Functional Morphology Final Paper:

Squirrel Forepaw Functionality Analysed via Ulna Morphology

So Chang* a, Diego Molina Ochoa** b

a
Department of Biology, University of Washington, Seattle, United States of America
b
Department of Biology, University of Washington, Seattle, United States of America
*
Corresponding author: sojungie@uw.edu
**
Corresponding author: diego.molina@comcast.net

Summary

1. While morphological research on hands in primate species is extensive, there is

limited understanding of the structural significance of the forelimbs of other
species.
2. Here we are analysing the morphological differences of various squirrel species
forelimbs depending on contrasting environments, with arboreal and ground
squirrel species being our main comparison groups. Ulnar morphology is being
used as a proxy for hand dexterity and strength.
3. Ulnar length and circumference, along with 2D geometric morphometric data,
were gathered from 95 squirrel specimens, and analysis was accomplished using
principal component analysis to take into consideration phylogeny.
4. Results showed that while linear measurements were significantly different
between arboreal and ground squirrels, differences between the two groups could
be associated primarily to phylogeny. (Insert stuff about morphometrics here)
5. While the results do point to significant differences in forelimb morphology
between squirrel species in different environments, more research is required to
find a casual link between environment and forelimb morphology, as well as how
forelimb morphology influences forepaw function.
Key words: Dexterity, Hands, Sciuridae,
 Introduction:

When considering the morphological traits that define the human species, there are three

that immediately stand out and set us apart from the majority of mammalian species: bipedalism, a

brain that is proportionally large when compared to body size, and exceptionally dexterous hands.

Of those three, the use of hands carries particular significance in the biological and socio-cultural

evolution of humanity; skillful hands fashioning the tools and carving the writing that would propel

the species into the realm of civilization. It is of little surprise that the evolution of hands has been

of particular interest among evolutionary biologists. There has been a great deal of research on

many aspects of the nature of hands amongst the order of primates, where they enjoy the use of

manipulatory forelimbs (Taylor).

Within the body of research, there is a problem where few species outside the primate

order have received attention when it comes to the morphology of hands. This is likely due to the

fact that primates hold a significant monopoly over the presence of opposable thumbs, a trait that

tends to be defining when it comes to describing hands. This bias can be seen even in naming

themes: the term “hand” tends to be limited to human-like appendages shared among primates,

while other species are defined as having “paws”. Paws bring to mind a very distinct morphology,

one that lacks the dexterity of human-like hands, despite the fact that many species show forepaw

structure that shares many characteristics to traditional primate hands.

This is not to say that research on non-primate forelimbs is completely lacking, indeed, the

importance of forelimbs in the mobility of other mammal species has led to a great deal of

investigation. Previous research have explored the muscular and skeletal differences between

arboreal squirrels, flying squirrels, prairie dogs, and rats (Peterka). The study divided up the

species into categories, looking at their habitat and their habitual movements. They found that

arboreal and flying squirrels had an extensive development of the muscles that flexes the forearms
and extends the digits. Through these results, we can see that in species that live in habitats that

involves climbing, there are adaptations in the muscles to optimize the action.

In this experiment, we hope to explore the question of whether there are skeletal

differences between ground, arboreal, and flying squirrels based on their habitual movements of

climbing, climbing and gliding, and running. The family of Sciuridae is great in the number of

species and employs a great number of habitats across the world, with 278 species inhabiting many

continents with the exception of Antarctica and Australia (Thorington). Since there is such a great

number of species, our research focuses primarily on the subfamilies of Sciurinae (arboreal and

flying) and Xerinae (ground).

Musculature patterns are associated with skeletal patterns based on the amount of muscle surface

area at the origins and the insertions of the muscle. The greater surface area the muscle attaches

with, the greater the force it can contract with. The muscle we are interested in is the flexor

digitorum profundus (figure 2). This particular muscle originates in 4 heads; the condylo-radial

head, central head, radial head, and the ulnar head (Washmuth). The condylo-radial head originates

at the “posterior portion of the medial epicondyle” on the humerus. The central head originates

from “the anterior portion of the medial epicondyle.” The radial head attaches to the medial portion

of the radius bone. The ulnar head attaches to the medial portion of the ulna. All these heads fuse

together at one tendon, which separates into 4 different tendons, each attached to the ends of the 4

phalanges apart from the thumb.

From our knowledge of anatomy, when a muscle contracts, it brings the parts of insertion

closer to the point of origin on the body. Therefore, the origin of the muscle should have a greater

surface area for a greater force. Following this thought, our experiment seeks to look at the

differences in the skeletal structure of the ulnar head origin to see if there are significant changes

between adaptations for different habitats in the squirrels.

 While primate hand and arm morphology is a great starting point to explore the

relationships between function and structure, expanding our perspective into the vast order of

Rodentia will help hone in on some important features that may be independent from other habits

and habitats that other mammalian research have yet to explore. By exploring how grip strength,

initiated by the flexor digitorum profundus, might affect structural patterns on the ulnar bone, we

will be able to uncover an insight to how different habitats affect muscle structure and thus skeletal

patterns. If we do indeed find significant evidence to answer our question, we will be able to make

room for future research to explore how different niches lead to significant variations within

different species.

The Sciuridae family offers significant potential with regards to researching trends in hand

morphology for two reasons: firstly, the Sciuridae family demonstrate very distinct manipulatory

appendage structures, that is, their forelimbs are not merely slightly different hindlegs, and have

distinct uses among squirrel species beyond locomotion, such as grasping and manipulating objects.

Second, squirrels are present in a wide range of environments, and have adapted to various

different ecological niches. Some squirrels are primarily tree climbing, others are exclusively

terrestrial, and a few have even evolved the ability to glide. This diversity in strategies will allow us

to see how hand morphology differs within a single phylogenetic family depending on environment

and behavior.

The objective of this research study is to compare the morphological trends of different skeletal

patterns among tree climbing, ground, and flying squirrels.

To analyze how hand morphology differs between squirrel species, we need a measurement

that can effectively describe the nature of a squirrel’s forepaw. Dexterity is a complex factor

involving skeletal, muscular, and neural factors, but even partial analysis can reveal trends about
hand control and morphology. Analyzing the size and shape of phalanges is a promising option, but

given the size and uniformity of such bones, such research was not be feasible in the timeframe

available for this project. Another option beyond analyzing bone structure of the phalanges is to

analyze the muscles responsible for finger movement and control: analyzing such muscles serves as

an effective measure of hand dexterity or gripping strength. Of the muscles involved in gripping

strength, the flexor digitorum profundus is particularly important, as it controls the flexion of the

four primary digits in humans and many other mammals, including squirrels. Such flexing action is

particularly important for arboreal and flying squirrels by offering a strong enough grip onto a

surface for climbing. With this in mind, we offer our hypothesis for this research study.

Hypothesis: Exchange of fine hand-motor control for grip strength may significantly influence hand

and arm musculoskeletal morphology differences between arboreal, flying, and terrestrial squirrels.

Null Hypothesis: Exchange of fine hand-motor control for grip strength has no significant effect on

hand and arm musculoskeletal morphology between arboreal, flying, and terrestrial squirrels

Prediction: Ulna of flying and tree-climbing squirrel species will have greater surface area and

circumference than those of ground squirrel.

Materials and Methods

For the sake of this research project, we have divided the specimens of the Sciuridae family

available from the Burke Museum Mammalogy collection based on whether they fit into one of

three categories: whether they are primarily tree-climbing, terrestrial, or gliding species.

We collected data on several factors for each species analyzed, focusing primarily on the

nature of the ulna. We made linear measurements of maximum length and circumference,

measured in millimeters with the use of calipers and labeled twines. This is to describe the size of

the ulna between different species. We also took photographs to make measurements through 2D

geometric morphometric analysis, in which we established shared landmarks along the anterior

surface of the ulna and measured them for all specimens. This describes the shape of the ulna. By

gathering both the size and shape data, we were able to compare the ulnas of different squirrel

species on a number of levels. We also recorded the overall mass of the specimen, measured in

grams, a value that was pre-recorded in each stored specimen in the Burke museum, along with

their sex. This allowed us to control for overall body size while comparing ulnas of different species
 Ulnar
Species Name N Habitat Mass(g) Ulnar Length(mm) circumference(mm)
Glaucomys sabrinus 5 Gliding 94.51-114.45 40.73-42.55 3.0-4.0
Glaucomys volans 2 Gliding 51.2-62 30.03-30.95 2.5-3.0
Ammospermophilus lecurus 5 Ground 82-120 22.58-25.6 4.0-5.0
Marmota caligata 5 Ground 3000-5600 66.8-87.74 16.0-20.0
Marmota flaviventris 5 Ground 829-2966 58.55-66.09 13.0-18.0
Marmota monax 4 Ground 1955-3885 67.32-76.47 16-18
Spermophilus beecheyi 5 Ground 130.4-622.4 26.97-45.07 6.5-10
Spermophilus columbianus 5 Ground 131.5-660 22.79-38.94 6.0-9.0
Spermophilus lateralis 5 Ground 130-174 26.29-28.73 6.0-8.0
Spermophilus saturatus 5 Ground 205-280 30.35-34.49 6.0-8.0
Tamias amoenus 5 Ground 44-51.2 19.94-21.63 4.0-5.0
Tamias minimus 5 Ground 26.85-31.2 16.54-18.75 3.5-4
Tamias ruficaudus 5 Ground 50-60 21.6-23-6 4.5-5
Tamias striatus 3 Ground 47.2-97.2 17.14-23.72 4.0-5.0
Sciurus aberti 5 Arboreal 560.4-765.5 52.52-53.8 9.5-10.5
Sciurus carolinesis 5 Arboreal 405.6-769.8 38.82-50.65 8.0-11.0
Sciurus griseus 5 Arboreal 543-833.9 56.34-60.01 10.0-12.0
Sciurus niger 5 Arboreal 503-803.5 43.19-56.9 10.0-11.5
Sciurus vulgaris 1 Arboreal 46.2 9
Tamiasciurus douglasii 5 Arboreal 118.5-222 32.67-35.51 6.0-7.5
Tamiasiurus hudsonicus 4 Arboreal 219.5-298 32.69-38.38 7.0-8.0

Data Analysis:

In order to measure and compare landmarks of the ulna across the different species, R-

studio was utilized through the package of geomorph. This allowed us to measure the differences

between the different structural morphologies of the ulna. For analysis of length and circumference,

we used the phylANOVA function in R to do a phylogenetic ANOVA test to compare the linear

measurements of species in different habitats while taking phylogeny into consideration. For

comparing geometric morphometrics, we will use (Insert geomorph info here)

(Insert image of landmarks here)

Results:

We measured the Ulnar lengths and circumference of 94 ulnar specimens representing 21

different squirrel species. We also took images of all specimens for the purpose of geometric

morphophology analysis, applying 29 landmarks utilising the tpsdig program.

Circumference

Discussion:

Our results show that there does exist significant difference in ulnar morphology between ground

and tree squirrels, however, this difference can be described completely through phylogenetic

momentum. Therefore, we cannot reject our null hypothesis, and cannot conclude that ulnar

morphology is dependent on habitat type.

 Our failure to reject the null hypothesis may be due to the phylogenetic constraints present

in our specimens: habitat type is conserved to a great degree phylogenetically, with ground

squirrels and arboreal squirrels forming very distinct blocs on the sciurid family tree. This alone

makes it difficult to determine if habitat type alone imposes sufficient pressure to change forelimb

morphology among squirrel species. To accomplish this, we would have needed an example of a

squirrel species that differed in habitat type to its nearest phylogenetic neighbour. For example, we

would have needed a ground squirrel who had an ancestor that was a tree squirrel, or vice-versa.

Had we had such a species in our study, and then observed that ulnar morphology remained

conserved between squirrels of similar habitats regardless of ancestry, then we could have

determined that some form of convergent evolution driven by habitat type had taken place.

Unfortunately, none of the species in this study matched this definition, but finding such an example

of convergent evolution could be a focus for future research on this subject.

A few patterns emerge when observing the data available: Ulnar circumference and length

tended towards low extremes among tree and flying squirrels, however, the opposite is not true for

ground squirrels, that is ground squirrels tended to vary significantly more at the mid point of ulnar

morphology. This may represent some contrasts between the selective pressures experienced by

ground and arboreal squirrels: lack of significant selective forces generates a wider spread of values

for a trait, as seen with ground squirrels, while directional selection pushes values towards an

narrower extreme, as seen with arboreal squirrels. What is curious is that the results seem to

contradict our initial prediction that arboreal squirrels will have larger and longer ulnas, and thus

have larger flexor digitorum for the purpose of climbing and clinging to trees. After all, muscle size

is positively correlated to strength, so it would make sense that, to have the strength to climb for

long periods of time, arboreal squirrels will have larger ulnas and musculature.

There may be several explanations for this discrepancy: perhaps smaller muscles are better

adapted for long-term, efficient muscles exertion. Thus, rather than selecting for raw strength,
arboreal squirrels with musculature that is suited for endurance have an adaptive advantage. There

is also the possibility that we overestimated the important of the flexor digitorum, the ulna, and the

forelimb in general with regards to climbing: climbing is a highly complex and dynamic form of

movement, utilizing muscles from the legs, the back, and arms simultaneously. Squirrels may

depend less on their ability to grasp for climbing and more on the power of back and leg muscles.

Thus, to maximize climbing ability, evolution has driven a reduction in forelimb size in favour of

more critical muscle regions.