This is the accepted version of the journal article: López-Torres, Sergi; Bertrand, Ornella C.; Lang, Madlen M.; [et al.]. «Cranial endocast of Anagale gobiensis (Anagalidae) and its implications for early brain evolution in Euarchontoglires». Palaeontology, Vol. 66, Issue 3 (May/June 2023), art. e12650. DOI 10.1111/pala.12650 This version is available at https://ddd.uab.cat/record/275489 under the terms of the license Page 1 of 78 1 2 3 Cranial endocast of Anagale gobiensis (Anagalidae) and its implications for early brain 4 5 6 evolution in Euarchontoglires 7 8 9 Sergi López-Torres1,2,3,4, Ornella C. Bertrand5, Madlen M. Lang6, Łucja Fostowicz- 10 11 Frelik7,8, Mary T. Silcox6, Jin Meng1,4 12 13 1Division of Paleontology, American Museum of Natural History, 10024-5192, New 14 15 16 York, NY, United States of America. JM ORCiD: 0000-0002-3385-8383 17 18 19 2Richard Gilder Graduate School, American Museum of Natural History, 10024-5192, 20 21 New York, NY, United States of America. 22 23 24 3University of Warsaw, Faculty of Biology, Biological and Chemical Research Centre, 25 26 27 Institute of Evolutionary Biology, Warsaw, Poland. SLT ORCiD: 0000-0002-0046- 28 29 1013 30 31 4New York Consortium in Evolutionary Primatology, New York, NY, United States of 32 33 34 America. 35 36 37 5Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, 38 39 Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès. 40 41 42 Barcelona, Spain. OCB ORCiD: 0000-0003-3461-3908 43 44 6Department of Anthropology, University of Toronto Scarborough, M1C 1A4, Toronto, 45 46 47 ON, Canada. MML ORCiD: 0000-0003-2604-4733. MTS ORCiD: 0000-0002-4174- 48 49 9435 50 51 52 7Department of Organismal Biology and Anatomy, The University of Chicago, 53 54 55 Chicago, IL, United States of America. ŁFF ORCiD: 0000-0002-1266-1178 56 57 8Institute 58 of Paleobiology, Polish Academy of Sciences, Warsaw, Poland. 59 60 1 Page 2 of 78 1 2 3 4 5 6 7 8 9 Corresponding author: Sergi López-Torres. Postal address: Żwirki i Wigury 101, 10 11 12 Warsaw 02-089, Poland. Telephone: +48-730-980-445. E-mail: s.lopez- 13 14 torres@uw.edu.pl. 15 16 17 Short running title: Cranial endocast of an anagalid 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 2 Page 3 of 78 1 2 3 ABSTRACT 4 5 6 Anagalids are an extinct group of primitive mammals from the Asian Palaeogene 7 8 9 thought to be possible basal members of Glires. Anagalid material is rare, with only a 10 11 handful of crania known. Here we describe the first virtual endocast of an anagalid, 12 13 based on the holotype of Anagale gobiensis (AMNH 26079; late Eocene, China), which 14 15 allows for comparison with published endocasts from fossil members of modern 16 17 18 euarchontogliran lineages (i.e. primates, rodents, lagomorphs). The endocast displays 19 20 traits often observed in fossorial mammals, such as relatively small petrosal lobules and 21 22 a low neocortical ratio, which would be consistent with previous inferences about use of 23 24 25 subterranean food sources based on heavy dental wear. In fact, Anagale gobiensis has 26 27 the lowest neocortical ratio yet recorded for a euarchontogliran. This species was 28 29 olfaction-driven, based on the relatively large olfactory bulbs and laterally expansive 30 31 32 palaeocortex. The endocast supports previous inferences that relatively large olfactory 33 34 bulbs, partial midbrain exposure and low encephalisation quotient are ancestral for 35 36 Euarchontoglires, although the likely fossorial adaptations of Anagale gobiensis may 37 38 also partly explain these traits. While Anagale gobiensis is a primitive mammal in many 39 40 41 aspects, some of its derived endocranial traits point towards a new, different trajectory 42 43 of brain evolution within Euarchontoglires. 44 45 46 47 48 49 Keywords: Anagalidae, Eocene, China, endocast, brain evolution 50 51 52 53 54 55 56 57 58 59 60 3 Page 4 of 78 1 2 3 INTRODUCTION 4 5 6 Anagalidae is an enigmatic and poorly studied group of primitive mammals known 7 8 from the Palaeogene of China and Mongolia (Hu, 1993; Li, 2016; López-Torres & 9 10 11 Fostowicz-Frelik, 2018a). Although anagalids were never common in the fossil record, 12 13 their species diversity drops substantially after the middle Palaeocene (Nongshangian 14 15 Asian Land Mammal Age [ALMA]), with only Anagale gobiensis Simpson, 1931 and 16 17 18 Zofiagale ergilinensis López-Torres & Fostowicz-Frelik, 2018a being known from the 19 20 late Eocene (Ulangochuian and Ergilian Asian Land Mammal Ages, respectively; 21 22 López-Torres & Fostowicz-Frelik, 2018a) and Anagalopsis kansuensis Bohlin, 1951, 23 24 possibly coming from the earliest Oligocene (Bohlin, 1951; but see Zhang & Wang 25 26 27 [2016] for an earlier estimate). No anagalids have been reported from the late 28 29 Palaeocene (Gashatan ALMA) and most of the Eocene (Bumbanian to Sharamurunian 30 31 ALMAs) (Li et al., 2016; López-Torres & Fostowicz-Frelik, 2018a). 32 33 34 There has been considerable debate about which genera should be included in 35 36 37 Anagalidae, which relates to the question of the group’s broader affinities. On the one 38 39 hand, workers have used Anagalidae as a convenient taxonomic label for a diversity of 40 41 forms that seem to share some similarities to Glires without clearly pertaining to one of 42 43 44 its modern lineages (Szalay & McKenna, 1971; Wang, 1975; Xu, 1976a, b; Huang & 45 46 Zheng, 1983). Glires is comprised of the extant orders of Rodentia and Lagomorpha, 47 48 and both comprehensive morphological (e.g. Li et al., 1989; Meng et al., 2003; 49 50 Marivaux et al., 2004; Meng, 2004) and molecular (e.g. Murphy et al., 2001; Huchon et 51 52 53 al., 2002; Poux et al., 2006) analyses have established the monophyly of this group. In 54 55 turn, Glires belongs, together with Euarchonta (Dermoptera, Scandentia and Primates; 56 57 Archonta of Asher and Helgen, 2010), in the higher-level clade Euarchontoglires 58 59 60 (Springer et al., 2004; O’Leary et al., 2013). On the other hand, a more restrictive sense 4 Page 5 of 78 1 2 3 was used by Hu (1993), who considered just seven of the 15 genera often included in 4 5 6 the group to be unquestionable anagalids (McKenna & Bell, 1997; Li, 2016). Here we 7 8 employ Hu’s (1993) more restrictive sense of anagalids. 9 10 11 In terms of their relationships to modern groups, historically anagalids had been 12 13 suggested to be closely related to Scandentia (Simpson, 1931), to Glires (Bohlin, 1951), 14 15 and later to Macroscelidea and Lagomorpha (McKenna, 1975). In modern cladistic 16 17 18 contexts, a diversity of possible relationships have been recovered. Meng et al. (2003), 19 20 included both Anagale and Anagalopsis in their phylogenetic analysis, in which 21 22 Anagalidae was recovered as sister to Glires. Similarly, Meng (2004) and Fostowicz- 23 24 25 Frelik (2017) placed Anagalidae as sister to Glires. In contrast, work by Asher and 26 27 colleagues (Asher et al., 2003, 2005, 2019; Asher, 2007) have recovered a diversity of 28 29 possible resolutions to the phylogenetic position of the group, including relationships to 30 31 32 Scandentia, a position on the lagomorph stem, a position at the base of Rodentia, as 33 34 sister group to the order Macroscelidea (elephant shrews), as an early diverging 35 36 placental lineage, or as unresolved. One factor likely contributing to this diversity of 37 38 results is the fact that most analyses have sampled only a small subset of the anagalids 39 40 41 known (even if the group is restricted to the subset included by Hu, 1993), and are 42 43 based on character matrices that were not designed to address the question of anagalid 44 45 relationships specifically. López-Torres and Fostowicz-Frelik (2018b) sampled the 46 47 48 entire group (including both Hu’s [1993] unquestionable anagalids, and anagalids sensu 49 50 lato) and employed characters that were specifically designed to tease apart anagalid 51 52 relationships. Their preliminary results placed most anagalid species on the Glires stem 53 54 55 (including Anagale), except for some species that were nested within Euarchonta. So, 56 57 while questions still remain about anagalid relationships, the evidence currently 58 59 available points to relationships with Euarchontoglires. As such, anagalids have the 60 5 Page 6 of 78 1 2 3 potential to be informative about primitive states in both Glires and Euarchontoglires. In 4 5 6 particular, Anagale gobiensis is one of the most basal anagalids according to López- 7 8 Torres and Fostowicz-Frelik’s (2018a) phylogenetic analysis, making it of particular 9 10 relevance to that perspective, in spite of its fairly late age (late Eocene). 11 12 13 The palaeoecology of anagalids remains largely unknown. Anagale gobiensis 14 15 has significantly worn-down cheek teeth and powerful forelimbs, which led McKenna 16 17 18 (1963) to suggest that they may have obtained fairly abrasive food below the surface of 19 20 the ground. This view is consistent with Bohlin’s (1951) suggestion that anagalids were 21 22 potentially fossorial animals. Also, the high degree of dental wear observed in anagalids 23 24 25 is similar to that observed in some extinct fossorial rodents such as Tsaganomys and 26 27 Cyclomylus (Bryant & McKenna, 1995; Wang, 2001). 28 29 30 This current paper presents the first endocranial data for any anagalid, deriving 31 32 from a virtual endocast of Anagale gobiensis (AMNH 26079; holotype) from the late 33 34 35 Eocene of Nei Mongol, China. In the last decade, numerous descriptions of virtual 36 37 endocasts of fossil euarchontogliran taxa have been published, in particular those of 38 39 primates (Silcox et al., 2009, 2010a; Kirk et al., 2014; Orliac et al., 2014; Harrington et 40 41 al., 2016, 2020; Ramdarshan & Orliac, 2016) and rodents (Bertrand & Silcox, 2016; 42 43 44 Bertrand et al., 2016a, 2017, 2018, 2019), but cranial endocasts have also been 45 46 generated for lagomorphs (López-Torres et al., 2020) and apatemyids (von 47 48 Koenigswald et al., 2009; Silcox et al., 2011). These studies have shed light on the early 49 50 51 brain evolution of modern mammalian lineages. For example, “plesiadapiform” 52 53 endocasts provide some data relevant to the earliest phases of primate brain evolution 54 55 (Silcox et al., 2009, 2010a, 2020; Orliac et al., 2014; Long et al., 2015), Rhombomylus 56 57 58 turpanensis Zhai, 1978 and “ischyromyids” are relevant to rodent brain evolution 59 60 (Meng et al., 2003; Bertrand & Silcox, 2016; Bertrand et al., 2016a, 2017, 2018, 2019) 6 Page 7 of 78 1 2 3 and Megalagus turgidus Cope, 1873 to lagomorph brain evolution (López-Torres et al., 4 5 6 2020). These endocasts have also been used to make broader-based inferences about 7 8 what might have been primitive in the form of the brain for Euarchonta, Glires and 9 10 Euarchontoglires (Silcox et al., 2010a; Bertrand et al., 2016a; López-Torres et al., 11 12 2020). However, groups that branched off nearer to the base of Euarchontoglires have 13 14 15 the potential to offer particularly salient insights into these earlier evolutionary stages 16 17 (Silcox et al., 2011). Therefore, the study of the anagalid brain is key for a holistic 18 19 understanding of the evolution of the brain in Euarchontoglires, and particularly Glires 20 21 22 (see Fig. 1). 23 24 25 26 27 MATERIALS AND METHODS 28 29 30 Specimen AMNH 26079 includes an almost complete cranium with a portion of the 31 32 33 right parietal missing and damaged premaxilla (Fig. 2). The specimen was described by 34 35 Simpson (1931) as the holotype of the new genus and species Anagale gobiensis and it 36 37 was later re-described by McKenna (1963), together with a fragmentary rostrum 38 39 40 (AMNH 26141). The cranium of Anagale gobiensis comes from the Ulan Gochu 41 42 Formation of Twin Oboes, Nei Mongol, China (Simpson, 1931) and is dated to the 43 44 Ulangochuian Asian Land Mammal Age. Early work considered the Ulangochuian to be 45 46 early Oligocene (Simpson, 1931; McKenna, 1963; Romer, 1966; Li & Ting, 1983; Tong 47 48 49 et al., 1995), but later work correlated it to the Chadronian North American Land 50 51 Mammal Age, reaching a consensus that the Ulangochuian is late Eocene in age (Wang, 52 53 1997; Meng & McKenna, 1998; Zhang & Li, 2000; Luterbacher et al., 2004; Wang et 54 55 56 al., 2007, 2019; Zhang & Wang, 2016), between 37.2 and 39.9 Ma (Wang et al., 2019). 57 58 59 60 7 Page 8 of 78 1 2 3 The cranium of Anagale gobiensis (AMNH 26079) was scanned at the X-ray 4 5 6 computed tomography scanner at the American Museum of Natural History. The 7 8 specimen was scanned with an 85.80 mm distance from the source. The energy settings 9 10 were 155 kv and 135 μA. A total of 1800 projections were generated, with a voxel size 11 12 of 0.021118 mm. The digital specimen was captured in 2905 slices, each 1371 by 1758 13 14 15 pixels. 16 17 18 The endocranial cavity of Anagale gobiensis was manually segmented in Avizo 19 20 9.0.1 (Visualization Science Group, 1995-2015) with a WACOM Cintiq 21UX tablet. A 21 22 new labelfield module was created in the segmentation editor and the inside of the 23 24 25 endocranial cavity was isolated using the pen tool for approximately every five slices, 26 27 with the intervening slices segmented with the interpolating tool. In all cases, 28 29 interpolation was checked to ensure it accurately followed the endocranial shape with 30 31 32 edits made when needed. When bone was missing, a straight line was traced from the 33 34 two nearest pieces of preserved bone. The cranium is practically undistorted, making the 35 36 resulting quantitative measurements highly reliable. 37 38 39 The endocast of Anagale gobiensis (Fig. 3) is compared to the sample of previously 40 41 published endocasts pertaining to early members of Euarchontoglires. In particular, the 42 43 44 comparative sample includes early Glires, that is, “ischyromyids” (Paramys, 45 46 Pseudotomus, Reithroparamys, Rapamys, and Ischyromys; Bertrand & Silcox, 2016; 47 48 Bertrand et al., 2016a, 2019), fossil sciuroids (Bertrand et al., 2017, 2018), 49 50 51 Rhombomylus (Meng et al., 2003) and a stem lagomorph (López-Torres et al., 2020); 52 53 Primates, including “plesiadapiforms” (Silcox et al., 2009, 2010a; Orliac et al., 2014) 54 55 and early Tertiary fossil euprimates (Kirk et al., 2014; Harrington et al., 2016, 2020; 56 57 58 Ramdarshan and Orliac, 2016); and the most primitive apatemyid known from 59 60 endocranial data (Labidolemur kayi; Silcox et al., 2011). A sample of extant rodents 8 Page 9 of 78 1 2 3 (Bertrand & Silcox, 2016; Bertrand et al., 2017, 2018, 2019) and extant lagomorphs 4 5 6 (López-Torres et al., 2020) were used to provide context to the fossil data in the 7 8 quantitative analyses. Also, a high-quality 1:1 cast of the endocast of Anagale gobiensis 9 10 was 3D-printed to facilitate examination and to validate our observations on the 11 12 computer screen. 13 14 15 Surface area, length and volumetric measurements were taken on the endocasts to 16 17 18 calculate several ratios using Avizo 9.0.1 (Table 1). The volume of the endocast of 19 20 Anagale gobiensis was obtained by isolating the more complete half of the endocast 21 22 using the ‘volume edit’ module and doubling its volume. The volume of the olfactory 23 24 25 bulbs was obtained by isolating the two olfactory bulbs, also using volume edit. The 26 27 petrosal lobules were re-segmented in the transverse plane (XZ dimension) using a 28 29 distinct labelfield module and the volume measured from the resulting reconstruction. 30 31 32 The term petrosal lobule is used here rather than alternatives like “paraflocculus”, 33 34 following Lang et al. (2022). The homology of the cerebellar structures that are housed 35 36 in the subarcuate fossa can vary between orders, which has implications for their 37 38 nomenclature and how we understand their functions between taxa. In Glires (Tan et al., 39 40 41 1995; Panezai et al., 2020) and Scandentia (Ni et al., 2018) the lobules are formed by 42 43 the paraflocculi, which are divided into the ventral and dorsal paraflocculus (Tan et al., 44 45 1995). In contrast, the petrosal lobules of primates are formed by only a portion of the 46 47 48 paraflocculi (Voogd and Barmack, 2006). While the dorsal and the ventral paraflocculi 49 50 are functionally linked to the petrosal lobules (Voogd et al., 2012), they sit within the 51 52 endocranial cavity adjacent to the flocculus and subarcuate fossa (Xiong et al., 2010). 53 54 55 As we are using endocasts and not neural tissue, any measurements obtained represent 56 57 the potential maximum volume of the tissue occupying the subarcuate fossa of the 58 59 petrosal bone and consequently, the term “petrosal lobule” is more accurate in 60 9 Page 10 of 78 1 2 3 describing what can be measured (Lang et al., 2022). To obtain the neocortical surface 4 5 6 area, the area above the rhinal fissure was selected, whereas the circular fissure and the 7 8 confluence of sinuses were excluded. We followed Jerison (2012) and Long et al. 9 10 (2015), who measured only one side of the neocortex (the more complete hemisphere), 11 12 excluding the superior sagittal sinus, and then doubled the area of the hemisphere. The 13 14 15 resulting data on relative neocortical surface area, relative olfactory bulb and petrosal 16 17 lobule volumes are presented in Table 1. The surface renderings of the endocasts 18 19 described in this paper are available from MorphoSource (Boyer et al., 2014). 20 21 22 23 24 25 We generated a series of bivariate plots as well as boxplots based on the data in 26 27 28 Table 1, combined with values collected from the literature (endocranial volume and 29 30 surface area, olfactory bulb volume, petrosal lobule volume, neocortical surface areas; 31 32 Gingerich & Gunnell, 2005; Silcox et al., 2009, 2010a, 2011; Kirk et al., 2014; Orliac et 33 34 35 al., 2014; Harrington et al., 2016, 2020; Ramdarshan & Orliac, 2016; Bertrand et al., 36 37 2019, 2021; López-Torres et al., 2020; see data in Table 1 and in López-Torres et al., 38 39 2022, tables S1-S4) using the software PAST (Hammer et al., 2001). Because the 40 41 specimen of Anagale gobiensis is the only specimen for which we have data (n = 1), it 42 43 44 is impossible to use statistical hypothesis testing. Instead, we follow a descriptive and 45 46 heuristic approach for our quantitative data. 47 48 49 Body mass estimation 50 51 52 For comparative purposes, we calculated the encephalisation quotient (EQ) using 53 54 equations from Jerison (1973) and Eisenberg (1981). For the calculation of EQs, it is 55 56 57 important to have an accurate estimation of the body mass of the animal in question. 58 59 López-Torres & Fostowicz-Frelik (2018a) reconstructed a body mass for Anagale 60 10 Page 11 of 78 1 2 3 gobiensis of 1756 g based on Conroy’s (1987) prosimian equation using M1 area. They 4 5 6 argued that even though Conroy’s equation uses a prosimian primate sample, this 7 8 equation has been used to estimate body masses in “plesiadapiforms” in the past 9 10 literature (e.g. Conroy, 1987; Rose et al. 1993; Bloch & Gingerich, 1998; Gingerich & 11 12 Gunnell, 2005; Fox et al., 2015; Silcox et al., 2017; Silcox & López-Torres, 2017) and 13 14 15 analagids have a similar generalized morphotype to other primitive euarchontoglirans, 16 17 such as “plesiadapiforms”. Our estimates based on cranial length (305.76 g—using 18 19 Kielan-Jaworowska’s [1983] equation; 444.99 g—using Thewissen & Gingerich’s 20 21 22 [1989] equation; 531.39 g—using Bertrand et al.’s [2016b] equation; 649.31 g—using 23 24 Millien & Bovy’s [2010] equation), are notably much lower than the dental estimate, 25 26 which highlights the challenge in finding the most appropriate equation for body mass 27 28 29 estimation for an extinct animal with no clear modern homologue. 30 31 32 Body masses based on the postcranial elements that actually support the body weight of 33 34 the animal could be argued to be more directly connected to an animal’s mass than 35 36 equations based on cranial or dental measurements (Hylander, 1985; Grabowski et al., 37 38 2015; Yapuncich et al., 2015). However, the fragmentary nature of the postcranial 39 40 41 skeleton of Anagale gobiensis limits the number of equations that could be used. From a 42 43 phylogenetic standpoint, postcranial equations based on samples of rodents (Moncunill- 44 45 Solé et al., 2014) and lagomorphs (Moncunill-Solé et al., 2015) should be relevant, 46 47 48 because Anagale gobiensis has been suggested to be closely related to Glires. From 49 50 Moncunill-Solé et al.’s (2014) rodent equation using the transversal width of the distal 51 52 end of the femur, Anagale gobiensis has an estimated body mass of 1464.68 g. A 53 54 55 lagomorph equation using the transversal width of the proximal end of the tibia 56 57 (Moncunill-Solé et al., 2015) gives a similar result, 1311.26 g. Although somewhat 58 59 lower, a similar value (1122.77 g) is yielded by Tsubamoto’s (2014) mammalian 60 11 Page 12 of 78 1 2 3 equation using trochlear width of the astragalus. Another comparable estimate (1430.89 4 5 6 g) was calculated from an equation based on tibial midshaft diameter of several 7 8 mammalian groups (“Insectivora”, Primates, Lagomorpha, Rodentia, Carnivora, 9 10 Proboscidea, and Artiodactyla) from Gingerich (1990). Because the tibia of Anagale 11 12 gobiensis has a very prominent tibial crest (Simpson, 1931), the tibial midshaft diameter 13 14 15 was measured immediately distal to the end of the tibial crest, where the cross-section 16 17 of the bone shaft is more circular. It is worth noting that similar estimates are obtained 18 19 when using Glires-specific equations (Moncunill-Solé et al., 2015) and more 20 21 22 generalized mammalian equations (Gingerich, 1990; Tsubamoto, 2014). Whereas all the 23 24 mentioned postcranial equations yield body mass estimates over 1 kg, Gingerich (1990) 25 26 also provides an equation based on tibial length (with a lower r2) which gives an 27 28 29 estimated body mass of 520.24 g, making it similar to the cranial length estimates. 30 31 However, long bone circumference has been found to be more strongly related to body 32 33 mass than length (Campione & Evans, 2012), so we consider tibial diameter to likely 34 35 better reflect how the animal distributes the weight through its limbs rather than the 36 37 38 length of the bone (or the length of the cranium); the r2 values from Gingerich (1990) 39 40 support this reasoning. Therefore, we have employed the estimates based on tibial 41 42 midshaft diameter (Gingerich, 1990), proximal tibial width (Moncunill-Solé et al., 43 44 45 2015), distal femoral width (Moncunill-Solé et al., 2014) and astragalar trochlear width 46 47 (Tsubamoto, 2014) in our discussion of relative brain size. Bertrand et al.’s (2016b) 48 49 rodent equation based on cheektooth (P4-M3) length actually provides similar values 50 51 52 (1311.48 g—left toothrow; 1348.56 g—right toothrow) to the mentioned postcranial 53 54 equations. However, Bertrand et al.’s (2016) cheektooth area equation gives much 55 56 higher body mass estimates (2235.59 g—left toothrow; 2374.41 g—right toothrow) than 57 58 the cheektooth length equation. This might be explained by the significantly wider M1 59 60 12 Page 13 of 78 1 2 3 in Anagale gobiensis with respect to the other cheek teeth than typically seen in modern 4 5 6 rodents, making cheektooth length probably more appropriate for estimating body mass 7 8 in Anagale gobiensis. It is also possible that the molar area increases with wear; a 9 10 common effect seen in fossil rodents and lagomorphs with high crowned teeth. The 11 12 teeth of AMNH 26079 are deeply worn, making estimates based on molar area not 13 14 15 completely reliable. On these bases, we consider the best estimate for a range of body 16 17 masses for AMNH 26079 to be 1122.77—1464.68 g. See Table 2 for a detailed list of 18 19 body mass estimates. 20 21 22 Institutional abbreviations 23 24 25 AMNH—American Museum of Natural History, New York, NY; AMNH F:AM—Frick 26 27 28 collection, American Museum of Natural History, New York; UW—University of 29 30 Wyoming, Laramie, WY. 31 32 33 34 35 36 37 38 39 40 41 RESULTS: DESCRIPTION AND COMPARISONS 42 43 44 Olfactory bulbs 45 46 47 Anagale gobiensis has pedunculated olfactory bulbs that are well separated from 48 49 the frontal lobes by a clearly demarcated circular fissure. The bulbs reach their full 50 51 52 length close to the midline (Fig. 3a). The olfactory bulbs account for at least 18.96% of 53 54 the total endocast length (Table 1). That proportion is greater than in early Oligocene 55 56 Ischyromys and sciuroid rodents (15.16-17.85%; Bertrand & Silcox, 2016; Bertrand et 57 58 59 al., 2017), within the range of early and middle Eocene Paramys, Pseudotomus, 60 13 Page 14 of 78 1 2 3 Reithroparamys, and Rapamys (15.58-22.06%; Bertrand & Silcox, 2016; Bertrand et 4 5 6 al., 2019), Eocene euprimates (11.7%-19.9%; calculated from data in Harrington et al., 7 8 2020) and Oligo—Miocene sciuroids (18.62-20.30%; Bertrand et al., 2018), and similar 9 10 to that of Palaeocene—Eocene “plesiadapiforms” (19.39-22.30%; Silcox et al., 2009, 11 12 2010; Orliac et al., 2014). The olfactory bulb length proportion is lower than in the 13 14 15 early Eocene rodent Paramys copei (22.06%; Bertrand et al., 2016a), the early 16 17 Oligocene lagomorph Megalagus turgidus (22.03%; López-Torres et al., 2020) and the 18 19 apatotherian Labidolemur kayi (31%; Silcox et al., 2011). With respect to the width of 20 21 22 the olfactory bulbs, Anagale gobiensis has the highest olfactory bulbs-to-cerebrum 23 24 width ratio (62.47%) compared to fossil rodents, stem lagomorphs, fossil euprimates 25 26 and “plesiadapiforms”, and comparable to those in Labidolemur (62.08%; Silcox et al., 27 28 29 2011). 30 31 32 The cribriform plate is damaged in AMNH 26079 (see Fig. 4a), so an accurate 33 34 reconstruction of the ventral aspect of the olfactory bulbs is impossible. However, for 35 36 the sake of acquiring a better estimate of the olfactory bulb volume, we reconstructed 37 38 the ventral aspect of the bulbs by placing straight lines between preserved bony 39 40 41 landmarks. When olfactory bulb volume is plotted against body mass, Anagale 42 43 gobiensis falls slightly above the fossil rodent morphospace, whereas Labidolemur falls 44 45 much higher than Anagale gobiensis (Fig. 5). The olfactory bulbs of Anagale gobiensis 46 47 48 account for 7.15% of the overall volume of the endocast (Table 1), which should be 49 50 considered a conservative estimate because they are clearly somewhat truncated 51 52 ventrally (Fig. 3c, d). This percentage is higher than in any fossil rodent or lagomorph 53 54 55 (Fig. 6). Among early Tertiary primates, only Plesiadapis cookei Jepsen, 1930 has been 56 57 reported to have a higher relative olfactory bulb volume (7.8%; Orliac et al., 2014) than 58 59 in Anagale gobiensis and that value probably represents an overestimate because the 60 14 Page 15 of 78 1 2 3 ventral aspect of the cerebrum is missing from the reconstruction (see Orliac et al., 4 5 6 2014: fig. S2). The apatotherian Labidolemur also shows a higher ratio (12.18-14.75%; 7 8 Silcox et al., 2011). 9 10 11 The olfactory bulbs in Anagale gobiensis are located above the M1 (Fig. 2), 12 13 similar to what is observed in Ischyromys typus (Bertrand and Silcox, 2016) and Me. 14 15 turgidus (López-Torres et al., 2020). In Cedromus wilsoni the olfactory bulbs are 16 17 18 located above the P4 (Bertrand et al., 2017) and in Paramys spp. above the M2 19 20 (Bertrand et al., 2016a). The location varies amongst primates, with their rostral extent 21 22 being over the molars in fossil euprimates (Harrington et al., 2016, 2020; Ramdarshan 23 24 25 & Orliac, 2016), above the M3 in Microsyops annectens and Plesiadapis tricuspidens, 26 27 but above the premolars in Ignacius graybullianus (Silcox et al., 2009, 2010a; Orliac et 28 29 al., 2014). 30 31 32 33 34 35 Cerebrum and midbrain 36 37 38 The presence of a well demarcated circular fissure, with no coverage of the 39 40 olfactory peduncles by the cerebrum, is consistent with little expansion of the frontal 41 42 43 lobes in Anagale gobiensis rostrally. In fact, the rhinal fissure can be traced rostrally 44 45 and is well separated from the circular fissure, so the anterior edges of the frontal lobes 46 47 of the neocortex are not even the anteriormost structure of the cerebrum, as they usually 48 49 50 are in other fossil euarchontoglirans (Fig. 3a). Uncovered olfactory peduncules also 51 52 occur in “ischyromyids” (Scott & Osborn, 1890; Bertrand and Silcox, 2016; Bertrand et 53 54 al., 2016a, 2019), the pseudosciurid Adelomys vaillanti (Dechaseaux, 1958), the 55 56 57 theridomyid Theridomys bonduelli (Dechaseaux, 1958) and the stem lagomorph Me. 58 59 turgidus (López-Torres et al., 2020). This observation has been made for 60 15 Page 16 of 78 1 2 3 “plesiadapiforms” as well (Silcox et al., 2009; Orliac et al., 2014). Early Eocene 4 5 6 euprimates (Kirk et al., 2014; Harrington et al., 2016, 2020; Ramdarshan & Orliac, 7 8 2016), the basal member of Glires Rh. turpanensis (Meng et al., 2003), the sciurid C. 9 10 wilsoni (Bertrand et al., 2017), the aplodontiid Altasciurus relictus (= Prosciurus 11 12 relictus in Bertrand et al., 2018), the sciurin Protosciurus cf. rachelae (Bertrand et al., 13 14 15 2018) and the apatotherian L. kayi (Silcox et al., 2011) all have a very short circular 16 17 fissure, so the cerebrum is closer to the olfactory bulbs (Fig. 7). 18 19 20 The rhinal fissure marks the separation between the palaeocortex and the 21 22 neocortex and provides information on the degree of development of the latter (Jerison, 23 24 25 2012; Long et al., 2015). The orbitotemporal canal has been used as a proxy landmark 26 27 for the rhinal fissure in some fossil taxa (e.g. Gurche, 1982; Novacek, 1982; Martin, 28 29 1990; Silcox et al., 2009, 2010a, 2011; Bertrand & Silcox, 2016), based on its 30 31 32 relationship to the rhinal fissure in lemurs (Martin, 1990) and at least some rodents 33 34 (Bertrand et al., 2017). When present on the endocast, the orbitotemporal canal is a 35 36 raised structure that runs along its lateral aspect (see Bertrand et al. [2019: fig. 4C-D]). 37 38 An orbitotemporal canal is not observable on the endocast of Anagale gobiensis (Fig. 39 40 41 8); based on the CT data it does not have an intraosseous path either (i.e., it is not 42 43 visible in cross-section in Fig. 4b). In Anagale gobiensis, the rhinal fissure is visible as a 44 45 weak sulcus running longitudinally, but positioned quite far dorsally, on both sides of 46 47 48 the endocast (Fig. 3a, d; Fig. 9). The position of the rhinal fissure in Anagale gobiensis 49 50 is more dorsal than in any other Palaeogene euarchontogliran for which the braincase is 51 52 known (Silcox et al., 2009, 2010a, 2011; Orliac et al., 2014; Bertrand and Silcox, 2016; 53 54 55 Bertrand et al., 2016a, 2017, 2018, 2019; López-Torres et al., 2020), making the 56 57 neocortex very limited in its extension. The neocortical surface area ratio for Anagale 58 59 gobiensis (10.98%) represents the lowest ever recorded in a fossil euarchontogliran 60 16 Page 17 of 78 1 2 3 (Fig. 10), with neocortical surface area ratios in other Palaeogene euarchontoglirans, 4 5 6 ranging between 16.25% (in Paramys delicatus; Bertrand et al., 2019) and 31.71% 7 8 (Protosciurus cf. rachelae; Bertrand et al., 2018, 2019). Although, these values are very 9 10 low, other Eocene mammals appear to have similar neocortical ratios such as the 11 12 taeniodont Ectoganus copei (8%) and the tillodont Trogosus hillsi (10%; see Bertrand et 13 14 15 al., 2022). On the other hand, notably higher values have been calculated for early fossil 16 17 euprimates (e.g. 44-46% for Rooneyia viejaensis; 41-43% for Microchoerus erinaceus; 18 19 Ramdarshan and Orliac, 2016; Harrington et al., 2020), making this group of mammals 20 21 22 the most neocorticalised among primitive euarchontoglirans (Fig. 7; Jerison, 2012; 23 24 Long et al., 2015). 25 26 27 The development of neocortical sulci is related to absolute brain size, with the 28 29 brain generally being lissencephalic (i.e., without infoldings) in species with brain 30 31 32 masses of less than 5 g (Macrini et al., 2007). With an endocast volume of 7798.04 mm3 33 34 and using a brain mass/cranial capacity ratio of 1.05:1 (Jerison, 1973), Anagale 35 36 gobiensis would have had an estimated brain mass of 7.43 g. Therefore, the brain of 37 38 Anagale gobiensis would have been large enough to be expected to exhibit some 39 40 41 infoldings. There are three weakly developed sulci that can be observed (Fig. 3a). One 42 43 is the lateral sulcus (Fig. 9), which runs parallel to the superior sagittal sinus/sulcus, a 44 45 feature generally also found in other fossil euarchontoglirans whose brains would have 46 47 48 weighed more than 5 g (Silcox et al., 2010b, 2011; Orliac et al., 2014; Bertrand et al., 49 50 2016, 2019; Harrington et al., 2016; López-Torres et al., 2020). Another weak sulcus is 51 52 located caudolateral to the frontal lobes, running obliquely caudolaterally, joining the 53 54 55 ventral neocortical edge in the rostral third of the neocortex. This is tentatively 56 57 identified as a presylvian sulcus. There is also a weak, convexly curved sulcus, lateral to 58 59 the lateral sulcus. This is tentatively identified as a suprasylvian sulcus. The last two 60 17 Page 18 of 78 1 2 3 sulci are identified based on their relative position in the endocast, and we acknowledge 4 5 6 that complex sulcal morphology and variation makes it difficult to establish meaningful 7 8 homologies. Finally, there is no Sylvian sulcus in Anagale gobiensis, making it similar 9 10 in this way to other fossil euarchontoglirans that are not euprimates (Harrington et al., 11 12 2016). Interestingly, the sulcal pattern observed in Anagale gobiensis is similar to that 13 14 15 seen in the endocast of the extant macroscelidean Rhynchocyon (Benoit et al., 2013, fig. 16 17 2e). Whereas this could potentially support phylogenetic proximity between anagalids 18 19 and macroscelideans, a hypothesis suggested by McKenna (1975) and Asher et al. 20 21 22 (2005), as noted above, we the balance of evidence to support a closer relationship to 23 24 Euarchontoglires. Alternatively, the similarity in sulcal pattern between Anagale 25 26 gobiensis and Rhynchocyon might reflect evolutionary convergence if anagalids lived in 27 28 29 a similar habitat (i.e. montane and lowland forests and closed-canopy woodlands; 30 31 Kingdon et al., 2013) or had a similar ecology (i.e. digging for food sources; Kingdon et 32 33 al., 2013) to macroscelideans. 34 35 36 Because the edge of the neocortex is located fairly far dorsally, the lateral aspect 37 38 of the endocast is mostly made up of exposed palaeocortex (i.e. piriform cortex, 39 40 41 olfactory cortex). The palaeocortex is inflated, expanding laterally beyond the 42 43 neocortical edge and overhanging ventrally. On the ventral side of the endocast, 44 45 between the olfactory bulbs and the optic chiasm, there is a pair of olfactory tubercles. 46 47 48 They are well-defined and ventrally protruding. 49 50 51 Caudally, the cerebrum does not cover the cerebellum in Anagale gobiensis at all 52 53 (Fig. 3a, Fig. 9). A pair of colliculi is exposed. One hypothesis is that these could be the 54 55 caudal (= inferior) colliculi, with the rostral (= superior) colliculi covered by the 56 57 58 transverse sinuses. Caudal colliculi that are visible on the surface of the endocast are 59 60 also observed in some specimens of Ischyromys (A. E. Wood no. 221 and AMNH 18 Page 19 of 78 1 2 3 F:AM 144638); Reithroparamys delicatissimus; Reithroparamys sciuroides; Rapamys 4 5 6 atramontis; Ad. vaillanti; the cylindrodontid Dolocylindrodon texanus; Plesiadapis 7 8 cookei; Pl. tricuspidens; Microsyops cf. elegans; one specimen of Microsyops 9 10 annectens (UW 14559); Ig. graybullianus; L. kayi, and in the extant sengi Rhynchocyon 11 12 (Wood, 1937; Gingerich & Gunnell, 2005; Silcox et al., 2009, 2010a, 2011, 2020; 13 14 15 Benoit et al., 2013; Orliac et al., 2014; Bertrand & Silcox, 2016; Bertrand et al., 2019). 16 17 Alternatively, it is possible that the visible colliculi in Anagale gobiensis are the rostral 18 19 colliculi and the caudal colliculi are not visible. In other primitive euarchontoglirans 20 21 22 (e.g. Me. turgidus, Paramys, C. wilsoni, Altasciurus aff. saskatchewaensis, one 23 24 specimen of Microsyops annectens; Silcox et al., 2010; Bertrand et al., 2016, 2017, 25 26 2018; López-Torres et al., 2020) the colliculi are not visible, but a patch of midbrain is 27 28 29 apparent between the transverse sinuses and the cerebellum. In Rh. turpanensis the 30 31 midbrain is very broadly exposed (Meng et al., 2003), but the colliculi are not apparent, 32 33 although this may be a product of preservation. This contrasts with the condition of 34 35 Eocene euprimates, in which the midbrain is never exposed (Kirk et al., 2014; 36 37 38 Harrington et al., 2016, 2020; Ramdarshan & Orliac, 2016). 39 40 41 Finally, the hypophyseal fossa (for the pituitary gland) is apparent as a clear, 42 43 nearly circular bulge on the ventral surface of the endocast, approximately in line 44 45 horizontally with the stem of the pathway for cranial nerve V3 (Fig. 3b). 46 47 48 49 50 51 Cerebellum and petrosal lobules 52 53 54 The cerebrum of Anagale gobiensis does not overlap onto the cerebellum, so the 55 56 57 dorsal surface of the cerebellum is broadly exposed on the endocast, exhibiting a vermis 58 59 that is separated from the lateral lobes by paramedian fissures (Fig. 3a, Fig. 9). The 60 19 Page 20 of 78 1 2 3 vermis has two clear fissures running coronally. Rostral to the lobe of the vermis is the 4 5 6 fissura prima (Dow, 1942). In lateral view (Fig. 3d), between the fissura prima and the 7 8 summit of the vermis, there is another sulcus that has not been identified. Caudoventral 9 10 to the summit of the vermis is the prepyramidal fissure (fissura suprapyramidalis in 11 12 Dow, 1942; Fig. 9). This fissure is continuous with the fissura parafloccularis (Larsell 13 14 15 and Dow, 1935; Dow, 1942; Maugoust and Orliac, 2021), which defines the rostral edge 16 17 of the petrosal lobules. No other fissure ventral to the prepyramidal fissure is observable 18 19 (Fig. 9), suggesting there is no visible fissura secunda. Excluding the petrosal lobules, 20 21 22 the cerebrum is clearly wider than the cerebellum. 23 24 25 The petrosal lobules are rounded with very short stems attaching them to the rest 26 27 of the cerebellum (see Figs. 3, 4c; Fig. 9). They are more anteriorly located than the 28 29 lateral lobes and vermis in dorsal view (Fig. 3a). When petrosal lobule volume is plotted 30 31 32 against body mass, Anagale gobiensis appears close to the “ischyromyid” morphospace 33 34 and just above Aplodontia rufa, an extant aplodontiid rodent (Fig. 11). The percentage 35 36 of petrosal lobule volume relative to the volume of the total endocast in Anagale 37 38 gobiensis (1.01%) is within the range of “ischyromyids” (Fig. 12; 0.62-2.05%; Bertrand 39 40 41 & Silcox, 2016; Bertrand et al., 2016a, 2019), but lower than in Me. turgidus (2.31 %; 42 43 López-Torres et al., 2020), Al. relictus (3.35%; Bertrand et al., 2018), C. wilsoni 44 45 (2.96%; Bertrand et al., 2017) and Protosciurus cf. rachelae (3.65-4.82%; Bertrand et 46 47 48 al., 2018). Comparing the percentage of petrosal lobule volume with “plesiadapiforms” 49 50 and apatemyids is not possible, because quantitative measures for the relative size of the 51 52 petrosal lobule have not yet been published for these groups. 53 54 55 56 57 58 59 60 20 Page 21 of 78 1 2 3 Brainstem and cranial nerves 4 5 6 Various casts of the openings for the cranial nerves can be seen on the ventral 7 8 9 surface of AMNH 26079 (Fig. 3b). The cast of the optic chiasm is visible on AMNH 10 11 26079, located just rostral to the level of the sphenorbital fissure. The sphenorbital 12 13 fissure in Anagale gobiensis serves as a passage for the ophthalmic veins and cranial 14 15 nerves III, IV, V1 and VI. The casts of these nerves show a faint contour and based on 16 17 18 the typical branching pattern of cranial nerves, the most medial cranial nerve in the 19 20 sphenorbital fissure would likely be the oculomotor nerve (CN III). Of the two other 21 22 casts, the more medial one may pertain to the trochlear nerve (CN IV), while the largest 23 24 25 and most lateral cast likely contained part of the trigeminal nerve (CN V) and abducens 26 27 nerve (CN VI) (Fig. 3b). Anagale gobiensis lacks a foramen rotundum or any evidence 28 29 of a separate passageway for V2, which means it also presumably passed through the 30 31 32 sphenorbital fissure, as observed in L. kayi (Silcox et al., 2011). This contrasts with the 33 34 situation in “ischyromyids”, C. wilsoni, and extant squirrels, in which there is a distinct 35 36 passageway for V2 on the endocast even though it ultimately does lead to the 37 38 sphenorbital fissure rather than to a distinct foramen rotundum (Bertrand et al., 2017; 39 40 41 2019). In Ap. Rufa, the sphenorbital fissure is not confluent with the foramen rotundum, 42 43 whereas in fossil aplodontiids, they are merged with each other (Bertrand et al., 2018). 44 45 With respect to stem lagomorphs, there is no separate foramen rotundum in Palaeolagus 46 47 48 haydeni (Wolniewicz and Fostowicz-Frelik, 2021). The foramen rotundum is variable in 49 50 its presence in primates; there is a separate passageway for V2 leading to a distinct 51 52 opening on the endocast of adapoids (Harrington et al., 2016) and Microsyops 53 54 55 annectens (Silcox et al., 2010), but not Ig. graybullianus or Microchoerus erinaceus 56 57 (Silcox et al., 2009; Ramdarshan & Orliac, 2016). The condition in Plesiadapis is a 58 59 matter of continuing debate (Bloch & Silcox, 2006; Orliac et al., 2014). Gurche (1978) 60 21 Page 22 of 78 1 2 3 considered the presence of a distinct foramen rotundum to be variable in Necrolemur 4 5 6 antiquus (see discussion in Harrington et al., 2020), which highlights the lability of this 7 8 character. 9 10 11 AMNH 26079 has a foramen ovale (Simpson, 1931) as is typical of mammals, 12 13 but unlike some rodents lacks a separate foramen ovale (Wahlert, 1974; Bertrand et al., 14 15 2018: Table S14). Casts of the internal auditory meatus, with passageways for cranial 16 17 18 nerves VII (facial) and VIII (vestibulocochlear), are visible on both sides located 19 20 rostroventral to the petrosal lobules (Fig. 3b-d). A cast of the jugular foramen, through 21 22 which the internal jugular vein and cranial nerves IX (glossopharyngeal), X (vagus) and 23 24 25 XI (accessory) would have run, is positioned directly ventral to the petrosal lobule (Fig. 26 27 3b, c; Fig. 9), in a similar position to the one seen in fossil rodents (Bertrand & Silcox, 28 29 2016; Bertrand et al., 2016a, 2017, 2018, 2019), early euprimates (Harrington et al., 30 31 32 2016, 2020; Ramdarshan & Orliac, 2016), “plesiadapiforms” (Silcox et al., 2009, 33 34 2010a; Orliac et al., 2014) and Labidolemur (Silcox et al., 2011). In Me. turgidus, the 35 36 jugular foramen is shifted more rostrally (López-Torres et al., 2020). Finally, there is a 37 38 cast of a single hypoglossal foramen for cranial nerve XII (hypoglossal) on each side of 39 40 41 the brainstem (Fig. 3b; Fig. 9), as seen in some “ischyromyids” (e.g. Paramys copei; 42 43 Pa. delicatus; Notoparamys costilloi; Pseudotomus horribilis; Pseudotomus petersoni; 44 45 Ra. atramontis; Re. delicatissimus; Re. sciuroides; Bertrand et al., 2016, 2019), C. 46 47 48 wilsoni (Bertrand et al., 2017); Al. relictus, Protosciurus cf. rachelae (Bertrand et al., 49 50 2018); stem lagomorphs (Me. turgidus and P. haydeni; López-Torres et al., 2020; 51 52 Wolniewicz and Fostowicz-Frelik, 2021); early euprimates (Harrington et al., 2016, 53 54 55 2020; Ramdarshan and Orliac, 2016), “plesiadapiforms” (Bloch and Silcox, 2006; 56 57 Silcox et al., 2009, 2010a; Orliac et al., 2014) and L. kayi (Silcox et al., 2011). 58 59 However, casts for two hypoglossal nerves (per side) are observed in some 60 22 Page 23 of 78 1 2 3 “ischyromyids” (e.g. Pseudotomus robustus [= Pseudotomus oweni in Bertrand et al., 4 5 6 2019], Is. Typus; Wahlert, 1974; Bertrand and Silcox, 2016; Bertrand et al., 2019). 7 8 9 10 11 12 Blood vessels 13 14 Anagale gobiensis has a pattern of venous sinuses on the endocast that is 15 16 17 basically similar to the arrangement inferred as primitive for therian mammals (Wible & 18 19 Rougier, 2004). The superior sagittal sinus is most prominent around the level of the 20 21 cerebral frontal lobes and fades away more caudally. This suggests that this sinus would 22 23 24 have partially been located deep in the meninges (Macrini et al., 2007). The superior 25 26 sagittal sinus is continuous with the transverse sinus. In Anagale gobiensis the 27 28 transverse sinus connects with the sigmoid sinus, which is continuous with the internal 29 30 31 jugular vein as observed for rodents (Fig. 9; Bertrand & Silcox, 2016; Bertrand et al., 32 33 2016a, 2017, 2018, 2019), early euprimates (Harrington et al., 2016, 2020; Ramdarshan 34 35 & Orliac, 2016), “plesiadapiforms” (Silcox et al., 2009, 2010a, 2020) and apatemyids 36 37 (Silcox et al., 2011). The jugular foramen is continuous with the cast of the inferior 38 39 40 petrosal sinus in Anagale gobiensis. This is also observed in early euprimates (Kirk et 41 42 al., 2014; Harrington et al., 2016, 2020; Ramdarshan & Orliac, 2016), fossil rodents, 43 44 Me. turgidus and Microsyops annectens (Silcox et al., 2010a; Bertrand & Silcox, 2016; 45 46 47 Bertrand et al., 2017, 2018, 2019; López-Torres et al., 2020). The inferior petrosal sinus 48 49 is not well marked in Ig. graybullianus (Silcox et al., 2009) and it is not preserved in Pl. 50 51 tricuspidens or L. kayi (Silcox et al., 2011; Orliac et al., 2014). There is a large 52 53 54 postglenoid foramen present (Simpson, 1931). However, the cast of the postglenoid 55 56 vein was not traced when reconstructing the endocast, because the vein would have 57 58 been very large and would have significantly affected the volume and surface area 59 60 23 Page 24 of 78 1 2 3 calculations. The orbitotemporal canal is not observable on the endocast, and as noted 4 5 6 above did not run intraosseously. 7 8 9 With respect to arteries, a small cast on the surface of the endocast lateral to the 10 11 pituitary gland is here identified as the cast for the carotid foramen (Fig. 3b), but it is 12 13 not clearly related to a pathway for a vessel on the endocast which makes interpreting 14 15 its contents somewhat ambiguous. Lateral to the carotid foramen, there is a larger cast 16 17 18 that corresponds to the piriform fenestra (posterior lacerate foramen of Simpson, 1931; 19 20 pyriform fenestra of McDowell, 1958). The piriform fenestra transmits various 21 22 neurovascular structures, including the postglenoid vein (Wible and Shelley, 2020). The 23 24 25 alisphenoid canal, which transmits the ramus infraorbitalis of the stapedial artery 26 27 (Wible, 1987), is visible and merges with the sphenorbital fissure (Fig. 3b-d). As Wible 28 29 (1987) discussed, there is a significant amount of variability among mammals in the 30 31 32 presence or absence of an alisphenoid canal, suggesting that it is a feature that arose 33 34 multiple times (see also Asher, 2001). Within Euarchontoglires it is present in 35 36 lagomorphs (Wible, 1987; Wolniewicz and Fostowicz-Frelik, 2021) and scandentians 37 38 (Wible, 1987), but absent in dermopterans (associated with the complete involution of 39 40 41 the internal carotid artery; Wible, 1993). In rodents, it is variably present (Wible, 1987), 42 43 but has been identified in the most primitive forms (Wahlert, 1974; Bertrand & Silcox, 44 45 2016). Although missing in “plesiadapiforms” (Silcox et al., 2009, 2010; Orliac et al., 46 47 48 2014), a clear alisphenoid canal is present in Labidolemur (Silcox et al., 2011) and has 49 50 been identified in the early euprimate Microchoerus erinaceus (Ramdarshan & Orliac, 51 52 2016). These various lines of evidence suggest that Anagale’s condition (presence of the 53 54 55 canal) is likely primitive for Glires, and possibly for Euarchontoglires. 56 57 58 59 60 Relative brain size and encephalisation quotient 24 Page 25 of 78 1 2 3 Figure 13 shows a bivariate plot of logarithmic endocranial volume against 4 5 6 logarithmic body mass. The range of body masses for Anagale gobiensis places this 7 8 species within the shared morphospace between extinct euprimates and extinct rodents. 9 10 11 The EQ estimates of Anagale gobiensis are calculated based on the range of body 12 13 mass estimates given above (Fig. 14). Anagale gobiensis has a range of EQs (Eisenberg, 14 15 1981) of 0.61—0.74 (Table 3). Compared to other early members of Euarchontoglires 16 17 18 (Fig. 14), Anagale gobiensis has EQ values within the range of “ischyromyids” (0.36— 19 20 0.76; Bertrand and Silcox, 2016; Bertrand et al., 2016a, 2019), Oligocene sciuroids 21 22 (0.99—1.33; Bertrand et al., 2017, 2018) and Eocene euprimates (0.37—1.53; Kirk et 23 24 25 al., 2014; Harrington et al., 2016, 2020; Ramdarshan & Orliac, 2016). Anagale 26 27 gobiensis has higher EQ values than Palaeocene and early Eocene “plesiadapiforms” 28 29 (0.14—0.61; Silcox et al., 2009, 2010a; Orliac et al., 2014; Long et al., 2015), the 30 31 32 Oligocene stem lagomorph Megalagus (0.39; López-Torres et al., 2020) and the late 33 34 Palaeocene—early Eocene apatemyid Labidolemur (0.36; Silcox et al., 2011). 35 36 However, Anagale gobiensis has a lower EQ than Oligocene sciuroids (0.99—1.33; 37 38 Bertrand et al., 2017, 2018) and the late Eocene apatemyid Carcinella (1.28; von 39 40 41 Koenigswald et al., 2009). 42 43 44 45 46 47 DISCUSSION 48 49 The EQ results and the plot of endocranial volume and body mass show that 50 51 52 Anagale gobiensis had a level of encephalisation similar to that of “ischyromyids” and 53 54 Eocene euprimates (Figs. 12 and 13). However, the endocast of Anagale gobiensis 55 56 differs considerably in the proportions of the subregions from the endocasts of fossil 57 58 59 euprimates in having much better developed regions related to olfaction (such as the 60 25 Page 26 of 78 1 2 3 palaeocortex, olfactory bulbs and olfactory tubercles), but a much smaller neocortex 4 5 6 (Fig. 7). Indeed, probably the most surprising feature of the endocranial morphology of 7 8 Anagale gobiensis is its extremely low degree of neocorticalisation, which is 9 10 particularly surprising in light of its late Eocene age and the trend of increasing 11 12 neocorticalisation in both Glires and Primates. At 10.98% of neocortical surface area 13 14 15 relative to the total endocranial surface area, Anagale gobiensis is the least 16 17 neocorticalised euarchontogliran known to date, although there are other Eocene 18 19 mammals that do also exhibit a low neocortical surface ratio (Fig. 8; see Bertrand et al., 20 21 22 2022). The neocortex in Anagale gobiensis is almost entirely restricted to the dorsal 23 24 face of the endocast, with very little ventral projection on the sides. The neocortex 25 26 encompasses regions responsible for the primary processing of visual, auditory, 27 28 29 somatosensory, motor, sensorimotor and prefrontal sensory information (Martin, 1990; 30 31 Kaas, 2012; Long et al., 2015), and for these reasons may be impacted based on aspects 32 33 of a species’ ecology, such as its lifestyle. The locomotor behaviour of Anagale 34 35 gobiensis is largely unknown. The high degree of dental wear, tendency towards 36 37 38 hypsodonty and the inference of adaptations for forelimb digging (McKenna, 1963) 39 40 could be interpreted as evidence that anagalids sourced their food from underground. A 41 42 smaller neocortex has been suggested to be linked to fossorial lifestyle (Bertrand et al., 43 44 45 2018; 2021), likely because of reduced visual requirements, so perhaps this low degree 46 47 of neocorticalisation could be an indication that Anagale gobiensis was actually 48 49 fossorial. This inferred pattern of behaviour could also help to explain the similarities 50 51 52 in the form of the endocast to the macroscelidean Rhynchocyon (Benoit et al., 2013). 53 54 However, this idea requires further testing based on assessment of the postcranial 55 56 material. 57 58 59 60 26 Page 27 of 78 1 2 3 Anagale gobiensis has complete exposure of one set of colliculi, tentatively 4 5 6 identified as the caudal colliculi (see discussion in the section ‘Cerebrum and midbrain’ 7 8 above). Whereas this is often seen as a primitive trait, it has been argued that an 9 10 expansion of this region could be associated with increased auditory acuity (Edinger, 11 12 1964). This may be critical in semi-fossorial species to avoid predators or when 13 14 15 visibility is reduced in subterranean environments (Begall et al., 2007). 16 17 18 Compared to other fossil euarchontoglirans, Anagale gobiensis has a proportionally 19 20 large volume of the brain dedicated to olfaction. The olfactory bulb volume ratio in 21 22 Anagale gobiensis is higher than in any fossil member of Glires and within the upper 23 24 25 range for plesiadapiforms (Fig. 6). Only apatemyids have proportionately much larger 26 27 olfactory bulbs. This pattern is also seen in the analysis relative to body mass (Fig. 5), in 28 29 which Labidolemur and Anagale gobiensis both sit above the range of variation for 30 31 32 other euarchontoglirans. Therefore, Anagale gobiensis was likely an animal that was 33 34 largely olfaction driven. This is also consistent with A. gobiensis having a laterally and 35 36 ventrally inflated palaeocortex (also known as olfactory cortex) and well-defined, 37 38 ventrally protruding olfactory tubercles; a greater emphasis in olfaction is supported by 39 40 41 larger palaeocortical structures (Barton et al., 1995). 42 43 44 The petrosal lobule volume ratio in Anagale gobiensis is much lower than that 45 46 observed in Megalagus and modern lagomorphs, within the rodent range (Fig. 12). 47 48 When considered in relation to body mass Anagale gobiensis is just below 49 50 51 “ischyromyid” morphospace, near the aplodontiid rodent Ap. rufa (Fig. 11). The 52 53 petrosal lobules play a role in the stabilisation of eye movements necessary for accurate 54 55 visual tracking (Rambold et al., 2002; Hiramatsu et al., 2008), suggesting that Anagale 56 57 58 gobiensis was probably living in an environment in which quick visual tracking was not 59 60 a selective pressure. Following McKenna’s (1963) observation that Anagale gobiensis 27 Page 28 of 78 1 2 3 was probably a forelimb scratching digger, this species might have spent some time in 4 5 6 burrows, where there is a lack of visual cues, or might have had a relatively slow mode 7 8 of locomotion in which quick visual tracking was not critical for survival. Bertrand et 9 10 al. (2018; 2021) deduced that a low petrosal lobule ratio might be linked to fossoriality 11 12 in aplodontiid rodents, which is consistent with the idea that Anagale gobiensis might 13 14 15 have been a digger. Additionally, Lang et al. (2022) found that fossorial taxa have 16 17 significantly smaller petrosal lobules relative to body mass compared to other locomotor 18 19 groups across Euarchontoglires. 20 21 22 As a potential outgroup to Glires and a lineage branching off a deep node in 23 24 25 Euarchontoglires, Anagale gobiensis has the potential to be crucially informative to the 26 27 brain evolution of these clades, with the proviso that its own patterns of behaviour may 28 29 have shaped certain aspects of its brain anatomy, particularly if it was fossorial. Anagale 30 31 32 gobiensis shares features with other primitive groups in the lineages of Rodentia, 33 34 Lagomorpha and Primates, which may therefore be relevant for reconstructing ancestral 35 36 stages in Euarchontoglires. As observed in all these groups, Anagale gobiensis has a 37 38 partially exposed midbrain and a relatively low EQ. However, it is not entirely clear if 39 40 41 these traits are symplesiomorphies inherited from the euarchontogliran ancestor or if 42 43 Anagale gobiensis has gone through a secondary reversal of these traits due to fossorial 44 45 adaptations. Anagale gobiensis has a higher relative volume of the olfactory bulbs than 46 47 48 in fossil members of Glires, or “plesiadapiforms”, but smaller than in Labidolemur; this 49 50 could suggest that large olfactory bulbs may be ancestral for Euarchontoglires but could 51 52 also reflect adaptations in Apatemyidae and Anagalidae. Anagale gobiensis also 53 54 55 displays small petrosal lobules, comparable proportionally to those of modern and 56 57 extinct fossorial rodents (see Bertrand et al., 2021; Lang et al., 2022), suggesting that 58 59 the reduction in petrosal lobule size in Anagale gobiensis might reflect some degree of 60 28 Page 29 of 78 1 2 3 fossorial behaviour. As the petrosal lobule (and the flocculus) has been classically 4 5 6 interpreted as being used for gaze stabilisation (Rambold et al., 2002; Hiramatsu et al., 7 8 2008), it is expected that animals that live in subterranean or low-light environments 9 10 would have small floccular fossae. There is still some uncertainty in determining 11 12 whether Glires ancestrally had small or large petrosal lobules, since “ischyromyids” are 13 14 15 fairly variable in this character (Bertrand et al., 2019). Based on a recent study, the 16 17 petrosal lobules were reconstructed as being small at the base of Euarchontoglires and 18 19 then increasing in proportion through time (Bertrand et al., 2021). As a late Eocene 20 21 22 taxon, this would lead one to expect that the petrosal lobule relative volume would be 23 24 higher than the value reconstructed for primitive Rodentia (there is insufficient data to 25 26 reconstruct the primitive lagomorph node) but is actually lower, consistent with the 27 28 29 inference about fossoriality. Finally, Anagale gobiensis is uniquely less neocorticalised 30 31 compared to other euarchontoglirans of similar age and even compared to Palaeocene 32 33 euarchontoglirans (Fig. 7). This suggests that Anagale gobiensis may have secondarily 34 35 decreased its neocortical area due to fossoriality or other ecological aspects and 36 37 38 therefore the ancestor of Glires and Euarchontoglires might have been more 39 40 neocorticalised than Anagale gobiensis. Low neocorticalisation is likely primitive for 41 42 Euarchontoglires as the Eocene “ischyromyids” and “plesiadapiforms” have less 43 44 45 expanded neocortices than more derived rodents and primates. 46 47 48 49 50 CONCLUSION 51 52 53 The present study reconstructs the first virtual endocast of an anagalid, a poorly 54 55 56 known group of Palaeogene euarchontoglirans from Asia, argued to be closely related to 57 58 modern members of Glires. The endocast of Anagale gobiensis sheds some new light on 59 60 the reconstruction of the palaeoecology and behaviour of the animal, which has 29 Page 30 of 78 1 2 3 remained largely unknown. Anagale gobiensis displays relatively small petrosal lobules, 4 5 6 and the lowest neocortical ratio recorded for a euarchontogliran. These traits might be 7 8 expected if anagalids were fossorial, which would be consistent with indications that 9 10 they may have sourced their food from below ground (McKenna, 1963). The evidence 11 12 from the endocast shows that Anagale gobiensis was likely very olfaction driven, based 13 14 15 on the relatively large olfactory bulbs and olfactory tubercles, and a laterally expansive 16 17 palaeocortex (= olfactory cortex; piriform cortex). This suite of traits means that 18 19 Anagale gobiensis does not follow the trend in other euarchontogliran lineages of 20 21 22 having significantly increased their degree of neocorticalisation by the end of the 23 24 Eocene. The expansion of the palaeocortex in Anagale gobiensis represents a different 25 26 pattern than seen in other fossil members of Euarchontoglires. 27 28 29 Despite its likely specialisation for fossoriality, Anagale gobiensis pertains to a very 30 31 32 primitive lineage of euarchontoglirans and its endocast may be informative with respect 33 34 to some aspects of the earliest stages of brain evolution in Euarchontoglires. The 35 36 endocast further supports relatively large olfactory bulbs, partial midbrain exposure, and 37 38 a relatively low EQ as ancestral traits for Euarchontoglires. The petrosal lobule volume 39 40 41 ratio falls within the fairly broad range of “ischyromyid” rodents, which is consistent 42 43 with previous studies showing that an increase in petrosal lobules occurred through time 44 45 in Euarchontoglires including rodents (Bertrand et al., 2022). However, the low ratio of 46 47 48 the petrosal lobules of Anagale gobiensis could be related to an adaptation to 49 50 fossoriality as previously hypothesised for rodents (Bertrand et al., 2021). Ultimately, 51 52 the relative size of the petrosal lobules has not been explored in other key early 53 54 55 euarchontoglirans, such as “plesiadapiforms” or apatemyids, which would be 56 57 fundamental to constrain what the ancestral state would be. 58 59 60 30 Page 31 of 78 1 2 3 The palaeoecological diversity of anagalids is currently unknown. Future work on 4 5 6 other known anagalid crania, such as those of Anagalopsis kansuensis (Bohlin, 1951), 7 8 Eosigale gujingensis (Hu, 1993), and Linnania lofoensis (Chow et al., 1973), will help 9 10 augment our understanding of the behaviour and endocranial morphology of this family. 11 12 Additionally, an in-depth study of the postcranial skeleton of Anagale gobiensis would 13 14 15 also be key to have the full picture of the locomotor ecology of this species and provide 16 17 further insight into this enigmatic group of mammals. 18 19 20 21 22 23 Author contributions. SLT, OCB, MTS, ŁFF and JM contributed to the conception and 24 25 design of the study. OCB and JM acquired the CT data, SLT and MML segmented out 26 27 28 the virtual specimen and collected the data. All authors interpreted the data. SLT drafted 29 30 the article. All authors revised it critically for important intellectual content and gave 31 32 final approval before submission. 33 34 35 36 37 38 Acknowledgements. The authors would like to thank Mao Fangyuan (Institute of 39 40 Vertebrate Paleontology and Paleoanthropology) for facilitating access to data. We also 41 42 43 thank Morgan Hill from the Microscopy and Imaging Facility (MIF) at the AMNH for 44 45 facilitating the scanning of specimens. We are grateful to Rémi Allemand and Gabriela 46 47 San Martin-Flores for help during the segmentation process, to Keegan Selig for 3D 48 49 50 printing of the endocast, and Peggy Brady for relevant discussions on the cranial 51 52 anatomy of Anagale gobiensis. We would also like to thank Robert Asher, Sally 53 54 Thomas, and three anonymous reviewers for helpful comments that significantly 55 56 57 improved this paper. This study was supported by a Kalbfleisch Postdoctoral Research 58 59 Fellowship to SLT, a Marie Skłodowska-Curie Actions: Individual Fellowship (H2020- 60 31 Page 32 of 78 1 2 3 MSCA-IF-2018-2020; No. 792611) to OCB, an NSERC CGS grant to MML, and an 4 5 6 NSERC Discovery Grant to MTS. The authors have no conflicts of interest to declare. 7 8 9 10 11 12 DATA AVAILABILITY 13 14 A 3D surface model of the endocast of Anagale gobiensis (AMNH 26079) is available 15 16 in MorphoSource: http://morphosource.org/permalink/?PXXX [login username: 17 18 19 anagale.reviewer@gmail.com; password: 12345]. Other data for this study are available 20 21 in the Dryad Digital Repository: https://datadryad.org/stash/share/XXXX [please note 22 23 that the data for this paper are not yet published and these temporary links should not be 24 25 26 shared without the express permission of the author] 27 28 29 30 31 32 REFERENCES 33 34 35 ASHER, R. J. 2001. Cranial anatomy in tenrecid insectivorans: Character evolution 36 37 across competing phylogenies. American Museum Novitates, 3352, 1-53. 38 39 40 ASHER, R. J. 2007. A web-database of mammalian morphology and a reanalysis of 41 42 placental phylogeny. BMC Evolutionary Biology, 7, 108. 43 44 45 ASHER, R. J. and HELGEN, K. M. 2010. Nomenclature and placental mammal 46 47 48 phylogeny. BMC Evolutionary Biology, 10, 102. 49 50 51 ASHER, R. J., MENG, J., WIBLE, J. R., MCKENNA, M. C., ROUGIER, G. W., 52 53 DASHZEVEG, D. and NOVACEK, M. J. 2005. Stem Lagomorpha and the 54 55 antiquity of Glires. 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The Tertiary chronostratigraphic researches and 4 5 6 Tertiary chronostratigraphic chart of China. Journal of Stratigraphy, 24, 120– 7 8 125. 9 10 11 ZHANG, Z. Q. and WANG, J. 2016. On the geological age of mammalian fossils from 12 13 Shanmacheng, Gansu Province. Vertebrata PalAsiatica 54: 351–357. 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 49 Page 50 of 78 1 2 3 FIGURE CAPTIONS 4 5 6 Figure 1. Phylogenetic hypothesis of euarchontogliran relationships. Based on Korth & 7 8 9 Emry (1991), Meng et al. (2003) and Silcox et al. (2010b); modified. Only groups 10 11 represented by fossil endocasts discussed in the manuscript are included. Scandentia 12 13 and Dermoptera are excluded because they are unknown from fossil skulls. 14 15 16 17 18 19 Figure 2. Transparent cranium with endocast of Anagale gobiensis (AMNH 26079) 20 21 from the Ulan Gochu Formation at Twin Oboes, Nei Mongol, China. (a) dorsal, (b) left 22 23 24 lateral, (c) right lateral and (d) ventral views. Scale bar represents: 1 cm. 25 26 27 28 29 30 Figure 3. Virtual endocast of Anagale gobiensis (AMNH 26079) in (a) dorsal, (b) 31 32 ventral, (c) right lateral and (d) left lateral views. Scale bar represents: 5 mm. 33 34 35 36 37 38 Figure 4. Coronal slices of the cranium of Anagale gobiensis (AMNH 26079) at the 39 40 level of the a) olfactory bulbs, b) main body of the cerebrum, and c) the petrosal 41 42 lobules. Scale bar represents: 1 cm. 43 44 45 46 47 48 Figure 5. Bivariate plot of logarithmic olfactory bulb volume against logarithmic body 49 50 51 mass for fossil Euarchontoglires. BM = Body mass. Data are given in Table S1. 52 53 54 55 56 57 Figure 6. Boxplot of olfactory bulb volume ratio of fossil Euprimates, Plesiadapiformes, 58 59 fossil Rodentia, extant Rodentia, stem Lagomorpha (represented by Megalagus 60 50 Page 51 of 78 1 2 3 turgidus), extant Lagomorpha, Apatemyidae (represented by Labidolemur kayi) and 4 5 6 Anagale gobiensis. Data are given in Table S4. 7 8 9 10 11 12 Figure 7. Endocast morphology and relationships for Euarchontoglires. The topology of 13 14 the tree is based on Fig. 1. 15 16 17 18 19 20 Figure 8. Interior of the cranium of Anagale gobiensis (AMNH 26079) sliced in the 21 22 sagittal plane. Scale bar represents: 1 cm. 23 24 25 26 27 28 Figure 9. Virtual endocast of Anagale gobiensis (AMNH 26079) in oblique caudal 29 30 view. Scale bar represents: 5 mm. 31 32 33 34 35 36 Figure 10. Boxplot of neocortical ratio of fossil Euprimates, plesiadapiforms, fossil 37 38 39 Rodentia, extant Sciuroidea, stem Lagomorpha (represented by Megalagus turgidus), 40 41 extant Lagomorpha and Anagale gobiensis. Data are given in Table S4. 42 43 44 45 46 47 Figure 11. Bivariate plot of logarithmic petrosal lobule volume against logarithmic body 48 49 mass for selected Euarchontoglires. BM = Body mass. Data are given in Table S2. 50 51 52 53 54 55 Figure 12. Boxplot of petrosal lobule volume ratio of fossil Euprimates, Ischyromyidae, 56 57 fossil Aplodontiidae (represented by Mesogaulus paniensis [*] and Altasciurus relictus 58 59 60 [**]), extant Aplodontiidae (represented by the only living species, Aplodontia rufa), 51 Page 52 of 78 1 2 3 fossil Sciuridae, extant Sciuridae, stem Lagomorpha (represented by Megalagus 4 5 6 turgidus), extant Lagomorpha and Anagale gobiensis. Data are given in Table S4. 7 8 9 10 11 12 Figure 13. Bivariate plot of logarithmic endocranial volume against logarithmic body 13 14 mass for fossil Euarchontoglires. BM = Body mass. Data are given in Table S1. 15 16 17 18 19 20 Figure 14. Encephalisation quotients (EQs) for Anagale gobiensis (AMNH 26079) 21 22 compared to fossil Euarchontoglires. EQs based on Eisenberg (1981). Abbreviations: 23 24 BoM↑, high body mass estimate used; BoM↓, low body mass estimate used; BrM↑, 25 26 27 high brain mass estimate used; BrM↓, low brain mass estimate used. Data are given in 28 29 Table S3. 30 31 32 33 34 35 TABLE CAPTIONS 36 37 38 Table 1. Quantitative measurements of the endocast of Anagale gobiensis (AMNH 39 40 26079). Calculations of EQ based on an estimated body mass of Anagale gobiensis of 41 42 1122.77—1464.68 g (see Tables 2 and 3). The total endocranial volume (TV) was 43 44 45 calculated by doubling the volume of the more complete left side. The neocortical 46 47 surface area (NS) was measured by doubling the area of the neocortex for the more 48 49 complete left hemisphere. 50 51 52 53 54 55 Table 2. Body mass estimations for Anagale gobiensis, AMNH 26079. The body mass 56 57 58 estimates indicated with an asterisk are argued to be the most likely to be correct (see 59 60 discussion in the text). 52 Page 53 of 78 1 2 3 4 5 6 Table 3. Calculation of encephalisation quotients (EQ) for Anagale gobiensis, AMNH 7 8 9 26079. For details on the body mass equations see Table 2. 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 53 Page 54 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Figure 1. Phylogenetic hypothesis of euarchontogliran relationships. Based on Korth & Emry (1991), Meng et 26 al. (2003) and Silcox et al. (2010b); modified. Only groups represented by fossil endocasts discussed in the 27 manuscript are included. Scandentia and Dermoptera are excluded because they are unknown from fossil skulls. 28 29 338x190mm (300 x 300 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 55 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Figure 2. Translucid cranium with endocast of Anagale gobiensis (AMNH 26079) from the Ulan Gochu 26 Formation at Twin Oboes, Nei Mongol, China. (a) dorsal, (b) left lateral, (c) right lateral and (d) ventral 27 views. Scale bar represents: 1 cm. 28 338x190mm (600 x 600 DPI) 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 56 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 Figure 3. Virtual endocast of Anagale gobiensis (AMNH 26079) in (a) dorsal, (b) ventral, (c) right lateral and 43 (d) left lateral views. Scale bar represents: 5 mm. 44 45 165x186mm (600 x 600 DPI) 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 57 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 Figure 4. Coronal slices of the cranium of Anagale gobiensis (AMNH 26079) at the level of the a) olfactory 29 bulbs, b) main body of the cerebrum, and c) the petrosal lobules. Scale bar represents: 1 cm. 30 165x108mm (600 x 600 DPI) 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 58 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Figure 5. Bivariate plot of logarithmic olfactory bulb volume against logarithmic body mass for fossil 27 Euarchontoglires. BM = Body mass. Data are given in Table S1. 28 29 165x101mm (600 x 600 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 59 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Figure 6. Boxplot of olfactory bulb volume ratio of fossil Euprimates, Plesiadapiformes, fossil Rodentia, 27 extant Rodentia, stem Lagomorpha (represented by Megalagus turgidus), extant Lagomorpha, Apatemyidae 28 (represented by Labidolemur kayi) and Anagale gobiensis. Data are given in Table S4. 29 165x98mm (600 x 600 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 60 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Figure 7. Endocast morphology and relationships for Euarchontoglires. The topology of the tree is based on 28 Fig. 1. 29 165x103mm (600 x 600 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 61 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 Figure 8. Interior of the cranium of Anagale gobiensis (AMNH 26079) sliced in the sagittal plane. Scale bar 40 represents: 1 cm. 41 42 165x170mm (600 x 600 DPI) 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 62 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 Figure 9. Virtual endocast of Anagale gobiensis (AMNH 26079) in oblique caudal view. Scale bar represents: 31 5 mm. 32 33 200x147mm (300 x 300 DPI) 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 63 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Figure 10. Boxplot of neocortical ratio of fossil Euprimates, plesiadapiforms, fossil Rodentia, extant 26 Sciuroidea, stem Lagomorpha (represented by Megalagus turgidus), extant Lagomorpha and Anagale 27 gobiensis. Data are given in Table S4. 28 165x93mm (600 x 600 DPI) 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 64 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Figure 11. Bivariate plot of logarithmic petrosal lobule volume against logarithmic body mass for selected 28 Euarchontoglires. BM = Body mass. Data are given in Table S2. 29 203x127mm (300 x 300 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 65 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Figure 12. Boxplot of petrosal lobule volume ratio of fossil Euprimates, Ischyromyidae, fossil Aplodontiidae 25 (represented by Mesogaulus paniensis [*] and Altasciurus relictus [**]), extant Aplodontiidae (represented 26 by the only living species, Aplodontia rufa), fossil Sciuridae, extant Sciuridae, stem Lagomorpha 27 (represented by Megalagus turgidus), extant Lagomorpha and Anagale gobiensis. Data are given in Table S4. 28 29 165x88mm (300 x 300 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 66 of 78 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 Figure 13. Bivariate plot of logarithmic endocranial volume against logarithmic body mass for fossil 28 Euarchontoglires. BM = Body mass. Data are given in Table S1. 29 203x127mm (600 x 600 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 67 of 78 1 2 3 Table 1. 4 5 Measurement (abbreviation); values in mm 6 Total length (TL) 41.14 7 Olfactory bulb length (OL) 7.80 8 Olfactory bulb width (OW) 15.38 9 10 Olfactory bulb height (OH) 8.42 11 Maximum neocortex height (NMH) 5.17 12 Cerebrum total length (CRML) 19.92 13 Cerebrum maximal width (CRMW) 24.62 14 Cerebrum maximal height (CRMH) 17.38 15 Cerebellum length (vermis) (CLML) 12.35 16 Cerebellum width (without petrosal lobules) (CLW) 17.42 17 18 Length ratios; values in % 19 OL/TL 18.96 20 CRML/TL 48.42 21 CLML/TL 30.02 22 CLW/CRMW 70.76 23 OW/CRMW 62.47 24 OW/CLW 88.29 25 26 NMH/CRMH 32.07 27 Surfaces (abbreviation); values in mm 2 28 Total endocast area (TS) 3827.58 29 Neocortical surface area (NS) 420.24 30 Neocortical surface area (one side) (NS1) 210.12 31 Volumes (abbreviation); values in mm3 32 Total endocast (TV) 7798.04 33 34 Olfactory bulbs (right and left sides; OV) 557.78 35 Petrosal lobules (right and left sides; PLV) 79.11 36 Surface and volume ratios; values in % 37 NS/TS 10.98 38 OV/TV 7.15 39 PLV/TV 1.01 40 Mass; values in mg 41 42 Olfactory bulb mass 531.22 43 Petrosal lobule mass 75.34 44 Encephalization quotients 45 Jerison's EQ 0.47-0.56 46 Eisenberg's EQ 0.61-0.74 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 68 of 78 1 2 3 Table 2. 4 5 Equation Group Anatomical Measurement Body Source 6 landmark mass (g) 7 Postcranial equations 8 Tibial midshaft 5.39 mm 1430.89* This paper 9 Gingerich Mammalia diameter1 10 11 (1990) Tibial length 58.60 mm 520.04 This paper 12 Tsubamoto Trochlear 13 (2014) Mammalia width of 5.89 mm 1122.77* This paper 14 astragalus 15 Moncunill-Solé Distal femoral 16 et al. (2014) Rodentia transversal 13.17 mm 1464.68* This paper 17 diameter 18 Moncunill-Solé Proximal tibial 19 20 et al. (2015) Lagomorpha transversal 13.33 mm 1311.26* This paper 21 diameter 22 Cranial equations 23 Kielan- 24 Jaworowska Mammalia Cranial length 57.6 mm 305.76 This paper 25 (1983) 26 Thewissen & Insectivora 27 Gingerich + Cranial length 57.6 mm 444.99 This paper 28 29 (1989) Tupaiidae 30 Millien & Bovy Rodentia Cranial length 57.6 mm 649.31 This paper 31 (2010) 32 Bertrand et al. Rodentia Cranial length 57.6 mm 531.39 This paper 33 (2016a) 34 Dental equations 35 López-Torres & 36 Conroy (1986) Prosimii M1 area 19.58 mm2 1756.11 Fostowicz-Frelik 37 (2018) 38 39 Left toothrow 15.52 mm 1311.48* This paper 40 Bertrand et al. Rodentia length 2 41 (2016a) Right toothrow 15.66 mm 1348.56* This paper 42 length2 43 1The tibial shaft diameter in Anagale gobiensis was measured immediately distal to the tibial crest. 44 45 2Toothrow length in Anagale gobiensis is treated as P4-M3 toothrow length to be comparable to rodents. 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 69 of 78 1 2 3 Table 3. 4 5 Body mass equation Jerison’s (1973) EQ Eisenberg’s (1981) EQ 6 Gingerich (1990) 0.48 0.62 7 Tsubamoto (2014) 0.56 0.74 8 Moncunill-Solé et al. (2014) 0.47 0.61 9 Moncunill-Solé et al. (2015) 0.50 0.66 10 Bertrand et al. (2016a) 0.50 0.66 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 1 Page 70 of 78 1 Total Olfactory 2 Order Superfamily Family Species Specimen number Endocrania Log Endocr l Volume Vol bulb volume Log OB Vol (mm3) Body mass (g) Log Body Mass Type of body mass equation used Source (mm3) 3 - - - - Anagalidae Anagalidae Anagale gobiensis (low body mass estimate) Anagale gobiensis (high body mass estimate) AMNH 26079 AMNH 26079 7798.04 7798.04 3.891985459 3.891985459 557.78 557.78 2.746462938 1122.77 3.0502908 Trocheal width of astragalus (Tsubamoto, 2014) 2.746462938 1464.67 3.165739786 Distal femoral transversal diameter (Moncunill-Solé et al., 2014) This paper This paper 4 Lagomorpha - Rodentia Sciuroidea Megalagidae Megalagus turgidus Aplodontiidae Altasciurus relictus FMNH UC 1642 USNM 437793 7052.78 957.45 3.848360337 2.981113836 280.1 34 2.447313109 2325.01 3.366424825 1.531478917 30.07 1.478133428 Width of occipital condyles (Moncunill-Solé et al., 2015) Cranial length (Bertrand et al., 2016) López-Torres et al. (2020) Bertrand et al (2019) Rodentia Sciuroidea Sciuridae Protosciurus cf. rachelae YPM 14736 4546.82 3.657707762 216.41 2.335277325 349.62 2.543596268 Cranial length (Bertrand et al., 2016) Bertrand et al (2019) 5 Rodentia Rodentia Sciuroidea - Sciuridae Cedromus wilsoni Ischyromyidae Paramys copei USNM 256584 AMNH 4756 3609.87 7526.65 3.557491562 3.876601721 106.97 455.45 2.029257936 2.65844166 268.89 2.429574651 1029.89 3.012790841 Cranial length (Bertrand et al., 2016) Cheektooth area (Bertrand et al., 2016) Bertrand et al (2019) Bertrand et al (2019) 6 Rodentia Rodentia - - Ischyromyidae Paramys delicatus Ischyromyidae Pseudotomus horribilis AMNH 12506 USNM 17159 12565.40 4.099176318 15188.20 4.181506307 595.51 808.92 2.7748876 2913.82 3.46446272 2.907905573 7466.70 3.873128748 Cranial length (Bertrand et al., 2016) Cranial length (Bertrand et al., 2016) Bertrand et al (2019) Bertrand et al (2019) Rodentia - Ischyromyidae Pseudotomus robustus USNM 17161 12063.00 4.081455328 717.06 2.855553074 5396.00 3.732071693 Cranial length (Bertrand et al., 2016) Bertrand et al (2019) 7 Rodentia Rodentia - - Ischyromyidae Pseudotomus petersoni Ischyromyidae Pseudotomus hians AMNH 2018 AMNH 5025 17014.90 4.230829401 13679.10 4.136057524 704.34 743.20 2.847784203 6644.56 3.822466227 2.871105117 3153.50 3.498792835 Cranial length (Bertrand et al., 2016) Cranial length (Bertrand et al., 2016) Bertrand et al (2019) Bertrand et al (2019) 8 Rodentia Rodentia Rodentia - - - Ischyromyidae Reithroparamys delicatissimus Ischyromyidae Rapamys atramontis Ischyromyidae Rapamys atramontis AMNH 12561 AMNH 128706 AMNH 128704 6433.98 7109.97 6006.47 4.136057524 3.851867768 3.778619312 206.31 224.62 226.06 2.314511859 843.33 3.498792835 2.351444556 1307.61 3.116477912 2.354219881 918.93 2.96328243 Cranial length (Bertrand et al., 2016) Cranial length (Bertrand et al., 2016) Cheektooth area (Bertrand et al., 2016) Bertrand et al (2019; 2021) Bertrand et al (2019) Bertrand et al (2019) 9 Rodentia Rodentia - - Ischyromyidae Ischyromys typus Ischyromyidae Ischyromys typus ROMV 1007 AMNH 12252 5578.07 5934.55 3.74648396 3.773387793 180.09 218.46 2.255489598 1342.23 3.127826941 2.339367953 1086.42 3.035997752 Cranial length (Bertrand et al., 2016) Cheektooth area (Bertrand et al., 2016) Bertrand et al (2019) Bertrand et al (2019) 10 Rodentia - Ischyromyidae Ischyromys typus AMNH F:AM 144638 7276.91 3.861947004 229.12 2.360063001 1109.01 3.044935462 Cheektooth area (Bertrand et al., 2016) Bertrand et al (2019) Primates - Microsyopidae Microsyops annectens UW 12362 5900 3.770852012 300.00 2.477121255 1710 3.23299611 Cranial length (Silcox et al., 2009a) Silcox et al (2010) Primates Paromomyoidea Paromomyidae Ignacius graybullianus USNM 421608 2140 3.330413773 120 2.079181246 253 2.403120521 Cranial length (Silcox et al., 2009a) Silcox et al (2009b) 11 Primates Primates Plesiadapoidea Plesiadapidae Plesiadapis tricuspidens Plesiadapoidea Plesiadapidae Plesiadapis cookei MNHN CR 125 UM 87990 5210 5000 3.716837723 3.698970004 136 - 2.133538908 - 6372 2239 3.804275767 Cranial length (Martin, 1990) 3.342422681 Long bone lengths and diameters (Gingerich & Gunnell, 2005) Orliac et al. (2014) Gingerich & Gunnell (2005) 12 Primates Adapoidea Notharctidae Notharctus tenebrosus AMNH 127167 7380 3.868056362 155 2.190331698 2641 3.421768401 M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) Primates Adapoidea Notharctidae Notharctus tenebrosus USNM V 23277 8060 3.906335042 180 2.255272505 2923 3.465828815 M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) Primates Adapoidea Notharctidae Notharctus tenebrosus USNM V 23278 7430 3.870988814 112 2.049218023 2244 3.351022853 M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) 13 Primates Primates Adapoidea Adapoidea Notharctidae Notharctidae Smilodectes gracilis Smilodectes gracilis USNM V 17994 USNM V 17996 8630 8990 3.936010796 3.953759692 178 150 2.250420002 2.176091259 1420 1303 3.152288344 3.114944416 M1 area (Dagosto & Terranova, 1992) M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) Harrington et al. (2016) Primates Adapoidea Notharctidae Smilodectes gracilis USNM V 21815 7440 3.871572936 92.2 1.964730921 1582 3.199206479 M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) 14 Primates Primates Adapoidea Adapoidea Notharctidae Adapidae Smilodectes gracilis Adapis parisiensis UM 32773 NHM M 1345 8420 8810 3.925312091 3.944975908 130 212 2.113943352 2.326335861 1547 1074 3.189490314 3.031004281 M1 area (Dagosto & Terranova, 1992) M1 area (Dagosto & Terranova, 1992) Harrington et al. (2016) Harrington et al. (2016) 15 Primates Primates - Omomyoidea - Rooneyia viejaensis Microchoeridae Microchoerus erinaceus TMM 40688-7 UM-PRR 1771 7234 4260 3.859378504 3.629409599 94 41 1.973127854 1.612783857 381 597 2.580924976 2.775974331 Cranial length (Silcox et al., 2009a) m1 area (Conroy, 1987) Kirk et al. (2014); Harrington et al. (2020) Ramdarshan & Orliac (2016) Primates Omomyoidea Microchoeridae Necrolemur antiquus MaPhQ 289 2360 3.372912003 46.6 1.668385917 144 2.158362492 Cranial length (Martin, 1990) Harrington et al. (2020) 16 Apatotheria - Apatotheria - Apatemyidae Apatemyidae Labidolemur kayi (assumption set 1) Labidolemur kayi (assumption set 2) USNM 530208/530221 USNM 530208/530221 607.79 605.52 2.78375355 2.782128492 74.05 74.05 1.869525063 1.869525063 74 74 1.86923172 1.86923172 Postcranial measurements (Armstrong et al., 2011) Postcranial measurements (Armstrong et al., 2011) Silcox et al (2011) Silcox et al (2011) 17 Apatotheria - Apatotheria - Apatemyidae Apatemyidae Labidolemur kayi (assumption set 3) Labidolemur kayi (assumption set 4) USNM 530208/530221 USNM 530208/530221 501.88 524.13 2.700599889 2.719439018 74.05 68.22 1.869525063 1.833911715 74 74 1.86923172 1.86923172 Postcranial measurements (Armstrong et al., 2011) Postcranial measurements (Armstrong et al., 2011) Silcox et al (2011) Silcox et al (2011) Table S1. Data for bivariate plots in Figures 5 and 13, including endocranial volume, olfactory bulb volume, and body mass for Anagale gobiensis, stem lagomorphs, fossil rodents, plesiadapiforms, fossil euprimates, and apatemyids. Information on how body mass was estimated for each species is included. 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 71 of 78 1 Petrosal 2 Body mass lobule Log Body Log PL Order Family Species Specimen 3 (g) volume mass volume 4 (mm3) - Anagalidae Anagale gobiensis (low body mass estimate) AMNH 26079 1122.77 77.98 3.0502908 1.8919832 5 - Anagalidae Anagale gobiensis (high body mass estimate) AMNH 26079 1464.67 77.98 3.1657398 1.8919832 6 Lagomorpha Leporidae Brachylagus idahoensis AMNH 92869 339.56 153.04 2.5309165 2.184805 7 Lagomorpha Leporidae Lepus americanus phaeonotus AMNH 97648 998.68 201.00 2.9994264 2.3031961 Lagomorpha Leporidae Lepus arcticus AMNH 42139 4003.10 270.10 3.6023964 2.4315246 8 Lagomorpha Leporidae Lepus americanus bairdii AMNH 99352 1396.22 231.02 3.1449539 2.3636496 9 Lagomorpha Leporidae Oryctolagus cuniculus AMNH 34816 1796.07 207.03 3.2543233 2.3160333 10 Lagomorpha Leporidae Poelagus marjorita AMNH 51052 2480.24 210.83 3.3944937 2.3239324 Lagomorpha Leporidae Romerolagus diazi AMNH 148172 1027.79 142.54 3.0119044 2.1539368 11 Lagomorpha Ochotonidae Ochotona pallasi AMNH 59712 223.85 59.32 2.3499571 1.7732011 12 Lagomorpha Ochotonidae Ochotona princeps princeps AMNH 120698 202.92 74.74 2.3073249 1.8735531 13 Lagomorpha Ochotonidae Ochotona princeps schisticeps AMNH 40547 155.10 71.80 2.1906118 1.8561244 Lagomorpha Megalagidae Megalagus turgidus FMNH UC 1642 2325.01 162.59 3.3664248 2.2110938 14 Rodentia Extant Aplodontiidae Aplodontia rufa AMNH 42389 1475.86 64.41 3.1690452 1.8089533 15 Rodentia Fossil Aplodontiidae Altasciurus relictus USNM 437793 30.07 32.08 1.4781334 1.5062208 16 Rodentia Fossil Aplodontiidae Mesogaulus paniensis AMNH F:AM 65511 266.48 46.27 2.4256646 1.6652995 Rodentia Extant Sciuridae Callosciurus sp. USNM 294865 379.50 123.91 2.5792118 2.0931064 17 Rodentia Extant Sciuridae Dremomys rufigenis USNM 488602 281.57 125.71 2.4495864 2.0993698 18 Rodentia Extant Sciuridae Lariscus insignis USNM 488570 194.45 113.00 2.2888079 2.0530784 Rodentia Extant Sciuridae Rhinosciurus laticaudatus USNM 488511 393.21 97.36 2.5946246 1.9883806 19 Rodentia Extant Sciuridae Aeromys tephromelas USNM 481190 971.69 166.10 2.9875277 2.2203696 20 Rodentia Extant Sciuridae Glaucomys volans AMNH 240290 63.30 33.73 1.8014037 1.5280163 21 Rodentia Extant Sciuridae Hylopetes spadiceus USNM 488639 80.57 17.93 1.9061734 1.2535803 Rodentia Extant Sciuridae Petaurista petaurista USNM 589079 2050.35 199.90 3.311828 2.3008128 22 Rodentia Extant Sciuridae Pteromys buechneri USNM 172622 86.73 37.53 1.9381693 1.5743786 23 Rodentia Extant Sciuridae Pteromyscus pulverulentus USNM 481178 248.25 49.50 2.3948893 1.6946052 24 Rodentia Extant Sciuridae Ratufa affinis USNM 488104 783.80 234.90 2.8942053 2.370883 Rodentia Extant Sciuridae Sciurus carolinensis AMNH 258346 437.91 163.26 2.6413849 2.2128798 25 Rodentia Extant Sciuridae Sciurus granatensis USNM 441999 242.05 131.63 2.3839051 2.1193549 26 Rodentia Extant Sciuridae Tamiasciurus hudsonicus USNM 549146 189.29 120.02 2.2771277 2.0792536 27 Rodentia Extant Sciuridae Eutamias minimus USNM 298500 34.17 34.80 1.533645 1.5415792 Rodentia Extant Sciuridae Funisciurus pyrropus USNM 294865 220.16 99.54 2.3427384 1.9979976 28 Rodentia Extant Sciuridae Heliosciurus rufobrachium USNM 378091 394.39 113.61 2.5959259 2.0554166 29 Rodentia Extant Sciuridae Paraxerus cepapi USNM 367956 182.66 44.66 2.2616435 1.6499187 30 Rodentia Extant Sciuridae Protoxerus stangeri USNM 435027 544.05 183.74 2.7356388 2.2642037 Rodentia Extant Sciuridae Xerus rutilus AMNH 179092 353.90 111.95 2.5488832 2.0490241 31 Rodentia Extant Sciuridae Marmota marmota AMNH 146619 4546.97 127.40 3.6577223 2.1051694 32 Rodentia Extant Sciuridae Cynomys ludovicianus AMNH114522 938.70 70.97 2.972525 1.8510748 33 Rodentia Extant Sciuridae Urocitellus richardsonii AMNH 15062 245.78 55.33 2.3905463 1.7429607 Rodentia Fossil Sciuridae Cedromus wilsoni USNM 256584 393.49 113.96 2.5949337 2.0567524 34 Rodentia Fossil Sciuridae Protosciurus cf. rachelae YPM 14736 349.62 150.42 2.5435963 2.1773056 35 Rodentia Ischyromyidae Ischyromys typus AMNH F:AM 144638 1109.01 116.58 3.0449355 2.0666241 36 Rodentia Ischyromyidae Ischyromys typus ROMV 1007 1401.80 91.12 3.1466861 1.9596137 Rodentia Ischyromyidae Paramys copei AMNH 4756 1029.89 89.99 3.0127908 1.9541943 37 Rodentia Ischyromyidae Paramys delicatus AMNH 12506 2704.83 129.30 3.43214 2.1115985 38 Rodentia Ischyromyidae Pseudotomus hians AMNH 5025 3153.50 142.40 3.4987928 2.15351 39 Rodentia Ischyromyidae Pseudotomus horribilis USNM 17159 4472.67 173.80 3.6505669 2.2400498 Rodentia Ischyromyidae Pseudotomus robustus USNM 17161 3911.71 74.80 3.5923667 1.8739016 40 Rodentia Ischyromyidae Pseudotomus petersoni AMNH 2018 6644.56 65.80 3.8224662 1.8182259 41 Rodentia Ischyromyidae Rapamys atramontis AMNH 128704 918.93 123.02 2.9632824 2.0899757 42 Rodentia Ischyromyidae Rapamys atramontis AMNH 128706 1057.82 108.09 3.0244118 2.0337855 Rodentia Ischyromyidae Reithroparamys delicatissimus AMNH 12561 843.33 148.86 2.9259975 2.172778 43 Primates - Rooneyia viejaensis TMM 40688-7 381 60.53 2.580925 1.7819685 44 Primates Adapoidea Adapis parisiensis NHM M 1345 1074 44.92 3.0310043 1.6524194 45 Primates Adapoidea Notharctus tenebrosus AMNH 127167 2641 103.90 3.4217684 2.0166114 Primates Adapoidea Notharctus tenebrosus USNM V 23277 2923 78.26 3.4658288 1.8935393 46 Primates Adapoidea Smilodectes gracilis USNM V 17996 1303 111.35 3.1149444 2.0466863 47 Primates Adapoidea Smilodectes gracilis USNM V 21815 1582 120.08 3.1992065 2.0794779 48 Primates Adapoidea Smilodectes gracilis UM 32773 (=MPM 2612) 1547 122.63 3.1894903 2.0885932 Primates Omomyoidea Microchoerus erinaceus UM-PRR1771 597 48.25 2.7759743 1.683536 49 Primates Omomyoidea Necrolemur antiquus MaPhQ 289 144 25.59 2.1583625 1.4081314 50 Table S2. Data for the bivariate plot in Figure 11, including petrosal lobule volume and body mass for Anagale gobiensis, living and fossil lagomorphs, and living and fossil rodents. 51 52 53 54 55 56 57 58 59 60 Page 72 of 78 1 2 Group Species Body mass Brain Jerison's Eisenberg's (g) mass (g) EQ EQ Source 3 Anagalidae Anagale gobiensis (low body mass estimate) 1122.77 7.4267 0.559656 0.74277404 This paper 4 Anagalidae Anagale gobiensis (high body mass estimate) 1464.67 7.4267 0.468346 0.61012735 This paper Stem lagomorph Megalagus turgidus 2325.01 6.71693 0.310803 0.39200494 López-Torres et al. (2020) 5 Extant Lagomorpha Brachylagus idahoensis 339.561082 4.90014 0.822835 1.18741999 López-Torres et al. (2020) 6 Extant Lagomorpha Lepus americanus phaeonotus 998.681398 9.73476 0.793465 1.06175485 López-Torres et al. (2020) Extant Lagomorpha Lepus americanus bairdii 1396.21846 9.1093 0.593178 0.7753435 López-Torres et al. (2020) 7 Extant Lagomorpha Lepus arcticus 4003.10484 15.191 0.488425 0.59304291 López-Torres et al. (2020) 8 Extant Lagomorpha Extant Lagomorpha Oryctolagus cuniculus Romerolagus diazi 1796.06978 1027.79098 8.91727 5.73354 0.490517 0.458422 0.62995199 López-Torres et al. (2020) 0.61219303 López-Torres et al. (2020) 9 Extant Lagomorpha Poelagus marjorita marjorita 2480.24151 11.2456 0.4983 0.6256518 López-Torres et al. (2020) 10 Extant Lagomorpha Extant Lagomorpha Ochotona pallasi Ochotona princeps princeps 223.84704 202.923758 2.03624 2.16221 0.452043 0.512629 0.67164444 López-Torres et al. (2020) 0.76691336 López-Torres et al. (2020) 11 Extant Lagomorpha Ochotona princeps schisticeps 155.100148 2.36176 0.670416 1.02201659 López-Torres et al. (2020) Fossil Sciuroidea Altasciurus relictus 30.07 0.91185 0.776951 1.32855846 Bertrand et al. (2019) 12 Fossil Sciuroidea Mesogaulus paniensis 266.48 3.30 0.65 0.96 Bertrand et al. (2018) 13 Fossil Sciuroidea Protosciurus cf. rachelae 349.62 4.3303 0.713063 1.02690931 Bertrand et al. (2019) Fossil Sciuroidea Cedromus wilsoni 268.89 3.43797 0.675003 0.99012925 Bertrand et al. (2019) 14 Ischyromyidae Paramys copei 1029.89 7.16824 0.57235 0.76422646 Bertrand et al. (2019) 15 Ischyromyidae Ischyromyidae Paramys delicatus Pseudotomus horribilis 2913.82 7466.70 11.967 14.465 0.47601 0.306295 0.59096189 0.35602181 Bertrand et al. (2019) Bertrand et al. (2019) 16 Ischyromyidae Pseudotomus robustus 5396.00 11.4886 0.30241 0.3595902 Bertrand et al. (2019) Ischyromyidae Pseudotomus petersoni 6644.56 16.2047 0.371028 0.43480043 Bertrand et al. (2019) 17 Ischyromyidae Pseudotomus hians 3153.50 13.0277 0.491469 0.60678718 Bertrand et al. (2019) 18 Ischyromyidae Rapamys atramontis 1307.61 6.7714 0.460741 0.605006 Bertrand et al. (2019) Ischyromyidae Rapamys atramontis 918.93 5.72045 0.493003 0.66355341 Bertrand et al. (2019) 19 Ischyromyidae Ischyromys typus 1342.23 5.31245 0.355197 0.46556207 Bertrand et al. (2019) 20 Ischyromyidae Ischyromyidae Ischyromys typus Ischyromys typus 1086.42 1109.01 5.65195 6.93039 0.43541 0.526586 0.57920863 0.6994881 Bertrand et al. (2019) Bertrand et al. (2019) 21 Plesiadapiformes Microsyops annectens 1710 5.61905 0.319429 0.41388806 Silcox et al. (2010) Plesiadapiformes Ignacius graybullianus 253 2.0381 0.416824 0.6140308 Silcox et al. (2009) 22 Plesiadapiformes Plesiadapis tricuspidens 6372 4.9619 0.116843 0.13732804 Orliac et al. (2014) 23 Plesiadapiformes Plesiadapis cookei 2200 4.7619 0.228653 0.28950948 Gingerich & Gunnell (2005) Apatemyidae Labidolemur kayi (low brain mass estimate) 74 0.47798 0.222762 0.35764366 Silcox et al. (2011) 24 Apatemyidae Labidolemur kayi (high brain mass estimate) 74 0.57884 0.269769 0.43311371 Silcox et al. (2011) 25 Euprimates Euprimates Notharctus tenebrosus Notharctus tenebrosus 2641 2923 7.02857 7.67619 0.298608 0.304691 0.37327879 0.3781876 Harrington et al. (2016) Harrington et al. (2016) 26 Euprimates Notharctus tenebrosus 2244 7.07619 0.3353 0.42395277 Harrington et al. (2016) 27 Euprimates Euprimates Smilodectes gracilis Smilodectes gracilis 1420 1303 8.21905 8.5619 0.529184 0.583951 0.69088059 0.76698416 Harrington et al. (2016) Harrington et al. (2016) 28 Euprimates Smilodectes gracilis 1582 7.08571 0.42436 0.54985188 Harrington et al. (2016) Euprimates Smilodectes gracilis 1547 8.01905 0.487509 0.63266649 Harrington et al. (2016) 29 Euprimates Adapis parisiensis 1074 8.39048 0.651377 0.86719808 Harrington et al. (2016) 30 Euprimates Rooneyia viejaensis 381 6.88952 1.070997 1.53313215 Kirk et al. (2014) Euprimates Microchoerus erinaceus 597 4.05714 0.466806 0.64755128 Ramdarshan & Orliac (2016) 31 Euprimates Necrolemur antiquus 144 2.36 0.70409 1.08482846 Harrington et al. (2020) 32 Extant Rodentia Extant Rodentia Acomys dimidiatus Acomys dimidiatus 76 50.5 0.97 0.884 0.444064 0.532189 0.7116141 0.87758969 Bertrand et al. (2019) Bertrand et al. (2019) 33 Extant Rodentia Aeromys tephromelas 904.59 10.92 0.95071 1.28101056 Bertrand et al. (2019) 34 Extant Rodentia Extant Rodentia Allactaga elater Allactaga sibirica 80 193 1.9 3.5 0.840431 0.85815 1.34196713 1.28834004 Bertrand et al. (2019) Bertrand et al. (2019) 35 Extant Rodentia Allactaga sibirica 106 2 0.732645 1.1470399 Bertrand et al. (2019) Extant Rodentia Aplodontia rufa 1475.86 7.52 0.471755 0.61424253 Bertrand et al. (2019) 36 Extant Rodentia Aplodontia rufa 982.00 7.60 0.626495 0.83931688 Bertrand et al. (2019) 37 Extant Rodentia Aplodontia rufa 870.00 8.40 0.750966 1.01463516 Bertrand et al. (2019) Extant Rodentia Aplodontia rufa 710.00 8.40 0.860508 1.17929641 Bertrand et al. (2019) 38 Extant Rodentia Aplodontia rufa 985.00 8.80 0.723934 0.96964939 Bertrand et al. (2019) 39 Extant Rodentia Extant Rodentia Aplodontia rufa Apodemus agrarius 887.00 28.5 8.30 0.632 0.732467 0.5582 0.98830156 0.95809098 Bertrand et al. (2019) Bertrand et al. (2019) 40 Extant Rodentia Apodemus agrarius agrarius 26.2 0.62 0.57936 1.00028465 Bertrand et al. (2019) Extant Rodentia Apodemus flavicollis 30 0.75 0.640042 1.09462762 Bertrand et al. (2019) 41 Extant Rodentia Apodemus flavicollis flavicollis 33 0.8 0.640478 1.0880893 Bertrand et al. (2019) 42 Extant Rodentia Apodemus sylvaticus 19.4 0.59 0.674285 1.18892265 Bertrand et al. (2019) Extant Rodentia Apodemus sylvaticus 22 0.564 0.592481 1.03552643 Bertrand et al. (2019) 43 Extant Rodentia Apodemus sylvaticus 21.6 0.59 0.627461 1.09807273 Bertrand et al. (2019) 44 Extant Rodentia Apodemus sylvaticus sylvaticus 19 0.6 0.695353 1.22785888 Bertrand et al. (2019) Extant Rodentia Arvicola amphibius 137.8 1.53 0.470125 0.7226398 Bertrand et al. (2019) 45 Extant Rodentia Arvicola amphibius 131 1.5 0.476802 0.73550456 Bertrand et al. (2019) 46 Extant Rodentia Extant Rodentia Arvicola amphibius Atherurus africanus 84.5 1600 1.086 17.2 0.463078 1.02232 0.73659833 1.3235932 Bertrand et al. (2019) Bertrand et al. (2019) 47 Extant Rodentia Atherurus africanus 3620 25.3 0.870171 1.06402308 Bertrand et al. (2019) Extant Rodentia Atherurus africanus 1925 17.8 0.934696 1.19458244 Bertrand et al. (2019) 48 Extant Rodentia Atherurus africanus 2250 23 1.087888 1.37526896 Bertrand et al. (2019) 49 Extant Rodentia Atlantoxerus getulus 251.00 3.75 0.771515 1.13716435 Bertrand et al. (2019) Extant Rodentia Brachytarsomys albicauda 300 2.47 0.450655 0.65599736 Bertrand et al. (2019) 50 Extant Rodentia Brachyuromys ramirohitra 94 1.4 0.555842 0.87758318 Bertrand et al. (2019) 51 Extant Rodentia Extant Rodentia Callosciurus sp. Callospermophilus lateralis 437.35 246.00 6.67 2.98 0.945942 0.621223 1.34110378 0.91693369 Bertrand et al. (2019) Bertrand et al. (2019) 52 Extant Rodentia Callospermophilus lateralis 217.00 3.10 0.701592 1.0446931 Bertrand et al. (2019) 53 Extant Rodentia Extant Rodentia Capromys pilorides Castor canadensis 7000 20500 11 44 0.243218 0.473583 0.28398474 0.51289644 Bertrand et al. (2019) Bertrand et al. (2019) 54 Extant Rodentia Castor canadensis 18000 53 0.622389 0.68021951 Bertrand et al. (2019) Extant Rodentia Castor canadensis 14500 45 0.610822 0.67775864 Bertrand et al. (2019) 55 Extant Rodentia Castor canadensis 22500 52 0.525847 0.56580027 Bertrand et al. (2019) 56 Extant Rodentia Castor canadensis 8000 38 0.768302 0.88873416 Bertrand et al. (2019) Extant Rodentia Castor canadensis 9750 48 0.850016 0.96973415 Bertrand et al. (2019) 57 Extant Rodentia Castor canadensis 12500 42 0.629707 0.70600983 Bertrand et al. (2019) 58 Extant Rodentia Extant Rodentia Castor canadensis Castor canadensis 16000 5500 38 43 0.482883 1.117491 0.53212012 1.32701226 Bertrand et al. (2019) Bertrand et al. (2019) 59 Extant Rodentia Castor canadensis 20000 40 0.437712 0.47486774 Bertrand et al. (2019) Extant Rodentia Castor canadensis 4180 25.48 0.795848 0.96339379 Bertrand et al. (2019) 60 Extant Rodentia Castor canadensis 5380 29.52 0.778594 0.92600288 Bertrand et al. (2019) Extant Rodentia Castor canadensis 14500 45 0.610822 0.67775864 Bertrand et al. (2019) Extant Rodentia Castor fiber 14300 41.4 0.56721 0.62997968 Bertrand et al. (2019) Page 73 of 78 1 Extant Rodentia Castor fiber 16900 38.8 0.475298 0.52175958 Bertrand et al. (2019) 2 Extant Rodentia Castor fiber 25000 45 0.424045 0.45290986 Bertrand et al. (2019) 3 Extant Rodentia Extant Rodentia Castor fiber Castor fiber 23100 20000 45.5 39 0.452075 0.426769 0.48552702 0.46299604 Bertrand et al. (2019) Bertrand et al. (2019) 4 Extant Rodentia Cavia aperea 163 2.7 0.741339 1.12621111 Bertrand et al. (2019) Extant Rodentia Cavia aperea 260 3 0.602432 0.88575955 Bertrand et al. (2019) 5 Extant Rodentia Cavia aperea 430 3.9 0.559061 0.7935473 Bertrand et al. (2019) 6 Extant Rodentia Cavia aperea 647 5.46 0.595258 0.82110433 Bertrand et al. (2019) Extant Rodentia Cavia aperea 460 4.2 0.575467 0.81298656 Bertrand et al. (2019) 7 Extant Rodentia Cavia aperea 792 4.7 0.447477 0.60857778 Bertrand et al. (2019) 8 Extant Rodentia Extant Rodentia Cavia aperea Cavia aperea f. porcellus 540 485 4.1 4.57 0.504544 0.604349 0.70483481 0.85063291 Bertrand et al. (2019) Bertrand et al. (2019) 9 Extant Rodentia Cavia porcellus 520 5.4 0.681538 0.95461038 Bertrand et al. (2019) 10 Extant Rodentia Extant Rodentia Cavia porcellus Cavia porcellus 650 348 5.8 6.4 0.630369 1.05716 0.86925456 1.52295195 Bertrand et al. (2019) Bertrand et al. (2019) 11 Extant Rodentia Cavia porcellus 648 4.4 0.479199 0.66094001 Bertrand et al. (2019) Extant Rodentia Cavia porcellus 493.1 3.8 0.496977 0.69869328 Bertrand et al. (2019) 12 Extant Rodentia Cavia porcellus 361 4.1 0.660804 0.94951842 Bertrand et al. (2019) 13 Extant Rodentia Cavia porcellus 324 3.8 0.658472 0.95335612 Bertrand et al. (2019) Extant Rodentia Cavia porcellus 214.94 3.32 0.757363 1.12849124 Bertrand et al. (2019) 14 Extant Rodentia Cavia porcellus 214.57 3.28 0.749103 1.11631729 Bertrand et al. (2019) 15 Extant Rodentia Extant Rodentia Cavia porcellus Cavia porcellus 432 456 4 4.23 0.571616 0.582979 0.81110464 0.82410254 Bertrand et al. (2019) Bertrand et al. (2019) 16 Extant Rodentia Cavia porcellus 675 4.54 0.481106 0.66167683 Bertrand et al. (2019) Extant Rodentia Cavia porcellus 971 4.28 0.355489 0.47662455 Bertrand et al. (2019) 17 Extant Rodentia Cavia porcellus 900 4.94 0.431721 0.58191876 Bertrand et al. (2019) 18 Extant Rodentia Cavia porcellus 500 5 0.647857 0.90992809 Bertrand et al. (2019) Extant Rodentia Cavia porcellus 700 4.73 0.489175 0.67106326 Bertrand et al. (2019) 19 Extant Rodentia Chaetodipus baileyi 31.2 0.61905 0.514588 0.87765641 Bertrand et al. (2019) 20 Extant Rodentia Extant Rodentia Chaetodipus californicus Chaetodipus fallax 26 20.3 0.57143 0.49524 0.536721 0.549048 0.92716396 0.96503229 Bertrand et al. (2019) Bertrand et al. (2019) 21 Extant Rodentia Chaetodipus formosus 15.3 0.40952 0.548725 0.98374461 Bertrand et al. (2019) 22 Extant Rodentia Extant Rodentia Chaetodipus hispidus Chaetodipus penicillatus 35.2 16.5 0.62857 0.41905 0.481936 0.533787 0.81505691 0.95191929 Bertrand et al. (2019) Bertrand et al. (2019) 23 Extant Rodentia Chaetodipus spinatus 19.1 0.4381 0.505935 0.89305603 Bertrand et al. (2019) Extant Rodentia Chinchilla chinchilla 520 7.8 0.984444 1.37888167 Bertrand et al. (2019) 24 Extant Rodentia Chinchilla chinchilla 425 6.4 0.924652 1.31355129 Bertrand et al. (2019) 25 Extant Rodentia Chinchilla chinchilla 320 6.9 1.205639 1.74708074 Bertrand et al. (2019) Extant Rodentia Chinchilla chinchilla 470 8.9 1.201996 1.69555726 Bertrand et al. (2019) 26 Extant Rodentia Chinchilla chinchilla 450 6 0.834291 1.18045343 Bertrand et al. (2019) 27 Extant Rodentia Extant Rodentia Chinchilla lanigera Chinchilla lanigera 432 380 5.2 5.2 0.743101 0.80978 1.05443604 1.15941349 Bertrand et al. (2019) Bertrand et al. (2019) 28 Extant Rodentia Chinchilla lanigera 385 5.05 0.779563 1.11512951 Bertrand et al. (2019) Extant Rodentia Chinchilla lanigera 460 5.1 0.698781 0.98719796 Bertrand et al. (2019) 29 Extant Rodentia Chinchilla lanigera 385 4.9 0.756408 1.08200685 Bertrand et al. (2019) 30 Extant Rodentia Chinchilla lanigera 370 5.32 0.843403 1.20981015 Bertrand et al. (2019) Extant Rodentia Chinchilla lanigera 385 5.5 0.849029 1.21449748 Bertrand et al. (2019) 31 Extant Rodentia Chinchilla lanigera 425 6.4 0.924652 1.31355129 Bertrand et al. (2019) 32 Extant Rodentia Chinchilla lanigera 500 6 0.777428 1.0919137 Bertrand et al. (2019) Extant Rodentia Colomys goslingi 58.2 1.372 0.75106 1.22626963 Bertrand et al. (2019) 33 Extant Rodentia Cricetomys emini 1000 6.6 0.53748 0.71914887 Bertrand et al. (2019) 34 Extant Rodentia Extant Rodentia Cricetomys emini Cricetulus griseus 80.50 36 2.70 0.67 1.189322 0.506024 1.89823371 0.85444836 Bertrand et al. (2019) Bertrand et al. (2019) 35 Extant Rodentia Cricetulus griseus 23.18 0.628 0.637019 1.10930398 Bertrand et al. (2019) Extant Rodentia Cricetus cricetus 450 2.85 0.396288 0.56071538 Bertrand et al. (2019) 36 Extant Rodentia Cricetus cricetus 297 2.2 0.404105 0.58865085 Bertrand et al. (2019) 37 Extant Rodentia Cuniculus paca 5635 35.8 0.915383 1.08516654 Bertrand et al. (2019) Extant Rodentia Cuniculus paca 6125 33.5 0.810033 0.95468735 Bertrand et al. (2019) 38 Extant Rodentia Cuniculus paca 3665 33.2 1.132471 1.38356067 Bertrand et al. (2019) 39 Extant Rodentia Extant Rodentia Cuniculus paca Cuniculus paca 9000 5000 37.3 26.1 0.696923 0.723018 0.79954745 0.86432599 Bertrand et al. (2019) Bertrand et al. (2019) 40 Extant Rodentia Cuniculus paca 3627 21.85 0.750539 0.9176163 Bertrand et al. (2019) 41 Extant Rodentia Extant Rodentia Cuniculus paca Cynomys ludovicianus 4559 939 48 6.88 1.414547 0.584242 1.70197277 0.78518563 Bertrand et al. (2019) Bertrand et al. (2021) 42 Extant Rodentia Cynomys ludovicianus 1200.00 6.40 0.461262 0.60934145 Bertrand et al. (2019) Extant Rodentia Dasymys incomtus 102.5 1.574 0.58971 0.92543077 Bertrand et al. (2019) 43 Extant Rodentia Dasyprocta leporina 3600 21.6 0.745675 0.91214686 Bertrand et al. (2019) 44 Extant Rodentia Dasyprocta leporina 2684 20 0.840552 1.04955646 Bertrand et al. (2019) Extant Rodentia Dasyprocta leporina 2390 18.4 0.835815 1.05215142 Bertrand et al. (2019) 45 Extant Rodentia Dasyprocta leporina 2004 17 0.868953 1.10743815 Bertrand et al. (2019) 46 Extant Rodentia Extant Rodentia Dasyprocta leporina Dasyprocta leporina 2880 2350 18.5 18 0.741647 0.826943 0.92150173 1.04221461 Bertrand et al. (2019) Bertrand et al. (2019) 47 Extant Rodentia Dasyprocta leporina 2550 17 0.739411 0.92658242 Bertrand et al. (2019) Extant Rodentia Dasyprocta leporina 3172 18.3 0.687665 0.84867109 Bertrand et al. (2019) 48 Extant Rodentia Dasyprocta leporina 2371 19.8 0.904232 1.13891342 Bertrand et al. (2019) 49 Extant Rodentia Dasyprocta leporina 2400 27 1.223041 1.53915485 Bertrand et al. (2019) Extant Rodentia Dasyprocta leporina 2370 19.8 0.904487 1.13926901 Bertrand et al. (2019) 50 Extant Rodentia Dasyprocta mexicana 1527 17.8 1.091608 1.41792736 Bertrand et al. (2019) 51 Extant Rodentia Dasyprocta mexicana 2300 20 0.932161 1.17659301 Bertrand et al. (2019) Extant Rodentia Dasyprocta punctata 3172 18.34 0.689168 0.8505261 Bertrand et al. (2019) 52 Extant Rodentia Dendromus mesomelas 14 0.51 0.725242 1.30830891 Bertrand et al. (2019) 53 Extant Rodentia Extant Rodentia Desmodillus auricularis Dicrostonyx groenlandicus 60 52.1 1.12 0.8983 0.600725 0.529613 0.97872564 0.87143772 Bertrand et al. (2019) Bertrand et al. (2019) 54 Extant Rodentia Dipodomys deserti 114.7 1.6 0.555944 0.86560134 Bertrand et al. (2019) Extant Rodentia Dipodomys heermanni 71.3 1.38095 0.659823 1.06210467 Bertrand et al. (2019) 55 Extant Rodentia Dipodomys merriami 41 1.05714 0.731791 1.22447097 Bertrand et al. (2019) 56 Extant Rodentia Dipodomys microps 65.7 1.12381 0.567208 0.91826748 Bertrand et al. (2019) Extant Rodentia Dipodomys ordii 60.2 1.39048 0.744137 1.21209601 Bertrand et al. (2019) 57 Extant Rodentia Dipodomys panamintinus 74 1.46667 0.683542 1.09742492 Bertrand et al. (2019) 58 Extant Rodentia Extant Rodentia Dipodomys spectabilis Dolichotis patagonum 146.6 5500 1.98095 15 0.58396 0.389823 0.89373706 0.46291125 Bertrand et al. (2019) Bertrand et al. (2019) 59 Extant Rodentia Dolichotis patagonum 5650 25.7 0.655963 0.77748522 Bertrand et al. (2019) 60 Extant Rodentia Extant Rodentia Dolichotis patagonum Dolichotis patagonum 5650 7880 25.66 33.5 0.654942 0.684212 0.77627512 0.79230113 Bertrand et al. (2019) Bertrand et al. (2019) Extant Rodentia Dolichotis patagonum 7500 32 0.675581 0.78501781 Bertrand et al. (2019) Extant Rodentia Dolichotis patagonum 5500 26 0.675693 0.8023795 Bertrand et al. (2019) Page 74 of 78 1 Extant Rodentia Dolichotis patagonum 7200 30 0.650919 0.75852531 Bertrand et al. (2019) 2 Extant Rodentia Dremomys rufigenis 418.43 5.59 0.815626 1.15993435 Bertrand et al. (2019) 3 Extant Rodentia Extant Rodentia Eliurus minor Eliurus myoxinus 38 59 1.46 1.69 1.063449 0.916716 1.78890741 1.49530947 Bertrand et al. (2019) Bertrand et al. (2019) 4 Extant Rodentia Epixerus ebii 605.00 9.77 1.114296 1.54430811 Bertrand et al. (2019) Extant Rodentia Erethizon dorsatum 5160 27.9 0.756742 0.90264773 Bertrand et al. (2019) 5 Extant Rodentia Erethizon dorsatum 3430 28.7 1.023419 1.2561431 Bertrand et al. (2019) 6 Extant Rodentia Erethizon dorsatum 4980 25.1 0.697186 0.83367903 Bertrand et al. (2019) Extant Rodentia Erethizon dorsatum 4980 27.2 0.755517 0.90342907 Bertrand et al. (2019) 7 Extant Rodentia Erethizon dorsatum 4640 27.1 0.789262 0.94846401 Bertrand et al. (2019) 8 Extant Rodentia Extant Rodentia Erethizon dorsatum Erethizon dorsatum 6200 6640 27.1 24 0.649958 0.549765 0.76537507 0.64429051 Bertrand et al. (2019) Bertrand et al. (2019) 9 Extant Rodentia Erethizon dorsatum 2800 30.77 1.257044 1.56496786 Bertrand et al. (2019) 10 Extant Rodentia Extant Rodentia Erethizon dorsatum Erethizon dorsatum 2725 3410 21.22 19.15 0.882813 0.685555 1.10115647 0.84179319 Bertrand et al. (2019) Bertrand et al. (2019) 11 Extant Rodentia Erethizon dorsatum 5000 24 0.664845 0.79478252 Bertrand et al. (2019) Extant Rodentia Eutamias sibiricus 110.00 2.60 0.929093 1.45083368 Bertrand et al. (2019) 12 Extant Rodentia Funisciurus anerythrus 230.00 4.03 0.878238 1.30240885 Bertrand et al. (2019) 13 Extant Rodentia Funisciurus carruthersi 107.50 4.00 1.45156 2.27034886 Bertrand et al. (2019) Extant Rodentia Funisciurus carruthersi 158.00 4.50 1.261627 1.92079554 Bertrand et al. (2019) 14 Extant Rodentia Funisciurus carruthersi 195.50 3.25 0.790012 1.1849762 Bertrand et al. (2019) 15 Extant Rodentia Extant Rodentia Funisciurus carruthersi Funisciurus carruthersi 168.00 195.00 4.10 4.65 1.103177 1.132266 1.67236033 1.69864337 Bertrand et al. (2019) Bertrand et al. (2019) 16 Extant Rodentia Funisciurus isabella 160.50 3.14 0.871917 1.32601361 Bertrand et al. (2019) Extant Rodentia Funisciurus lemniscatus 154.00 3.22 0.918137 1.4003511 Bertrand et al. (2019) 17 Extant Rodentia Funisciurus pyrropus 258.75 4.47 0.899855 1.32350915 Bertrand et al. (2019) 18 Extant Rodentia Funisciurus pyrropus 301.15 4.34 0.789315 1.14866046 Bertrand et al. (2019) Extant Rodentia Funisciurus substriatus 186.10 3.70 0.930784 1.40095071 Bertrand et al. (2019) 19 Extant Rodentia Galea spixii 672 6.2 0.658981 0.90659495 Bertrand et al. (2019) 20 Extant Rodentia Extant Rodentia Geomys bursarius Geomys pinetis 192.4 313.5 1.48571 2.30476 0.365037 0.408287 0.54814874 0.59249563 Bertrand et al. (2019) Bertrand et al. (2019) 21 Extant Rodentia Gerbillus gerbillus 95 3.4 1.340364 2.11464913 Bertrand et al. (2019) 22 Extant Rodentia Extant Rodentia Gerbillus nanus Gerbillus pyramidum 18 79 0.52 1.14 0.62487 0.508526 1.10758371 0.81271013 Bertrand et al. (2019) Bertrand et al. (2019) 23 Extant Rodentia Gerbillus pyramidum 70 1.2 0.580476 0.93558568 Bertrand et al. (2019) Extant Rodentia Gerbillus pyramidum 72.3 1.08 0.511234 0.82212203 Bertrand et al. (2019) 24 Extant Rodentia Gerbillus pyramidum 79 1.059 0.472394 0.75496494 Bertrand et al. (2019) 25 Extant Rodentia Gerbillus pyramidum 57.9 1.004 0.551516 0.90079599 Bertrand et al. (2019) Extant Rodentia Gerbillus pyramidum 59.4 0.981 0.529725 0.86365821 Bertrand et al. (2019) 26 Extant Rodentia Glaucomys volans 63.97 1.91 0.983817 1.59570287 Bertrand et al. (2019) 27 Extant Rodentia Extant Rodentia Glaucomys volans Glaucomys volans 52.00 64.00 1.89 1.92 1.115728 0.986233 1.83609099 1.59956877 Bertrand et al. (2019) Bertrand et al. (2019) 28 Extant Rodentia Glis glis 118 1.4 0.477294 0.74166916 Bertrand et al. (2019) Extant Rodentia Glis glis 148 1.9 0.556541 0.85120624 Bertrand et al. (2019) 29 Extant Rodentia Grammomys dolichurus 45 1.18 0.767447 1.27579124 Bertrand et al. (2019) 30 Extant Rodentia Graphiurus murinus 17.7 0.551 0.66962 1.18830042 Bertrand et al. (2019) Extant Rodentia Heliosciurus gambianus 209.80 3.53 0.819207 1.22270988 Bertrand et al. (2019) 31 Extant Rodentia Heliosciurus rufobrachium 354.98 5.79 0.943314 1.35705735 Bertrand et al. (2019) 32 Extant Rodentia Heliosciurus rufobrachium isabellinus 280.90 4.84 0.922494 1.3490281 Bertrand et al. (2019) Extant Rodentia Heliosciurus rufobrachium rufobrachium 334.00 5.03 0.853794 1.23352259 Bertrand et al. (2019) 33 Extant Rodentia Heterocephalus glaber 39 0.52 0.372228 0.62501495 Bertrand et al. (2019) 34 Extant Rodentia Extant Rodentia Heterogeomys cherriei Heteromys desmarestianus 405 77.1 3.19048 1.07619 0.476079 0.487957 0.67859969 0.78116679 Bertrand et al. (2019) Bertrand et al. (2019) 35 Extant Rodentia Heteromys pictus 40.2 0.71429 0.501025 0.83949778 Bertrand et al. (2019) Extant Rodentia Hoplomys gymnurus 637 3.84762 0.423875 0.585334 Bertrand et al. (2019) 36 Extant Rodentia Hybomys univittatus 56.8 1.149 0.63933 1.04562782 Bertrand et al. (2019) 37 Extant Rodentia Hydrochoerus hydrochaeris 31000 85 0.693466 0.72960221 Bertrand et al. (2019) Extant Rodentia Hydrochoerus hydrochaeris 24130 61.4 0.59248 0.63438196 Bertrand et al. (2019) 38 Extant Rodentia Hydrochoerus hydrochaeris 29500 76 0.640991 0.67673739 Bertrand et al. (2019) 39 Extant Rodentia Extant Rodentia Hydrochoerus hydrochaeris Hydrochoerus hydrochaeris 10000 12000 55 53 0.957594 0.816664 1.09052985 0.91824086 Bertrand et al. (2019) Bertrand et al. (2019) 40 Extant Rodentia Hydrochoerus hydrochaeris 13000 53 0.774021 0.86543142 Bertrand et al. (2019) 41 Extant Rodentia Extant Rodentia Hydrochoerus hydrochaeris Hydrochoerus hydrochaeris 17000 28000 61 52 0.7443 0.454178 0.8167187 0.48126166 Bertrand et al. (2019) Bertrand et al. (2019) 42 Extant Rodentia Hydrochoerus hydrochaeris 24031 61.4 0.594114 0.63631487 Bertrand et al. (2019) Extant Rodentia Hydrochoerus hydrochaeris 28500 75 0.647342 0.68509522 Bertrand et al. (2019) 43 Extant Rodentia Hydrochoerus hydrochaeris 27670 52.21 0.459649 0.48746314 Bertrand et al. (2019) 44 Extant Rodentia Hydrochoerus hydrochaeris 26700 75 0.676266 0.71898164 Bertrand et al. (2019) Extant Rodentia Hydrochoerus hydrochaeris 34300 73 0.556538 0.58140676 Bertrand et al. (2019) 45 Extant Rodentia Hydrochoerus hydrochaeris 33000 75.5 0.590692 0.61875897 Bertrand et al. (2019) 46 Extant Rodentia Extant Rodentia Hydrochoerus hydrochaeris Hydrochoerus hydrochaeris 54000 29500 82 76.02 0.461242 0.641159 0.46678548 0.67691547 Bertrand et al. (2019) Bertrand et al. (2019) 47 Extant Rodentia Hydrochoerus hydrochaeris 28000 75 0.655065 0.6941274 Bertrand et al. (2019) Extant Rodentia Hydrochoerus hydrochaeris 32500 71 0.561197 0.5884906 Bertrand et al. (2019) 48 Extant Rodentia Hylopetes spadiceus 84.22 2.02 0.863155 1.37330095 Bertrand et al. (2019) 49 Extant Rodentia Hypogeomys antimena 875 8 0.712465 0.96223001 Bertrand et al. (2019) Extant Rodentia Hystrix brachyura 22000 43 0.441432 0.47571905 Bertrand et al. (2019) 50 Extant Rodentia Hystrix cristata 7036.5 36.5 0.804234 0.93869344 Bertrand et al. (2019) 51 Extant Rodentia Hystrix cristata 15000 32 0.424607 0.47002106 Bertrand et al. (2019) Extant Rodentia Hystrix cristata 10000 37 0.6442 0.73362917 Bertrand et al. (2019) 52 Extant Rodentia Hystrix hybrid 13500 37 0.526862 0.5875294 Bertrand et al. (2019) 53 Extant Rodentia Extant Rodentia Hystrix javanica Hystrix sp. 23000 15000 20 37.5 0.199293 0.497587 0.21410473 0.55080593 Bertrand et al. (2019) Bertrand et al. (2019) 54 Extant Rodentia Ictidomys tridecemlineatus 200.00 2.20 0.526685 0.7887427 Bertrand et al. (2019) Extant Rodentia Ictidomys tridecemlineatus 115.00 2.40 0.832458 1.29589468 Bertrand et al. (2019) 55 Extant Rodentia Ictidomys tridecemlineatus 139.00 2.16 0.660443 1.0145664 Bertrand et al. (2019) 56 Extant Rodentia Ictidomys tridecemlineatus 153.00 2.30 0.657511 1.00330033 Bertrand et al. (2019) Extant Rodentia Jaculus jaculus 73 1.85 0.87009 1.39825918 Bertrand et al. (2019) 57 Extant Rodentia Jaculus orientalis 140 2.64 0.802633 1.23237832 Bertrand et al. (2019) 58 Extant Rodentia Extant Rodentia Jaculus orientalis Lagidium viscacia 98 2460 2.5 12.4 0.965244 0.552477 1.51952406 0.69407208 Bertrand et al. (2019) Bertrand et al. (2019) 59 Extant Rodentia Lagidium viscacia 2252 16 0.756341 0.95608003 Bertrand et al. (2019) 60 Extant Rodentia Extant Rodentia Lagidium viscacia Lagidium viscacia 2350 3845 15.2 14.8 0.698308 0.488877 0.88009233 0.59526903 Bertrand et al. (2019) Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 6630 20 0.458601 0.5375079 Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 6385 21 0.493833 0.5803301 Bertrand et al. (2019) Page 75 of 78 1 Extant Rodentia Lagostomus maximus 5965 17.2 0.423339 0.49986368 Bertrand et al. (2019) 2 Extant Rodentia Lagostomus maximus 3930 18 0.585933 0.71235567 Bertrand et al. (2019) 3 Extant Rodentia Extant Rodentia Lagostomus maximus Lagostomus maximus 3765 2940 16.5 16.3 0.552765 0.644486 0.67405095 0.79962332 Bertrand et al. (2019) Bertrand et al. (2019) 4 Extant Rodentia Lagostomus maximus 7000 17 0.375882 0.43888551 Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 1575 12.2 0.732826 0.94983263 Bertrand et al. (2019) 5 Extant Rodentia Lagostomus maximus 1990 14.5 0.744655 0.94949262 Bertrand et al. (2019) 6 Extant Rodentia Lagostomus maximus 1395 12 0.781871 1.02204778 Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 1500 11.1 0.688907 0.8959635 Bertrand et al. (2019) 7 Extant Rodentia Lagostomus maximus 1265 11.8 0.820919 1.08046297 Bertrand et al. (2019) 8 Extant Rodentia Extant Rodentia Lagostomus maximus Lagostomus maximus 1125 1095 13.2 11.7 0.993392 0.896597 1.31824439 1.19204948 Bertrand et al. (2019) Bertrand et al. (2019) 9 Extant Rodentia Lagostomus maximus 1440 13 0.8292 1.0815086 Bertrand et al. (2019) 10 Extant Rodentia Extant Rodentia Lagostomus maximus Lagostomus maximus 1225 1510 12.7 12.6 0.902758 0.778529 1.19085226 1.01205117 Bertrand et al. (2019) Bertrand et al. (2019) 11 Extant Rodentia Lagostomus maximus 1335 11.9 0.798534 1.04704583 Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 1325 11.9 0.802567 1.05288777 Bertrand et al. (2019) 12 Extant Rodentia Lagostomus maximus 1115 12.3 0.931214 1.236507 Bertrand et al. (2019) 13 Extant Rodentia Lagostomus maximus 1075 11 0.85343 1.13612284 Bertrand et al. (2019) Extant Rodentia Lagostomus maximus 1030 11.2 0.894202 1.19396981 Bertrand et al. (2019) 14 Extant Rodentia Lagostomus maximus 1125 11.2 0.842878 1.1185104 Bertrand et al. (2019) 15 Extant Rodentia Extant Rodentia Lagostomus maximus Lagostomus maximus 1230 3854 11.7 8.8 0.829408 0.290229 1.09378244 0.35333192 Bertrand et al. (2019) Bertrand et al. (2019) 16 Extant Rodentia Lagurus lagurus 15 0.4 0.543122 0.97505099 Bertrand et al. (2019) Extant Rodentia Lariscus insignis 324.71 4.65 0.803933 1.16378115 Bertrand et al. (2019) 17 Extant Rodentia Lemmus lemmus 21 0.7 0.75863 1.33024192 Bertrand et al. (2019) 18 Extant Rodentia Lemmus trimucronatus 32 1.126 0.920252 1.56675928 Bertrand et al. (2019) Extant Rodentia Lemmus trimucronatus 48 1.312 0.817186 1.35235337 Bertrand et al. (2019) 19 Extant Rodentia Lemniscomys striatus 53.3 1.02 0.592259 0.9729643 Bertrand et al. (2019) 20 Extant Rodentia Extant Rodentia Lemniscomys striatus Lemniscomys striatus 53 53.4 1.007 1.014 0.586926 0.588036 0.96458432 0.96590028 Bertrand et al. (2019) Bertrand et al. (2019) 21 Extant Rodentia Lemniscomys striatus 57 0.99 0.549563 0.89859225 Bertrand et al. (2019) 22 Extant Rodentia Extant Rodentia Lophuromys sikapusi Lophuromys sikapusi 63.5 70 1.2 1.13 0.619643 0.546615 1.00554974 0.88100985 Bertrand et al. (2019) Bertrand et al. (2019) 23 Extant Rodentia Macrotarsomys bastardi 28.5 0.8 0.706582 1.21277339 Bertrand et al. (2019) Extant Rodentia Macrotarsomys ingens 65 1.7 0.864203 1.40012853 Bertrand et al. (2019) 24 Extant Rodentia Malacomys longipes 98 1.285 0.496136 0.78103537 Bertrand et al. (2019) 25 Extant Rodentia Malacothrix typicus 124 0.5 0.164891 0.25533646 Bertrand et al. (2019) Extant Rodentia Marmota bobak 5333.00 11.97 0.31761 0.37797396 Bertrand et al. (2019) 26 Extant Rodentia Marmota marmota 4547 14.47 0.427247 0.51415569 Bertrand et al. (2021) 27 Extant Rodentia Extant Rodentia Marmota marmota Marmota marmota 5000.00 4050.00 16.00 17.00 0.44323 0.542341 0.52985501 0.65797149 Bertrand et al. (2019) Bertrand et al. (2019) 28 Extant Rodentia Marmota marmota 2950.00 17.00 0.670636 0.83187007 Bertrand et al. (2019) Extant Rodentia Marmota marmota 3500.00 17.00 0.598056 0.73301597 Bertrand et al. (2019) 29 Extant Rodentia Marmota sibirica 1890.00 18.10 0.962206 1.2313222 Bertrand et al. (2019) 30 Extant Rodentia Mastomys coucha 21.8 0.7132 0.753814 1.31834294 Bertrand et al. (2019) Extant Rodentia Mastomys natalensis 63 0.9 0.4672 0.75858696 Bertrand et al. (2019) 31 Extant Rodentia Meriones crassus 122 1.36 0.453416 0.70292259 Bertrand et al. (2019) 32 Extant Rodentia Meriones libycus 93 1.51 0.603826 0.95405725 Bertrand et al. (2019) Extant Rodentia Meriones meridianus 50 1.2 0.727261 1.20010242 Bertrand et al. (2019) 33 Extant Rodentia Meriones shawi 140 1.48 0.449961 0.69087875 Bertrand et al. (2019) 34 Extant Rodentia Extant Rodentia Meriones unguiculatus Mesocricetus auratus 50 100 1.3 1.4 0.787866 0.53327 1.30011096 0.83830664 Bertrand et al. (2019) Bertrand et al. (2019) 35 Extant Rodentia Mesocricetus auratus 125 1.12 0.367373 0.56856416 Bertrand et al. (2019) Extant Rodentia Mesocricetus auratus 87 1.32 0.551969 0.87620263 Bertrand et al. (2019) 36 Extant Rodentia Mesocricetus brandti 80 1 0.442332 0.70629849 Bertrand et al. (2019) 37 Extant Rodentia Microdipodops megacephalus 13.6 0.47619 0.690444 1.24806379 Bertrand et al. (2019) Extant Rodentia Microdipodops pallidus 12.9 0.50476 0.758246 1.37570417 Bertrand et al. (2019) 38 Extant Rodentia Micromys minutus 5.5 0.267 0.710047 1.36746979 Bertrand et al. (2019) 39 Extant Rodentia Extant Rodentia Micromys minutus Micromys minutus 6.1 5.95 0.282 0.278 0.699676 0.701354 1.33776522 1.34331267 Bertrand et al. (2019) Bertrand et al. (2019) 40 Extant Rodentia Microtus agrestis 42.5 0.9 0.608192 1.01510173 Bertrand et al. (2019) 41 Extant Rodentia Extant Rodentia Microtus agrestris Microtus arvalis arvalis 30.5 20.5 0.532 0.4 0.449003 0.440559 0.76701635 0.77381475 Bertrand et al. (2019) Bertrand et al. (2019) 42 Extant Rodentia Microtus oeconomus stimmingi 45 0.68 0.442258 0.73520173 Bertrand et al. (2019) Extant Rodentia Microtus pennsylvanicus 23.7 0.6606 0.660201 1.14788839 Bertrand et al. (2019) 43 Extant Rodentia Microtus pennsylvanicus 22.9 0.6464 0.661044 1.15212065 Bertrand et al. (2019) 44 Extant Rodentia Microtus pennsylvanicus 25.2 0.7166 0.687317 1.18991317 Bertrand et al. (2019) Extant Rodentia Microtus pennsylvanicus 27.9 0.7394 0.662435 1.13869445 Bertrand et al. (2019) 45 Extant Rodentia Mus minutoides 10.4 0.327 0.567485 1.04524464 Bertrand et al. (2019) 46 Extant Rodentia Extant Rodentia Mus minutoides Mus musculus 5 18 0.26 0.43 0.737025 0.516719 1.42892822 0.91588653 Bertrand et al. (2019) Bertrand et al. (2019) 47 Extant Rodentia Mus musculus 24 0.45 0.445954 0.77469555 Bertrand et al. (2019) Extant Rodentia Mus musculus 20.85 0.43 0.468259 0.82149484 Bertrand et al. (2019) 48 Extant Rodentia Mus musculus 16 0.36 0.468124 0.83662065 Bertrand et al. (2019) 49 Extant Rodentia Mus musculus 24.32 0.475 0.46657 0.80975835 Bertrand et al. (2019) Extant Rodentia Mus musculus 24 0.5 0.495504 0.86077284 Bertrand et al. (2019) 50 Extant Rodentia Mus musculus 18.92 0.424 0.492774 0.87040041 Bertrand et al. (2019) 51 Extant Rodentia Mus musculus domesticus 12 0.36 0.567636 1.03510224 Bertrand et al. (2019) Extant Rodentia Mus musculus musculus 20 0.5 0.559885 0.98510535 Bertrand et al. (2019) 52 Extant Rodentia Mus triton 8.4 0.36 0.720865 1.34775421 Bertrand et al. (2019) 53 Extant Rodentia Extant Rodentia Muscardinus avellanarius Myocastor coypus 13 3300 0.5 23 0.747217 0.841671 1.35496099 1.03586316 Bertrand et al. (2019) Bertrand et al. (2019) 54 Extant Rodentia Myocastor coypus 3380 19 0.684224 0.84067883 Bertrand et al. (2019) Extant Rodentia Myocastor coypus 5000 23 0.637143 0.76166658 Bertrand et al. (2019) 55 Extant Rodentia Myocastor coypus 5300 18.7 0.498191 0.59313333 Bertrand et al. (2019) 56 Extant Rodentia Myocastor coypus 3300 21 0.768482 0.94578811 Bertrand et al. (2019) Extant Rodentia Myocastor coypus 7450 15.64 0.331674 0.38558131 Bertrand et al. (2019) 57 Extant Rodentia Myocastor coypus 7530 15.6 0.328466 0.38156733 Bertrand et al. (2019) 58 Extant Rodentia Extant Rodentia Myocastor coypus Myocastor coypus 8475 5510 17.55 17.43 0.341383 0.452423 0.39330422 0.53718029 Bertrand et al. (2019) Bertrand et al. (2019) 59 Extant Rodentia Myocastor coypus 3800 14.77 0.49175 0.59926029 Bertrand et al. (2019) 60 Extant Rodentia Extant Rodentia Myocastor coypus Myocastor coypus 5700 5000 18.35 23 0.465606 0.637143 0.55152289 0.76166658 Bertrand et al. (2019) Bertrand et al. (2019) Extant Rodentia Myocastor coypus f domestica 3500 18.4 0.647308 0.79338199 Bertrand et al. (2019) Extant Rodentia Myodes glareolus 35 0.593 0.456402 0.77218117 Bertrand et al. (2019) Page 76 of 78 1 Extant Rodentia Myodes glareolus 20 0.7 0.783839 1.37914749 Bertrand et al. (2019) 2 Extant Rodentia Myoprocta pratti 780 9.9 0.952249 1.29646279 Bertrand et al. (2019) 3 Extant Rodentia Extant Rodentia Mystromys albicaudatus Neotamias minimus 130 37.05 1.47 1.45 0.469671 1.073591 0.72489336 1.80917233 Bertrand et al. (2019) Bertrand et al. (2019) 4 Extant Rodentia Oenomys hypoxanthus 92 1.15 0.463211 0.73243606 Bertrand et al. (2019) Extant Rodentia Oenomys hypoxanthus 92 1.166 0.469656 0.74262647 Bertrand et al. (2019) 5 Extant Rodentia Oenomys hypoxanthus 92 1.162 0.468044 0.74007887 Bertrand et al. (2019) 6 Extant Rodentia Oenomys hypoxanthus 178 1.48 0.383088 0.57839667 Bertrand et al. (2019) Extant Rodentia Ondatra zibethicus 1500 4.8 0.297906 0.38744367 Bertrand et al. (2019) 7 Extant Rodentia Ondatra zibethicus 1500 7.6 0.471684 0.61345248 Bertrand et al. (2019) 8 Extant Rodentia Extant Rodentia Ondatra zibethicus Ondatra zibethicus 1600 1032 8.1 5.7 0.481441 0.454494 0.62332006 0.6067737 Bertrand et al. (2019) Bertrand et al. (2019) 9 Extant Rodentia Ondatra zibethicus 900 5.33 0.465804 0.62785972 Bertrand et al. (2019) 10 Extant Rodentia Extant Rodentia Ondatra zibethicus Orthogeomys heterodus 900 630 5.33 3.9619 0.465804 0.439708 0.62785972 0.60766874 Bertrand et al. (2019) Bertrand et al. (2019) 11 Extant Rodentia Orthogeomys hispidus 542.1 3.65714 0.448877 0.62689966 Bertrand et al. (2019) Extant Rodentia Otomys irroratus 57.7 0.97143 0.534862 0.87380724 Bertrand et al. (2019) 12 Extant Rodentia Otomys irroratus 66 1.62 0.815153 1.31925081 Bertrand et al. (2019) 13 Extant Rodentia Paraxerus cepapi 138.13 2.91 0.892084 1.37101409 Bertrand et al. (2019) Extant Rodentia Paraxerus cepapi 223.80 3.22 0.714727 1.06195425 Bertrand et al. (2019) 14 Extant Rodentia Paraxerus poensis 100.00 2.73 1.041145 1.63669391 Bertrand et al. (2019) 15 Extant Rodentia Extant Rodentia Pelomys campanae Pelomys fallax 91 121 1.23 1.46 0.499075 0.489447 0.78974948 0.7592181 Bertrand et al. (2019) Bertrand et al. (2019) 16 Extant Rodentia Pelomys fallax 110 1.406 0.502425 0.78456621 Bertrand et al. (2019) Extant Rodentia Pelomys fallax 111 1.412 0.501519 0.78265536 Bertrand et al. (2019) 17 Extant Rodentia Perognathus flavus 8 0.28571 0.591126 1.10897111 Bertrand et al. (2019) 18 Extant Rodentia Perognathus longimembris 8 0.29524 0.610831 1.14593682 Bertrand et al. (2019) Extant Rodentia Perognathus parvus 17.3 0.44762 0.552378 0.98181421 Bertrand et al. (2019) 19 Extant Rodentia Perognathus parvus 17 0.58 0.724179 1.28875588 Bertrand et al. (2019) 20 Extant Rodentia Extant Rodentia Peromyscus californicus Peromyscus californicus 55 51.2 1.0292 1.0295 0.585161 0.614093 0.95919356 1.011676 Bertrand et al. (2019) Bertrand et al. (2019) 21 Extant Rodentia Peromyscus eremicus 20.1 0.6506 0.726092 1.27709688 Bertrand et al. (2019) 22 Extant Rodentia Extant Rodentia Peromyscus eremicus Peromyscus gossypinus 20 27.2 0.6416 0.68 0.718444 0.619679 1.26408718 1.06709428 Bertrand et al. (2019) Bertrand et al. (2019) 23 Extant Rodentia Peromyscus leucopus 25.5 0.63 0.599484 1.03699267 Bertrand et al. (2019) Extant Rodentia Peromyscus leucopus 19.7 0.6372 0.720779 1.26953778 Bertrand et al. (2019) 24 Extant Rodentia Peromyscus leucopus 22.9 0.6221 0.636193 1.10880918 Bertrand et al. (2019) 25 Extant Rodentia Peromyscus maniculatus bairdii 16.5 0.5182 0.660088 1.17715637 Bertrand et al. (2019) Extant Rodentia Peromyscus maniculatus bairdii 18 0.5188 0.623428 1.10502774 Bertrand et al. (2019) 26 Extant Rodentia Peromyscus maniculatus gracilis 22.1 0.6885 0.721074 1.25987811 Bertrand et al. (2019) 27 Extant Rodentia Extant Rodentia Peromyscus maniculatus gracilis Peromyscus polionotus 26 13.5 0.6909 0.4903 0.648936 0.714426 1.12101077 1.2920811 Bertrand et al. (2019) Bertrand et al. (2019) 28 Extant Rodentia Peromyscus polionotus 13.1 0.4867 0.723616 1.31146123 Bertrand et al. (2019) Extant Rodentia Petaurista petaurista 1096.65 11.73 0.898077 1.19389145 Bertrand et al. (2019) 29 Extant Rodentia Petinomys setosus 41.86 1.44 0.983395 1.64307785 Bertrand et al. (2019) 30 Extant Rodentia Podomys floridanus 36.5 0.8896 0.665698 1.12298233 Bertrand et al. (2019) Extant Rodentia Podomys floridanus 39.2 0.8917 0.636115 1.06773123 Bertrand et al. (2019) 31 Extant Rodentia Praomys jacksoni 49 0.87 0.534449 0.88317959 Bertrand et al. (2019) 32 Extant Rodentia Protoxerus aubinni 525.00 8.03 1.006816 1.40927342 Bertrand et al. (2019) Extant Rodentia Protoxerus stangeri 767.23 9.28 0.90281 1.23057362 Bertrand et al. (2019) 33 Extant Rodentia Protoxerus stangeri 690.60 10.25 1.069449 1.46848724 Bertrand et al. (2019) 34 Extant Rodentia Extant Rodentia Pteromys buechneri Pteromys nitidus 106.37 1600.00 2.22 11.80 0.81127 0.701359 1.26982654 0.9080465 Bertrand et al. (2019) Bertrand et al. (2019) 35 Extant Rodentia Pteromyscus pulverulentus 195.44 3.45 0.838463 1.25767647 Bertrand et al. (2019) Extant Rodentia Rattus norvegicus 274 2.18 0.422651 0.6191501 Bertrand et al. (2019) 36 Extant Rodentia Rattus norvegicus 278 2.3 0.441608 0.64626339 Bertrand et al. (2019) 37 Extant Rodentia Rattus norvegicus 197 1.61 0.389361 0.58370812 Bertrand et al. (2019) Extant Rodentia Rattus norvegicus 291 2.27 0.422704 0.61662328 Bertrand et al. (2019) 38 Extant Rodentia Rattus norvegicus 305 2.36 0.425843 0.61916299 Bertrand et al. (2019) 39 Extant Rodentia Extant Rodentia Rattus norvegicus Rattus norvegicus domestica 448 324 2.36 2.7 0.329135 0.467862 0.46584468 0.67738461 Bertrand et al. (2019) Bertrand et al. (2019) 40 Extant Rodentia Rattus norvegicus norvegicus 202 2.6 0.618311 0.92531202 Bertrand et al. (2019) 41 Extant Rodentia Extant Rodentia Rattus rattus Rattus rattus 160 200 2.24 1.59 0.622739 0.38065 0.94727069 0.57004586 Bertrand et al. (2019) Bertrand et al. (2019) 42 Extant Rodentia Rattus rattus 150 1.92 0.557364 0.85166454 Bertrand et al. (2019) Extant Rodentia Rattus rattus alexandrinus 150 1.57 0.455761 0.69641319 Bertrand et al. (2019) 43 Extant Rodentia Rattus rattus rattus 217 1.68 0.380803 0.56702727 Bertrand et al. (2019) 44 Extant Rodentia Ratufa affinis 1074.27 11.73 0.910274 1.21185368 Bertrand et al. (2019) Extant Rodentia Ratufa bicolor 1440.00 12.00 0.765415 0.99831563 Bertrand et al. (2019) 45 Extant Rodentia Ratufa bicolor 1400.00 12.00 0.779999 1.0193454 Bertrand et al. (2019) 46 Extant Rodentia Extant Rodentia Ratufa indica Ratufa indica 1935.00 1010.00 11.40 11.60 0.596551 0.938386 0.76214184 1.25468597 Bertrand et al. (2019) Bertrand et al. (2019) 47 Extant Rodentia Rhinosciurus laticaudatus 507.38 4.17 0.535657 0.75157065 Bertrand et al. (2019) Extant Rodentia Sciurus carolinensis 592.55 7.67 0.886828 1.23084865 Bertrand et al. (2019) 48 Extant Rodentia Sciurus carolinesis 469.00 7.58 1.025185 1.44635923 Bertrand et al. (2019) 49 Extant Rodentia Sciurus carolinesis 466.00 7.48 1.016019 1.43407177 Bertrand et al. (2019) Extant Rodentia Sciurus carolinesis 535.00 7.53 0.932431 1.303431 Bertrand et al. (2019) 50 Extant Rodentia Sciurus granatensis 336.99 6.02 1.01647 1.46763308 Bertrand et al. (2019) 51 Extant Rodentia Sciurus niger 365.00 6.90 1.103905 1.5849926 Bertrand et al. (2019) Extant Rodentia Sciurus niger 365.00 6.50 1.03991 1.49310897 Bertrand et al. (2019) 52 Extant Rodentia Sciurus niger 703.00 10.20 1.051862 1.44254078 Bertrand et al. (2019) 53 Extant Rodentia Extant Rodentia Sciurus niger Sciurus niger 770.00 650.00 10.50 9.20 1.01873 0.999896 1.38822874 1.37881758 Bertrand et al. (2019) Bertrand et al. (2019) 54 Extant Rodentia Sciurus niger 580.00 8.95 1.049893 1.4593558 Bertrand et al. (2019) Extant Rodentia Sciurus niger cinereus 328.00 7.20 1.237416 1.79003169 Bertrand et al. (2019) 55 Extant Rodentia Sciurus vulgaris 350.00 7.50 1.234112 1.77715693 Bertrand et al. (2019) 56 Extant Rodentia Sciurus vulgaris 323.00 6.10 1.059212 1.53389219 Bertrand et al. (2019) Extant Rodentia Sciurus vulgaris 287.00 5.81 1.091978 1.59447751 Bertrand et al. (2019) 57 Extant Rodentia Sciurus vulgaris 316.00 5.89 1.037872 1.50529536 Bertrand et al. (2019) 58 Extant Rodentia Extant Rodentia Sciurus vulgaris Sciurus vulgaris 327.00 361.00 6.23 5.85 1.072901 0.942855 1.55237835 1.35480068 Bertrand et al. (2019) Bertrand et al. (2019) 59 Extant Rodentia Sciurus vulgaris 389.00 6.00 0.919822 1.31481126 Bertrand et al. (2019) 60 Extant Rodentia Extant Rodentia Sigmodon hispidus Spalax leucodon 148 122 1.18 3 0.345641 1.000182 0.52864388 1.55056454 Bertrand et al. (2019) Bertrand et al. (2019) Extant Rodentia Spalax leucodon 180 2.63 0.675681 1.01936323 Bertrand et al. (2019) Extant Rodentia Spalax leucodon 214 1.9 0.434705 0.64792133 Bertrand et al. (2019) Page 77 of 78 1 Extant Rodentia Spermophilus citellus 290.00 2.58 0.481539 0.70261927 Bertrand et al. (2019) 2 Extant Rodentia Spermophilus suslicus 224.00 2.30 0.510364 0.75826179 Bertrand et al. (2019) 3 Extant Rodentia Extant Rodentia Stochomys longicaudatus Tachyoryctes splendens 65 282 1.27 2.1 0.64561 1.04597837 0.399366 0.58386148 Bertrand et al. (2019) Bertrand et al. (2019) 4 Extant Rodentia Tachyoryctes splendens 174 2.29 0.601847 0.910131 Bertrand et al. (2019) Extant Rodentia Tachyoryctes splendens 206 2.4 0.563299 0.84182994 Bertrand et al. (2019) 5 Extant Rodentia Tachyoryctes splendens 218 2.24 0.506175 0.75346847 Bertrand et al. (2019) 6 Extant Rodentia Tamias striatus 80.00 2.70 1.194297 1.90700593 Bertrand et al. (2019) Extant Rodentia Tamias striatus 97.00 2.90 1.127404 1.77607697 Bertrand et al. (2019) 7 Extant Rodentia Tamias striatus 75.00 2.22 1.02537 1.64468415 Bertrand et al. (2019) 8 Extant Rodentia Extant Rodentia Tamias striatus Tamiasciurus hudsonicus 62.00 256.61 2.31 4.90 1.212072 1.97023032 0.99299 1.46134237 Bertrand et al. (2019) Bertrand et al. (2019) 9 Extant Rodentia Tamiasciurus hudsonicus 125.00 4.80 1.574455 2.43670355 Bertrand et al. (2019) 10 Extant Rodentia Extant Rodentia Tamiasciurus hudsonicus Tamiasciurus hudsonicus 159.00 248.00 4.10 5.02 1.144634 1.74190655 1.040495 1.53491504 Bertrand et al. (2019) Bertrand et al. (2019) 11 Extant Rodentia Tamiasciurus hudsonicus 183.00 4.71 1.196733 1.80335808 Bertrand et al. (2019) Extant Rodentia Tamiasciurus hudsonicus 169.00 4.21 1.12828 1.70970351 Bertrand et al. (2019) 12 Extant Rodentia Tamiasciurus hudsonicus 183.00 3.97 1.008712 1.52002793 Bertrand et al. (2019) 13 Extant Rodentia Tamiasciurus hudsonicus 159.00 4.80 1.340059 2.03930523 Bertrand et al. (2019) Extant Rodentia Thamnomys venustus 82.5 1.291 0.559398 0.89130263 Bertrand et al. (2019) 14 Extant Rodentia Thamnomys venustus 82.5 1.284 0.556365 0.88646985 Bertrand et al. (2019) 15 Extant Rodentia Extant Rodentia Thamnomys venustus Thomomys bottae 82.5 185.2 1.35 0.584963 0.93203606 1.40952 0.355281 0.53492508 Bertrand et al. (2019) Bertrand et al. (2019) 16 Extant Rodentia Thomomys talpoides 90.6 1.00952 0.410827 0.65030421 Bertrand et al. (2019) Extant Rodentia Thryonomys gregorianus 3500 13.15 0.462614 0.56700941 Bertrand et al. (2019) 17 Extant Rodentia Thryonomys gregorianus 3500 13.15 0.462614 0.56700941 Bertrand et al. (2019) 18 Extant Rodentia Thryonomys swinderianus 4500 12 0.356737 0.42961442 Bertrand et al. (2019) Extant Rodentia Urocitellus columbianus 482.00 3.52 0.467939 0.65891969 Bertrand et al. (2019) 19 Extant Rodentia Urocitellus columbianus 529.00 3.57 0.4456 0.62338973 Bertrand et al. (2019) 20 Extant Rodentia Extant Rodentia Urocitellus parryii Urocitellus parryii 878.00 958.00 5.63 5.74 0.500249 0.6754564 0.481078 0.64561897 Bertrand et al. (2019) Bertrand et al. (2019) 21 Extant Rodentia Urocitellus parryii 482.00 3.95 0.524851 0.73905857 Bertrand et al. (2019) 22 Extant Rodentia Extant Rodentia Urocitellus parryii Urocitellus richardsonii 756.00 354.00 4.98 3.08 0.48924 0.6675459 0.502342 0.72281254 Bertrand et al. (2019) Bertrand et al. (2019) 23 Extant Rodentia Urocitellus richardsonii 361.00 3.31 0.534169 0.76755496 Bertrand et al. (2019) Extant Rodentia Urocitellus richardsonii 246 3.01 0.627154 0.92574645 Bertrand et al. (2021) 24 Extant Rodentia Xerus inauris 638.00 6.41 0.705366 0.97394253 Bertrand et al. (2019) 25 Extant Rodentia Xerus rutilus 354 5.72 0.933601 1.34337023 Bertrand et al. (2021) Extant Rodentia Zapus hudsonius 19.3 0.6961 0.798302 1.40810193 Bertrand et al. (2019) 26 Extant Rodentia Zapus hudsonius 15.2 0.705 0.948796 1.70176556 Bertrand et al. (2019) 27 Extant Rodentia Zygogeomys trichopus 545 3.62857 0.443781 0.6195511 Bertrand et al. (2019) Table S3. Encephalization quotients, brain mass, and body mass for Anagale gobiensis, living and fossil lagomorphs, living and fossil rodents, plesiadapiforms, fossil euprimates, and apatemyids in Figure 14. 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 78 of 78 1 2 Group Species Specimen number NS*/ OV/T PV2/T Source TS V V 3 Euprimates Notharctus tenebrosus AMNH 127167 30.88 2.1 1.41 Harrington et al. (2016); Bertrand et al. 2022 4 Euprimates Notharctus tenebrosus USNM V 23277 30.43 2.23 0.97 Harrington et al. (2016); Bertrand et al. 2022 5 Euprimates Notharctus tenebrosus USNM V 23278 29.42 1.51 NA Harrington et al. (2016); Bertrand et al. 2022 Euprimates Smilodectes gracilis USNM V 17994 30.59 2.06 NA Harrington et al. (2016); Bertrand et al. 2022 6 Euprimates Smilodectes gracilis USNM V 17996 30.85 1.67 1.24 Harrington et al. (2016); Bertrand et al. 2022 7 Euprimates Smilodectes gracilis USNM V 21815 31.37 1.24 1.61 Harrington et al. (2016); Bertrand et al. 2022 Euprimates Smilodectes gracilis UM 32773 (=MPM 2612) 32.84 1.54 1.46 Harrington et al. (2016); Bertrand et al. 2022 8 Euprimates Adapis parisiensis NHM M 1345 30.93 2.4 0.51 Harrington et al. (2016); Bertrand et al. 2022 9 Euprimates Rooneyia viejaensis TMM 40688-7 43.77 0.94 0.84 Kirk et al. (2014); Ramdarshan & Orliac (2016); Bertrand et al. 2022 Euprimates Microchoerus erinaceus UM-PRR 1771 41.34 0.96 1.13 Ramdarshan & Orliac (2016); Bertrand et al. 2022 10 Euprimates Necrolemur antiquus MaPhQ 289 NA 1.94 1.09 Harrington et al. (2020); Bertrand et al. 2022 11 Plesiadapiformes Microsyops annectens UW 12362 21.40 5.09 - Silcox et al. (2010); Long et al. 2015 12 Plesiadapiformes Ignacius graybullianus USNM 421608 19.72 5.53 - Silcox et al. (2009b); Long et al. 2015 Plesiadapiformes Plesiadapis cookei UM 87990 - 7.8 - Gingerich & Gunnell (2005) 13 Plesiadapiformes Plesiadapis tricuspidens MNHN CR 125 19.93 4.9 - Orliac et al. (2014) 14 Fossil Aplodontiidae Altasciurus relictus USNM 437793 29.69 3.55 3.35 Bertrand et al. (2019; 2021) Fossil Aplodontiidae Mesogaulus paniensis AMNH F:AM 65511 26.49 3.29 1.33 Bertrand et al. (2018) 15 Fossil Sciuridae Protosciurus cf. rachelae YPM 14736 30.59 4.76 3.31 Bertrand et al. (2019) 16 Fossil Sciuridae Protosciurus cf. rachelae YPM 14737 30.95 3.65 2.96 Bertrand et al. (2019) Fossil Sciuridae Cedromus wilsoni USNM 256584 29.59 2.96 3.16 Bertrand et al. (2019) 17 Ischyromyidae Paramys copei AMNH 4756 17.10 6.05 1.20 Bertrand et al. (2019) 18 Ischyromyidae Paramys delicatus AMNH 12506 16.25 4.74 1.03 Bertrand et al. (2019) Ischyromyidae Pseudotomus horribilis USNM 17159 18.89 5.33 1.14 Bertrand et al. (2019) 19 Ischyromyidae Pseudotomus robustus USNM 17161 22.92 5.94 0.62 Bertrand et al. (2019) 20 Ischyromyidae Pseudotomus petersoni AMNH 2018 22.89 4.14 0.39 Bertrand et al. (2019) Ischyromyidae Pseudotomus hians AMNH 5025 23.02 5.43 1.04 Bertrand et al. (2019) 21 Ischyromyidae Reithroparamys delicatissimus AMNH 12561 17.33 3.21 2.31 Bertrand et al. (2019; 2021) 22 Ischyromyidae Rapamys atramontis AMNH 128706 20.24 3.16 1.52 Bertrand et al. (2019) 23 Ischyromyidae Rapamys atramontis AMNH 128704 21.77 3.76 2.05 Bertrand et al. (2019) Ischyromyidae Ischyromys typus ROMV 1007 21.18 3.23 1.63 Bertrand et al. (2019) 24 Ischyromyidae Ischyromys typus AMNH 12252 18.45 3.68 - Bertrand et al. (2019) 25 Ischyromyidae Ischyromys typus AMNH F:AM 144638 23.03 3.15 1.60 Bertrand et al. (2019) Extant Aplodontiidae Aplodontia rufa AMNH 42389 25.94 2.58 0.82 Bertrand et al. (2018) 26 Extant Sciuridae Sciurus carolinensis AMNH 42389 35.41 3.18 2.03 Bertrand et al. (2019) 27 Extant Sciuridae Sciurus granatensis AMNH 42389 35.39 2.69 2.08 Bertrand et al. (2019) Extant Sciuridae Tamiasciurus hudsonicus USNM 549146 36.30 2.50 2.33 Bertrand et al. (2019) 28 Extant Sciuridae Eutamias minimus USNM 298500 34.93 3.36 2.29 Bertrand et al. (2019) 29 Extant Sciuridae Funisciurus pyrropus USNM 294865 36.44 3.22 2.19 Bertrand et al. (2019) Extant Sciuridae Heliosciurus rufobrachium USNM 378091 35.85 2.45 1.87 Bertrand et al. (2019) 30 Extant Sciuridae Paraxerus cepapi USNM 367956 35.62 2.62 1.46 Bertrand et al. (2019) 31 Extant Sciuridae Protoxerus stangeri USNM 435027 38.02 2.31 1.89 Bertrand et al. (2019) Extant Sciuridae Aeromys tephromelas USNM 481190 35.19 2.85 1.45 Bertrand et al. (2019) 32 Extant Sciuridae Glaucomys volans AMNH 240290 35.69 3.49 1.68 Bertrand et al. (2019) 33 Extant Sciuridae Petaurista petaurista USNM 589079 35.28 1.64 1.62 Bertrand et al. (2019) Extant Sciuridae Hylopetes spadiceus USNM 488639 36.73 3.30 0.85 Bertrand et al. (2019) 34 Extant Sciuridae Petinomys setosus USNM 488674 35.76 3.10 - Bertrand et al. (2019) 35 Extant Sciuridae Pteromyscus pulverulentus USNM 481178 33.82 2.81 1.37 Bertrand et al. (2019) 36 Extant Sciuridae Pteromys buechneri USNM 172622 34.11 1.75 1.61 Bertrand et al. (2019) Extant Sciuridae Rhinosciurus laticaudatus USNM 488511 34.10 3.88 2.22 Bertrand et al. (2019) 37 Extant Sciuridae Callosciurus sp. USNM 294865 38.79 3.28 1.77 Bertrand et al. (2019) 38 Extant Sciuridae Lariscus insignis USNM 488570 33.88 4.73 2.32 Bertrand et al. (2019) Extant Sciuridae Dremomys rufigenis USNM 488602 36.82 3.96 2.14 Bertrand et al. (2019) 39 Extant Sciuridae Ratufa affinis USNM 488104 37.09 1.64 1.91 Bertrand et al. (2019) 40 Extant Sciuridae Xerus rutilus AMNH 179092 38.52 3.28 1.87 Bertrand et al. (2021) Extant Sciuridae Marmota marmota AMNH 146619 33.18 2.57 0.84 Bertrand et al. (2021) 41 Extant Sciuridae Cynomys ludovicianus AMNH114522 34.83 2.53 0.98 Bertrand et al. (2021) 42 Extant Sciuridae Urocitellus richardsonii AMNH 15062 36.57 3.35 1.75 Bertrand et al. (2021) Leporidae Brachylagus idahoensis AMNH 92869 32.01 2.80 2.97 López-Torres et al. (2020) 43 Leporidae Lepus americanus phaeonotus AMNH 51052 35.11 3.58 1.97 López-Torres et al. (2020) 44 Leporidae Lepus arcticus AMNH 97648 41.98 2.55 1.69 López-Torres et al. (2020) Leporidae Lepus americanus bairdii AMNH 42139 31.84 2.76 2.42 López-Torres et al. (2020) 45 Leporidae Oryctolagus cuniculus AMNH 99352 35.21 2.33 2.21 López-Torres et al. (2020) 46 Leporidae Poelagus marjorita AMNH 34816 33.23 3.04 1.79 López-Torres et al. (2020) Leporidae Romerolagus diazi ? 28.08 3.18 2.37 López-Torres et al. (2020) 47 Ochotonidae Ochotona pallasi AMNH 120698 33.71 2.71 2.77 López-Torres et al. (2020) 48 Ochotonidae Ochotona princeps princeps AMNH 59712 30.97 2.68 3.29 López-Torres et al. (2020) 49 Ochotonidae Ochotona princeps schisticeps Stem lagomorph Megalagus turgidus AMNH 40547 FMNH UC 1642 31.84 2.49 18.98 3.97 2.90 2.31 López-Torres et al. (2020) López-Torres et al. (2020) 50 Apatemyide Labidolemur kayi (low end of range) USNM 530208/530221 - 12.18 - Silcox et al. (2011) 51 Apatemyide Labidolemur kayi (high end of range) USNM 530208/530221 - 14.75 - Silcox et al. (2011) Anagalidae Anagale gobiensis AMNH 26079 10.98 7.15 1.01 This paper 52 Table S4. Data for boxplots in Figures 6, 10, 12. NS*, neocortical surface (calculation of NS varies among taxa; taxa coming from Bertrand et al. [2022] include the superior sagittal sinus, whereas the other valu 53 54 55 56 57 58 59 60