Estimating Body Mass of Fossil Rodents
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Estimating body mass of fossil rodents Matthijs Freudenthal & Elvira Martín-Suárez Freudenthal, M. & Martín-Suárez, E. 2013. Estimating body mass of fossil rodents. Scripta Geologica, 145: 1-130, 8 appendices including one on-line, 4 tables, 5 figures. Leiden, November 2013. M. Freudenthal, Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avda. Fuen- tenueva s/n, E-18071 Granada, Spain; Naturalis Biodiversity Center, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands ([email protected]). E. Martín-Suárez, Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avda. Fuen- tenueva s/n, E-18071 Granada, Spain. Keywords – Rodentia, body mass. Reconstructing the body mass of a fossil animal is an essential step toward understanding its palaeoecol- ogical role. Length × width (L×W) of the first lower molar (m1) is frequently used as a proxy for body mass in fossil mammals. However, among rodents, Muroidea have no premolar and an elongated m1, whereas other groups have a premolar and a m1 that is not elongated. This leads to an overestimation of body mass in muroids and/or an underestimation in other rodents. To solve this problem we assembled data of upper and lower tooth row length and body mass in extant rodents, and calculated regression equations for all rodents, rodents with premolars, rodents without premolars and for taxonomic groups at superfamily or family level. Data for complete tooth rows in fossil rodents are scarce, so we took the sum of the lengths of the (three or four) cheek teeth as an approximation of tooth row length. We estimate body mass of the fossil rodents, using the regression equations of the extant taxa. Contents Introduction ................................................................................................................................................................. 1 Material and methods ........................................................................................................................................... 3 Fossil data ..................................................................................................................................................................... 3 Recent data ................................................................................................................................................................... 4 Taxonomic groups ................................................................................................................................................... 5 Use of (natural) logarithms ................................................................................................................................ 5 Correction factors ..................................................................................................................................................... 6 Analysis of L/W of m1 per taxonomic group .......................................................................................... 6 Analysis of toothrow length vs body mass .............................................................................................. 8 Synopsis ...................................................................................................................................................................... 13 Discussion .................................................................................................................................................................. 13 Conclusions .............................................................................................................................................................. 14 Acknowledgements ............................................................................................................................................. 14 References .................................................................................................................................................................. 14 Note: Appendix 8 (a+b) is available digitally at the Scripta Geologica website; all other appendices are printed herein .................................................................................... 15 Introduction Reconstructing the body mass of a fossil animal is an essential step toward under- standing its palaeoecological role (Hopkins, 2008). Legendre (1989) used length × width (L×W) of m1 (first lower molar) as a surrogate for tooth area and based body mass esti- mates on log (L×W). Among rodents, Muroidea have no premolar and an elongated m1, 2 Freudenthal & Martín-Suárez. Estimating body mass of fossil rodents Scripta Geol., 145 (2013) Figure 1. Length/width diagram of m1 in >2300 fossil rodent populations (measurements of population means in 1 mm units). Regression lines diverge. Figure 2. Length/width diagram of m1 in >2300 fossil rodent populations (logarithms of population means in 0.1 mm units). Regression lines are parallel. Freudenthal & Martín-Suárez. Estimating body mass of fossil rodents Scripta Geol., 145 (2013) 3 whereas other groups have a premolar and a m1 that is not elongated. According to Hopkins (2008), this leads to an overestimation of body mass in murids (and/or an un- derestimation in other rodents). Figures 1 and 2 give linear and logarithmic mean length/mean width plots of m1 of a large number of fossil rodent populations, in which the muroid group and the non-muroid group are clearly distinguished. The all-rodents regression equation of Legendre is based on 61 species with premolar and 154 species without premolar. With a different sample composition, the regression equation may change considerably. Evidently, body mass estimates on the basis of m1 area for Muroidea and other rodents cannot be based on the same regression equation. Hopkins (2008) investigated tooth row length and area as proxies for body mass in rodents. One of her conclusions is that the results for both proxies are good for all rodents and that they are better for individual clades. “Within species, toothrow measurements are poor predictors of individual body mass, because the complete adult dentition is generally erupted early in life, whereas body mass continues to change as an animal reaches full adult mass” (Hopkins, 2008, p. 238). To this we might add that body mass may vary considerably throughout the year and with changing habitat conditions. Tooth row length in extant rodents is frequently reported, but data for individual tooth measurements are scarce. We assembled data for tooth row length and body mass of extant rodents, and calculated least squares linear regression lines for the entire data set and for a number of subgroups. On the other hand, tooth row length is only available for a limited number of fossil rodent species and, therefore, we looked for another parameter that would permit body mass estimates of fossil rodents. We calculated the sum of the lengths of the cheek teeth (LRsum and URsum) as a surrogate for (alveolar) tooth row length and used the regres- sion lines of extant rodents to estimate body mass of the fossil species. In order to test the validity of m1 area for body mass estimates, as did Legendre (1989), we will analyze the relationship L/W of fossil m1 in various taxonomic groups and signal the taxa that are most different from the mean. Then we discuss several correction factors that should be applied to the body mass results. Finally, we analyze the reliability of body mass estimates of fossil rodents, obtained from the regression equations of extant rodents for which tooth row length and body mass are known. Ide- ally, body mass estimates should be equal when using the upper or the lower tooth row, but this is frequently not the case and we investigate the causes of these differences. Material and methods Abbreviations used herein are listed in Table 1. Fossil data Tooth row lengths (LRsum and URsum) are calculated from our database of tooth measurements of fossil rodents which contains about 2650 populations from 948 locali- ties. Of these, 1600 populations contain data for all lower cheek teeth and complete up- per rows can be composed for 1485 populations. Some 1270 populations have data for all upper and lower cheek teeth, and permit comparison of LRsum and URsum. The values of LRsum and URsum are used to estimate body mass, using the regression lines obtained from the Recent material. 4 Freudenthal & Martín-Suárez. Estimating body mass of fossil rodents Scripta Geol., 145 (2013) Table 1. Abbreviations used. LTR lower tooth row length (recent) UTR upper tooth row length (recent) LRsum sum of lengths of lower teeth (fossil) URsum sum of lengths of upper teeth (fossil) L, W length, width LxW length x width ~ tooth area p4, m1, m2, m3 lower teeth P3, P4, M1, M2, M3 upper teeth WithP rodents with premolar WithoutP rodents without premolar InfMass mass estimate based on LRsum SupMass mass estimate based on URsum MeanMass (InfMass + SupMass)/2 SEE standard error of estimate &PE percentage error of estimate df degrees of freedom RGM Rijksmuseum van Geologie en Mineralogie, now Naturalis Biodiversity Center, Leiden Tooth row length for the fossil species (LRsum and URsum) is calculated as the sum of the mean lengths of the three molars per population, plus the mean length of the premolar in taxa that have a premolar. In Sciuridae, Gliridae, Eomyidae and Therido- morpha, the first and second molars are often not distinguished. In these cases length of m1,2, resp. M1,2 is counted twice, thus the mean length of m1,2 replaces