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Estimating Head and Body Length in Fossil Rodents

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Estimating Head and Body Length in Fossil Rodents Estimating head and body length in fossil rodents Matthijs Freudenthal & Elvira Martín-Suárez Freudenthal, M. & Martín-Suaréz, E. Estimating head and body length in fossil rodents. Scripta Geologica, 149: 1-158, 4 figs., 8 tables, Leiden, August 2015. M. Freudenthal, Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avda. Fuen- tenueva s/n, E-18071 Granada, Spain, and Naturalis Biodiversity Center, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands ([email protected]); E. Martín-Suárez, Departamento de Estratigrafía y Pale- ontología, Universidad de Granada, Avda. Fuentenueva s/n, E-18071 Granada, Spain. Key words – body mass, teeth, tooth row length. We present estimates for head and body length (HBL) of fossil rodents. We assembled HBL data and tooth row length data (LTR, UTR) for a large number of extant rodents, and calculated regression lines of HBL on LTR and UTR for all rodents together (all-rodents equation), and for separate taxonomic groups (family equations). In fossil rodents, data for complete tooth rows are scarce, therefore we use the sum of the lengths of the teeth (LRsum, URsum) as a surrogate for tooth row length. The relation between this parameter and real tooth row length (LTR, UTR) is calculated on the basis of a number of populations for which both parameters are available. We estimate HBL of fossil rodents, using LRsum and URsum and the regression lines of extant rodents, and we compare the results for lower and upper tooth row when both are available. For each species we calculate HBL through the all-rodents equation and through the family equation. We consider the amount of difference between these two values as a measure of the reliability of this method. Contents Introduction ................................................................................................................................................................. 1 Material and methods ........................................................................................................................................... 2 Validity of the regression equations ............................................................................................................. 5 Comparison of InfHbl and SupHbl ............................................................................................................... 7 Comparison of HBL from family equations and all-rodents equation .................................... 8 Relation between measured rows and sum of tooth lengths ....................................................... 11 Conclusions ............................................................................................................................................................... 12 Acknowledgements .............................................................................................................................................. 12 References .................................................................................................................................................................. 13 Note: Appendix 9a is available digitally at the Scripta Geologica website; all other appendices are printed herein Introduction Freudenthal & Martín-Suárez (2013) published an approach to body mass estimates of fossil rodents on the basis of the parameters LRsum and URsum, that is, the sum of the mean lengths of the individual teeth that constitute the lower and upper tooth rows. Since then we have considerably augmented our data base of fossil rodent measure- ments and recent rodent tooth row lengths, and we have added and corrected some body mass data for extant rodents. Besides body mass, head and body length (HBL) is a second parameter commonly found in studies on Recent mammals, and here we use it to further test the validity of 2 Freudenthal & Martín-Suaréz. Estimating head and body length in fossil rodents. Scripta Geol., 149 (2015) LRsum and URsum for estimating the body size of fossil rodents. We have created a data base for HBL of extant rodents, which contains data for almost 80 % of the known species. We also assembled tooth row length data (LTR, UTR) for a large number of extant rodents, and calculated regression lines of HBL on LTR and UTR for all rodents to- gether, and for separate taxonomic groups. In fossil rodents, data for complete tooth rows are scarce, therefore we use the sum of the lengths of the teeth (LRsum, URsum) as a surrogate for tooth row length. The relation between this parameter and real tooth row length (LTR, UTR) is calculated on the basis of a number of populations for which both paramaters are available. We estimate HBL of fossil rodents, using LRsum and URsum, and the regression lines of extant rodents. The results for seven taxonomic groups are given in Appendices 1-7. Material and methods Abbreviations – Abbreviations used herein are listed in Table 1. Recent data – We assembled data for upper and lower tooth row length (UTR and LTR) from several main sources, including Mammal Species Accounts (http://www. science.smith.edu/ departments/Biology/VHAYSSEN/msi/), African rodent database (http://projects.biodiversity.be/africanrodentia/; Hollister, 1919) and a large number of Table 1. Abbreviations used. LTR lower tooth row length (Recent) UTR upper tooth row length (Recent) UTRalv upper tooth row alveolar length (Recent) UTRcr upper tooth row crown length (Recent) LRsum Sum of lengths of lower teeth (fossil) URsum Sum of lengths of upper teeth (fossil) L, W Length, Width LxW length × width ~ tooth area p4, m1, m2, m3 lower teeth P3, P4, M1, M2, M3 upper teeth InfMass mass estimate based on LRsum SupMass mass estimate based on URsum MeanMass (InfMass + SupMass)/2 HBL Head and body length InfHbl HBL estimate based on LRsum SupHbl HBL estimate based on URsum MeanHbl (InfHbl + SupHbl)/2 linhbl HBL based on linear data loghbl HBL based on detransformed log data SEE standard error of estimate &PE Percentage error of estimate df degrees of freedom log natural logarithm RGM Rijksmuseum van Geologie en Mineralogie, now Naturalis Biodiversity Center, Leiden, the Netherlands Freudenthal & Martín-Suaréz. Estimating head and body length in fossil rodents. Scripta Geol., 149 (2015) 3 smaller sources (see Appendix 8b). We have tooth row data for 1035 lower rows and 2767 upper rows, and HBL data for 1793 of the 2277 Recent rodent species. Our taxonomic basis is MSW3 (Mammal species of the World 3) as found on the internet in 2005 (http:// www.bucknell.edu/msw3/). Simple linear least squares regression lines were calculat- ed from our LTR, UTR and HBL data, and these are used to estimate head and body length of fossil rodents. HBL values used in the calculations are the midpoint of the minimum and maximum values found for each species; tooth row values are the mid- point per population. In both cases we use the midpoint, because mean or median are commonly not available. HBL data in cm are given in Appendix 8a, literature references in Appendix 8b. Our taxonomic groups represent different taxonomic levels; some are families (or even subfamilies, according to different authors) and others are higher taxonomic units. Eomyidae are classified in the Geomyoidea; therefore, Recent Geomyoidea are used as an equivalence for Eomyidae. The equivalence between Recent and fossil groups is given in Table 2. Theridomorpha has no recent equivalent. Fossil data – Tooth row lengths (LRsum and URsum) are calculated from our data- base of tooth measurements of fossil rodents which contains about 3450 populations from 1133 localities. Of these, 2187 populations contain data for all lower cheek teeth and complete upper rows can be composed for 2055 populations. Some 1755 popula- tions, representing 753 species, have data for all upper and lower cheek teeth, and per- mit comparison of LRsum and URsum. The values of LRsum and URsum are used to estimate HBL using the regression lines obtained from the Recent material. 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 Table 2. Equivalence between Recent and fossil groups. Column 2 (= MSW3) gives the number of species recognized in Mammal Species of the World (Wilson & Reeder, 2005). Column 4 gives the number of fossil species used in this work, for which both upper and lower tooth row could be composed, discarding open nomenclature (sp.). Recent MSW3 fossil Column 4 all-rodents 2277 all-rodents 745 Sciuridae 279 Sciuridae 48 Gliridae incl. Graphiurinae 28 Gliridae 168 Geomyoidea 100 Eomyidae 70 Castoridae 2 Castoridae 2 Dipodidae 51 Dipodidae 7 Nesomyidae 61 Arvicolidae 151 Arvicolidae 14 Cricetidae 530 Cricetidae 285 Muridae 730 Muridae 78 Spalacidae 36 Spalacidae 5 Hystricomorpha 290 Hystricomorpha Theridomorpha 66 Various 19 Various 4 Freudenthal & Martín-Suaréz. Estimating head and body length in fossil rodents. Scripta Geol., 149 (2015) of m1,2, resp. M1,2, is counted twice, thus the mean length of m1,2 replaces the mean length of both m1 and m2. Uncommonly, in Gliridae, a P3 may be present. For calcula- tion of URsum we ignored this element, because it is hardly ever found, if present it is usually very small, its presence may vary within a species, it is probably shed early in life and is hardly ever functional, and for most species it
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