Figures and references for Ariel Marcy’s Major Grant Proposal

Variation in pocket gopher (Thomomys) digging morphology as a

determining factor in species distribution in northeastern California

Fig 1: Distribution map of Thomomys genus pocket gophers, for reference, top border is
Oregon, right border is Nevada (from Thaeler 1968)

Fig 2: PCA illustrating how genera and two species of Thomomys spread on axes of size and
claw- or tooth-digging traits (From Lessa and Stein 1992)
Fig 3. Measurements for morphometric analysis (from Lessa and Stein 1992)

Table 1: Linear morphometrics measurements, abbreviations, grouped by expected

changes for each digging mode
Measurement Abbr.
Humerus length
Deltoid process
Width of epicondyles
Length of ulna
Olecranon process
Basilar length of skull
Zygomatic breadth
Claw-digging
HUML Decreased (higher forces experienced in arms)*
DELTW Increased (greater muscle attachment)2
EPICW Increased (greater muscle attachment)2
ULENG Increased (due to increase in OLCER)*
OLCER Increased (greater lever for triceps)*
BAL Decreased (dorsal flattening leads to greater
surface area for muscle attachment)*
ZB Increased (more surface area for jaw muscles)*

Measurement Abbr. Tooth-digging

Rostral width RW Decreased (smaller cross-area of incisors decrease
R)1
Rostral breadth RD Increased (elongated to increase procumbency)2
Width of incisors INCW Decreased (smaller cross-area of incisors)1
Length of diastema DIAST Increased (indicator of procumbency)2
Vertical width of incisors INCL Increased (indicator of procumbency)2
Distance from tip of incisor to BASE Increased (indicator of procumbency)2
base of first molar
Rationales for predictions were adapted from Lessa 1990 (denoted by 1), Lessa and Stein 1991
(denoted by 2), or extensions were made from Stein 2000, which was not specifically discussing
differences between modes of digging but subterranean adaptations in general (denoted by *)
 Fig. 4. PCA of my preliminary linear measurements PC1 is correlated with size.

Fig 5. Scatter plot of P21 versus deltoid width shows that PC2 is moderately correlated to
variation in the deltoid process.
 Fig 6: Landmarks for rodent mandible (From Zelditch and Swiderski 2009)

Fig 7: PCA results of landmark-based GM of a rodent cranium (From Zelditch and Swiderski
2009)

References

1. Anderson, R. P., Peterson, A. T. and Gomez-Laverde, M. 2002. “Using niche-based

GIS modeling to test geographic predictions of competitive exclusion and competitive
release in South American pocket mice.” Oikos 98: 3 – 16

2. Barnosky, A.D., 1982. “Locomotion in Moles (Insectivora, Proscalopidae) from the

Middle Tertiary of North America.” Science 216, 183-185.

3. Blois, J., 2008. “Small mammal response to the Pleistocene-Holocene transition in

northern California.” Journal of Vertebrate Paleontology 28, 53A -54A.

4. Cameron, G.N., 2000. Community Ecology of Suberranean Rodents. In: Lacey E.A.,
Patton J.L., Cameron G.N., (Eds.), Life Underground. The University of Chicago
Press, Chicago, pp 227-256.

5. Cartini, A. and O’Higgins. 2004. “Patterns of morphological evolution in Marmota

(Rodentia, Sciuridae): geometric morphometrics of the cranium in the context of
marmot phylogeny, ecology and conservation.” Biological Journal of the Linnean
Society, 82, 385–407.
6. Cox, G.W. and Hunt, J., 1994. “Pocket gopher herbivory and mortality of Ocotillo on
Stream Terrace, Bajada, and hillside sites in the Colorado Desert, Southern
California.” The Southwestern Naturalist 39, 364-370.

7. Eviner, V.T. and Chapin, F.S., 2003. “Gopher-plant-fungal interactions affect

establishment of an invasive grass.” Ecological Society of America 84, 120-128.

8. Hickman, G.C., and Brown L.N.,1973. “Mound- building behavior of the southeastern
pocket gopher.” Journal of Mammology 54, 786- 790.

9. Lessa, Enrique P., and Barbara R. Stein., 1992. "Morphological constraints in the
digging apparatus of pocket gophers." Biological Journal of the Linnaen Society 47,
439-53.

10. Lessa, Enrique P., and Charles S. Thaeler, Jr., 1989. "A reassessment of
morphological specializations for digging pocket gophers." Journal of Mammology
70, 689-700.

11. Lessa, Enrique P., 1990. "Morphological Evolution of Subterranean Mammals:

Integrating Structural, Functional, and Ecological Perspectives." In Liss A.R. (Ed.),
Evolution of Subterranean Mammals at the Organismal and Molecular Levels pp 211-
30.

12. Luna, F. and Antinuchi, C.D., 2006. “Cost of foraging in the subterranean rodent
Ctenomys talarum: effect of soil hardness.” Canadian Journal of Zoology 84, 661-
667.

13. Miller, M.A. and Gross, M.M., 1998. “Locomotor advantages of Neandertal skeletal
morphology at the knee and ankle.” Journal of Biomechanics 31, 355-361.

14. Patton, personal communication

15. Rayfield, E.J., 2007. “Finite Element Analysis and Understanding the Biomechanics
and Evolution of Living and Fossil Organisms.” Annual Review of Earth and
Planetary Sciences 35, 541-576.

16. Romanach et al, 2005. “Effects of species, sex, age, and habitat on geometry of
pocket gopher foraging tunnels.” Journal of Mammalogy 86, 750-756.

17. Stein, B.R., 1993. Comparitive Hind Limb Morphology in Geomyine and
Thomomyine Pocket Gophers. Journal of Mammalogy, 74, 86-94.

18. Stein, B.R., 2000. Morphology of Subterranean Rodents. In: Lacey E.A., Patton J.L.,
Cameron G.N., (Eds.), Life Underground. The University of Chicago Press, Chicago,
pp 20-61.
19. Thaeler, Charles S. Jr., 1968. An Analysis Of The Distribution Of Pocket Gopher
Species In Northeastern California. PhD. Thesis, University of California Press,
Berkeley and Los Angeles.

20. Weijs WA, Hillen B. 1985. “Cross-sectional areas and estimated intrinsic strength of
the human jaw muscles.” Acta Morphol. Neerlando-Scand 23, 267–74.

21. Zeldich M.L, Swiderski, H.D.S., and Fink, W.L., 2004. “Geometric Morphometrics
for Biologists: APrimer.” Elsevier, San Diego.

22. Zelditch, M.L., and Swiderski, H.D.S., 2009. “2009 Berkeley Geometric
Morphometrics Workshop.” Berkeley, California.