1

Lab 1 Early chordates and external morphology of fish

Order of lab (divided to ensure staggered use of specimens and timely completion)
 Odd numbered groups Start at 1.1 Even numbered groups Start at 1.4

Introduction
Relationships among early chordates and jawless fishes is still an active area of investigation.
One study (Satoh and colleagues, 2014) suggests that the urochordates are the sister group to
vertebrates. Fish inhabit a variety of habitats, from fresh water to sea water and some tend to live
primarily in the water column (PELAGIC fish) or on or near the bottom (BENTHIC fish). There is a great
deal of variation in the fish body plan, with adaptations to their environments and niches.

Taxonomy from Integrated Taxonomy Information System January 2023

(https://www.itis.gov)

Subphylum Cephalochordata
Order Amphioxiformes Branchiostoma sp. (amphioxus)

Subphylum Urochordata
Class Ascidiacea Order Enterogona Corella sp. , Ciona sp. (sea squirts)
Class Thaliacea Order Salpidae Salpa sp. (salp)

Subphylum Vertebrata
Infraphylum “Agnatha”1
Class Myxini Order Myxiniformes yxine sp. (hagfish)
Class Petromyzontida or Cephalaspidomorphi Order Petromyzontiformes Entosphenus
sp. (lamprey)

Infraphylum Gnathostomata
Superclass/Class Chondrichthyes
Subclass Elasmobranchii Order Squaliformes Squalus sp. (dogfish shark)

“Osteichthyes”2
Superclass Actinopterygii (ray finned fishes)
Class Holosteii Order Amiiformes Amia calva ( bowfin)
Class Teleostei
Order Perciformes Perca sp. (perch),
Micropterus sp. (bass)
Order Cypriniformes, Cyprinus sp.(carp)

1Although recent molecular evidence supports monophyly of the extant agnathans (i.e., the cyclostomes), the agnathans are
paraphyletic when the extinct jawless fishes are included (see lecture material). Hence, although some classifications consider
Agnatha a superclass, strictly cladistic classifications no longer use Agnatha as a formal taxonomic name.
2 Although generally no longer used as formal taxonomic name, this is a monophyletic group (i.e., a clade) that includes all remaining
vertebrates.

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Specimens
 Branchiostoma sp. (formerly known as Amphioxus) slides and whole specimens in slide boxes at the head
of each bench
 Corella sp. plastomount at head of each bench
 Other tunicates on the demo bench
 Salp: on the demo bench
 Hagfish: preserved whole specimens at the head of each bench
 Lamprey: preserved whole specimens at the head of each bench
 Perch (Perca): preserved whole specimens at the head of each bench
 Fish scale slides in a slide box at the head of each bench
 Fish specimens to illustrate diversity and for keying on the demonstration bench
 Ganoid scales of the gar on the demo bench

Video playlist to watch before lab

 Branchiostoma (Amphioxus) from Shape of Life

 Hagfish from Smithsonian Institute

 Lamprey predation of paddlefish from National Geographic

Useful word roots

a: Gr. Prefix meaning ‘without’ (agnathan) epi: Gr. On or above (epibranchial)

actin: Gr. Ray (actinotrichia, Actinoptypergii) ex: L. Out of or away (excurrent)

gnath: Gr. Jaw (agnathan)
atri: L. Entrance (atrium)
myo: Gr. Muscle (myomeres)
branch: Gr. Gill (branchial basket)
oper: L. To cover (operculum)
bucc: L. Cheek (buccal cavity)
pteryg: Gr. Wing, fin (Actinoptypergii
ceph: Gr. Head (cephalochordate) Sarcoptyergii)
chord: L. String (chordate) sarco: G. Flesh ( Sarcoptyergii)
cten: Gr. Comb (ctenoid)

endo: Gr. Within (endostyle)

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Anatomical definitions associated with directions

CAUDAL (posterior) - toward the tail LATERAL - toward the side.

CRANIAL (anterior, rostral) - toward the head MEIDIAL - toward the midline.
DORSAL - toward the back MEDIAN - on the midline.
VENTRAL - toward the belly PROXIMAL - part of structure nearest a point of
reference. e.g., with reference to the shoulder, the
DISTAL - part of a structure furthest from a point of
wrist is the distal end and the
reference.

2.1 Cephalochordates

 Examine the diagrams, images and slides of Branchiostoma sp (“amphioxus” adult, whole, and cross
sections). Identify the structures in bold in the description below. The nerve cord is more easily viewed
in cross section.

Feeding: Cephalochordates are marine chordates that bury in the sand and feed on
microorganisms and phytoplankton. They exhibit some specialization related to their mode of feeding.
The CIRRI of the buccal region prevent entry of particles that are too large. The VELUM ensures that
water flow is one way. Mucus secreted by the ENDOSTYLE is moved up the walls of the pharynx by
cilia. Food particles adhere to the mucus, From the pharynx, the muscous moves to the
EPIBRANCHIAL GROOVE and into the gut. Note that the digestive system is divided into the
FOREGUT, MIDGUT and HINDGUT, with a MIDGUT CAECUM. Filtered water passes out of the slits
to the ATRIUM and leaves via the ATRIOPORE.
Support and locomotion: The body is supported by the NOTOCHORD. The notochord prevents
the body from shortening when the V shaped MYOMERES ( blocks of striated muscle fibers that run
the length of both sides of the body) contract. The myomeres occupy a great deal of space and as a
result the coelom is small. The myomeres contract in segments (myotomes). This allows the animal to
swim as it produces lateral undulations that causes water to move toward the posterior and propels the
body forward. The notochord helps to keep the body stiff as the animal burrows. The NERVE CORD
(also referred to as a “spinal cord”) is most identifiable in the cross section. Motor neurons form the
nerve cord innervate the myomeres.
Circulation and gas exchange: Cephalochordates lack a heart. Blood flows anteriorly in the
ventral region, through the gills and then posteriorly through the DORSAL AORTA. Blood is supplied
to the myomeres and organs through a series of arteries. The capillary beds of the tissues converge
on veins which lead into a ventral aorta that carries blood toward the anterior. This general pattern is
common to the other chordates we will examine in the course. Gas exchange occurs across the pharynx
and the across thin flaps of the body wall.
Excretion: The excretory system of cephalochordates is composed of paired NEPHRIDIA (not
usually visible) that open into the ATRIUM and the BUCCAL region

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 What features enable you to determine from where along the anterior to posterior axis, a cross
section is taken in Branchiostoma sp. ?

Figure 1.1: Longitudinal section of cephalochordates (modified from Pough et al., 2009)

A fin ray

nerve cord
notochord
myoseptum
dorsal aorta myomere
epibranchial groove
atrium
gill slit
gill bar
midgut caecum

gonad

lymph space endostyle

metapleural fold

Figure 1.2: Cross section through the pharynx of an adult cephalochordate (modified from
Feduccia, 1975) B
nerve cord
myotome
notochord

area of dorsal aorta

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students of Biology slitin Winter 2021 only.
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2.2 Tunicates
 Examine the images of larval ascidians and the whole and plastomount ascidians
(Ciona sp and Corella sp) and the adult salps ( demonstration bench) .

Tunicates include the salps, larvaceans and ascidians. They are called “tunicates” due to the
presence of the outer covering or TUNIC.
Ascidians may be solitary or colonial and are benthic as adults. They are hermaphrodites but
generally do not self-fertilize. Most shed gametes into the water column where fertilization occurs. The
pelagic tadpole larvae display many of the chordate characteristics. However, when larvae settle to the
bottom and adhere to the substrate, they undergo metamorphosis wherein the notochord and nerve
cord are absorbed with only remnants remaining.
Ascidians feed on plankton suspended in the water column. Water enters the INCURRENT
SIPHON and passes into the BRANCHIAL CAVITY. Water is drawn into the cavity via cilia on the
BRANCHIAL BASKET or PHARYNX. Note the net like structure of the branchial basket. Food particles
(phytoplankton) are trapped in and consumed along with mucus that is continually produced by the
ENDOSTYLE. Food and mucus pass into the ESOPHAGUS which leads to the digestive tract.
Undigested food and water are expelled through the EXCURRENT SIPHON.
Salps are marine pelagic tunicates and may exist as colonies. They have a transparent
cylindrical body with incurrent and excurrent siphons at opposite ends of the long axis. A few pharyngeal
slits are present. Salps move via jet propulsion. The water filled tunic is surrounded by circular muscle
bands that cause water to be expelled from the excurrent siphon when they contract. Water enters via
the incurrent siphon when these muscles relax,

1. Locate the bolded structures above on the Corella sp. plastomount. What differentiates
the incurrent siphon from the excurrent siphon?
2. Which of the adult characters are visible in the image of the ascidian larva in your image
package?

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Figure 1.3: Ascidian tadpole larva and adult (modified from Pough et al., 2009)

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1.3 Jawless fish

 Examine the whole specimens of hagfish and lamprey ( end of bench) and the slides of the whole
ammocoete larva.

Hagfish and lamprey are jawless fishes in the Subphylum Vertebrata. Both lineages have
undergone loss of bone and scale independently. The understanding of the phylogenetic relationships
among them and other chordates has undergone several revisions in the past century.

Hagfish are deep-sea marine predators and scavengers that lack vertebrae and retain the
notochord in the adult. They lack a larval stage and much about their life cycle is unknown. They use
teeth like projections composed of keratin on the tongue to rasp flesh from prey. Hagfish have
rudimentary eyes and three pairs of sensory BARBELS. They have a large NARES/NOSTRIL that
leads to an OLFACTORY SAC. They are well known for their production of mucous that likely deters
predators and helps in their escape from them. Their gills are held in BRANCHIAL POUCHES with
internal and external openings.

Figure 1.4: Hagfish whole (modified from Pough et al., 2009)

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Lamprey have one or two fleshy DORSAL FINS and a caudal fin (not supported by bony
structures). They have well developed eyes and NOSTRIL/NARES is located on the dorsal surface of
the head. Lamprey have prominent sense organs. The LATERAL LINE system is anterior to the EYE.
External GILL SLITS are visible.

About 50% of lampreys are parasites of other fish as adults while the larvae, called
AMMOCOETE, are free living. The parasitic adults have a pronounced BUCCAL FUNNEL /ORAL
HOOD which contains many spines composed of keratin (which are often called teeth) for attaching to
host species
Lamprey can tolerate variations in salinity which has contributed to the ability of some species
to become invasive species.

Figure 1.5: Lamprey external morphology (modified from Wischnitzer, 1972 )

The ammocoete larvae look a great deal like the adult cephalochordates. They bury in the
sediment and feed by straining food particles though pharyngeal gill slits with a current generated by
muscular rather than ciliary action. The remaining lamprey species also filter feed as larvae but do not
feed at all as adults.

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A fin ray
nerve cord
notochord
myoseptum
dorsal aorta myomere
epibranchial groove
atrium
gill slit
midgut caecum gill bar

gonad
lymph space endostyle
Figure 1.6: Longitudinal section of an ammocoete larvae (modified from Feduccia, 1975)
metapleural fold

B
nerve cord
myotome
notochord

area of dorsal aorta

external gill slit

gill bar
gill slit

sub-pharyngeal gland

lingual muscles

Figure 1.7: Cross sections through the pharynx of an ammocoete larvae (modified from
Feduccia, 1975)

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1. Compare the external morphology of the lamprey and hagfish.

2. Locate the nares/nostril, mouth and sensory barbels on the hagfish.
3. Locate the bolded structures on the adult lamprey.
4. In the ammocoete larval microscope slides, identify the MYOTOMES, NERVE CORD,
NOTOCHORD, and area of the DORSAL AORTA. Sections through the pharynx will show the
GILL BARS and GILL SLITS.
5. Examine the cross section series from anterior to posterior in relation to the longitudinal section.
What features are diagnostic of where the section was taken from on the anterior to posterior
axis?

1.4 External morphology and body shape of jawed fish

 Examine figures of the whole specimens of the dogfish and perch. Use the external morphology key o label
the features described below where applicable.

The fish body is divided into three more or less distinct regions, the HEAD, the TRUNK,
separated from the TAIL by the urogenital and anal openings. While some fish are harder to categorize,
the shape of the body and the position of the MOUTH can be an indication of the feeding habits and
habitat. The mouth of the perch is TERMINAL, situated at the anterior end of the body, like many
midwater predators. Fish whose mouths are on the ventral surface (i.e. SUBTERMINAL) are usually
bottom feeders, and those whose mouths are turned upwards (i.e. SUPERIOR) are usually surface
feeders. Examine this linked material on fish mouths from the Florida Museum. In bony fish the
OPERCULUM, a bony plate, covers the GILLS. The ANUS and urogenital openings are located on the
ventral midline just anterior to the ANAL FIN.
In some groups, such as the eels, the gills are much further back on the head and the operculum
is small. Eels rely more heavily on a ‘swallowing” movement to move water past the gills.
Chondrichthyans have septal gills which are each protected in a chamber formed by individual flaps
formed from gill septa.
In the perch there are two pairs of NARES (nostrils). Water passes into the anterior pair, over
the olfactory epithelium, and out the posterior pair. The incurrent naris is quite obvious, surrounded by
a fleshy lip. The excurrent naris ( less obvious) appears as a small break in the skin medial to the
anterior edge of the eye. The dogfish NARES are on the ventral surface, cranial and ventral to the eyes.
The shark has SPIRACLES, small openings caudal from the eyes. The spiracles allow water to pass
over the gills even when the mouth is closed. On the ventral surface of the shark, cranial to the mouth
are some of the more obvious SENSORY PITS.
The EYES are large and lidless, situated laterally on the head. Their fields of view are
independent so the fish does not have binocular vision; however they do provide a wide field of view,
an important adaptation for animals with the head fused to the pectoral girdle which prevents them from
turning it independently of the body.
The LATERAL LINE, an organ of pressure reception (sound waves and water turbulence) can
be seen as a distinct line of perforations through the scales of the LATERAL LINE SCALE ROW which,
in perch, runs the length of the body from the back of the skull to the CAUDAL FIN. The lateral line
tends to be less obvious in the shark. It extends into the head region in a complicated network.

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Figure 1.8: External morphology of the a) perch, Perca sp. and b) the dogfish shark(male) ,
Squalus sp. (‘b’ modified from Feduccia,1975)

Fins enable the fish to control its movements and in some fish are used to generate movement
Fins are generally named for their location : PECTORAL, PELVIC, DORSAL FINS ( single or multiples,
spiny or soft) ANAL and CAUDAL. Not all fish have each fin type. The yellow perch has spines on its
anterior dorsal fin. The spines in the fins of higher teleosts serve a defensive function, making the fish
a less attractive mouthful for a predator. Several species of spiny teleosts have evolved potent venoms
associated with their fin spines.
Sarcopterygian fishes such as lungfish have lobed pectoral fins. Male dogfish have CLASPERS,
projections from the pelvic fin that aid in sperm transfer during mating. Several lineages of fishes,
including the salmonids (salmon, trout and their relatives) have a fleshy ADIPOSE FIN which lacks the
bony structural supports of the other fins. The pectoral and pelvic fins are more functional for turning,
braking and for sculling movements, which assist in fine control of the fish's attitude and position, than
they are for producing lift, which is their primary function in sharks and more primitive bony fishes.

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Fish body shapes range widely. Many are somewhat laterally compressed like the perch. Fish
that constantly swim in the water column in pursuit of prey or that launch themselves in a surprise sit
and wait attack tend to be the most streamlined. The body tends to be COUNTERSHADED (i.e. darker
on the back than the belly). The flatfish takes lateral compression to the extreme being laterally
flattened, lying on one side. Pelagic flatfish larvae metamorphose when they settle to ocean floor
(benthos) such that both eyes are on one side of their heads. Some fish like the skates and rays are
dorso-ventrally flattened
Some key body measurements are reflective of functional traits of fish. Fishes with larger body
depth are generally better at turning and accelerating and their predators must have large gapes to
capture and eat them. Fish with deeper heads can eat larger prey. Fish with wide and deep CAUDAL
PEDUNCLES (part of the trunk posterior to the dorsal and anal fins) tend to be specialized for have
faster starts while fish with narrow tapered caudal peduncles tend to excel at sustained swimming
because there is less drag force created by the lateral motion of swimming

Figure 1.9: Key body measurements for studying the external morphology of fish . Lechêne et
al (2018). PLOS ONE. 13. e0209025. 10.1371/journal.pone.0209025. CC BY.

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Fin number, shape and placement are adapted to habitat and feeding habit. Broad categories of
fish include (but are not limited to), lie in wait predators, rover predators, and bottom dwelling fish. The
caudal fin of the rover, and lie in wait predators is usually forked. Lie in wait predators tend to have an
elongate body with dorsal and anal fins far back on the body. This enables the fish to generate a great
deal of thrust. The relatively even distribution of fins onthe rover predators gives stability and
maneuverability. Bottom dwelling fish tend to be dorsoventrally flattened ( depressed like the skates
and rays though some with flatfish such as halibut and flounder being flattened laterally. The latter
have extremely large pectoral fins that flat and/or undulate to generate movement. Fish with the rattail
shape tend to have large pectoral fins. Eel shaped fish have small or absent paired fins and long dorsal
and anal fins. This is an adaptation for entering small spaces such as holes in reefs.

 Examine the images of various fish, making note of the body shape and the types and
locations of their fins.

Figure 1.10: Selected fish body shapes (modified from Moyle and Cech, 1982).

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1. Locate the bolded structures in the previous descriptions on the images of the perch and dogfish
where applicable ( remember that you have a key to the diagrams.)
2. Compare the fins of the perch, codfish, trout and pike (in your image package) with respect to
type, number and relative position on the body axis.
3. What would you predict about the perch’s habitat and feeding habit, given its body shape and
fin placement?
4. Describe the positions of the pectoral, pelvic and anal fins with respect to each other and the
whole body using proper anatomical terminology. Compare the location of the pelvic fin on the
perch and the dogfish.
5. What do you notice about the symmetry of the perch fin as compared to that of the shark fin?
6. The perch is compressed- what does this mean? How does the shape of the fish relate to
movement through a density fluid such as water?
7. How is countershading of fish likely to be adaptive?

1.5 Scales

Scales are derived from dermal bone which is produced by ossification of the dermis ( a layer
of skin). Originally scales consisted of two bony layers overlain by a layer of DENTINE with an outer
layer of hard dense, highly mineralized material. In the bony fishes, the surface layers were lost,
leaving the bony basal plates. In shark-like fishes, the bony portions were lost, leaving the tooth-like
enamel and dentine outer layers. The primitive scale types, where they are found among fossil fishes,
are always RHOMBIC in shape and they always join edge to edge rather than overlapping. This
complete armor covering underwent reduction very early in the evolutionary history of all groups of
fishlike vertebrates. The scales are much reduced in living bony fish compared to their primitive state.
In all teleosts (large group of actinopterygians), the scales consist only of a thin LAMELLAR BONE
which is underlain by a fibrous connective tissue layer.
Scale types can be grouped according to two different growth patterns. Chondrichthyans have
PLACOID scales. As the fish grows there is an increase in the number of these scales. Other scale
types include the GANOID, CTENOID and CYCLOID scales. These scales grow by concentric
addition, a pattern called CYCLOMORIAL GROWTH. Separate units are deposited in rings around the
edge of each scale, which enlarges as the fish grows. Some fish species have lost their scales or have
scales that are small and embedded or on certain locations of the body. Eels have cycloid scales deeply
embedded in the skin. Most catfish lack scales. Why do you think this might be the case?
Among living fish, GANOID SCALES are found only on gars, bichirs and reedfish. They retain
more of the primitive structure, consisting of a basal plate of lamellar bone and a hard, shiny outer layer
of GANOINE, a highly mineralized, dense, enamel-like substance. They articulate edge to edge, barely
overlapping one another. Ganoid scales have a small foramen which allows passage of blood vessels
and nerves to the overlying skin. In early bony fish ganoid scales formed the skeletal supports for the
fins. In the extant gars, these scales form a protective exoskeleton.
PLACOID SCALES or DENTICLES are unique to the chondrichthyans. They are tooth like
structures composed of a basal plate embedded in the skin with a spine projecting through the
epidermis. The bony portions of the primitive fish scale have been lost leaving a PULP CAVITY which
opens on the undersurface of the basal plate and projects into the spine. A layer of DENTINE forms

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most of the basal plate and the core of the spine. This is covered by ENAMEL which, like ganoine, is a
dense, highly mineralized, glassy substance. Placoid scales may be separate or join edge to edge.

 Examine the diagrams and slides of the scale types .Be able to identify the scale type,
characteristics and be able to name the listed fish that possess them.

Figure 1.11: Structure of ganoid and placoid scales.

CTENOID SCALES are associated with the more advanced teleosts which have spiny fins. The
exposed posterior portion of ctenoid scales is covered with minute spine like projections (CTENI). The
anterior portion of the scale is overlain by adjacent scales. Scale growth starts at the FOCUS, and is
periodic. During periods of growth, numerous, fine, concentric ridges, the CIRCULI, are laid down. In
winter, when there is little to no growth, the scale margin is resorbed, leaving a visible band. The
complete spring circuli of each year are called the ANNULI. These are sometimes used by fisheries
biologists to age fish, especially when other methods are unavailable.
CYCLOID SCALES, like ctenoid scales, consist only of a thin plate of bone. They lack cteni,
however, and the circuli are continuous around the entire scale. Although there are quite a few
exceptions, cycloid scales are generally found among the more primitive teleosts which also lack spines
in the fins.

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Figure 1.12: Structure of ctenoid and cycloid scales

1.6 Fish identification exercise/fish diversity

Dichotomous keys are useful tools for the identification of species and other taxa. They are
usually prepared by taxonomists for a particular geographic area. An identification key for one
geographical area will not necessarily be useful for another. You will use a very simple dichotomous
key prepared for a few of our lab specimens to learn how to use them. We have chosen species that
can be readily separated on the basis of the external characteristics that you have observed in this lab.

 Use the key provided to identify the fish species present. Note that this key is prepared for
these species only. You may be required to identify unknown specimens using a similar key on an
exam. This key was constructed to identify the specimens in this exercise in particular and would
not be useful in a general sense. Record your steps and the species name of each lettered
specimen and have them checked by a TA. Identify the Linnean taxonomy associated with each of
these specimens.

 As you work through the key, compare the external morphology of these fish to that of the perch. Note
the presence or absence of operculi and describe the gills, fins, tail and general body shape. Do any
of these seem to correlate with the diet or habitat?

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Key for selected fish in the Biology 2210 lab

1 A. Jaws and paired fins present ………………………………...........2

B . Jaws and paired fins absent……………………………………..10

2 A. Placoid scales present and bony operculum absent ………….......................................….3

B. Placoid scales absent and bony operculum present (can be reduced in size) ..…………...5

3 A. Dorsoventrally flattened ………………………………………....Raja sp., skate

B. Not dorsoventrally flattened …………………………………….4

4 A. Multiple gill slits …………………………………………………..Squalus sp. , dogfish

B. Single gill opening……………………………………………….Hydrolagus sp., ratfish

5 A. Fins with bony rays ………………………………………………6

B. Fins lacking bony rays………………………………………...…Protopterus sp. , lungfish

6 A. Laterally flattened ….. …………………………………………..Hippoglossus sp.,halibut

B. Not laterally flattened …………………………………………...7

7 A. Ganoid scales present, jaws prolonged with sharp needle like teeth
…..................................Lepisosteu sp., gar
B. Ganoid scales absent, specimen lacks scales or has cycloid or ctenoid scales…………………..8

8 A. Pelvic fin present ………………………………………………... 9

B. Pelvic fin absent, long dorsal fin which is continuous with the caudal fin and coming to a point
………………………………………………………..…….Anguilla sp., eel

9 A. Single dorsal fin, adipose fin on the back between the dorsal and caudal fins, distinct pelvic
axillary process… …………………………………………….Oncorhynchus sp., Trout
B. Three dorsal fins, distinct barbell near tip of lower jaw ….. .Gadus sp. , cod

10 A. Fleshy barbells around small mouth, single low median dorsal and anal fin continuous with
caudal fin…………………………………………….. …………Myxine sp., hagfish
B. Mouth consisting of a cup with horny teeth.. ……………… Petromyzon sp., lamprey

©2023 Joy Stacey and the Department of Biological Sciences. For the individual study of students of Biology 2210 in Winter 2021 only.