BIOLOGY OF SAUROPOD

DINOSAURS
Vertebrate Paleontology
Chan Cheuk Yan Claudia Student ID: 97191026-4
ccycmbbs@akane.waseda.jp
Introduction

Sauropods, or Sauropoda are part of a clade called Saurischian dinosaur, which is another

name for “lizard-hipped” dinosaurs. Their name, sauro- + -pod, directly translates to “lizard-

footed”. These quadruped reptiles have extremely lengthy necks, long tails, small heads and

four pillar-like extremities. They are also herbivorous creatures. They are best known to us to

be the largest dinosaur and terrestrial animal to ever walk this Earth and some of them can

reach up to 30 meters long, weighing at least 30 tons. Some well-known examples of genera

that fall into this clade include Braichiosaurus, Diplodocus, Apatosaurus, Brontosaurus and

Mamenchisaurus.

This clade of dinosaurs has become one of the most recognizable to mankind purely due to

their magnificent size, and their biology had been a major research subject amongst

paleontologists since the 19th century. They first started to appear in the late Triassic Period

(Mesozoic Era) and their population was at its peak by the late Jurassic period, which was

around 180 million years ago. They lived on for another few hundred million years until the

late Cretaceous in the same era, before being replaced by Titanosaurs. Hence, they are

deemed one of the most successful terrestrial animals of all time.

I have chosen to research about Sauropod dinosaurs because it has always been a mystery to

paleontologists and mankind about how this magnificent creature evolved to become so

enormous. The Sauropods have also been my favorite type of dinosaur as I have always been

fascinated about the significance of the length of their necks and what was its purpose.

Therefore, I would like to look into this aspect as well, and about how they evolved to have

such a characteristic.
Subject and Method

In this essay, the main subject would be to discuss the biology of Sauropods and the reasons

for the evolution of their gigantism.

I would firstly explore the various types of the most well-known types of Sauropods and

discuss the characteristics of each. I would then dive into Sauropods’ anatomical features and

their lifestyle, the purposes of them and the pros and cons of its gigantic body size. Finally, I

will discuss the hypothesis that have been set by paleontologists over the years of how the

Sauropod dinosaurs came to grow to such a size.

In order to do this, I would be using the internet, such as various research papers and

scientific journals from NCBI and Science Daily, articles from organizations such as National

Geographic and the American Museum of Natural History, journals from institutions such as

UC Berkeley, and Encyclopedia websites such as Britannica and Wikipedia to obtain external

secondary sources of data to conduct most of my research about the topic. The method of this

research paper would mostly be qualitative as I would be generating a few hypotheses that

could explain the Sauropod’s gigantic body size.

Types of Sauropods and their characteristics

The Sauropoda group of dinosaurs had evolved into several major subgroups, namely

Cetiosauridae, Brachiosauridae (including Brachiosaurus), Camarasauridae (including

Camarasaurus), Diplodocidae (including Diplodocus and Apatosaurus) and Titanosauridae. I

will discuss seven various types of famous Sauropods.

Firstly, there is the long-necked Alamosaurus. This group of sauropods are believed to weigh

around 33 tons and were up to 69 feet long. They were discovered in a region near Kirtland

Shale, New Mexico. They lived approximately 70 to 65 million years ago during the late

Cretaceous. They went extinct eventually during the Cretaceous mass extinction event during

the Mesozoic era.

Another well-known

Sauropod is the

Argentinosaurus. They were

discovered in Argentina, with

which they adopted their

name from. They are known to be arguably the largest and heaviest terrestrial animal to ever

walk this Earth. They were estimated to weigh between 66-97 tons and were around 98-118

feet in length. However, when they were initially hatched from their eggs, juvenile

Argentinosaurus were actually only the size of a football. This species of sauropods lived on

until the late Cretaceous period, which is around 97 to 93 million years ago.

Next is the Apatosaurus, which lived 150 millions years ago during the Late Jurassic period.

They were known as Brontosaurus as well in various parts of the world. They were first

discovered in Oklahoma, Wyoming, Utah, New Mexico and Colorado in the US. They

weighed between 16 to 22 tons and were around 69 to 75 feet in length. This species had

similar skull structure to the Diplodocus, hence, scientists often confused the Apatosaurus

with the Diplococus. Their diets consisted of plants alone, hence their heads and brains were

consequently very small (herbivores had smaller brains than carnivores usually).
The Brachiosaurus is one of the most

famous dinosaurs to mankind. The

renown sauropod lived through the

Jurassic period around 154 million years

ago. They too were found in the US. Their

bodily dimensions were similar to the

Apatosaurus, with their length estimated

to be about 66 to 69 feet and weighed

between 35 to 58 metric tons. Brachiosaurus’ necks were believed to made of 13 cervical

vertebrae, which allowed it to reach plants as high as 9 meters off the ground.

Another species were the Camarasaurus, which translates to “chambered lizard”. The name

implies that Camarasaurus’ spine had large holes in them, which was a unique trait to this

type of dinosaur. They lived between 155 to 145 million years ago during the Late Jurassic

era. Again, they were discovered in the US at the Morrison Formation. They were estimated

to have a length of around 75 feet and weighed around 51 tons.

Diplodocus is another very well-known dinosaur to us. Their name roughly means “double

beam” from neo-Latin, referencing to their double-beamed tail bone. They were the first

dinosaur fossil to ever be excavated by museum paleontologists in 1897. Diplodocus lived in

the late Jurassic era around 104 million years ago. They were again found in the US. They

measured an estimated length of 79 feet in length and a weight of 12 metric tons. They had an

especially long tail entirely made up of a total of 80 caudal vertebrae and had a special whip-

like function to defend itself. The spikes that lined up on its back had the similar purpose of

self-defense too.
Finally, a type of Sauropods called Saltasaurus stands out from the rest, as they were

relatively small and had short, stubby limbs, together with a rather short neck. They were

discovered in Argentina and this special sauropod lived during the late Cretaceous era. Even

though they were still larger than modern day animals, there were considerably tinier than its

sauropod counterparts. They were only 42 feet in length and weighed seven tons.

Description of Sauropods’ anatomy and lifestyle

Most Sauropods have a shared body plan and their body parts were all designed in a way

such that drove the sauropods body size up beyond any land animals that have ever stepped

foot on Earth. This will be discussed more in depth later in this essay. This part will merely

describe the appearance of the dinosaurs and their livelihood and these will be compared to

modern day animals.

Firstly, Sauropod dinosaurs had a small head balanced on an extremely long neck that is often

found at over 15 meters long, which is almost six times longer than that of the modern-day

giraffe. In their necks, there were 19 cervical vertebrae that all contain air sacs, whereas

almost all mammals nowadays are limited to only seven cervical vertebrae. However, the
abundance of cervical vertebrae made its backbone extremely inflexible. Sauropods’ giant

bodies and quadrupedal stance helped to provide a stable base for the neck and head.

In turn for the long neck, the head and consequently the brain had evolved to be very small

and light. By minimizing the size of their heads and by reducing its function to solely

harvesting tools to get plants into the body, the Sauropods could spend less energy to lift their

head and so that the power could be saved for their metabolism and movements of their

enormous bodies.

Though, by having a small head, it caused Sauropods to apparently lose their ability to orally

process and masticate food. On the other hand, a small head could also have reduced the

development of dense neck muscles, which in turn reduces the mass of their neck and this

allowed the subsequent continuous elongation of the neck.

Their nostrils were located high up on the skull rather than at the end of the snout like other

terrestrial vertebrates. Sauropods also did not have ears that stuck out like ours, but their head

housed the inner ear.

Sauropods did not seem to have chewed their food due to the small size of their heads.

Resembling the Galapagos giant tortoise, Sauropods tend to gather and swallow plant fodder

whole without much oral processing. They were also suspected by Steve Salisbury, a

paleontologist working at the University of Queensland, to have turtle-like beaks to protect

and hold their teeth together. Another discovery was that Sauropods replace their teeth quite

often. Each sauropod species also had different types of teeth that largely depended on their

diet. For example, Camarasaurus had spoon-shaped teeth, while Diplodocus had pencil-

shaped teeth. The two types of dinosaurs had different diets. The Diplodocus ate plants that
grew low to the ground or soft aquatic vegetation, while the Camarasaurus ate leaves that

were from higher branches, which is supposedly tougher. There were also presumably more

wind-blown particles consumed by the Camarasaurus as they browse high. This made the

Camarasaurus’ teeth wear at a higher rate than the

Diplodocus’ teeth.

Camarasaurus Diplodocus

The neck is said to be held horizontally to feed on plants over a wide area instead of standing

vertically straight up. This is because if the necks were vertical and were used to browse high

trees, the energy needed to create the arterial blood pressure for pumping blood towards the

head would consume half of the sauropod’s energy intake. The heart would also need to be

15 times larger than a whale to be able to pump blood all the way up to the head structures.

However, sauropods may raise their necks when being alert.

The significance of having such a long neck allowed Sauropods to sweep out large feeding

areas moving their body, which gave Sauropods an advantage in terms of searching for food.

This definitely reduced the energy cost needed for moving their huge body while looking for
food. However, Sauropods could not retract their necks above their shoulders to explore

higher feeding grounds.

The neck can also act as a radiator that can deal with large amounts of heat produced by their

large body mass, which presumably performs immense amounts of metabolism. This is

because the long necks were predicted to have cooled down the blood vessels on the way to

the brain, which avoids heated blood entering the head of Sauropods.

Furthermore, Sauropods were quadrupedal, and they had thick pillar-like legs, just like the

modern-day African elephant, to support their enormous trunk and torso. Their hind legs

were thick and ended in a club-like feet with five toes, but only the inner three bore claws.

The forelimbs, on the other hand, were similar to modern day sauropods. There were

slenderer and ended in pillar-like hands that were built for supporting an enormous amount of

weight. In the case of the forelimbs, the Manus bones were arranged in fully vertical columns

with reduced finger bones. The fingers might not even be visible in later sauropods such as

the Eusauropods due to extreme modification of the front feet. The metacarpals bones were

semicircular and horseshoe-shaped. Unlike elephants however, the front feet of Sauropods

did not consist of any fleshy padding on the back, making them concave in appearance.

Finally, the bottom and sides of the forefoot were often covered in spiny scales according to

evidence from Portugal.

Another notable feature that the Sauropod’s body possess were a system of air sacs that were

shown by indentations in most of their spine. These air sacs greatly reduced the overall

weight of their long necks. They also allow a single-direction airflow through their stiff lungs

and allow Sauropods to get enough oxygen for their body. This respiratory system is similar

to that of birds, which scientist hypothesized to have contributed to Sauropod’s gigantism.

Sauropods had very long tapering and often whip-like tails. There were mainly three

functions. Firstly, they acted as a communicating tool between the species. Secondly, the

long tail was for balancing with the long neck. Lastly, it can be used for attacking enemies

and also defending themselves from predators.

In terms of their lifestyle, I would be discussing their living habitat, feeding habits and

herding rituals.

Sauropods were widespread geographically and were found in nearly all continents except

Antarctica. They were once thought to spend most of their time in shallow waters as this

would help support their ponderous bodies. However, after considerable evidence, including

a study in the 1950s conducted by a paleontologist called Kermack, it was shown that if

sauropods were to be submerged in merely a several meters of water, the water pressure can

fatally collapse the sauropod’s lungs and trachea. Also, in 2004, paleontologist named

Henderson said that due to the existence of air sacs in their spines, sauropods must be

buoyant and hence cannot be submerged under water. This indicates that they were better

equipped for living on solid ground.

Regarding their feeding habits, Sauropods were dominantly herbivorous, and their longs

necks enable them to foliage from even the tallest trees in similar manner as modern giraffes.

Sauropods may require up to 100,000 calories a day as they weigh more than 10 African

elephants combined. Therefore, they needed to eat as much food as they could as fast as

possible. The way in which sauropods were able to do that will be discussed later as this
contributed hugely to how they grew so enormous. Additionally, several types of sauropods

were able to coexist because they fed on various types of plants.

Apparently, Sauropods swallowed stones to help break down plant matter they ate, which

were usually swallowed whole without chewing. Sauropods lack enzymes that were required

to breakdown cellulose in plants. The rocks swallowed and also the microorganisms such as

bacteria residing in their digestive tracts can.

Finally, sauropods were oviparous, which meant that they laid eggs. Each herding season,

they would lay a similar range of number of eggs as large sea turtles today, which does not

exceed 200 to 400 eggs at maximum at one time. Their herds were separated by age because

young sauropods had different diets from adult counterparts. Size difference also played a

role. The segregation of the herd happened as soon as after hatching. Hence, scientists

predicted that there was minimal parental care towards the juvenile. There have also been

opposite arguments about this matter. It is still unknown today why there were segregated or

what was the level of parental care like.

How have sauropods grown so enormous?

The most notable feature of Sauropods was definitely their size. They were the largest

terrestrial animal to ever exist on this Earth, with the only competitor being the modern-day

blue whale, but unlike them, sauropods lived on land.

Moreover, the largest dinosaurs all belong to this group. The longest one is called

Supersaurus at 34 meters long, which was way longer than the longest terrestrial animal alive
today – the reticulated python, which is only at seven meters long. The tallest sauropod is

called Sauroposeidon at 18 meters high, as compared to the giraffe, which is only at a mere

five meters tall. The heaviest of them all was the Argentinosaurus, which weighed a

staggering 120 metric tons. The largest land animal alive today, the Savanah elephant, only

weight about 10.4 metric tons. Even the so-called dwarf sauropods i.e five meters long, were

considered among the largest animals in their ecosystem.

Sauropods actually started out to be small animals of only 10kg in body mass from the early

Late Triassic era, but as they evolved, they grew to a body mass of an astounding 100,000 kg.

The large size obviously had its significance. This feature protected sauropods against

predators, helped them cover a greater range of acceptable foods, increased their resistance to

climatic extremes and even allowed them to live longer.

However, having such a large body definitely has its disadvantages. Firstly, it would increase

the sauropod’s exposure to predation. The demand for resources undoubtedly rises as well.

Other issues that a large body size may poses include circulatory problems. As mentioned

before, the size of the heart must be big enough to pump blood all the way up to the brain.

For example, the Brachiosaurus’ head is located 8 meters above the heart. Hence, it is

estimated that the heart would need to be at least 200kg to achieve that purpose. A very

complex respiratory system is also needed to supply enough oxygen to sustain the large body.

Finally, there could be a problem of overheating.

Despite all of these obstacles, it still did not obstruct the evolution of sauropods into the most

gigantic terrestrial animals. Sauropods’ bodies could double its body mass in less than a week
since birth and attain a staggering 80 metric tons in just 23 years. Human infants, on the other

hand, take 2.5 years at least to double its weight.

Studies have been done to verify if the Cope’s rule was in operation in Sauropods road to

gigantism. The Cope’s rule is named after an American Paleontologist named Edward

Drinker Cope, and the rule postulates that the population lineage of a species increase in body

size over evolutionary time. In other dinosaurs, they showed a gradual body size increase

over time, but Sauropods alone showed a rapid size increase in the Triassic Period, which

shuts down the possibility of the Cope’s rule playing a part in Sauropod’s evolution.

It is later discovered that sauropod’s growth dynamics is large due to a mechanism called the

Evolutionary Cascade. Basically, the sauropods’ ancestors had a unique mix of physiological

and anatomical traits that are fueled by positive feedback and this led sauropods to evolve to

its size. The traits include: having a high basal metabolic rate, a bird-style respiratory system

that include unidirectional airflow throughout the lungs, the production of many small

offspring and having limited oral processing of food.

This mix of traits led to five different evolutionary cascades:

1) Head and neck anatomy

As mentioned above, sauropod’s have a tiny head balanced on an extremely long

head. Firstly, the lack of oral processing of food meant that the head did not have to

carry a massive set of chewing muscles to deal with the increase amount of plant

fodder. As compared to modern animals, chewing muscles and their heads have to

grow large as their body size increases so as to support the amount of food needed to

be consumed. Without the need of developing large mastication muscles, this allow
 sauropods to develop a relatively small head that require less energy to carry.

Consequently, this allowed their necks to elongate. As the neck gets longer, browsing

for food becomes easier as they can reach the food that is span across a rather large

area with their long necks without having to move their body much, meaning that they

do not have to use that much energy. This ultimately led to an evolution of faster food

intake, which allowed the body to grow quickly.

Their extremely long neck allowed more efficient food uptake compared to other

herbivores as they can cover a much bigger feeding envelop that are out of reach for

other animals. This greatly increase the food source available to sauropods and it

allowed them to intake more energy than other animals, hence, this led to their body

growing bigger.

2) Reproduction

There was a retention of plesiomorphic oviparous mode of reproduction. This allowed

faster population recovery and a greater reproductive output of large dinosaurs. This

lowered the risk of extinction of the species. Sauropods also produced numerous but

small offspring each season much like modern day sea turtles. This saved sauropods a

great deal of energy for sauropods as they do not need to focus on just one or two

offspring that require a lot of care. This method of production was said to have

removed boundaries that kept other animals small by comparison.

On the other hand, mammals such as elephants and giraffes, which are considered

quite large already in modern days, they reproduce in a vastly different way. These

big mammals usually have to carry one offspring inside of them for long periods of
 time and the need for parental protection and milk means that the mother had to waste

more energy even after the birth of their child. Long pregnancies not only drain the

animal’s energy, the birthing process itself could also lead to potentially fatal

complications. These are the reproductive threshold that mammals could not cross,

and for mammals to grow larger, the mother would have to carry their child for a

longer period of time and there needs to be more parental care as well. The mammoth,

one of the largest terrestrial mammals, demonstrated that without changing the

method of reproduction, mammals could only get that much bigger, and with the

deadly consequence of extinction as well.

By adopting external birth and development via eggs, sauropods could avoid all these

constrains that mammals had that restricted their size – it saves sauropods a lot of

energy.

3) Mode of feeding

The time between the acquisition of food and swallowing in sauropods is very short.

This helped sauropods gain a high rate of food intake to sustain the large body size,

which is unique to this creature. There are many evolutionary specializations that

support their increased food intake rates. This includes fast tooth replacements,

widening gaps through broader jaws and the loss of cheeks. These reasons all helped

in producing a selective advantage of obtaining more energy from the environment for

bodily growth for sauropods, given that there was a large gut capacity in the dinosaurs

to deal with that high input of poorly chewed food. Furthermore, sauropods were

highly selective eaters. They always chose more nutritious plant parts such as the

seeds and young shoots, which further assisted their growth.

4) Bird-like lung

Sauropods are said to have a respiratory system that was similar to the modern-day

birds. There are two features of the bird-like lung. Firstly, it contains a system full of

air-sacs or cavities which provide fresh air to the lung tissues continuously and

secondly, there is a cross-current gas exchange mechanism going on in the lung

tissue. In sauropods, this system comes in the form of pneumatized bones. The axial

skeleton including the ribs are all filled with air spaces and the pneumatization would

extend backwards along their backbone as sauropods continued to evolve. This

system allows a dramatic increase in oxygen in take (around twice as much oxygen

per unit air volume as other animals). Furthermore, this lowers the cost of breathing

and reduces the gravitational impact on the sauropod’s respiratory system. When

tissues are supplied with more oxygen, there would be faster cell metabolism. In other

words, the organism would work more efficiently with the same amount of resources.

Hence, the growth rate of sauropods became really high.

5) Metabolism

In order to have such a high growth rate, sauropods need to have a remarkably high

basal metabolic rate (BMR) to run their huge bodies. This would supposedly cause

overheating if the BMR is that high. However, according to calculations, their body

temperature does not increase with the rise in size. Sauropods are said to have an

average body temp of 28 degrees Celsius, which is well below today’s endothermic
 vertebrates. Scientists constantly question where all the energy needed for growth

came from if having an insanely high BMR does not do the job.

Actually, sauropods have evolved so that they were able to conserve energy very well

and made use of resources available better than other animals. For instance, they save

their energy by firstly decreasing their body density. By having air sacs in their

vertebrae and ribs, this has greatly lightened their bodies, saving them the energy

from moving around their massive bodies. Their cost of respiration was also reduced

due to the bird-like respiratory system. This reduces the waste of energy that is

otherwise put on breathing muscles that are present in other animals. Increased

oxygen intake through each breath also means that the frequency of breathing can be

lowered, further minimizing the energy consumed from breathing. Finally, the long

neck that facilitates the easy browsing of food obviously saves energy that is

otherwise wasted on locomotion to look for food.

Other than the above intrinsic biological factors, there were also extrinsic factors that might

have contributed to sauropods’ gigantism.

1) Increased atmospheric oxygen content

Scientists have speculated that the oxygen content of the atmosphere in the Jurassic

period when the Sauropods developed was 30% above the oxygen level on Earth

today. This is indeed possible because for sauropods such as the Apatosaurus in the

Late Jurassic period, their respiratory system definitely could not have delivered

enough oxygen to the tissues with today’s level of oxygen. However, this prediction

was superseded by the likelihood of the presence of a bird-like lung in sauropods.

2) Increased carbon dioxide levels in the atmosphere

In the Mesozoic period, the carbon dioxide level was higher than it is today. This

would hypothetically increase plant productivity and hence increase the amount of

food available for animals, including sauropods. However, research shows that while

plant productivity may have increased, there would be a decrease in protein content

and increase in nonstructural carbohydrate content in the plants. This means that food

quality for sauropods. Therefore, it has been concluded that increased carbon dioxide

level does not contribute much to gigantism.

3) More nutritious food

Sauropods tend to ingest pre-angiosperm flora such as Ginko, cycads, ferns and

conifers, which is more nutritious than plants consumed by other herbivores. These

plants offer higher energy yields and hence enhance their growth.

4) Preference in habitat

Sauropods like the giant Paralititian chose to inhabit certain areas such as mangrove

environments. Mangroves are second most productive environments after tropical

rainforest, so it can provide a large resource base for evolution of huge body sizes.

While this hypothesis may be applicable to Paralititan, a type of Titanosaurian

sauropod, it does not fully explain sauropod gigantism in general.

Most of these hypotheses about the impact of extrinsic factors have been superseded and

hence it is safe to say that the evolutionary cascade hypothesis seems the most plausible in

explaining how sauropods grew to such an extraordinary size.

Conclusion

In conclusion, Sauropods were the largest terrestrial animals to have ever existed. They were

herbivores with well-developed quadrupeds, a long neck, a small head and a whip-like tail.

After much research, it has been discovered that there could be both extrinsic and intrinsic

factors that contributed to their gigantism, with the latter being the dominant reason.

The evolutionary cascade is the best explanation to date for sauropod gigantism. The

evolutionary cascades is the unique mix of traits of sauropods ancestors in the physiological

and anatomical aspect. The mix of traits fuelled positive feedback and it resulted in the

evolutionary selection of a gigantic body size in sauropods. There are five paths of

evolutionary cascades driven by the traits of sauropods ancestors. The physiological and

anatomical features of sauropod ancestors allow the sauropods to have advantages in

obtaining a lot of food and oxygen. Furthermore, their features also allow the conservation of

energy in sauropods that can used for growth. These features include an avian style

respiratory system, a high BMR, the lack of mastication, a long neck and having a huge

number of small offspring every season that were produced in a oviparous manner. Out of all,

the long neck of sauropods one of the major evolutionary advantage that drove the sauropods’

high growth rate, as the neck facilitates the sauropods in obtaining a huge amount of food

efficiently to sustain their bodies. All these reasons combined allowed evolutionary cascade
in sauropods in the Late Triassic and Early Jurassic eras, causing the selection for a huge

body size.

Scientists and palaeontologists are still constantly trying to find out how their body size

evolved to such extent. It is especially difficult as the dinosaur remains are still very scarce

and limited and most of the estimation in their anatomy is based on computer generated

images. Finding out about their physiology and the amount of metabolism sauropods need

poses even a bigger challenge for researches.

In addition, there needs to be future investigation in whether the sauropod body size could be

influenced by the extrinsic factors such as atmosphere oxygen and carbon dioxide levels.

There also needs to be more research on whether there are other extrinsic factors that are

currently unknown that could have also played a part.

Nevertheless, sauropods were one of the most successful large terrestrial animals on the earth

and they dominated the globe from the middle Jurassic period onward, until they finally

became extinct at the end of the Cretaceous period around 65 million years ago due to major

environmental changes.

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