Background: High morphological diversity can occur in closely related animals when selection favors morphologies that are subject to intrinsic biological constraints. A good example is subterranean rodents of the genus Thomomys, one of the most taxonomically and morphologically diverse mammalian genera. Highly procumbent, tooth-digging rodent skull shapes are often geometric consequences of increased body size. Indeed, larger-bodied Thomomys species tend to inhabit harder soils. We used geometric morphometric analyses to investigate the interplay between soil hardness (the main extrinsic selection pressure on fossorial mammals) and allometry (i.e. shape change due to size change; generally considered the main intrinsic factor) on crania and humeri in this fast-evolving mammalian clade. Results: Larger Thomomys species/subspecies tend to have more procumbent cranial shapes with some exceptions, including a small-bodied species inhabiting hard soils. Counter to earlier suggestions, cranial shape within Thomomys does not follow a genus-wide allometric pattern as even regional subpopulations differ in allometric slopes. In contrast, humeral shape varies less with body size and with soil hardness. Soft-soil taxa have larger humeral muscle attachment sites but retain an orthodont (non-procumbent) cranial morphology. In intermediate soils, two pairs of sister taxa diverge through differential modifications on either the humerus or the cranium. In the hardest soils, both humeral and cranial morphology are derived through large muscle attachment sites and a high degree of procumbency. Conclusions: Our results show that conflict between morphological function and intrinsic allometric patterning can quickly and differentially alter the rodent skeleton, especially the skull. In addition, we found a new case of convergent evolution of incisor procumbency among large-, medium-, and small-sized species inhabiting hard soils. This occurs through different combinations of allometric and non-allometric changes, contributing to shape diversity within the genus. The strong influence of allometry on cranial shape appears to confirm suggestions that developmental change underlies mammalian cranial shape divergences, but this requires confirmation from ontogenetic studies. Our findings illustrate how a variety of intrinsic processes, resulting in species-level convergence, could sustain a genus-level range across a variety of extrinsic environments. This might represent a mechanism for observations of genus-level niche conservation despite species extinctions in mammals.
Photographs of Northern California Thomomys pocket gopher lateral humeri for 2D Geometric Morphometrics
Zipped folder containing photographs of lateral humerus specimens of genus Thomomys gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically.
Humerus_Lateral.zip
Photographs of Northern California Thomomys pocket gopher anterior humeri for 2D Geometric Morphometrics
Zipped folder containing photographs of anterior humerus specimens of genus Thomomys gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically.
Humerus_Anterior.zip
Landmark coordinates and slider identifications for each specimen view for geometric morphometrics
Landmark coordinates created by tpsdig2 software (Rohlf, Stony Brook) in TPS files the file format of landmark coordinates read in by geomorph, the R package for Geometric Morphometrics (GMM). Semilandmarks or "sliders" are indicated in the associated .CSV files for each view. Please read the ReadMe file for how to use R to combine the TPS and CSV files to recreate the collection of landmarks and semilandmarks we used for our GMM analyses.
Landmark Coordinates.zip
Photographs of Northern California genus Thomomys subgenus Megascapheus pocket gopher lateral crania for 2D Geometric Morphometrics
Zipped folder containing photographs of lateral crania specimens of genus Thomomys subgenus Megascapheus gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically. Combine with the subgenus Thomomys file of the same view before proceeding with analyses.
Crania_Lateral_Megascapheus.zip
Photographs of Northern California Thomomys subgenus Thomomys pocket gopher lateral crania for 2D Geometric Morphometrics
Zipped folder containing photographs of lateral crania specimens of genus Thomomys subgenus Thomomys gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically. Combine with the subgenus Megascapheus file of the same view before proceeding with analyses.
Crania_Lateral_Thomomys.zip
Photographs of Northern California genus Thomomys subgenus Megascapheus pocket gopher ventral crania for 2D Geometric Morphometrics
Zipped folder containing photographs of ventral crania specimens of genus Thomomys subgenus Megascapheus gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically. Combine with the subgenus Thomomys file of the same view before proceeding with analyses.
Crania_Ventral_Megascapheus.zip
Photographs of Northern California Thomomys subgenus Thomomys pocket gopher ventral crania for 2D Geometric Morphometrics
Zipped folder containing photographs of ventral crania specimens of genus Thomomys subgenus Thomomys gophers from our Northern California study region housed in the UC Berkeley Museum of Vertebrate Zoology's (MVZ) mammal collections. Abbreviated taxonomic identification and the MVZ catalog numbers are given in the filenames of each specimen photo. Photos with a number at the end of the filename are photos taken of the same specimen at different times during the digitization process to assess operator error. Photos with "copy" at the end of the filename are photo files that were digitally duplicated to assess landmarking error specifically. Combine with the subgenus Megascapheus file of the same view before proceeding with analyses.
Crania_Ventral_Thomomys.zip
Geo-referrenced NRCS Soil Data for Northern California genus Thomomys pocket gophers
This CSV file contains the soil information extracted from the US Department of Agriculture (USDA)'s Natural Resource Conservation Service (NRCS) STATSGO soil database. The soil characteristics of clay, silt, sand, bulk density, linear extensibility are given as a weighted average for both a depth of 20cm and of 1m; the total soil depth for the area is also reported. These characteristics are associated with all unique specimen localities in our study area. Unique ID refers to the specimen's unique ID number in the Arctos collaborative museum database. The CatNum ID is the Museum of Vertebrate Zoology (MVZ) catalog number we use to identify specimens throughout our analyses. For more information about this dataset, please see Marcy et al (2013) PLOS ONE doi: http://dx.doi.org/10.1371/journal.pone.0064935
GeoreferrencedNRCSSoilData.csv