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Daniel Perret, Department of Biology, Stanford University


The mountain beaver, Aplodontiarufa, is an understudied endemic rodent that
displays several primitive characteristics, including the inability to produce
hypertonic urine and relatively simple teeth. Aplodontiarufa is the only extant
species in its genus and family, Aplodontiidae. The mountain beaver, part of an
ancient North American rodent lineage, is now confined to an endemic range within
the Pacific Northwest and northern California. At least two of the seven extant
subspecies are considered threatened or endangered.

My objective is to broadly understand the interplay between the environment and

the ecology and evolution ofA. rufa. I propose to accomplish my objective using two
approaches. First, I will determine the environmental constraintsonthe spatial
distribution of this species by compiling climatic and spatial data in a Geographic
Information System (GIS) analysis. Second, I will measure and quantify the
phenotypic differences between and within subspecies of A. rufa through
morphometric analyses. The integration of these data sets will allow me to
understand how the environment sculpts the persistence of the lineage through
time, and how this has influenced within-species divergence. My data will also shed
light on the climatic variables that likely played a role in the extirpation of
Aplodontiarufa from northern California during the Pleistocene-Holocene transition.

The results from my research will form the foundation of my Honors Thesis in
Ecology and Evolution at Stanford University. I hope also to open the door to deeper
investigation into the greater effects and implications of A. rufaecology. My results
will thus serve as a foundation for subsequent studies of the molecular
phylogenetics of the species and its survival into the future.


Aplodontiarufa is the only remaining species in its genusand once-diverse family,
Aplodontiidae. The species exists in a series of disjunct populations, comprising the
seven different subspecies distributed throughout the mesicPacific Northwest and
into parts of the mostly montane northern California coast (Fig. 1). These subspecies
have been defined on a mostly geographical basis (Hall 1981). My objectives are (1)
to characterize the environmental variables that define the distribution ofA. rufa,
and (2) to determine how (or if) these variables correlate with the morphology of
the seven subspecies.

The mountain beaver is one of the most primitive extant
rodent species, possessing an apomorphic suite of skull
features and jaw musculature not shared by other
rodents (Carraway&Verts 1993). Additionally, the species
retains the unusual feature of being unable to produce
hypertonic urine, a result of simplified renal anatomy.
Thus, A. rufa must drink large amounts of water dailyȄ
approximately one third of its body massȄalmost all of
which is accounted for in urinary water loss (Pfeiffer et al.
1965). Because of this, I hypothesize that the spatial
distribution of the species is controlled by a limited
number of environmental factors pertaining to water
availabilityȄfor example, annual seasonal rainfall or soil
moisture content. However, my preliminary data suggest
that the distribution of A. rufa is not controlled by a single
climatic variable, but by many such variables. Using GIS, I
will conduct a geostatistical analysis, correlatingmuseum
specimen locality data with environmental data,in order
to perform variable-specific niche modeling to determine
which environmental factorsbest explain the present
distribution of A. rufa.
Figure 1. Current distribution of
I hypothesize that substantial morphological variation
Aplodontiarufa, with subspecies
boundries delineated. From exists between A. rufa subspecies, especially along a
north-to-south axis. Many mammals demonstrate body
size clines with latitude and elevation, consistent with
Bergmannǯs Rule. Preliminary work by Samantha Hopkins (University of Oregon)
suggests species-level morphological variation between
Aplodontiasubspecies,especially along a north-to-south axis (S. Hopkins, personal
communication). I predict that this variation is associated with an environmental
gradient or cline, which my data is intended to reveal. 

c  !  

Aplodontiarufa is a particularly interesting study organism for several reasons. It is
the lone survivor of the most ancient extant rodent lineage and possesses a number
of unusual traits not shared by other mammals. In addition to its need for copious
daily fresh water, other such traits include a simple tooth structure (Aplo = simple, -
dontia = tooth), primitive reproductive anatomy, and a simplified zygomasseteric
system believed to be ancestral to other rodent variants. Aplodontiarufa also
maintains an unusual life history characterized by slow development and small
litter sizes of two or three, resulting in smalllocal population sizes and slow intrinsic
rates of growth. Mountain beavers prefer an unusual diet of succulent plants that
are toxic to other species, including alder leaves and young conifer trees. The
girdling of conifer trees by Aplodontiarufacan cause extensive damage to
commercial tree nurseries in the Pacific Northwest, resulting in economic losses and
the loss of up to 25 % of new seedlings.Thus, in spite of their conservation status,A.
rufa populations are often controlled extensively by foresters. (Carraway&Verts
1993, Cambell& Evans 1988, Motobu 1978)

These aspects of A. rufa together make the species one of considerable conservation
concern. Currently, at least two of seven subspecies are listed as threatened or
endangered. The species is believed to be very sensitive to climatic aridification,
making it an indicator species for climate change. Despite this, no research thus far
has specifically targeted environmental variables controllingA. rufa distribution, and
little is known about the role of the species in the old-growth temperate forests in
which it is most often found.

Aplodontiahas persisted
virtually unchanged
morphologicallyin the
fossil record for
approximately 6 million
years, except for trending
towards greater
crowned dentition
correlated with an
abrasive diet) (McGrew
1941, Rensberger 1982).
Until the last 2 million
years, Aplodontia
occupied an area that
extended south into
southern California and Figure 2. Approximate Miocene distribution of Aplodontiarufa. Blue
east to the Great Plains. points represent A. rufa fossil localities, based on data from MioMap.

However, with mid-

Pliocene climate change and the uplift of the Rocky Mountains and Cascade Ranges,
the geographic range of Aplodontia contracted, consistent with its tracking ahighly
habitat-specific niche into its current distribution (Shotwell 1958). Aplodontiarufa
now occupies a disjunct distribution in the Pacific northwest, relict from a much
wider Miocene distribution (Fig. 2). This region is home to many endemics, many of
which are descendents of ancient lineages of plants and animals. For example, more
than 60% of terrestrial and freshwater mollusks species are found in the Pacific
northwest are endemic to the biome. Over 100 species of plants are endemic to the
region and several ancient amphibian lineages are found nowhere else (Carstons et
al. 2005, Hadly, personal communication). These data suggestthat the Pacific
northwestbiome may be a Miocene-era relict habitat essential to the survival of A.
rufa and other such relictual species, underscoring the mountain beaverǯs
importance as a community-wide indicator species. Thus, determing the factors
behind the modern distribution of A. rufa may shed light on the nature of past
climate change in the Pacific Northwest, as well as offer a model for the effects of
future climate change on this biome and its species.


I will use A. rufaosteologicalspecimens from the Museum of Vertebrate Zoology
(MVZ) at the University of California, Berkeley, for my morphological analysis. The
MVZ houses 360 A. rufaspecimens, representing the four California sub-species:A. r.
californica, A. r. humboldtiana, A. r. phaea, and A. r. nigra. Specimens are categorized
by the county of collection, which roughly corresponds with geographically-defined
subspecies. There are an additional 32 and 36 specimens at the California Academy
of Sciences and Humboldt State University, respectively, which I will use to bolster
my sample size to at least 400 specimens. Using linear morphometric techniques, I
will quantify variation in form between subspecies, using standard cranial and post-
cranial measurementsas per Lessa& Stein 1992. This initial linear morphometric
analysis will open the door for further geometric morphometric analysis if
necessary, which allows for the discrimination of more subtle differences in shape.

For this first stage of my project, I will focus only on California subspecies of A.
rufa,in part to demonstrate the feasibility of my project and to initially test my
hypotheses about environmental controls on subspecies distribution and
morphologic variation. Fortuitously, most of the habitat diversity observed between
subspecies occurs in California, making it an apt initial study area. Subsequent
stages of my project (no funding yet requested) will include an extensive latitudinal
sampling of the species through the entirety of its range and into Canada.

Using GIS and relevant spatial statistics, I will perform niche-modeling analysis to
determine which environmental variablesare correlated with the spatial
distribution of A. rufa. I will also apply GIS analyses to see how morphological
differences between subspecies play out spatially and across environmental




My biggest resource is the Hadly Lab at Stanford University. Professor Hadly is
advising me on this project, providing technical and conceptual support. Hadly Lab
members also provide advice and help, and represent a great pool of collective
knowledge. Several members of the lab have performed similar work, and are able
to help me iron out the details in my analyses. The Hadly Lab is also connected to
the MVZ at Berkeley, giving me expedited access to a large collection of A. rufa

I have begun linear morphometric measurements on specimens at the MVZ, with the
aim of quantifying my measurement error and ironing out procedural details. My
current error for 10 measurements over 40 replicates is <2%. I have also begun
training with the GIS applications needed for my spatial distribution analysis and
begun the introduction to GIS tutorial offered by Branner Library. Once I have
achieved competency with the programs, GIS staff at Branner Library and members
of the Hadly Lab will help me with my analysis. Relevant coursework includes
Bio101 (Ecology), Bio136 (Evolutionary Paleobiology) and Anthro206 (Human
Origins, provided an introduction to osteology).

My budget reflects a modest investment in travel to the museums in northern
California (MVZ, Cal Academy and Humboldt State University) that house the
specimens I will need to measure. While travel to Berkeley and the Cal Academy can
be achieved by public transit, travel to Humboldt State University will involve use of
a private vehicle. I am also asking for funds to cover costs for the digital calipers I
will use to measure the specimens, as well as costs associated with printing a poster
to present my preliminary results at a meeting next year.

$  % 

Stanford to MVZ (BART &Caltrain, $16 $160
per trip x 10 trips)
Stanford to California Academy of $80
Science (Bart &Caltrain, $16 per trip x
5 trips)
Lunch at MVZ/Cal Academy ($10 x 15 $150
Stanford to Humboldt State (618 mi RT $315
x 0.51/mile)
Per diem at Humboldt State (x 2 days) $184
cubtotal Travel Î 
Digital Calipers $100
Poster Printing $100
cubtotal cupplies Î
Total Î