(a) Reconstruction of the skeleton based in the partial skeleton IPS56468 from Abocador de Can Mata. Missing elements are based on extant giant flying squirrel Petaurista petaurista and are colored …
(a–b) Right humerus (IPS56468f) in cranial and caudal views. (c–d) Right femur (IPS56468b) in cranial and caudal views. (e–f) Right tibia (IPS56468a) in cranial and caudal views. (g–h) Lumbar …
Note that some elements, such as the femora, pelvis and tail vertebrae are in anatomical connection. This scheme has been drawn based on multiple pictures taken during the preparation process. Not …
(a to c) Partial left hemimandible (IPS56468j) in lateral, medial and dorsal views. (d to e) Partial right hemimandible (IPS56468i) in lateral and medial views. A caudal vertebra and a bone fragment …
Articulated bones are shown on top and disarticulated ones are shown below. (a) Miopetaurista neogrivensis. (b) Petaurista petaurista, large-sized flying squirrel, subtribe Pteromyina. (c) Hylopetes …
All elements are scaled to femur length and shown in anterior view. Humerus (a–e) and articulated femur and tibia (f–j) of: (a,f) the xerin ground squirrel Xerus erythropus; (b,g) the callosciurin …
(a to e) Proximal end of the humerus in lateral view. (f to j) Distal end of the humerus in anterior view. (k to o) proximal end of the femur in anterior view. (p to s) Proximal end of the tibia in …
Virtual reconstruction based on µCT scan data from specimens IPS56468h (see Figure 6—figure supplement 1) and IPS88677 (see Figure 6—figure supplement 2). Specimen IPS56468h was used as the basis …
The partial skull (IPS56468h) belongs to the same individual as the partial skeleton IPS56468 from Abocador de Can Mata locality ACM/C5-D1. Fossil specimens are shown on the left, whereas …
Fossil specimens are shown on the left, whereas tridimensional models based on µCT scan data are shown on the right. (a) Dorsal view. (b) Ventral view. (c) Lateral (left) view. (d) Lateral (right) …
The analysis is based on 38 taxa, 105 morphological characters and 3345 base pairs (see Materials and Methods and Appendix 1.1). Purple circles at the nodes indicate posterior probabilities higher …
Based on 58 extant taxa and 3345 base pairs (see Materials and Methods and Appendix 1.2). The extinct Miopetaurista neogrivensis and Douglassciurus jeffersoni are used as calibration points. Purple …
Temporal ranges of purported flying squirrel genera in Europe, Asia and North America. The 95% highest posterior density (HPD) intervals for flying squirrel divergence as derived from total evidence …
(a) Schematic geological map of the Vallès-Penedès Basin (Catalonia, Spain) showing the area enlarged in Figure 9b. The inset shows its location within the Iberian Peninsula. (b) Situation of the …
The squirrel is shown reducing speed just before landing on a tree branch. Skeleton reconstruction based in the partial skeleton IPS56468 from Abocador de Can Mata. Coat pattern and color are based …
Miopetaurista neogrivensis is compared to Petaurista petaurista (large-sized flying squirrel; subtribe Pteromyina); Hylopetes sagitta (small-sized flying squirrel; subtribe Glaucomyina); and Sciurus …
Virtual reconstruction based on µCT scan data from specimens IPS56468h and IPS88677. Specimen IPS56468h was used as the basis for the reconstruction, with missing elements taken from IPS88677. A …
Middle ear cavity anatomy in Miopetaurista neogrivensis (left; IPS88677) and the extant giant flying squirrel Petaurista petaurista (right; ZMA13418) reconstructed from µCT scan data. Note the …
This list includes the catalogue numbers (preceded by the acronym ‘IPS’) of the various bones and bone fragments belonging to the partial skeleton of a single individual of Miopetaurista …
Catalogue no. | Region | Description |
---|---|---|
IPS56468a | leg | right tibia |
IPS56468b | leg | right femur |
IPS56468c | leg | left femur |
IPS56468d | leg | distal half of the left tibia |
IPS56468e | arm | left humerus |
IPS56468f | arm | right humerus |
IPS56468g | arm | distal half of the right radius, with damaged epiphysis |
IPS56468h | cranium | almost complete cranium, laterally compressed in an oblique angle |
IPS56468i | cranium + tail | partial right mandible (angular process broken), caudal vertebra (probably corresponding to the mid part of the tail) |
IPS56468j | cranium | partial left mandible (articular process broken, all other processes with minor damage) |
IPS56468k | pelvic girdle | partial right coxal (proximal end of the ilium and part of the pubis damaged) |
IPS56468l | pelvic girdle | partial left coxal (missing most of the pubis and ischium, extensive damage in the ilium) |
IPS56468m | trunk | lumbar vertebrae (L3–L4) in anatomical connection |
IPS56468n | trunk | lumbar vertebrae (L5–L6) in anatomical connection |
IPS56468o | neck | partial axis (only part of the vertebral body is preserved) and partial cervical vertebra (C3) |
IPS56468p | trunk | thoracic vertebra (T1?) |
IPS56468q | tail | four caudal vertebrae (mid part of the tail) that articulate with one another |
IPS56468r | trunk | seven rib fragments |
IPS56468s | ankle | left calcaneus and left navicular |
IPS56468t | ankle | left astragalus |
IPS56468u | trunk | three partial thoracic vertebrae (T2–T4?) that articulate with one another |
IPS56468v | indeterminate | associated bone fragments (may not belong to M. neogrivensis) |
IPS56468w | indeterminate | associated bone fragments (may not belong to M. neogrivensis) |
IPS56468x | foot? | six distal phalanges which are not assigned to any particular ray or side; attribution to the foot is tentative |
IPS56468y | trunk | two sternebrae that articulate with one another |
IPS56468z | foot | left metatarsals 2–4 in anatomical connection |
IPS56468aa | thoracic girdle | right clavicle (with minor damage in its acromial end) and partial left clavicle (acromial end missing) |
IPS56468ab | foot | complete right metatarsal 3, and partial rigth metatarsals 4 (proximal end missing), 2 and 4 (only distal half preserved) |
IPS56468ac | foot | four proximal phalanges and one partial proximal phalanx (distal half); they are not assigned to any particular ray or side |
IPS56468ad | foot? | seven intermediate phalanges and five fragments; they are not assigned to any particular ray or side and attribution to the foot is tentative |
IPS56468ae | hand | four proximal phalanges; they are not assigned to any particular ray or side |
IPS56468af | ankle | left intermediate and medial cuneiform |
IPS56468ag | ankle | right navicular |
IPS56468ah | wrist | right scapholunate and dorsal end of the right pisiform |
This list includes the additional material of Miopetaurista neogrivensis from locality ACM/C5-D1 and the approximately stratigraphicaly equivalent localities ACM/C8-Af and ACM/C6-A5, as well as from …
Catalogue no. | Locality | Anatomical element |
---|---|---|
IPS43480 | ACM/C5-D1 | L m1 |
IPS43505 | ACM/C5-D1 | L m3 |
IPS43675 | ACM/C5-D1 | R P4 |
IPS43677 | ACM/C5-D1 | partial left mandible with p4–m3 (only part of the mandibular body preserved) |
IPS43724 | ACM/C5-D1 | right maxillary fragment with partial P4-M2 |
IPS77856 | ACM/C5-D1 | R P4 |
IPS77857 | ACM/C5-D1 | R P4 |
IPS77858 | ACM/C5-D1 | R M1/M2 |
IPS77859 | ACM/C5-D1 | R M1/M2 |
IPS77860 | ACM/C5-D1 | fragment of R M1/M2 |
IPS77861 | ACM/C5-D1 | fragment of R M1/M2 |
IPS77862 | ACM/C5-D1 | L M1/M2 |
IPS77863 | ACM/C5-D1 | broken L M1/M2 |
IPS77864 | ACM/C5-D1 | fragment of L M1/M2 |
IPS77865 | ACM/C5-D1 | fragment of L M1/M2 |
IPS77866 | ACM/C5-D1 | L M3 |
IPS77867 | ACM/C5-D1 | R M3 |
IPS77868 | ACM/C5-D1 | L dp4 |
IPS77869 | ACM/C5-D1 | fragment of L dp4 |
IPS77870 | ACM/C5-D1 | R dp4 |
IPS77871 | ACM/C5-D1 | L p4 |
IPS77872 | ACM/C5-D1 | R p4 |
IPS77873 | ACM/C5-D1 | R p4 |
IPS77874 | ACM/C5-D1 | L m1 |
IPS77875 | ACM/C5-D1 | L m1 |
IPS77876 | ACM/C5-D1 | L m1 |
IPS77877 | ACM/C5-D1 | L m2 |
IPS77878 | ACM/C5-D1 | L m2 |
IPS77879 | ACM/C5-D1 | broken L m1/m2 |
IPS77880 | ACM/C5-D1 | abraded L m1/m2 |
IPS77881 | ACM/C5-D1 | fragment of L m1/m2 |
IPS77882 | ACM/C5-D1 | fragment of L m1/m2 |
IPS77883 | ACM/C5-D1 | R m1 |
IPS77884 | ACM/C5-D1 | lingually abraded R m1 |
IPS77885 | ACM/C5-D1 | broken R m1/m2 |
IPS77886 | ACM/C5-D1 | R m3 |
IPS77887 | ACM/C5-D1 | fragment of upper molar |
IPS77888 | ACM/C5-D1 | fragment of lower molar |
IPS78179 | ACM/C5-D1 | L M1/M2 |
IPS85340 | ACM/C5-D1 | fragment of left mandible with m1–m3 and associated p4 |
IPS85410 | ACM/C6-A5 | partial cranium (includes the dorsal half of the skull as well as part of the right zygomatic arch and maxillary bone with damaged P4–M3) |
IPS87560 | ACM/C8-B sector | partial left mandible with p4–m3 (incisor and coronoid process broken, p4 damaged) |
IPS88677 | ACM/C8-Af | partial cranium (dorsoventrally crushed) |
List of cranial elements used in the reconstruction of the complete skull indicating catalog no. of each bone or bone fragment and, when appropriate, which side (left or right) was mirrored.
Cranial element | Side | Catalog. no |
---|---|---|
alisphenoid | left | IPS88677 |
auditory bulla | left | IPS56468h |
basisphenoid | left | IPS56468h |
basioccipital and occipital condyle | left | IPS56468h |
cheek teeth | right | IPS56468h |
frontal | – | IPS56468h |
frontal (orbital part) | left | IPS88677 |
hamular process of pterygoid | right | IPS56468h |
incisor | left | IPS56468h |
jugal | left | IPS88677 |
jugal (posterior half) | right | IPS88677 |
maxillary | right | IPS56468h |
occipital (right half) | right | IPS88677 |
palatines | – | IPS88677 |
parietal | – | IPS56468h |
postorbital process of frontal | right | IPS88677 |
pterygoid | left | IPS56468h |
pterygoid fossa (alisphenoid) | – | IPS56468h, IPS88677 |
premaxillary | right | IPS56468h |
premaxillary (dorsal fragments) | right, left | IPS88677 |
squamosal | left | IPS56468h |
squamosal (zygomatic process) | right | IPS88677 |
zygomatic plate (maxillary) | left | IPS88677 |
Miopetaurista neogrivensis is considered as a calibration point for the divergence between Petaurista and all the other Pteromyina in node dating analysis and is considered as a tip date in the …
Fossil | Locality | Node on phylogeny | Max. age | Min. age | Taxonomic refs. | Age refs. |
---|---|---|---|---|---|---|
Acritoparamys atavus | Bear Creek, Montana, USA | maximum age for the root (oldest-known rodent) | 56.0 Ma | 55.8 Ma | Janis et al., 2008; Korth, 1994 | Janis et al., 2008 |
Erlianomys combinatus | AS-1, Nuhetingboerhe, Erlian Basin, Inner Mongolia, China | maximum age for the root (oldest-known Myodonta) | 54.8 Ma | 53.9 Ma | Li and Meng, 2010; Wu et al., 2012 | Wu et al., 2012 |
Eogliravus wildi | Mas de Gimel, Hérault, France | maximum age for the root (oldest-known Gliridae) | 50.7 Ma | 47.4 Ma | Daams and Bruijn, 1995; Dawson, 2003; Hartenberger J-L and Hartenberger, 1971; Storch and Seiffert, 2007; Vianey-Liaud, 1985 | Daams and Bruijn, 1995; Dawson, 2003 |
Spurimus selbyi | Upper Teppe Formation, Wyoming, USA | minimum age for the root (oldest-known Aplodontiidae) | 46.3 Ma | 45.7 Ma | Flynn and Jacobs, 2008; Korth, 1994 | Flynn and Jacobs, 2008 |
Douglassciurus jeffersoni | Renova Formation, Pipestone Springs, Montana, USA | Douglassciurus/other Sciuridae | 36.6 Ma | 35.8 Ma | Emry and Korth, 1996; Emry and Thorington, 1982; Goodwin, 2008 | Goodwin, 2008 |
Heteroxerus sp. | Canales, Loranca Basin, Spain | Xerinae/Sciurinae | 26.4 Ma | 25.0 Ma | Álvarez-Sierra et al., 1999 | Álvarez-Sierra et al., 1999 |
Sciurus olsoni | NB29A, Truckee Formation, Nevada, USA | Sciurus/ Tamiasciurus | 13.6 Ma | 10.3 Ma | Emry et al., 2005 | Goodwin, 2008; Emry et al., 2005; Woodburne, 2004 |
Miopetaurista neogrivensis | Abocador de Can Mata ACM/C5- D1, Vallès-Penedès Basin, Spain | Petaurista/other Pteromyina | 11.6 Ma | this work | this work |
IPS, acronym for the collections of the Institut Català de Paleontologia Miquel Crusafont. All other specimens are kept in the collections of the Naturalis Biodiversity Center (Leiden, the …
Catalogue no. | Taxon | Element | Voltage (kV) | Current (µA) | Filter | Projections | Voxel size (µm) |
---|---|---|---|---|---|---|---|
IPS56468h | Miopetaurista neogrivensis | skull | 130 | 72 | 1 mm | 1880 | 39.71 |
IPS88677 | Miopetaurista neogrivensis | skull | 130 | 72 | 1 mm | 1880 | 39.71 |
ZMA13418 | Petaurista petaurista | skull | 100 | 100 | 0.5 mm | 1290 | 13.17 |
IPS56468ah | Miopetaurista neogrivensis | carpals | 80 | 124 | not used | 651 | 10.67 |
056.054.b | Hylopetes sagitta | carpals | 100 | 100 | not used | 558 | 11.32 |
RMNH.MAM.130 | Petaurista petaurista | carpals | 79 | 125 | not used | 482 | 13.17 |
27291 | Sciurus vulgaris | carpals | 59 | 167 | not used | 550 | 13.04 |
Molecular model specifications for total evidence and node dating phylogenetic analyses.
Analysis software | Genes | Model | Gamma distribution | Invariant |
---|---|---|---|---|
BEAST | cytb codon 1, 16S, 12S | GTR | yes | yes |
BEAST | cytb codon 2, irbp codon 2 | HKY | no | yes |
BEAST | cytb codon 3 | GTR | yes | yes |
BEAST | irbp codon 1 | HKY | no | no |
BEAST | irbp codon 3 | HKY | yes | no |
MrBayes | cytb codon 1, 16S, 12S | GTR | yes | yes |
MrBayes | cytb codon 2, irbp codon 2 | HKY | yes | yes |
MrBayes | cytb codon 3 | GTR | yes | yes |
MrBayes | irbp codon 1 | HKY | yes | no |
MrBayes | irbp codon 3 | GTR | yes | no |
Reconstruction of the skeleton of Miopeaturista neogrivensis.
GenBank accession numbers for all the sequences used in phylogenetic analyses.
Accession numbers for all the species considered in total evince analysis and/or node dating analysis. We retrieved the four genes more frequently used in previous phylogenetic studies of the Sciuridae: 12S, 16S, cytochrome b, and irbp (Mercer and Roth, 2003).
Morphological character matrix for total evidence phylogenetic analysis.
See Appendix 2 for character description and coding.
Cheek tooth measurements of Miopetaurista neogrivensis.
Cheek tooth measurements of Miopetaurista neogrivensis from localities ACM/C5-D1, ACM/C8-Af, ACM/C6-A5, as well as sector ACM/C8-B (material is listed in Tables 1–2). Measurement method follows Casanovas-Vilar et al., 2015: L, mesiodistal length; W, buccolingual width; w1, buccolingual width measured at the mesial margin of the tooth (only for m1 and m2); w2, buccolingual width measured at the distal margin of the tooth (only for m1 and m2). Estimated measurements (due to minor damage to the cheek teeth) are within parentheses. Abbreviations for dental elements follow Smith and Dodson, 2003. IPS, acronym for the collections of the Institut Català de Paleontologia Miquel Crusafont. All measurements are in mm.
Cranial measurements of Miopetaurista neogrivensis and extant flying squirrels.
Cranial measurements for Miopetaurista neogrivensis and some extant flying squirrels. The specimens of the extant species are housed in the Naturalis Biodiversity Center (Leiden, the Netherlands), except for Glaucomys sabrinus, which comes from the collections of the Institut Català de Paleontologia Miquel Crusafont. Cranial measurements for the virtually reconstructed skull of M. neogrivensis are also included. Estimated measurements are in parentheses. Acronyms and definitions of the measurements follow Bertrand et al., 2016 and Nicolas et al., 2008: BNAS, maximum breadth of the nasals; BRCA, maximum breadth of the braincase; BULL, maximum length of the auditory bulla; CTL, length of the upper cheek tooth row; DIA, length of the diastema (from the alveolus of the incisor to the alveolus of the P3); HEBA, henselion–basion length; HEPA, henselion–palation length; IF, length of the incisive foramen; INTE, smallest interorbital breadth; LNAS, maximum length of the nasals; PALA, smallest palatal breadth; RB, maximum rostrum breadth; RH, mediosagittal projection of rostrum height at the anterior border of the M1; SL, maximum length of the skull. The virtual reconstruction of the cranium of M. neogrivensis is shown in Figure 6, for the original crania see Figure 6—figure supplements 1–2. All measurements are in mm.
Mandibular measurements of Miopetaurista neogrivensis and extant flying squirrels.
Mandibular material of Miopetaurista neogrivensis is listed in Tables 1–2. The specimens of the extant species are housed in the Naturalis Biodiversity Center (Leiden, the Netherlands), except for Glaucomys sabrinus which comes from the collections of the Institut Català de Paleontologia Miquel Crusafont. Estimated measurements are in parentheses. Acronyms and definitions of the measurements are as follows: DIA1, length of the diastema measured from the alveolus of the incisor to the anterior alveolus of the p4; DIA2, length of the diastema from the anterior alveolus of the p4 to the projection of the incisor; DDIA, depth of the mandibular corpus at the lowermost point of the diastema; DM1, depth of the mandibular corpus below the anterior border of the m1; DM3, depth of the mandibular corpus below the anterior border of the m3; LAJ, length from the most ventral point of the angular apophysis to the anterior margin of the articular condyle; LCJ, length from the tip of the coronoid apophysis to the anterior margin of the articular condyle; LTL, length of the lower cheek tooth row; ML, maximum length of the mandible; RH, maximum height of the mandibular ramus. The mandibular remains of M. neogrivensis are shown in Figure 3. All measurements are in mm.
Postcranial measurements of Miopetaurista neogrivensis and extant squirrels.
Postcranial measurements for Miopetaurista neogrivensis (for material list see Table 1) and some extant squirrels. The specimens of the extant species are housed in the Naturalis Biodiversity Center (Leiden, the Netherlands), except for Glaucomys sabrinus and Sciurus carolinensis which come from the collections of the Institut Català de Paleontologia Miquel Crusafont. Only right elements were measured. Acronyms and definitions of the measurements are based in Samuels and Van Valkenburgh, 2008: BI, brachial index (RL/HL), indicating the proportions of the distal and proximal elements of the forelimb; CI, crural index (TL/FL), indicating the proportions of the distal and proximal elements of the forelimb; FEB, epicondylar mediolateral breadth of the distal femur; FGT, mediolateral breadth of the greater trochanter of the femur; FL, femoral length from the tip of the femoral head to the tip of the medial epicondyle; FMD, greater diameter of the femur at midshaft; FRI, femoral robusticidty index (FMD/FL); HEB, epicondylar mediolateral breadth of the distal humerus; HL, humeral length from the tip of the head to the tip of the most distal point of the trochlea; HMD greater diameter of the humerus at midshaft; HRI, humeral robusticity index (HMD/HL); IM, intermembral index [(HL +RL)/(FL +TL)], indicating the length of the forelimb relative to the hindlimb; TL, tibial length from the medial condyle to the tip of the medial malleolus; TMD, greater diameter of the tibia at midshaft; TRI, tibial robusticity index (TMD/TL); RL, radius lenght. All measurements are in mm. Specimens shown in Figure 5 are: Miopetaurista neogrivensis IPS56468; Petaurista petaurista RMNH.MAM.130; Hylopetes sagitta 056.054.b; Callosciurus prevostii 056.099 .j; Urocitellus paryii 065.238a; Xerus erythropus 26483.