EEB 3273/Schwenk

LAB 1: DIVERSITY OF LIVING VERTEBRATES

Reading: Homberger & Walker—Preface; Ch. 1; p. 14

Introduction
The evolutionary history of vertebrates spans approximately 500 millions years. Although this may seem like
an immense time span to us, vertebrates actually occupy a relatively brief period of geologic time when one
considers that the earth is approximately 4.6 billion years old. Time spans of this magnitude are difficult for us
to comprehend so it helps if we compress the history of the earth into one calendar year. Using this scale, the
earth came into existence on January 1st. December 31, midnight, is the present time. The first evidence of life,
fossil bacteria, appeared on April 4. It is not until November 20 (520 mybp) that we find fossil evidence of the
first vertebrates. Jawed fish evolved on Nov. 30, the first amphibians invaded land on Dec. 3, and early reptiles
appeared on Dec. 7. The first mammals evolved on Dec. 16 and the oldest known Homo sapiens only appeared
around 11:45 p.m. on Dec. 31! Despite the recent origin of vertebrates, various lineages (both living and
extinct) have diversified into a remarkable array of morphological forms.
There are seven ‘classes’ of living vertebrates. Their presumed relationships are shown on the accompanying
phylogeny. The named groups are all natural or monophyletic groups. In older classifications not all taxa
(formally recognized groups in a classification) are monophyletic. The example of the Reptilia will be
discussed in lecture or lab. Another example is the Osteichthyes. As originally conceived, the Class
Osteichthyes represented all of the bony fish (fish with bony, internal skeletons) and only these fish. This
includes the ray-finned fishes (Actinopterygia), the coelacanths (Actinistia) and the lungfishes (Dipnoi) on the
phylogeny, plus many extinct groups. However, if you follow these lineages back in time to the point on the
phylogeny where they share a common ancestor, you will see that not all of the groups that eventually evolved
from that common ancestor are included in the “Osteichthyes”. One additional important lineage also evolved
from this common ancestor – the Tetrapoda (terrestrial, limbed vertebrates), the group that includes human
beings. A group that contains an ancestor and some, but not all, of its descendants (such as leaving tetrapods
out of the Osteichthyes) is called a paraphyletic group. Compare the paraphyletic groups discussed with some
monophyletic groups (e.g., Mammalia and Sauropsida) and be sure you understand the difference. In modern
usage a name for a group that is paraphyletic is put in quotation marks. So the old use of Osteichthyes to mean
only the bony fish excluding their tetrapod descendants would be written “Osteichthyes.” In order to restrict
our classifications to monophyletic groups, Osteichthyes (without quotes) is now recognized to include the
tetrapods, just as Reptilia includes the birds (“Reptilia” represents the old use of the term, meaning reptiles
exclusive of birds, sometimes called the non-avian reptiles for accuracy).

Today’s Lab:
Use the phylogeny, classification and list of diagnostic traits, in conjunction with the specimens on the tables, to
distinguish the different vertebrate groups (‘Classes’). Become familiar with examining specimens of all kinds,
including skins, skulls, skeletons, microscope slides, wet-mounts, etc. THESE SPECIMENS ARE
EXTREMELY VALUABLE—PLEASE HANDLE THEM WITH CARE! Each number in the list below
represents a ‘station’, i.e., part of one of the lab benches where the specimens for that section are laid out.

(1) Phylum Chordata—Non-Vertebrate Chordates + Agnatha (jawless fishes)

In addition to the representatives of the various vertebrate groups, there are also specimens of non-
vertebrate chordates. Vertebrates are generally considered to be a ‘subphylum’ within the phylum
Chordata. The group of chordates most closely related to vertebrates is the Urochordata (also called
‘ascidians,’ ‘sea squirts’, or ‘tunicates’). Adult tunicates don’t look much like vertebrates, but their larvae
do (see microscope slide under dissecting scope). The larvae are called ‘tadpoles’ and they have a mouth,
a pharynx with pharyngeal bars for filter-feeding and a muscular tail stiffened by a notochord that is

EEB 3273—Lab 1—page 1

 used for swimming; (you won’t be able to see these things with our dissecting scopes). We believe that
the ancestral vertebrates looked something like a tunicate larva that eventually evolved a head and a
skeleton, etc. The most distantly related chordate group is the Cephalochordata (= “head-chord”) calle.
Until recently, cephalochordates were thought to be the group most closely related to craniates, but
molecular studies have confirmed that it is the tunicates, instead. Cephalochordates include a little fish-
like animal that burrows in the sand and filter-feeds called Branchiostoma, but most often people refer to
it by its old or common name, Amphioxus. This animal also has a mouth leading to a pharynx supported
by pharyngeal bars and a muscular tail stiffened by a notochord, with a dorsal, hollow (tubular) nerve cord
above it. It also has segmented body musculature and probably looks very much like an early or ancestral
vertebrate. Note, however, that it lacks a head—the notochord ends at the very front end with nothing in
front of it. In vertebrates with heads, the notochord ends at the back or the brain/skull Compare the
amphioxus to the larval lamprey (a vertebrate) at the next station—called the ammocoetes larva (it is a
lamprey even though it has a different name – don’t ask me why). Note how similar. The main
differences are that the larval lamprey has more of an actual head with a brain and an eye. Also note that
the number of its pharyngeal bars is greatly reduced.
Look at the non-vertebrate chordate specimens (and also the jawless vertebrates, the hagfish and
lampreys) we have in the lab. Try to observe the following where you can:
• Mouth
• Pharynx
• *Pharyngeal bars (pharyngeal skeleton)
• *Pharyngeal slits (open from pharyngeal pouches) (function as gills in the lamprey and
other fishes)
• *Muscular tail (post anal)
• *Notochord that runs length of tail, stiffening it for swimming
• Note: water passes into the mouth and out the pharyngeal slits; food particles are trapped in the
pharynx and moved into the gut for digestion; this is known as filter- or suspension-feeding

*key chordate character (shared by all chordates, including vertebrates)

Vertebrate Diversity
Familiarize yourself with the names and characteristics of the living groups of vertebrates. Note that most
of the vertebrates you are familiar with have mouths with moveable jaws. These are known as the ‘jawed
vertebrates’ or Gnathostomata (gnatho = jaw; stoma = mouth). The earliest vertebrates, like their closest
chordate relatives, lacked jaws. The earliest fishes were all jawless and there was quite a lot of early
jawless fish diversity. However, since the gnathostomes evolved, they appear to have out-competed the
jawless fishes and only lampreys and hagfish remain. Lampreys and hagfish are ‘primitive’ in their lack
of jaws, their reduced, cartilaginous skeletons and in other aspects of their anatomy, but they are also very
specialized and ‘derived’ in other traits. Both groups are now parasitic or exist on detritus, and therefore
have a very reduced morphology. These groups will be discussed further in lecture.
The jawed vertebrates include most of the fishes you are familiar with as well as the Tetrapoda
(terrestrial, limbed vertebrates). The living fish that are most closely related to the tetrapod vertebrates are
the lungfish, or Dipnoi. They have lungs for air-breathing, as well as gills, and also have robust, muscular
fin bases with internal, bony skeletal supports.
Several tetrapod groups (e.g., cetacean and pinniped mammals) have secondarily returned to the water and
modified their limbs into fins (an example of a ‘reversal’ in character evolution). However, their fish-like
bodies and aquatic traits are all secondarily acquired adaptations to life in the water and these animals are
still ‘tetrapods’, i.e., they belong within the Tetrapoda because they descended from limbed, terrestrial
ancestors.

EEB 3273—Lab 1—page 2

(2) Chondrichthyes (cartilaginous fishes, sharks, rays, skates + chimaera or ratfish)
Characteristics of Chondrichthyes (cartilaginous fishes):
paired fins
cartilaginous internal skeleton
dermal denticles
males have claspers on anal fins
external gill openings uncovered or covered by skin only
no air bladder
two-chambered heart
mouth ventral
(3) Osteichthyes (bony fishes) not including tetrapods
Characteristics of Osteichthyes not including tetrapods (Actinoptyerygia, Actinistia, Dipnoi):
paired fins
some or all of skeleton is bone
most with bony scales in skin
bony operculum covers gills
lungs or swimbladder
mouth usually terminal
(4) Amphibia (Anura/frogs, Urodela/salamanders, Gymnophiona/caecilians)
Note: Although the entire group, including all fossil species, is called the Amphibia, the living species are often
grouped together under the name Lissamphibia.
Characteristics of Lissamphibia (the living amphibian groups—caecilians, salamanders, frogs):
paired limbs (lost in caecilians and some salamanders)
limbs laterally placed
moist, glandular skin
pedicellate teeth
"amphibian" papilla of ear
lay eggs in moisture or are live-bearing
three-chambered heart
respiration by gills, lungs, skin, and/or mouth lining
(5) Reptilia (crocodylians, birds, turtles, tuatara, snakes, lizards)
Characteristics of non-avian reptiles (reptiles except for birds):
paired limbs (or secondarily lost as in snakes)
limbs laterally placed
larteral, undulatory movement
dry, scaly skin, often with osteoderms (bones within scales)
partially (most) or completely (crocs) divided ventricle in heart
lay amniotic egg or live-bearing (some squamates)
Characterisitics of Aves (Reptilia—birds):
forelimbs modified as wings
hind limbs under body
body bends dorso-ventrally instead of laterally
feathers
synsacrum
completely divided ventricle in heart (i.e., four-chambered heart)
lay amniotic egg

EEB 3273—Lab 1—page 3

(6) Mammalia (monotremes, marsupials, eutherians/’placentals’)
Characterisitics of Mammalia:
well-developed limbs
limbs ventral, beneath body (sagittal plane)
vertical (dorso-ventral) flexion during locomotion
hair (pelage)
single bone in lower jaw (dentary)
mammary glands
four-chambered heart
chorioallantoic or vitelline placenta and live-birth (except for monotremes, which lay an amniotic egg)

EEB 3273—Lab 1—page 4

 A CLASSIFICATION OF LIVING VERTEBRATES

CHORDATA
CEPHALOCHORDATA and UROCHORDATA (non-vertebrate chordates)
VERTEBRATA (chordates with vertebrae)
AGNATHA (hagfishes, slimehags)
Petromyzontiformes (lampreys)
Myxiniformes (hagfish)
GNATHOSTOMATA (jawed vertebrates)
CHONDRICHTHYES (cartilaginous fishes)
Elasmobranchii (sharks, rays & skates)
Holocephali (chimaeras or ratfish)
OSTEICHTHYES (bony fish and tetrapods)
ACTINOPTERYGIA (ray-finned fishes incl. teleosts)
SARCOPTERYGIA (fleshy-finned fish and tetrapods)
Actinistia (coelacanths)
Dipnoi (lungfish)
TETRAPODA (limbed vertebrates)
LISSAMPHIBIA (living amphibians)
Gymnophiona (caecilians)
Anura (frogs)
Urodela (salamanders)
AMNIOTA (tetrapods with an amniote egg)
SAUROPSIDA (reptiles, including birds, and their ancestors)
REPTILIA (living reptiles and birds)
DIAPSIDA (lepidosaurs and archosaurs)
Testudines (turtles; also Chelonia)
Lepidosauria (tuatara, lizards, snakes, amphisbaenians)
ARCHOSAURIA (crocodilians and birds; also incl. dinos.)
Crocodylia (alligators, crocodiles and gharials)
Aves (birds)
SYNAPSIDA (mammals and their ancestors)
MAMMALIA (living mammals)
Monotremata (monotremes)
Theria (marsupials and placentals)
Metatheria (marsupials)
Eutheria (placentals like us)

EEB 3273—Lab 1—page 5

CEPHALOCHORDATA (amphioxus, lancelet)

UROCHORDATA (tunicates, sea squirts)

EEB 3273—Lab 1—page 6

VERTEBRATA: AGNATHA (larval lamprey = ‘ammocoetes’ or ‘ammocoetes larva’)

EEB 3273—Lab 1—page 7