ability to survive desiccation of their habitats during the
African dry season

Class Actinopterygii, subclass Cladista: bichirs and Reedfish
In poorly oxygenated water, bichirs are obligate air breathers and will drown if denied access to the
surface
Bichers are unique in that they use their dorsally placed spiracles to exhale (not inhale) spent air from
the highly vascularized and invaginated lungs
Spiracles serve no apparent aquatic respiratory function
 Polypterids are unique in that they inhale through their mouths by recoil aspiration
o Use the elastic energy stored in their integumentary scale jacket during exhalation to power
inhalation of atmospheric air
Referred to as flagfins: because of the 5-18 dorsal finlets each consist of a vertical spine to which are
attached horizontal rays, giving them a flag and pole appearance
In all other ray-finned fishes, the dorsal fin rays emerge as vertical bony elements from the body of the
fish
Pectoral fin is love-shaped but constructed differently from the lobe fins of lungfishes and
crossopteygians

Class Actinopterygii, Subclass Chondrostei, Order Acipenseriformes: sturgeons and paddlefishes
Although considered primitive actinopterygians, the extant acipenseriform sturgeons and paddlefishes
are highly derived
Share a number of characteristics such as cartilaginous skeleton, heterocercal tail, reduced squamation,
more fin rays than supporting skeletal elements, unique jaw suspension, and a spiral valve intestine
Skeletons are secondarily so ancestral

Acpineseridae
All species spawn in in freshwater, although some species move seasonally between marine and
freshwater and some are technically anadromous
Sturgeons can be identified by the four barbels in from of the ventrally located mouth, five rows of bony
scutes on a body, heterocercal tail, elongate snout, a single dorsal fin situated near the tail, no
brachiostegal rays, and largely cartilagionous endoskeleton, including an unconstructed notochord
Protrusible mouth can be extended very rapidly
They are exceptionally long-lived
Fecundity is relatively high

Polyodontidae
Date back to at least to the Early Cretaceous
Larvae similar to those of sturgeons, and retain the heterocercal tail, unconstructed notochord, largely
cartilaginous endoskeleton, spircle, spiral valve intestine, and two small barbels
Bony scutes are missing and the body is essentially naked except for patched of minute scaled
Not benthic swimmers but instead move through the open waters

Subclass Neopterygii, Order Lepisosteiformes: the gars
Gars and Bowfin
Traditionally considered members of the order Holostei
Holosteans are paraphyletic
Neopterygians
o Shared jaw, tail, and dermal armor characteristic
Gars have entirely ossified skeletons
Primitiveness is evident in their hinged, diamond shaped, interlocking ganoidlike scaled and abbreviate
heterocercal caudal fin





Order Amiiformes: the bowfin
Considered more derived than the cars
Retains the abbreviate heterocercal tail and rudimentary spiral valve intestine of more primitive groups,
but has teleost-like amphicoelous vertebrae as well as cycloid scales (scale type in which the ganoid and
dentine layer have been lost
Distinct among all living fishes in possessing a single, medial gular plate on the underside of the head
Incapable of surviving in warm, deoxygenated water without access to atmospheric oxygen
As in gars, bowfin gulp air and pass it to a highly vascularized gas bladder

Conclusions and Summary
Teleosts are the most successful fishes alive today, but a few highly derived species of several primitive
groups represent the successful fishes of the past
Cephalochordae lancelets are arguably fishes that lack most chordate structure
Lampreys and hagfishes are jawless fishes that are probably convergently similar
Coelacanths were though to have gone extinct until a live on was captured
o With lobe fins, diphycercal tail, hollow spines, a specialized notochord, jointed skull, young born
alive, and tetrapod-like locomotion
Lungfishes lack jaw teeth but have unusual toothplates on the mouth roof and floor
The most primitive actinopterygian fishes are the highly derived, relict, chondrostean sturgeons and
paddlefishes
Bichirs and reedfish of Africa have been variously placed with the lungfishes, lobefins, and rayfins
because they have larvae with gills, lobelike fins, ganoid scaled, and a modified heterocercal tail
Two living orders represent close ancestors of teleosts
o Lepisosteiform gars
Breath atmospheric oxygen via a highly vascularized gas bladder
Interlocking ganoid scaled
o Bowfin
Closer to the teleosts
Can also breath atmospheric air and are predaceous
Cycloid scales, biconcance vertebra, a large gular plate, an elongate dorsal fin, and the
males guard the young for an extended period

Lecture 6
Teleost perfect bone
o Inhabit the widest range of habitat types and show the greatest variation in body plans and
foraging and reproductive habits
o Aside from the wide range of habitat, there are flying, walking, and immobile telesots, and
annual telests that emerge from testing eggs when it rains and then breed and die

Teleostean phylogeny
Modern teleosts arose during four major radiations that produced the subdivision osteoglossomorpha,
Elopomorpha, Otocephala, and Euteleostei
Synapomorphic characters that unite the teleosts
o Numerous bones of the tail and skull
o Uroneural bones
o Mobile premaxillary bone rather than having the premaxilla fused to the braincase
Essential for upper jaw protrusion and allows a fish to shoot its mouth forward during prey
capture
 Advances in locomotion and feeding apparently lead to their success

A survey of living teleostean fishes
Subdivision Osteoglossomorpha
Bony tongues
Most primitive living telesosts
Arawana

Subdivision Elopomorpha
Leptocephalus pointed head
Typically willowleaf or ribbon shaped
Thin and fragile appeareance, this effect is heightened by a lack of red blood cells, which makes them
translucent
Reduction in the number of uroneural bones in the tail as compared to osteoglossomorphs

Subdivision Otocephala, Superorder Clupeomorpha

Subdivision Euteleostei

Subdiviosion Euteleostei, Superorder Protacanthopterygii
Undergone repeated revision
o Orders
Argentiniforms
Deepsea inhabitants
Salmoniformes
Salmonoidei
Osmeroidei

Neognathi
Remaining euteleost, are collectively placed in the unranked taxonomic category Neognathi
o Order
Esociformes
Pikes and mud-minnows viewed as the sister group of all higher euteleosts
because they share a distinct type of tooth attachment and a skeleton made up of
acellular bone
Neoteleostei

Superorder Stenopterygii
Stomiiforms
o All deepsea fishes of the mesopelagic and bathypelagic regions
o Often characterized by long teeth uniquely attached to jaw bones, large mouths, histologicall
unique photophores that include a duct, and a peculiar ventral adipose fin ahead of the anal fin
in some

Superorder Ateleopodomorpha
Jellynose fishes
Bulbous-headed, elongate species that swim just above the bottom in deep water
Skeleton is largely cartilaginous

Superorder Cyclosquamata
Fishes more advanced than ateleopodomorphs are sometimes referred to as eurypterygians
Most primitive superorder in this group is the cyclosquamata
Aulopiforms
o Include truly bizarre giganturic telescope fishes which undergo a spectacular motomorphosis

Superorder Scopelomorpha
All fishes above the level of the cyclosquamata have lost the fifth pharyngeal toothplate and the muscle
that lift its
Advanced groups are often termed the Ctenosquamata in reference to the predominance of ctenoid
scaled among them

Acanthimorpha: the spiny teleosts
Neoteleost above scopelomorphs possess true fine spines and are termed acanthomorphs
Appearance of true fin spines, rather than hardened segmented rays, marks a major evolutionary step in
the evolution of bony fishes
Locomotion was improved by the strengthening of vertebral accessories, providing body stiffening and
better attachment for muscles
Superorder Lampriomorpha
Most primitive members of a major taxon retain certain ancestral traits but possess others indicative of
advanced status
Lampriforms lack true spines, but the mailla helps move the premaxilla and bears no teeth

Superorder Polymixiomorpha
Beardifishes
Possess advanced characters such as four to six true spines in the dorsal fin and four spines in the anal
fin, and their pelvic fins are located fairly forward on the body

Superorder Paracnthopreygii
Many paracanthopterygians have sonic muscles on their gas bladders and produce souns

Summary
Teleosts aros in the early meozoic and radiated as modern osteoglossomorphs, elopomorphs,
otocephalanas, and euteleosts
Osteoglosommorphs
o Tropical
o Freshwater
o Tongue teeth bite against the mouth roof
o Produce and detect weak electric fields
Elopomorphs
o Ribbon-shaped leptocephalus larva
o Tenpounders, tarpons, bonefishes, spiney eels, and true ells
o Predominantly marine that occur from very shallow to very great depths
Otocephala
o Clupeomorpha
Generally small, schooling fishes of pelagic marine and occasionally freshwater habitats
Otophysic ear-to-gas-bladder connection and by bony scutes on the belly
 Herrings and anchovies are axceedingly important fisheries species
o Ostariophysi
Milkfishes, minnows, suckers, characins, loaches, catfishes, and S.A. knifefishes
Modified anterior vertebrae, making up the weberian apparatus that aids in hearing
(absent in more primitive anotophysi)
Produce and respond to alarm substances
Cypriniforms posses pharyngeal jaws and dentition used in manipulating and crushing prey
and vegetation
Characines are highly successful S American and African fishes such as piranhas and tetras
Euteleostei
o Protacanthipterygians
Loosely related marine, freshwater, and diadromous fishes
Deepsea argentiniforms, salmons, and smelts
Whitefishes, graylings, salmons, and trouts salmoniforms
o Adipose fine, triangular flap at the base of the pelvic fin and configuration
of the last three vertebrae
o Many salmonids undergo extensive migrations between fresh and salt
water during their lives
Osemeriforms are generally small, silvery, elongate, water column dwelling fishes
such as FW smelts, deepsea barreleyes and slick heads and southern hemisphere
Salamanderfish and galaxiids
Euteleosts more advanced than protacanthoptrygians are referred to as neognaths more advanced
than the esociform pikes the remaining: Neoteleostei
Superorders of neoteleosts share similarities in the articulation between the skull and first cervical
vertebra, two muscles that move the pharyngeal jaws, and jaw teeth that can be depressed posteriorly
First four superorders of neoteleosts pelagic fishes
o Stenopterygians
Deepsea fishes with long teeth, large mouths and peculiar photophores including
bristlemouths and marine hatchetfishes
o Ateleopodomorph jellynose
Swim just above the bottom and have cartilaginous skeletons
o Cyclosquamates
Primarily deepsea forms such as the bizarre giganturid tesecopefishes and tripodfishes but
include the shallow water lizard fishes
o Scopelomorphs are primarily lanternfishes
Neoteleosts above scopelomorphs possess true fin spines and are termed acanthomorphs
Other acanthomorph advances:
o Strengthened vertebral accessories and tail structures that improved swimming
o Pharyngeal tooth diversification
o Improved jaw protrusion
o Lampriomorphs (tube-eyes oarfish and polymixiomorphs (beardifshes) world longest teleost
Superorder Paracanthopterygii consists of FW cavefishes and trout-perches, but mostly of marine,
benthic, nocturnal fishes, including very deepsea ophidiiforms, commercially important cods,
acoustically active toadfishes, anatomicall specialized batfishes and handfishes, and the diverse
bathypelagic lophiiform angler fish



Lecture 7
Introduction
Jaw mobility and protrusibility are maximal
Pharyngeal dentition and action reach their highest level of development
Series Mugilomorpha
Are a family of nearshore, catadromous fishes of considerable economic importance and of some
taxonomic controversy
Separated spiny and soft dorsal fins and spines in the pelvic and anal fins inclusion with other
acanthopterygians
Primitive in that some have cycloid scales or intermediate between cycloid and ctenoid and the pelvic
girdle lacks any direct ligamentous or bony connection to the cleithral region of the pectora girdle
Frequently leapf from the water
Frequency of such jumping increases when dissolved oxygen levels are low

Series Aterninomorpha
Silversides, needlefishes, sauries, flyingfishes, halfbeaks, killifishes, topminnows, and livebeares
Unique way of protruding the jaw
Premaxilla does not articulate directly with the maxilla
Terminal or superior mouths

Series Percomorpha
Most advanced euteleostean clade
Common: anteriorly placed pelvic girdle that is connected to the pectoral girdle directly or by a ligament

Series Percomorpha, Order Perciformes, the perchlike fishes
Larges order in the Percomorpha

Suborder Percoidea
Largest perciform order
In contrast with lower telests such as ostariphysans and prtacanthopterygains are characterized by
o Presence of spines in the dorsal, anal and pelvic fins
o Two dorsal fins
o Ctenoid scales
o Pelvic fins in the abdominal position
o Laterally placed and vertically oriented pectoral fins
o Maxilla excluded from the gape
o Physoclistous gas bladder
o Absence of orbitosphenoid, mesocoracoid, epipleural, and epicentral bones
o Acellular bone
o Nerver more than 17 principal caudal fin rays

Suborder Elassomatoidei
Strong convergence with the true sunfishes
Suborder Labroidei
Cichlidae, Pomacentridae< Labridae, Scaridae

Suborder Zoarcoidei
Elongate fishes
 Occupy benthic habitats ranging from tidepools to abyssal depths

Subotder Notothenioidei
Ice0fishes
Restricted to high latitude of the southern hemisphere
Production of a variety of glycoprotein antifreezes

Suborder Trachinoidei
Mostly benthic

Suborder Blennioidei
Small, benthic, marine fishes of tropical and subtropical regions
Possess long dorsal and anal fins and fleshy flaps termed cirri on some part of the head

Suborders Icosteoidei, Gobiesocoidei, and Callionymoidei

Suborder Gobioidei
Usually small, benthic or sand-burrowing fishes, mostly marine but about 10% inhabit freshwater
Lack a gas bladder

Suborder Kurtoidei
Peculiar manner in which males care for the eggs
Males have a hooklike growth of the supraoccipital crest on top of their heads to which the eggs are
attached and where they are carried until hatching

Suborder Acanthuroidei
With one exception, contains a group of medium-sized, compressed fishes with small mouths that
usually form shoals over coral reefs or in nearby habitats

Suborder Xiphioidei
Includes some of the fastest and largest predators in the sea
No pelvic fins, a single caudal keel, and relatively stiff, sharklike pectoral, dorsal, and anal fins in the
swordfish

Suborder Scombroidei
Include some of the larges and most economically valuable predators in the sea
Non-protactile mouth in more advanced groups

Suborder Stromateoidei
Tropical and warm temperate fishes of the open sea that often associate as juveniles with floating or
swimming objects
Thick-walles sac in the pharyngeal region that contains teeth made from hardened papillae

Suborder Anabantoidei
Labyrinth fishes because of a complexly folded, auxiliary breathing structure deried from the
epibranchial of the first gill arch located above the gill chamber


Suborder Channoidei
Snake heads
Highly predatory freshwater fishes
Suprabranchial breahing organ reminiscent of that of the anabantoids and are primarily swamp dwellers

Suborder Caproidei
Were though to be pre-perciforms
Medium-sized, reddish, deep-bodied, rhomboid, schooling fishes of moderate depths

Series Percomorpha: advanced percomorph orders flatfishes and twisted jaws
Order Pleuronectiforms
Flatfishes are distinctive compressed acanthopterygians that all share certain features, most noticeably
a marked asymmetry that includes having both eyes on the same side of the head in juveniles and adults

Order Tetraodontiforms
Pinnacle of teleostean evolution is reached among the highly derived fishes of the order
Tetraodontiformes
Common pattern of four teeth in the outer jaws of puffers
High degree of fusion or loss of numerous bones in both the head and body
Tend to eat animals that are generally unavailable to most other reef fishes, such as sponges, sea
urchins, hard corals and jellyfishes



























Summary