Fish or Mammals?

Case study
Background
Cetaceans (sih-TAY-shuns) are a group of animals made up of about 90 different species, including
porpoises, dolphins, and whales. Like fish, Cetaceans spend their whole lives in the water. But, like
mammals, they need to come to the surface to breathe air.

Through the centuries, scientists have used multiple lines of evidence to classify Cetaceans. As new
lines of evidence have become available, we have been able to understand Cetaceans’ relationship
with other animals at finer levels of detail. Fairly recently, scientists were finally able to identify Ceta-
ceans’ closest living relative—the animal with which they share the most recent common ancestor.

Follow along to see what evidence scientists used. Analyze it to learn what it showed them.

pygmy right whale harbor porpoise

common dolphin

tiger shark Atlantic bluefin tuna

elk

house cat sea otter

sockeye salmon

© 2016 University of Utah Updated July 19, 2017 1

Evidence from Anatomy
Before the early 1700s, people disagreed about whether cetaceans should be classified as mam-
mals or as fish. The table below lists some of the observations about mammal, cetacean, and fish
anatomy that were available 300 years ago.

Mammals Cetaceans Fish

Internal skeleton Internal skeleton Internal skeleton

Warm-blooded Warm-blooded Cold-blooded

Lungs Lungs Gills

Forelimb bones (mouse) Flipper bones (whale) Pectoral fins (shark)

Live birth Live birth Lays eggs (with a few exceptions)

Mammary glands/milk Mammary glands/milk No mammary glands/milk

Adults have bare skin, but Skin is usually covered with

Skin is covered with fur
embryos have fur scales, sometimes bare

4-chambered heart 4-chambered heart 2-chambered heart

Brain anatomy (cat) Brain anatomy (whale) Brain anatomy (shark)

Flippers and a tail for moving in Fins and a tail for moving in
Four limbs for moving on land
water water

Embryos have 4 limb buds. The Embryos have 4 fin buds. The
Embryos have 4 limb buds. The
front two become flippers, and front two become pectoral fins,
front two become forelimbs, and
the rear two are absorbed back and the rear two become pelvic
the rear two become hindlimbs.
into the body. fins.

© 2016 University of Utah Fish or Mammals? Case study 2

Evidence from Fossils
In 1839, an anatomist determined that a fossil previously thought to be a dinosaur was actually an
ancient whale that shared characteristics with mammals. The fossil, which he called Zeuglodon,
revealed two key details:

• The Zeuglodon fossil looked different from any living whale, showing that whale species have
changed over time.
• The Zeuglodon fossil had teeth with two roots. Reptiles (including dinosaurs) have teeth with
one root. Most land mammals have teeth with two roots.

0 1 meter
Skeleton redrawn from Gidley (1913)
Tooth inset from Lucas (1901)
Zeuglodon

You can see that Zeuglodon shares a lot of characteristics with modern orca:

Modern Orca The orca spends its life entirely in the water. It can be found in all of
the world’s oceans. It grows to be about 20 feet (6 meters) long, and it hunts fish,
seals, and other animals for food.

Powerful tail muscles

anchored to spine
Nostrils

Ears well suited for Pelvis & internal

0 1 meter
hearing underwater femur bones

Image drawn from Clement & Schlader

© 2016 University of Utah Fish or Mammals? Case study 3

Evidence from Fossils (cont.)
Paleontologists have uncovered a series of fossils with anatomical characteristics that are in be-
tween those of whales and land mammals. This evidence shows how the ancestors of Cetaceans
changed over time.

Dorudon With flippers and tiny hindlimbs, Durodon wouldn’t Discovered: 1845
have been able to move on land. Its fossils, which are an estimated
36–40 million years old, have been found in coastal areas around
the world. At 16 feet (5 meters) long, Dorudon lived during the
same time period as Zeuglodon, but was much smaller in size.

Powerful tail muscles

anchored to spine

Nostrils

Pelvis & small,

external hind legs
0 1 meter
Ears well suited for
hearing underwater

Image modified from Gingerich et al (2009)

Rodhocetus probably spent time both in the water and on land. It Discovered: 1994
probably moved speedily through water but quite awkwardly on
land. Its fossils, which are about 46-47 million years old, have been
found in modern-day Pakistan.
Powerful tail muscles
Nostrils anchored to spine

Ears could hear well in

water and pretty well in air

0 1 meter Webbed toes

Image modified from Gingerich (2010)

© 2016 University of Utah Fish or Mammals? Case study 4

Evidence from Fossils (cont.)

Ambulocetus means “walking whale.” Ambulocetus probably Discovered: 1993

moved comfortably both in the water and on land. Its fossils,
which are about 49 million years old, have been found in
modern-day Pakistan.
Tail muscles helped to
push the animal
Nostrils through the water.

Hooved toes

Ears could hear pretty 0 1 meter

well in water and air

Image from Thewissen & Dillard (2014). Used with permission.

Pakicetus probably moved swiftly on land and spent most of its Discovered: 1983
time there. However, molecular test results suggest that it ate
mainly fish and other animals that lived in the water. Its fossils, which are about
50 million years old, have been found in modern-day Pakistan.

Nostrils Small attachments

for tail muscles

Ears could hear

somewhat in water but
better in air.

Hooves

0 1 meter Image from Thewissen & Dillard (2014). Used with permission.

© 2016 University of Utah Fish or Mammals? Case study 5

Fossil ankle bones
Modern Cetaceans do not have ankles or ankle bones. But fossil whales do. The anatomy of fossil
feet and ankles showed that ancestral whales had hooves. Animals with hooves are called ungulates
(UHN-gyoo-litz).

Ungulates are further divided by whether they have an odd or even number of toes. Odd-toed
ungulates have 1 or 3 toes—for example horses and rhinoceroses. Even-toed ungulates have 2 or 4
toes—for example pigs, deer, camels, and hippopotamuses.

Ankle bones from Ankle bones from

Fossil ankle bones odd-toed ungulates
even-toed ungulates

Rodhocetus Pakicetus Pig Deer Rhinoceros Horse

(Ancient)

Pakicetus and pig bones © 2016 J. G. M. 'Hans' Thewissen

Evidence from Embryos

Notice how the position of the nostrils and the shape of the hindlimbs in the series of fossil whales
changed over time.

Similar changes happen over time in the nostrils and hindlimbs of developing dolphin embryos:

Younger Older

Nostril
Nostril
Nostril
Nostril
Forelimb Nostril Forelimb Forelimb Forelimb
Forelimb

Hindlimb Hindlimb (No Hindlimb)

Drawings from the Digital Library of Dolphin Development:

http://web.neomed.edu/web/anatomy/DLDD/index.html

© 2016 University of Utah Fish or Mammals? Case study 6

DNA Evidence: Comparing Amino Acid Sequences
Whales make milk to feed their babies. Caseins are nutritional proteins that are found in milk. Since
all mammals make milk, they all have genes that code for casein. In the mid-1990s, one group of
researchers decided to investigate which mammals are most closely related to whales by looking at
the amino acid sequences of casein proteins:

68.4%

63.4%

83.4%

76.9%

66.5%

66.7%

38.4%

DNA Evidence: Transposons

Transposons are chunks of
DNA from viruses that get
inserted randomly into a cell’s
genome. They are commonly
found in the DNA of most
living things.

The neat thing about trans-

posons is that once they pop
into an animal’s germline (the
cells that give rise to eggs and
sperm), they are passed to all
of its offspring.

At this point, the transposons

have usually been inactivated.
They do not code for proteins,
and they cause no harm. But organisms that carry a transposon continue to pass it to their offspring,
for thousands or even millions of generations.

Only the descendants of the original ancestral animal will have the transposon in that particular
place in their genome.

© 2016 University of Utah Fish or Mammals? Case study 7

DNA Evidence: Transposons (cont.)
The tree shows the relationships among groups of ani-
mals with hooves (ungulates).

Even-toed ungulates can be further divided into rumi-

nants (ROOM-in-entz), which share a certain kind of
stomach anatomy, and another group that contains pigs
and camels.

To find out where cetaceans fit on this tree, researchers looked for 5 different transposons in specific
locations in the genomes of several ungulates. The table shows what they found.

© 2016 University of Utah Fish or Mammals? Case study 8

References
Clement, N. & Schlader, R. The Idaho Virtualization Laboratory (IVL) at the Idaho Museum of Natural History and Port
Townsend Marine Science Center, Orca Bone Atlas. Rendering of a 3D Model. Retreived August 15, 2016 from http://
www.ptmsc.org/boneatlas/

Gidley, J. W. (1913). A recently mounted zeuglodon skeleton in the United States Museum. Proceedings of The United
States National Museum, 44, 649-654. Retrieved from http://www.biodiversitylibrary.org/part/52500#/summary

Gingerich P. D. (2010, December 27). Research on the Origin and Early Evolution of Whales (Cetacea). Retreived August
15, 2016 from http://www-personal.umich.edu/~gingeric/PDGwhales/Whales.htm

Gingerich P.D., ul-Haq M., von Koenigswald W., Sanders W.J., Smith B.H., Zalmout I.S. (2009). New Protocetid Whale
from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and Sexual Dimorphism. PLoS ONE 4(2):
e4366. doi:10.1371/journal.pone.0004366

Lucas, F.A. (1901). Animals of the past. London. Retrieved from http://www.biodiversitylibrary.org/item/48943#page/107/
mode/1up

Rogers, A.R. (2011). The Evidence for Evolution. Chicago, IL, USA: The University of Chicago Press.

Thewissen, J.G.M., Cooper, L.N., George, J.C. & Bajpai, S. (2009). From land to water: the origin of whales, dolphins, and
porpoises. Evolution: Education and outreach 2, 2, 272-288. doi: 10.1007/s12052-009-0135-2 (open access)

Thewissen, J.G.M. & Dillard, J. (2014). The Walking Whales: From Land to Water in Eight Million Years. University of
California Press. Used with permission.http://www-personal.umich.edu/~gingeric/PDGwhales/Whales.htm

Thewissen, J.G.M., Williams, E.M., Roe, L.J. & Hussain, S.T. (2001). Skeletons of terrestrial cetaceans and the relationship
of whales to artiodactyls. Nature 413, 277-281. doi: 10.1038/35095005

Gatesy, J., Hayashi, C., Cronin, M. A. & Arctander, P. (1996). Evidence from milk casein genes that
cetaceans are close relatives of hippopotamid artiodactyls. Molecular Biology and Evolution 13, 7, 954-963.

Nikaido, M., Rooney, A. P. & Okada, N. (1999). Phylogenetic relationships among cetartiodactyls based on insertions of
short and long interspersed elements: hippopotamuses are the closest extant relatives of whales. Proceedings of the
National Academies of Science, 96, 10261-10266.

Rogers, A. R. (2011). The Evidence for Evolution. Chicago, IL, USA: The University of Chicago Press.

Accession numbers for casein amino sequences (GenPept/UniProtKB/Swiss-Prot):

Q27952.1 (fin whale; Balaenoptera physalus)
F5CIN1 (pygmy right whale; Caperea marginata)
P02668.1 (cow; Bos taurus)
P02670.2 (goat; Capra hircus)
P11841.2 (pig; Sus scrofa)
Q28441.1 (hippopotamus; Hippopotamus amphibius)
P79139.1 (camel; Camelus dromedarius)
P11840.2 (water buffalo; Bubalus bubalis)
P06796.2 (mouse; Mus musculus)

© 2016 University of Utah Fish or Mammals? Case study 9