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Smithsonian Institution

Scholarly Press

Skeletal Anatomy of the 

North American Pangolin 

Patriomanis americana

(Mammalia, Pholidota) from the 

Latest Eocene of Wyoming (USA)

Timothy J. Gaudin, Robert J. Emry, and Jeremy Morris

sm iths onian contri b uti ons to paleo b io logy • nu m b er 9 8
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s m i t h s o n i a n c o n t r i b u t i o n s t o p a l e o b i o l o g y • n u m b e r 9 8

Skeletal Anatomy of the 

North American Pangolin 

Patriomanis americana

(Mammalia, Pholidota) from the 

Latest Eocene of Wyoming (USA)

Timothy J. Gaudin, Robert J. Emry, and Jeremy Morris

washington d.c.

2016
ABSTRACT

Gaudin, Timothy J., Robert J. Emry, and Jeremy Morris. Skeletal Anatomy of the North American Pangolin Patriomanis americana (Mammalia, Pholidota) from the Latest Eocene of Wyoming (USA). Smithsonian Contributions to 

Paleobiology, number 98, viii + 102 pages, frontispiece + 52 figures, 1 appendix with 13 tables, 2016.—Patriomanis 

americana is the only pangolin (Mammalia, Pholidota), living or extinct, known from the Western Hemisphere. 

It derives from latest Eocene (Chadronian North American Land Mammal Age) deposits from central Wyoming 

and western Montana. Since its initial description more than 40 years ago based on a partial skeleton, several 

nearly complete skeletons have been discovered, together including nearly every bone in the skeleton. This taxon 

is thus not only the most completely preserved fossil pangolin but is also among the best preserved of any Eocene 

mammal taxon. In the present study we have prepared a detailed, bone-by-bone description of the osteology of 

Patriomanis, comparing it with other well-known fossil pangolin skeletons (Eomanis, Euromanis, Cryptomanis, 

and Necromanis), as well as representatives of the three extant pangolins genera (Manis, Smutsia, Phataginus). We 

provide a catalog of all known Patriomanis specimens and their provenance and an extensive series of measurement 

tables incorporating the comparative taxa. We analyze the alpha-level taxonomy of the genus, concluding that all 

specimens should be kept in a single species, Patriomanis americana, based on currently available fossil material. 

We summarize the taxonomic and phylogenetic position of Patriomanis and discuss its implications for the biogeographic history of the order Pholidota. We analyze the paleobiology of Patriomanis, concluding that it was likely a 

myrmecophagous, arboreal animal with a prehensile tail. Last, we discuss its paleoecology, suggesting that its late 

appearance in the Eocene record, during a time of global cooling, may imply that earlier pangolins are waiting to be 

discovered in the Eocene record of Asia and North America. KEY WORDS: Eocene, fossils, morphology, osteology, 

pangolins, Patriomanis, Pholidota, skeleton, skull.

Cover image: Drawing of Patriomanis americana skeleton (detail from Figure 1).

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Library of Congress Cataloging-in-Publication Data

Names: Gaudin, Timothy J., author. | Emry, Robert J., author. | Morris, Jeremy, 1981—author. | Smithsonian 

Institution Scholarly Press, publisher.

Title: Skeletal anatomy of the North American Pangolin Patriomanis americana (Mammalia, Pholidota) from the latest 

Eocene of Wyoming (USA) / Timothy J. Gaudin, Robert J. Emry, and Jeremy Morris.

Other titles: Smithsonian contributions to paleobiology; no. 98.

Description: Washington, D.C.: Smithsonian Institution Scholarly Press, 2016. | Series: Smithsonian contributions 

to paleobiology, ISSN 0081-0266; number 98 | Includes bibliographical references.

Identifiers: LCCN 2016024058

Subjects: LCSH: Pangolins, Fossil—North America. | Extinct mammals—North America.

Classification: LCC QE882.P45 G38 2016 | DDC 569/.31—dc23 | SUDOC SI 1.30:98 LC record available at 

https://lccn.loc.gov/2016024058

ISSNs: 1943-6688 (online); 0081-0266 (print)

ZooBank Registration: 8 September 2016 (LSID: urn:lsid:zoobank.org:pub:0E6025B4-6E40-452A-A333-

25D39FE616A4)

Publication date: 22 September 2016

The paper used in this publication meets the minimum requirements of the American National Standard for 

Permanence of Paper for Printed Library Materials Z39.48–1992.
Contents

LIST OF FIGURES vi

INTRODUCTION 1

Institutional Abbreviations 3

Other Abbreviations 3

Catalog of Specimens 3

Acknowledgments 6

DESCRIPTION 7

Axial Skeleton: Skull 7

Premaxilla 7

Nasal 7

Maxilla 8

Palatine 10

Lacrimal 11

Jugal 11

Frontal 12

Parietal 12

Interparietal 12

Squamosal 13

Ethmoid and Vomer 15

Orbitosphenoid and Presphenoid 16

Alisphenoid 16

Pterygoid 17

Basisphenoid and Basioccipital 18

Petrosal 19

Ectotympanic/Entotympanic/Ear Ossicles 22

Occipital 22

Endocranium 22

Mandible 23
i v • 

Axial Skeleton: Vertebral Column 24

Atlas 24

Axis 26

Cervical Vertebrae (Exclusive of Atlas and Axis) 27

Thoracic Vertebrae 28

Lumbar Vertebrae 31

Caudal Vertebrae 32

Axial Skeleton: Ribs and Sternum 36

Ribs 36

Sternum 37

Hind Limb 38

Pedal Unguals 38

Pedal Intermediate Phalanges 39

Pedal Proximal Phalanges 40

Metatarsals 41

Prehallux 43

Entocuneiform 43

Mesocuneiform 44

Ectocuneiform 44

Cuboid 45

Navicular 46

Astragalus 47

Calcaneus 49

Fibula 51

Tibia 53

Patella 55

Femur 55

Pelvis and Sacrum 58

Forelimb 62

Scapula 62

Clavicle 64

Humerus 64

Ulna 68

Radius 71

Scapholunar 73

Triquetrum 75

Pisiform 75

Trapezium 76

Trapezoid 76

Capitate 77

Hamate 78

Metacarpals 79

Manual Proximal Phalanges 81

Manual Intermediate Phalanges 82

Manual Unguals 83

DISCUSSION 83

Species Comparisons Based on Size and Stratigraphy 83

Phylogeny, Taxonomy, and Biogeographic Origins 85

Paleobiology 85

Paleoecology, Climate, and Geographic Distribution 87
n u m b e r 9 8 • v

APPENDIX: TABLES OF MEASUREMENTS 89

A1. Skull and mandible 90

A2. Atlas and axis 90

A3. Thoracic and lumbar vertebrae 91

A4. Metatarsals and pedal phalanges 92

A5. Astragalus 93

A6. Calcaneus 94

A7. Tibia and fibula 94

A8. Patella and femur 95

A9. Scapula 95

A10. Humerus, radius, and ulna 96

A11. Scapholunar 97

A12. Metacarpals 97

A13. Manual phalanges 98

REFERENCES 99
Figures

Frontispiece. Life reconstruction of Patriomanis americana

foraging for termites in the treetops viii

1. Skeleton of Patriomanis americana in right lateral 

and dorsal view 4

2. Partially prepared skeleton of P. americana 5

3. Skull of P. americana in right and left lateral views 

with reconstruction 8

4. Skull of P. americana in dorsal and ventral views 

with reconstructions 9

5. Skulls and mandible of modern pangolins 10

6. Skull of Phataginus tricuspis and skull fragment of Patriomanis 

americana 13

7. Close up of right auditory region of P. americana 

in ventral view 15

8. Close up of left auditory region of Smutsia gigantea

in ventral view 17

9. Mandible of P. americana in lateral, medial, and 

occlusal views 24

10. Atlas of P. americana in anterior, lateral, and posterior 

views with reconstruction 25

11. Atlas and axis of P. americana in multiple views 25

12. Axis of P. americana in anterior and ventral views 

with reconstruction 26

13. Cervical vertebra (C4 or C5?) of P. americana in anterior, 

dorsal, and lateral views 27

14. Thoracic vertebrae of P. americana in anterior view 28

15. Thoracic vertebrae of P. americana in dorsal view 29

16. Thoracic vertebrae of P. americana in lateral view 29
n u m b e r 9 8 • vii

17. Penultimate lumbar vertebra (L6) of P. americana in anterior view 31

18. Lumbar vertebrae (L6 and L7) of P. americana in dorsal view 31

19. Lumbar vertebrae (L6 and L7) of P. americana in left lateral view 32

20. Caudal vertebrae of P. americana in dorsal views 34

21. Distal caudal vertebrae and chevrons of P. americana 35

22. Ribs of P. americana 36

23. Sternebrae of P. americana in ventral and lateral views 37

24. Pes of P. americana in dorsal view 38

25. Ungual phalanx II of right pes of P. americana in medial 

and ventral views 39

26. Proximal phalanges III (of left manus and right pes) 

of P. americana in proximal and distal views 40

27. Metapodials of P. americana in distal view 41

28. Distal tarsals of P. americana (left entocuneiform, left 

mesocuneiform, and right ectocuneiform) 43

29. Cuboid and navicular of P. americana 45

30. Right astragalus of P. americana 47

31. Calcaneus of P. americana 49

32. Right tibia and fibula and left pes of Phataginus tricuspis 50

33. Right tibia and fibula of Patriomanis americana in 

posterior and anterior views 52

34. Right tibia of P. americana in proximal and distal views 53

35. Right patella of P. americana in posterior view 55

36. Right femur of P. americana in posterior and anterior views 56

37. Left femur of P. americana in proximal and distal views 57

38. Pelvis, sacrum, and left femur of Phataginus tricuspis 58

39. Right pelvis and sacrum of Patriomanis americana 59

40. Sacrum of P. americana in dorsal and lateral views 61

41. Scapula of Patriomanis americana and extant pangolin Smutsia 

temminckii in lateral view 63

42. Right humerus of P. americana in anterior and posterior views 65

43. Right humerus of P. americana in proximal and distal views 66

44. Forelimb skeleton of modern pangolins 67

45. Right ulna of P. americana in posterior and anterior views 69

46. Right ulna of P. americana in lateral, medial, and distal views 70

47. Radius of P. americana in posterior and anterior views 71

48. Left radius of P. americana in proximal, distal, and lateral views 72

49. Right manus of P. americana in dorsal view 73

50. Left scapholunar of P. americana in proximal and distal views 74

51. Triquetrum and pisiform of P. americana 75

52. Capitate and hamate of P. americana 77
Frontispiece. Life reconstruction of Patriomanis americana foraging for termites in the treetops. The tree is from th

genus Carya, which, according to Prothero (1994), was present in this part of Wyoming during the Chadronian North 

American Land Mammal Age (latest Eocene). Illustration by Julia Morgan Scott for Smithsonian Institution.
INTRODUCTION

One of the more remarkable and unexpected findings to emerge from the 

past 100 years of detailed study of the North American Cenozoic fossil mammal 

record is the discovery of a fossil pangolin in latest Eocene deposits of central 

Wyoming (Emry, 1970). The pangolins themselves, relegated to their own order, 

the Pholidota, are an extraordinary group of placental mammals. Rare, secretive, solitary, and largely nocturnal, the animals in this group comprise only 

eight living species distributed across sub-Saharan Africa, the Indian subcontinent, Southeast Asia, Indonesia, and the westernmost Philippines (Corbet and 

Hill, 1991; Nowak, 1999; Gaubert and Antunes, 2005, Schlitter, 2005; Gaudin 

et al., 2009). Thus, they are relatively poorly known even in the parts of the 

globe they frequent and virtually unheard of outside those areas. Their external 

appearance is truly bizarre, for they are the only mammals to be covered in 

an epidermal armor of large overlapping keratinous scales, earning them such 

nicknames as “perambulating artichoke.” And their list of oddities does not 

end there. They have enormously enlarged salivary glands producing copious, 

sticky saliva; a portion of their stomach is converted into a cornified “gizzard” 

for grinding their preferred food, ants and termites; their tongue is remarkably 

elongated, taking its origin from a xiphisternal cartilage that may itself extend 

far back into the abdominal cavity; and one species, the long-tailed pangolin 

(Phataginus tetradactyla), has the most caudal vertebrae of any living mammal, 

its prehensile organ formed around 47–49 vertebral elements (Flower, 1885; 

Kingdon, 1974, 1997). 

Pangolins are fossorially adapted for excavating ant and termite nests, and 

most species are either arboreal or semiarboreal/scansorial, with the exception 

of the two terrestrial African ground pangolins (Smutsia spp.). Their terrestrial 

progression is generally slow (Kingdon, 1974; Heath 1992a, 1992b; Nowak, 

1 Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, 615 McCallie Avenue, Chattanooga, 

Tennessee 37403-2598 USA.

2 Department of Paleobiology, MRC 121, National Museum of Natural History, Smithsonian 

Institution, P.O. Box 37012, Washington, D.C. 

20013, USA.

3 Present address: Department of Biology, University of Utah, 257 South 1400 East, Rm. 201, 

Salt Lake City, Utah 84112-0840, USA.

*Correspondence: timothy-gaudin@utc.edu

Manuscript received 15 January 2016; accepted 

17 March 2016.

Skeletal Anatomy of the North American 

Pangolin Patriomanis americana 

(Mammalia, Pholidota) from the Latest 

Eocene of Wyoming (USA)

Timothy J. Gaudin,1* Robert J. Emry,2 and Jeremy Morris1,3
2 • 

of Natural History (F:AM 78999) and two later recovered 

by Emry from same vicinity (see below) housed at Smithsonian’s Natural History Museum (USNM-P 299960, 

494439), both relatively complete but in combination representing almost every skeletal element in the body. 

Articulated, well-preserved skeletons are a rarity in 

vertebrate paleontology, complete skeletons even more 

so. These remains of Patriomanis not only represent the 

most complete known fossil pholidotan skeleton but are 

also among the best-known skeletons of any fossil mammal from the Eocene. Such finds are, of course, invaluable 

for understanding phylogenetic relationships among living 

and extinct mammals, although in this case, an exhaustive phylogenetic treatment of Patriomanis and other living and extinct pangolins has already been conducted by 

Gaudin et al. (2009). That work is an essential companion 

to the present study and is frequently cited in the material 

that follows. However, these Patriomanis skeletons are significant beyond what can be ascertained from them concerning pangolin phylogenetic relationships. They show us 

the skeletal morphology of a pangolin significantly more 

primitive than that of any alive today and hence should 

prove valuable in any subsequent investigations of morphological evolution in the group. To the degree that 

function can be inferred from these remains, they provide 

insight into the paleobiology of Patriomanis and into paleobiological aspects of pangolin evolution. Finally, given 

the relative rarity of complete fossil mammal skeletons, 

especially from the early parts of the Cenozoic, they have 

the potential to yield insight into broader aspects of skeletal evolution and supraordinal phylogeny across placental 

mammals as a whole, including, for example, their putative relationship to Carnivora in the supraordinal clade 

Ferae (see O’Leary et al., 2013, and references therein). 

The primary goal of the present study is to describe 

in detail the skeletal anatomy of the North American fossil pangolin Patriomanis americana. We describe all the 

known material of Patriomanis and compare its anatomy 

to that of the other fossil pangolins. As has been noted 

elsewhere (e.g., Gaudin et al., 2006; Gaudin, 2010), the 

fossil record of Pholidota is poor, with very few extinct 

taxa (McKenna and Bell, 1997; Gaudin et al., 2009). That 

said, the taxa that do exist are reasonably well known, 

including Eomanis (middle Eocene of Germany; Storch, 

1978, 2003), Euromanis (middle Eocene of Germany; 

Storch and Martin, 1994; Gaudin et al., 2009), Cryptomanis (late Eocene of China; Gaudin et al., 2006), and 

Necromanis (Oligocene-Miocene of central Europe; Koenigswald and Martin, 1990; Koenigswald, 1999; see also 

Storch, 2003; Gaudin et al., 2009; Hoffmann et al., 2009; 

1999). Given their slow movements, alongside the low 

diversity and abundance at which their populations exist, 

and their sluggish reproductive rate (litter size typically 

one, occasionally two or three in Asiatic species, birthed 

once a year in all but African tree pangolins, which may 

breed two or three times in a year; Kingdon, 1974, 1997; 

Heath, 1992a, 1992b, 1995; Nowak, 1999), they do not 

seem like a group that would be likely to spread quickly 

or to have a broad, global distribution. Indeed, the presence of different species of myrmecophagous mammals 

on different continents, with vermilinguans restricted to 

South and Central America, aardvarks and aardwolves 

in Africa, and echidnas and numbats in Australia, has 

become an iconic example of convergent evolution in 

undergraduate textbooks (e.g., Vaughan et al., 2011), 

even at the introductory level (e.g., Solomon et al., 2011). 

Extant pangolins are the only one of these groups to be 

broadly distributed on two continents (although vermilinguans do extend their range into southern parts of Central 

America, and aardvarks were distributed into Europe and 

Asia historically; Holroyd and Mussell, 2005). Until the 

recent report of the new extinct taxon Cryptomanis from 

the late Eocene of northern China, the Patriomanis localities lay on the other side of the world from the only other 

known Paleogene pangolins, the middle Eocene Eomanis 

and Euromanis from central Europe (Gaudin et al., 2006, 

2009). To the present day, all but one of the Paleogene 

pangolin records (the exception being several isolated 

ungual phalanges from the Fayum of Egypt of questionable value; see discussion in Gaudin et al., 2009) pertain to 

localities in the midlatitudes of the Northern Hemisphere, 

outside the range not only of the modern pangolins but 

of all other extant myrmecophagous mammals (although 

these areas were characterized by a subtropical climate in 

the Eocene; Prothero, 1994). Thus, the presence of pangolins throughout northern Laurasia and into western North 

America in the Eocene is a startling finding. 

Emry (1970, 2004) describes the somewhat circuitous 

route by which the first partial remains of a creature from 

the modern day midlatitudes of North America were recognized as pertaining to this group from the Old World 

tropics. He notes that the original identification was based 

largely on postcranial remains, although the type specimen included a well-preserved braincase (Emry, 1970). 

He subsequently reported on the remains of an additional 

specimen that preserved the front end of the skull, which 

demonstrated that this fossil pangolin was edentulous 

like its modern relatives (Emry, 2004). There are, in fact, 

three primary specimens, including the incomplete type 

housed in the Frick Collections at the American Museum 
n u m b e r 9 8 • 3

L left

L1, L2, L3, ... lumbar vertebrae

m., mm. muscle, muscles

mc metacarpal

MML maximum mandibular length

mt metatarsal

mya million years ago

NALMA North American Land Mammal 

Age

R right

T1, T2, T3, ... thoracic vertebrae

Catalog of Specime

There are six specimens of Patriomanis, five of which 

comprise multiple skeletal elements, two of those representing nearly complete skeletons. All but one are housed 

in the Department of Paleobiology at the U.S. National 

Museum of Natural History of the Smithsonian Institution. The remaining specimen, the holotype, is part of the 

Frick Collection at the American Museum of Natural History. The type and four of the five other specimens come 

from three different sites in the Chadronian NALMA (latest Eocene) deposits of the White River Formation in the 

Flagstaff Rim area of Natrona County, Wyoming (Emry, 

1970, 1973, 2004). The remaining specimen is from Chadronian NALMA strata at Pipestone Springs, Montana. 

For the convenience of the reader, a complete catalog of 

specimens and their provenance is provided below.

F:AM 78999 (holotype; see Emry, 1970; Figure 1, skull): 

White River Formation, Flagstaff Rim, Natrona 

County, Wyoming; Chadronian NALMA, latest Eocene; stratigraphic level, 5 m above Ash F, is in the late 

middle Chadronian part of the section (Emry, 1973, 

2004; Unit 21 of Emry, 1973); assigned to Chadronian Biochron Ch3, 34.8–35.8 mya, in Rose (2008) 

and Janis et al. (2008, appendix 1; = middle Chadronian)—Braincase (endocranium visible, endocast 

available), L and R mandibular fragments, incomplete 

atlas and axis, cervical vertebrae fragments, 8 thoracic 

vertebrae (3 articulated with ribs, 1 partial anterior, 4 

partial posterior), 7 caudal vertebra (1 compete anterior, 6 partial), rib fragments (including 3 mediolaterally expanded ribs), 2 pelvic fragments (one including 

the pubic symphysis, the other preserving the left ischium ventral to the obturator foramen), fragments of 

L femur, partial L patella, partial L and R tibia, proximal R fibula, R calcaneus, L cuboid, R proximal mt I 

and V, 2 distal mt, partial R scapula, partial L and R 

Hoffmann, 2011, for new, undescribed material). Because 

Patriomanis is more complete than any of the other fossil 

pangolin taxa, however, and the whole group Pholidota 

is rather small, we also compare Patriomanis to each of 

the three extant pangolin genera, the African tree pangolins (two species in genus Phataginus), the African ground 

pangolins (two species in the genus Smutsia), and the 

Asian pangolins (four species in the genus Manis; Gaudin 

et al., 2009). Last, some comparisons are included, where 

appropriate, to the metacheiromyid palaeanodonts Metacheiromys and Palaeanodon and the enigmatic “edentate”

Eurotamandua. According to Gaudin et al. (2009), Palaeanodonta is the sister taxon to Pholidota (see also Matthew, 1918; Emry, 1970; Rose, 1979; Rose et al., 2005; 

O’Leary et al., 2013), forming a more inclusive clade 

Pholidotamorpha, whereas Eurotamandua falls within 

Pholidota proper, although the latter is a more debatable

conclusion. It is hoped that these broad-based comparisons will yield additional insight into the evolution of skeletal anatomy in pangolins. We also examine in brief the 

paleobiological implications of the Patriomanis remains, 

as well as their potential to inform us about the Paleogene 

history of pangolins and the terrestrial ecosystems of the 

late Eocene.

Institutional Abbreviat

AMNH American Museum of Natural History, 

New York, NY

CM Carnegie Museum of Natural History, Pittsburgh, PA

F:AM Frick Collection, American Museum of Natural History, New York, NY

FMNH Field Museum of Natural History, Chicago, 

IL

SMF Senckenberg Museum, Frankfurt am Main, 

Germany

USNM National Museum of Natural History, 

Smithsonian Institution, Washington, D.C.

UTCM University of Tennessee at Chattanooga Natural History Museum, Chattanooga, TN

Other Abbreviati

C1, C2, C3, ... cervical vertebra

Ca1, Ca2, Ca3, ... caudal vertebra

char(s). character(s)

CN cranial nerve

GL greatest length

GSL greatest skull length
4 • 

anteriorly, including first 5 in articulation, the remainder more posterior, many composed only of centra or 

partial centra), 4–5 chevron bones, 11 ribs with proximal end preserved (including 2 L, 4 R), many rib fragments, 3 sternebrae, complete pelvis, L and R femur, L 

and R patella, L and R tibia, R fibula, partial L fibula, 

complete R pes (except entocuneiform), L pes (missing astragalus, ungual phalanx I, damaged calcaneus, 

cuboid, mt I and II), miscellaneous sesamoids, partial 

L and R scapula, R humerus, R ulna, L and R radius, 

complete R manus (triquetrum damaged), L manus 

(missing hamate, mc III, proximal phalanx II, intermediate phalanx IV, damaged triquetrum), miscellaneous sesamoids, ?hyoid element.

USNM-P 460256: Renova Formation, Climbing Arrow 

Member, Pipestone Springs Local Fauna, Jefferson 

River Basin, Montana; Chadronian NALMA, latest 

Eocene; assigned to Chadronian Biochron Ch3 (= 

middle Chadronian), 34.8–35.8 mya in Rose (2008) 

and Janis et al. (2008, appendix 1)—Partial R ulna.

USNM-P 494439 (Figure 2): White River Formation, 

Flagstaff Rim, Natrona County, Wyoming; Chadronian NALMA, latest Eocene; stratigraphic level 5 m 

above Ash F (at the same stratigraphic level and only 

3–4 m away from the spot where the type was discovered; Emry, 1973, 2004; Unit 21 of Emry, 1973) 

assigned to Chadronian Biochron Ch3 (= middle Chadronian), 34.8–35.8 mya in Rose (2008) and Janis 

et al. (2008, appendix 1)—Partial skull (including premaxilla, rostrum, partial L and R dentary, complete L 

humerus, partial L and R ulna, partial L and R radius, 

partial R scapholunar, R pisiform, L hamate, manual 

phalanges (including 2 proximal phalanges, 1 partial 

and 1 complete intermediate phalanx, 4 ungual phalanges [digits I, II, III, V]).

USNM-P 299960 (Figure 1, postcrania): White River Formation, Flagstaff Rim, Natrona County, Wyoming; 

Chadronian NALMA, latest Eocene; stratigraphic 

level near base of claystone immediately below Ash 

A (Unit 4 of Emry, 1973) or about 16 m below Ash 

B, which is radiometrically dated at 35.9 Ma (Swisher 

and Prothero, 1990; Prothero and Swisher, 1992). This 

specimen occurred almost exactly at the early Chadronian–middle Chadronian transition as defined by Prothero and Emry (1996, 2004); assigned to Chadronian 

Biochron Ch2-3, 34.8–36.6 mya in Rose (2008) and 

Janis et al. (2008, appendix 1) but, given stratigraphic 

position, likely in the older part of this range—Skull 

fragments (including frontal bone with 8 grooves on 

underside for ethmoturbinals; L posterolateral corner 

of skull showing parietal, squamosal, occipital, and 

petrosal; basisphenoid–basioccipital with sella turcica 

dorsally and groove for musculotubal canal ventrally; 

R junction of parietal, squamosal, and occipital, and 

3 miscellaneous fragments), partial axis, 3 cervical 

vertebrae (1 complete, 2 centra only), 11 thoracic vertebrae (9 anterior, 2 posterior, range from partial to 

nearly complete), 5 lumbar vertebrae (3 partial, last 

3 in articulation, last 2 nearly complete), complete 

sacrum (3 sacrals), 28 caudal vertebrae (9 complete 

FIGURE 1. Drawing of skeleton of Patriomanis americana in right lateral (top) and dorsal (bottom) views based on the skull of the holoty

(F:AM 78999) and the skeleton of USNM-P 299960. Drawn by Larry Isham and digitized by Mary Parrish for Smithsonian Institution.
n u m b e r 9 8 • 5

FIGURE 2. Partially prepared skeleton 

Patriomanis americana (in block), USNM-P 

494439: (a) dorsal view with more superficial 

elements of block removed to show position of 

skull; (b) full specimen in dorsal view; (c) line 

drawing of full specimen in dorsal view with 

skeletal elements highlighted. Abbreviations: 

ast, astragalus; cav, caudal vertebrae; clc, calcaneus; cu, cuboid; cv, cervical vertebrae; fib, 

fibula; fm, femur; hu, humerus; il, ilium; ip, 

intermediate phalanx; isch, ischium; lv, lumbar 

vertebra; mc, metacarpal; mt, metatarsal; mt 

V, fifth metatarsal; pat, patella; pp, proximal 

phalanx; rad, radius; s, scapula; sk, skull; sv, 

sacral vertebra; tib, tibia; tv, thoracic vertebra; 

ul, ulna. Gray areas represent matrix. Scale 

bar = 5 cm. Photos (a), (b) by James DiLoreto; 

illustration (c) by Tim Gaudin and Julia Morgan Scott for Smithsonian Institution.
6 • 

New York, NY; John Flynn and Meng Jin, Department of 

Vertebrate Paleontology, American Museum of Natural 

History, New York, NY; Bruce Patterson, Larry Heaney, 

and Bill Stanley, Division of Mammals, Field Museum 

of Natural History, Chicago, IL; Gerhard Storch, Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt 

am Main, Germany; Norbert Micklich and Connie Kurz, 

Hessisches Landesmuseum Darmstadt, Darmstadt, Germany; Ken Rose, Johns Hopkins University, Baltimore, 

MD; Richard Thorington, Linda Gordon, and Helen 

Kafka, Department of Mammalogy, National Museum of 

Natural History, Washington, D.C.; and Eberhard “Dino” 

Frey, Staatliches Museum für Naturkunde, Karlsruhe, Germany. We especially thank John Flynn and Meng Jin of 

the American Museum of Natural History for providing 

access to the type material of Patriomanis (and Cryptomanis). Special thanks are also owed to Gerhard Storch, who 

provided access to undescribed material of Necromanis for 

the Gaudin et al. (2009) study and who also provided us 

casts of several specimens of Eomanis waldi and the type 

of Euromanis krebsi and showed us his three-dimensional 

X-ray photographs of Eurotamandua joresi. His material 

help and insightful comments were critical to the success 

of this project. 

All of the USNM specimens of Patriomanis from 

Flagstaff Rim were collected between 1971 and 1991, 

and during that interval Emry was assisted in the field by 

various staff of the museum and volunteers; among these, 

he especially thanks Frederick Grady, USNM preparator, 

who in 1975 found USNM 299960 and helped to collect 

it, and Wang Banyue of the Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, who helped collect 

USNM 494439 in 1991. For help with the preparation 

and casting of Patriomanis and Cryptomanis specimens, 

we thank Fred Grady, Pete Kroehler, and Steve Jabo of the 

Department of Paleobiology at the National Museum of 

Natural History. 

This work was greatly improved by the comments of 

Ken Rose; we thank him for his excellent and thorough 

review. We thank Julia Morgan Scott, Mary Parrish, and 

the late illustrator Larry Isham for their skillful work in 

the preparation of illustrations. Mary Parrish and Larry 

Isham prepared Figure 1, and Larry Isham prepared Figures 24b, 33, 36, 39b, 42, 45, 47a,b, and 49a (T.J.G. 

prepared Figure 48c). Julia Morgan Scott prepared the 

frontispiece and the remaining drawings and formatted 

and labeled all the figures. B. Ammons and L. Boykin 

assisted with the formatting of the tables, and L. Boykin 

assisted with the preparation of the figure captions and 

other editing tasks. For housing T. Gaudin during his 2002 

and R petrosals, partial L and R orbital walls), partial 

atlas and axis, at least 5 thoracic vertebrae (perhaps 

1 or 2 more, last 3 in articulation with lumbars), 6 

articulated lumbar vertebrae (lumbars 3–5 damaged, 

6 articulated with sacrum), partial sacrum (3 sacrals), 

26 caudal vertebrae (include fragment of first, 14 anterior and mid-caudals in articulation likely starting 

with Ca3, 11 posterior), 11 ribs (4 L, 7 R) plus multiple rib fragments, partial L pelvis, nearly complete L 

femur, damaged R femur, L patella, damaged L and R 

tibia, damaged R fibula, distal L fibula, L calcaneus, L 

astragalus, L and R mesocuneiform, L and R ectocuneiform, partially articulated L pes (including cuboid, 

isolated tarsal [?mesocuneiform or prehallux], mt I–V, 

proximal phalanges II–V, intermediate phalanges ?IV 

and ?V), articulated R pes (including calcaneus, astragalus, cuboid, mt I–V, proximal phalanx II–III, intermediate phalanx II–IV, ungual phalanges I and III), 

partial L and R scapula, damaged R humerus, L and 

R ulna, L and R radius, partial L manus (including mt 

I, proximal phalanx II, intermediate phalanx II, ungual phalanx IV), partial R manus (including triquetrum, trapezium, capitate, hamate, mc I–IV, proximal 

phalanges I–IV, intermediate phalanx II–IV, ungual 

phalanges I–IV), miscellaneous isolated phalanges 

(1 proximal, 1 intermediate, 1 ungual).

USNM-P 531556: White River Formation, Flagstaff Rim, 

Natrona County, Wyoming; Chadronian NALMA, 

latest Eocene; stratigraphic level just below Ash D 

(Unit 17 of Emry, 1973) assigned to Chadronian Biochron Ch3 (= middle Chadronian), 34.8–35.8 mya

in Rose (2008) and Janis et al. (2008, appendix 1)—

Complete R mt V, incomplete ungual phalanx, complete L humerus, partial R humerus, partial R ulna, 

partial L radius, complete R radius, miscellaneous 

fragments.

USNM-P 531557: White River Formation, Flagstaff Rim, 

Natrona County, Wyoming; Chadronian NALMA, 

latest Eocene; stratigraphic level 8 m [25 feet] below 

Ash D (Unit 17 of Emry, 1973)—Distal fibula, distal R 

mt II, left capitate, 3 ungual phalanges, miscellaneous 

fragments.

Acknowledgme

We thank the following institutions and individuals for 

providing access to the specimens that were used for comparative purposes in this study: Ross MacPhee, Richard 

Monk, Nancy Simmons, and Eileen Westwig, Department 

of Mammalogy, American Museum of Natural History, 
n u m b e r 9 8 • 7

Since the publication of Emry’s (2004) description, 

additional preparation of the skull from USNM-P 494439 

has been undertaken, particularly in the orbital wall, posterior palate, nasopharynx, basicranium, auditory region, 

and occiput (Figures 3, 5–7). Therefore, the following 

description will focus mainly on these areas, although the 

skull bones and portions of the mandible already described 

by Emry (1970, 2004; Rose and Emry, 1993) will be given 

at least cursory treatment. For terminology we largely follow that of Wible and Gaudin (2004) and Wible (2010).

Premaxilla

The edentulous premaxilla of Patriomanis has been 

described in detail by Emry (2004), so we will limit our 

discussion here to comparisons with a somewhat wider 

variety of fossil and extant pangolins than those considered in the previous work. Emry (2004) has already noted 

the strong resemblance of the premaxilla in Patriomanis 

to that of Manis javanica. As in all the modern forms, the 

prominent dorsal process of the premaxilla is oriented posterodorsally; that is, the anterior edge extends anteroventrally to posterodorsally (Figure 3c). In the only other fossil 

pangolin that preserves the premaxilla, Eomanis waldi, the 

leading edge of the bone is roughly vertical and distinctly 

concave anteriorly, giving the whole dorsal process a C 

shape in lateral view (Storch, 2003; Rose et al., 2005). This 

morphology is also present in the palaeanodont Metacheiromys and in the enigmatic edentate mammal Eurotamandua, making it a synapomorphy of Pholidotamorpha 

(as defined by Gaudin et al., 2009) that is subsequently 

modified in patriomanids and manids. Emry (2004:131) 

described the incomplete posteromedian process on the 

palatal portion of the premaxilla as “essentially like that of 

M. javanica.” However, the form of the anterior edge of the 

maxilla, with its rather short median V-shaped notch (Figure 4b), makes it likely that Patriomanis did not possess an 

elongate median process like that found in M. javanica, M. 

crassicaudata, and the two extant African genera (Figure 

5b; Gaudin et al., 2009) and, moreover, that the incisive 

foramen lay between the premaxilla and maxilla, rather 

than being completely enclosed in the former, a derived 

condition found in Smutsia, M. pentadactyla, and in some, 

but not all, specimens of M. javanica (Gaudin et al., 2009).

Nasal

Emry (2004) describes these elements in detail, and 

there is little to add here. The notch described by Emry 

between the medial and lateral anterior processes of the 

period of sabbatical study and again during the summer 

of 2004, special thanks are owed to the Department of 

Paleobiology at the National Museum of Natural History. 

T. Gaudin’s work on this project was supported by a sabbatical grant from the University of Chattanooga Foundation and by NSF RUI Grant DEB 0107922 and NSF AToL 

Grant 0629959.

DESCRIPTION

Axial Skeleton: Skul

Figures 3–9

The skull of Patriomanis has been described and illustrated in several previous publications. Emry (1970:466) 

described the type specimen (F:AM 78999), an uncrushed 

isolated braincase in excellent condition, broken through 

“at the level of the cribriform plate, which is also missing.” It displays much of the basicranial and endocranial 

anatomy, although the former is not everywhere well preserved and the latter can be observed only through the 

anterior opening into the braincase or the foramen magnum, limiting which structures are visible. The same skull 

was illustrated in Rose and Emry (1993: fig. 7.1, right 

lateral view), where it was used to show the separation 

between the alisphenoid and parietal bones at the back 

of the orbit and the enclosure of the foramen ovale by the 

alisphenoid in pangolins. Emry (2004) described the other 

good skull for Patriomanis, that of USNM-P 494439. He 

noted (p. 131) that “the skull is distorted,” mainly through 

some mediolateral crushing, and that “some of the individual bones are broken and displaced, especially in the 

interorbital region and the dorsal part of the braincase.” 

His description focused almost entirely on the bones of 

the rostrum (premaxilla, maxilla, nasal) and the mandible since these were elements missing from the type and, 

more importantly, these elements showed that Patriomanis was edentulous like the modern forms. This skull was 

also used to code characters in Gaudin and Wible’s (1999) 

cladistic phylogenetic analysis of pangolins (although it 

was misattributed to USNM-P 299960). The other Patriomanis specimen with well-preserved postcrania, USNMP 299960, includes only disarticulated skull fragments, 

including a portion of the frontal bone, marked on the 

inside with grooves to accommodate the ethmoturbinals; 

the left posterolateral corner of the skull, including portions of the parietal, squamosal, petrosal, and occipital; 

and portions of the basisphenoid and basioccipital.
8 • 

manids, the posterior portions of the nasals together form 

a narrow V-shaped or U-shaped extension of variable 

width that deeply indents the anterior edge of the frontal 

(Figure 5a).

Maxilla

Emry (2004) provides a very extensive description 

of the maxilla in Patriomanis, noting its characteristic 

pholidotan morphology, as indicated by the complete 

loss of the upper dentition, and the presence of both an 

alveolar sulcus and a median longitudinal palatal trough 

(Figure 4b,d). All three are basal apomorphies of Pholidota (Gaudin et al., 2009). Additional preparation of the 

posterior reaches of the palate failed to reveal any remains 

of the palatine bones, so the location and form of both 

the maxillary–palatine suture and the palatine foramina 

cannot be ascertained. Similarly, further preparation in the 

anteroventral orbital wall yielded only a few isolated bone 

fragments, leaving it unclear whether Patriomanis, like 

nasal is quite deep (Figures 3, 4a,c), like that of Manis

and Smutsia, whereas the notch is present but much shallower in Phataginus (Figure 5a,c) and Eomanis waldi, the 

only other fossil pangolin that preserves the anterior nasal 

region (Gaudin et al., 2009). Emry (2004) notes that the 

posterior parts of both nasals are missing in Patriomanis. It appears, however, that on the left side of USNM-P 

494439 the upper sutural edge of the maxilla, that is, 

the portion in contact with the nasal, is preserved intact. 

The right maxilla is not as well preserved (Figure 3) yet 

in outline appears roughly symmetrical with the left in 

dorsal view (Figure 4a,c). In addition, the matrix underlying the missing right nasal bears a ridge that may represent a cast of the underside of the frontonasal suture. If 

the posterior region of the nasal is reconstructed on this 

basis (Figure 4c), it would appear to flare posteriorly and 

form a broad, shallow U-shaped suture with the frontal. 

This is very reminiscent of the posterior nasal morphology illustrated by Koenigswald and Martin (1990: fig. 

3a,b) for E. waldi and Necromanis edwardsi. In extant 

FIGURE 3. Skull of Patriomanis american

(a) USNM-P 494439 in right lateral view; (b) 

USNM-P 494439 in left lateral view; (c) reconstruction of the skull in left lateral view. Abbreviations: 

as, alisphenoid; ef, ethmoid foramen; f, frontal; iof, 

infraorbital foramen; ipa, interparietal; l, lacrimal; 

lf, lacrimal foramen; mx, maxilla; n, nasal; nc, 

nuchal crest; oc, occipital; occ, occipital condyle; 

of, optic foramen; os, orbitosphenoid; p, parietal; 

pal, palatine; pgf, postglenoid foramen; pm, premaxilla; pt, pterygoid; sf, supraorbital foramen; sq, 

squamosal; zp, zygomatic process of squamosal. 

Stippled areas represent matrix. Scale bar = 1 cm. 

Illustrations by Julia Morgan Scott for Smithsonian 

Institution.
n u m b e r 9 8 • 9

FIGURE 4. Skull of Patriomanis american

(a) USNM-P 494439 in dorsal view; (b) USNM-P 

494439 in ventral view; (c) reconstruction of the 

skull in dorsal view; (d) reconstruction of the skull 

in ventral view. Abbreviations: as, alisphenoid; asu, 

alveolar sulcus; bo, basioccipital; bs, basisphenoid; 

ctpp, caudal tympanic process of petrosal; ef, ethmoid foramen; eo, exoccipital region of occipital 

bone; er, epitympanic recess; f, frontal; fc, fenestra 

cochleae; fev, fenestra vestibuli; fm, foramen magnum; fo, foramen ovale; hf, hypoglossal foramen; 

hy, hyoid fragment; if, incisive foramen; iof, infraorbital foramen; ipa, interparietal; ips, opening for

inferior petrosal sinus; jf, jugular foramen; met, 

median trough; mx, maxilla; n, nasal; nc, nuchal 

crest; occ, occipital condyle; of, optic foramen; os, 

orbitosphenoid; p, parietal; pf/cf, piriform fenestra/

carotid foramen; pff, primary facial foramen; pgf, 

postglenoid foramen; pm, premaxilla; pop, paraoccipital process of petrosal (= mastoid process of 

Evans and Christensen, 1979); pr, promontorium 

of petrosal; prs, presphenoid; pt, pterygoid; ptc, 

posttympanic crest; ptp, posttympanic process of 

squamosal; sof/fr, fused sphenorbital fissure and 

foramen rotundum; sq, squamosal; stmn, stylomastoid notch; zp, zygomatic process of squamosal; 

zpm, zygomatic process of maxilla. Stippled areas 

represent matrix. Scale bar = 1 cm. Illustrations by 

Julia Morgan Scott for Smithsonian Institution.
1 0 • 

modern pangolins, lacked an orbital exposure of the maxilla (Gaudin, 2004; this study shows that an orbital wing 

of the maxilla is also missing in Metacheiromys and vermilinguan anteaters). The infraorbital foramen lies toward 

the back of the facial process of the maxilla in Patriomanis (Figure 3). It is displaced forward in Smutsia and even 

more so in E. waldi, where it is located near the anteroposterior midpoint of the bone (Gaudin et al., 2009).

Palatine

The only portion of the palatine that is preserved in 

USNM-P 494439 is a portion of perpendicular process (= 

orbital wing of Gaudin et al., 2009). We believe there are 

fragments preserved of the perpendicular process on both 

the left and right sides, although it is difficult to determine which portions of these fragments pertain to the 

FIGURE 5. Skulls and mandible of modern pangolins: (a) Phataginus tricuspis, UTCM 1695, in dors

view; (b) Manis javanica, USNM 198852, in ventral 

view; (c) Phataginus tricuspis, UTCM 1695, in right 

lateral view; (d) mandible of Phataginus tricuspis, CM 

86715, in right dorsolateral view. Abbreviations: as, 

alisphenoid; asu, alveolar sulcus; bo, basioccipital; bs, 

basisphenoid; cf, carotid foramen; cp, condylar process; cpf, caudal palatine foramen; eam, external auditory meatus; ec, ectotympanic; ef, ethmoid foramen; f, 

frontal; fev, fenestra vestibuli; fo, foramen ovale; hf, 

hypoglossal foramen; if, incisive foramen; iof, infraorbital foramen; jf, jugular foramen; l, lacrimal; met, 

median trough; mf, mental foramen; mndf, mandibular foramen; mnp, mandibular prong; ms, mandibular symphysis; mx, maxilla; mxf, maxillary foramen; 

n, nasal; nc, nuchal crest; oc, occipital; occ, occipital 

condyle; of, optic foramen; os, orbitosphenoid; p, 

parietal; pal, palatine; pet, petrosal; pgf, postglenoid 

foramen; pm, premaxilla; prs, presphenoid; pt, pterygoid; so, supraoccipital region of occipital bone; sof, 

sphenorbital fissure; spf, sphenopalatine foramen; sq, 

squamosal; stmf, stylomastoid foramen; zp, zygomatic 

process of squamosal; zpm, zygomatic process of maxilla. Scale bars = 1 cm. Panels (b) and (c) modified from 

Gaudin et al. (2009). Illustrations by Julia Morgan 

Scott for Smithsonian Institution.
n u m b e r 9 8 • 1 1

accommodates the origin of the inferior oblique muscle in 

other mammals (Wible and Gaudin, 2004).

It is noteworthy that the lacrimal foramen is missing 

in all modern pangolins, and the bone itself is absent in 

both Manis and Smutsia (Figure 5; Jollie, 1968; Gaudin 

et al., 2009; the lacrimal is indistinguishably fused to the 

maxilla in these genera according to Grassé, 1955). In this 

regard, Patriomanis is more similar to Eomanis waldi and 

nonpholidotan eutherians (Evans and Christensen, 1979; 

Novacek, 1993). Interestingly, although the bone is absent, 

the lacrimal fenestra is at least variably present in all extant 

species of Manis and Smutsia, whereas it is absent in both 

species of Phataginus, despite the fact that both species 

retain a separate lacrimal element (Jollie, 1968; Gaudin 

et al., 2009). Whether E. waldi possessed this opening is 

unknown (Gaudin et al., 2009). In Phataginus, the orbital 

portion of the lacrimal is much larger than its almost nonexistent facial exposure (Figure 5c). The facial exposure 

is also small in the palaeanodont Metacheiromys and 

the enigmatic edentate Eurotamandua, whereas the large 

facial process of the lacrimal in Patriomanis is more like 

the condition in cingulate and vermilinguan xenarthrans 

(Gaudin and Branham, 1998; Gaudin, 2004).

Jugal

Emry (2004) describes a sutural surface for the jugal 

carried dorsally on the zygomatic process of the maxilla, 

although the bone itself is not known to be preserved 

in any Patriomanis specimen. Because of this, it is not 

known definitively whether Patriomanis had a complete 

zygomatic arch like E. waldi (Storch, 2003) and some M. 

pentadactyla (Emry, 2004; we also observed a complete 

zygoma in one specimen of M. javanica, USNM 14448) 

or an incomplete zygoma like all remaining modern pangolins (Figure 5). However, we are reconstructing a complete zygoma on the basis of the presence of the jugal bone 

(Figures 3 and 4; but see discussion below of the zygomatic process of the squamosal). The jugal is present in 

E. waldi and many, if not most, of those M. pentadactyla specimens with a complete zygomatic arch (it is presumably synostosed to the maxilla and squamosal in the 

remaining examples). The other extant pangolin species 

lack a jugal and are characterized by an incomplete arch 

(except for the aforementioned specimen of M. javanica, 

with both a jugal and complete zygoma, and M. crassicaudata FMNH 98232, with the jugal attached to maxilla but 

an incomplete zygoma). Weber (1928) suggests the bone is 

fused early in ontogeny to the squamosal, whereas Grassé 

(1955) asserts that it is fused to the maxilla.

palatine and which to the pterygoid. In modern pangolins, 

the suture between the perpendicular process of the palatine and the pterygoid lies at or just posterior to the level 

of the sphenorbital fissure (Figure 5; Jollie, 1968; Gaudin 

et al., 2009: char. 376). There is a crack in approximately 

this position on the larger, better-preserved palatine fragment from the left side, suggesting that roughly the anterior 

two-thirds of this piece is from the palatine’s perpendicular 

process. Moreover, the preserved morphology suggests that 

the palatine of Patriomanis has a low, anteroposteriorly 

elongate exposure in the ventral wall of the orbit (Figure 3) 

as in extant pangolins (Novacek and Wyss, 1986; Gaudin, 

2004). The left palatine fragment ends anteriorly at a broken surface marked by two grooves that presumably led 

to foramina, the upper being the sphenopalatine foramen 

and the lower the caudal palatine foramen (Jollie, 1968). 

These foramina were likely completely enclosed within the 

perpendicular process of the palatine, as they are in extant 

taxa (Jollie, 1968). 

Although the anterior parts of the perpendicular process and the entire horizontal process are missing, the 

palatine certainly possessed its normal contacts with the 

horizontal process of the maxilla and at least the base of 

the maxilla’s zygomatic process if an orbital exposure of 

the maxilla was indeed lacking (see above). In addition, 

USNM-P 494439 preserves dorsal contacts between the 

perpendicular process and the frontal and orbitosphenoid 

and, as noted, likely a posterior contact with the pterygoid 

(Figure 3), as in extant pangolins (Figure 5; Jollie, 1968). 

The orbital portion of the palatine is unknown in other 

fossil pangolins.

Lacrimal

The lacrimal of USNM-P 494439 is preserved only on 

the left side of the skull (Figure 3b,c). In lateral view it is a 

triangular element with its apex pointing anteriorly. It has a 

large facial process extending forward from the prominent, 

rounded orbital rim that crosses the bone dorsoventrally 

and a smaller semicircular orbital exposure that houses a 

small lacrimal foramen immediately posterior to the orbital 

rim. It contacts the zygomatic process of the maxilla anteroventrally and the frontal both dorsally and posteriorly. It 

could conceivably have had a small, posteroventral contact 

with either the maxilla or, more likely, the palatine within 

the orbit, but that portion of the orbital wall is badly damaged on both sides of the skull and does not permit a more 

definitive assessment. There is a small pit between lacrimal 

and frontal within the orbit, near the posteroventral corner of the former. This is likely the lacrimal fenestra that 
1 2 • 

of no record of its contents in pangolins, but in xenarthrans 

and carnivorans, this is an opening that houses the frontal diploic vein (Evans and Christensen, 1979; Wible and 

Gaudin, 2004), whereas in humans it transmits the supraorbital nerve (a branch of the ophthalmic nerve, CN V1) and 

its accompanying vessels (Pick and Howden, 1977). 

The frontal of Patriomanis is not marked by a postorbital process or temporal lines on the skull roof. These 

features are also absent in all known members of Pholidota, 

including Eurotamandua (Gaudin et al., 2009). This likely 

indicates that the temporalis muscle was strongly reduced in 

Patriomanis, although perhaps not to the same degree as in 

the modern forms, where Doran and Allbrook (1973:997; 

see also Weber, 1928) note “the almost total absence of 

masticatory muscles.” Patriomanis also lacks a postorbital 

constriction and has instead a weak interorbital constriction in dorsal view (Figure 4c). This is the condition in all 

extant pangolins (Figure 5; except for some S. gigantea), 

whereas in other fossil taxa, namely, Necromanis, Eomanis, 

and Eurotamandua, a separate postorbital constriction is 

present (Gaudin and Wible, 1999; Gaudin et al., 2009).

Parietal

Although the parietals are preserved in their entirety 

in the type, Emry (1970) says little about them, apart 

from noting their sutural contacts with the frontal anteriorly and the occipital and interparietal posteriorly. In 

addition, they contact the squamosal laterally at an elongate suture (length ~30% GSL; Table A1). This suture 

is similarly elongated in extant Phataginus and some S. 

temminckii (Figure 5c; length >25% GSL), whereas the 

suture is much shorter in other Smutsia, in Manis, and in 

E. waldi, Eurotamandua, and other out-groups (Gaudin 

et al., 2009; Table A1). The frontoparietal suture is situated at or just behind the level of the zygomatic process 

of the squamosal (Figures 3c, 4c), as in all other pangolins except E. waldi (and Eurotamandua; Gaudin et al., 

2009). The parietals themselves are relatively flat dorsally 

and in their anteroposterior profile and strongly convex 

laterally, and their lateral margins are slightly convex in 

dorsal view. USNM-P 494439 adds no real information 

to the description of the parietal since the bones are badly 

cracked and distorted on the left, and much of the medial 

and anterior portions are entirely missing on the right.

Interparietal

Emry (1970) notes the presence of a large, triangular interparietal element in Patriomanis that is suturally 

Frontal

Only the posterodorsal regions of the frontal bone 

are preserved in the type (F:AM 78999), from the skull 

roof immediately behind the interorbital constriction to 

the frontoparietal suture (Emry, 1970), which, in turn, lies 

just anterior to the level of the zygomatic process of the 

squamosal. Consequently, Emry (1970) has little to say 

about its morphology. The frontal is damaged on both 

sides of USNM-P 494439, especially on the right, where 

large chunks are missing from the skull roof and dorsal 

orbital wall, whereas on the left the bone is present in its 

entirety but cracked and distorted. Nevertheless, its morphology and sutural contacts can be largely reconstructed 

on the basis of the available material (Figures 3, 4).

On the rostrum, the frontal contacts the nasals anteriorly (see above), the maxilla anterolaterally, and the lacrimal laterally (Figures 3c, 4c). The frontal has an extensive 

exposure in the medial wall of the orbit, being by far the 

largest component of the orbital mosaic, as is the case in 

modern manids (Figure 5; Jollie, 1968; Novacek and Wyss, 

1986). Within the orbit it contacts the lacrimal anteriorly, 

the palatine and orbitosphenoid laterally, and the squamosal and parietal posteriorly (Figure 3c). The dorsal portion 

of the alisphenoid closely approaches, but appears not to 

contact, the frontal (Figure 3c) because of the presence of 

a small orbitosphenoid–squamosal contact (Gaudin and 

Wible, 1999; Gaudin et al., 2009; see also fig. 7.1 in Rose 

and Emry, 1993). A similar condition prevails in E. waldi, 

Phataginus, S. gigantea, and some S. temminckii, whereas 

in other S. temminckii and in Manis the orbitosphenoid 

and squamosal share a broad sutural contact (Figure 5b; 

Gaudin and Wible, 1999; Gaudin et al., 2009). Because of 

poor preservation (see above), it is not possible to determine whether there was also an anterolateral contact with 

the maxilla in the orbital wall. 

The orbital surface of the frontal is smooth except for 

a single relatively large ethmoid foramen located entirely 

within the frontal but close to the frontal–orbitosphenoid 

suture, anterodorsal to the optic foramen (Figures 3c, 4d). 

In the older extinct forms Eomanis and Eurotamandua, 

the ethmoid foramen actually lies on the suture between 

the frontal and orbitosphenoid, as it does in the extant species M. pentadactyla and M. javanica and variably so in M. 

crassicaudata, S. temminckii, and P. tricuspis (Gaudin et al., 

2009). As noted above, the frontal also participates in the 

lacrimal fenestra. In addition, just below the dorsal rim of 

the orbit, midway between the lacrimal and ethmoid foramina, is a very small supraorbital foramen. This foramen is 

also known in modern pangolins (Gaudin, 2004). We know 
n u m b e r 9 8 • 1 3

(Figure 4b), but given its rodlike shape and the fact that 

it is broken at either end, we interpret this as a displaced 

portion of the hyoid arch (probably stylohyal) rather than 

a portion of the mandible.

The distal tip of the zygomatic process resembles the 

morphology in modern pangolins and differs from that 

in E. waldi, Eurotamandua, and metacheiromyid palaeanodonts, in which the process is laterally and anteriorly 

directed, relatively short, and inflated at its base and terminates in a sharp point (Gaudin et al., 2009; see also 

Storch, 2003: fig. 1). The weathered condition of the tip in 

both the type and USNM-P 494439 leaves open the possibility that the zygomatic process is broken off in both 

specimens and thus incompletely preserved. Although its 

shape is like that of modern pangolins, the likely presence of a jugal bone (see above), lacking in modern forms, 

raises the possibility of a complete zygoma in Patriomanis. However, if a complete zygoma were present, a more 

distinct from the occipital (Figure 4c), whereas in modern pangolins the interparietal is fused to the occipital 

(marked as supraoccipital region of occipital in Figure 5; 

Jollie, 1968). The condition is unknown in other fossil 

pangolins except E. waldi, where a separate interparietal 

is not apparent (Gaudin, unpublished data). As discussed 

by Rose and Emry (1993), Novacek and Wyss (1986) had 

suggested the lack of an interparietal was a derived feature 

linking xenarthrans and pholidotans, before adding a note 

acknowledging its presence in pangolins based on Emry 

(1970) and Jollie (1968). The apical angle of the interparietal in Patriomanis is slightly obtuse (approximately 

100°), as in modern Manis (where it is fused to the supraoccipital in adults, as noted by Jollie, 1968), whereas in 

Phataginus and some specimens of either Smutsia species, 

it forms a right angle or even a slightly acute angle (Figure 

5a,c; Gaudin and Wible, 1999; Gaudin et al., 2009). 

Squamosal

The squamosals are preserved nearly intact in the 

type specimen (Emry, 1970) and on the right in USNM-P 

494439, whereas on the left the bone is broken and distorted, and its posterodorsal corner is missing (Figure 3). 

The zygomatic process is the only portion of the squamosal where the morphology is questionably known. 

The best-preserved process is on the left side of USNM-P 

494439, although there is also a substantial process preserved on the right in the type. The latter is described by 

Emry (1970:468) as follows: “The zygomatic processes 

are small, directed downward, and, although somewhat 

weathered, appear to have ended in a rather sharp point. 

There was apparently no glenoid articular surface.” The 

description fits the morphology of USNM-P 494439 well 

enough, although the zygomatic process in that specimen 

is somewhat more elongated ventrally and comes to a narrower point. The absence of a distinct glenoid fossa would 

match the morphology in modern pangolins, where the 

mandibular condyle “[contacts] lightly with the ventromedial side of the zygomatic process” (Emry, 1970:468); that 

is, it has a ligamentous attachment to the inner base of 

the process (and perhaps to the lateralmost portion of the 

alisphenoid) but does not attach to the moderately rugose, 

somewhat expanded distal end of the process (see P. tricuspis, UTCM 1695, Figure 6a; note that the distal tip of 

the zygomatic process is scored erroneously as an anteroposteriorly convex “glenoid fossa” in Gaudin and Wible, 

1999: char. 51, and in Gaudin et al., 2009: char. 356). The 

left zygomatic process of USNM-P 494439 has a rodlike 

bone fragment attached by matrix to this “glenoid” region 

FIGURE 6. Skull of Phataginus tricuspis and skull fragment 

Patriomanis americana: (a) skull of Phataginus tricuspis (UTCM 

1695) in ventrolateral view; (b) skull fragment of Patriomanis 

americana (USNM-P 299960) showing cast of ethmoturbinals in 

dorsal view. Abbreviations: eb, ectotympanic bulla; etb, ethmoturbinal; ja, jaw articulation; mand, mandible; nsp, nasal septum; zp,

zygomatic process of squamosal. Scale bars = 1 cm. Photos by Tim 

Gaudin, composed and labeled by Julia Morgan Scott for Smithsonian Institution.
1 4 • 

As in these taxa, the first arch is undoubtedly the porus 

acousticus, the arch that would accommodate the external 

auditory meatus. Dorsal and medial to the ventromedial 

edge of the porus is a small shelf that forms the squamosal 

contribution to the roof of the epitympanic recess. The 

first process immediately behind the porus and carrying 

an anteroventral facet for the missing posterior crus of 

the ectotympanic is likely what Wible et al. (2004) and 

Wible (2008) label the “posttympanic crest,” although in 

Patriomanis it abuts both the tympanohyal and the large, 

blocky paroccipital process of the petrosal (sensu Wible 

and Gaudin, 2004; equal “mastoid process” of Evans and 

Christensen, 1979) and is enlarged and rugose laterally, 

as if it may have served as a site for muscle attachment. 

In Zalambdalestes and Solenodon, it has the tympanohyal 

contact but is separated from the paroccipital process by 

the stylomastoid notch for cranial nerve VII, and it lacks 

any lateral expansion (Wible et al., 2004; Wible, 2008). 

The arch behind the posttympanic crest is semicircular 

and is well lateral to the stylomastoid notch of Patriomanis. It accommodates a significant lateral exposure of 

the paroccipital process of the petrosal, that is, a lateral 

mastoid exposure, a feature missing in modern pangolins (as noted above; Novacek and Wyss, 1986; Patterson 

et al., 1992) but present in E. waldi, Eurotamandua, and 

palaeanodonts (Gaudin et al., 2009). However, there is a 

ventral groove on the right paroccipital process of USNMP 494439 that leads toward the back edge of this arch, 

which may imply that the arch is a kind of auxiliary stylomastoid notch. Immediately behind this second arch is 

a second process, smaller than the posttympanic crest and 

much smaller in USNM-P 494439, where it is preserved 

only on the right side and even then may be damaged distally, than in the type, where it is preserved apparently 

intact on the right but not the left. This process abuts the 

paroccipital process medially and posteriorly; hence, we 

identify it as the posttympanic process of the squamosal 

on the basis of its similarity to the morphology in Zalambdalestes and Solenodon (Wible et al., 2004; Wible, 2008). 

We make this conclusion despite the fact that in Patriomanis there is a tall, anteroposteriorly narrow arch separating the posttympanic process from the lower end of the 

nuchal crest on the occipital bone. This arch is not present 

in either Zalambdalestes or Solenodon. This third arch in 

the sequence accommodates a second, smaller lateral mastoid exposure and could conceivably have housed a tendon of insertion for the sternomastoid (the cleidomastoid 

is absent in modern pangolins; see Windle and Parsons, 

1899), given that sternomastoid and cleidomastoid insert 

anterolaterally elongated zygomatic process with a facet 

for the jugal would be expected, like that of E. waldi. The 

ventrally directed process in the two Patriomanis specimens shows no trace of a facet for the jugal as preserved. 

Thus, the evidence for a complete zygoma in Patriomanis 

is ambiguous, although we have chosen to reconstruct its 

presence in Figures 3 and 4.

The posterolateral surface of the zygomatic process is 

marked by a clearly delineated depression, the postglenoid 

fossa. The lateral margin of the fossa is defined by a strong 

ridge that passes from the tip of the zygomatic process (as 

preserved) posterodorsally, arching over the porus acousticus to terminate at the base of a prominent squamosal 

posttympanic crest (see below). In the middle of the postglenoid fossa is a large opening, the postglenoid foramen 

(Figures 3a,c, 4b,d, and 7), which in cingulate xenarthrans 

and other placental mammals transmits the capsuloparietal emissary vein (Evans and Christensen, 1979; Wible 

and Gaudin, 2004; this vein is one of the major drainage 

routes for the transverse sinus in eutherians; see Wible, 

1990). In USNM-P 494439, there is an additional small 

foramen of unknown function medial to the postglenoid 

foramen (Figure 7). The postglenoid foramen is present 

in all pholidotans (except Eurotamandua; Gaudin et al., 

2009) and in Palaeanodon but not Metacheiromys (Patterson et al., 1992), but its position varies. In Patriomanis 

and Phataginus tetradactyla, it lies posterior to the base 

of the zygomatic process. In E. waldi, Smutsia, M. pentadactyla, and some M. crassicaudata it lies on the lateral 

aspect of the process itself, whereas in M. javanica, some

M. crassicaudata, and P. tricuspis it lies on the posterior 

aspect of the zygomatic process (Figure 5; Gaudin and 

Wible, 1999; Gaudin et al., 2009).

The portion of the squamosal bordering the ear region 

is quite different from that in modern manids. Patriomanis lacks the postauditory inflation of the squamosal 

characteristic of the extant taxa (Figure 5; Gaudin and 

Wible, 1999: char. 48, figs. 2, 4; Gaudin et al., 2009: char. 

353; see also illustrations in Emry, 1970: figs. 9B, 10A), 

an inflation that largely hides the mastoid portion of the 

petrosal from external view (Novacek and Wyss, 1986; 

Patterson et al., 1992). Instead, the posteroventral squamosal margin, that is, the postglenoid area, is marked by 

three arches and two processes (Figures 3, 4b,d, and 7). 

This arrangement is quite reminiscent of that described 

for the late Cretaceous eutherian Zalambdalestes by Wible 

et al. (2004) and, to a lesser extent, that of the extant West 

Indian eulipotyphlan Solenodon paradoxus (Hispaniolan 

solenodon) described by Wible (2008). 
n u m b e r 9 8 • 1 5

they are covered by matrix and would be difficult to expose 

without damaging more superficial elements. The fragment 

from USNM-P 299960 preserves impressions of four ethmoturbinal scrolls on each side of the skull, but extant taxa 

are known to possess six ethmoturbinals (Grassé, 1955). 

Modern pangolins are one of several placental orders in 

which a separate mesethmoid ossification makes a sizable 

contribution to the osseous nasal septum (Novacek and 

Wyss, 1986; Novacek, 1993). This morphology is also 

known in carnivorans, the sister taxon to Pholidota in 

molecular-sequence-based placental phylogenies (Meredith 

et al., 2011) or combined morphology- and molecularbased phylogenies (O’Leary et al., 2013) but not xenarthrans (but see Ferigolo, 1981), a group often allied with 

Pholidota in exclusively morphology-based phylogenies 

(Gaudin and McDonald, 2008; O’Leary et al., 2013).

on the posterior surface of the posttympanic process in 

Solenodon (Wible, 2008). 

Like most placental mammals, the squamosal contacts the parietal dorsally, the occipital (i.e., exoccipital) 

posteriorly, the frontal anteriorly, and the alisphenoid and 

petrosal medially. As noted above, there is also a small 

anterior contact with the orbitosphenoid.

Ethmoid and Vomer

Neither the ethmoid nor vomer is available for study in 

any of our Patriomanis specimens, apart from some impressions of the ethmoturbinals on the underside of a fragment 

of the frontal bone preserved in USNM-P 299960 (Figure 

6b). The type is broken posterior to these elements since 

the entire rostrum is absent, whereas in USNM-P 494439 

FIGURE 7. Close-up of right auditory region of Patriomanis americana, USNM-P 494439, in ventral view: (a) pencil drawing; (b) labeled line drawing. Abbreviations: bo, basioccipital; bs, basisphenoid; cc, cochlear canaliculus; cp, crista parotica; ctpp, caudal tympanic process of petrosa

eo, exoccipital; er, epitympanic recess; fc, fenestra cochleae; fev, fenestra vestibuli; fi, fossa incudis; 

fs, facial sulcus; ips, inferior petrosal sinus; jf, jugular foramen; nc, nuchal crest; pag, promontorial 

artery groove; pff, primary facial foramen; pgf, postglenoid foramen; pop, paraoccipital process of 

petrosal (= mastoid process of Evans and Christensen, 1979); pr, promontorium of petrosal; pt, 

pterygoid; ptc, posttympanic crest; ptp, posttympanic process of squamosal; stf, stapedius fossa; sq, 

squamosal; stmn, stylomastoid notch; th, tympanohyal; ttf, m. tensor tympani fossa; ?, unknown 

foramen. Stippled areas represent matrix. Scale bar = 1 cm. Illustrations by Julia Morgan Scott for 

Smithsonian Institution.
1 6 • 

are missing from the type specimen. The left alisphenoid 

of USNM-P 494439 is damaged posteriorly, with a broad 

transverse crack running through the foramen ovale, and 

the posterior reaches of the bone appear to be lost. The 

right alisphenoid is not preserved in USNM-P 494439.

The main body of the alisphenoid is nearly transversely oriented, extending from its lateral contact with 

the squamosal along the inner base of the zygomatic process to its medial contact with the pterygoid (Figure 4d). 

At its anterior end, it becomes more vertical, forming the 

semicircular lateral margin of the sphenorbital fissure (= 

anterior lacerate foramen of Emry, 1970), as well its lateral and ventral walls. The floor of the fissure contacts 

the orbitosphenoid–presphenoid medially. It appears to be 

broken anteriorly but must have extended far enough forward to contact the perpendicular process of the palatine. 

As noted previously, there is a crack that may represent 

the palatine–pterygoid suture—it extends as far back as 

the posteriormost point on the sphenorbital fissure’s lateral margin. If it is indeed the suture, then there is a clear 

orbital contact between alisphenoid and palatine. The 

dorsalmost portion of the alisphenoid extends upward 

between the orbitosphenoid and palatine, terminating 

where the orbitosphenoid and squamosal briefly contact. 

As noted by Rose and Emry (1993), there is no alisphenoid–parietal contact. The contact is absent in all extant 

manids as well but is present in E. waldi and metacheiromyid palaeanodonts (Gaudin et al., 2009). The posterior 

margin of the alisphenoid appears to contact the petrosal in USNM-P 494439, but this contact is likely due to 

postmortem breakage and distortion of the skull. In the 

type, where the posterior edge appears unbroken and the 

specimen is less distorted, there seems to be no contact 

with the petrosal, unless it is with the lateralmost extremity of the posterior edge. Instead, the alisphenoid forms 

the anterior edge of a confluent carotid foramen (= median 

lacerate foramen of Emry, 1970, comprising the medial 

half of the opening) and piriform fenestra (Figure 4d). The 

carotid foramen–piriform fenestra is present in a newborn 

M. javanica, but the fenestra is obliterated by an extensive 

suture between the alisphenoid and petrosal in the adult 

(Figure 5b; Jollie, 1968; see also Figure 8).

As in other mammals, several of the large orbital 

foramina are associated with the alisphenoid (Figure 4d). 

These have been described by Emry (1970), who notes 

that the foramen rotundum (for the maxillary nerve, CN 

V2) is absent in Patriomanis, becoming confluent instead 

with the sphenorbital fissure (for CN III, IV, V1, VI and 

several major orbital arteries and veins), as in modern 

Manidae (Figure 5; Jollie, 1968; a separate foramen 

Orbitosphenoid and Presphenoid

Emry (1970) makes no mention of the orbitosphenoid 

since the type preserves no more than the posterodorsalmost fragments of the bone. Our additional preparation 

in the orbital region of USNM-P 494439, however, has 

revealed a sizable orbitosphenoid exposure in the medial 

wall of the orbit, as in modern pangolins (Jollie, 1968; 

Novacek, 1993: fig. 9.22C; Gaudin, 2004). The exposed 

surface is triangular with its apex dorsad (Figures 3c, 4d). 

In the middle of the bone is a large optic foramen, and 

the posteroventral portion of the orbitosphenoid forms 

the medial wall of the sphenorbital fissure. The optic foramen is similarly large in E. waldi, Phataginus, and S. temminckii but is relatively smaller in Manis, S. gigantea, and 

various out-groups (Gaudin et al., 2009).

The orbitosphenoid is sutured to the frontal along 

its anterodorsal edge and to the palatine along its ventral 

edge (where dorsal connections between the palatine and 

the frontal and alisphenoid clearly exclude any connection 

between orbitosphenoid and maxilla or pterygoid). Its posterodorsal edge is sutured to the frontal, squamosal, and 

alisphenoid. The connection to the squamosal is lacking 

in the out-groups and in Eurotamandua but is present in 

E. waldi and all of Manidae and is particularly broad in 

members of the genus Manis (Figure 5b; Gaudin et al., 

2009). The right and left orbitosphenoids are fused in the 

ventral midline to form a rodlike presphenoid element that 

was likely exposed in the roof of the nasopharynx (visible 

in USNM 494439 but hidden in ventral view by matrix and 

bone fragments in Figure 4b). As in M. javanica (Figure 

5b; Jollie, 1968), the presphenoid would be sutured to the 

basisphenoid posteriorly and the palatine laterally. It may 

also have had a posterolateral connection to the pterygoid 

and an anterior connection to the vomer, but this cannot be 

confirmed with the material presently in hand.

Alisphenoid

Despite the fact that most of the left and right alisphenoid bones are preserved in the type specimen (F:AM 

78999), Emry’s (1970) description focuses on the foramina in and around these elements rather than a description 

of the bones themselves. USNM-P 494439 provides additional information beyond what was preserved in the type, 

in that it retains intact the anteriormost portion of the 

bone on the left side of the skull, that is, the portion forming the lateral wall of the sphenorbital fissure, as well as 

some vestige of the bone’s ventral connections to the palatine and pterygoid, whereas the palatines and pterygoids 
n u m b e r 9 8 • 1 7

Pterygoid

The pterygoid bones are not mentioned by name in 

Emry’s (1970) description. Emry (1970:468) does note 

that the “sphenoidal wings that extend the narial channel backward to the basioccipital in Manis were apparently also present in Patriomanis, but the ventral part has 

been broken away.” These so-called basisphenoid wings 

(Gaudin and Wible, 1999) were shown by Gaudin et al. 

(2009; see description of char. 347) to be tympanic processes of the pterygoid, not parts of the basisphenoid. In 

USNM-P 494439, the tympanic process of the pterygoids 

appears to be partially preserved (Figures 3, 4), and as in 

the type, the pterygoids appear to terminate just in front 

of the basisphenoid–basioccipital suture (the suture is preserved in USNM-P 494439; see below), where they are 

rotundum is retained in palaeanodonts; Gaudin et al., 

2009). The sphenorbital fissure lies between the orbitosphenoid and alisphenoid. Rose and Emry (1993) note 

that the foramen ovale (for the mandibular nerve, CN V3) 

lies wholly within the alisphenoid in Patriomanis. It is 

located centrally within the main body of this element. 

This is also the condition in palaeanodonts, Phataginus, 

and M. crassicaudata, whereas a squamosal contribution 

to the lateral margin of the foramen ovale is known to 

occur in S. temminckii, M. pentadactyla, and some specimens of S. gigantea and M. javanica (Gaudin et al., 2009). 

In some modern pangolins, the foramen ovale is also situated more posteriorly, even with the anterior edge of the 

ectotympanic (M. pentadactyla, M. crassicaudata, some 

specimens of S. temminckii and P. tricuspis; Gaudin et al., 

2009). 

FIGURE 8. Close-up of left auditory region of Smutsia gigantea, AMNH 53858, in ventral view: (a) pencil drawin

(b) labeled line drawing. Abbreviations: as, alisphenoid; bo, basioccipital; bs, basisphenoid; cf, carotid foramen; cp, crista 

parotica; cs, sulcus for internal carotid artery; en, entotympanic; eo, exoccipital; er/es, epitympanic recess/epitympanic 

sinus; fc, fenestra cochleae; fev, fenestra vestibuli; fo, foramen ovale; fs, facial sulcus; ips, inferior petrosal sinus; jf/hf, confluent jugular foramen and hypoglossal foramen; pe, petrosal (paroccipital or “mastoid” exposure); pr, promontorium 

of petrosal; pt, pterygoid; sq, squamosal; stmf, stylomastoid foramen; th, tympanohyal. Scale bar = 1 cm. Modified from 

Gaudin and Wible (1999). Illustrations by Julia Morgan Scott for Smithsonian Institution.
1 8 • 

2004; = anterior portion of greater petrosal nerve in Pick 

and Howden, 1977; Evans and Christensen, 1979). This 

groove is also present on the left side of USNM-P 494439. 

In both the type and USNM-P 494439, it originates posterior to the tympanic process of the pterygoid, opposite the 

inferior petrosal sinus opening identified by Emry (1970). 

As Emry (1970) notes, this origin, which is shared with 

modern manids, is posterior to its more typical position in 

other eutherians, where it emerges from the carotid foramen (see, e.g., Homo, Pick and Howden, 1977 [emerges 

from foramen lacerum, homolog of carotid foramen as 

used here]; Dasypus, Wible, 2010; Solenodon, Wible, 

2008; in Canis it emerges from a separate hiatus Fallopii 

just lateral to the carotid foramen, Evans and Christensen, 

1979). The anterior termination of the groove, the pterygoid canal itself, is not preserved in Patriomanis. 

Emry (1970) labels three openings associated with the 

lateral edge of the basisphenoid–basioccipital complex in 

his fig. 9: the foramen for the inferior petrosal sinus, the 

jugular foramen (following usage of Wible and Gaudin, 

2004; = posterior lacerate foramen of Emry, 1970), and 

the hypoglossal foramen (following usage of Wible and 

Gaudin, 2004; = condylar foramen of Emry, 1970). He 

notes in the text, however, that the foramen assigned to 

the inferior petrosal sinus is continuous with a groove 

emerging from the hypoglossal foramen, a groove that 

would normally house either the internal carotid artery, 

the inferior petrosal sinus, or both but that would normally originate in the jugular rather than the hypoglossal foramen. The jugular foramen has been assigned to an 

opening lateral to this groove in Emry’s (1970) fig. 9. The 

morphology is indeed difficult to interpret. There appear 

to be openings on the left side of USNM-P 494439 (Figure 

4b,d) that correspond to those described by Emry (1970). 

There is a clearly marked, semiovate notch medial and 

slightly anterior to the basioccipital tuber that mirrors the 

inferior petrosal sinus opening (Figure 7). A second notch 

in the lateral margin of the basioccipital lies a short distance farther posteriorly and medially. This notch would 

appear to be the remnant of the hypoglossal foramen. 

There is no indication of the jugular foramen on the basioccipital; however, there is a vertical groove on the posteromedial surface of the petrosal. It houses the cochlear 

canaliculus (Figure 7), which typically opens in or near the 

jugular foramen (see, e.g., Canis, Evans and Christensen, 

1979; Dasypus, Wible, 2010); hence, this groove almost 

certainly forms the lateral margin to the jugular foramen. 

The petrosal groove is displaced laterally relative to the 

inferior petrosal sinus and hypoglossal foramina, which 

would again be consistent with Emry’s (1970) assessment 

fused to the lateral edge of the basisphenoid. This placement would suggest that in Patriomanis the auditory tube 

was displaced posteriorly and faced directly mediad, as 

in modern pangolins (Gaudin and Wible, 1999; Gaudin 

et al., 2009), and not anteromedially, as in most eutherians. 

This tympanic process of the pterygoid also likely formed 

the medial margin of the carotid foramen (= median lacerate foramen of Emry, 1970), whereas the alisphenoid and 

petrosal form the anterior and posterior margins of this 

opening, which, as noted above, is confluent laterally with 

the piriform fenestra.

The pterygoid hamulus is not preserved in any Patriomanis specimen. As was just noted, the pterygoid is fused 

to the basisphenoid posteromedially. It also was likely 

sutured to the palatine anteriorly and the alisphenoid 

dorsally and may also have attached to the presphenoid 

anteromedially.

Basisphenoid and Basioccipital

Emry (1970:468) describes these elements together, 

noting that they form a basicranial plate that is “rather 

narrow and decreases in width anteriorly.” Although the 

suture between the two is fused in the type, it is preserved 

in USNM-P 494439, where it assumes a shallow V shape 

with its vertex pointing anteriorly, and in a fragment 

from USNM-P 299960, where it is straight, the difference in shape perhaps ontogenetic. It is located roughly 

where Emry (1970) suggests, at the midpoint of the ear 

region, terminating laterally at the anteromedial rim of 

the purported foramen for the inferior petrosal sinus just 

anterior to a large pair of basioccipital tubera (Figure 

4c,d). Gaudin and Wible (1999) recognize small crests 

that extend posterolaterally from the tubera as tympanic 

processes of the basioccipital (= basioccipital “wings” of 

Gaudin and Wible, 1999). Although the processes are 

more extended ventrally and anteroposteriorly elongated 

in the modern forms, this characterization does not seem 

unreasonable. Immediately posterior to the basioccipital 

tubera in Patriomanis is a pair of shallow fossae. On the 

basis of the morphology in other eutherian mammals (see, 

e.g., Homo, Pick and Howden, 1977; Canis, Evans and 

Christensen, 1979; Euphractus, Wible and Gaudin, 2004; 

Solenodon, Wible, 2008), the basioccipital tubera likely 

served as attachment points for the longus capitis muscles, 

and the concavities immediately behind likely served as 

attachment sites for the rectus capitis ventralis muscles. 

Emry (1970) identifies a narrow groove on the ventrolateral margin of the basisphenoid as housing the vidian 

nerve (= nerve of the pterygoid canal in Wible and Gaudin, 
n u m b e r 9 8 • 1 9

better than the right, which is missing the entire ventral 

exposure of the pars canalicularis. Emry (1970) suggests 

that both petrosals have been displaced. He described 

several major foramina associated with the petrosal (see 

above) but not the bones themselves. Both left and right 

petrosals are much better preserved in USNM-P 494439 

and have undergone extensive preparation as part of this 

study. A close-up illustration of the right petrosal is provided (Figure 7) since it is slightly more complete than 

the left.

The ventral surface of the petrosal is dominated by a 

globose promontorium. A similar morphology is present 

in E. waldi and Metacheiromys, but in modern pangolins 

the promontorium tends to be strongly reduced, with the 

lateral wall of the petrosal pars cochlearis (i.e., the portion that houses the cochlea and sacculus) nearly flat in 

ventral view (Figure 8; Gaudin and Wible, 1999; Gaudin 

et al., 2009). On the right side of USNM-P 494439 there 

is a weak groove originating at the ventral margin of the 

fenestra vestibuli and traveling medially across the promontorium, nearly transverse to its long axis. If anything, 

the groove is even less in evidence on the left side. Modern pangolins lack a transpromontorial internal carotid 

or stapedial artery (Gaudin et al., 2009)—indeed, the stapedial system is missing entirely (Bugge, 1979). Instead, 

as noted above, the internal carotid travels in a carotid 

sulcus along the ventromedial edge of the petrosal. In contrast, the promontorial groove, weak though it is, suggests 

that at least a vestige of the stapedial arterial system may 

have been retained in Patriomanis (Figure 7). The promontorium also bears a distinct, dorsally situated pit along 

its anterolateral margin. This is almost certainly a tensor 

tympani fossa, that is, an attachment point for the tensor 

tympani muscle. The anterior edge of the fossa is likely 

formed by a very narrow epitympanic wing of the petrosal anterior to the promontorium. An epitympanic wing is 

present in modern pangolins as well, ranging from a small 

shelf like that of Patriomanis (e.g., P. tricuspis UTCM 

1695) to a prominent process (e.g., M. javanica USNM 

198852, FMNH 33550; see also Palaeanodon, Patterson 

et al., 1992).

 Immediately lateral and slightly anterior to the promontorium is a deep groove curving anteromedially, the 

facial sulcus, which houses the facial nerve (CN VII) and 

represents part of the ventral exposure of the pars canalicularis of the petrosal (i.e., the portion housing the semicircular canals and utricle). The anterior half of this sulcus 

is roughly horizontal and is bounded anterolaterally by a 

robust crista parotica, a condition found in some modern 

pangolins (P. tricuspis, some specimens of P. tetradactyla,

of the type specimen. Unfortunately, the morphology in 

both specimens is quite different from that in modern pangolins (Figures 5, 8), and the relevant features are not preserved in any other fossil pangolins. In the modern taxa, 

the internal carotid artery typically travels in a clearly 

marked sulcus on the ventromedial edge of the petrosal, 

the sulcus terminating anteriorly at the carotid foramen 

(Figure 8). If an inferior petrosal sinus foramen is present (as in Manis sp., S. gigantea, and some S. temminckii;

Gaudin et al., 2009), it is generally found at the end of a 

dorsal canal, hidden in ventral view, that travels dorsal to 

the carotid sulcus and opens in the anterior margin of the 

jugular foramen (Figure 8). Moreover, the tympanic processes of the basioccipital (likely homologous at least in 

part with the basioccipital tubera of Patriomanis) form the 

medial margin of the jugular foramina. The groove emerging anteriorly from the hypoglossal foramen, when present, connects with the jugular foramen rather than passing 

medial to this opening (this groove is absent in Phataginus but present in Manis sp., S. temminckii, and those 

S. gigantea in which the jugular and hypoglossal foramina are not fused; Gaudin, unpublished data). Because of 

the difference in morphology between Patriomanis and 

the extant species, Gaudin et al. (2009) coded the inferior petrosal sinus foramen as being absent in the former. 

Ultimately, the identity of this “inferior petrosal sinus” 

opening cannot be conclusively established—it could, for 

instance, represent a patent basicochlear commissure, as 

in the armadillo Dasypus novemcinctus (Wible, 2010), an 

interpretation not considered by either Emry (1970) or 

Gaudin et al. (2009). That said, we now believe Emry’s 

(1970) assignment of this opening to the inferior petrosal 

sinus is the most probable interpretation. We also accept 

his assignment of the remaining openings to the jugular 

and hypoglossal foramina, respectively.

Neither the type nor USNM-P 494439 preserves the 

posterior edge of the basioccipital, so the morphology 

of the ventral margin of the foramen magnum cannot be 

ascertained. The basisphenoid contacts the presphenoid 

anteriorly, the pterygoid and possibly the palatine laterally, and the basioccipital posteriorly. The basioccipital 

contacts the basisphenoid anteriorly and the exoccipital 

posterolaterally. Neither element appears to have a lateral connection with the petrosal, although both closely 

approach its medial margin (Figure 7).

Petrosal

Although both left and right petrosals are present in 

the type specimen, neither is well preserved. The left is 
2 0 • 

The lateral surface of the crista parotica slopes 

anterodorsally, where it abuts a shelf extending medially 

from the inner wall of the squamosal, just dorsal to the 

porus acousticus. The shelf is actually separated from the 

crista by a narrow gap anteriorly (which may be a preservational artifact), whereas posteriorly the two are sutured. 

Together they form the roof to a triangular depression, the 

epitympanic recess (as noted above; Figure 7). The epitympanic recess houses the articulation between the mallear head and the body of the incus in basal eutherians 

(e.g., Lepticitis, Novacek, 1986; Zalambdalestes, Wible 

etal., 2004; Solenodon, Wible, 2008; Maelestes, Wible etal., 

2009; see also extant Canis, Evans and Christensen, 1979; 

Solenodon, Wible, 2008). In modern pangolins, this area is 

not fully roofed over, opening rather into a sinus between 

the squamosal and petrosal, the epitympanic sinus, which 

is continuous with the inflation in the posterolateral corner of the squamosal (Figure 8; Jollie, 1968; Emry, 1970; 

Gaudin et al., 2009). An epitympanic sinus has also been 

reported for E. waldi (Rose et al., 2005; Gaudin et al., 

2009) and Palaeanodon (Patterson et al., 1992). The latter authors suggested the presence of an epitympanic sinus 

might be considered a morphological synapomorphy of 

a cohort Edentata including xenarthrans, pangolins, and 

palaeanodonts, although its morphology is not uniform 

(e.g., it expands anterior to the porus acousticus in xenarthrans, posteriorly in pangolins, and in both directions in 

Palaeanodon), and it is absent in Patriomanis, in dasypodine and tolypeutine armadillos, and in mylodontid sloths 

(Patterson et al., 1989, 1992; Gaudin, 1995, 2004). The 

participation of the squamosal bone in the roof of the epitympanic recess or sinus is considered a unique synapomorphy of Pholidotamorpha (Pholidota + Palaeanodonta) 

by Gaudin et al. (2009).

Along the posterior wall of the epitympanic recess 

in Patriomanis lies a small circular pit that straddles the 

petrosal–squamosal suture and faces anteriorly. This 

depression represents the fossa incudis, which in modern manids is more medially located and faces laterally 

(Gaudin et al., 2009).

As in all mammals, there are two large foramina in the 

posterolateral and posteroventral walls of the promontorium. The more lateral of the two is the fenestra vestibuli, 

which in life accommodates the footplate of the stapes. As 

preserved on the right side of USNM-P 494439 (the only 

place where it is fully prepared), it is oval in shape, with 

a stapedial ratio of 1.63 (length/width; see Segall, 1970). 

In P. tricuspis (UTCM 1695) the ratio is somewhat lower, 

1.50. It is deeply recessed and positioned midway between 

but slightly ventral to the primary facial foramen and the 

and M. pentadactyla), although in most the crista parotica 

is reduced (Smutsia; Figure 8) or the facial sulcus has been 

converted to a closed canal (M. crassicaudata, M. javanica, 

some specimens of M. pentadactyla, and P. tetradactyla; 

Gaudin and Wible, 1999; Gaudin et al., 2009). The posterior portion of the facial sulcus extends ventrally toward 

a well-defined stylomastoid notch that lies posterolateral 

to the small, cylindrical tympanohyal. The notch itself 

forms two-thirds of a circle, open medially, with its lateral 

and posterior walls formed by the paroccipital process of 

the petrosal. In extant Manidae, the tympanohyal bends 

medially at its distal extremity to fuse with the lateral surface of the petrosal, enclosing the exit of the facial nerve 

(CN VII) in a complete stylomastoid foramen (Figure 8; 

Gaudin and Wible, 1999; Gaudin et al., 2009). 

Near the anterior terminus of the facial sulcus is an 

opening, the primary facial foramen, in the lateral wall of 

the petrosal’s pars cochlearis (Figure 7). The space immediately lateral to this foramen, the cavum supracochleare, 

houses the geniculate ganglion of the facial nerve (Evans 

and Christensen, 1979; Wible, 2010). This space is open 

ventrally, although shielded somewhat in ventral view by 

the crista parotica itself, which bends medially toward its 

termination on the lateral wall of the pars cochlearis just 

anterior to the cavum supracochleare, and by the dorsal 

rim of the tensor tympani fossa. This dorsal rim stands 

out as a distinct ridge from the lateral surface of the pars 

cochlearis. In other eutherian mammals, the cavum supracochleare is completely floored ventrally by the petrosal, 

creating a secondary facial foramen posterior to the cavum 

and an anterior hiatus Fallopii that transmits the greater 

petrosal nerve (e.g., the extinct Late Cretaceous Zalambdalestes, Wible et al., 2004; Solenodon, Wible, 2008; some 

Dasypus, Wible, 2010). This is also the morphology in 

P. tricuspis (UTCM 1695) and S. gigantea (AMNH 53858), 

although the morphology varies in extant pangolins; for 

example, as noted above, many extant forms enclose the 

facial sulcus in a canal. In Patriomanis, there is a small 

ventral groove where the crista parotica meets the pars 

cochlearis that likely accommodated the greater petrosal 

nerve.

The posterior half of the facial sulcus is nearly twice 

the transverse width of the anterior half, largely because of 

the presence of a sizable pit along its medial edge. This pit, 

which faces laterally, anteriorly, and ventrally, is the stapedius fossa for the stapedius muscle. In modern pangolins 

the stapedius fossa is shielded from view ventrally by the 

contact between the tympanohyal and the pars cochlearis 

(Figure 8), but in P. tricuspis (UTCM 1695) it is somewhat 

more anteriorly directed than it is in Patriomanis (Figure 7).
n u m b e r 9 8 • 2 1

2010), it likely housed the tympanic nerve, a branch of 

the glossopharyngeal nerve (CN IX). The closest analog 

we found to this process is on the right side of a specimen 

of M. pentadactyla (FMNH 75878), where the caudal 

tympanic process of the petrosal is lower and blunter and 

rounded distally and situated farther laterally but appears 

to be flanked by the same two grooves. In other specimens we examined from extant species, the caudal tympanic process appeared as a low knob or flange and was 

often not clearly delineated by neural grooves. With the 

exception of a few specimens of Phataginus (P. tetradactyla FMNH 62209 and P. tricuspis FMNH 42680 but not 

P. tricuspis UTCM 1695), these knobs or flanges tended 

to be positioned at the lateral edge of the jugular foramen, 

as in M. pentadactyla (FMNH 32511), rather than at its 

anterior margin, as in Patriomanis.

The grooves flanking the caudal tympanic process 

of the petrosal in Patriomanis are roofed by a horizontal plate that also extends between the base of the caudal tympanic process and the dorsal rim of the fenestra 

cochleae. This plate represents the roof of the posttympanic sinus (Wible, 2010) and is a feature missing in modern pangolins because of the lateral reorientation of the 

fenestra cochleae. The junction of this roof with the promontorium is marked by a very small, shallow “hairline” 

groove that extends from the medial side of the fenestra 

cochleae opening to the medial side of the pars cochlearis, 

where it connects to a small foramen of unknown function 

just anterior to the foramen for the cochlear canaliculus. 

Both the roof of the posttympanic sinus and the caudal 

tympanic process are part of the pars canalicularis of the 

petrosal.

Posterior and medial to the base of the caudal tympanic process is a broad vertical groove that extends dorsally into the braincase. In the anterior wall of this groove 

is a foramen, the opening for the cochlear canaliculus 

(= cochlear aqueduct; see Evans and Christensen, 1979) 

that transmits the perilymphatic duct of the inner ear. In 

other extant eutherians (e.g., Homo, Pick and Howden, 

1977; Canis, Evans and Christensen, 1979; Dasypus, 

Wible, 2010), the cochlear canaliculus opens into the lateral wall of the jugular foramen, so we identify this groove 

as the lateral wall to the jugular foramen as well, despite 

its lateral position relative to modern pangolins. As noted 

above, the medial margin of the jugular foramen is likely 

formed by the tympanic process of the basioccipital. 

Although several portions of the ventral exposure 

of the pars canalicularis have already been described, 

including the facial sulcus, cavum supracochleare, crista 

parotica, and the caudal tympanic process, the largest 

stapedius fossa. The ventral edge of the fenestra forms a 

strong ridge that is continuous anteriorly with the dorsal 

ridge of the tensor tympani fossa, and there is a lower, 

broader ridge orthogonal to both that separates the anterior fenestra vestibuli from the posterior tensor tympani 

fossa. In P. tricuspis (UTCM 1695) and other modern 

forms, the strong ventral rim of the fenestra vestibuli does 

not extend anterior to the opening, and the opening itself 

is only slightly recessed. 

The second large opening, situated at the back of the 

promontorium, is labeled here the fenestra cochleae (Figure 

7; following Gaudin and Wible, 1999; Gaudin et al., 2009). 

It is actually, in all likelihood, the aperture for the cochlear 

fossula, which would contain an opening inside for the 

attachment of the round window, the true fenestra cochleae 

(see, e.g., Wible, 2010), as in the modern M. javanica (Jollie, 

1968). However, since we have been unable to prepare the 

interior of this opening and cannot assess its morphology, 

we will retain the older usage in our description and figures. 

The fenestra cochleae faces mainly posteriorly and laterally 

to a small degree. It is separated from the fenestra vestibuli 

by a broad, superficially convex crista interfenestralis. This 

morphology differs strongly from that of modern pangolins, in which the crista interfenestralis is a narrow strut, 

the two fenestrae are in close proximity, and the fenestra 

cochleae faces almost directly lateral (Figure 8; Gaudin and 

Wible, 1999; Gaudin et al., 2009). The arrangement in 

Patriomanis is clearly less derived, resembling the condition 

in E. waldi and the palaeanodont Metacheiromys (Gaudin 

et al., 2009). The crista interfenestralis in Patriomanis 

bears a small, transverse ridge that is continuous anteriorly 

with the ventral rim of the fenestra vestibuli. Its function 

is unknown.

Part of the difficulty in preparing the fenestra cochleae 

stems from the fact that it is shielded posteriorly by a 

large, bulbous process with a rather square outline, the 

caudal tympanic process of the petrosal. The caudal tympanic process is present on both the left and right sides of 

the skull in USNM-P 494439 but is not in evidence in the 

type specimen, although we suspect its absence in the latter is due to postmortem damage. The process appears to 

be freestanding because it is bounded anteromedially and 

laterally by grooves. The latter is larger and deeper and 

undercuts the caudal tympanic process dorsally. On the 

basis of comparisons with other eutherians (Canis, Evans 

and Christensen, 1979; Dasypus, Wible 2010), this groove 

likely housed the auricular branch of the vagus nerve (CN 

X). The anteromedial groove does not undercut the caudal 

tympanic process. Again on the basis of comparisons with 

other eutherians (Evans and Christensen, 1979; Wible, 
2 2 • 

enigmatic edentate Eurotamandua (Gaudin et al., 2009). 

Gaudin and Wible (1999) demonstrate that the entotympanic element present in Smutsia (Figure 8) and some 

specimens of P. tricuspis (as well as palaeanodonts; Gaudin 

et al., 2009) is almost certainly absent in Patriomanis.

Occipital

The occipital bone, including both exoccipital and 

supraoccipital regions, has been thoroughly described by 

Emry (1970), so we will not repeat that description here 

but instead make some broader taxonomic comparisons. 

Emry (1970) notes that a strong lambdoid or nuchal crest 

is present in Patriomanis but lacking in modern pangolins. The plesiomorphic condition is displayed not only by 

Patriomanis but also by E. waldi, Eurotamandua, and the 

metacheiromyid palaeanodonts (Gaudin et al., 2009). The 

latter authors also note that in some pangolins, the exoccipital bone is laterally constricted immediately anterior to 

the occipital condyle, forming a discrete condylar “neck.” 

This feature is present in E. waldi and the extant giant pangolin (S. gigantea), as well as the species in the genus Manis 

(Figure 5b), but is not observed in Patriomanis, the extant 

genus and species Phataginus and S. temminckii, and metacheiromyid palaeanodonts (Gaudin et al., 2009). Last, the 

transversely oval foramen magnum of Patriomanis (Emry, 

1970) is known to have quite a varying distribution among 

pholidotamorphs, with Metacheiromys, S. gigantea, and 

some S. temminckii and M. javanica sharing the condition 

in Patriomanis, whereas a more circular foramen magnum 

is present in Palaeanodon, Phataginus, M. crassicaudata, 

and M. pentadactyla (Gaudin et al., 2009).

Endocranium

The matrix was removed from the interior of the type 

skull of Patriomanis, and an endocast was made. This 

endocast is described briefly in Emry (1970). The removal 

of this matrix has allowed us limited access to assess the 

bony anatomy of the walls of the endocranial cavity. Access 

is limited in the sense that we can address only what is visible through the foramen magnum or the anterior opening where the skull is broken just behind the presumptive 

location of the cribriform plate. Gaudin and Wible (1999) 

and Gaudin et al. (2009) have already noted several phylogenetically relevant features. Patriomanis is plesiomorphic in its retention of venous grooves on the endocranial 

surface (including those for the transverse, sigmoid, and 

superior petrosal sinuses), these grooves being lost in modern Manidae. In addition, the flooring of the middle cranial 

portion has not. It is manifested as a bulky, trapezoidal 

paroccipital process (following Wible and Gaudin, 2004; 

sometimes called the “mastoid process”; see Wible and 

Gaudin, 2004, for discussion). The paroccipital process 

has a nearly straight, elongate medial edge that is notched 

anteriorly. This notch, along with the tympanohyal, outlines the stylomastoid notch. The anterior and posterior 

edges of the paroccipital process approach one another 

laterally, lending the process its trapezoidal shape. The lateral edge is formed into two processes that are exposed in 

lateral view: an anterior process that is framed by the arch 

of the posttympanic crest of the squamosal and a posterior process that lies in the arch between the posttympanic 

process of the squamosal and the lower end of the nuchal 

crest of the exoccipital bone. The ventral surface of the 

paroccipital process is complex. It posteromedial portion 

is triangular, narrowing laterally, and its surface is anteroposteriorly convex. It ends at a transverse crest that marks 

its ventralmost extent. This crest is separated by a groove 

from a second similar but more anterior crest. The groove 

is broad medially, where it backs against the stylomastoid 

notch, then narrows laterally as it approaches the anterior 

margin of the posttympanic process. As noted above, this 

groove may be involved in transmitting the facial nerve. In 

front of the anterior crest, but raised dorsally, is the rather 

shapeless anterior lateral prong of the paroccipital process 

that is exposed beneath the posttympanic crest. In almost 

all modern Manidae, the paroccipital region of the petrosal is covered by the squamosal in lateral view (Gaudin 

et al., 2009) because of its expansion to accommodate 

internal sinuses (see above). However, there is at least one 

specimen of S. gigantea (AMNH 53858; Figure 8, labeled 

“pe”) in which a small “mastoid exposure” of the paroccipital process is present. This exposure is triangular and 

extends farther dorsally than in Patriomanis and is simpler 

in its morphology. It abuts the anterior edge of the nuchal 

crest, as in Patriomanis, but lacks a posttympanic process 

or posttympanic crest of the squamosal, and the paroccipital process lies behind the enclosed stylomastoid foramen 

and does not participate in transmitting the facial nerve.

Ectotympanic/Entotympanic/Ear Ossicles

None of these elements is preserved in Patriomanis. 

Emry (1970) speculates that the ectotympanic element 

resembled the simple crescent-shaped form in the extant 

genus Manis, a form also present in E. waldi and the 

Paleocene palaeanodont Palaeanodon, as opposed to the 

inflated, bullate form evident in modern African pangolins, 

as well as the Eocene palaeanodont Metacheiromys and the 
n u m b e r 9 8 • 2 3

symphysis, the distal tip of the anterolateral mandibular 

“prong,” and most of its posterior reaches. There are also 

two small fragments from the left and right dentaries of 

the type specimen. The largest of these preserves the ventral margin of the bone from the posteriormost part of the 

symphysis to the beginning of the ascending ramus. Emry 

(2004) has already drawn attention to its characteristic 

pangolin apomorphies, including its edentulous nature, its 

spout-like symphysis with an unfused, sutural attachment 

between the left and right dentaries, and the presence of 

the distinctive anterolateral bony prong in the symphyseal 

area, a feature that, according to Doran and Allbrook 

(1973:888), “gives attachment to the lower lip” in modern 

pangolins. Emry (2004) has also noted plesiomorphic features, like the retention of a weak coronoid process, lacking in the modern taxa. Emry (2004) also draws attention 

to the greater overall robusticity of the horizontal ramus 

relative to modern pangolins. The ramus is very thin transversely but deeper dorsoventrally than in extant manids. 

However, it should be noted that all known pholidotans, 

including the modern forms, Necromanis, Patriomanis, 

E. waldi, and Eurotamandua, have a maximum depth of 

the horizontal ramus less than 10% of overall maximum 

mandibular length. 

One feature not described by Emry (2004) but illustrated here (Figure 9b) is a small bone fragment that we 

believe pertains to the right dentary of USNM-P 494439. 

It is not attached to the dentary, and indeed, we cannot 

fit it precisely to the existing broken posterior end of the 

bone. However, what we take to be its broken dorsal edge 

roughly matches the shape of the broken posteroventral 

edge of the right dentary. That would make this fragment 

a preserved angular process of the dentary. Note that it 

is robust and hooked posteriorly, with a convex leading 

edge, concave trailing edge, and rounded tip. The angular 

process is missing in Necromanis and modern manids but 

is retained in E. waldi and Eurotamandua. In E. waldi

it forms either a short triangular process reminiscent of 

the coronoid in Patriomanis (see Storch, 2003: fig. 1, SMF 

MEA 263) or a blunt semicircular and slightly inflated 

process (type specimen, SMF Me 84, Storch, 1978), but 

in neither case is it as strong or posteriorly hooked as 

that of Patriomanis. The angular process is hooked in 

Eurotamandua (see especially Storch and Habersetzer, 

1991: fig. 3a,b), but it is pointed at the tip instead of 

rounded, giving it a less robust appearance.

A few features of the symphyseal region not described 

by Emry (2004) are distinctive in Patriomanis. As in living pangolins, there are two mental foramina on the anterior dentary. The more posterior opens laterally on the 

fossa by the sphenoid complex is plesiomorphic, in contrast to the derived squamosal floor found in manids. One 

of the cranial fragments from USNM-P 299960 preserves 

a clear, ovate hypophyseal fossa on the dorsal surface of 

the basisphenoid, flanked laterally by grooves for the internal carotid artery, those grooves in turn lying medial to a 

depression that presumably represents the cavum epiptericum. This same fragment also has a clear groove running 

along the side of the basioccipital. This groove presumably 

accommodated the inferior petrosal sinus.

Patriomanis represents an intermediate condition in 

the development of a bony tentorium, that is, an osseous 

lamina formed within the sheet of connective tissue separating the cerebrum from the cerebellum. In most mammals (e.g., Homo, Pick and Howden, 1977; Erinaceus, 

Gaudin et al., 2009), this partition is composed entirely of 

membranous connective tissue. In Patriomanis, a partial 

bony tentorium is present in the form of an outgrowth 

from the inner surfaces of the squamosal and parietal 

bones, running posterodorsal to anteroventral along the 

anterior edge of the petrosal. In modern forms, the tentorium is a broad bony septum that extends all the way 

from the anteroventral edge of the petrosal to the roof of 

the cranial cavity, attaching again to the inner walls of the 

squamosal and parietal (anterior to their contact with the 

occipital). Curiously, an extensive bony tentorium like 

that of modern pangolins is also found in the most basal 

living carnivorans, Nandinia (Gaudin et al., 2009), and 

other extant Carnivora (Evans and Christensen, 1979; 

Wyss and Flynn, 1993; Homberger and Walker, 2004),

with the Carnivora likely representing the sister taxon to 

Pholidotamorpha (Meredith et al., 2011; O’Leary et al., 

2013). In both Nandinia and Patriomanis, the groove for 

the superior petrosal sinus passes between the petrosal 

and the tentorium, whereas in modern pangolins, there is 

a foramen piercing the tentorium that accommodates this 

vessel (Gaudin et al., 2009). 

Mandible

The most completely preserved mandibular remains 

known for Patriomanis are those from USNM-P 494439 

described in detail by Emry (2004). These remains include 

a nearly complete left dentary (Figure 9a,c) figured by 

Emry (2004: fig. 10.5A,B,D) in medial, lateral, and dorsal 

views, lacking only the posteriormost portions, including 

the condyle and angle. It also includes a much less complete right dentary, illustrated by Emry only in dorsal view 

(Emry, 2004: fig. 10.5D) but here in both medial and dorsal views (Figure 9b,c). It is missing the anterior part of the 
2 4 • 

(Smutsia), and it has a convex profile in E. waldi and a 

concave profile in M. javanica, M. pentadactyla, and some 

specimens of S. gigantea.

Axial Skeleton: Vertebral Colu

Figures 10–21; Ta

Atlas

Although not described by Emry (1970), a nearly 

complete atlas is available from the type specimen of 

Patriomanis (F:AM 78999; Figure 10a–c). The specimen 

is nearly complete ventrally but is missing the transverse 

processes laterally and the left anterior articular surface. It 

is also missing portions of the lamina, although the right 

side of the lamina is nearly complete. A second atlas is 

available from USNM-P 494439. It has not been prepared 

completely free of the matrix, but it preserves a complete 

transverse process and anterior articular facet on the left 

side (Figure 11).

In cranial view, the atlas is wider transversely than it 

is tall dorsoventrally (Table A2). This is the case in extant 

pangolins as well, although the discrepancy is somewhat 

less in the African forms. The dorsally bowed lamina is 

capped by a short, conical neural spine, whereas the ventral arch is nearly horizontal. As in other therian mammals 

(see, e.g., the early metatherian Mayulestes, Muizon, 1998; 

the early eutherian Maelestes, Wible et al., 2009; or the 

extant Canis, Evans and Christensen, 1979), the anterior 

articular facet for the occipital condyle is strongly concave 

dorsoventrally and faces strongly mediad and slightly craniad. The facet is broader dorsally than ventrally, taking 

on an inverted teardrop shape (Figure 10a). The left facet 

is widely separated ventrally from its counterpart on the 

right, the separation (~13 mm) equaling nearly 60% of 

the maximum width of the atlas (excluding the transverse 

process; Table A2). In Smutsia, M. javanica, and M. crassicaudata, this separation is much narrower, comprising less 

than 40% of the maximum atlas width (Gaudin et al., 

2009). On its caudal surface, the atlas of Patriomanis 

bears a pair of posteromedially facing articular facets for 

the axis, situated at the intersection of the dorsal and ventral arches and the transverse process (Figure 10b). The 

surface contour of the facets is nearly flat, although in ventral view one can detect a slight transverse concavity. In 

outline, the facets take on the shape of a rounded equilateral triangle. The dorsolateral edge of this triangle forms 

a raised rim because of the presence of a short groove 

immediately anterior to this edge. The groove terminates 

horizontal ramus a short distance posterior to the anterolateral bony prong. The second, more anterior foramen 

typically perforates the base of the prong in extant forms, 

whereas in Patriomanis it is located well anterior and ventral to the prong, on the ventral surface of the symphysis. 

The morphology of the prong is itself unusual. In Patriomanis it arises from the dorsal edge of the symphysis, 

whereas in the extant manids it is separated by a groove 

from the dorsal rim and attaches much farther ventrally 

on the lateral sides of the symphysis. Finally, the symphysis itself is elongate with a straight ventral profile in lateral 

view in Patriomanis (Figure 9a). The symphysis is much 

shorter in E. waldi, Eurotamandua, and some individual 

specimens of the two species of African ground pangolins 

FIGURE 9. Mandible of Patriomanis americana, USNM-P 49443

(a) left mandible in lateral view; (b) right mandible in medial view; 

(c) left and right mandibles in occlusal view. Abbreviations: ap, 

angular process; crp, coronoid process; mand, mandible; mf, mental foramen; mnp, mandibular prong; ms, mandibular symphysis. 

Scale bar = 1 cm. Illustrations by Julia Morgan Scott for Smithsonian 

Institution.
n u m b e r 9 8 • 2 5

it a deepened appearance in lateral view. In contrast, the 

transverse process of the atlas tends to be flattened dorsoventrally and expanded anteroposteriorly in Eomanis and 

Smutsia (Gaudin et al., 2009). The anterior opening of the 

transverse canal perforates the base of the transverse process, emptying into an open groove that travels anteriorly 

and dorsally onto the surface of the lamina, eventually 

reaching the lateral vertebral foramen. Although the latter 

seems to form only a partially closed foramen in the type 

(Figure 10), it is completely enclosed in bone in USNM-P 

494439. In Manis, the groove connecting the transverse 

and lateral vertebral foramina is partially closed over by 

at the posterior opening to the transverse canal. The latter 

opening is thus situated at or near the lateral edge of the 

posterior articular facet, as in the extant Phataginus and 

Smutsia, whereas in Manis this opening is situated more 

medially, in a position dorsal to the midpoint of the facet.

In lateral view (Figure 10c), the height of the atlas 

is slightly greater than its anteroposterior length, as it is 

in most modern pangolin species except the two in the 

genus Smutsia, in which the atlas is anteroposteriorly 

elongated (Table A2). There is a prominent transverse 

process extending posterolaterally and slightly ventrad 

from the lateral surface of the bone in USNM-P 494439 

(Figure 11). It extends at roughly a 30° angle from the 

lateral surface of the atlas in dorsal view and is flattened 

in a vertical plane, orthogonal to its long axis. From the 

side, it appears quite deep dorsoventrally, flaring toward 

its distal end before terminating in two rounded distal 

points. The process appears similarly deep in Manis (and 

variably so in Phataginus; see Gaudin et al., 2009), with 

M. javanica showing a distal bifurcation reminiscent of 

the condition in Patriomanis. The transverse process is 

flattened either anteroposteriorly or in a plane oblique 

to the anteroposterior axis in these modern taxa, giving 

FIGURE 10. Atlas of Patriomanis americana: (a) atlas of F:

78999 (holotype) in anterior view; (b) in left lateral view; 

(c) in posterior view; (d) reconstruction of the atlas in left lateral 

view. Abbreviations: aas, anterior articular surface; lvf, lateral 

vertebral foramen; ns, neural spine; pas, posterior articular surface; tf, transverse foramen; tp, transverse process. Scale bar = 

1 cm. Illustrations by Julia Morgan Scott for Smithsonian 

Institution.

FIGURE 11. Atlas and axis of Patriomanis americana, USNM

494439: (a) axis in posterolateral view, axis in ventral view; (b) atlas 

in dorsal view, axis in dorsolateral view. Abbreviations: aas, anterior 

articular surface; atl, atlas; ax, axis; c, centrum; lvf, lateral vertebral 

foramen; ns, neural spine; pas, posterior articular surface; pz, posterior zygapophysis; tf, transverse foramen; tp, transverse process. 

Stippled areas represent matrix; the cross-hatched areas show broken surfaces. Scale bar = 1 cm. Illustrations by Julia Morgan Scott 

for Smithsonian Institution.
2 6 • 

is completely separate from the anterior articular facets 

(Gaudin et al., 2009). In Patriomanis, the anterior articular 

facets are ovate, somewhat broader than they are tall, and 

have a slightly convex surface contour in both the transverse and dorsoventral planes (Figure 12a). The outline of 

these facets varies substantially among other pangolins. 

They are broader than tall in Cryptomanis, Necromanis, and Manis, whereas the maximum transverse width 

and maximum dorsoventral height are nearly equal in the 

extant African forms (Gaudin et al., 2009). The surface 

contour is transversely convex not only in Patriomanis but 

also in Cryptomanis and Smutsia. It is concave transversely 

in Phataginus, M. javanica, and M. pentadactyla, whereas 

in Necromanis and M. crassicaudata, it is convex medially 

and convex laterally; that is, it is largely convex but with 

a small concave depression immediately ventral and lateral 

to the dens (Gaudin et al., 2009).

The centrum is low and broad, as in other pangolins. 

In ventral view (Figure 12b), it is almost square, its transverse width nearly equal to its anteroposterior length. This 

shape resembles the condition in Cryptomanis, Necromanis, and M. javanica but stands in contrast to the condition in other extant manids, in which the centrum is wider 

than it is long, even when the dens is included in the length 

measurement (Gaudin et al., 2009; Table A2). The ventral 

surface of the centrum is marked by a prominent midline 

crest flanked by deep lateral excavations. The midline 

crest broadens posteriorly to form a raised triangular area. 

This kind of surface relief is missing in other pangolins 

with the exception of Cryptomanis and some M. javanica 

(Gaudin et al., 2009), in which the excavations tend to be 

weaker and more laterally situated and the midline crest 

is broader.

a bridge of bone connecting the anteromedial edge of 

the transverse process to the lateral surface of the atlas 

(Gaudin et al., 2009). Such a bridge is also variably present in Smutsia and Eomanis, but Phataginus resembles the 

condition in Patriomanis (Gaudin et al., 2009). 

In dorsal view, the lamina of the atlas appears to be 

relatively broad, with parallel anterior and posterior edges 

(Figure 11b). However, the middle portion of the posterior 

edge is missing, leaving open the possibility that there is 

some sort of notch or concavity in this area. Patriomanis lacks the prominent depressions anterolateral to the 

neural spine that are found in extant Manis and S. gigantea (Gaudin et al., 2009). These depressions likely house 

the origins of the m. rectus capitis dorsalis (see Homo, 

Pick and Howden, 1977; Canis, Evans and Christensen, 

1979). Patriomanis also lacks any indication of an alar 

notch between the anterior edge of the transverse process 

and the lamina. Such a notch is evident in Phataginus, M. 

crassicaudata, and S. temminckii and is variably present in 

S. gigantea. The ventral arch, seen in ventral view, is narrower anteroposteriorly than the dorsal arch. It has a gently concave anterior margin, whereas the posterior margin 

is marked by a weak median spine flanked by lateral concavities. The ventral surface is otherwise unmarked.

Axis

Portions of the axis are preserved in all three of the 

most important specimens of Patriomanis, the type, F:AM 

78999; USNM-P 494439; and USNM-P 299960. The 

centrum is preserved in both the type and in USNM-P 

299960, but only USNM-P 494439 preserves any part of 

the neural arch. The partially prepared axis in this specimen preserves the posterior zygapophysis, the base of the 

transverse process, and portions of the neural spine. The 

axis of USNM-P 299960 is illustrated in Figure 12, and 

that of USNM-P 494439 is shown in Figure 11; these will 

largely serve as the basis for the following description.

The dens is reminiscent of that found in primitive therians (e.g., Mayulestes, Muizon, 1998; Maelestes, Wible 

et al., 2009). It is short and broad and thumb shaped in 

dorsal or ventral view, although slightly broader at its base 

than at its tip (Figure 12b). It bears a roughly triangular 

articular surface on its ventral surface that contacts a narrow corresponding surface on the floor of the atlantal vertebral canal. This facet abuts the anterior articular facets 

for the atlas along its posterolateralmost margins, resembling the condition in Cryptomanis, Necromanis, and 

most extant Manis (variably present in M. pentadactyla), 

whereas in the African pangolins the facet on the dens 

FIGURE 12. Axis of Patriomanis americana: (a) axis of USNM

299960 in anterior view (dorsal portion reconstructed on the basis 

of USNM-P 494439); (b) in ventral view; (c) reconstruction of the 

axis in ventral view. Abbreviations: aas, anterior articular surface;

c, centrum; de, dens; pz, posterior zygapophysis; tp, transverse process. Cross-hatched areas represent broken surfaces. Scale bar = 1 

cm. Illustrations by Julia Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 2 7

process is broken on both sides but is more complete on the 

left, missing roughly the distal half of the process. On both 

sides, the base of the transverse process is clearly perforated by a transverse foramen for the vertebral artery. The 

lack of a transverse foramen in the type specimen might be 

used to confirm its identity as a seventh cervical, although 

it should be noted that in all modern manids, the transverse 

foramen is present in C7 (Gaudin et al., 2007). The transverse process attaches to the side of the centrum and the 

base of the pedicle, in contrast to that of the type, in which 

it is attached more dorsally, at the junction of the pedicle 

and lamina. Furthermore, the shape and orientation of the 

transverse process are quite different in the two specimens. 

In the type it is directed dorsolaterally (in anterior view) 

As noted above, the only specimen that preserves any 

portion of the neural arch is USNM-P 494439 (Figure 11). 

In this specimen the posterior zygapophysis is preserved 

in its entirety. Its surface faces ventrolaterally, and it is 

characterized by a concave surface contour like that of P. 

tricuspis. In contrast, the surface contour is convex in all 

other extant manids. Only the base of the transverse process and the base of the neural spine are present in USNMP 494439, but there is enough of each process in place to 

demonstrate that the former extends posterolaterally from 

the neural arch and that the latter extends anteriorly to 

cover portions of the neural arch of the preceding atlas, as 

in most living pangolins (Gaudin et al., 2009).

Cervical Vertebrae (Exclusive of Atlas and Axis)

The remaining cervical vertebrae are poorly represented in the available material of Patriomanis. Only the 

type (F:AM 78999) and USNM-P 299960 preserve any of 

the cervicals other than the axis and atlas, and the former 

includes only fragments of these vertebrae, most notably a 

sizable fragment of C6 or C7 described and illustrated in 

Emry (1970). The latter specimen preserves what appears 

to be a nearly complete C4 or C5, along with two additional isolated cervical centra that likely represent C6 and 

C7 (Figures 1, 13). No complete cervical series is known 

for any fossil pangolin (Storch, 1978; Koenigswald and 

Martin, 1990; Storch and Martin, 1994; Gaudin et al., 

2006). Although Storch (1978) states that the cervical 

count is “probably” seven in Eomanis waldi, it is difficult to tell from his illustrations of the type material, and 

indeed, in his reconstruction (Storch, 1978: plate 5, fig. 1) 

he illustrates only five or six cervicals. Nevertheless, given 

the near-universal presence of seven cervicals among modern mammals, including all eight species of modern pangolins, it is likely that Patriomanis (and the other fossil 

pangolin taxa) also possessed seven cervicals.

The cervical preserved in USNM-P 299960 (Figure 13) 

differs from that described by Emry (1970) in a number 

of respects. It is nearly complete bilaterally, unlike that 

from the type. The centrum is even more depressed, with 

rounded or squared off lateral edges, in contrast to the 

type, in which the lateral edge of the centrum appears to 

form a kind of lateral keel (Emry, 1970: fig. 11). The cervical centra in Cryptomanis are much wider and flatter than 

those of Patriomanis, which more closely resemble the condition in Necromanis and extant manids. The centrum in 

USNM-P 299960 is roughly the same relative length as that 

from the type. The neural arch bears a tall, vertical neural spine, a feature missing from the type. The transverse 

FIGURE 13. Cervical vertebra (C4 or C5?) of Patriomanis americana, USNM-P 299960: (a) in anterior view; (b) in dorsal vie

(c) in right lateral view. Abbreviations: az, anterior zygapophysis;

c, centrum; ns, neural spine; ped, pedicle; pz, posterior zygapophysis; tf, transverse foramen; tp, transverse process. Stippled areas represent matrix; the cross-hatched areas show broken surfaces. Scale 

bar = 1 cm. Illustrations by Julia Morgan Scott for Smithsonian 

Institution.
2 8 • 

the anterior zygapophyseal facets are usually concave, and 

the posterior zygapophyseal facets are convex, with an 

orientation similar to that of USNM-P 299960. In Necromanis the zygapophyseal facets are flatter, as in Patriomanis (Gaudin et al., 2009).

Thoracic Vertebrae

Emry (1970) described only one fragment of a posterior thoracic vertebra (= “dorsal” vertebra) from the 

type specimen (F:AM 78999). However, as reported in a 

footnote from that work, additional thoracics were later 

recovered, including parts of an anterior thoracic with a 

partial centrum, three thoracics still in articulation with 

one another and their corresponding ribs, and three other 

posterior thoracics. USNM-P 494439 preserves more thoracics, including the ultimate and penultimate still in articulation with the lumbar vertebrae but not fully prepared 

free of the matrix, as well as four free thoracics and perhaps one or two isolated thoracics still mostly embedded 

in matrix. The most complete thoracic series derives from 

USNM-P 299960, which preserves at least 13 fragmentary 

to nearly complete thoracic vertebrae from the anterior, 

middle, and posterior parts of the series (Figure 1). We 

have illustrated a representative, fairly complete thoracic 

vertebra from each of these regions in Figures 14 (anterior view), 15 (dorsal view), and 16 (lateral view). In our 

reconstruction (Figure 1), we estimate that at least 15 thoracic vertebrae were present in Patriomanis, which accords

FIGURE 14. Thoracic vertebrae of Patriomanis americana, USNM-P 299960, in anterior view: (a) anterior thoracic (T4 or T5

vertebra; (b) middle thoracic vertebra; (c) postdiaphragmatic thoracic vertebra. Abbreviations: az, anterior zygapophysis; c, centrum; 

crf, capitular rib facet; mp, metapophysis; ns, neural spine; ped, pedicle; tp, transverse process; trf, tubercular rib facet. Stippled areas 

represent matrix; the cross-hatched areas show broken surfaces. Scale bar = 1 cm. Illustrations by Julia Morgan Scott for Smithsonian 

Institution.

and is cylindrical in cross section. In USNM-P 299960, it 

is oriented ventrolaterally (in anterior view); it is dorsoventrally compressed and anteroposteriorly expanded. In 

dorsal view it has anterior and posterior extensions proximally, and the distal end is likely oriented posterolaterally 

(Figure 13). Both types of extensions are also present in 

most cervicals in living manids.

In extant pangolins and in Cryptomanis (but not in 

Necromanis; Gaudin et al., 2009) the processes that bear 

the anterior and posterior zygapophyseal articulations 

extend far anterior and posterior to the centra of their 

corresponding vertebra (Gaudin et al., 2009). The anterior processes, for example, typically extend to roughly 

one-third the length of the preceding centrum. The processes in Patriomanis are much shorter, extending only 

slightly beyond the margins of the centra in lateral view 

(Figure 13c). The anterior zygapophyseal facet in USNMP 299960 is preserved only on the left side of the vertebra. 

It is egg shaped in outline, narrowing medially (Figure 

13b), and faces dorsomedially (Figure 13a). The surface 

contour is almost flat (slightly convex), with the posterior 

zygapophyseal facet lacking even this slight curvature and 

facing ventrolaterally. Emry (1970) describes a similar 

surface contour for the zygapophyseal facets in the more 

posterior cervical preserved in the type specimen (F:AM 

78999), but the orientation of the facets differs, with the 

posterior facet roughly horizontal and the anterior facet 

facing “outward and upward” (Emry, 1970:472). In 

extant pangolins, Cryptomanis, Eomanis, and Euromanis, 
n u m b e r 9 8 • 2 9

well with the condition in most modern pangolin species, 

which have between 14 and 16 thoracics, although some 

Phataginus tetradactyla have only 13 and Smutsia temminckii has only 11 (Gaudin et al., 2009). Eomanis waldi 

has 13 thoracics (Storch, 1978, asserts there are 12, but 

better preserved specimens recovered subsequently show 

the presence of an extra vertebra; Gaudin et al., 2009), 

but only 12 are preserved in Euromanis krebsi (Storch and 

Martin, 1994; Gaudin et al., 2009).

As is evident in the reconstruction (Figure 1), the neural spines of the thoracic vertebrae are fairly uniform in 

height, although in the more anterior thoracics the spines 

are slightly longer and less broad anteroposteriorly (Figure 

16). This condition also characterizes modern and extinct 

pangolins, with the exception of Eomanis and Euromanis 

(Gaudin et al., 2009). The distal tips of the spines take the 

form of a flat, anteroposteriorly elongate lenticular surface, 

especially in the more posterior vertebrae (Figures 15, 16). 

The neural spines are posterodorsally inclined in the anterior thoracics and nearly vertical in the posterior thoracics, 

but there is no real anticlinal vertebra, which is often present in other placental mammals (Slijper, 1946; for a classic example, see Canis in Evans and Christiansen, 1979). 

An anticlinal vertebra is present in the Asian pangolins 

Manis javanica and M. crassicaudata but is not in evidence 

in other extant forms or in Eomanis or Euromanis. The 

diaphragmatic vertebra is not preserved in Patriomanis, so 

FIGURE 16. Thoracic vertebrae of Patriomanis americana, USNM-P 299960, in lateral view: (a) anteri

thoracic (T4 or T5?) vertebra in left lateral view; (b) middle thoracic vertebra in right lateral view; (c) postdiaphragmatic thoracic vertebra in left lateral view. Abbreviations: az, anterior zygapophysis; c, centrum; crf,

capitular rib facet; crf 2, second posterior capitular rib facet; mp, metapophysis; ns, neural spine; ped, pedicle; 

pz, posterior zygapophysis; rf, rib facet; tp, transverse process; trf, tubercular rib facet. Stippled areas represent 

matrix; the cross-hatched areas show broken surfaces. Scale bar = 1 cm. Illustrations by Julia Morgan Scott 

for Smithsonian Institution.

FIGURE 15. Thoracic vertebrae of Patriomanis americana, USNMP 299960, in dorsal view: (a) anterior thoracic (T4 or T5?) vertebr

(b) middle thoracic vertebra; (c) postdiaphragmatic thoracic vertebra. Abbreviations: az, anterior zygapophysis; mp, metapophysis; 

ns, neural spine; pz, posterior zygapophysis; tp, transverse process; 

trf, tubercular rib facet. Stippled areas represent matrix; the crosshatched areas show broken surfaces. Scale bar = 1 cm. Illustrations 

by Julia Morgan Scott for Smithsonian Institution.
3 0 • 

illustrated (Figures 14b, 15b, and16b) has a smaller tubercular facet but is attached to a much larger, more distinct 

metapophysis. In the postdiaphragmatic thoracic vertebra 

illustrated (Figures 14c, 15c, and 16c), the diapophysis is 

low, extending laterally and somewhat posteriorly from the 

side of the pedicle rather than the junction of the pedicle 

and lamina. It has also lost its association with the metapophysis, which is instead attached to the zygapophysis. 

Indeed, apart from the small tubercle extending dorsolaterally from the top of the zygapophyseal facet, which pertains solely to the metapophysis, it is not clear how much 

of the large process forming the lateral wall to this facet is 

composed of metapophysis and how much is composed of 

zygapophysis (the same could be said for many, if not most, 

therian mammals). It should be noted that the aforementioned “small tubercle” is apparently the broken remnant 

of a much larger, distally squared off metapophysis that 

extends a fair distance dorsally and laterally away from the 

zygapophyseal facet. The metapophyses are preserved in 

their entirety in the penultimate and ultimate thoracic vertebrae of USNM-P 494439. The disposition of these processes in Patriomanis is similar to that in other pangolins, 

as regards the separation of diapophysis and metapophysis 

in the posterior thoracic vertebrae. Although metapophyses 

are not well developed in the Messel pangolins or in extant 

Manis except for the posteriormost thoracics, they extend 

nearly to the front of the thoracic series in Cryptomanis 

and Smutsia and to the middle of the series in Phataginus. 

The thoracic centra have a reniform cross section, 

with a dorsal depression for the capacious vertebral canal. 

The centra in the anterior thoracics are more depressed 

dorsoventrally and shorter anteroposteriorly and become 

progressively longer and deeper in more posterior thoracic vertebrae (Figures 14, 16), although the transverse 

dimension is the greatest in all (Table A3). As is evident in 

Table A3, the dimensions of the thoracic centra are very 

similar in Patriomanis and other extinct and extant pangolins, with the exception of the African tree pangolins 

(Phataginus), in which the centra are somewhat longer 

anteroposteriorly. The typical thoracic centrum bears part 

of two capitular facets for the heads of corresponding ribs, 

an anterior facet on the anterior dorsolateral margin of 

the centrum and a second on the posterior dorsolateral 

edge. However, in the more posterior centra, the posterior facet is lost, and the rib articulates entirely on one 

centrum rather than between two adjacent centra. It is 

unclear where this transition takes place, although in both 

USNM-P 299960 and 494439 there are at least two vertebrae with a single capitular facet. This is also the case in 

Cryptomanis, whereas in modern pangolins there may be 

its position relative to that in other pholidotans cannot be 

ascertained, although it is clear from USNM-P 494439 that 

there are at least two thoracics behind the diaphragmatic 

vertebra. In all other pholidotans, the diaphragmatic vertebra is in the posterior part of the thoracic series, with 

two to three postdiaphragmatic thoracics in the extant 

taxa (Gaudin et al., 2006, 2009) and three in Cryptomanis 

(Gaudin et al., 2006), Eomanis, and Euromanis (Gaudin 

et al., 2009). It should be noted that the specific vertebra 

involved may vary among extant forms, even within species, and the diaphragmatic transition (i.e., the transition 

from horizontal, thoracic-type zygapophyses [“tangential” 

in terminology of O’Leary et al., 2013] to more upright, 

lumbar-type zygapophyses [“centric” in terminology of 

O’Leary et al., 2013]) may involve two vertebrae in some 

individuals (Gaudin et al., 2009). 

The pedicles of the thoracic vertebrae are low and relatively narrow anteroposteriorly in the anterior thoracic 

vertebrae, becoming anteroposteriorly broader in more 

posterior thoracics (Figure 16). When viewed dorsally, 

the thoracic laminae are roughly rectangular in shape, 

broad transversely but even longer in their anteroposterior 

dimensions (Figure 15). As noted above, the zygapophyseal facets have a tangential orientation in the anterior 

thoracics; that is, they are approximately horizontal and 

parallel to the surface of the laminae, although with a 

slight dorsomedial to ventrolateral slant. The facets are 

flat and ovate in outline, with the long axes of the anterior 

facets running posteromedial to anterolateral. Consonant 

with Emry’s (1970) observations on the type specimen, the 

zygapophyseal facets of the postdiaphragmatic thoracic 

vertebrae in Patriomanis begin to take on the “enrolled” 

or “embracing” morphology so typical of pangolins 

(Frechkop, 1949; Rose et al., 2005; Gaudin et al., 2006, 

2009), with strongly concave anterior facets contacting 

the cylindrical posterior facets ventrally and laterally and 

barely overlapping their dorsal surfaces (Figures 14–16). 

Euromanis and Eomanis are the only pangolins, living or 

extinct, to lack these enrolled zygapophyses (Rose et al., 

2005; Gaudin et al., 2009).

The anteriormost thoracic vertebra illustrated (probably T4 or T5; Figures 14a, 15a, and 16a) bears a robust 

diapophysis (= dorsally situated transverse process bearing 

an articular facet for the tubercle of the rib; see Homberger 

and Walker, 2004) extending laterally from the junction 

of the lamina and pedicle. It is marked distally by an oval, 

concave facet for the rib tubercle. A small ridge extends 

dorsal to the tubercular facet that represents a rudimentary 

metapophysis (= mammillary process of Homberger and 

Walker, 2004). The diapophysis of the midthoracic vertebra 
n u m b e r 9 8 • 3 1

vertebrae, which matches the count from the other patriomanid genus Cryptomanis (Gaudin et al., 2006), whereas 

the older taxa Euromanis and Eomanis have only six lumbars each (Storch and Martin, 1994; Gaudin et al., 2009), 

as do most modern manids, with the exception of Smutsia 

gigantea and some Phataginus tetradactyla, which have 

only five (Gaudin et al., 2009). 

The neural spines of the lumbar vertebrae are essentially upright, with their height roughly one and a half 

times their anteroposterior length (Figure 19; Table A3) 

and a flat, lenticular dorsal surface (Figure 18). The spines 

are roughly equivalent in height throughout the series. The 

lumbar neural spines exhibit similar dimensions in Smutsia gigantea and Eomanis. They are somewhat taller in 

S. temminckii and Cryptomanis and somewhat shorter in 

extant Manis, and in Phataginus the length of the neural spines actually exceeds their height (Table A3; Gaudin 

et al., 2009). The laminae and pedicles of the lumbar vertebrae are much like those described above for the posterior thoracics, the laminae broad and rectangular, the 

pedicles low and anteroposteriorly elongate. The posterior attachment of the pedicles is located well anterior to 

the posterior edge of the centrum and ventrolateral to the 

dorsal edge of the centrum. It is separated from the posterodorsolateral corner of the centrum by a deep groove 

for the spinal nerve (Figure 19b). This groove was first 

described in Cryptomanis (Gaudin et al., 2006) and is also 

present in Necromanis (Gaudin et al., 2009), but the condition is unknown in either Eomanis or Euromanis and 

is clearly absent in living pangolins (Gaudin et al., 2009). 

Like the posterior thoracic vertebrae, the lumbar vertebrae bear large metapophyseal–zygapophyseal processes 

one (Smutsia), two (Manis), or three (Phataginus) thoracic 

centra with a single capitular facet.

Lumbar Vertebrae

The type specimen (F:AM 78999) preserves but a 

single fragment of vertebral arch from the lumbar series 

(Emry, 1970). USNM-P 494439 preserves six lumbar vertebrae in series, including the first lumbar, and USNMP 299960 also preserves at least six, including the last 

two, both of which are well preserved and illustrated in 

Figures 17–19. We estimate Patriomanis had seven lumbar 

FIGURE 17. Penultimate lumbar vertebra (L6) of Patriomanis americana, USNM-P 299960, in anterior view. Abbreviations: az, anterior zygapophysis; c, centrum; mp, metapophysis; ns, neural spi

ped, pedicle; tp, transverse process. Scale bar = 1 cm. Illustration by 

Julia Morgan Scott for Smithsonian Institution.

FIGURE 18. Lumbar vertebrae of Patriomanis americana, USNM-P 299960, in dorsal view: (a) penultimate vertebra (L6); (b) last lumbar vertebra (L7). Abbreviations: az, anterior zygapophysi

mp, metapophysis; ns, neural spine; pz, posterior zygapophysis; tp, transverse process. Scale bar = 

1 cm. Illustrations by Julia Morgan Scott for Smithsonian Institution.
3 2 • 

the height of the neural spines. The metapophyses are much 

shorter in Eomanis and Euromanis and somewhat taller in 

modern pangolins, in which they are nearly two-thirds the 

height of the neural spines. The transverse processes in the 

lumbar vertebrae represent pleurapophyses, that is, fused 

lumbar ribs (Homberger and Walker, 2004). They extend 

laterally from the junction between the pedicle and centrum (Figures 17, 19). In dorsal view, the transverse processes are wing shaped, with a straight to slightly concave 

anterior border and a convex posterior margin, and they 

are directed laterally and somewhat anteriorly (Figure 

18). This morphology is generally similar to that found in 

Manis javanica, although in some individuals of this species the anterior lumbar transverse processes are oriented 

more directly lateral in dorsal view, as is typically the case 

in M. crassicaudata, both species of Phataginus, Smutsia 

temminckii, and some individuals of S. gigantea (Gaudin 

et al., 2009). In some S. gigantea, these anterior transverse 

processes are actually directed posterolaterally, a condition apparently also present in Cryptomanis, judging from 

the preserved bases of the lumbar transverse processes. 

Although the last lumbar transverse process of Patriomanis is slightly shorter than that of the preceding vertebra, 

this extinct genus lacks the dramatic reduction of the last 

lumbar transverse process in modern pangolins, in which 

it may be greatly shortened in length (some M. javanica 

and P. tricuspis), anteroposterior breadth (S. temminckii, 

P. tetradactyla, some S. gigantea, and P. tricuspis), or even 

reduced to a small, rodlike element (M. crassicaudata, M. 

pentadactyla, and some S. gigantea; Gaudin et al., 2009). 

The lumbar centra resemble those of the posterior 

thoracic vertebrae described above, although they are 

larger than the thoracic centra. They are wide and low, 

with a reniform cross section (Figure 17) indented dorsally 

by the transversely ovate vertebral canal. In Patriomanis, 

as in Necromanis, Cryptomanis, and some extant taxa 

(M. crassicaudata, M. pentadactyla, and S. gigantea), the 

anteroposterior length of the centra exceeds the dorsoventral depth by roughly a third (Table A3), whereas in other 

extant taxa (M. javanica, S. temminckii, and both species 

of Phataginus) the centra are more elongated, their length 

exceeding their depth by at least 1.5- fold. 

Note that the sacral vertebrae are described with the 

pelvis below.

Caudal Vertebrae

Emry (1970) describes three caudal centra from the 

type specimen (F:AM 78999). In actuality, the type specimen includes at least four additional caudal vertebrae, 

extending anterodorsally and slightly laterally from the 

anterior half of the laminae. These processes form the lateral margins to the strongly enrolled or embracing anterior 

zygapophyses, which contact the cylindrical posterior zygapophyses along their ventral, lateral, and dorsal surfaces. 

The illustrated lumbar vertebrae from USNM-P 299960 

(Figures 17–19) lack large free-standing metapophyses 

extending beyond the zygapophyseal facets, carrying only 

short, laterally directed tubercles, but these are evidently 

broken remnants of larger processes. The lumbar metapophyses of USNM-P 494439 are like those described 

above for the posterior thoracics, extending a considerable 

distance dorsally and laterally, ending in squared distal 

tips. These metapophyses reach a height roughly one-half 

FIGURE 19. Lumbar vertebrae of Patriomanis americana, USNM

299960, in left lateral view: (a) penultimate vertebra (L6); (b) last 

lumbar vertebra (L7). Abbreviations: c, centrum; mp, metapophysis; ns, neural spine; ped, pedicle; pz, posterior zygapophysis; spn 

g, groove for spinal nerve; tp, transverse process. Scale bar = 1 cm. 

Illustrations by Julia Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 3 3

themselves separated by deep U-shaped or V-shaped 

notches in dorsal view (Figure 20a). These enrolled zygapophyses are present in at least the first five caudals of 

USNM-P 299960. In modern manids, they are present in 

the anterior one-quarter to one-half of the caudal vertebrae (Gaudin, unpublished data), whereas in Cryptomanis they are known to be present in the first five caudals 

(Gaudin et al., 2006). The pedicles in the Patriomanis caudals are considerably lower than those in the lumbar vertebrae, and the vertebral canal is correspondingly smaller. 

The freestanding portions of the metapophyses are lower 

than in the lumbar vertebrae, leaving the neural spine 

much higher than the metapophyses, in contrast to the 

anterior caudals of modern manids, where the low neural spine is roughly the same height as the metapophyses. 

The centrum is shaped much as in the lumbar vertebrae, 

although as Emry (1970) noted, the centrum is roughly as 

wide as it is long (in fact, width slightly exceeds length in 

Ca1 from USNM-P 299960), and it bears blunt ventral 

articular processes for the chevron bones. These processes 

are paired, are situated at the anterior and posterior margins of the centra, and are connected to one another by a 

pair of longitudinal crests that cross the ventral surface of 

the centrum (Emry, 1970: fig. 13). 

The primary difference between the lumbar vertebrae and anterior caudals, however, is the presence in the 

latter of greatly elongated transverse processes. As illustrated in Rose and Emry (1993: fig. 7.4) and our skeletal 

reconstruction (Figure 1), the tips of the caudal transverse 

processes actually extend laterally to a point even with 

the dorsal edge of the ischium. These processes are even 

broader in Manis (e.g., M. javanica, USNM 198852) and 

Smutsia (e.g., S. temminckii, AMMH 168955), extending 

well lateral to the ischium, whereas in Phataginus (e.g., 

P. tricuspis, CM 16206), they are large but do not extend 

to the lateral edge of the ischium (Gaudin et al., 2009). 

The transverse processes of the anterior caudal vertebrae 

are even smaller in Eomanis (e.g., SMF MEA 263) and

Euromanis (Storch and Martin, 1994). The first caudal 

of Patriomanis has a thick, rectangular transverse process with a muscular fossa extending along its anterior 

edge for much of its length, ending laterally in a muscular 

boss (Figure 20a). It is oriented laterally and slightly posterior, with a straight or slightly convex distal edge that 

slants anterodorsolaterally. In succeeding vertebrae, the 

transverse processes become progressively narrower both 

mediolaterally and anteroposteriorly and have rounded 

(Ca2) or tapered tips (Ca3; Figure 1). 

The caudal vertebrae from the middle of the caudal 

series differ strongly from the more anterior elements. 

including one nearly complete anterior caudal and a partial caudal from the posterior part of the series. However, 

two other specimens have a much more complete caudal 

sequence. USNM-P 494439 preserves at least parts of 27 

caudal vertebrae, with nearly 20 in series from the middle 

toward the anterior part of the sequence. We estimate this 

specimen had at least 31 caudals. USNM-P 299960 preserves 28, with nine complete caudal vertebrae clustered 

mostly anteriorly and many of the remainder comprising 

only centra or even partial centra, particularly toward the 

posterior end of the tail. The first five caudal vertebrae 

are present in sequence, separated by a gap from a more 

posterior sequence of six, with another gap separating a 

more posterior sequence of seven, a still more posterior 

sequence of eight following another gap, a couple more 

posterior caudals partially preserved, and the last few 

caudals likely missing. The specimen also preserves four 

or five partial chevron bones. We estimate that USNMP 299960 had at least 34 caudals (despite the fact that 

only 31 are reconstructed in Figure 1, which also does not 

match exactly the sequence of gaps described above) and 

perhaps as many as 36. This many caudal vertebrae would 

mean that Patriomanis had a tail nearly as long as the 

extant African tree pangolins, which include the species 

with the largest number of caudals known for any extant 

mammal, Phataginus tetradactyla, with 47–49 caudal vertebrae (Flower, 1885; Gaudin, unpublished data), and a 

second species, P. tricuspis, with 35–39 caudals (Gaudin, 

unpublished data). In the other living pangolins, the number of caudals ranges from 24 to 29 (Flower, 1885; Gaudin 

et al., 2009; Gaudin, unpublished data; Flower, 1885, 

reports that some M. javanica have 30 caudal vertebrae). 

Gaudin et al. (2006) report 17 preserved caudals in the 

other patriomanid genus, Cryptomanis, but the series is 

much less complete in this specimen than in Patriomanis, 

and the actual number of caudals is hard to estimate. The 

tail is even less completely known in Necromanis (Koenigswald and Martin, 1990). The tail in Eomanis waldi is 

well preserved, with between 21 and 24 caudal vertebrae 

(Storch, 1978; Gaudin, unpublished data), but these are 

much shorter in the anteroposterior dimension than in the 

caudal vertebrae of other pholidotans, leaving this taxon 

with the shortest tail among known pangolins. 

The anterior caudal vertebrae are reminiscent of the 

lumbars in many respects. The neural spines are tall, vertical, and broad anteroposteriorly, but with a rounded rather 

than a flat terminus. After Ca4 the neural spines become 

notably shorter and more posteriorly inclined. The wide, 

rectangular laminae support embracing zygapophyses like 

those of the lumbar vertebrae, the zygapophyseal processes 
3 4 • 

USNM-P 299960 the twinned transverse processes begin 

in the 17th of 27 preserved vertebrae and are present 

in the next 7 preserved caudals (this section almost certainly incorporates a gap of at least 2 caudals), whereas in 

USNM-P 494439 the double processes begin with the 8th 

preserved caudal and include 14 more preserved caudals, 

including 11 consecutive vertebrae. In the more anterior 

examples, the transverse processes are anteroposteriorly 

broad, almost platelike, and diverge into separate processes only at the tips, but in more posterior vertebrae 

the transverse processes are deeply divided into two triangular, distally tapering and slightly recurved processes 

(Figure 20b). Double transverse processes have been noted 

in the two other patriomanid genera. In Cryptomanis the 

tips of the processes are broken, but a distinct gap can be 

seen between the bases of the doubled processes in at least 

one vertebra (Gaudin et al., 2006: fig. 5B). Double transverse processes have been described but not illustrated in 

one of the few preserved caudals of Necromanis (Storch 

The vertebral centra take on a cylindrical shape, as noted 

by Emry (1970), losing their reniform cross section and 

becoming more elongated, such that anteroposterior length 

is roughly one and a half to two times centrum transverse 

width, which is, in turn, roughly equal to centrum depth. 

The neural spines, although incomplete, are clearly much 

smaller than those from the more anterior caudals (Emry, 

1970), with their bases restricted to the posterior half of 

the laminae. Although strong zygapophyses persist and 

form synovial joints at least through the midpoint of the 

tail (e.g., they are present in 16 of 27 preserved vertebrae 

in USNM-P 494439), they no longer possess the enrolled 

morphology, forming nearly flat surfaces facing dorsomedially in the case of the anterior zygapophyses and ventrolaterally in the smaller posterior zygapophyses (Figure 

20b). Similar changes in centrum shape and zygapophyseal articulations are evident in modern pangolins.

The most striking feature of the mid-caudals in Patriomanis is the presence of double transverse processes. In 

FIGURE 20. Caudal vertebrae of Patriomanis americana: (a) first caudal of USNM-P 299960 in dorsal vie

(b) middle caudals of USNM-P 494439 in dorsal view, anterior toward right. Abbreviations: az, anterior zygapophysis; c, centrum; mp, metapophysis; ns, neural spine; pz, posterior zygapophysis; tp, transverse process. 

Stippled areas represent matrix; the cross-hatched areas show broken surfaces. Scale bars = 1 cm. Illustrations 

by Julia Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 3 5

and the zygapophyses are nonfunctional and reduced, the 

anterior being more robust and persisting farther posteriorly. They also retain ventral bosses or facets for the 

attachment of chevron bones nearly to the very end of the 

sequence. They lack, however, the robust transverse processes of the modern pangolins’ distal caudals. Instead, 

they take on an hourglass shape with very short, rounded 

transverse processes near the anterior and posterior margins of the centra (Figure 19a), much like the posterior 

and Martin, 1994). Storch (1978) describes and illustrates 

divided transverse processes on at least one caudal vertebra in Eomanis, but they are considerably shorter than 

those in Patriomanis and do not appear to be present in 

other Eomanis caudals, although several succeeding vertebrae do have strongly hourglass-shaped centra.

Double transverse processes very similar to those of 

Patriomanis are present in semiarboreal or arboreal vermilinguan anteaters, in which they have been associated 

with the presence of a prehensile tail (Hirschfeld, 1976; 

Gaudin and Branham, 1998). Such twinned transverse 

processes are not found, however, in other mammalian 

taxa with prehensile tails (Argot, 2003; Youlatos, 2003; 

Organ, 2010), nor are they present in modern pangolins 

with prehensile tails, that is, in the African tree pangolins 

in the genus Phataginus (Gaudin et al., 2006), or in the 

scansorial pangolins of the genus Manis (Gaudin, unpublished data). Double processes are also known to occur 

on the caudals of semiaquatic mammals (e.g., pantolestids [Rose and Koenigswald, 2005], beavers, otters, and 

aquatic sloths [Muizon and McDonald, 1995]).

The posteriormost caudal vertebrae in modern pangolins are quite robust. Robust transverse processes (single, 

not double) extend nearly to the very tip of the tail; for 

example, in M. javanica USNM 198852 they are found 

in 28 of 29 vertebrae, in Phataginus tetradactyla USNM 

481406 they are present in 46 of 49 vertebrae, and in 

Smutsia gigantea AMNH 53848 they are present in all 

27 caudal vertebrae (there are actually double transverse 

processes found only on Ca27). Similarly, chevron articulations are found almost all the way to the tip of the tail: 

to Ca23/24 in M. javanica USNM 198852, to Ca44/45 

in Phataginus tetradactyla USNM 481406, and to Ca25 

in Smutsia gigantea AMNH 53848. The centra of these 

vertebrae are cylindrical like those of the mid-caudals but 

become not only smaller but also somewhat more dorsoventrally compressed posteriorly, and of course, they 

lose progressively their neural spines and zygapophyseal 

articulations. The zygapophyseal processes themselves 

persist, the posterior disappearing first; for example, in 

Phataginus tricuspis CM 16206, the last vestiges of the 

posterior zygapophyses are lost on the 36th of 37 preserved caudal vertebrae (the specimen is likely missing the 

last one or two caudals), whereas there are very small 

vestiges of the anterior zygapophyses on Ca37. The terminal caudals present in Patriomanis, as noted above, are 

the least well preserved, and many lack complete centra 

(Figure 21a). As in Patriomanis’s living relatives, the centra become smaller and somewhat more depressed posteriorly, the neural spines and vertebral canals disappear, 

FIGURE 21. Distal caudal vertebrae and chevrons of Patrioman

americana, USNM-P 299960: (a) distal caudal vertebrae in ventral 

view, with posteriormost preserved caudal at upper left, progressively more anterior caudals to the right; lower left caudal anterior 

to vertebra at upper right, lower right vertebra representing the most 

anterior in the sequence; (b) chevrons in left lateral view, anterior 

to the left, with leftmost representing the first chevron in the caudal 

series (note single pair of dorsal facets). Scale bars = 1 cm. Photos 

by Tim Gaudin, composed and labeled by Julia Morgan Scott for 

Smithsonian Institution.
3 6 • 

other chevrons, the first has but a single pair of dorsal 

facets near its posterior edge.

Axial Skeleton: Ribs and Ster

Figures 

Ribs

Although Emry (1970) does not describe this portion of the axial skeleton in the type specimen, the type 

does preserve a few rib fragments, including a proximal 

end of a left rib and three fragments that are derived from 

mediolaterally expanded rib shafts (Figure 22a), like those 

described for Cryptomanis (Gaudin et al., 2006). USNM-P 

494439 preserves additional ribs, including five fairly large 

fragments prepared free, plus three left and seven right ribs 

preserved in situ in the matrix. USNM-P 299960 includes 

11 ribs for which the proximal ends are intact (two from 

the left, four from the right, five indeterminate), plus many 

rib fragments. Three nearly complete ribs from the middle 

of the series are illustrated in Figure 22b.

caudals of Cryptomanis (Gaudin et al., 2006) and Eomanis (Storch, 1978), although perhaps somewhat more 

robust than either. This morphology is very reminiscent 

of the pattern in vermilinguan anteaters with a prehensile 

tail (e.g., Tamandua mexicana, CM 76827), as well as 

prehensile-tailed marsupials, primates, and carnivorans 

(Argot, 2003; Youlatos, 2003; Organ, 2010), although 

in the latter the centra tend to be somewhat thinner and 

elongated anteroposteriorly.

Although Emry (1970) notes that no chevron bones 

were preserved with the type, a few chevron bones have 

been recovered from the other Patriomanis specimens 

(Figure 21b). These are generally rectangular, with their 

dorsoventral depth much greater than their anteroposterior length. They have robust processes that extend far 

ventrally beneath the hemal canal. Most are hourglass 

shaped in lateral view, expanding anteroposteriorly at the 

distal and proximal ends. The first, however, tapers distally and is recurved posteriorly in lateral view. It strongly 

resembles the condition in M. javanica USNM 198852, in 

which the first chevron is tapered and recurved posteriorly 

and the remainder are more hourglass shaped. Unlike the 

FIGURE 22. Ribs of Patriomanis american

(a) large fragment from anterior, transversely 

expanded rib in F:AM 78999 (holotype), shown in 

anterior view, with proximal end pointed upward; 

(b) three nearly complete ribs from the middle of 

the series in USNM-P 299960, shown in posterior 

view—the ribs on the left and right have their proximal ends pointing upward, whereas in the middle 

specimen the proximal end is pointed downward. 

Abbreviations: cap, capitulum (= head); ne, neck; 

rs, rib shaft; tub, tubercle. Scale bars = 1 cm. Photos by Tim Gaudin, composed and labeled by Julia 

Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 3 7

2006) but differs markedly from the morphology in modern pangolins. In Phataginus, the sternebrae are dorsoventrally flattened, whereas in Manis, S. temminckii, and 

some S. gigantea (the remainder of S. gigantea resemble 

Phataginus), the sternebrae have a deep anteroventral 

process, and the heads of the ribs overlap ventrally the 

posterior portions of the sternebrae (Gaudin et al., 2006, 

2009). In Eomanis (Storch, 1978), the individual sternebrae are wider transversely than they are long, as in living 

manids but not patriomanids (Gaudin et al., 2006), and 

they appear flattened dorsoventrally, as in the modern species of Phataginus (Gaudin et al., 2009). The manubrium 

is preserved in the type of Eomanis waldi (Storch, 1978), 

and a partial manubrium and xiphisternum are preserved 

in Cryptomanis (Gaudin et al., 2006). The reader is 

referred to the original descriptions and to Gaudin et al. 

(2009) for more information on these elements. 

Because there are three transversely expanded fragments in the type from three different ribs, it can be 

inferred that both the first and second rib shafts were 

expanded transversely relative to more posterior ribs, as is 

the case in Eomanis, Euromanis, the extant genera Smutsia and Manis, and at least the first rib in Cryptomanis

(Gaudin et al., 2009). The capitular facets of the ribs are 

convex both anteroposteriorly and transversely, more so 

in the former plane, and are separated by a distinct neck 

of varying length from flatter tubercular facets, which are 

supported on a distinct tubercular boss and face dorsomedially (like the capitulum) and somewhat posteriorly. 

There is a small muscular fossa on the proximal and posterior surface of the shaft, just ventral to the tuberculum. 

The shaft itself is flattened anteroposteriorly and of varying curvature. Cryptomanis has very similar ribs, as does 

Eomanis, whereas in Euromanis the ribs are more cylindrical, and in modern pangolins the rib shafts are wider 

anteroposteriorly than they are mediolaterally (Gaudin, 

unpublished data).

Sternum

The only remnant of the sternum of Patriomanis 

appears to be a set of isolated bones from USNM-P 299960 

(Figure 23). These include two articulated sternal elements, 

likely representing the penultimate (i.e., the sternebrae to 

which the xiphisternum is attached) and antepenultimate 

elements, and an isolated, more anterior sternebra. The 

two anterior sternebrae (i.e., the isolated and antepenultimate elements) are hourglass shaped in ventral view, with 

paired rib facets on their lateral surfaces at both anterior 

and posterior ends, the latter adjoining nearly identical facets on the anterior end of the next sternebra. The 

transverse widths of the anterior sternebrae are nearly 

equivalent to their anteroposterior lengths (e.g., 46.4 mm/

52.2 mm = 0.89 in the antepenultimate segment), but what 

is more striking is the depth of the elements, which is nearly 

two-thirds their length. This depth gives the lateral rib facets of the adjacent sternebrae, when taken together, a deep 

U shape, with the opening of the U at the dorsal end (Figure 23b). The more posterior sternebra is taken to be the 

penultimate sternebra because it is twice the length of the 

anterior element and bears an extra pair of rib facets near 

its anteroposterior midpoint. It would then presumably 

have had a third pair of rib facets, along with a median 

distal articulation for the xiphisternum at its semicircular 

terminus, but these remain covered by matrix. 

The morphology of the sternebrae in Patriomanis 

closely resembles that of Cryptomanis (Gaudin et al., 

FIGURE 23. Sternebrae of Patriomanis americana, USNM

299960, including the penultimate (far left), antepenultimate, and a 

more anterior sternebra: (a) in ventral view; (b) in left lateral view. 

Abbreviation: ccf, costal cartilage facet. Scale bar = 1 cm. Photos 

by Tim Gaudin, composed and labeled by Julia Morgan Scott for 

Smithsonian Institution.
3 8 • 

32c; Grassé, 1955; Guth, 1958; Gebo and Rasmussen, 

1985; Koenigswald, 1969, 1999; Koenigswald and Martin, 1990; Gaudin et al., 2006, 2009) with the exception of 

Eomanis and Euromanis (Storch, 1978; Storch and Martin, 

1994). The fissure measures as much as half the length of 

the ungual in dorsal view (Gaudin, unpublished data), as it 

does in the extant pangolin species. The unguals terminate 

in rather blunt rounded tips, a characteristic in which they 

differ from the living manids, where the unguals terminate 

in sharp points (Figure 32c). In dorsal view, the unguals 

are “waisted” in appearance, with a distinct constriction 

anterior to their base. The unguals expand slightly anterior to this constriction before narrowing again toward 

the tips (Figure 24). A very similar morphology is reported 

in Necromanis (Koenigswald, 1969, 1999; Koenigswald 

and Martin, 1990) and Cryptomanis (Gaudin et al., 2006) 

and is observed in Eomanis, Euromanis, and Manis pentadactyla but is not present in other extant manid species 

(Gaudin et al., 2009).

In lateral view, the unguals are roughly triangular, elevated proximally and tapering distally with a flat ventral 

surface and a gently convex dorsal surface (Figure 25a). 

The proximal surface is deeply concave in lateral view as in 

Hind Li

Figures 24–40; T

Pedal Unguals

As noted by Emry (1970), very little of the pes is preserved in the type specimen of Patriomanis (F:AM 78999). 

None of the pedal unguals are preserved. However, some 

of the pedal unguals are preserved in USNM-P 494439. 

Some are apparently lost in this specimen, and a few have 

been prepared free of the matrix. A few pedal unguals are 

also available from USNM-P 531557. A virtually complete set of left and right pedal unguals has been preserved 

with USNM-P 299960 (Figure 24), and hence, the description that follows is based largely on that specimen.

The five unguals are broadly similar in size, the discrepancy between the largest and smallest being less in 

Patriomanis than in the modern pangolin species. As in 

the modern species, the third ungual is the largest, and the 

first and fifth are the smallest (Table A4). 

The pedal unguals are deeply fissured (Figures 24, 25), 

as they are in all extinct and extant pholidotans (Figure 

FIGURE 24. Pes of Patriomanis america

(a) right pes of USNM-P 299960 in dorsal view; 

(b) labeled outline drawing of same. Abbreviations: 

I–V, digits I–V; ast, astragalus; clc, calcaneus; cu,

cuboid; ecc, ectocuneiform; enc, entocuneiform; 

ip, intermediate phalanx; mec, mesocuneiform; mt, 

metatarsal; nav, navicular; pp, proximal phalanx; 

prh, prehallux; up, ungual phalanx. Scale bar = 

1 cm. Illustration (a) by Julia Morgan Scott (JMS) 

for Smithsonian Institution (SI); (b) by Larry Isham, 

digitized by JMS for SI.
n u m b e r 9 8 • 3 9

its proximal end. The left and right foramina are equivalent in size. These foramina connect to the grooves lying 

above the subungual process and presumably conveyed 

blood vessels toward the vascular foramina at the proximal ends of those grooves. Distally, the subungual process bears a short, low, rounded median crest flanked by 

two shallow concavities. This crest doubtless served as the 

insertion point for the tendons of the deep digital flexors 

(i.e., the tendons of the m. flexor fibularis, following Jouffroy, 1966). The subungual process of Patriomanis closely 

resembles that of Cryptomanis and the extant species M. 

pentadactyla and M. javanica but differs significantly from 

the narrower, strongly constricted subungual process of 

Phataginus (Gaudin et al., 2009). 

Pedal Intermediate Phalanges

These elements are not preserved in the type (Emry, 

1970), but several intermediate phalanges are preserved 

and have been prepared in the right and left pes of USNMP 494439. However, as a complete right and left series is 

available in USNM-P 299960 (Figure 24), the description 

that follows is based on that specimen. 

As was the case with the ungual phalanges, the third 

intermediate phalanx is the longest, and the fifth is the shortest, although the second intermediate phalanx is approximately equal in length to the third in Patriomanis (Table

A4). In dorsal view the intermediate phalanges are cylindrical and about half again as long as they are wide. They are 

slightly constricted in the middle and expanded at either 

end. Distally, they bear strongly convex, subequal medial 

and lateral condyles for articulation with the ungual phalanges. Immediately proximal to these condyles are paired 

dorsolateral concavities, presumably sites of origin for the 

collateral ligaments of the interphalangeal joint (based on 

anatomy of humans and dogs; Pick and Howden, 1977; 

Evans and Christensen, 1979). These concavities form distinct indentations behind the condyles visible in dorsal view 

(Figure 24). Although the concavities themselves are present in extant manids, they do not form dorsal indentations 

behind the condyles (Figure 32c; Gaudin et al., 2009). 

The proximal articular surface is deeply concave in 

lateral view. It bears a single dorsal process and paired 

ventral processes that embrace the condyles of the proximal phalanges. The articular surface is approximately as 

wide as it is tall. The articular surface resembles those of 

the ungual phalanges in that it is divided by a central ridge 

into medial and lateral concavities that receive the medial 

and lateral condyles of the proximal phalanges. These 

concavities are flanked by well-developed lateral ridges 

in Patriomanis, a point of resemblance to Cryptomanis 

modern pangolins, with dorsal and ventral extensions that 

wrap around the distal articular surface of the intermediate 

phalanx. The ungual process is divided from the subungual 

process by a deep longitudinal groove on both the medial 

and lateral sides of the bone. A large vascular foramen is 

located at the proximal end of this groove (Figure 25a). 

The proximal articular surface is ovate, slightly broader 

transversely than it is tall. In this respect it resembles Cryptomanis and M. pentadactyla but differs markedly from 

other modern species in which the pedal unguals are compressed laterally (e.g., M. javanica, Phataginus tricuspis). 

The articular surface is divided by a prominent median 

ridge into two dorsoventrally elongate concave surfaces 

that receive the medial and lateral condyles of the intermediate phalanges. 

In ventral view the pedal unguals have an extensive 

subungual process that occupies the proximal half of the 

bone (Figure 25b). The process takes the form of a rounded 

triangular platform separated by the aforementioned 

groove from the ungual process. The subungual process 

is perforated by two large oval subungual foramina at 

FIGURE 25. Ungual phalanx II of right pes of Patriomanis americana (USNM-P 299960): (a) in medial view; (b) in ventral vie

Abbreviations: sunf, subungual foramen; sunp, subungual process; vf, vascular foramen. Scale bar = 0.5 cm. Photos by Tim 

Gaudin, composed and labeled by Julia Morgan Scott for Smithsonian Institution.
4 0 • 

The proximal articular surface itself is concave, although 

lacking the dorsal and ventral proximal prongs present in 

the other phalanges (Figure 26b). The lateral edge of the 

proximal end is actually slightly convex in lateral view. This 

articular surface also lacks the median keel that divides the 

proximal articular surface in the other phalanges. The facet 

is divided in its ventral half by a narrow midline depression 

that receives the keel of the distal metatarsals. This gives the 

articular surface a horseshoe-like shape, a shape that resembles that of Cryptomanis but differs from the extant species 

in which the entire articular surface is deeply divided by the 

metatarsal keel. In lateral view the proximal phalanges of 

Patriomanis are again shaped much like the intermediate 

phalanges, compressed in the middle with a triangular base 

and pulley-shaped condyles.

The first proximal phalanx differs somewhat in 

its morphology from the other four. It is more slender 

(Figure 24), with its maximum length more than twice its 

lacking in modern pangolin species (Gaudin et al., 2009). 

In lateral view the intermediate phalanges are expanded 

proximally and distally and constricted in the middle, as 

in dorsal view. The base of the bone is triangular, and the 

condyles are pulley shaped.

Pedal Proximal Phalanges

Again, these elements are not preserved in the type 

(Emry, 1970). Proximal phalanges are preserved and have 

been prepared in the right and left pedes of USNM-P 

494439 and USNM-P 299960. As before, the following 

description is based largely on the complete set available 

from the latter specimen (Figure 24). 

The relative lengths of the proximal phalanges are more 

variable among living manids than was the case for the 

other phalanges. The fifth proximal phalanx is the shortest 

in all extant species, and the first is the longest in the majority of extant species (Gaudin et al., 2009). However, in 

M. pentadactyla the proximal phalanges become progressively shorter laterally, whereas in P. tricuspis the third proximal phalanx is approximately equal in length to the second 

and fourth (Figure 32c), and in M. javanica, S. gigantea, 

and Cryptomanis the third is decidedly longer than the second and fourth. In Patriomanis the first through third proximal phalanges are of approximately equal length, with the 

fourth shorter and the fifth shorter still (Table A4). 

In dorsal view the proximal phalanges of Patriomanis 

closely resemble the intermediate phalanges, with a cylindrical shaft that is constricted in the middle and expanded 

at either end (Figure 24). As in Cryptomanis, the proximal 

phalanges are slightly longer than the intermediate phalanges in Patriomanis (Table A4). In every living species 

except Manis javanica the opposite is the case (Figure 32c). 

Indeed, in S. gigantea, M. pentadactyla, and M. crassicaudata the proximal phalanges are compressed proximodistally to the point that they no longer have a cylindrical 

shaft. The width of the proximal phalanges measured at 

the distal condyles in these taxa is as great as or greater 

than the overall length of the bone. The proximal phalanges have two subequal pulley-shaped distal condyles 

(Figures 24, 26d). These phalanges in Patriomanis lack a 

dorsal process at their proximal end that would serve to 

embrace the more proximal metatarsals, and hence, the 

metatarsal–phalangeal joint appears somewhat less stable

than the interphalangeal joints. In manids, a large distal 

keel stabilizes the metatarsal–phalangeal joint (Figure 

32c). As noted in subsequent descriptions, this keel is not 

developed dorsally in Patriomanis or in Cryptomanis or 

Necromanis (Gaudin et al., 2006, 2009). 

FIGURE 26. Proximal phalanges of Patriomanis americ

(USNM-P299960): (a) proximal phalanx III of left manus in proximal view; (b) proximal phalanx III of right pes in proximal view; 

(c) proximal phalanx III of left manus in distal view; (d) proximal 

phalanx III of right pes in distal view. Scale bars = 1 cm. Photos 

by Tim Gaudin, composed and labeled by Julia Morgan Scott for 

Smithsonian Institution. 
n u m b e r 9 8 • 4 1

as its sagittal mobility. Such a keel is present on only the 

ventral half of the distal condyles of the metatarsals in 

Patriomanis (Figure 27b) and Cryptomanis (Gaudin et al., 

2006) and is not as robust as the keel present in the modern species. With the exception of metatarsal I, the metatarsal articulations with the tarsals are flat in Patriomanis. 

They have a similar morphology in Cryptomanis (Gaudin 

et al., 2006) and apparently in Necromanis (Koenigswald 

and Martin, 1990). In extant pangolins the distal tarsals 

show extensive overlap onto the dorsal surfaces of both 

metatarsals I and III. They overlap metatarsals II and IV 

to a lesser degree (Figure 32c). 

The shafts of the metatarsals of Patriomanis are wide 

and flat, their width exceeding their depth by 20% or more 

in all five digits. In Cryptomanis, the metatarsals are even 

wider, with all but digit I 50% wider than deep. Among 

modern pangolins, the depth at midshaft of the metatarsals 

is as great as or greater than the width for at least some of 

the digits, although the pattern varies among the living species (digits II–IV deepened in Phataginus and S. temminckii, 

digits I–IV in S. gigantea, M. pentadactyla, and M. crassicaudata, and only digit I in M. javanica). In dorsal view 

the metatarsals are roughly five times longer than wide, 

as in most manids, and most are slightly expanded both 

proximally and distally. The distal condyles appear slightly 

widened transversely and evenly convex in dorsal view. The 

condyles are distinctly narrower than the epicondyles (Figure 24), in contrast to the condition in the Manidae (Figure 32c; Gaudin et al., 2009). Immediately proximal to the 

condyles of metatarsals II–V is a midline depression on the 

dorsal surface. On their ventral surface each metatarsal has 

depth measured at the proximal end (its deepest point). In 

this respect it resembles the condition in Cryptomanis and 

P. tricuspis but differs from other extant species in which 

the first proximal phalanx is shorter and deeper (Gaudin 

et al., 2009). The ventral edge of the first proximal phalanx is gently concave in lateral view. In living pangolins 

(including P. tricuspis) the ventral edge is gently convex 

for roughly two-thirds its length, then indented sharply 

just proximal to the distal condyles. No such indentation 

is present in Patriomanis.

Metatarsals

In the type specimen, the only preserved portions of 

the metatarsals are the proximal ends of the right first and 

fifth metatarsals and the distal ends of the second and third 

metatarsals. In USNM-P 531556 the entire right fifth metatarsal is preserved, and USNM-P 531557 includes a distal 

end of what appears to be the second metatarsal on the right 

side. In USNM-P 494439 all five metatarsals are preserved 

intact on the left foot, although they remain embedded in 

the matrix ventrally. On the right pes, the third metatarsal 

is missing, and the other four are damaged to a certain 

extent. USNM-P 299960 preserves a complete series on 

the right (Figure 24). The first and second metatarsals are 

badly damaged on the left side. The left fifth metatarsal is 

also slightly damaged proximally and laterally.

As was the case for the proximal phalanges, there is 

some variation among pholidotans in the relative lengths 

of the metatarsals. In Patriomanis, the first metatarsal is 

the shortest, and the third is the longest (Table A4). These 

proportions are similar to those of Cryptomanis and of 

extant Smutsia and Phataginus, except that in the latter 

two genera the third and fourth metatarsals are subequal 

in length. Among Manis species the metatarsals exhibit 

several different patterns of relative lengths. The overall length of the digits relative to body size seems to be 

greater in Patriomanis and Cryptomanis (Gaudin et al., 

2006) than in extant manids, a difference reflected in the 

lengths of the metatarsals. The length of the third metatarsal in Patriomanis is 35% that of the tibia, whereas in 

modern pangolins the third metatarsal is 25% or less of 

tibial length (Table A4). 

Accompanying a shortening of the digits in the modern pangolins is an apparent increase in mobility of both 

the metatarsal–tarsal joints and the metatarsal–phalangeal 

joints. At the distal end of the metatarsals, extant pangolins have a robust median keel that extends dorsal and 

ventral to the shaft (Figure 32c). This keel increases the 

lateral stability of the metatarsal–phalangeal joint as well 

FIGURE 27. Metapodials of Patriomanis americana in distal v

(USNM-P 299960): (a) left metacarpal III; (b) left metatarsal III. 

Abbreviations: das, distal articular surface; k, ventral keel. Scale 

bar = 1 cm. Photos by Tim Gaudin, composed and labeled by Julia 

Morgan Scott for Smithsonian Institution. 
4 2 • 

Cryptomanis (Gaudin et al., 2009). It is broadly expanded 

dorsally and over the plantar tubercle ventrally and sharply 

constricted in between. In Manidae the facet lacks either the 

constriction or the ventral expansion or both and takes on a 

triangular or quadrangular appearance.

Metatarsal IV. The proximal end of the fourth 

metatarsal bears two medial facets for metatarsal III. It 

is deeply excavated on its lateral surface. Situated in the 

dorsal portion of this excavation is a kidney-bean-shaped 

facet for metatarsal V. In dorsal view the proximal end 

of the bone is elevated along its lateral edge, as is the 

third metatarsal (Figure 24). The dorsal surface itself is 

fairly flat, as in Cryptomanis, but contrasts with the condition in modern manid species in which this region of 

the fourth metatarsal is either markedly concave (Manis, 

S. gigantea) or deeply grooved (Phataginus, S. temminckii; 

Gaudin et al., 2009). In proximal view the articular facet 

for the cuboid is broad dorsally. It narrows ventrally but 

is slightly expanded at its ventral extremity. This ventral 

expansion helps to enclose a foramen formed between 

the proximal articular surfaces of the third and fourth 

metatarsals. This foramen is connected by a groove in 

the medial surface of the cuboid to a second foramen that 

opens more proximally between the dorsal surfaces of the 

cuboid and ectocuneiform. The arrangement in Patriomanis is very similar to the condition in M. crassicaudata. 

The latter species appears to be the only modern pangolin 

with the foramen between metatarsal III and IV, although 

the foramen between the cuboid and ectocuneiform is 

more widely distributed among the living species. Both 

foramina are absent in Cryptomanis (Gaudin et al., 2006, 

2009). Given that the foramen between cuboid and ectocuneiform appears likely to have accommodated a ligament (see ectocuneiform description below), the foramen 

between metatarsals III and IV is likely related to the network of tarsal and metatarsal ligaments as well.

Metatarsal V. The proximal end of the fifth metatarsal is characterized not only by a broad plantar tubercle 

on its ventral surface but also by a prominent lateral process. 

This process lies at the same level as the proximal articular 

facet for the cuboid but is separated from this articular facet 

by a well-marked groove. The process is somewhat elongated 

dorsally and distally in lateral view. An even more prominent 

lateral process is present on metatarsal V in Cryptomanis 

and in modern pangolins (Figure 32c). The dorsal surface of 

metatarsal V is weakly concave at its proximal end in Patriomanis (Figure 24). Metatarsal V articulates with the cuboid 

proximally. The cuboid facet is D shaped in the type (F:AM 

78999) and in USNM-P 299960, with a straight medial edge 

and a curved lateral margin. It is more elongate transversely 

in USNM-P 531556. The cuboid facet is contiguous on its 

a distal condyle bearing the aforementioned midline keel 

(Figure 27b). In addition, a robust plantar tubercle extends 

ventrally from the proximal end of each metatarsal. 

Metatarsal I. In dorsal view the shaft of the first 

metatarsal is slightly bowed medially (Figure 24). At its 

proximal end the first metatarsal articulates laterally with 

metatarsal II and proximally with the entocuneiform. The 

proximal articular facet for the entocuneiform is triangular 

in proximal view. It is broad ventrally and narrows dorsally. It is strongly convex, the narrow dorsal portion overlapping extensively onto the dorsal surface of the shaft, as 

noted above. In all of these features Patriomanis resembles 

Cryptomanis (Gaudin et al., 2006) and the extant pangolin 

species. On the medial side of its proximal end, the first 

metatarsal bears a low, rugose proximodistally elongate 

ridge. A similar ridge is present in Cryptomanis and most 

extant manids (absent only in Smutsia gigantea).

Metatarsal II. Metatarsal II is the only metatarsal 

that is not markedly expanded proximally in dorsal view 

(Figure 24), a feature in which Patriomanis resembles Cryptomanis (Gaudin et al., 2006) and Necromanis (Koenigswald 

and Martin, 1990) but differs from extant manids (Figure 

32c). On its proximal end it articulates medially with the first 

metatarsal and entocuneiform, proximally with the mesocuneiform, and laterally with the third metatarsal. The latter 

articulation is actually formed by two distinct facets, one 

carried dorsally on the shaft and a second carried ventrally 

on the plantar tubercle. Separate dorsal and ventral contacts 

between the second and third metatarsals are also present in 

Cryptomanis, M. javanica, and M. crassicaudata (Gaudin 

et al., 2009). The proximal articular surface is roughly rectangular, elongated dorsoventrally with indentations along 

its medial and lateral edges. A similar indentation along the 

lateral margin of the proximal articular surface is present in 

P. tricuspis, S. gigantea, M. javanica, and M. crassicaudata. 

Metatarsal II of Patriomanis bears a large depression on the 

posteromedial surface at its proximal end. Such a depression 

is also found in Cryptomanis (Gaudin et al., 2009), although 

it is not present among living pangolins, suggesting it may be 

a patriomanid feature.

Metatarsal III. This metatarsal articulates at its 

proximal end with metatarsal II medially, the ectocuneiform 

proximally, and metatarsal IV laterally. As was the case with 

its medial connection, its lateral connection to metatarsal IV 

is composed of a dorsal facet on the shaft and a ventral facet 

on the plantar tubercle, a point of resemblance to Cryptomanis, Manis javanica, and M. crassicaudata (Gaudin 

et al., 2009). In dorsal view the proximal articular surface 

is elevated along its lateral edge, as in manids (Figure 24). In 

proximal view, the articular surface for the ectocuneiform 

is vaguely dumbbell shaped, resembling the condition in 
n u m b e r 9 8 • 4 3

Cryptomanis the prehallux is a rodlike element (Gaudin 

et al., 2006). However, in Patriomanis, as in extant pangolins, the prehallux is laterally compressed. In medial view 

it is roughly triangular in shape, narrow proximally and 

expanded distally, as in Smutsia, M. javanica, and P. tetradactyla. The prehallux of Patriomanis is convex medially 

and concave laterally and slightly twisted about its long 

axis, such that the dorsodistal corner is rotated medially 

approximately 20o. It lacks any definitive articular facets 

on its lateral side, and there are no obvious articular facets 

on the medial side of the entocuneiform. There is a small 

boss on the proximomedial surface of the navicular that 

may have served as a ligamentous attachment point. By 

contrast in some extant species there are well-developed 

articular facets on the proximal end of the prehallux for 

the navicular (M. javanica, M. crassicaudata) or the navicular and entocuneiform (Phataginus; Gaudin et al., 2009). 

Entocuneiform

Although missing from the type and from the right 

side of USNM-P 299960, a nearly complete left entocuneiform is preserved in the latter specimen. In addition, both 

left and right entocuneiforms are intact and available for 

study in USNM-P 494439.

medial edge with the facet for metatarsal IV. However, the 

two facets are separated by a sharp ridge and are oriented 

at an obtuse angle to one another, with the cuboid facet facing mostly proximally, as well as dorsally and medially, and 

the facet for metatarsal IV facing mostly medially, as well as 

proximally and dorsally. The latter facet is L shaped, with 

one leg extended dorsoventrally and the other extending distally and dorsally. It is convex along its long axis. The morphology of these proximal articulations is generally similar in 

Patriomanis and manids, although the angle between cuboid 

and metatarsal IV facets and the shape of these facets varies 

somewhat among Patriomanis and the various extant species. In Cryptomanis, the division between the facets for the 

cuboid and metatarsal IV is much less marked, and the former facet is damaged so that its shape cannot be determined.

Prehallux

In USNM-P 299960 a sesamoid bone is preserved on 

the medial side of the right pes, abutting the entocuneiform 

(Figure 24). A similar element is present in Cryptomanis 

(Gaudin et al., 2006), all extant manid species (Figure 

32c), and certain fossil armadillos (Scott, 1903–1904) and 

is termed by Szalay and Schrenk (1998) a “prehallux.” 

We will follow the nomenclature of the latter authors. In 

FIGURE 28. Distal tarsals of Patriomanis americana: (a) left entocuneiform in medial vi

(USNM-P 299960); (b) left entocuneiform in lateral view (USNM-P 299960); (c) left mesocuneiform in proximal view (USNM-P 299960); (d) left 

mesocuneiform in distal view (USNM-P 299960); 

(e) right ectocuneiform in proximal view (USNMP 494439); (f) right ectocuneiform in distal view 

(USNM-P 494439). Abbreviations: mcf, mesocuneiform facet; mIIf, metatarsal II facet; naf, navicular facet; plp, plantar process. Scale bar = 0.5 cm

Photos by Tim Gaudin, composed and labeled by 

Julia Morgan Scott for Smithsonian Institution.
4 4 • 

the navicular and metatarsal II, respectively (Figure 28c,d), 

and smaller articulations medially and laterally with the 

entocuneiform and ectocuneiform. It has a rugose, nearly 

square exposure on the dorsal surface of the tarsus (Figure 24). In Cryptomanis, Manis, and Smutsia, the dorsal 

surface of the mesocuneiform is elongated mediolaterally 

(Gaudin et al., 2009). The navicular facet is ovate, slightly 

narrower ventrally than dorsally, with its dorsoventral axis 

somewhat longer than its greatest transverse diameter (ratio 

of depth/width = 1.3; Figure 28c). Among living pangolins 

it most closely resembles Phataginus. In Cryptomanis, the 

facet is more quadrangular, and in Smutsia it is elongated 

dorsoventrally and indented along its medial edge, whereas 

in Manis it is both elongated dorsoventrally (ratio of greatest depth/width ≥ 1.8) and L shaped, with a distinctive 

ventrolateral indentation (Gaudin et al., 2009). The facet 

for metatarsal II in Patriomanis is also wider dorsally than 

ventrally, giving it a rounded triangular appearance (Figure 

28d). It closely resembles the facet for metatarsal II in living pangolins. On the medial surface of the mesocuneiform 

is a small teardrop-shaped facet for the entocuneiform. The 

facet lies in the dorsal and proximal corner of the medial 

surface and tapers ventrally, reaching approximately to the 

midpoint of the bone. On the lateral surface of the mesocuneiform is an ovate facet for the ectocuneiform. This facet 

is situated in the middle of the lateral surface and covers 

roughly half of its dorsoventral extent. The facet contacts 

both the proximal and distal margins of the lateral surface. 

The ectocuneiform facet resembles that of Cryptomanis 

and Manis but differs from the African pangolin species in 

which the facet is elongated dorsoventrally, occupying at 

least 75% of the dorsoventral extent of the lateral mesocuneiform (Gaudin et al., 2009).

Ectocuneiform

The ectocuneiforms are not preserved in the type, but 

both left and right ectocuneiforms are well preserved in 

USNM-P 494437 and USNM-P 299960.

Like the mesocuneiform, it is a blocky element, but it 

is substantially larger than the mesocuneiform. Its dorsal 

exposure on the tarsus is rugose and slightly wider than 

it is tall in a proximodistal direction (ratio = 1.2; Figure 

24). In living pangolins the width of the dorsal surface of 

the ectocuneiform ranges from one and a half to two times 

greater than its height, whereas in Cryptomanis this surface 

is nearly square (Gaudin et al., 2009). The proximal end of 

the bone bears a robust plantar process that is separated 

by a distinct neck from the body of the bone (Figure 28e). 

This process is approximately circular in cross section. In 

The morphology of this bone in Patriomanis is similar 

to that encountered in most extant pangolin species. It is a 

triangular element, broad distally and tapering proximally 

in medial view (Figure 28a). It is strongly compressed 

mediolaterally and bears a large concavity on its medial 

surface and a prominent boss on its ventrodistal corner. 

The distal articular facet for metatarsal I is strongly concave and, as noted previously, overlaps the dorsal surface 

of the shaft of the metatarsal (Figure 24). In distal view this 

articular facet is oval in outline, much like that of Cryptomanis and Phataginus (Gaudin et al., 2009). In other 

extant species the facet takes on a triangular (Smutsia, 

M. crassicaudata) or sigmoid shape (M. javanica, M. pentadactyla; Gaudin et al., 2009). Articular surfaces for three 

different bones, navicular, mesocuneiform, and metatarsal 

II, are present on its lateral side (Figure 28b). In Patriomanis the facets for navicular and mesocuneiform are contiguous and situated proximally along the dorsal edge of the 

lateral surface. These facets are almost circular and nearly 

the same size, the latter being slightly elongated dorsoventrally and hence slightly larger. The morphology of these 

facets is similar to that of Cryptomanis but differs notably 

from that of Phataginus, on the one hand, in which the 

mesocuneiform facet is greatly elongated dorsoventrally 

and clearly larger than the navicular facet, and Smutsia and 

Manis, on the other hand, in which it is the navicular facet 

that is enlarged along a dorsoventral axis and two separate 

and smaller mesocuneiform facets are present. The facet 

for metatarsal II lies along the distal edge of the entocuneiform in Patriomanis, well separated from the facet for the 

mesocuneiform. However, the shape and position of the 

facet differ somewhat in USNM-P 299960 and USNM-P 

494439. In the former it is a round facet situated just ventral to the dorsoventral midline of the bone (Figure 28b). In 

the latter the facet begins at approximately the same place 

but appears to have a long low extension running along the 

distal edge toward its dorsal termination. In extant manids

this facet is typically ovoid as in USNM-P 299960 but is 

situated near the dorsal edge of the entocuneiform, contiguous with the ventral edge of the mesocuneiform facet. 

It occurs in a similar position in Cryptomanis, where the 

facet is elongated proximodistally.

Mesocuneiform

The mesocuneiforms are missing from both the type 

and USNM-P 494437. However, both right and left mesocuneiforms are preserved in USNM-P 299960.

The mesocuneiform in Patriomanis is small and blocky. 

It has broad articular surfaces proximally and distally for 
n u m b e r 9 8 • 4 5

2009). In extant dogs, a similar foramen is present and 

appears to accommodate a branch of the proximal extensor retinaculum (Evans and Christensen, 1979).

Cuboid

The left cuboid is preserved intact in the type. In 

USNM-P 494439, both left and right cuboids are present, although neither has been prepared free of the matrix 

on its ventral surface. The right cuboid is well preserved 

in USNM-P 299960, but the left is badly damaged, with 

nearly the entire ventral half of the bone eroded away.

As is typical for eutherian mammals (e.g., Pick and 

Howden, 1977; Evans and Christensen, 1979; Ji et al., 

2002; Kielan-Jaworowska et al., 2004), the cuboid is by 

far the largest element in the distal tarsal row (Figure 24). 

Its maximum height and width in Patriomanis are more 

than twice that of the next largest distal tarsal, the ectocuneiform. The cuboid articulates medially with the ectocuneiform and distally with metatarsals IV and V. At its 

proximal end it bears a complex series of articulations for 

the more proximal tarsals, including facets for the navicular, astragalus, and calcaneus (Figure 29a). As in Cryptomanis (Gaudin et al., 2006), Necromanis (Koenigswald 

and Martin, 1990), and living pangolins (Figure 32c), the 

dorsal surface of the cuboid is irregular, with a flat distal 

edge and a sharply angled proximal edge formed by the 

contact of the proximally and laterally oriented calcaneal 

facet and the proximally and medially oriented astragalar 

facet. On its ventral surface the cuboid carries a prominent plantar process (Figure 29a). The process is elongated 

transversely at its base and is oriented perpendicular to 

the proximodistal axis of the cuboid, in contrast to the 

condition in some extant manids in which this process 

Cryptomanis and living pangolins the process is also present, although it is reduced in Phataginus and S. temminckii, 

and in M. pentadactyla, Smutsia, and Phataginus it lacks a 

distinct neck separating it from the body of the ectocuneiform. The ectocuneiform articulates proximally with the 

navicular and distally with metatarsal III. The proximal 

articular surface is in the shape of a rounded triangle in 

USNM-P 299960. In USNM-P 494439, the facet takes on 

more of an L shape, with a marked ventromedial concavity (Figure 28e). The distal facet is more angular and also 

varies somewhat in its shape between the two specimens. 

In USNM-P 299960, it is trapezoidal, almost triangular, 

with its dorsal width more than twice its ventral width. In 

USNM-P 494439 the ventral width of the facet is greater, 

with the dorsal end only one and a half times wider than 

the ventral end. The two ends of the facet are separated by 

sharp medial and lateral indentations (Figure 28f), perhaps 

a less developed form of the distinctive T-shaped metatarsal facet of the ectocuneiform in Cryptomanis and in 

M. javanica and M. crassicaudata. In other extant manids 

this facet is quadrangular or trapezoidal as in USNM-P 

299960 (Gaudin et al., 2009). On its medial edge the ectocuneiform articulates with the mesocuneiform, as described 

above. On its lateral surface, the ectocuneiform of Patriomanis has two discrete facets for the cuboid. The more 

proximal facet is localized in the dorsal half of the lateral 

surface, and the more distal facet extends along the entire 

distal edge of the lateral surface. The proximal and distal 

facets are separated by a shallow groove that manifests 

itself on the dorsal surface of the tarsus as a large foramen 

between the ectocuneiform and cuboid (Figure 24). This 

foramen is present in Necromanis (Koenigswald and Martin, 1990) and most extant manids (except Smutsia and P. 

tricuspis) but is absent in Cryptomanis (Gaudin et al., 2006, 

FIGURE 29. Tarsals of Patriomanis america

(USNM-P 299960): (a) right cuboid in medial view, 

dorsal to the left and proximal toward the top of 

photo; (b) left navicular in proximal view, lateral 

to the left, dorsal toward the bottom of the photo. 

Abbreviations: astf, astragalar facet; cuf, cuboid 

facet; ecf1, first (proximal) ectocuneiform facet; 

ecf2, second (distal) ectocuneiform facet; naf, navicular facet; plp, plantar process. Scale bar = 0.5 cm. 

Photos by Tim Gaudin, composed and labeled by 

Julia Morgan Scott for Smithsonian Institution. 
4 6 • 

one modern species, P. tetradactyla, there is no contact 

between the navicular and cuboid. In addition, the shape 

and proportions of the astragalar and navicular facets are 

variable. As is the case with most other pedal elements, 

the anatomy of this region of the cuboid in Patriomanis is 

most similar to that of Cryptomanis.

Navicular

The navicular is not preserved in the type. In USNM-P 

494439, the distal surface of the right navicular is visible 

within the matrix. Both the left and right naviculars are 

fully prepared and well preserved in USNM-P 299960.

The navicular articulates proximally with the astragalar head and laterally with the cuboid. On its distal surface 

it articulates with the entocuneiform, mesocuneiform, and 

ectocuneiform. In Patriomanis, as in Cryptomanis and living manids (Figure 32c; Gaudin et al., 2006, 2009), the 

navicular carries a prominent proximal extension along 

its medial edge. This extension wraps around the medial 

side of the astragalar head (Figure 24). As noted by Rose 

and Emry (1993), the head of the astragalus in Patriomanis lacks the distinctive concavity so characteristic of 

extant pangolins. Correspondingly, the astragalar facet of 

the navicular lacks any proximal convexity (Figure 29b). 

It is evenly concave both transversely and dorsoventrally 

and wider transversely than it is deep dorsoventrally. It 

is roughly ovate in shape with a rounded medial margin, 

although it is squared off along its lateral edge, where it 

articulates with the cuboid. The astragalar facet of the 

navicular in Cryptomanis is similar, albeit deeper dorsoventrally. As described in Gaudin et al. (2006:151), the 

facet in Cryptomanis bears a “weak proximal extension” 

on its dorsal margin that “may foreshadow the distinctive 

peg and socket articulation of modern manids.” No such 

proximal extension is present in Patriomanis. The cuboid 

facet is narrow proximodistally and elongate dorsoventrally, much like the corresponding navicular facet on 

the cuboid described previously (Figure 29b). The distal 

surface of the navicular is rugose and carries a prominent 

knob on its proximomedial portion. This knob in all likelihood served as a point of ligamentous attachment for 

the prehallux. As noted above, in some living manids the 

prehallux forms a synovial joint with the navicular. As in 

Cryptomanis, the cuneiform facets are contiguous, separated from one another by low ridges. The mesocuneiform 

and ectocuneiform facets face distally, are quadrangular 

in shape, and are roughly equivalent in size. The mesocuneiform facet is convex, whereas the ectocuneiform facet 

is concave. The latter is a feature that is apparently unique 

tilts either proximomedially (M. crassicaudata, also Cryptomanis) or proximolaterally (S. temminckii, P. tricuspis, 

and M. pentadactyla, some S. gigantea and M. javanica;

Gaudin et al., 2009). 

The distal facet of the cuboid in Patriomanis is more 

elongate dorsoventrally in its medial portion, where it 

articulates with metatarsal IV, than in its lateral portion, 

where it articulates with metatarsal V. This is the case in 

Cryptomanis and all extant pangolins as well. 

The calcaneal facet of the cuboid is strongly convex 

both dorsoventrally and proximodistally. It is ovate in 

shape, with its long axis oriented proximomedial to distolateral. It tapers to a point that extends ventrally, medially, and proximally slightly behind the astragalar facet. 

The calcaneal facet is particularly large in Patriomanis 

and Cryptomanis relative to that of extant manids. Its 

maximum length is approximately 80% of the maximum 

height of the cuboid bone itself. In modern pangolins the 

length of the calcaneal facet is only 50%–65% of the 

height of the cuboid (Gaudin et al., 2009). The astragalar 

facet is contiguous laterally with the calcaneal facet, with 

the ridge separating the two forming the highest point on 

the cuboid. It is ovate, with its long axis running dorsoventrally, and much smaller than the calcaneal facet. The 

astragalar facet lies at a slightly obtuse angle to the calcaneal facet, facing proximally and medially, as noted above, 

and is strongly concave both transversely and dorsoventrally. The astragalar facet is contiguous medially with a 

contact surface for the navicular. This latter articular surface is difficult to discern in the type since the relevant 

area is somewhat damaged in this specimen but is clearly 

evident in USNM-P 299960 (Figure 29a). The navicular 

facet is remarkably narrow proximodistally, although its 

dorsoventral length is greater than that of the astragalar 

facet. It faces medially, as does the facet for the ectocuneiform with which it shares its distal edge. The latter facet 

is one of two on the medial surface of the cuboid for the 

ectocuneiform. The upper facet is the larger of the two. It 

is oval, extends from the dorsal edge for somewhat more 

than half of the dorsoventral length of the medial surface, 

and is separated by a deep groove from the second ectocuneiform facet. This second facet is a small semicircular 

articular surface that is carried on a short process situated at the dorsal and distal corner of the medial cuboid 

surface (Figure 29a). The morphology of the medial surface of the cuboid and its astragalar, navicular, and ectocuneiform facets is highly variable among extant pangolin 

species (Gaudin et al., 2009). As noted above, some species lack the ectocuneiform–cuboid foramen and have 

only one facet for the ectocuneiform medially. In at least 
n u m b e r 9 8 • 4 7

dorsoventral plane, and extends narrowly onto the medial 

side of the neck (Figure 30). It has a width to height ratio 

greater than that of most other pangolins (Table A5). In 

Cryptomanis (Gaudin et al., 2006), the astragalar head is 

more convex dorsoventrally and deeper, extending farther 

ventrally than is the case in Patriomanis, with a broader 

exposure on the medial side of the neck. In Necromanis

(Helbing, 1938; Koenigswald and Martin, 1990), the 

astragalar head is convex dorsoventrally but flat transversely. The astragalar head of Euromanis krebsi (Storch 

and Martin, 1994) is very primitive, with an almost 

spherical shape. The head of Patriomanis appears more 

primitive than that of Cryptomanis in several respects. 

The ventrolateral edge of the head of Cryptomanis bears 

a distinct facet for the cuboid (Gaudin et al., 2006) that is 

reduced and less clearly demarcated in Patriomanis. The 

head of Cryptomanis (Gaudin et al., 2006) also possesses 

a weak concavity at its dorsal margin leading to a weak 

groove descending along the lateral edge of the head, 

which Gaudin et al. (2006) suggest foreshadows the concavity found in extant pangolins. Both features are absent 

in Patriomanis. 

The astragalar neck of Patriomanis is short, parallel 

sided, and more medially situated than is the case in most 

modern pangolins (Figure 30). A similar morphology is 

to Patriomanis among manids and patriomanids. The 

entocuneiform facet is decidedly smaller than the other 

two and directed distally and medially. The relative sizes 

and shapes of the cuneiform facets are difficult to discern 

in extant manids because of their tendency to fuse with 

one another, but they are clearly organized quite differently from those of Patriomanis and Cryptomanis and, 

indeed, vary considerably among the various extant species (Gaudin et al., 2009). 

Astragalus

Although no astragalus is preserved in the type (Emry, 

1970), both left and right astragali are preserved in USNMP 494439. However, only the left has been prepared free 

of the matrix. In addition, a well-preserved, fully prepared 

right astragalus is available from USNM-P 29960. The latter has been figured previously by Rose and Emry (1993).

The most obvious difference between the astragalus 

of Patriomanis and living manids is the presence of a 

primitive convex head in the fossil form (Rose and Emry, 

1993). In extant Manidae, the astragalar head bears 

a large lateral concavity that occupies almost its entire 

distal surface (Rose and Emry, 1993). The astragalar 

head of Patriomanis is hemispherical, is compressed in a 

FIGURE 30. Right astragalus of Patriomanis america

(USNM-P 299960): (a) in dorsal view; (b) in plantar view; 

(c) in lateral view; (d) in distal view. Abbreviations: afo, 

astragalar foramen; ef, ectal facet; ff, fibular facet; fdft, fact for 

the m. flexor digitorum fibularis tendon; h, head; ltro, lateral 

trochlea; mtro, medial trochlea; suf, sustentacular facet; vap, 

ventral astragalar process. Scale bars = 1 cm. Illustrations by 

Julia Morgan Scott for Smithsonian Institution.
4 8 • 

Jouffroy, 1966) and is a diagnostic character of the family Patriomanidae (Szalay and Schrenk, 1998; Gaudin 

et al., 2006), although it is also present in Necromanis 

(Helbing, 1938; Gaudin et al., 2009). 

The fibular facet of Patriomanis is L shaped in lateral 

view with a deep pit ventral to the proximal end of the 

facet and a ventrally projecting arm that extends from 

the distal portion of the facet (Figure 30c). This morphology closely resembles that of Cryptomanis (Gaudin 

et al., 2006) but differs from the more U- or C-shaped 

facet of extant manids (Gaudin et al., 2009). The facet 

faces laterally and slightly proximally, so that it is less 

vertical than the fibular facet of Cryptomanis (Gaudin 

et al., 2006). It has a dorsal border that forms a sharp 

edge with the lateral trochlea. The more primitive fibular 

facet in Euromanis krebsi extends farther proximally, is 

broader and more rounded distally, and is much flatter 

than that in Patriomanis. The medial side of the astragalar body bears a strong process projecting medially from 

its ventral surface (labeled “vap” in Figure 30b,c). A 

similar process is present in Necromanis (Helbing, 1938; 

Koenigswald and Martin, 1990); both are much smaller 

than the prominent process in the same location in Cryptomanis (Gaudin et al., 2006). This process is absent in 

living manids. As noted in Gaudin et al. (2006), its function is unknown.

The ventral surface of the astragalus bears an elongate concave ectal facet that tapers along its long axis 

in a distolateral direction (Figure 30b). The distolateral 

extension of this facet abuts the ventral arm of the fibular facet. This facet is similar in shape to the ectal facet 

in Necromanis (Helbing, 1938; Koenigswald and Martin, 

1990) and Cryptomanis (Gaudin et al., 2006) but is less 

concave than the latter. The ectal facet in extant manids 

is ovate and shortened anteroposteriorly (Gaudin et al., 

2009). The sustentaculum in Patriomanis is an ovate facet 

with its long axis oriented proximodistally (Figure 30b). 

It is located in the center of the astragalus, resembling 

in this aspect the primitive placental condition (Szalay, 

1977). The sustentaculum of Necromanis (Helbing, 1938; 

Koenigswald and Martin, 1990) is quite similar, whereas 

in Cryptomanis (Gaudin et al., 2006) the sustentaculum 

is larger, is more elongated distally, has migrated to the 

lateral side of the neck, and is separated by merely a shallow groove from the navicular facet. The sustentaculum in 

living manids may be dorsoventrally or transversely elongated and laterally or more centrally located and may lie 

distinct from the navicular and cuboid facets or may abut 

these facets (Gaudin et al., 2009). In many living species it 

is reduced in size, to the point of being completely absent 

present in Cryptomanis (Gaudin et al., 2006), Necromanis (Helbing, 1938; Koenigswald and Martin, 1990), and 

Euromanis krebsi (Storch and Martin, 1994). In the fossil forms, the lateral edge of the neck is in line with the 

central trochlear depression. In extant pangolins the head 

lies much closer to the lateral edge of the astragalus, well 

lateral of the central trochlear depression (Figure 32c). 

The neck of all four fossil forms is narrow relative to the 

astragalar body, with a ratio of minimum neck width 

to maximum body width smaller than that of all extant 

manids except Manis crassicaudata (Table A5). Cryptomanis has a lower ratio than Patriomanis because of 

a distinct constriction halfway down the neck (Gaudin 

et al., 2006), and Euromanis krebsi has the lowest ratio 

of any pangolin owing to its small, cylindrical neck. In 

lateral view, the astragalar neck of Patriomanis, Necromanis, and Euromanis krebsi has an angular appearance, 

with a flat planar surface separated from the dorsal surface of the neck by a weak crest. In Cryptomanis this area 

is rounded and smooth. 

The astragalar trochlea of Patriomanis has a rough 

hourglass shape, similar to that in most pangolins (Figure 30a). The trochlea is formed by two rounded swellings oriented proximodistally and separated by a central 

depression. The lateral trochlea forms a half circle with 

its lateral border defined by a sharp ridge. The medial 

trochlea has a more rounded medial edge that is interrupted by a steep concave depression at its distomedial 

corner. This depression is also present in Cryptomanis (Gaudin et al., 2006) and Necromanis (Helbing, 

1938; Koenigswald and Martin, 1990) but is absent in 

Euromanis krebsi (Storch and Martin, 1994) and living manids (Figure 32c). The trochlea in Patriomanis is 

highly symmetrical compared to that of other pangolins, 

with a maximum proximodistal depth nearly equal for 

the medial and lateral trochleae (Table A5). A patent 

astragalar foramen is located proximal and slightly lateral to the central trochlear depression (Figure 30a; see 

also Rose and Emry, 1993). This primitive feature is also 

present in Cryptomanis (Gaudin et al., 2006) and Necromanis (Helbing, 1938; Koenigswald and Martin, 1990; 

Rose and Emry, 1993) but is rudimentary or absent 

in extant manids (Gaudin et al., 2009). An additional 

articular facet lies ventral to the medial trochlea at the 

proximal end of the astragalar body. This facet is narrow and slightly convex proximodistally and wide and 

concave transversely, with its central furrow directed 

in a proximal and medial direction (Figure 30b). This 

facet most likely served as the roof of a bursal cavity for 

the tendon of the m. flexor fibularis (Humphry, 1870; 
n u m b e r 9 8 • 4 9

in some specimens of M. pentadactyla and M. crassicaudata (Gaudin et al., 2009).

Calcaneus

Only the distal portion of the right calcaneus is preserved in the type specimen of Patriomanis (Emry, 1970). 

However, a complete right calcaneus and a partial left 

calcaneus are preserved in USNM-P 299960. A complete 

left calcaneus is preserved in USNM-P 494439. The following description is based on the two complete calcanei 

(Figure 31; Table A6).

The calcaneus of Patriomanis (Emry, 1970) is similar 

to that of Cryptomanis (Gaudin et al., 2006) and Necromanis (Helbing, 1938; Koenigswald and Martin, 1990) 

but differs significantly from that of most extant manids. 

The most obvious difference between the fossil and modern forms is the arrangement of the calcaneal facets. The 

ectal facet of Patriomanis (Emry, 1970) is oblong, with 

a proximodistal length approximately twice that of its 

transverse width (Figure 31a,b). However, it is less elongate proximodistally than the ectal facet in Cryptomanis

(Gaudin et al., 2006) and Necromanis (Helbing, 1938; 

Koenigswald and Martin, 1990). Most extant manids 

have an ectal facet that is wider, flatter, and more circular. Proximally, the ectal facet in Patriomanis is convex proximodistally, is flat transversely, and is directed 

medially and slightly dorsally. The distal portion of the 

ectal facet is flat and directed distomedially and slightly 

dorsally (Emry, 1970). This pattern is also present in 

Cryptomanis (Gaudin et al., 2006). In extant Manidae, 

the ectal facet is oriented more directly dorsad. The ectal 

facet extends well proximal to the fibular facet in Patriomanis (Emry, 1970), Cryptomanis (Gaudin et al., 2006), 

and Necromanis (Helbing, 1938; Koenigswald and Martin, 1990). In living pangolins, with the exception of M. 

pentadactyla, the proximal edge of the ectal facet lies 

even with or distal to the fibular facet. The ectal facet is 

much larger than the fibular facet, as it is in all pangolins 

except P. tricuspis, although this discrepancy is often 

slight in extant forms. The fibular facet in Patriomanis

(Emry, 1970), Cryptomanis (Gaudin et al., 2006), and 

Necromanis (Helbing, 1938; Koenigswald and Martin, 

1990) is ovate, with its long axis oriented in a distolateral direction. The shape and orientation of the facet 

vary within Patriomanis. In USNM-P 494439, the fibular facet is narrow proximodistally, convex transversely, 

and directed dorsally, whereas in USNM-P 299960 it is 

flat, nearly circular in outline, and directed dorsally and 

slightly distally (Figure 31a,b).

FIGURE 31. Calcaneus of Patriomanis americana: (a) left calcane

of USNM-P 494439 in dorsal view; (b) right calcaneus of USNM-P 

299960 in dorsal view; (c) left calcaneus of USNM-P 494439 (cast) 

in plantar view; (d) left calcaneus of USNM-P 494439 (cast) in distal view. Abbreviations: apt, anterior plantar tubercle; clct, calcaneal tuber; cuf, cuboid facet; ef, ectal facet; ff, fibular facet; perp, 

peroneal process; sup, sustentacular process and facet. Scale bars = 

1 cm. Illustrations (a), (b) by Julia Morgan Scott (JMS) for Smithsonian Institution (SI); photos (c), (d) by Tim Gaudin, composed and 

labeled by JMS for SI.
5 0 • 

The anterior plantar tubercle lies on the ventral 

surface of the calcaneus medial to the cuboid facet in 

all pangolins, including Patriomanis (Emry, 1970). 

In Patriomanis (Emry, 1970), Cryptomanis (Gaudin 

et al., 2006), and Necromanis (Helbing, 1938; Koenigswald and Martin, 1990), it also lies proximal to the 

cuboid facet (Figure 31c), whereas its position is coincident with or distal to the cuboid facet in Euromanis 

krebsi (Storch and Martin, 1994) and all living manids 

except S. gigantea (Gaudin et al., 2009). The tubercle 

The calcaneal tuber in both extant and fossil pangolins is large and expanded at its tip. In Patriomanis, 

the tuber is nearly cylindrical, but the dorsoventral 

axis remains longer than the transverse axis, whereas 

in Cryptomanis (Gaudin et al., 2006) the diameter of 

the tuber is significantly greater transversely than it 

is dorsoventrally. The tuber in living manids either 

resembles that of Patriomanis or is expanded obliquely 

along a dorsomedial to ventrolateral axis (Gaudin 

et al., 2009).

FIGURE 32. Right tibia and fibula and left pes of Phataginus tricuspis (CM 16206): (a) tibia and fibula in anterior view; (b) tibia a

fibula in posterior view; (c) pes in dorsal view (unguals reconstructed on the basis of other specimens). Abbreviations: I–V, digits I–V; ast,

astragalus; clc, calcaneus; clcf, fibular calcaneal facet; cu, cuboid; cyf, cyamelle facet; ecc, ectocuneiform; enc, entocuneiform; fib, fibula;

gmtp, groove for m. tibialis posterior tendon; ip, intermediate phalanx; lco, lateral condyle; lie, lateral intercondylar eminence; lplm, lateral process of lateral malleolus of fibula; mco, medial condyle; mec, mesocuneiform; mie, medial intercondylar eminence; mma, medial 

malleolus of tibia; mt, metatarsal; nav, navicular; pp, proximal phalanx; ppf, posterior process of fibula; prh, prehallux; tib, tibia; ttb, tibial 

tuberosity; up, ungual phalanx. Scale bars = 1 cm. Illustrations by Julia Morgan Scott for Smithsonian Institution; parts (a), (b) modified 

from Gaudin et al. (2006), (c) modified from Gaudin et al. (2009).
n u m b e r 9 8 • 5 1

large sustentacular process is present in all African pangolins except P. tricuspis, which bears a weak process, 

whereas in Asian pangolins the process is absent (with 

the exception of FMNH 57338, a juvenile Manis crassicaudata specimen). The sustentacular facet in Manidae is 

also variable. It may be distally elongated, transversely 

broadened, ovate, triangular, or entirely absent (Gaudin 

et al., 2009).

Fibula

As noted by Emry (1970), only the proximal portion of the right fibula is preserved in the type specimen. 

A distal portion of the left fibula is preserved in USNMP 531557. An additional proximal right fibula and distal 

left fibula, although damaged, are available from USNMP 494439. However, USNM-P 299960 preserves nearly 

complete right and left fibulae (Figure 30), although both 

are damaged at the proximal and distal ends. 

The fibula in Patriomanis is significantly shorter 

than the tibia (Figure 33). In fact, the ratio of fibula to 

tibia length is lower in Patriomanis than in Cryptomanis

(Gaudin et al., 2006) and all living manids (Table A7). 

Emry (1970) describes the fibular head of Patriomanis as flattened laterally and elongated anteroposteriorly. 

In Cryptomanis, the head is expanded transversely and 

does not extend as far anteriorly or posteriorly (Gaudin 

et al., 2006). Among extant manids, the fibular head is 

somewhat variable in shape, but most possess a posterior 

process that extends farther posteriorly from the shaft and 

an anterior process that extends less far anteriorly than is 

the case in Patriomanis (Gaudin et al., 2009). Emry (1970) 

identifies a medially and dorsally facing proximal tibial 

facet that rests on an anterior projection of the head, displaced well anterior to the shaft. In Cryptomanis (Gaudin 

et al., 2006), the proximal tibial facet is more in line with 

the shaft, rather than being displaced anteriorly, and faces 

more posteriorly. In both the type and USNM-P 299960, 

there appears to be a second medially directed facet on the 

posterior portion of the head. This is likely a sesamoid facet 

for articulation with the cyamelle, similar to that in Cryptomanis (Gaudin et al., 2006) and some extant manids. 

This cyamelle facet is not described in Emry (1970), and 

Gaudin et al. (2006) assert on this basis that it is absent, 

but more thorough examination has revealed its presence 

(Figure 33a). The posterior process of the proximal fibula 

extends only slightly posterior to the sesamoid facet. This 

process consists of a lateral and a medial tubercle with a 

midline ridge in between that extends farther distally than 

in Cryptomanis (Gaudin et al., 2006) before merging with 

is positioned at the same level as the distal edge of the 

sustentacular facet in USNM-P 494439 and is slightly 

distal to the sustentacular facet in USNM-P 299960. 

The tubercle is freestanding in all five fossil pangolin 

genera, whereas it is sessile in living manids. In Patriomanis (Emry, 1970), the tubercle is less prominent than 

in the other four fossil forms. A distinct groove is evident medial to the tubercle, on the ventral surface of the 

sustentacular process, allowing for the passage of the 

m. flexor fibularis. This groove is present in patriomanids and Necromanis but is absent in Euromanis krebsi

(Storch and Martin, 1994) and the extant Manidae. 

The peroneal process in Patriomanis (Emry, 1970) is a 

knoblike eminence projecting from a dorsomedial ridge 

midway between the ectal and cuboid facets (Figure 

31a,c). This process is highly reduced relative to that 

in Cryptomanis (Gaudin et al., 2006) and most extant 

Manidae but not as reduced as in Necromanis (Helbing, 

1938; Koenigswald and Martin, 1990), in which it is 

nearly absent.

The cuboid facet in Patriomanis (Emry, 1970) forms 

an irregular circle, indented along its medial margin by 

a depression that separates it from the anterior plantar 

tubercle (Figure 31d). The facet is slightly concave transversely, is flat dorsoventrally, and is directed distomedially and somewhat ventrally. The facet is similar in shape 

to that of Cryptomanis but is less symmetrical than the 

circular facet of Euromanis krebsi (Storch and Martin, 

1994). Modern African pangolins possess a cuboid facet 

similar to that of Patriomanis (Emry, 1970), although 

in Smutsia it tends to be more elongated transversely, 

whereas Asian pangolins possess a transversely ovate 

facet that is strongly concave transversely and directed 

more distally. The sustentacular facet in Patriomanis

(Emry, 1970) is circular, is directed dorsally and slightly 

distomedially, and lies opposite the distal portion of the 

ectal and fibular facets (Figure 31a,b). It covers almost 

the entire dorsal surface of the sustentacular process, 

similar to the condition in Euromanis krebsi (Storch and 

Martin, 1994). The sustentacular facet is similar in shape 

to that of Necromanis (Helbing, 1938; Koenigswald and 

Martin, 1990) and Euromanis krebsi (Storch and Martin, 1994) but is smaller and less elongated distally than 

the sustentacular facet of Cryptomanis (Gaudin et al., 

2006). In Necromanis (Helbing, 1938; Koenigswald and 

Martin, 1990) and, to a lesser degree, in Cryptomanis

(Gaudin et al., 2006), the sustentacular process extends 

farther medially than the sustentacular facet. Among 

extant manids, the sustentacular process is usually positioned more distally but is variable in morphology. A 
5 2 • 

shaft is compressed anteroposteriorly and bowed laterally. 

In modern manids, the shape is variable—it is cylindrical 

in some taxa, whereas in others the shaft is compressed 

transversely; additionally, the shaft may be straight or 

bowed medially or laterally (Gaudin et al., 2009).

A lateral process is present on the lateral malleolus 

of all pholidotans (Figure 32a,b; Gaudin et al., 2006). In 

Patriomanis, the lateral process is rectangular in anterior 

view (Figure 33b). It is thin with a proximodistal length 

greater than its transverse width. In extant Manidae, the 

lateral process is usually even more elongated proximodistally. The lateral malleolus in Patriomanis is V shaped in 

anterior view and compressed along an anteromedial to 

posterolateral axis. In Cryptomanis (Gaudin et al., 2006), 

the lateral malleolus is less compressed transversely with 

a squared terminus. The distal tibial facet in Patriomanis

is triangular and flat. The facet faces more anteriorly in 

Patriomanis and Cryptomanis (Gaudin et al., 2006) than 

the medial edge of the fibula. In Cryptomanis (Gaudin 

et al., 2006), a similar morphology is present, although 

in this genus the lateral tubercle is larger and the medial 

tubercle is less pronounced than in Patriomanis, and there 

is a deep pit in between the lateral tubercle and the median 

ridge. In extant Manidae, a single large posterior process 

is present that may extend distally well down the shaft 

of the fibula (Figure 32b; Gaudin et al., 2009). As in all 

pholidotans (Gaudin et al., 2006), an eminence lies on the 

anterolateral surface of the proximal fibula. This eminence 

extends well forward of the tibial facet in Patriomanis

and Cryptomanis (Gaudin et al., 2006) but not in extant 

manids (Gaudin et al., 2009). 

On the basis of its proximal end, Emry (1970) 

describes the fibular shaft in Patriomanis as thin, straight, 

and cylindrical. The complete fibulae preserved in USNMP 299960 confirm this description for the entire length of 

the fibular shaft. In Cryptomanis (Gaudin et al., 2006), the 

FIGURE 33. Right tibia and fibula of Patriomanis americ

(USNM-P 299960): (a) in posterior view; (b) anterior view. 

Abbreviations: adp, anterior distal tibial process; astf, astragalar facet; clcf, fibular calcaneal facet; cnc, cnemial crest; cyf, 

cyamelle facet; fib, fibula; gmtp, groove for m. tibialis posterior tendon; lco, lateral condyle; lie, lateral intercondylar eminence; lplm, lateral process of lateral malleolus of fibula; mco, 

medial condyle; mie, medial intercondylar eminence; mma, 

medial malleolus of tibia; popf, popliteal fossa; ppf, posterior 

process of fibula; tib, tibia; ttb, tibial tuberosity. Scale bar = 

1 cm. Drawn by Larry Isham, digitized and labeled by Julia 

Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 5 3

Emry (1970) notes that the tibia of Patriomanis is 

expanded transversely at the distal and proximal ends like 

that of other pangolins (Figure 33). Its shape particularly 

resembles that of Cryptomanis (Gaudin et al., 2006). The 

most obvious difference between these two fossil forms 

and extant pangolins is the presence of a robust cnemial 

crest on the proximal portion of the tibial shaft (Figure 

33b). As stated in the description of the type specimen, 

the cnemial crest in Patriomanis extends farther down the 

shaft than in extant pangolins (Emry, 1970). In fact, the 

crest extends approximately two-thirds down the length 

of the tibial shaft, where it forms a distinct, rounded angle 

before rejoining the shaft. Distal to this, the cnemial crest 

blends imperceptibly into a weak, rounded ridge that is 

in line with the anterior edge of the medial malleolus. A 

sharp cnemial crest is also present in Cryptomanis (Gaudin 

et al., 2006), but in this genus it terminates halfway down 

the tibial shaft in a prominent tubercle. In extant Manidae (except M. pentadactyla), the cnemial crest is greatly 

reduced, weak, and rounded if present at all (Figure 32a; 

Gaudin et al., 2009). Emry (1970) describes a prominent 

fossa lateral to the cnemial crest in Patriomanis. This 

fossa is also present in Cryptomanis (Gaudin et al., 2006), 

although it is less deep. This fossa is absent in most living 

manids, although a weak fossa is present in M. pentadactyla (Gaudin et al., 2009).

Additional surface markings are present on the tibial shaft that distinguish Patriomanis (Emry, 1970) and 

Cryptomanis (Gaudin et al., 2006) from living pangolins. 

Emry (1970) describes and illustrates a well-developed 

popliteal depression in Patriomanis immediately below 

the medial condyle on the posterior surface of the proximal tibia. The fossa is deep, narrow, and elongated 

in extant manids, in which it faces more medially (Gaudin 

et al., 2009). 

As noted in Gaudin et al. (2006), the astragalar and 

calcaneal facets of the fibula in all pangolins lie anterior 

and distal to the pit for the astragalo-fibular ligament. The 

astragalar facet in Patriomanis has a flat surface. Although 

angular, it is roughly C shaped, with a distal moiety that 

is less vertical than in Cryptomanis (Gaudin et al., 2006), 

which has an L-shaped astragalar facet that is markedly 

concave at its proximal extremity. The astragalar facet in 

most living pangolins is C shaped but less angular than 

in Patriomanis. The calcaneal facet is ovate and narrow 

transversely and is directed posteriorly and slightly distally 

and laterally. It is similar in shape to that in other pangolins except those in the genus Manis, in which the calcaneal facet is compressed proximodistally. Along its medial 

edge, the distal half of the calcaneal facet abuts the distal 

portion of the astragalar facet. From there it extends proximally nearly as far as the astragalar facet, much like the 

condition in Phataginus (Gaudin et al., 2009). In Cryptomanis (Gaudin et al., 2006), S. temminckii, and Manis, 

the proximal edge of the calcaneal facet is well distal to 

that of the astragalar facet. As noted by Gaudin et al. 

(2006, 2009), the calcaneus does not contact the fibula in 

S. gigantea.

Tibia

Most of the right and left tibiae are preserved in the 

type specimen and were described by Emry (1970). The 

right and left tibiae are also available from USNM-P 

494439, although both are damaged. A complete pair of 

tibiae is available from USNM-P 299960 (Figures 33, 34).

FIGURE 34. Right tibia of Patriomanis america

(USNM-P 299960): (a) in proximal view; (b) in distal view. Abbreviations: adp, anterior distal tibial 

process; cyf, cyamelle facet; gmtp, groove for m. 

tibialis posterior tendon; lco, lateral condyle; lie, 

lateral intercondylar eminence; ltrf, facet for lateral 

trochlea of astragalus; mco, medial condyle; mie, 

medial intercondylar eminence; mma, medial malleolus of tibia; mml, pit for attachment of medial 

meniscal ligament; mtrf, facet for medial trochlea of 

astragalus; pcl, pit for attachment of posterior cruciate ligament. Scale bar = 1 cm. Illustrations by Julia 

Morgan Scott for Smithsonian Institution. 
5 4 • 

Emry, 1970) and Cryptomanis (Gaudin et al., 2006). The 

medial intercondylar eminence in Patriomanis (Emry, 

1970) is elongated anteroposteriorly, as in all pangolins, 

but is narrower transversely than in other forms. It is continuous anteriorly with the tibial tuberosity. The lateral 

intercondylar eminence is well developed but smaller than 

the medial intercondylar eminence (Figures 33, 34a), also 

a characteristic of all pangolins. In Patriomanis (Emry, 

1970), the lateral intercondylar eminence is smooth and 

rounded and lies at the posteromedial extent of the lateral 

tibial condyle.

As related by Emry (1970), the proximal tibia bears 

a facet for the cyamelle (Figures 33a, 34a), a sesamoid 

bone that forms within the tendon of the mm. popliteus and peroneus longus (Jouffroy, 1966). This sesamoid 

facet projects posteriorly and slightly laterally from the 

posterolateral edge of the lateral tibial condyle. It is convex proximodistally and transversely and is elongated 

in a dorsolateral direction, whereas in Cryptomanis

(Gaudin et al., 2006) the facet is nearly flat and deeper 

proximodistally. The orientation of this facet is variable 

in extant Manidae. The facet for the cyamelle is small 

compared to the femoral articular surfaces. The proximal 

fibular facet (Emry, 1970) is ovate and elongated anteroposteriorly. It is directed laterally and somewhat distally 

and anteriorly, similar to the condition of extant Manis

and Cryptomanis (Gaudin et al., 2006, 2009) and differing from the proximal fibular facet in extant African 

pangolins, which is not visible in anterior view (Gaudin 

et al., 2009). 

The distal end of the tibia in Patriomanis is triangular 

(Figure 34b; Emry, 1970) but much less expanded transversely than in other pangolins. In fact, Patriomanis has a 

ratio of distal maximum transverse width to anteroposterior depth less than that of all other pangolins (Table A7). 

The distal end of the tibia has undergone approximately 

50° of torsion (clockwise on right tibia, counterclockwise 

on left tibia) relative to a transverse axis across the tibial 

head. Among other pangolins, only Cryptomanis (Gaudin 

et al., 2006), Smutsia gigantea, and Manis javanica, each 

with approximately 45° of torsion, have undergone such 

a large degree of twisting of the distal tibia (Gaudin et al., 

2009).

The medial malleolus of Patriomanis (Emry, 1970) is 

large and occupies a larger portion of the distal tibial surface than in extant manids. However, Cryptomanis possesses an even more robust malleolus (Gaudin et al., 2006). 

The posteromedial surface of the malleolus in Patriomanis (Emry, 1970) and Cryptomanis (Gaudin et al., 2006) 

is impressed by a broad groove for the passage of the 

distally for almost one-third the length of the shaft. It is 

bounded laterally by a sharp crest that marks the lateral 

edge of the shaft, descending from the lower margin of 

the proximal fibular facet to the midshaft region, where 

it nearly contacts a similar crest rising proximally from 

the distal fibular facet. A popliteal fossa is also present in 

Cryptomanis (Gaudin et al., 2006) and M. pentadactyla

but is absent in other living manids. The posterior surface 

of the distal tibial shaft bears a broad, shallow concavity. 

In Cryptomanis (Gaudin et al., 2006), the concave posterior distal surface is interrupted by a rounded median 

ridge, creating two separate concavities. In most extant 

manids, the posterior distal tibial surface is convex (Figure 32b). The shaft itself is curved in medial view, such 

that the proximal and distal ends of the tibia lie posterior 

to the middle portion of the shaft. A similar condition 

prevails in Cryptomanis, whereas the tibial shaft is nearly 

straight in living pangolins.

As noted by Gaudin et al. (2006), both Patriomanis 

and Cryptomanis possess a robust tibial tuberosity that 

forms an elevated U-shaped eminence at its distal extent, 

where it abuts the proximal cnemial crest (Figure 33b). 

The tibial tuberosity in extant manids (except M. crassicaudata) is horizontally ovate or rectangular, positioned 

on the proximal portion of the anterior tibia but not 

extending distally (Figure 32a). Emry (1970:481) describes 

the medial tibial condyle in Patriomanis as “slightly convex” (anteroposteriorly), whereas that of Cryptomanis is 

flat (Gaudin et al., 2006). The medial condyle is concave 

transversely by virtue of a lappet forming much of the 

medial surface of the well-developed medial intercondylar eminence (Figures 33a, 34a). This lappet is bounded 

anteriorly by a discrete pit that has been interpreted by 

Gaudin et al. (2006) as a site for the attachment of the 

cranial tibial ligament of the medial meniscus (Figure 34a). 

The medial lappet is bounded posteriorly by a fossa for the 

posterior cruciate ligament. The fossa for the anterior cruciate 

ligament is located on the opposite side of the medial intercondylar eminence anterior and medial to the lateral condyle. All three of these fossae are present in all pangolins. 

As Emry (1970) also points out, the lateral tibial condyle 

is more convex than the medial condyle. The medial condyle is large and ovate, with its long axis oriented anteroposteriorly, whereas the lateral condyle is smaller and has 

an irregular outline, with its maximum width and depth 

roughly equivalent (Figure 34a). In contrast, the lateral 

condyle of Cryptomanis is widened transversely (Gaudin 

et al., 2006). Although the lateral condyle is elevated relative to the medial condyle in all pangolins, this characteristic is more pronounced in Patriomanis (Figure 33; 
n u m b e r 9 8 • 5 5

crassicaudata) but less so than the remarkably robust 

patella of Cryptomanis (Gaudin et al., 2006; Table A8). 

The patella in Patriomanis is sharply pointed distally, 

with angular lateral margins and a rugose anterior surface. Both the distal point and the surface rugosities are 

more strongly developed in Cryptomanis (Gaudin et al., 

2006). The patella in Euromanis krebsi (Storch and 

Martin, 1994) and extant manids (Gaudin et al., 2009) 

is more ovate or rectangular and has a smooth anterior 

surface with rounded edges. The posterior surface of 

the patella in Patriomanis bears an articular facet that is 

divided nearly in half by a smooth, elevated ridge that is 

absent in all other pangolins (Figure 35). The rectangular 

articular surface is nearly flat proximodistally, whereas 

other pangolins possess a more concave articular surface. 

The medial facet is weakly concave transversely, whereas 

the lateral facet is almost flat. As in Cryptomanis (Gaudin 

et al., 2006) and in contrast to living pangolins, the medial 

portion of the patellar facet extends farther distally than 

the lateral portion.

Femur

Although no femur is preserved in the type specimen (Emry, 1970), right and left femora are preserved in 

USNM-P 494439, the left still articulated in the acetabulum of the pelvis. Complete left and right femora (Figures 

36, 37) are also preserved in USNM-P 299960 and have 

been prepared free of the matrix; thus, the description is 

largely based on these specimens.

m. tibialis posterior tendon (Figures 33a, 34b; inferred 

by analogy to Smutsia gigantea and Manis pentadactyla; 

see Humphry, 1870; Jouffroy, 1966; Gaudin et al., 2006). 

In living pangolins, this groove is more pronounced, in 

some taxa virtually enclosed in a bony tunnel, and may 

be accompanied by a second weaker and more anterior 

groove, presumably for the tendon of the m. flexor tibialis (Figure 32b; Jouffroy, 1966; Gaudin et al., 2009). The 

distal tibia bears two articular surfaces for the astragalar 

trochlea (Figure 34b). These surfaces are separated by a 

smooth ridge that is less pronounced than in Cryptomanis (Gaudin et al., 2006). The anterior distal tibial process 

that lies at the anterior end of this ridge (Figures 33b, 34b) 

is also less pronounced in Patriomanis than in Cryptomanis (Gaudin et al., 2006) but is larger than that in extant 

manids. The lateral trochlear facet is slightly concave 

anteroposteriorly and flat transversely, whereas this facet 

is concave transversely in Cryptomanis (Gaudin et al., 

2006) and living pangolins. The concave medial trochlear 

facet is nearly centered on the distal end of the tibia and 

is slightly larger than the lateral facet. Patriomanis (Emry, 

1970) is the only pangolin in which the anteroposterior 

depth of the medial trochlear facet exceeds that of the lateral facet (Figure 34b; Gaudin et al., 2009). The medial 

facet has a small anterior indentation that is present in 

Cryptomanis but absent in all living manids except Manis 

pentadactyla. 

The distal fibular facet in Patriomanis (Emry, 1970) is 

a slender, transversely elongated surface that faces laterally and slightly posteriorly, similar to that in Cryptomanis (Gaudin et al., 2006). In both fossil species, the facet 

faces more posteriorly than in extant manids (Gaudin 

et al., 2009). The facet is not visible in distal view in either 

fossil form, whereas it can be seen in distal view in living 

pangolins (Gaudin et al., 2009). 

Patella

Although no patellae are preserved in the type specimen, a left patella is available from USNM-P 494439. 

Left and right patellae are also available from USNM-P 

299960 (Figure 35). 

The patella in Patriomanis is roughly diamond shaped 

in anterior view. Like that of Cryptomanis (Gaudin 

et al., 2006), its height and width are roughly equivalent, whereas in extant manids and Euromanis krebsi

(Storch and Martin, 1994), the patella is distinctly elongated proximodistally (Table A8). The patella is deeper 

anteroposteriorly than that of Euromanis krebsi (Storch 

and Martin, 1994) and most living manids (except Manis 

FIGURE 35. Right patella of Patriomanis americana (USNM

299960) in posterior view. Abbreviations: lff, lateral femoral facet;

mff, medial femoral facet. Scale bar = 1 cm. Illustration by Julia 

Morgan Scott for Smithsonian Institution.
5 6 • 

compressed anteroposteriorly, similar to the condition in 

Cryptomanis (Gaudin et al., 2006) and Euromanis krebsi 

(Storch and Martin, 1994). 

The greater trochanter in Patriomanis is directed 

proximolaterally and is relatively narrow transversely 

compared to other pangolins (Table A8). Indeed, Patriomanis and Cryptomanis (Gaudin et al., 2006) differ 

from modern pangolins in having a greater trochanter 

with an anteroposterior depth greater than its transverse width. The greater trochanter in Patriomanis is 

The femoral head in Patriomanis is nearly hemispherical (Figure 36) but is slightly compressed along 

an anteromedial to posterolateral axis (Figure 37a). As 

in all pangolins, the head extends farther proximally 

than the greater trochanter. A fovea lies on the posteromedial surface of the head as in Cryptomanis (Gaudin 

et al., 2006) and Euromanis krebsi (Storch and Martin, 

1994), although it is more medially positioned in the 

latter. A fovea is absent in living manids (Figure 38b,c; 

Gaudin et al., 2009). The femoral neck in Patriomanis is 

FIGURE 36. Right femur of Patriomanis americana (USNM-P 299960): (a) in posterior vie

(b) in anterior view. Abbreviations: fvc, fovea capitis; gt, greater trochanter; h, head; icfo, intercondylar fossa; itc, intertrochanteric crest; lco, lateral condyle; lec, lateral epicondyle; lt, lesser trochanter; mco, medial condyle; mec, medial epicondyle; pg, patellar groove; trf, trochanteric fossa; tt, 

third trochanter. Scale bar = 1 cm. Illustrations drawn by Larry Isham, digitized and labeled by Julia 

Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 5 7

farther laterally. By considering the position of the third 

trochanter, pholidotans can be arranged in a series: Eomanis and Euromanis (Storch, 1978; Storch and Martin, 

1994) have the most proximally situated third trochanter, 

positioned well proximal to the midshaft. In Cryptomanis

(Gaudin et al., 2006), the third trochanter is just proximal to midshaft, whereas in Patriomanis it is just distal to 

midshaft. In Necromanis (Koenigswald and Martin, 1990; 

Koenigswald, 1999), it is well distal to midshaft. Finally, 

extant manids bear a third trochanter that is reduced in 

size and located at the distal end of the femur (Table A8; 

Emry, 1970; Gaudin et al., 2009). A thin, elevated ridge 

extends proximally from the third trochanter in Patriomanis and forms the lateral edge of the femur, ending just 

before it reaches the level of the lesser trochanter. A similar 

ridge extends distally from the third trochanter, terminating at the lateral epicondyle. 

The femoral shaft in Patriomanis is more compressed 

anteroposteriorly than that of extant pangolins with the 

exception of Smutsia gigantea (Table A8). The femoral 

shaft is also somewhat compressed in Necromanis (Koenigswald, 1969, 1999; Koenigswald and Martin, 1990) and 

Euromanis krebsi (Storch and Martin, 1994). The femoral shaft in Cryptomanis, however, is significantly broader 

and more flattened than other pangolins (Gaudin et al., 

2006). The femoral shaft in Patriomanis has a strongly 

convex anterior face, a flat posterior face, with a sharp 

lateral edge and a rounded medial margin connecting the 

medial epicondyle and the lesser trochanter. Patriomanis

lacks any evidence of a supracondylar tuberosity like that 

which is prominently developed in Cryptomanis (Gaudin 

et al., 2006). Cryptomanis (Gaudin et al., 2006) also has 

an elevated ridge crossing the posterior distal surface of 

the femur that is absent in Patriomanis.

connected to the femoral head by a narrow ridge (Figure 

37a). The comparable ridge is much broader in Cryptomanis (Gaudin et al., 2006) and Euromanis krebsi 

(Storch and Martin, 1994). In the latter, an additional 

ridge extends distally from the greater trochanter on 

the anterior surface of the femur. The posterior surface 

of the proximal femur is impressed by a deep, ovate, 

proximodistally elongated trochanteric fossa (Figure 

36a). This fossa is also present in Necromanis (Koenigswald, 1969, 1999; Koenigswald and Martin, 1990) and 

Cryptomanis (Gaudin et al., 2006) but is much reduced 

in the latter. Living pangolins lack a trochanteric fossa 

(Figure 38b,c; Gaudin et al., 2009). In Patriomanis, the 

trochanteric fossa is bordered posteriorly by a robust 

intertrochanteric ridge that extends from the greater trochanter to the lesser trochanter, although it is much less 

pronounced near the latter. This ridge is less developed 

in Cryptomanis (Gaudin et al., 2006) and is absent in 

extant manids (Gaudin et al., 2009). The lesser trochanter in Patriomanis projects posteromedially and lies posterior to the femoral neck (Figures 36, 37a). It is similar 

in morphology to that of Necromanis (Koenigswald, 

1969, 1999; Koenigswald and Martin, 1990). In Cryptomanis (Gaudin et al., 2006), the lesser trochanter is 

more pronounced and is connected to the femoral head 

by a blunt ridge that is not present in Patriomanis. In 

extant pangolins the lesser trochanter tends to be more 

medially oriented (Figure 38b,c; Gaudin et al., 2009).

The femur in Patriomanis bears a robust, freestanding third trochanter that is nearly semicircular in outline 

with a rugose tubercle on its posterior surface (Figure 36). 

A similar morphology is present in Cryptomanis (Gaudin 

et al., 2006), although in this taxon the third trochanter 

is more robust, is broader proximodistally, and extends 

FIGURE 37. Left femur of Patriomanis americana (USNM-P 299960): (a) in proximal vie

(b) in distal view. Abbreviations: fvc, fovea capitis; gt, greater trochanter; h, head; icfo, intercondylar fossa; itc, intertrochanteric crest; lco, 

lateral condyle; lec, lateral epicondyle; lt, lesser 

trochanter; mco, medial condyle; mec, medial 

epicondyle; pg, patellar groove; trf, trochanteric 

fossa. Scale bars = 1 cm. Illustrations by Julia 

Morgan Scott for Smithsonian Institution
5 8 • 

condyle is much wider than the lateral condyle (Table

A8). In posterior view, the medial condyle in Patriomanis

appears ovate, compressed transversely, and convex transversely and proximodistally. The lateral condyle is narrow 

transversely and rectangular in outline. It is convex proximodistally but flat transversely (Figure 36a). In Cryptomanis (Gaudin et al., 2006), the medial and lateral condyles 

have nearly the same shape but are wider transversely. 

The patellar articular surface in Patriomanis is concave transversely, tapers slightly at its proximal end, and 

is inclined mediad so that it points toward the femoral 

head (Figures 36b, 37b). The medial edge of the facet is 

in line with the lateral edge of the femoral head and is 

slightly more elevated than the lateral edge. In Cryptomanis (Gaudin et al., 2006) and African tree pangolins (Figure 38b), the patellar surface is wider and less concave. In 

Manis, the patellar surface is almost flat. The intercondylar fossa in Patriomanis is deep and directed slightly laterally and has a uniform width throughout its extent (Figure 

37b). In Cryptomanis (Gaudin et al., 2006), the fossa is 

broader and widens anteriorly.

Pelvis and Sacrum

The type specimen preserves only two small fragments 

of the pelvis, including the pubic symphysis and a fragment of the right acetabulum and pubis, but does not preserve the sacrum (Emry, 1970). However, a partial pelvis 

and sacrum are preserved in USNM-P 494439, although 

these have not been prepared free of the matrix. A complete pelvis and sacrum are preserved in USNM-P 299960 

(Figure 39) and have been illustrated by Rose and Emry 

(1993).

The most obvious difference between the pelvis of 

Patriomanis and living manids is the presence of a large 

gluteal fossa. The ilium in Patriomanis is expanded medially and dorsally to form a large, deeply concave gluteal 

fossa that covers most of the lateral surface of the ilium. 

Cryptomanis (Gaudin et al., 2006) also possesses a large 

gluteal fossa. A weaker gluteal fossa is apparently present in Necromanis (Koenigswald and Martin, 1990). In 

Euromanis krebsi (Storch and Martin, 1994) and living 

pangolins (Figure 38a; Gaudin et al., 2009), however, the 

dorsolateral surface of the ilium is either flat or slightly 

convex and does not form an expanded surface.

The gluteal fossa is bordered laterally by a sharp, 

narrow lateral iliac crest (Figure 39b). The anterior end 

of the lateral iliac crest is confluent with a prominent, 

laterally projecting cranial ventral iliac spine. The posterior termination of the lateral iliac crest lies at the ventral 

The distal end of the femur in Patriomanis is deeper 

than in other pangolins, with a maximum depth nearly 

equivalent to its maximum width (Figure 37b; Table A8). 

As in Cryptomanis (Gaudin et al., 2006) and Smutsia 

gigantea but unlike in other modern pangolins, the medial 

FIGURE 38. Pelvis, sacrum, and left femur of Phataginus tricus

(CM 16206): (a) pelvis and sacrum in left lateral view; (b) femur 

in anterior view; (b) femur in posterior view. Abbreviations: acet, 

acetabulum; fsp, femoral spine; gt, greater trochanter; h, head; icfo, 

intercondylar fossa; il, ilium; ipe, iliopubic eminence; isch, ischium; 

issp, ischial spine; istb, ischial tuberosity; lco, lateral condyle; lec, 

lateral epicondyle; lilc, lateral iliac crest; lt, lesser trochanter; mco, 

medial condyle; mec, medial epicondyle; obf, obturator foramen; pg, 

patellar groove; pub, pubis; spn f, spinal nerve foramen; sv2, second

sacral vertebra; tf, trochanteric fossa; tt, third trochanter. Scale bar = 

1 cm. Illustrations by Julia Morgan Scott for Smithsonian Institution; (a) modified from Gaudin et al. (2009), (b) and (c) modified 

from Gaudin et al. (2006).
n u m b e r 9 8 • 5 9

rounded and often poorly demarcated. The gluteal fossa 

in Patriomanis is bordered dorsally and medially by an 

elevated dorsal iliac crest (= medial dorsal flange of ilium 

in Gaudin et al., 2009) that is slightly folded over laterally 

along its upper edge. A weak cranial dorsal iliac spine lies 

just medial and ventral to the dorsal iliac crest near its 

edge of the femoral spine. In Euromanis krebsi (Storch 

and Martin, 1994) the lateral iliac crest is developed to 

a similar extent, although it is less sharp than that of 

Patriomanis, and it appears to extend somewhat farther 

posteriorly, reaching the anterior margin of the acetabular eminence. The lateral iliac crest of extant manids is 

FIGURE 39. Right pelvis of Patriomanis americana (USNM-P 299960): (a) pelvis and sacral vertebrae 

lateral view; (b) pelvis in ventrolateral view. Abbreviations: acet, acetabulum; cadis, caudal dorsal iliac spine; 

cvis, cranial ventral iliac spine; dic, dorsal iliac crest (= medial dorsal flange of ilium in Gaudin et al., 2009); 

fsp, femoral spine; glf, gluteal fossa; il, ilium; ipe, iliopubic eminence; isch, ischium; issp, ischial spine; istb, 

ischial tuberosity; lilc, lateral iliac crest; mp, metapophysis of first sacral vertebra; obf, obturator foramen; 

pub, pubis; sv, sacral vertebra; tp, transverse processes of third and fourth sacral vertebrae. Scale bar = 1 cm. 

Illustration (a) by Julia Morgan Scott (JMS) for Smithsonian Institution (SI); (b) drawn by Larry Isham, 

digitized and labeled by JMS for SI.
6 0 • 

posteriorly and ventrally (Figure 39). The posterodorsal limb of the lunate surface is undercut ventrally by a 

deep fossa that separates it from the lateral surface of the 

ischium. In both respects it closely resembles the condition 

in Euromanis krebsi (Storch and Martin, 1994). In living 

pangolins, the two arms of the lunate surface approach one 

another more closely and may even contact one another, 

while the opening between them rotates either posteriorly 

(Manis, Smutsia) or ventrally (Phataginus; Figure 38a; 

Gaudin et al., 2009). 

The ischium in Patriomanis is flattened transversely, 

although, as in other pangolins, its lateral surface is 

marked by a rounded ridge connecting the posterodorsal 

arm of the acetabular lunate surface to the ischial tuberosity. The element appears deeper dorsoventrally than that 

of other pangolins because of the position and orientation 

of the ischial spine. The spine is erect, pointing dorsally. 

It is situated along the dorsal margin of the ischium, well 

anterior to the ischial tuberosity, lying over the anterior 

portion of the obturator foramen (Figure 39). It is connected by a crest to the ischial tuberosity and is roughly 

in line with the body of the ischium and the tuberosity, 

the latter being only weakly flared laterally. Gaudin et al. 

(2006) described a similar condition in Cryptomanis and 

noted that in modern pangolins the ischial spine is displaced posteriorly and inflected medially, whereas the 

ischial tuberosity is more strongly flared laterally (Figure 

38a). Euromanis krebsi (Storch and Martin, 1994) shows 

a mixture of these conditions, with an anteriorly situated, 

medially inflected ischial spine and an ischial tuberosity 

that is not flared laterally. 

As described by Emry (1970), the pubis extends ventrally and posteriorly from the iliopubic eminence toward 

the pubic symphysis. Its shaft is flattened transversely and 

flares strongly toward its posterior end in both a dorsal and ventral direction (Figure 39b). This is similar to 

the condition in other pangolins (Gaudin et al., 2009), 

although in Euromanis krebsi (Storch and Martin, 1994) 

the pubis is not strongly flattened, and in neither Euromanis nor Eomanis (Storch, 1978; Storch and Martin, 1994) 

is the pubis so strongly flared distally, whereas in modern 

pangolins the pubis is more strongly downturned ventrally 

as it approaches the symphysis (Figure 38a). All pangolins possess a short pubic symphysis. In Patriomanis, the 

pubic symphysis is extremely short and bears a triangular 

ossification on its posterior edge (Emry, 1970). In ventral 

view, the anterior edge of the pubic symphysis is narrow 

and V shaped, and the posterior edge takes the form of a 

broad, shallow U. Eomanis (Storch, 1978) displays a similar morphology. In extant manids, however, both edges 

anterior end (not visible in Figure 39 because it lies on the 

medial surface of the ilium). Posteriorly, the dorsal iliac 

crest terminates at an elevated caudal dorsal iliac spine 

that projects slightly mediad. Euromanis krebsi (Storch 

and Martin, 1994) exhibits a weak caudal dorsal iliac 

spine but lacks a distinct dorsal iliac crest or cranial dorsal iliac spine. Living manids lack all of these structures 

(Gaudin et al., 2009). 

The cranial surface of the ilium in Patriomanis bears 

a triangular iliac crest that reflects the roughly triangular shape of the body of the ilium. The ventrolateral iliac 

surface is flat, with sharp edges separating it from the 

dorsolateral and medial surfaces. The auricular surface is 

flat, oval, and rugose and covers the posterior half of the 

medial face of the ilium. This articular surface shares a 

broad border with the ventral edge of the ilium, whereas 

it contacts only the dorsal edge of the ilium at the caudal 

dorsal iliac spine and the sloping dorsal ridge behind it. 

The auricular surface terminates posteriorly just anterior 

to the leading edge of the acetabulum, resembling the condition in Cryptomanis (Gaudin et al., 2006) and Euromanis krebsi (Storch and Martin, 1994). In Phataginus, the 

sacroiliac attachment extends posteriorly to the center of 

the acetabular fossa (Figure 38a), whereas in Manis and 

Smutsia it extends even farther posteriorly (Gaudin et al., 

2009). The body of the ilium runs nearly parallel to the 

sacrum (Figure 39a), the angle between the two approximating 10°, whereas in extant pangolins the dorsal margin of the pelvis is steeply inclined in an anterodorsal to 

posteroventral orientation (Figure 38a), with the angle 

between the pelvis and sacrum at least twice as large as 

that of Patriomanis (Gaudin et al., 2009).

The ventral junction of the ilium and pubis is marked 

by a robust, anteroposteriorly broad iliopubic eminence 

(Figure 39b). This process is well developed in all pangolins. The femoral spine is rounded and rather indistinct 

and lies on the acetabular eminence, nearly in line vertically with the iliopubic eminence. The femoral spine takes 

on a similar appearance in Cryptomanis and Necromanis 

(Koenigswald and Martin, 1990), although its position is 

difficult to compare because the iliopubic eminence is not 

preserved in available material of either taxon. In extant 

pangolins (except for M. crassicaudata and some M. pentadactyla; Gaudin et al., 2009), however, the femoral spine 

is often more robust and is situated well anterior to the 

iliopubic and acetabular eminences, displaced forward 

along the lateral iliac crest (Figure 38a).

The acetabular fossa in Patriomanis is broad and 

ovate, with a U-shaped articular (lunate) surface for the 

femoral head interrupted by a wide opening that faces 
n u m b e r 9 8 • 6 1

are nearly equal in height with the neural spines (Gaudin 

et al., 2009). The sacral neural spines and metapophyses are distinct and separate from one another in fossil 

pangolins, whereas in some modern pangolins these elements fuse to varying degrees (Gaudin et al., 2006, 2009). 

Gaudin et al. (2006) described the ventral surface of the 

sacral vertebrae in Cryptomanis, noting that the centra are 

“bulbous” with anteroposteriorly elongated ventral sacral 

foramina. They contrast this with the condition in extant 

forms, in which the centra have a flat surface and circular sacral foramina. In this respect the sacral vertebrae 

are broadly U shaped, with the posterior curvature being 

much shallower. 

As in other pangolins, the obturator foramen in 

Patriomanis is relatively small and ovate, its maximum 

diameter only slightly more than 1.25 times the maximum diameter of the acetabulum (Figure 39b). By comparison, the diameter of the obturator foramen in the 

opossum Caluromysiops (Gaudin, unpublished data), the 

basal carnivoran Nandinia, and the palaeanodont Metacheiromys is approximately twice that of the acetabulum 

(Gaudin et al., 2009). The foramen is ovate, with its long 

axis running anteroposteriorly, similar to the condition in 

Manis or Smutsia. In Phataginus the opening is more circular, whereas in Eomanis and Euromanis (Storch, 1978; 

Storch and Martin, 1994) it is more triangular, tapering 

anteriorly. 

The sacrum in Patriomanis comprises three fused vertebrae (Figure 40). As in Euromanis krebsi (Storch and 

Martin, 1994), Cryptomanis (Gaudin et al., 2006), and 

Phataginus (Gaudin et al., 2009), which also have three 

sacrals, only the anterior two sacral vertebrae are attached 

to the pelvis, and the attachment is restricted to the ilium 

(Figures 39a, 40b). In Manis and Smutsia, the sacrum 

incorporates three to four vertebrae, at least three are 

attached to the pelvis, and the attachment extends posteriorly to include the anterior part of the ischium (Gaudin 

et al., 2009). In fossil pangolins, the iliosacral junction is 

sutural, whereas the joint fuses in adults of extant taxa 

(Gaudin et al., 2009). The transverse process of the first 

sacral vertebrae and the anterior portion of the second are 

expanded dorsoventrally in Patriomanis to accommodate 

the auricular surface. In contrast, the transverse process 

of the third sacral vertebrae and the posterior portion of 

the second are thin dorsoventrally but greatly expanded 

anteroposteriorly to form a winglike structure. The lateral 

edges of these latter processes overlie the medial edge of 

the ischium. In most extant manids, the transverse process 

of the last sacral vertebra is cylindrical or rod shaped, is 

unexpanded anteroposteriorly, and extends far laterally 

beyond the body of the ischium (Gaudin et al., 2009). In 

living pangolins this transverse process also bears a process on its ventral surface for the sacrospinous ligament 

(Gaudin et al., 2009) that is absent in Patriomanis. The 

neural spines of the sacrum in Patriomanis are tall and 

robust, and the metapophyses are short, such that the 

heights of the former are more than twice that of the latter 

(Figures 39a, 40b). Euromanis krebsi (Storch and Martin, 

1994) is similar, although the neural spines are somewhat 

lower and broader anteroposteriorly. In extant manids, 

the sacral metapophyses tend to be elongated so that they 

FIGURE 40. Sacrum of Patriomanis americana, USNM-P 29996

(a) in dorsal view; (b) in left lateral view. Abbreviations: aur s, auricular surface; az, anterior zygapophysis; c, centrum; mp, metapophysis; ns, neural spine; pz, posterior zygapophysis; spn f, foramen for 

spinal nerve; tp, transverse process. Scale bar = 1 cm. Illustrations 

by Julia Morgan Scott for Smithsonian Institution.
6 2 • 

(Figure 41d), being marked instead in USNM-P 494439 

by a series of jagged points that might be the result of 

postmortem breakage or damage incurred in preparation 

(Figure 41c). In both Euromanis and Eomanis (Storch and 

Martin, 1994; Storch, 2003) the scapula is much narrower 

and more triangular in appearance, largely because of the 

narrower supraspinous and infraspinous fossae. The latter is much less expanded dorsally, especially in Eomanis, 

so that the angle between the posterior and dorsal borders is approximately 75o. Only the region immediately 

around the glenoid is preserved in the known specimens of 

Cryptomanis (Gaudin et al., 2006) and Necromanis (Koenigswald and Martin, 1990), so the overall shape of the 

scapula cannot be compared to that of Patriomanis.

The supra- and infraspinous fossae of extant pangolins typically extend dorsally to contact the dorsal border of the scapula, whereas in Patriomanis there is a ridge 

marking the upper limit of these depressions that lies 

below the dorsal border, meaning that a small sliver of 

scapula would have been exposed above the infraspinatus and supraspinatus muscles. Eomanis and Euromanis 

(Storch and Martin, 1994; Storch, 2003) resemble the 

extant taxa in this regard. The scapular spine in Patriomanis is tall and folded over posteriorly along its lateral 

edge. When viewed distally, the height of the scapular 

spine is nearly equivalent to the maximum width of the 

glenoid fossa, much higher than that in modern pangolins, where scapular spine height ranges from 65% to 80% 

of the maximum width of the glenoid (Table A9; Gaudin 

et al., 2009). The spine is also more elongated than in 

modern forms, extending close to, but not quite reaching, 

the dorsal border of the scapula. In extant manids, the 

spine ends at a noticeably more ventral point. With respect 

to the height and length of the scapular spine, Euromanis 

and Eomanis more closely resemble Patriomanis than the 

condition in Manidae (Storch and Martin, 1994; Storch, 

2003; Gaudin et al., 2009). As noted above, the acromial 

end of the scapular spine is damaged in both the type 

specimen and USNM-P 299960. The preserved portion 

of the acromion in the latter specimen (Figure 41a; Rose 

and Emry, 1993: fig. 7.5) closely resembles the morphology of the acromion in Manis javanica (USNM 198852; 

see Rose and Emry, 1993: fig. 7.5), in which the acromion 

is short, is bent slightly forward at its tip, and bears what 

appears to be a rudimentary metacromion process along 

its posterior margin. However, the damage to the lateral 

surface leaves open the possibility that a better-developed 

acromion and a better-developed metacromion were present in Patriomanis. Other living pangolins tend either to 

resemble the condition in M. javanica or to exhibit even 

in Patriomanis resemble the condition in modern forms 

rather than that of Cryptomanis.

Forelim

Figures 41–52; Ta

Scapula

A complete scapula is not preserved. However, the 

morphology of nearly the entire element can be reconstructed on the basis of the available material, with the 

exception of the anterior edge (Figure 41c). Contrary to 

the initial report in Emry (1970), there is a substantial portion of the right scapula preserved in the type specimen. 

This specimen includes the entire scapular spine but for 

its ventral tip, as well as the central portion of the dorsal 

border and most of the glenoid, although the anterior and 

posterior scapular borders, along with most of the supraspinous and infraspinous fossae, are broken away (Figure 

41b). In USNM-P 299960, a portion of the right scapula 

is also preserved, including nearly the whole ventral portion of the bone with an intact glenoid articulation, most 

if not all of the posterior border, and the ventral reaches 

of the scapular spine (Figure 41a). Once again, the ventral 

tip of the scapular spine has been worn away, so that the 

morphology of the acromion process cannot be unequivocally reconstructed. This specimen has been illustrated in 

Rose and Emry (1993: fig. 7.5). USNM-P 299960 also 

preserves the glenoid region from the left scapula. Portions of both the left and right scapula are preserved in 

USNM-P 494439, including, most importantly, the entire 

dorsal border posterior to the scapular spine, the posterior 

angle, and the dorsal portions of the posterior border on 

the left side (Figure 2). Neither scapula has been prepared 

free from the matrix in USNM-P 494439.

The scapula in modern pangolins tends to be broad 

and rectangular in shape (Figure 41d). Indeed, it is almost 

square in some taxa, with the maximum anteroposterior 

length roughly equal to the maximum dorsoventral height. 

More particularly, the supraspinous fossa is generally 

rectangular, whereas the infraspinous fossa is triangular, 

broad dorsally but narrowing ventrally, the straight posterior border of the scapula forming an acute (roughly 60o) 

angle with the dorsal border. The shape of the supraspinous fossa cannot be determined in Patriomanis, but the 

infraspinous fossa can be reconstructed in its entirety, and 

its shape resembles that of the modern forms. The posterior angle is more rounded than in extant manids and 

lacks the suprascapular ossification present in these taxa 
n u m b e r 9 8 • 6 3

extremely elongated acromion process, extending ventrally far past the glenoid to lie lateral to the proximal 

humerus (although not contacting the humerus), much 

as in the palaeanodont Metacheiromys (Simpson, 1931). 

less development of the acromion, with a blunt rounded 

process that lacks even the rudiment of a metacromion 

(Figure 41d). In contrast, both Euromanis and Eomanis (Storch and Martin, 1994; Storch, 2003) possess an 

FIGURE 41. Scapula of Patriomanis americana and extant pangolin Smutsia temminckii: (a) right scapula 

P. americana, USNM-P 299960, in lateral view; (b) right scapula of P. americana, F:AM 78999 (holotype), 

in lateral view; (c) composite reconstruction of right scapula of P. americana in lateral view; (d) left scapula 

(reversed image) of S. temminckii (AMNH 168955) in lateral view. Abbreviations: ap, acromion process; cop, 

coracoid process; g, glenoid cavity; igt, infraglenoid tubercle; isf, infraspinous fossa; por, posterior ridge of 

infraspinous fossa; scn, scapular notch; ssc, suprascapular ossification; ssf, supraspinous fossa; ssp, spine of 

scapula; tmf, teres minor fossa. Cross-hatched areas represent broken surfaces. Scale bars = 1 cm. Illustrations 

(a)–(c) by Julia Morgan Scott (JMS) for Smithsonian Institution (SI); photo (d) by Tim Gaudin, composed and 

labeled by JMS for SI.
6 4 • 

process (Figure 41a,c). A similar morphology exists in 

most modern pangolins, where a low knob or rugosity is 

often developed in this area, contrary to claims by Horovitz (2003) that no vestige of the process can be found in 

living pangolins. 

Although most of the anterior edge of the scapula is 

lacking in the available Patriomanis specimens, USNMP 299960 preserves enough of the ventral portion of the 

supraspinous fossa to show that it extended well anterior 

to the anterior edge of the glenoid and that the ventral 

edge of the supraspinous fossa was marked by a low, 

broad scapular notch like that in many extant pangolins 

(Figure 41a,c). 

Clavicle

Although a clavicle is known to occur in both Eomanis (Storch, 1978) and Euromanis (Storch and Martin, 

1994), it has not been recovered in either Necromanis or

Cryptomanis and is definitely lacking in living Manidae 

(Gaudin et al., 2009). No vestige of the clavicle has been 

recovered with any of the known Patriomanis specimens. 

Given the completeness of this material, there is strong 

negative evidence that the clavicle was absent in Patriomanis as well.

Humerus

In the type specimen, only the distal portion and a partial shaft of the humerus are preserved (Emry, 1970). However, a complete left and partial right humerus are available 

from USNM-P 531556. A damaged right humerus is available from USNM-P 494439, and the essentially complete 

left humerus is also present, although still in place, incompletely prepared, on the underside of the block; a nearly 

complete right humerus is available from USNM-P 299960 

(Figures 38, 39) and has been illustrated by Rose and 

Emry (1993).

The humeral head in Patriomanis is strongly convex 

both transversely and longitudinally and ovoid, with its 

long axis tilted slightly in a distomedial direction as in 

other pangolins (Figure 42b; Gaudin et al., 2006, 2009). 

The head is flanked by prominent greater and lesser tubercles and is separated from the former in proximal view 

by a deep, anteroposteriorly elongated fossa (Figure 43a). 

This fossa, which presumably accommodates the tendon 

of origin of the m. biceps brachii passing laterally to insert 

on the “outer surface ... of the glenoid cup” (Humphry, 

1870:37), is also present and even more sharply defined in 

modern Manis but is absent in living African pangolins. 

Neither Eomanis nor Euromanis shows any evidence of a 

separate metacromion, however, in strong contradistinction to Metacheiromys. 

The posterior border of the scapula in Patriomanis is 

marked for most of its length by an elevated lateral crest 

that becomes sharper ventrally (Figure 41a,c). A prominent 

secondary scapular spine is present in Eomanis and Euromanis (Storch and Martin, 1994; Storch, 2003; Gaudin 

et al., 2009) but is lacking in living pangolins (Figure 41d). 

Instead, the posterior border is marked laterally by a low, 

rounded eminence in these taxa. The sharp, ventral portion of the posterior ridge in Patriomanis is marked by a 

shallow but distinct, dorsoventrally elongate fossa along 

its posterior surface (Figure 41a,c). This fossa is likely a 

site for muscle origin—in other placental mammals (e.g., 

Canis, Evans and Christensen, 1979; Tenrec, Neveu and 

Gasc, 2002) this area serves as the site of origin for the 

m. teres minor and the m. triceps brachii caput longum. 

Modern pangolins are known to possess both a scapular 

insertion of the m. triceps brachii long head (Humphry, 

1870; Windle and Parsons, 1899) and a distinct m. teres 

minor (Windle and Parsons, 1899). In Gaudin et al. (2009: 

char. 203) this depression is called a “teres minor fossa” 

because of its position and its relatively small size, but in 

fact, it could have accommodated the origin of the teres 

minor along with the long head of the triceps in whole or 

in part. This depression is also present in Euromanis and 

Eomanis but is absent in Necromanis and extant pangolins (Gaudin et al., 2009). At the ventral terminus of the 

posterior ridge, the scapula of Patriomanis is marked by 

a prominent infraglenoid tuberosity (Figure 41a,c), a feature also present in Cryptomanis and living manids but 

not well developed in Euromanis or Eomanis (Storch and 

Martin, 1994; Storch, 2003).

The glenoid fossa in Patriomanis is generally egg 

shaped in outline, narrowing slightly at its anterior end. It 

is gently concave both transversely and anteroposteriorly, 

with its anterior terminus extending somewhat farther 

ventrally than its posterior end in lateral view. It differs in 

only minor respects from the glenoid morphology of other 

extinct and extant pangolins. For example, in Cryptomanis and in Smutsia, the outline of the glenoid is straighter 

along the medial edge and more convex along the lateral 

edge, whereas in Euromanis the glenoid outline is more 

rounded and does not narrow anteriorly as in other pangolins. Cryptomanis (Gaudin et al., 2006) is described as 

having a rudiment of the coracoid process in the form of 

a small but distinct tubercle. There is a rugosity on the 

anterior margin of the glenoid in Patriomanis that likely 

represents an even less developed version of the coracoid 
n u m b e r 9 8 • 6 5

pits are present on the posterior end of the greater tubercle 

in USNM-P 531556 (Figures 42b, 43a). The proximal pit 

is elongated along the posterior one-third of the proximal 

tubercular crest and is likely an insertion surface for the 

m. supraspinatus (based on analogy with the dog; Evans 

and Christensen, 1979; see also Humphry, 1870). The 

larger, circular distal pit is likely an insertion for the m. 

infraspinatus and possibly the m. teres minor (Humphry, 

1870; Evans and Christensen, 1979). 

The lesser tubercle in Patriomanis projects from the 

anteromedial edge of the head and forms a prominent 

knob (Figures 42, 43a). It is blunt with a broad base that 

is elongated distally, posteriorly, and slightly laterally from 

its proximal tip. The process is compressed orthogonal to 

this axis. In Necromanis (Koenigswald and Martin, 1990; 

Koenigswald, 1999), the lesser tubercle is of similar size 

and position. In modern manids (Figure 44a,b), the lesser 

tubercle is larger and more elongated, is positioned more 

medially at its base, and does not extend as far proximally 

as in Patriomanis and Necromanis. A distinct U-shaped 

groove is present on the anterior surface of the proximal 

humerus in Patriomanis. This groove is formed by the 

anterior extensions of the greater and lesser tubercles and 

The greater tubercle in Patriomanis is elongated 

along a posterolateral to anteromedial axis, beginning at 

the midpoint of the head and extending far anteriorly to 

form a freestanding process (Figure 43a). It is flat laterally 

and forms a sharp lateral crest along its proximal margin that is continuous with the proximal end of the pectoral crest at its most anterior and medial point (Figure 

42a). It extends proximally just beyond the level of the 

lesser trochanter but not as far as the humeral head. The 

greater tubercle is similar in Necromanis (Koenigswald, 

1969, 1999; Koenigswald and Martin, 1990) and extant 

manids, although in the living forms it typically does not 

extend as far anteriorly or medially. In Cryptomanis, the 

greater tubercle is divergent posterolaterally and extends 

farther posteriorly than in Patriomanis (Gaudin et al., 

2006). (Note that Gaudin et al.’s description of the proximal humerus contains a lapsus. The fragment identified as 

the proximal right humerus is, in fact, the left proximal 

humerus. The greater tuberosity [= greater tubercle] has 

been correctly identified and described, but in the illustration of this element in their fig. 6 the proximal humerus 

should be transposed horizontally, so that the broken deltopectoral crest points toward the greater tubercle.) Two 

FIGURE 42. Right humerus of Patriomanis america

(USNM-P 299960): (a) in anterior view; (b) in posterior view. 

Abbreviations: aer, ascending entepicondylar ridge; br, bicipital ridge; cpt, capitulum; dc, deltoid crest; dpc, deltopectoral 

crest; eec, entepicondyle; eef, entepicondylar foramen; gtu, 

greater tubercle; h, head; ispf, fossa for m. infraspinatus; ltu, 

lesser tubercle; olfo, olecranon fossa; pbt, pulley for tendon of 

m. biceps brachii; rcfo, confluent radial and capitular fossa; 

scr, supinator crest; sspf, fossa for m. supraspinatus; tr, trochlea. Scale bar = 1 cm. Illustrations drawn by Larry Isham, 

digitized and labeled by Julia Morgan Scott for Smithsonian 

Institution.
6 6 • 

(Storch, 2003) a robust, broad deltopectoral platform is 

present that differs from the narrower, more elongate crest 

of the patriomanids and Necromanis. A distal pulley is 

present in Manis and Smutsia gigantea but is absent in 

Eomanis and the other modern African species (Gaudin 

et al., 2009). Like Necromanis (Koenigswald and Martin, 

1990; Koenigswald, 1999), Patriomanis possesses a prominent deltoid crest extending from roughly the midpoint 

of the deltopectoral crest laterally and proximally, eventually turning posteriorly and continuing proximally before 

terminating at a point very close to the posterior edge of 

the greater tubercle (Figure 42). A similar deltoid crest is 

present in Manis and Smutsia but is absent in Phataginus 

(Gaudin et al., 2009). 

The humeri of USNM-P 299960 and USNM-P 

531556 confirm Emry’s (1970) surmise, based on the type, 

of a crest extending from the posterior wall of the entepicondylar foramen to the lesser tubercle. The proximal 

portion of this crest is composed of a prominent bicipital 

ridge like that found in Cryptomanis (Gaudin et al., 2006)

and Necromanis (Koenigswald and Martin, 1990; Koenigswald, 1999), extending distally from the lowermost edge 

of the lesser tubercle to the midshaft, where it is marked 

by a low, proximodistally elongate tuberosity. The lower 

edge of this tuberosity is joined by a crest ascending from 

the posterior wall of the entepicondylar foramen (Figure 

42). The latter is sharp edged distally but becomes more 

rounded proximally. Together the bicipital ridge and this 

allows for the passage of the tendon of origin of the m. 

biceps brachii (Figure 43a; Humphry, 1870; Evans and 

Christensen, 1979). This groove is present in Necromanis

(Koenigswald and Martin, 1990; Koenigswald, 1999), but 

it is strongly reduced or absent in extant Manidae (Gaudin 

et al., 2009).

The highly derived humeral shaft of fossil pangolins is 

very distinct from that of living manids. As described by 

Emry (1970) in the type specimen, the deltopectoral crest 

in Patriomanis extends from the greater tubercle to the 

proximal end of the entepicondylar bridge (Figure 42a) 

and is longer than that of any extant manid (Figure 44a,b; 

Table A10; Gaudin et al., 2009). Although the deltopectoral crest is incomplete in the type, the complementary 

portions in subsequently collected specimens confirm the 

reconstruction by Emry (1970). The deltopectoral crest 

projects farther medially than the shaft and is strongly 

folded over medially to form a deep concavity for the 

m. biceps brachii (Gaudin et al., 2006). Just proximal to 

the junction with the entepicondylar bridge, the deltopectoral crest ends abruptly in a pulley that accommodates 

the distal tendon of the m. biceps brachii (Figure 42a; 

Emry, 1970). Cryptomanis (Gaudin et al., 2006) and Necromanis (Helbing, 1938; Koenigswald and Martin, 1990; 

Koenigswald, 1999) also possess a robust deltopectoral 

crest with a distal pulley for the biceps tendon. In extant 

manids, the deltopectoral crest tends to be less robust than 

in patriomanids and Necromanis, whereas in Eomanis 

FIGURE 43. Right humerus of Patriomanis americana (USNM-P 299960): (a) in proximal vie

(b) in distal view. Abbreviations: bg, bicipital groove; cpt, capitulum; eec, entepicondyle; ftbb, fossa 

for tendon of m. biceps brachii; gtu, greater tubercle; h, head; ltu, lesser tubercle; olfo, olecranon 

fossa; scr, supinator crest; sspf, fossa for m. supraspinatus; tr, trochlea. Scale bar = 1 cm. Illustrations 

by Julia Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 6 7

FIGURE 44. Forelimb skeleton of modern pangolins: (a) left humerus of Phataginus tricuspis (CM 16206) in anterior vie

(b) left humerus of Manis javanica (USNM 198852) in anterior view; (c) left radius of P. tricuspis (CM 16206) in anterior view; 

(d) left ulna of Smutsia gigantea (AMNH 53858) in anterior view; (e) left ulna of S. gigantea (AMNH 53858) in lateral view; (f) 

right manus of P. tricuspis (CM 16206) in dorsal view (unguals reconstructed on the basis of other specimens). Abbreviations: I–V,

digits I–V; ancp, anconeal process; atb, anterior tubercle; cap, capitate; ce, capitular eminence (= anterior proximal medial process 

of Gaudin et al., 2009); cpt, capitulum; crp, coronoid process; dpc, deltopectoral crest; dt, dorsal tubercle; eec, entepicondyle; eef, 

entepicondylar foramen; een, entepicondylar notch; gtu, greater tubercle; h, head; hm, hamate; iob, interosseous border; ip, intermediate phalanx; ltu, lesser tubercle; mc, metacarpal; obb, oblique border; olec, olecranon; pis, pisiform; pp, proximal phalanx; 

raf, radial facet; rcfo, confluent radial and capitular fossa; rn, radial notch; rtu, radial tuberosity; sclu, scapholunar; scr, supinator 

crest; sn, sigmoid notch; stp, styloid process; tpd, trapezoid; tpm, trapezium; tr, trochlea; trq, triquetrum; up, ungual phalanx; utb, 

ulnar tuberosity. Scale bars = 1 cm. Parts (a), (b), (c), and (f) modified from Gaudin et al. (2009) by Julia Morgan Scott (JMS) for 

Smithsonian Institution (SI); (d) and (e) created by JMS for SI.
6 8 • 

ascending entepicondylar crest form the elevated posterior 

margin of the trough for the m. biceps brachii. In extant 

pangolins, the bicipital ridge is less pronounced. 

The supinator crest in Patriomanis is damaged in both 

specimens, but enough is preserved to confidently infer its 

morphology. It begins in the middle of the posterior surface of the shaft and curves laterally to join the lateral edge 

of the ectepicondyle. The crest has a thin, sharp lateral 

margin and carries a weak distal concavity on its anterior 

surface just proximal to the anteroposteriorly e expanded 

ectepicondyle (Figure 42a). However, the supinator crest 

in Patriomanis, Necromanis (Koenigswald and Martin, 

1990; Koenigswald, 1999), and extant Manidae (Figure 

44a,b; Gaudin et al., 2009) is weakly developed compared 

to that in Cryptomanis (Gaudin et al., 2006) and Eomanis 

(Storch, 2003; Rose et al., 2005), in which the supinator 

crest is enlarged and flared laterally.

As described by Emry (1970), the trochlea in Patriomanis has a weakly concave distal margin in posterior 

view. It is strongly convex anteroposteriorly. As is typical for placental mammals (e.g., Homberger and Walker, 

2004), the posterior surface of the trochlea is deeply concave in distal view (Figure 43b). In contrast, the anterior 

surface is slightly convex. The medial edge of the trochlea 

is sharp. It forms a small medial lappet along its distal surface that extends a short distance under the entepicondyle 

and represents the distalmost point on the humerus. Necromanis (Koenigswald and Martin, 1990; Koenigswald, 

1999) and Manis (Gaudin et al., 2009) have a similarly 

shaped trochlea with a similar medial projection. The 

trochlea of Cryptomanis also has a slightly concave distal 

edge but lacks a medial projection (Gaudin et al., 2006). 

Emry (1970) describes the capitulum in Patriomanis as 

spherical and unkeeled, similar to that in living manids. 

A lateral extension is present on the capitulum in Patriomanis that covers a small part of the anterior and distal 

ectepicondylar surface (Figure 43b).

The entepicondylar bridge is subcylindrical, slightly 

flattened anteroposteriorly, lying anterior to a welldeveloped entepicondylar foramen (Figure 42a). This 

foramen is present in all pangolins (Figure 44a,b), with the 

exception of Smutsia temminckii. The entepicondylar process in Patriomanis (Emry, 1970) is prominent, thickened 

anteroposteriorly, and tapered medially, with a rugose surface. It does not extend as far medially as the entepicondyle of Cryptomanis (Gaudin et al., 2006), Necromanis 

(Koenigswald and Martin, 1990; Koenigswald, 1999), or 

extant pangolins (Table A10), so that the overall epicondylar width of the humerus is small in Patriomanis compared to that of other pangolins (Table A10). The distal 

edge of the entepicondylar process is irregular but roughly 

straight in Patriomanis, whereas Cryptomanis, Necromanis, and all extant manids possess a strongly concave 

entepicondylar notch (Figure 44a,b; Gaudin et al., 2009). 

In Cryptomanis (Gaudin et al., 2006) and Necromanis

(Koenigswald and Martin, 1990; Koenigswald, 1999), the 

entepicondyle is flared distally.

The olecranon fossa in Patriomanis is large and 

expanded transversely, so that it is wider than the posterior trochlear surface (Figure 42b), resembling Necromanis (Helbing, 1938) in this respect. In Cryptomanis

and extant Manidae, however, the olecranon fossa is substantially narrower (Gaudin et al., 2009). The olecranon 

fossa extends farther proximally in USNM-P 299960, in 

which it forms a somewhat triangular depression, than in 

USNM-P 531556, in which the olecranon fossa takes the 

form of a flattened oval. The radial and coronoid fossae 

are confluent, forming a large, shallow, roughly circular 

depression that lies proximal to the capitulum and the lateral half of the trochlea (Figure 42a). Cryptomanis and 

Necromanis display much the same condition, whereas in 

living manids the radial and coronoid fossae are more laterally situated, lying directly over the capitulum (Figure 

44a,b; Gaudin et al., 2009).

Ulna

Only a small part of the right ulna is present in the type 

specimen (Emry, 1970). However, complete right and left 

ulnae are available from USNM-P 494439. An additional 

complete right ulna is preserved in USNM-P 299960 (Figures 45, 46). A partial right ulna is also preserved in both 

USNM-P 531556 and USNM-P 460256. The latter is the 

only Patriomanis specimen known from outside the Flagstaff Rim localities in Wyoming. It was recovered from 

Chadronian age deposits in Pipestone Springs, Montana. 

The ulna in Patriomanis is shorter relative to the humerus 

than in any of the extant pangolins (Table A10; Gaudin 

et al., 2009). 

All pangolins possess a robust olecranon process 

(Gaudin et al., 2006). In Patriomanis, the olecranon is 

roughly rectangular in shape, deep anteroposteriorly, with 

a relatively straight posterior edge and a slight medial 

inflection (Figures 45, 46a,b). The olecranon of Cryptomanis is also deep but has no medial inflection, bending 

posteriorly instead, with both anterior and posterior edges 

inclined in this direction (Gaudin et al., 2006). In Necromanis (Koenigswald, 1969, 1999; Koenigswald and Martin, 1990) the olecranon is longer, less deep, and angled 

both medially and posteriorly, whereas in Eomanis and 
n u m b e r 9 8 • 6 9

Euromanis the olecranon is more similar to that of Patriomanis, except that it has a stronger medial inflection and 

is actually bent somewhat anteriorly (Storch, 1978; Storch 

and Martin, 1994). Among living manids the olecranon 

has a more pronounced medial inflection than in any fossil 

pangolin, so that the process tends to be broader transversely than it is deep (Figure 44d,e). It is also posteriorly 

bent in most modern forms, but its length varies, so that in 

some taxa (Phataginus) the olecranon is shorter than that 

of Patriomanis, whereas in others it is longer (M. crassicaudata and M. pentadactyla; Gaudin et al., 2009).

FIGURE 45. Right ulna of Patriomanis americana (USNM

299960): (a) in posterior view; (b) in anterior view. Abbreviations: 

ancp, anconeal process; atb, anterior tubercle; crp, coronoid process; 

iob, interosseous border; olec, olecranon; raf, radial facet; rn, radial 

notch; sn, sigmoid notch; stp, styloid process; utb, ulnar tuberosity. 

Scale bar = 1 cm. Drawn by Larry Isham, digitized and labeled by 

Julia Morgan Scott for Smithsonian Institution.

As indicated by Emry (1970), the semilunar notch in 

Patriomanis is transversely expanded (Figure 45b). The 

anconeal process of Patriomanis (Emry, 1970) is wider 

than the coronoid process and extends farther anteriorly 

(Figures 45b, 46a). It extends laterally beyond the lateral 

edge of the shaft, almost as far laterally as the radial notch 

(Figure 45b). It is very similar in these respects to other 

pangolins (Figure 44d,e; Koenigswald, 1969, 1999; Koenigswald and Martin, 1990; Storch and Martin, 1994; Szalay and Schrenk, 1998; Storch, 2003; Gaudin et al., 2006, 

2009). The medial edge of the anconeal process is slightly 

higher than the lateral and is connected to the proximal 

end of the olecranon by a distinct ridge. The coronoid process in Patriomanis is ovate, with its long axis oriented in 

a posteroproximal to anterodistal direction. The coronoid 

process is directed medially, as in all pangolins (Gaudin 

et al., 2006). It flares much farther medially than the shaft, 

and its lateral edge is well medial to the midline of the 

anconeal process (Figure 45b). The articular surface for 

the humeral trochlea has a complex shape. It is deeply concave proximodistally. Its proximal portion is strongly convex transversely and faces anteriorly and distally, whereas 

the distal portion is flat transversely, facing anteriorly and 

proximally. As noted by Emry (1970), the radial notch is 

contiguous with the humeral articulation. It is expanded 

transversely, narrow proximodistally, with a rounded lateral margin and a tapering medial extremity (Figures 45b, 

46a). It extends far lateral to the ulnar shaft and is supported by a distinct lateral flange, as in other pangolins 

(Figure 44d; Koenigswald, 1969, 1999; Koenigswald and 

Martin, 1990; Szalay and Schrenk, 1998). It is flat and 

faces directly anteriorly, as in Cryptomanis (Gaudin et al., 

2006), except for the most medial portion, which is curved 

anteriorly. In Necromanis the radial notch is more strongly 

concave (Koenigswald and Martin, 1990). In modern pangolins the radial notch faces more laterally. As described 

by Emry (1970), a distinct ulnar tuberosity lies distal to 

the semilunar notch on the anterior surface of the shaft 

just distal to the coronoid process (Figure 45b), a feature 

present in Necromanis, Cryptomanis, Eomanis, and all living pangolins (Figure 44d; Gaudin et al., 2006, 2009). The 

ulnar tuberosity serves as the site of insertion for the m. 

brachialis muscle in humans and for the mm. brachialis 

and biceps brachii in Canis (Evans and Christensen, 1979).

The ulnar shaft in Patriomanis (Emry, 1970) is flattened transversely and is concave on its medial and lateral sides (Figure 46a,b). The medial concavity begins just 

distal to the tip of the olecranon, where it occupies the 

posterior third of the process. It expands anteroposteriorly 

and deepens below the semilunar notch, then becomes 
7 0 • 

narrower and less deep toward the midshaft region, before 

expanding and deepening again in the distal third of the 

ulnar shaft, ending just proximal to the styloid process. 

The distal portion of this medial concavity is not present 

in most modern pangolins but is present in Cryptomanis 

and Necromanis (Koenigswald and Martin, 1990; Koenigswald, 1999; Gaudin et al., 2006). This medial concavity 

likely accommodates the m. flexor digitorum profundus 

(Jouffroy, 1966; Gaudin et al., 2006). A deep, sharp-e 

edged lateral concavity extends distally from the level of 

the anconeal process, becoming gradually less concave 

distally to terminate approximately three-fourths of the 

way down the shaft (Figure 41a). In Cryptomanis (Gaudin 

et al., 2006), this concavity extends much farther distally, 

whereas in the Manidae it terminates farther proximally. 

This concavity likely housed the mm. extensor digitorum 

profundus and abductor pollicis longus (Humphry, 1870; 

Jouffroy, 1966; Gaudin et al., 2006). The posterior surface of the ulna is flat proximally but tapers distally to 

form a narrow ridge (Figure 45a). The anterior surface 

of the ulnar shaft is marked by a sharp-e edged interosseous border that begins at the base of the radial notch and 

extends almost to the anterior tubercle at the distal end of 

the ulna. In its middle portion it is slightly folded over laterally, and it carries a slight concavity on its anteromedial 

surface. A second, very slight concavity on the anterolateral surface at the distal end represents the distal radial 

facet. Cryptomanis (Gaudin et al., 2006) has a very similar 

morphology. The ulnar shaft is straight and does not taper 

distally as it does in some modern pangolins (Figure 44e), 

but rather, as in Cryptomanis (Gaudin et al., 2006) and 

Eomanis (Storch, 2003), maintains a uniform depth.

The distal end of the ulna in Patriomanis, as in Cryptomanis (Gaudin et al., 2006), is flattened transversely and 

deep anteroposteriorly (Figure 46c). In fact, the distal end 

of the shaft in Patriomanis is deeper relative to maximum 

ulnar length than in all extant pangolins (Table A10). As in 

other pangolins (Gaudin et al., 2006), the distal end of the 

ulna in Patriomanis bears two distinct tubercles. The anterior tubercle lies on the anterior edge of the distal surface. A 

smaller posterior tubercle is situated midway up the lateral 

edge of the distal surface, just proximal to the lateral edge 

of the styloid process. The anterior tubercle represents the 

distal termination of the interosseous border that begins at 

the base of the coronoid process. The styloid process itself is 

capped by an ovate articular surface, its long axis oriented 

in an anterolateral to posteromedial direction. The styloid 

articular surface is thus wider than it is deep (Figure 46c), 

as it is in other therians (Gaudin et al., 2009). In contrast, 

in Cryptomanis and most living pangolins (with the sole 

FIGURE 46. Right ulna of Patriomanis americana (USNM

299960): (a) in lateral view; (b) in medial view; (c) in distal view. 

Abbreviations: ancp, anconeal process; atb, anterior tubercle; crp, 

coronoid process; iob, interosseous border; laco, lateral concavity 

for mm. extensor digitorum profundus and abductor pollicis longus; meco, medial concavity for m. flexor digitorum profundus; olec, 

olecranon; ptb, posterior tubercle; raf, radial facet; rn, radial notch; 

sas, styloid articular surface; sn, sigmoid notch; stp, styloid process; 

utb, ulnar tuberosity. Scale bars = 1 cm. Illustrations by Julia Morgan 

Scott for Smithsonian Institution.
n u m b e r 9 8 • 7 1

but similar to that of Necromanis (Helbing, 1938; Emry, 

1970). The proximal articulations were identified by Emry 

(1970). They include, on the proximal surface, a central 

capitular facet, a lateral shelf for articulation with the lateral extension of the capitulum, and a medial excavation 

for the trochlea, as well as an ulnar facet on the posterior 

surface and a sesamoid facet on the lateral surface (Figures 47a, 48a,c). In USNM-P 299960 the anterior edge 

of the proximal radius is deeply indented lateral to the 

capitular eminence (Figure 47b,c). This indentation is less 

well developed in USNM-P 531556 and is absent in the 

type (Emry, 1970). This indentation is also missing in Necromanis (Helbing, 1938) but is variably present in living 

pangolins. The capitular eminence is strongly developed in 

Cryptomanis, Necromanis (Helbing, 1938; Koenigswald 

and Martin, 1990; Gaudin et al., 2006), and extant manids 

(Figure 44c) except for S. gigantea. The triangular trochlear facet in Patriomanis (Emry, 1970) is wide, covering 

approximately one-third of the radial head. It is similarly 

well developed in Cryptomanis (Gaudin et al., 2006) and 

Necromanis (Helbing, 1938), whereas in extant manids 

the trochlear facet covers a much smaller portion of the 

proximal surface (Gaudin et al., 2009). The sesamoid 

facet identified by Emry (1970) in Patriomanis and Necromanis is small relative to that in modern pangolins. It also 

faces directly lateral in the two fossil genera, whereas in 

extant forms it is inclined distolaterally so that it becomes 

exception of M. pentadactyla), the styloid articular surface 

is compressed transversely. The articular surface is slightly 

convex both transversely and anteroposteriorly, and covers 

the posterior half of the distal ulnar surface, with its distalmost projection lying along its posterior edge. The articular 

surface faces distally and somewhat anteriorly, forming a 

45° angle with the long axis of the ulna. The styloid process is shorter proximodistally than in living manids (Table

A10) but extends farther distally than in Cryptomanis, in 

which the styloid articular surface forms a steep angle relative to the long axis of the ulna.

Radius

The type specimen preserves only partial radii, including the left proximal and left and right distal ends described 

and illustrated in Emry (1970). However, complete radii are 

available from USNM-P 494439 and USNM-P 299960, 

although the right radius of USNM-P 299960 is slightly damaged distally, missing its pseudostyloid process (Figures 47, 

48). A complete right radius and a partial left radius are also 

available from USNM-P 531556. 

The radius in Patriomanis is shorter relative to humeral 

length than in any extant pangolin (Table A10). The radial 

head is also transversely expanded more so than in modern 

manids (Figure 48a; Emry, 1970), with a depth to width 

ratio lower than that of all living pangolins (Table A10) 

FIGURE 47. Radius of Patriomanis americana: (a) rig

radius of USNM-P 299960 in posterior view; (b) right radius 

of USNM-P 299960 in anterior view; (c) left radius of USNMP 299960 in anterior view. Abbreviations: ce, capitular eminence (= anterior proximal medial process of Gaudin et al., 

2009); das, distal articular surface; dt, dorsal tubercle; duf, 

distal ulnar facet; iob, interosseous border; obb, oblique border; pstp, pseudostyloid process; puf, proximal ulnar facet; 

rtu, radial tuberosity; sef, sesamoid facet; stp, styloid process. 

Cross-hatched areas represent broken surfaces. Scale bar = 

1 cm. Illustrations (a), (b) drawn by Larry Isham, digitized 

and labeled by Julia Morgan Scott (JMS) for Smithsonian 

Institution (SI); (c) by JMS for SI. 
7 2 • 

Emry (1970, based on Humphry, 1870, and Windle and 

Parsons, 1899), however, the biceps does not have a radial 

attachment, at least in Manis. Jouffroy (1966:266, discussion of m. pronator teres) implies that such an attachment 

is present in S. gigantea. The process may also serve as a 

site of origin for the very large m. flexor digitorum profundus (Humphry, 1870;Windle and Parsons, 1899; Jouffroy, 

1966) of living pangolins.

The proximal radial shaft is transversely broad and 

anteroposteriorly compressed, reflecting the shape of the 

radial head. Distally, the shaft becomes increasingly compressed transversely and expanded anteroposteriorly (Figures 47, 48c), as noted by Emry (1970). At its deepest point, 

approximately three-fourths of the way down the shaft, its 

depth is three times greater than its width. The lateral compression of the radial shaft is much more pronounced in 

Patriomanis than in Cryptomanis (Gaudin et al., 2006), 

Necromanis (Helbing, 1938; Koenigswald and Martin, 

1990), or living manids. As in other pangolins, the anterior 

surface is marked by a very sharp and elevated oblique border, which originates on the proximal quarter of the shaft 

and extends its entire length, curving sharply mediad as it 

approaches the distal margin of the radius (Figure 47b,c). 

The oblique border terminates in a weak tubercle, which, 

following the terminology of MacPhee (1994), we identify 

as the dorsal tubercle. A weaker, rounded ridge extends 

straight distally from the oblique border near the point 

where it begins to curve medially. This ridge terminates 

at the base of a prominent pseudostyloid process (Figure 

47c; terminology of MacPhee, 1994; equal to “anteriorly 

directed tubercle” of Emry, 1970:480). The pseudostyloid process is rectangular, slightly wider than long, and is 

directed anteriorly, distally, and laterally. It is tilted somewhat so that its lateral edge is more elevated than its medial 

edge, and it extends farther distally than any other part 

of the radius. A prominent pseudostyloid process is also 

present in Necromanis (Koenigswald and Martin, 1990). 

In Cryptomanis (Gaudin et al., 2006), the median ridge 

extending distally from the oblique border is much more 

pronounced than in Patriomanis, but the region where a 

pseudostyloid process would be located is not preserved. 

The oblique border of living pangolins is, at best, weakly 

bifurcated distally, and there is no pseudostyloid process 

present (Figure 44c). Rather, the dorsal tubercle at the 

medial termination of the oblique crest is much more 

prominently developed than in Patriomanis. 

The posterior surface of the radial shaft bears a sharp 

interosseous border distally (Figure 47a). This crest bifurcates proximally near the midshaft. The medial edge and 

more prominent lateral edge of the bifurcated interosseous 

visible in a proximal view of the radial head. The proximal ulnar facet in Patriomanis (Emry, 1970) is narrow 

proximodistally and elongated and convex transversely 

(Figure 47a). It abuts the sesamoid facet along its lateral 

margin, in contrast to the condition in extant manids, in 

which the two facets are clearly separate (Gaudin et al., 

2009). Patriomanids and Necromanis lack a prominent 

radial tuberosity (Gaudin et al., 2009), but this process 

is strongly developed on the medial edge of the proximal 

radius in all modern manids except S. temminckii (Figure 44c). The function of this process in living manids is 

not entirely clear. In other placental mammals, the radial 

tuberosity serves as an insertion point for the m. biceps 

brachii (e.g., Evans and Christensen, 1979; Neveu and 

Gasc, 2002; Homberger and Walker, 2004). According to 

FIGURE 48. Left radius of Patriomanis americana (USNM

299960 ): (a) in proximal view; (b) in distal view; (c) in lateral view. 

Abbreviations: ce, capitular eminence; cpf, capitular facet; dt, dorsal tubercle; duf, distal ulnar facet; iob, interosseous border; obb, 

oblique border; pstp, pseudostyloid process; puf, proximal ulnar 

facet; rtu, radial tuberosity; sef, sesamoid facet; slf, scapholunar 

facet; stp, styloid process; trof, trochlear facet. Scale bar = 1 cm. Illustrations (a), (b) by Julia Morgan Scott for Smithsonian Institution 

and (c) by T. J. Gaudin.
n u m b e r 9 8 • 7 3

extremity of the distal radial articular surface. In living 

pangolins the styloid process is robust and forms a distinct 

process, confluent with, but set off by, indentations from 

the rest of the distal articular surface. It extends in either a 

ventral (Phataginus) or ventrolateral (Manis and Smutsia) 

direction from the medial or anteromedial margin of that 

surface (Gaudin et al., 2009). 

Scapholunar

Although Emry (1970) reports finding only two manual proximal phalanges and an ungual phalanx (probably 

from digit III) in the type, several more pieces of the manus 

have since been recovered from the type specimen, including several carpals from both the left and right side (partial right scapholunar, right pisiform, left hamate), two 

proximal phalanges, at least one complete intermediate 

phalanx, and unguals from at least four different digits (I, 

II, III, and V). There are also a partial right manus largely 

prepared free from the matrix and a partial left manus still 

in the matrix in USNM-P 494439. The former includes 

the triquetrum, trapezium, capitate, hamate, metacarpals I 

and IV, proximal phalanges I and II, intermediate phalanx 

II, and unguals I, II, IV, and V. The latter includes most of 

digits II, III, and IV. However, the descriptions of the various elements in the manus that follow are based largely 

on USNM-P 299960, except where otherwise noted. This 

specimen includes an intact, articulated right manus (Figure 49), as well as much of the left manus. It is the only 

specimen with an intact scapholunar element (Figure 50).

border are separated by a weak depression. In Necromanis 

(Helbing, 1938) and in extant pangolins, the interosseous 

border is similar in length but is undivided proximally. 

Multiple small, tubercular rugosities are situated around 

the proximal margin of the aforementioned posterior 

radial fossa in Patriomanis. These rugosities are also present in modern Manis but are absent in the extant African 

species. The distal ulnar facet lies near the distal margin 

of the posterior radial surface. Its medial edge is formed 

by the interosseous border. It is triangular and flat, as in 

all pholidotans, and is oriented posteriorly and somewhat 

laterally (Figures 47a, 48c).

Emry (1970:480) described the distal end of the 

radius as “strongly expanded and heavy ... [with] a single 

concave, transversely oval articular surface.” The articular surface is tilted distolaterally and slightly posteriad. 

In distal view (Figure 48b), the long axis of the articular 

surface appears twisted 45° relative to the long axis of the 

head (twisted counterclockwise on the right, clockwise on 

the left). Cryptomanis (Gaudin et al., 2006) likely shares 

a similar morphology judging by the preserved portions 

of the distal articulation, whereas in extant manids the 

distal articular surface is either circular (S. gigantea) or 

compressed transversely, so that its long axis is perpendicular to that of the radial head (Gaudin et al., 2009). 

As noted by Gaudin et al. (2006), this difference in orientation of the distal radius in the modern taxa may be 

related to the adoption of a pedolateral stance. The styloid 

process in Patriomanis is rudimentary, taking the form of 

a low, rounded tubercle situated near the posteromedial 

FIGURE 49. Right manus of Patriomanis americana (USNM-P 299960): (a) in dorsal vie

(b) labeled line drawing of the same. Abbreviations: I–V, digits I–V; cap, capitate; etb, extensor

tubercle; hm, hamate; ip, intermediate phalanx; 

mc, metacarpal; pp, proximal phalanx; sclu, 

scapholunar; tpd, trapezoid; tpm, trapezium; up, 

ungual phalanx. Scale bar = 1 cm. Illustration (a) 

drawn by Larry Isham, digitized by Julia Morgan 

Scott (JMS) for Smithsonian Institution (SI); (b) 

by JMS for SI.
7 4 • 

scaphoid and lunate are already fused, perhaps indicating 

that this embryo represents too late a stage to be informative regarding the composition of the adult bone. 

The scaphoid and lunate elements of the carpus are 

also fused in Cryptomanis (Gaudin et al., 2006) and 

Necromanis (Koenigswald and Martin, 1990) but not in 

Eomanis (Gaudin et al., 2009), although the evidence in 

the case of the latter is somewhat equivocal. In Patriomanis, as in the other Manoidea, the fused scapholunar 

element is clearly present, constituting the largest element 

by far in the carpus. The bone is half again as wide as it is 

deep in proximal view, much like the condition in modern 

pangolins (Figures 44f, 49, 50; Table A11). It articulates 

proximally with the radius and distally with the trapezium, trapezoid, capitate, and hamate. The proximal surface is dominated by the large articular facet for the radius 

(Figure 50a). This facet is much deeper dorsopalmarly in 

its lateral half than it is in its medial half, with the lateral 

portion roughly ovoid in outline and the medial portion 

tapering slightly toward the medial end, so that the facet 

resembles a rounded, sideways L in shape. The medial 

and lateral halves of the facet are separated in part by a 

strong concavity along the palmar margin. This concavity is also present in extant Manidae but is not present in 

Cryptomanis or Necromanis, in which the palmar margin 

is roughly straight (Gaudin et al., 2009). It is interesting that despite marked differences in the shape of the 

distal facet of the radius in modern pangolins versus 

Patriomanis, the shape of the proximal facet of the scapholunar does not generally reflect these differences but is 

instead generally similar among Patriomanis and most 

extant taxa.

In dorsal view, the scapholunar is very broad transversely and short proximodistally, with just a small sliver 

of a nonarticular dorsal surface exposed (Figure 49). In 

contrast to Cryptomanis and extant pangolins (Gaudin 

et al., 2009), there is no articular facet along the lateral 

edge for the triquetrum. The palmar surface of the bone is 

dominated by a large, rounded, smooth-surfaced palmar 

tuberosity. This protrusion covers the entire lateral half 

of the bone. It is ovate, with a transverse diameter much 

greater than its proximodistal extent. This is also the case 

in Cryptomanis, but in modern pangolins the palmar 

tuberosity tends to be more proximodistally elongated 

(Gaudin et al., 2009); in some taxa (e.g., Manis javanica, 

USNM 198852), the proximodistal diameter of the palmar tuberosity is nearly half the maximum width of the 

entire scapholunar.

In distal view, the surface of the scapholunar bears three 

facets separated from one another by prominent ridges 

In modern pangolins, the scaphoid and lunate elements of the carpus are fused into a single bone, the 

scapholunar. It is possible that this element is actually a 

scaphocentrolunate, that is, that an embryologically separate centrale element forms an additional contribution 

to the adult element. However, in the manus of a 7.6-cm 

embryo of Phataginus tricuspis illustrated by Szalay and 

Schrenk (1998: fig. 25), there is no indication of a separate centrale element. It should be noted, however, that the 

FIGURE 50. Left scapholunar of Patriomanis americana (USNM

299960): (a) in proximal view; (b) in distal view. Abbreviations: cpf,

capitate facet; hmf, hamate facet; pas, proximal articular surface;

patb, palmar tuberosity; tpdf, trapezoid facet; tpmf, trapezium facet. 

Scale bar = 1 cm. Photos by Tim Gaudin, composed and labeled by 

Julia Morgan Scott for Smithsonian Institution.
n u m b e r 9 8 • 7 5

In dorsal view, the triquetrum is wide transversely 

and short proximodistally, much like modern pangolins. 

It has a quadrate outline in USNM-P 299960, whereas in 

USNM-P 494439 it is tapered at either end, giving it a 

triangular shape (Figure 51a). Its dorsal surface is convex 

transversely and slightly concave proximodistally, and it 

bears a strong, smooth-surfaced knob at its lateral end.

In distal view, the triquetrum bears a single, distally 

facing facet for the hamate. This facet is roughly triangular with a medial base and is gently concave.

Pisiform

The pisiform is not preserved in USNM 494439 but is 

known from two right pisiforms, one preserved with the 

type specimen (F:AM 78999) and the other with USNMP 299960 (Figure 51b). The bone is elongated along its 

proximodistal axis, with a roughly rectangular outline. It 

has a blunt distal tip that is incised along its medial surface with a deep groove. This groove may very well have 

accommodated a tendon attaching the m. flexor carpi 

ulnaris to the outer metacarpals. In humans and dogs, the 

pisiform serves as the insertion point for the m. flexor carpi 

(Figure 50b). The facets are all strongly concave in both 

the transverse and dorsopalmar planes. Along its lateral 

edge, the scapholunar bears a crescent-shaped facet that 

faces distolaterally to articulate with the hamate. Medial to 

this is a larger facet with a roughly triangular outline (elongated in the dorsopalmar plane, apex on the palmar side) 

that contacts the capitate. Medial to this is a transversely 

wide, triangular facet (apex medial) for the trapezium 

and trapezoid. The trapezoid occupies the bulk of this last 

facet, with perhaps the medial third articulating with the 

trapezium, although there is no clear demarcation between 

these two regions on the surface of the facet itself. Both the 

capitate and trapezium–trapezoid facets face mainly distad, 

although the former is tilted slightly lateral and the latter 

slightly medial. The articulation between the scapholunar 

and hamate is missing in M. javanica and the extant African 

pangolins (Gaudin et al., 2009), and in all extant pangolin 

species the ridge separating the capitate and trapezium–

trapezoid facet is lacking (Gaudin et al., 2009).

Triquetrum

Although not figured with the remainder of the manus 

(Figure 49), an isolated, slightly damaged right triquetrum 

is preserved in USNM-P 299960 (Figure 51a), along with 

a partial left triquetrum. There is also a complete right 

triquetrum that has been prepared free of the matrix in 

USNM-P 494439 (Figure 51a). The former is somewhat 

more compressed in the dorsopalmar plane than the latter, 

but they are otherwise quite similar.

The triquetrum articulates proximally with the ulna 

and pisiform and distally with the hamate. Of the two 

articular facets on the proximal surface, the facet for the 

styloid process of the ulna is situated more dorsal and 

faces more proximally, whereas the pisiform facet lies in 

a more palmar position. Both facets are elongated in the 

transverse plane, and both are relatively flat in USNMP 494439, although the pisiform facet is slightly concave 

medially and slightly convex laterally in USNM-P 299960. 

The ulnar facet is roughly triangular or semicircular in outline, with the apex pointed dorsally. The ulnar facet is also 

elongated transversely in Eomanis, the extant pangolins 

in the genus Phataginus, and the extant species Smutsia 

gigantea, but in living Asian pangolins and S. temminckii, 

the transverse width is less than or equal to dorsopalmar 

depth (Gaudin et al., 2009). The ulnar and pisiform facets 

are contiguous along their entire transverse extent. They 

are situated well lateral to the medial edge of the bone, as 

in the extant pangolin genera Manis and Smutsia (Gaudin 

et al., 2009). 

FIGURE 51. Triquetrum and pisiform of Patriomanis america

(a) right triquetrum of USNM-P 299960 in dorsal view on right, 

right triquetrum of USNM-P 494439 in dorsal view on left; (b) right 

pisiform of USNM-P 299960 in medial view. Abbreviations: dsu, 

dorsal surface; hmf, hamate facet; tqf, triquetrum facet; ulf, ulnar 

facet. Scale bar = 1 cm. Photos by Tim Gaudin, composed and 

labeled by Julia Morgan Scott for Smithsonian Institution.
7 6 • 

facet for the second metacarpal. The upper facet is small, 

flat, and triangular; the lower facet is ovoid and concave. 

The distal surface carries the facet for the first metacarpal. 

This facet is compressed transversely and teardrop shaped, 

with a rounded palmar portion narrowing to a dorsolateral point. The outline of the distal facet for metacarpal I 

varies considerably among modern pangolins, even within 

species. In M. javanica (USNM 198852), the facet resembles that of Patriomanis, but in M. pentadactyla (USNM 

308733) and S. temminckii (AMNH 244096) the facet is 

ellipsoidal, with a rounded dorsal margin, and in S. gigantea (AMNH 53851) it is almost triangular and is broader 

transversely than it is deep in a dorsopalmar plane.

Trapezoid

The trapezoid is known only from USNM-P 299960 

(Figure 49). As in other pangolins, it is a transversely 

elongated element that articulates proximally with the 

scapholunar, medially with the trapezium, laterally with 

the capitate, and distally with the second metacarpal. In 

Patriomanis, as in Cryptomanis (Gaudin et al., 2006) 

and Eomanis (Gaudin et al., 2009), the dorsal surface 

of the trapezoid is rectangular in outline, with the distal 

and proximal articular surfaces roughly parallel to one 

another. In modern pangolins, the dorsal surface of the 

trapezoid is trapezoidal, with the distal and proximal surfaces converging on the medial side, largely because of the 

proximomedial tilt of the former (Gaudin et al., 2009). 

The proximal surface of the trapezoid carries a large 

articular facet for the scapholunar. This facet is unusual 

in that it has a concave surface contour, in contrast to the 

condition in other known living and fossil pangolins, in 

which the facet is convex (Gaudin et al., 2009). This facet 

is roughly triangular in outline, narrowing at its palmar 

end. The distal surface is also triangular and bears two 

facets. On its medial edge, there is a small rectangular 

facet facing distomedially for the trapezium. This facet 

has a very similar orientation in Cryptomanis, whereas 

in extant African pangolins (Smutsia and Phataginus), 

the facet faces directly mediad (Gaudin et al., 2009). In 

Asian pangolins (Manis), the trapezium actually overlaps 

the trapezoid proximally, so that the trapezium facet faces 

proximally and medially (Gaudin et al., 2009). The bulk 

of the distal surface is occupied by a triangular facet for 

metacarpal II. This facet is concave in its dorsolateral corner, but the dorsomedial and palmar corners form part of 

a flat or slightly convex, crescent-shaped surface. This sort 

of complex, concavo-convex metacarpal II facet is common among pholidotans.

ulnaris (Pick and Howden, 1977; Evans and Christensen, 

1979), as well as the site of origin of several ligaments 

and, in humans, the m. abductor digiti minimi. However, 

in Smutsia gigantea, the insertion of the flexor carpi ulnaris is continued by a tendon or ligament that passes from 

the pisiform onto the ulnar border of the fourth metacarpal (Jouffroy, 1966). The presence of the distal groove 

in Patriomanis suggests it may have possessed a similar 

arrangement, although neither S. gigantea nor any other 

extant pangolin was observed to have a distal groove on 

the pisiform. The proximal end of the pisiform is roughly 

triangular (apex lateral) and slightly elongated transversely. It is marked by two articular facets separated by 

a sharp crest. One facet faces more toward the proximal 

side of the carpus and articulates with the ulna, and the 

other faces more toward the distal side of the carpus and 

articulates with the triquetrum (Figure 51b). The ulnar 

facet is ovate transversely, and the triquetral facet is semicircular in outline, with the flat side facing proximally in 

dorsal view. The pisiform has not been described in any 

other fossil pangolin. 

Trapezium

The trapezium has been recovered from USNM-P 

494439 and USNM-P 299960. It has a roughly triangular outline in dorsal view, with the base facing laterally and the apex medially, and its proximodistal length 

is greater than its transverse width (Figure 49). In Manis 

javanica and M. pentadactyla, the trapezium is deeper on 

the medial side, giving it a more trapezoidal outline, but 

the bone resembles that of Patriomanis in other extant 

pangolins (Figure 44f). The trapezium articulates with the 

scapholunar proximally, the trapezoid and second metacarpal laterally, and the first metacarpal distally. The dorsal surface is curved in Patriomanis, in such a manner that 

the dorsalmost point is at the distolateral corner, and the 

dorsal surface curves in a palmar direction toward both 

its medial and proximolateral corners. This dorsal point 

actually overlaps the dorsal surface of the first metacarpal, much as was the case with the entocuneiform and first 

metatarsal. The proximal surface of the trapezium bears 

a small, convex facet for the scapholunar that occupies 

the proximolateral corner of the bone. In most living pangolins (except S. gigantea), the scapholunar facet is much 

larger, covering the entire proximal surface of the trapezium, and in every living species except M. crassicaudata 

the facet is concave rather than convex (Gaudin et al., 

2009). Along its lateral surface, the trapezium bears two 

facets: an upper facet for the trapezoid and a lower, larger 
n u m b e r 9 8 • 7 7

facet has a roughly triangular outline, although there is a 

steplike indentation approximately midway along its lateral margin. The facet takes on a similar shape in Eomanis

and Cryptomanis, whereas in all modern pangolins except 

M. javanica, the facet is more L shaped, with a very deep 

and abrupt lateral indentation restricting the width of the 

palmar portion of the facet.

The capitate is roughly semicircular in outline in both 

medial and lateral views. It articulates with the trapezoid 

medially, but this facet is actually visible in proximal 

view because the trapezoid slightly overlaps the capitate 

proximally, so that the facet on the capitate faces proximomedially (Figure 52a). The overlap of the trapezoid 

on the capitate is also evident in Eomanis, Cryptomanis, 

and some M. javanica, but in other extant pangolins the 

reverse is the case, with the capitate showing a slight proximal overlap of the trapezoid (Gaudin et al., 2009). On its 

lateral surface, the capitate bears a small, triangular facet 

for the hamate near its dorsal and distal corner. Eomanis

and Cryptomanis are similar, but in extant pangolins this 

triangular facet is displaced in a palmar direction, so that 

Capitate

A left capitate is available from USNM-P 531557, 

and a right capitate is available from USNM-P 494439, in 

addition to the left and right capitates of USNM-P 299960 

(Figures 49, 52a,b). The element lies between the scapholunar dorsally, the hamate laterally, the trapezoid medially, 

and metacarpal III distally. Its maximum width is virtually 

identical to that of the trapezoid, but its dorsal surface is 

slightly deeper proximodistally, especially along its lateral 

border with the hamate (Figure 49). The palmar surface is 

marked by a prominent, rounded tuberosity (Figure 52b). 

The facet for the scapholunar that covers the proximal surface of the bone is strongly convex both transversely and 

in the dorsopalmar plane. It is fairly narrow compared to 

that of other pangolins, its maximum width roughly half 

of its maximum dorsopalmar depth (Figure 52a). In Cryptomanis and Eomanis, the width of the facet is more than 

65% of its depth, in extant African pangolins more than 

75% of its depth, and in extant Asian pangolins between 

85% and 100% of its depth (Gaudin et al., 2009). The 

FIGURE 52. Capitate and hamate of Patrioman

americana: (a) left capitate of USNM-P 299960 in 

proximal view; (b) in distal view; (c) right hamate 

of USNM-P 494439 in proximal view; (d) in distal 

view. Abbreviations: das, distal articular surface; 

hmf, hamate facet; mIIIf, metacarpal III facet; mI

metacarpal IV facet; mVf, metacarpal V facet; pas,

proximal articular surface; patb, palmar tuberosity; 

slf, scapholunar facet; tpdf, trapezoid facet; tqf, triquetrum facet; unp, unciform process. Scale bar = 

1 cm. Photos by Tim Gaudin, composed and labeled 

by Julia Morgan Scott for Smithsonian Institution.
7 8 • 

extending in a distal, palmar, and lateral direction at its 

base, before taking a distinct turn in the proximal direction, extending outward as a proximally, laterally, and 

palmarly oriented prong (Gaudin et al., 2009).

The dorsal surface of the hamate in Patriomanis is 

trapezoidal in outline, with the proximal and distal edges 

approaching one another laterally (Figure 49). Its transverse width is nearly half again greater than that of the 

capitate or trapezoid and is also somewhat greater than 

its maximum proximodistal height. Among the African 

pangolins (Smutsia and Phataginus) and M. javanica, the 

dorsal surface takes on a more pentagonal shape and is 

more compressed transversely, such that the maximum 

proximodistal height equals or exceeds its width (Figure 

44f). In M. crassicaudata and M. pentadactyla, the dorsal 

surface is irregularly shaped but low, with its transverse 

width much greater than its height (Gaudin et al., 2009). 

The proximal surface of the hamate bears two facets: 

a small, more medially directed facet for the scapholunar 

and a larger, more laterally sloping facet for the triquetrum (Figure 52c). Both are convex mediolaterally and in 

a dorsopalmar plane. The former is narrow transversely, 

and the latter is narrower in a dorsopalmar plane. The distal articular surface is quite complex (Figure 52d). It bears 

a rectangular, transversely narrow, distomedially oriented 

facet for metacarpal III. More laterally is an ovate facet, 

gently concave both transversely and in a dorsopalmar 

plane and more elongate in a dorsopalmar than transverse 

direction. This articulates with metacarpal IV. Last, there is 

a third, even more laterally situated facet for metacarpal V. 

This facet is strongly concave in the dorsopalmar plane. 

Transversely, it is relatively flat in USNM-P 494439, 

whereas it is concave in the type specimen, F:AM 78999. 

In both specimens it is broadly triangular, narrowing at its 

palmar end, where it overlaps onto the base of the unciform process. The distal hamate surface is basically similar 

to that of Patriomanis in Eomanis but shows numerous 

differences in modern pangolins. In Phataginus, the facet 

for metacarpal III is vertical and is carried on the medial 

rather than the distal surface of the hamate (see, e.g., 

Gaudin et al., 2009: fig. 5a). In Phataginus and Manis, the 

facet for metacarpal IV becomes much broader, so that its 

width equals or exceeds its dorsopalmar depth, and the 

facet comes to occupy most of the distal hamate surface. 

Moreover, in Manis this facet, which is generally concave 

as in Patriomanis, bears a distinct, convex eminence in its 

dorsomedial corner. This eminence is present but smaller 

in the African genera and Patriomanis and is located 

between the facets for metacarpals III and IV. The facet 

for metacarpal V is directed distolaterally in M. javanica 

the facet comes to lie near the midpoint of the lateral surface (Gaudin et al., 2009).

The distal articular surface for metacarpal III has what 

Gaudin et al. (2009) term a “dumbbell shape” outline—it 

is roughly rectangular, compressed transversely, with relatively straight dorsal and palmar margins, but both lateral and medial margins are indented (Figure 52b). The 

outline is similar in Eomanis and Cryptomanis (Gaudin 

et al., 2009), whereas it differs in several respects in modern pangolins. In all extant taxa, the palmar edge is narrower than the dorsal edge. This is especially the case in 

Manis and Smutsia, where the distal capitate facet takes 

on a roughly triangular outline (Gaudin et al., 2009). In 

Phataginus, the palmar surface is only slightly narrower, 

whereas the medial border is indented and the lateral 

border is extended laterally into a blunt point, giving the 

whole articulation a bilobate appearance (Gaudin et al., 

2009). The surface contour of the facet is complicated in 

pangolins. It is concave in a dorsopalmar plane in all taxa. 

In Phataginus, the facet is then convex transversely, especially its dorsal lobe. In Patriomanis, Cryptomanis, Manis, 

and Smutsia, the facet has a midline eminence that runs in 

a dorsopalmar direction and is flanked by two concavities, 

so that the overall transverse contour of the facet is concave medially and laterally and convex in the middle. In 

the modern taxa, there is a rounded knob at the dorsal end 

of this midline eminence that appears to form a stop, preventing hyperextension at the carpometacarpal joint. The 

fairly complex surface contour of the distal capitate facet 

exhibited by Patriomanis is likely the primitive condition 

for pangolins, rather than the somewhat simpler shape in

Phataginus. 

Hamate

The hamate is the second largest element in the carpus, behind only the fused scapholunar element. It has 

been recovered from the type specimen (F:AM 78999), 

as well as USNM-P 494439 and USNM-P 299960 (Figures 49, 52c,d). It articulates with the triquetrum and 

scapholunar proximally, the capitate medially, and the 

three lateral metacarpals (III, IV, V) distally. The hamate 

is blocky but carries a prominent unciform process that 

attaches to its distal, palmar, lateral corner and extends 

in a distal, palmar, and lateral direction toward a blunt, 

rounded tip (Figure 52c,d). A prominent unciform process is also found in Eomanis and in all extant pangolins except M.

pentadactyla. In the modern taxa (excep

M.pentadactyla and some specimens of S. gigantea), however, the unciform process takes on a more complex shape
n u m b e r 9 8 • 7 9

waldi also has an elongated third metacarpal (Table A12; 

Gaudin et al., 2009), but its relatively short proximal and 

intermediate phalanges give it an appearance more like 

that of modern manids than the patriomanids.

Once again reminiscent of the condition in the pes, 

an apparent increase in the mobility of the metacarpophalangeal joints accompanies the shortening of the manual digits in modern pangolins. At the distal end of the 

metacarpals, extant manids have the same type of robust 

median keel that characterizes the metatarsals described 

above, extending dorsal and ventral to the metacarpal 

shaft (Figure 44f). This keel increases the lateral stability 

of the metacarpal–phalangeal joint as well as its sagittal 

mobility. Like Cryptomanis (Gaudin et al., 2006), Patriomanis has a keel extending only along the ventral and 

palmar surfaces of the distal condyles of the metacarpals, 

and this ventral keel is not as robust as that present in 

the modern species (Figure 27a). Eomanis and Euromanis

resemble the patriomanids is this respect (Gaudin et al., 

2009). In Necromanis, however, the median distal keel, 

although still restricted to the ventral half of the condyle, 

is elongated, thereby more closely resembling the robust 

condition of the keel in modern pangolins (Gaudin et al., 

2009). The dorsal surfaces of the distal condyles of metacarpals II–IV in Cryptomanis are divided by a shallow 

median groove into two “hemicondyles” (Gaudin et al., 

2006).

The metacarpal anatomy of Patriomanis is very reminiscent of the metatarsals in other respects as well. The 

shafts of the metacarpals tend to be relatively wide and 

flat or cylindrical, their minimum midshaft width equal 

to or exceeding their minimum midshaft depth. This mirrors the condition in metacarpals II–V of Cryptomanis, 

where width exceeds depth by an even greater amount 

(Gaudin et al., 2006; Table A12). Modern manids tend to 

have deeper metacarpals, with the midshaft depth exceeding the width for all digits, except in Phataginus and some 

M. javanica, in which the width is greater than or equal 

to the depth for only metacarpal III (Gaudin et al., 2009). 

The distal condyles appear slightly elongate transversely 

and are distinctly narrower than the epicondyles (Figure 

49), in contrast to the condition in the Manidae (Figure 

44f; Gaudin et al., 2009). Immediately proximal to the 

condyles of metacarpals II–V is a midline depression on 

the dorsal surface. This fossa is also present in Cryptomanis and Necromanis but is missing in modern pangolins (Gaudin et al., 2006, 2009). Nevertheless, it can 

be inferred from the myology of living forms that these 

depressions represent insertion sites for the m. extensor 

digitorum longus (Jouffroy, 1966). 

and M. crassicaudata, in contrast to the distally oriented 

facet of Patriomanis, whereas in the extant African taxa 

it has distinct distally and distolaterally facing facets for 

metacarpal V. Additionally, the facet for metacarpal V is 

not tapered at its palmar end in the extant pangolins but is 

of relatively uniform width throughout. 

Metacarpals

The metacarpals are not preserved in the type specimen, and USNM-P 531557 preserves only the distal portion of metacarpal IV. In USNM-P 494439, the first and 

fourth metacarpals from the right manus are intact and 

have been prepared free of the matrix. Metacarpals II 

and III are preserved on the left side but remain partially 

embedded in the matrix ventrally. USNM-P 299960 preserves a complete series on the right (Figure 49). The third 

metacarpal is missing on the left side. 

The third is the largest metacarpal in Patriomanis, in 

terms of its length, width, and depth (Table A12). This is 

also the case among extant manids (except S. temminckii, 

in which metacarpal IV is longest), whereas in Cryptomanis the second metacarpal is longer and equal in depth 

to the third (Table A12; Gaudin et al., 2006). However, 

if the ratio between width and length is considered, it 

becomes evident that the third and sometimes the second 

(Cryptomanis; Gaudin et al., 2006) or fourth metacarpals 

are greatly enlarged in other pangolins (Figure 44f). In 

Patriomanis the digits are relatively uniform in terms of 

metacarpal width versus depth, with values between 20% 

and 22% for all but metacarpal V (29%). In Phataginus, 

M. javanica and M. crassicaudata, and Cryptomanis, the 

ratio of width to depth for the third metacarpal exceeds 

30%, and in Smutsia and M. pentadactyla it exceeds 40%. 

The fifth is the shortest metacarpal in Patriomanis, but the 

first is the narrowest, and the first and fifth are of equivalent midshaft depth (Table A12). There is considerable 

variation among pholidotans regarding the relative sizes 

of the fifth and first metacarpals, although these are invariably the smallest metacarpal elements. In Phataginus, 

Smutsia, and M. javanica, the first metacarpal is shortest, 

whereas in M. pentadactyla and M. crassicaudata, the fifth 

is shorter (Table A12; Gaudin et al., 2009). As was the 

case with the pes, the overall length of the digits relative 

to body size seems to be greater in Patriomanis and Cryptomanis (Gaudin et al., 2006) than in extant manids, as 

reflected in the lengths of the metacarpals. The length of 

the third metacarpal in Patriomanis is 28% that of the 

humerus, whereas in modern pangolins the third metacarpal is 25% or less of humeral length (Table A12). Eomanis 
8 0 • 

crescent shaped, with its long axis oriented basically proximodistally. It is strongly concave, receiving a similarly 

shaped, strongly convex knob from metacarpal III. This 

morphology is similar to that in Cryptomanis, whereas 

in extant manids the articular surfaces between metacarpals II and III are flat (Gaudin et al., 2009). The proximal 

articulation for the trapezoid is triangular in outline, narrowing toward the palmar tuberosity. Its surface contour 

matches that of the distal trapezoid facet described above. 

In dorsal view, the proximal end of the metacarpal II shaft 

in Patriomanis lacks a concavity that is found in Cryptomanis and extant Manidae. 

Metacarpal III. The third metacarpal articulates 

proximally with the capitate, the hamate, and metacarpals II and IV (Figure 49). The articulation with metacarpal II has been described above. The articulation with 

the metacarpal IV is carried on the lateral surface and 

is narrow and generally rectangular in outline, with its 

long axis oriented in a dorsopalmar plane. The facet 

extends from the dorsal edge of the bone nearly to its palmar edge. A facet for the hamate is situated on a lateral 

extension of the proximal end of the bone. This extension slightly overlaps metacarpal IV proximally. The 

overlap is even more pronounced in Cryptomanis and the 

extant African pangolins (Figure 44f) and is also present in Eomanis (Gaudin et al., 2009). In Manis, there 

is no overlap, although a hamate facet is present on the 

third metacarpal. The hamate facet itself is like that of 

most other pangolins, narrow transversely and elongated 

in a dorsopalmar plane, facing proximally and laterally. 

However, in Phataginus and some S. gigantea, the hamate 

facet takes on a very different shape, with a much shorter 

dorsopalmar extent (Gaudin et al., 2009). The proximal 

facet for the capitate in Patriomanis is rectangular in outline, slightly narrower transversely than in the dorsopalmar plane. It has a saddle-shaped surface contour and is

concave transversely, with a strongly convex, semicircular 

profile in medial or lateral view. It is similar in Cryptomanis and modern manids, but in the latter group the 

dorsopalmar profile is sigmoid rather than semicircular 

(Gaudin et al., 2009). The sigmoid profile is created by 

a dorsal shelf that extends forward from the convex, 

palmar portion of the facet. This shelf presumably acts 

as a “stop” to prevent hyperextension at the capitate–

metacarpal II joint and is presumably an adaptation to 

resist large forces generated at this joint when excavating termite nests. It is interesting to note that the stop is 

missing only in S. temminckii, which is the least fossorial of the living pangolins (Kingdon, 1974, 1997; Heath, 

1992b; Nowak, 1999). The capitate facet in some modern 

The central metacarpals (II, III, and IV) of Patriomanis 

differ from those of other pholidotans in two additional 

aspects. In all three, the narrowest portion of the shaft 

in dorsal view lies proximal to the midpoint of the shaft, 

instead of at the midpoint. In addition, the shafts of these 

metacarpals are bowed dorsally, with a concave palmar 

profile. This dorsally bowed shaft can also be observed 

in metacarpals III and IV of Cryptomanis and metacarpal 

II in M. javanica but is otherwise unknown among other 

pholidotans.

Metacarpal I. In dorsal view the shaft of the first 

metatarsal is slightly bowed laterally toward its distal 

end (Figure 49). At its proximal end, metacarpal I articulates exclusively with the trapezium—M. pentadactyla is 

the only pholidotan that we have observed to retain an 

articulation between metacarpals I and II. The outline of 

the proximal articular facet for the trapezium takes on the 

form of an equilateral triangle, with a dorsal and palmar 

point, as well as a strong medial tuberosity. The facet is 

similar in shape in Cryptomanis, but in extant manids 

the medial tuberosity is small or absent, so that the facet 

takes on a more ovate or transversely narrow triangular 

outline. The proximal facet is strongly convex in the dorsopalmar plane, with the trapezium overlapping onto the 

dorsal surface of the metacarpal shaft. At the distal end, 

the condyle for the proximal phalanx is relatively broad in 

comparison to modern pangolins. The condyle is convex 

in a dorsopalmar plane, in contrast to the condition in 

M. crassicaudata and M. pentadactyla, in which the distal condyle has a flattened profile in medial view and the 

metacarpal–phalangeal joint is immobile. These two modern species have similarly immobile joints on digits II, III, 

and IV, whereas in M. javanica only the third metacarpal–

phalangeal joint is immobilized. Other pangolins retain 

mobile metacarpal–phalangeal joints. In all Manis species 

as well as P. tetradactyla, the median keel is absent on the 

metacarpal I condyle, but it is retained in Patriomanis and 

other pangolins.

Metacarpal II. The second metacarpal articulates 

with the trapezium, trapezoid, and metacarpal III at its 

proximal end (Figure 49). The facet for the trapezium is 

carried on the medial surface of the bone and is crescent 

shaped, being elongated in a distodorsal to proximopalmar direction. The facet extends from the dorsal edge of 

the medial surface to approximately the midpoint of the 

medial surface, an arrangement shared in S. gigantea and 

P. tricuspis, although the shape of the facet is more triangular in these taxa. In other extant manids, the facet extends 

almost to the palmar edge of the bone (Gaudin et al., 

2009). On the lateral surface, the facet for metacarpal III is 
n u m b e r 9 8 • 8 1

a large concave pit near its dorsal edge (Gaudin et al., 

2009).

As was the case with metacarpal III, metacarpal IV 

tends to be T shaped proximally in dorsal view in modern 

pangolins, with medial and lateral extensions at its proximal end that abut the adjacent metacarpals, although 

these are less prominent than in metacarpal III (Figure 

44f). These are much less developed in patriomanids relative to extant pangolins, with only a weak extension evident on the lateral side (Figure 49).

Metacarpal V. The lateralmost metacarpal articulates only with the hamate proximally and metacarpal IV 

medially. All pholidotans, including Patriomanis, have a 

lateral tubercle projecting from the lateral margin of this 

bone at its proximal end (Figures 44f, 49), although it 

should be noted that this feature is present in early eutherians (Kielan-Jaworowska et al., 2004) and is thus likely 

a primitive feature for all placentals. In African manids, 

this tubercle appears to be displaced proximally relative to 

the other pholidotans, so that it comes to lie proximal to 

the distal edge for the hamate. In these taxa, the metacarpal V–hamate articulation is L shaped, with a vertical and 

horizontal moiety (Gaudin et al., 2009, describe this as a 

“peg-and-socket” articulation). In Patriomanis and Manis, 

the lateral tubercle lies slightly distal to the distal edge 

of the hamate, and the proximal facet on metacarpal V 

is either flat (Manis) or slightly convex in a dorsopalmar 

direction (Patriomanis). At the distal end of the bone, 

the median keel is reduced or absent in Manis, but it is 

retained in Patriomanis, Cryptomanis, and the extant 

African species.

Manual Proximal Phalanges

Emry (1970: fig. 20) illustrates and briefly describes 

two manual proximal phalanges recovered from the type 

specimen. Although he does not specify, these are the 

proximal phalanges from the second and third digits, the 

former the longer of the two. In addition, proximal phalanx IV is now known from the type specimen. Manual 

proximal phalanges I and II are preserved and have been 

prepared in the right manus of USNM-P 494439, and the 

left pes preserves proximal phalanges II–V still embedded 

in matrix. USNM-P 299960 includes a complete series of 

proximal phalanges from the left and right manus (Figure 

49), except for digit III on the right. The following description is based largely on the complete set of elements available from the latter specimen. 

The relative lengths of the proximal phalanges are 

variable among pholidotans. The fifth proximal phalanx 

pangolins (Smutsia, M. crassicaudata, and M. pentadactyla; Gaudin et al., 2009) extends onto the palmar surface 

of the metacarpal shaft, allowing for increased plantar 

flexion at this joint. Such movement is prevented by the 

large palmar tuberosity of the metacarpal in Patriomanis, 

Cryptomanis, and other modern pangolins.

The proximal shaft of metacarpal III in modern 

manids has a distinctive T shape in dorsal view, with not 

only the aforementioned lateral extension abutting or 

overlapping metacarpal IV at its proximal end but also a 

corresponding medial extension contacting metacarpal II 

(Figure 44f). This second extension is missing in both 

Patriomanis (Figure 49) and Cryptomanis (Gaudin et al., 

2006). Patriomanis bears a small extensor tubercle on the 

medial side of the shaft of metacarpal III, situated just 

distal to the midshaft region. A similar tubercle is found 

on the lateral side of the second metacarpal. Cryptomanis manifests a similar morphology (Gaudin et al., 2009), 

but extensor tubercles are largely lacking in Eomanis and 

modern pangolins. However, in M. crassicaudata and

M. pentadactyla, the dorsal surface of metacarpal III is 

marked by a median crest that extends the entire length 

of the shaft. This crest bears a prominent tubercle at its 

proximal terminus (Gaudin et al., 2009). 

Metacarpal IV. This element articulates with 

the hamate and with metacarpals III and V proximally. 

The facet for metacarpal III in Patriomanis is rectangular, elongate in a dorsopalmar plane. It is slightly convex 

proximodistally and nearly flat to slightly concave in its 

dorsopalmar aspect. It is similar in Smutsia and Phataginus, although in the former the convexity tends to be better developed. In Manis, it is strongly convex in both the 

proximodistal and dorsopalmar aspects (Gaudin et al., 

2009). The facet for metacarpal V shows extensive shape 

variation among the pholidotans in which it is known. 

In Patriomanis it is teardrop shaped, with its long axis 

running in a dorsodistal to proximopalmar direction. The 

facet is concave in Patriomanis and Cryptomanis, whereas 

in most modern pangolins (except S. gigantea) it is convex 

at least in the dorsopalmar plane (Gaudin et al., 2009). 

The hamate facet is relatively simple in Patriomanis, as it 

is in Cryptomanis. It is rectangular in outline, is narrowest 

transversely, and is weakly convex in both the transverse 

and dorsopalmar planes. The facet varies considerably 

among extant Manidae. In African pangolins and M. 

javanica, it is triangular in outline, narrowing toward the 

palmar margin. In M. crassicaudata and M. pentadactyla, it takes on a semicircular shape, with the dorsal edge 

forming the base of the semicircle. Additionally, in all the 

extant species, the facet, although generally convex, bears 
8 2 • 

and IV, the proximal phalanges are slightly asymmetrical, 

the latter extending farther proximally on its medial side, 

the former extending farther proximally on its lateral side. 

Proximal phalanx III is marked in most extant pangolins 

(all but M. javanica; Gaudin et al., 2009) by a narrow 

transverse eminence that links the two distal condyles at 

their dorsopalmar midpoints. This eminence is lacking in 

patriomanids (Figure 26c).

Manual Intermediate Phalanges

There are no manual intermediate phalanges preserved 

in the type (Emry, 1970), but all five are preserved in the 

matrix surrounding the left manus of USNM-P 494439, 

along with several free elements from the right side. A 

complete right and left series is also available in USNM-P 

299960 (except for the left intermediate phalanx IV; Figure 49). The description that follows is based largely on 

that specimen. 

As was the case with the proximal phalanges, the 

manual intermediate phalanges are cylindrical in dorsal 

view, slightly constricted in the middle, and expanded at 

either end. In USNM-P 299960, the second is the longest, 

whereas in USNM-P 494439 the third is longest, as it is in 

all extant pangolin species. The fifth intermediate phalanx 

is the shortest in all pangolins except P. tricuspis, in which 

the second is marginally shorter than the fifth (Table A13). 

The manual intermediate phalanges are roughly equivalent in length to the proximal phalanges in Patriomanis 

(Table A13). In most instances, the intermediate phalanx 

is much longer (up to twice as long; Table A13) than the 

corresponding proximal element in living pangolins. This 

morphology is also evident in digit III of Cryptomanis 

(Gaudin et al., 2006: fig. 9). This appears to be due largely 

to the dramatic shortening and widening of the proximal 

phalanges that is so pronounced in Cryptomanis, Smutsia, and several Asian species, as described above. In these 

taxa, the proximal phalanges are often compressed proximodistally to the point that they no longer have a cylindrical shaft. 

The proximal and distal articular surfaces are basically identical to those described above for the pedal intermediate phalanges. As in those elements, the proximal end 

comprises two shallow articular fossae for the proximal 

phalangeal condyles and dorsal and paired ventral processes that embrace the interphalangeal joint, although to 

a lesser extent than in modern forms. The dorsal processes 

are particularly elongated proximally in extant manids, 

so that they overshadow the proximal articular surface in 

dorsal view. The distal condyles are paired, subequal, and 

is the shortest in all extant species except M. tetradactyla 

(proximal phalanx I is slightly shorter) and Patriomanis, 

whereas in Cryptomanis the third is the shortest (Table

A13). The longest is more variable. In Patriomanis, the 

second is generally longest, as it is in M. crassicaudata and 

M. pentadactyla. The third is longest in the extant African species and M. javanica, whereas the first is longest in 

Cryptomanis (Table A13). 

The proximal phalanges of Patriomanis have a 

roughly cylindrical shaft that is constricted in the middle 

and expanded at either end in dorsal view, although in digits III and IV the elements are very broad, with a proximal 

width nearly 80% of their maximum length. In Cryptomanis, proximal phalanges II–IV are similarly short and 

broad, their width actually equaling or exceeding their 

length. Likewise, dramatic shortening and broadening of 

the proximal phalanges occurs in the extant genus Smutsia 

(digits II–V) and the extant species M. crassicaudata and

M. pentadactyla (digits III–V; Table A13). As in Cryptomanis, Euromanis, and Eomanis, the width of the proximal phalanges is equal to or exceeds their depth, whereas 

in extant manids these bones are uniformly deepened, 

so that their dorsopalmar depth exceeds their transverse 

width (Gaudin et al., 2009).

At their proximal ends, the manual proximal phalanges resemble their pedal counterparts. They have a concave proximal facet that is divided ventrally by a median 

depression that in turn receives the keel of the distal metatarsals, giving the articular surface a horseshoe-like shape 

(Figure 26a). A similar morphology is present in Cryptomanis, whereas in extant species the entire articular surface is deeply divided by a median groove for the enlarged 

metatarsal keel. The absence of this keel in Patriomanis 

and other fossil pangolins (Gaudin et al., 2006, 2009) 

leaves the metatarsal–phalangeal joints appearing somewhat less stable than the interphalangeal joints. 

At the distal ends, the proximal phalanges of Patriomanis are capped by a pair of subequal, pulley-shaped distal condyles, as in all other pangolins (Figure 26c). Lateral 

to each of these condyles are well-marked fossae (Figure 

49), likely for the origin of collateral ligaments bracing the 

interphalangeal joints (based on anatomy of humans and 

dogs; Pick and Howden, 1977; Evans and Christensen, 

1979). These fossae are also present in Cryptomanis and 

Necromanis but are missing in Eomanis and extant pangolins (Figure 44f; Gaudin et al., 2009).

As was the case in the pes, the first manual proximal 

phalanx is more elongate and slender than the other proximal phalanges (Table A13). This is true not just of Patriomanis but of all living and extinct pangolins. On digits II 
n u m b e r 9 8 • 8 3

to their pedal unguals, nor are waisted manual unguals 

found in other extant manids (Gaudin et al., 2009). 

The proximal articular surface is as Emry (1970) 

described it and as it is described above for the pedal 

unguals. It has two strong concavities for medial and lateral distal condyles of the intermediate phalanges. The 

median crest on the subungual process is enlarged relative 

to that of the pedal unguals, taking the form of a large 

tubercle. In other respects, the subungual process and vascular foramina follow the pattern described above for the 

pedal unguals. 

DISCUSSION

Species Comparisons Based on Size and Strati

To date, all the specimens of Patriomanis have been 

assigned at least implicitly to a single species, Patriomanis 

americana. Given that the holotype specimen is relatively 

incomplete (Emry, 1970) and subsequent investigations 

of Patriomanis have tended to focus only on a portion of 

the anatomy, either the skull (Gaudin and Wible, 1999; 

Emry, 2004) or other relatively circumscribed portions of 

the skeleton (Rose and Emry, 1993), this approach has 

made sense. However, now that a complete assessment has 

been made of the available material, it is reasonable to ask 

whether more than one species can be recognized. 

There do not appear to be geographic grounds for 

recognizing more than one species. Patriomanis is known 

from only two locales, the Pipestone Springs Local Fauna 

in the Jefferson River Basin of southwestern Montana 

(USNM-P 460256) and the Flagstaff Rim area in Natrona 

County in central Wyoming (F:AM 78999 [holotype]; 

USNM-P 299960, 494439, 531556, 531557; see the Catalog of Specimens section of the Introduction). The two are 

roughly 500 km apart, certainly close enough to fall in the 

same species range if compared to the ranges of the extant 

forms (see, e.g., Kingdon, 1997). Moreover, the Pipestone 

Springs specimen is assigned to Chron 3 of the Chadronian NALMA (Janis et al., 2008; Rose, 2008), which is 

the case with all but one of the Flagstaff Rim specimens.

There are several positive arguments, however, that 

could be advanced for recognizing USNM-P 299960 as a 

second species relative to the other five specimens. USNMP 299960 derives from Emry’s (1973) Unit 4 of the White 

River Formation in the Flagstaff Rim area. The other 

specimens at that locality derive from Units 17 and 21 

(see the Catalog of Specimens section). Janis et al. (2008) 

assign all the strata below Ash B at the Flagstaff Rim 

strongly convex, flanked by pits for the interphalangeal 

collateral ligaments that are visible in dorsal view (Figure 

49), in contrast to the condition in extant taxa, in which 

the distal condyles are parallel and the fossae for the collateral ligaments are oriented laterally (Figure 44f). 

Manual Unguals

Emry (1970) described a single manual ungual in the 

type specimen of Patriomanis (F:AM 78999). Since that 

report, three additional unguals have been identified in the 

type, so that unguals from digits I, II, III, and V are now 

known. Manual unguals I, II, IV, and V have been prepared free from the right manus of USNM-P 494439, and 

manual unguals II and IV on the left are partially exposed 

in the matrix. A virtually complete set of left and right 

manual unguals has been preserved with USNM-P 299960 

(Figure 49); hence, the description that follows is based 

largely on that specimen. 

The third ungual is the largest, with the first and fifth 

nearly equivalent in size and smaller than the remaining 

unguals (Table A13). Among modern pangolins, the first 

ungual phalanx is notably smaller than the fifth in Manis 

and Smutsia and is dramatically reduced in Phataginus, in 

which its length is less than half that of the fifth (Figure 

44f; Gaudin et al., 2009). Although ungual phalanx III 

is the largest manual ungual in Patriomanis, its length is 

only about 10% greater than the next largest ungual, so 

that overall, the unguals are of a fairly uniform length, 

as in the pes (Figure 49). However, in Eomanis, Manis, 

and Phataginus, the third ungual is greatly enlarged, being 

at least 30% longer than the next largest element (Figure 

44f; Gaudin et al., 2009). In Smutsia, the second, third, 

and fourth unguals are all quite large. Consequently, the 

middle digit does not appear to be emphasized to the same 

degree 

The anatomy of the manual unguals is virtually identical to that described above for the pedal unguals. They 

carry the characteristic deep fissures found in other extinct 

and extant pholidotans (Grassé, 1955; Guth, 1958; Gebo 

and Rasmussen, 1985; Koenigswald, 1969, 1999; Koenigswald and Martin, 1990; Gaudin et al., 2006, 2009) with 

the exception of Eomanis and Euromanis (Storch, 1978; 

Storch and Martin, 1994). As in the pes, the fissure occupies a little more than half the length of the ungual, as 

in other eupholidotans. The unguals are blunt tipped and 

waisted. Necromanis (Koenigswald, 1969, 1999; Koenigswald and Martin, 1990), Cryptomanis (Gaudin et al., 

2006), and Manis pentadactyla also have waisted manual 

unguals, but Eomanis and Euromanis do not, in contrast 
8 4 • 

or even sexual dimorphism, as male pangolins are known 

to be larger than females (Heath, 1992a, 1992b; Nowak, 

1999). Moreover, it is difficult to know what the expected 

magnitude of interspecific size differences might be on the 

basis of comparisons with extant taxa. In two of the three 

modern genera, Phataginus (Kingdon, 1997) and Manis

(Heath, 1992a, 1995), the species broadly overlap in body 

size, and in Smutsia the giant pangolin is nearly twice the 

size of the ground pangolin (Kingdon, 1997). In the end, 

there are not enough specimens of Patriomanis to reliably 

discriminate among the various possibilities; indeed, distinguishing among individual differences, species-level differences, and gender-based differences remains a difficult 

problem for extinct taxa in general (see discussion in Pujos 

et al., 2012).

In the same stratigraphic interval occupied by Patriomanis (from just below Ash A to just below Ash G), 

the small ungulate artiodactyl Leptomeryx also shows a 

distinct increase in size (Emry, 1973: fig. 18; Emry and 

Gawne, 1986: fig. 5; Heaton and Emry, 1996: figs. 2–4, 6). 

However, unlike Patriomanis, Leptomeryx is represented 

by very large samples, which conclusively show that 

the increase in size of Leptomeryx from lower to higher 

stratigraphic levels is an evolutionary increase, with the 

whole population becoming larger, and not intrapopulation variation. This evidence suggests that the increase 

in size of Patriomanis through the same interval might 

also be evolutionary change rather than intrapopulation 

variation. The authors of the Leptomeryx studies faced 

the same question that we are faced with here regarding 

Patriomanis: whether to recognize these successive populations as distinct chronospecies when they are separated 

only by time and size and lack other obvious morphologic 

differences. Those authors declined to name new species 

of Leptomeryx, just as we decline to name a new species 

of Patriomanis here. 

In an attempt to get a different perspective on morphological variation among the various Patriomanis 

specimens, we looked at character polymorphism in the 

Gaudin et al. (2009) phylogenetic analysis. The number of characters that were coded as polymorphic for 

extant pangolin species in that analysis ranged from 7 

(M. crassicaudata) to 56 (S. gigantea). There did appear 

to be some correlation between polymorphisms and the 

number of specimens available for study. For example, 

the 305 postcranial characters in the study (out of a 

total of 395 characters) were coded in M. crassicaudata on the basis of only two specimens. It is noteworthy, then, that although Patriomanis was coded on the 

basis of more specimens than M. crassicaudata for the 

site, including Unit 4, to Chron 2/3 of the Chadronian 

NALMA, and those levels between Ash B and F, including Unit 17, to Chron 3, with the two specimens from 

Unit 21 (holotype, F:AM 78999, and USNM-P 4894439) 

found only 5 m above Ash F, which would presumably 

also be in Chron 3 or, at the latest, the base of Chron 4. 

At first glance the clustering of all the Patriomanis in or 

around Chron 3 might further confirm the idea that they 

can be lumped into the same species. However, USNMP 299960 sits some 45–46 m below the level containing 

USNM-P 531556, which in turn lies another 35 m below 

the holotype and USNM-P 494439 (Emry, 1973). Chrons 

2 and 3 together span a period of roughly 2 million years, 

from 36.6 to 34.8 mya (Janis et al., 2008; Rose, 2008). 

We think it unlikely that the Patriomanis remains from the 

Flagstaff Rim area span the entire range from the base of 

Chron 2 to the top of Chron 3, but it is not unreasonable 

to assume that either USNM-P 299960 comes from Chron 

2 and the others from Chron 3 or at least that USNM-P 

299960 comes from the base of Chron 3 and the others 

from substantially later in Chron 3 or even early Chron 4. 

Since Chron 3 itself is 1 million years in duration, it seems 

likely that the 80-m stratigraphic interval between Units 4 

and 21 represents a time period of about 1 million years. 

This would certainly be enough time to recognize USNMP 299960 as a chronospecies distinct from the type and 

USNM-P 494439, with the status of USNM-P 531556 less 

clear but closer to the latter than the former.

There is also some morphological support for recognizing USNM-P 299960 as a separate species. It is the 

smallest specimen of Patriomanis. There is only one complete skeletal element, the radius, held in common among 

USNM-P 299960, 531556, and 494439, the three specimens that define the stratigraphic range of Patriomanis at 

Flagstaff Rim. The greatest length of the radius is 48.9 mm 

in USNM-P 299960, 52.9 mm in USNM-P 531556, and 

53.5 mm in USNM-P 494439 (Table A10), the last being 

9% longer than the first, and the three specimens show 

an increase in size (at least for this element) through time. 

Although USNM-P 531556 preserves completely only the 

humerus (which is larger than USNM-P 299960, 63.4 

versus 61.1 mm; Table A10), radius, and metatarsal V, 

more extensive comparisons can be carried out between 

USNM-P 299960 and USNM-P 494439, including the 

calcaneus, astragalus, most of the elements in the manus, 

the tibia, and the ulna. In every instance, the bones from 

USNM-P 494439 are between 8% and 29% larger than 

their counterparts in USNM-P 299960, with most being 

15%–25% larger. Of course, it is possible that these size 

differences represent random interindividual size variation 
n u m b e r 9 8 • 8 5

et al. (2009), the two together constituting a clade identified as the superfamily Manoidea. Eomanidae is in turn 

the sister taxon to this clade, forming a more inclusive 

clade called Eupholidota.

Gaudin et al. (2009) assert that the order Pholidota 

likely has its biogeographic origins in Europe, with the 

oldest pangolins (Eomanis, Euromanis, and Eurotamandua) coming from the middle Eocene (ca. 45 mya) Messel 

deposits of Germany. The origin of Pholidotamorpha is 

less clear, as palaeanodonts have a North American origin 

(Rose and Lucas, 2000), as does the extant sister taxon 

to Pholidota, the Carnivora (Flynn and Wesley-Hunt, 

2005), leaving the origin of Pholidotamorpha equivocal, 

although almost certainly located somewhere in Laurasia. 

It would appear that Patriomanis arrived in North America from the west, most likely over the Bering land bridge, 

as it is linked to the European birthplace of the order by 

the slightly older, late Eocene (ca. 40 mya) pangolin from 

northern China, Cryptomanis (Gaudin et al., 2006). This 

further suggests that pangolins may have been distributed 

across all the Laurasian continents in the latter half of the 

Eocene epoch, although they are known to persist only 

in Europe after the Eocene, in the form of the OligoceneMiocene genus Necromanis.

Paleobiolog

As has been noted elsewhere (Gaudin et al., 2006, 

2009), Patriomanis is quite similar anatomically to the 

slightly older patriomanid from northern China, Cryptomanis. Although it is generally smaller bodied and less 

robust in its skeletal construction, Patriomanis has many 

of the features presented as indicative of a fossorial lifestyle in Cryptomanis (Gaudin et al., 2006). Like the former, Patriomanis has a large gluteal fossa on the ilium and 

a large femoral greater trochanter and third trochanter 

(located somewhat more distad on the femoral midshaft 

region) with a robust intertrochanteric ridge, although the 

lesser trochanter is smaller and the femoral shaft is more 

cylindrical. The tibia has a prominent cnemial crest that 

is slightly longer but less robust than that of Cryptomanis; the fibula, with its cylindrical shaft, is sturdy but less 

robust than in Cryptomanis, and the patella is less massive. Like Cryptomanis, Patriomanis has a well-developed, 

fairly elongate olecranon on the ulna, but it is less flared 

along its proximal posterior edge. The humerus has a 

very strong deltopectoral crest that is longer but lower 

than that of Cryptomanis. The entepicondylar process is 

massive but less elongate than in Cryptomanis, and the 

supinator crest is well developed but not as prominent, 

postcranial data, only 10 polymorphisms were found. Of 

these, five were “false” polymorphisms, in the sense that 

the codings were not based on actual variation among 

specimens, but rather on the fact that the fossil specimens 

provided enough information to rule out one or more 

states but not enough to assign a definitive coding among 

remaining states. Thus, the various specimens of Patriomanis show very few morphological differences from one 

another, apart from size, when compared to modern species. This morphological uniformity is a strong argument 

against recognizing separate species on morphological 

grounds.

In summary, although there are reasonable grounds 

for recognizing USNM-P 299960 as a separate chronospecies from the other Patriomanis specimens on the 

basis of stratigraphic data, there is much less support 

for recognizing it on morphological grounds. The available specimens of Patriomanis are morphologically quite 

uniform, apart from a clear size difference between the 

geologically older, smaller USNM-P 299960 and the 

geologically younger, larger USNM-P 494439, perhaps 

with USNM-P 531556 intermediate in both age and size. 

There are not enough specimens, however, to establish 

this as a real size trend, as opposed to random individual 

variation or some form of sexual dimorphism. Therefore, 

until additional specimens are recovered, we choose to 

retain all the specimens in the single species Patriomanis 

americana.

Phylogeny, Taxonomy, and Biogeographic O

These aspects of the biology of Patriomanis have 

been thoroughly investigated and discussed by Gaudin 

et al. (2009), so we will offer no more than a summary of 

their results in the present study. Patriomanis is assigned 

by Gaudin et al. (2009) to the superorder Pholidotamorpha, which in turn encompasses two orders, Pholidota 

and Palaeanodonta. Pholidota is divided into three families, Eomanidae, Patriomanidae, and Manidae, with the 

genera Eurotamandua and Euromanis included in the 

order but not assigned to a particular family. Eomanidae 

is monotypic, accommodating only the genus Eomanis, 

whereas Manidae includes all the extant pangolins. The 

family Patriomanidae, to which, of course, Patriomanis is 

assigned, also includes the extinct Asian pangolin Cryptomanis, and Hoffmann (2011) suggests that the midCenozoic European pangolin Necromanis should also be 

assigned to this family, although its familial affinities are 

not resolved in the Gaudin et al. (2009) study. The Patriomanidae represents the sister taxon to Manidae in Gaudin 
8 6 • 

extended nearly to the end of the caudal series, although 

they lacked the large transverse processes present in the 

terminal caudals of Phataginus. However, the morphology 

of the terminal caudals in Patriomanis was very similar to 

that of nonpholidotan extant mammals with prehensile 

tails, including vermilinguans, opossums, primates, and 

carnivorans (see caudal vertebrae description above). 

The prehensile tail demonstrates that Patriomanis was 

almost certainly an arboreal forest dweller. Additionally, 

it may have climbed in a different manner from its close 

relative Cryptomanis, with its longer digits but lack of a 

prehensile tail.

Unlike Cryptomanis, Patriomanis is represented by 

well-preserved skull material, including an edentulous rostrum and mandible (Emry, 1970, 2004). This morphology 

is a clear indication that like the modern forms, Patriomanis was myrmecophagous, feeding primarily on ants 

and termites. The older pangolin genus Eomanis is also

edentulous and was also presumably myrmecophagous, 

although its preserved stomach contents contain more sand 

and plant remains than insect cuticle (Koenigswald et al., 

1981). Like the living pangolins, the fossil forms lack both 

temporal lines and a frontal postorbital process (Gaudin 

et al., 2009) and therefore likely had the strongly reduced 

temporal fossa and concomitantly reduced temporalis 

muscle of the extant pangolins (note that in Eurotamandua temporal lines are also absent, but muscle scars on the 

braincase indicate a somewhat larger temporalis; Gaudin 

et al., 2009). However, both Patriomanis and Eomanis

(Storch, 1978) preserve at least a small coronoid and angular process on the mandible, features missing in the modern forms. Moreover, the horizontal ramus of the mandible 

is somewhat more robust in Patriomanis than in the living taxa. We interpret this to mean that Patriomanis and 

Eomanis had somewhat better developed masticatory musculature and likely a wider gape than is the case in living 

pangolins. 

The only fossil pangolin that preserves any portion of 

the xiphisternum is Cryptomanis, and it appears to possess the same elongated osseous xiphisternum found in the 

modern taxa, although, of course, we cannot say for certain whether it possessed the elongate cartilaginous extension found in extant forms (Grassé, 1955; Kingdon, 1974) 

or whether the tongue musculature attached to the xiphisternum or indeed any part of the sternum as in the modern taxa (Chan, 1995). In modern forms, this elongation 

of the tongue posteriorly facilitates the great protrusibility 

of the tongue during feeding.

Finally, we might make mention of the epidermal 

scales. Of course, the most recognizable feature of living 

especially at its proximal end. Patriomanis also has a very 

tall scapular spine, much better developed than in living manids (the condition is unknown in Cryptomanis; 

Gaudin et al., 2009).

As in Cryptomanis, all these features are more robust 

in Patriomanis than in the extant forms, whereas the 

digits are considerably less robust than in the modern 

taxa, lacking the deeply keeled metapodials and greatly 

enlarged manual third digit and claw of the living manids. 

Indeed, the manual third digit of Patriomanis is weaker 

than that of Cryptomanis. Thus, both patriomanids share 

the characteristic of having more fossorially adapted 

proximal limb skeletons, that is, in the girdles and long 

bones, and less fossorially adapted distal limb skeletons 

in the manus and pes. Gaudin et al. (2006) suggest that 

there is a progressive transfer of digging adaptations from 

the proximal end of the limb to the distal end in pangolin evolution. This may be the case, but Patriomanidae is 

the sister taxon to Manidae and not necessarily its direct 

ancestor. Moreover, Eomanis, an older taxon and the sister taxon to both Manidae and Patriomanidae, is in some 

respects more like the modern manids than are patriomanids, with a weak gluteal fossa and third trochanter 

on the femur (albeit at the proximal rather than distal 

end) and a larger third manual ungual (Gaudin et al., 

2009: chars. 128, 146, 303). It is therefore possible that 

the patriomanids are simply divergently specialized for 

digging relative to the living pangolins. This possibility 

would be consistent with Gaudin et al.’s (2006) suggestion that perhaps patriomanids dug in a different manner 

than the living taxa.

Cryptomanis is described as having a scansorial, semiarboreal mode of locomotion, lacking the prehensile tail 

of the arboreal African tree pangolins but with elongate 

digits, especially in the pes, that would have provided 

some grasping ability, although there is no evidence for 

opposability in the digits (Gaudin et al., 2006). The digits 

in Patriomanis are both absolutely and relatively shorter 

than those of Cryptomanis, but the tail is considerably 

longer. In fact, the evidence suggests that, unlike Cryptomanis, Patriomanis had a prehensile tail (see caudal vertebrae description above). It had a caudal vertebral count 

(31–36) commensurate with that of the modern prehensile-tailed African tree pangolin Phataginus tricuspis (35–

39) and greater than that of any living pangolin that lacks 

a prehensile tail. In addition, it had very large transverse 

processes in the anterior caudal vertebrae, very large and 

elaborate double transverse processes in the mid-caudals 

reminiscent of those in prehensile-tailed vermilinguan 

anteaters, and ventral articulations for chevrons that 
n u m b e r 9 8 • 8 7

epoch was the warmest period of the Cenozoic era, with 

the Paleocene-Eocene Thermal Maximum (PETM) at the 

very beginning of this epoch representing the warmest 

single point in the past 65 million years. As noted by Blois 

and Hadly (2009:198), the PETM was characterized by 

the “lowest equator-to-pole temperature gradient” and 

“the greatest extent of tropical and subtropical forests 

ever achieved.” These conditions persisted throughout 

the early Eocene, creating the kinds of habitats in northern Europe, Asia, and North America in which pangolins 

would certainly have thrived. However, from the beginning of the middle Eocene onward, there was a steady 

decline in global temperatures, culminating in a drastic 

drop known as the Terminal Eocene Event, when the 

northern climates became substantially drier and cooler 

(although still warmer than the present-day climate). In 

the late Cenozoic of western North America in particular, the tropical and subtropical forests of the Eocene were 

beginning to transition to the more open, grassy habits 

of the Oligocene epoch (Wing and Sues, 1992; Prothero, 

1994; Blois and Hadley, 2009). 

The fossil record of pangolins seems an ill fit in some 

regards when compared to this background of climate 

and vegetational change. As noted above, the oldest pangolins do not appear in the fossil record until the middle 

Eocene of Europe (Eomanis, Euromanis, Eurotamandua; 

Storch, 1978, 1981; Storch and Martin, 1994; Gaudin 

et al., 2009), when forests were still widespread in northern Laurasia but climatic deterioration was already well 

underway (Wing and Sues, 1992; Prothero, 1994; Blois 

and Hadley, 2009). The climate continued to become 

cooler and drier almost continually as, first, Cryptomanis 

made its appearance in the late Eocene of northern China 

(Gaudin et al., 2006) and, finally, Patriomanis made its 

appearance in North America in the latest Eocene (Rose, 

2008). One might imagine that the warm tropical climes 

of the early Eocene would have been much more hospitable for the origin and spread of early pangolins than the 

middle and especially the late Eocene. 

We offer two potential explanations for this apparent 

discrepancy between pangolin history and Eocene climatic 

history. Let us note at the outset that we do not imagine 

these to be the only possible explanations; although these 

appear plausible to us, other scenarios or contributing factors could certainly be imagined. That said, one possibility is to explain the discrepancy as the result of historical 

constraint. If the origin of the order is close to the age of 

its oldest fossils in the middle Eocene approximately 45 

mya, then the arrival of Patriomanis in North America 

occurs only 9–10 million years later. This is a long time, 

pangolins is their external covering of large, sharp-edged, 

overlapping scales that serve as an important defense mechanism, both from predators and from the often aggressive 

social insects that they feed upon (Kingdon, 1974, 1997; 

Heath, 1992a, 1992b, 1995; Nowak, 1999). Kingdon 

(1997) has speculated that the scaly covering evolved from 

an ancestral pangolin that had smaller scales only on its 

prehensile tail, like a modern opossum or murid or anomalurid rodents. From there the scales enlarged and spread 

to cover the dorsum, flanks, and limbs of the modern taxa. 

There is only one fossil pangolin for which scales are preserved, Eomanis waldi, from the fossil lagerstätten at Messel, but they are preserved as several isolated elements and 

reveal nothing about the extent of the scaly covering of 

the animal (Koenigswald et al., 1981). Storch (1978: plate 

5, fig. 2) reconstructs the scaly covering much like that of 

the modern forms, covering the dorsum, flanks, and top 

of the head, although not covering the distal two-thirds of 

the tail or the distal portions of the limbs. In our reconstruction of Patriomanis (frontispiece), we have chosen 

to reconstruct the scales as Storch (1978) did for Eomanis, but in line with Kingdon’s (1997) suggestion we have 

drawn a scale covering for the entire tail. We have also 

reconstructed more elongate pinnae for the ears and more 

and longer hair between the scales in Patriomanis than 

are found in any modern pangolin species. The longer 

ears and hair were intended as a kind of external reflection of its substantially more primitive skeletal anatomy, 

thus befitting an animal of its advanced geological age. It 

should be recognized, however, that all such reconstructions of the external anatomy are largely conjectural for 

the extinct pangolins. 

Paleoecology, Climate, and Geographic Distri

One of the curious aspects of the appearance of Patriomanis in the latest Eocene of North America is how the 

timing of its arrival relates to the paleoecology of this 

region. Modern pangolins are tropical creatures, their 

present distribution almost entirely encompassed within 

the belt between the Tropics of Cancer and Capricorn in 

Africa and South Asia (Macdonald, 2006). All but one of 

the extant species are forest dwellers, with only the African ground pangolin Smutsia temminckii habitually found 

in more open savannahs and grasslands (Kingdon, 1974, 

1997; Heath, 1992a, 1992b, 1995; Nowak, 1999). Certainly, no modern pangolin could survive in the cool, dry, 

continental climes and open, scrubby habitat of today’s 

Wyoming. However, the world of the Eocene was dramatically different from the modern world. The Eocene 
8 8 • 

Eocene (and Paleocene?) than is currently indicated by 

known fossils but that this results primarily from the 

incompleteness of their fossil record. The latter could be 

due to the difficulty in preserving, finding, and recognizing fossil pangolins. Patterson (1978) asserts that pangolins are less than ideal candidates for preservation, given 

their solitary lifestyle, preference for forested habitats, 

and lack of teeth, the most durable and readily recognizable part of mammalian skeletons. With the loss of teeth 

comes also a thinner and presumably less durable skull 

than that of comparably sized mammals. Pickford and 

Senut (1991) have also noted that mammalian paleontologists may overlook pangolin remains in the field or in the 

lab because of their edentulous nature and the difficulty of 

identifying fragmentary skulls or even well-preserved but 

isolated postcranial remains. This hypothesis is tantalizing to pangolin enthusiasts (an admittedly small group) 

because it suggests that more fossils remain to be found in 

Eocene, and perhaps even Paleocene, deposits in Europe, 

northern Asia, and North America. Nothing would form a 

more fitting ending for this monograph than that it should 

lead to the discovery of new fossil pangolins that will augment our understanding of this small but interesting and 

unusual group of mammals.

but given the limitations noted in the Introduction—slow 

locomotory speeds, low reproductive rate, and low population densities, along with the apparently limited dispersal abilities of myrmecophagous mammals in general—it 

does not seem to be an unreasonable time lag to account 

for dispersal half a planet away. However, recent phylogenetic analyses postulate a much earlier origin for the 

pholidotan lineage. These analyses suggest an origin either 

in the vicinity of the Paleocene–Eocene boundary or in the 

early Paleocene. The former, from Meredith et al. (2011), 

is based on their estimate for the age of Carnivora, the 

extant sister group to Pholidota (their published estimate 

for Pholidota is much younger [25 mya, late Oligocene] 

but refers only to the origin of Manidae; note that their 

figs. 1 and 2 suggest that Ferae [= Carnivora + Pholidota] 

may have an origin in the Paleocene or even the Cretaceous). The latter, from O’Leary et al. (2013), is based 

on ghost lineage analysis and the age of the extinct sister taxon to Pholidota, Palaeanodonta. The extra 10 or 

20 million years provided for the origin of Pholidota by 

these hypotheses make the historical constraint hypothesis 

much less compelling. 

Another possibility is that pangolins existed over 

much broader swaths of Laurasia much earlier in the 
Appendix: Tables of Measurements
9 0 • 

TABLE A2. Measurements of the atlas and axis in Patriomanis americana and living pangolin

Measurement

Patriomanis 

americana†, 

F:AM 78999, 

USNM-P 2999

Manis 

pentadactyla,

USNM 3087

Phataginus 

tricuspis,

USNM 5377

Smutsia 

gigantea,

AMNH 5384

Atlas

Maximum transverse width, excluding transverse process (mm) 22.4a 26.1 21.7 41.1

Maximum dorsoventral height (mm) 16.4a 17.4 13.0 24.2

Ratio of height versus width 0.73 0.67 0.60 0.59

Maximum anteroposterior length, excluding transverse process (mm) 14.4a 14.4 9.1 26.1

Width versus length 1.50 1.90 2.40 1.60

Axis

Maximum anteroposterior length, including dens (mm) 19.2b 15 8.9 22.3

Maximum transverse width, measured just behind anterior 

articular facets (mm)

16.7b 19.1 14.2 27.1

Length versus width 1.1 0.79 0.63 0.82

†Extinct.

aBased on only F:AM 78999. 

bBased on only USNM-P 299960.

TABLE A1. Measurements of the skull and mandible in Patriomanis americana and related living and extinct (†) pangolins. A dash (—

indicates data are not available. GSL = greatest skull length; MML = maximum mandibular length.

Measurement

Patriomanis 

americana†, 

F:AM 78999 

(type); USNM-

494439

Manis 

pentadactyla, 

FMNH 3251

Phataginus 

tricuspis, 

USNM 537785

CM 86715

Smutsia 

gigantea, 

AMNH 5385

Eomanis 

waldi†, 

SMF MEA 263

Pohl specimen

Skull

GSL (mm) 92a 89.6 68.6,b 69.6c 149.5 64.4d

Nasal length (mm) — 28.10 24.1c 55.40 26d

Length of parietal-squamosal suture (mm) 27.1e 21.1 22.5c 32.2 13.5d

Length of parietal-squamosal suture versus GSL ~0.3 0.24 0.30 0.22 0.21

Mandible

MML (mm) >67a,f 57.9 46.9b 110.0 50.1g

Depth of horizontal ramus of the mandible (mm) 6.3 4.9 3.6b 5.7 4.9g

Depth of horizontal ramus of the mandible 

versus MML

<0.09 0.08 0.08 0.05 0.1

aEstimate.

bBased on USNM 537785.

cBased on CM 86715.

dBased on Pohl specimen.

eBased on only F:AM 78999. 

fBased on only USNM-P 494439. 

gBased on SMF MEA 263.
n u m b e r 9 8 • 9 1

TABLE A3. Measurements of the thoracic and lumbar vertebrae in Patriomanis americana and related living and extinct (†) pangolins

A dash (—) indicates data are not available.

Measurement

Patriomanis 

americana†, 

USNM-

299960

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53848

Cryptomanis 

gobiensis†, 

AMNH

26140

Eomanis 

waldi†, 

SMF ME

263 (cast)

Thoracic vertebraea ultimate or 

penultimate 

vertebra

T16 (of 16) T14 (of 14) T14 (of 14) posteriormost 

preserved 

vertebra

—

Maximum transverse width of centrum (mm) 17.1 14.1 9.4 28.6 ~22 —

Maximum dorsoventral height of centrum (mm) 9.3 8.2 5.6 17.0 ~11 —

Ratio of height versus width 0.54 0.58 0.60 0.59 0.50 —

Maximum anteroposterior length of centrum 11.6 10.5 9.6 21.6 16.2 —

Width versus length 0.68 0.74 1.02 0.76 0.74

Lumbar vertebraea L7 L3 L3 L3 L6 L4

Neural spine height (mm) ~13 10 4.7 28.3 ~20 10.4

Neural spine anteroposterior length (mm) 8.4 9.8 8.2 17.6 8.2 6.5

Height versus length 1.5 1.02 0.57 1.6 2.4 1.6

 Maximum dorsoventral height of centrum (mm) 11.6 8.4 6.6 19.3 13.2b —

Maximum anteroposterior length of centrum 

(mm)

15 12.3 11 25 18.4b —

Length versus height 1.29 1.46 1.67 1.3 1.39b —

aAbbreviations: T14, T16 = 14th and 16th thoracic vertebrae; L3, L4, etc. = third lumbar vertebra, fourth lumbar vertebra, etc.

bMeasurement taken at L3, not L6.
9 2 • 

TABLE A4. Measurements of the metatarsals (mt) and pedal phalanges in Patriomanis americana and related living and extinct (†

pangolins. A dash (—) indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-

299960

Manis 

pentadactyla, 

USNM 3087

Phataginus 

tricuspis 

USNM 537785

CM 16206a

Smutsia 

gigantea, 

AMNH 53848,

53851b

Cryptomanis 

gobiensis†,

AMNH

26140

Eomanis 

waldi†,

SMF Me 84

(holotype), 

15643c

Greatest length of mt

Digit I (mm) 16.4 12.5 10.9 16.7 30.8 8.1

Digit II (mm) 26.6 14.4 14.3 22.6 36.6 15.2

Digit III (mm) 28.1 16 14.6 24.3 39.5 16.5

Digit III versus greatest length of tibia 0.35 0.24 0.25 0.2 0.43 0.31

Digit IV (mm) 27.4 15.6 14.9 24.8 38.7 13.6

Digit V (mm) 20.6 10 12.2 22.1 — —

Greatest length of proximal phalanges

Digit I (mm) 11.1 6.0 7.1 11.6 19.8 5.8

Digit II (mm) 11.2 4.2 6.3 7.3 16.2 6.3

Digit III (mm) 11.1 3.7 6.4 11.9 17.1 5.5

Digit IV (mm) 10.5 3.3 6.2 8.5 15.4 3.7

Digit V (mm) 9.0 3.2 5.8 6.6 14.1 2.4

Greatest length of intermediate phalanges 

(Ratio of intermediate/proximal 

phalanx length)

Digit II (mm) 10.9 (1.0) 5.7 (1.4) 8.9 (1.2) 9.8 (1.3) 13.8 (0.9) 4.3 (0.7)

Digit III (mm) 11 (1.0) 7.3 (2.0) 8.6 (1.3)

10.5 (1.3)

10.5 (0.9) 12.6 (0.7) 6.3 (1.1)

Digit IV (mm) 9.8 (0.9) 6.9 (2.1) 8.6 (1.4)

10.7 (1.4)

10.6 (1.2) 14.2 (0.9) 4.0 (1.1)

Digit V (mm) 8.2 (0.9) 5.9 (1.8) 8.0 (1.2) 7.6 (1.2) 10.9 (0.8) —

Greatest length of ungual phalanges

Digit I (mm) 10.1 8.5 4.3 14.9 13.0 6.2

Digit II (mm) 11.4 10.8 12.1 20.4 15.0 9

Digit III (mm) 13.1 13.3 14.9 25.0 15.5 11.1

Digit IV (mm) 12.0 11.0 15.0 22.3 16.2 7.4

Digit V (mm) 9.6 8.7 10.8 15.8 — 4.1

a All measurements from USNM 537785 except for intermediate phalanges; first number under intermediate phalanges from USNM 537785, second number from CM 

16206.

bMetatarsals from 53851, phalanges from 53848.

cMetatarsals: digit I from holotype, digits II–IV from 15643; phalanges: digits I, II from holotype, digits III–V from SMF Me 15643.
n u m b e r 9 8 • 9 3

TABLE A5. Measurements of the astragalus in Patriomanis americana and related living and extinct (†) pangolins. A dash (—) indicate

data are not available.

Measurement

Patriomanis 

americana†, 

USNM-P 2999

Patriomanis 

americana†,

USNM-P 4944

Manis 

pentadactyla, 

USNM 3087

Manis 

crassicaudata, 

FMNH 24440

Phataginus 

tricuspis, 

USNM 5377

Maximum width of head (mm) 10.9 12.2 8.7 12.1 6.3

Maximum height of head (mm) 8 8.0 7.3 12.3 4.4

Height versus width of head 1.36 1.53 1.19 0.98 1.43

Minimum neck width (mm) 10.2 9.9 8.2 10.0 6.0

Maximum body width (mm) 17.2 18.0 13.4 19.3 8.5

Neck width versus body width 0.59 0.55 0.61 0.52 0.71

Maximum proximodistal depth of lateral 

trochlea (mm)

9.5 — 8.8 13.8 6.3

Maximum proximodistal depth of medial 

trochlea (mm)

9.4 — 6.6 8.0 4.0

Depth of medial trochlea versus depth of 

lateral trochlea

1.01 — 1.33 1.73 1.58

Maximum proximodistal length (mm) 18.5 21.5 14.5 22.6 10.3

Measurement

Smutsia gigantea, 

AMNH 5385

Cryptomanis gobiensis†, 

AMNH 2614

Necromanis franconica†, 

SMF M3379a+

Euromanis krebsi†, 

SMF 94/1 (cast

Maximum width of head (mm) 18.1 15.1 12.0 6.0

Maximum height of head (mm) 17.3 11.9 9.5 4.3

Height versus width of head 1.05 1.27 1.26 1.40

Minimum neck width (mm) 18.0 12.7 12.5 5.4

Maximum body width (mm) 27.5 25.4 22.5 14.1

Neck width versus body width 0.65 0.50 0.56 0.38

Maximum proximodistal depth of lateral 

trochlea (mm)

18.6 13.2 15.6 10.2

Maximum proximodistal depth of medial 

trochlea (mm)

12.5 11.1 12.9 9.3

Depth of medial trochlea versus depth of 

lateral trochlea

1.49 1.19 1.21 1.10

Maximum proximodistal length (mm) 28.2 25 23.9 15.5
9 4 • 

TABLE A6. Measurements of the calcaneus in Patriomanis americana and related living and extinct (†) pangolin

Measurement

Patriomanis 

americana†, 

USNM-P 2999

Patriomanis 

americana†, 

USNM-P 4944

Manis 

pentadactyla, 

USNM 3087

Manis 

crassicaudata, 

AMNH 24440

Phataginus 

tricuspis, 

USNM 5377

Maximum proximodistal length (mm) 30.0 33.0 22.0 35.0 16.5

Maximum proximodistal length of tuber (mm) 15.0 18.0 11.0 17.0 8.5

Proximodistal length of tuber as a percentage 

of proximodistal length

0.50 0.55 0.50 0.49 0.51

Measurement

Smutsia gigantea, 

AMNH 5385

Cryptomanis gobiensis†, 

AMNH 2614

Necromanis franconica†, 

SMF M3379

Eomanis waldi†, 

SMF Me 15643

Maximum proximodistal length (mm) 45.0 42.4 39.2 18.9

Maximum proximodistal length of tuber (mm) 16.7 22.0 21.0 7.9

Proximodistal length of tuber as a percentage 

of proximodistal length

0.37 0.52 0.54 0.42

TABLE A7. Measurements of the tibia and fibula in Patriomanis americana and related living and extinct (†) pangolins. A dash (—

indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-P 29996

USNM-

494439a

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53858

Cryptomanis 

gobiensis†,

AMNH

26140

Eomanis 

waldi†,

SMF Me

84 (type)

Tibia

Maximum length (mm) 79.6, 85.7 67.3 59.0 133.1 92.5 49.5

Maximum transverse width of distal end (mm) 16.9, 18.9 16.3 14.1 39.6 25.7 —

Maximum anteroposterior depth of distal 

end (mm)

11.7, 13.9 8.7 5.7 21.2 14.0 —

Anteroposterior depth versus transverse 

width of distal end

1.44, 1.36 1.87 2.47 1.87 1.84 —

Fibula

Maximum fibula length (mm) 68.0 58.8 53.8 118.0 80.2 42.6

Fibula length versus tibia length 0.85 0.87 0.91 0.89 0.87 0.86

aFirst measurement for tibia and all fibula measurements from USNM-P 299960; second measurement for tibia from USNMP 494439.
n u m b e r 9 8 • 9 5

TABLE A8. Measurements of the patella and femur in Patriomanis americana and related living and extinct (†) pangolins. A dash (—

indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-

299960

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53858

Cryptomanis 

gobiensis†,

AMNH

26140

Euromanis 

krebsi†, 

SMF Me

94/1

Eomanis 

waldi†, 

Pohl 

specimen

Patella

Maximum patella height (mm) 15.5 18.7 10.4 32.3 21.4 11.8 —

Maximum patella width (mm) 13.7 11.5 7.7 23.5 18.4 9.4 —

Patella width versus patella height 1.13 1.63 1.35 1.37 1.16 1.26 —

Maximum patella depth (mm) 8.4 8.1 4.8 14.8 14.2 5.4 —

Patella depth versus patella height 0.56 0.43 0.46 0.46 0.66 0.46 —

Femur

Maximum femur length (mm) 85.2 72.8 51.4 149.8 103.0 78.0 47.3, 50.3a

Proximodistal length of greater 

trochanter (mm)

10.4 12.3 9.8 28.7 15.2 — —

Length of greater trochanter versus 

femur length

0.12 0.17 0.19 0.19 0.15 — —

Distance from head to third 

trochanter (mm)

49.2 54.8 42.4 114.5 48.0 31.5 18

Distance from head to third trochanter 

versus femur length

0.58 0.75 0.82 0.76 0.47 0.40 0.38

Width at narrowest point on shaft (mm) 13.3 9.7 6.7 25.3 18.8 — —

Depth at narrowest point on shaft (mm) 8.5 6.7 5.2 16.7 10.7 — —

Depth versus width at narrowest point 

on shaft

1.56 1.45 1.29 1.51 1.76 — —

Maximum width of distal end (mm) 23.7 21.6 15.4 48.4 35.3 22.1 16.5

Maximum depth of distal end (mm) 23.4 17.0 11.4 40.4 27.2 — —

Depth versus width at distal end 1.01 1.27 1.35 1.20 1.30 — —

Maximum width of medial condyle (mm) 10.0 9.5 6.4 21.6 15.2 — 7.5

Maximum width of lateral condyle (mm) 8.2 8.7 5.6 14.8 12.0 — 6.5

Width of lateral condyle versus width of 

medial condyle

1.22 1.09 1.14 1.46 1.27 — 1.15

aSecond measurement from SMF Me 1573.

TABLE A9. Measurements of the scapula in Patriomanis americana and related living and extinct (†) pangolins. A dash (—) indicate

data are not available.

Scapula measuremen

Patriomanis 

americana†,

USNM-

299960

Patriomanis 

americana†,

F:AM 

78999

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53858

Manis 

pentadactyla, 

USN

308733

Euromanis 

krebsi†,

SMF Me

94/1

Eomanis 

waldi†,

Pohl 

specimen

Dorsoventral lengtha (mm) — 61.1 36.6 102.4 50.8 — 36.5

Height of scapular spineb (mm) 10.5 — 4.4 14.3 7.2 9 —

Maximum transverse width of glenoid 

fossa (mm)

11 — 6.6 19 9.3 9.20 —

Height of spine versus width of glenoid 0.95 — 0.67 0.75 0.78 0.98 —

aMeasured from glenoid to dorsal border along spine.

bIn distal view.
9 6 • 

TABLE A10. Measurements of the humerus, radius, and ulna in Patriomanis americana and related living and extinct (†) pangolins

A dash (—) indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-

299960

Patriomanis 

americana†,

USNM-

531556

Patriomanis 

americana†,

USNM-

494439

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53858

Necromanis 

franconica†,

SMF Me

94/1

Eomanis 

waldi†,

Pohl 

specimen

Humerus

Length of deltopectoral crest (mm) 61.6 63.4 — 41.0 26.5 86.4 66.9 17.0

Maximum humerus length (mm) 76.9 81.0 — 62.3 48.4 121.2 84.2 47.3

Length of deltopectoral crest 

versus humerus lengt

0.79 0.78 — 0.66 0.55 0.71 0.79 0.47

Epicondyle width (mm) 28.8 29.4 — 28.8 22.0 59.4 44.0 18.0

Epicondyle width versus 

humerus length

0.37 0.36 — 0.46 0.45 0.49 0.52 0.38

Width of entepicondylar 

process (mm)

8.8 8.9 — 10.9 9.2 18.7 13.6 6.8

Width of entepicondylar 

process versus epicondyle 

width

0.31 0.30 — 0.38 0.42 0.31 0.3 0.4

Radius

Maximum radius length (mm) 48.9 52.9 53.5 42.1 39.7 81.8 60.2 33.5

Radius length versus humerus 

length

0.64 0.65 — 0.68 0.82 0.67 0.90 0.71

Maximum depth of head (mm) 7.1 6.9 — 7.7 5.2 14.8 — —

Maximum width of head (mm) 12.0 12.5 12.0 1.4 8.1 20.9 — —

Width versus depth of head 0.59 0.55 — 0.68 0.64 0.71 — —

Maximum depth of distal shaft 

(mm)

10.60 9.30 10.70 10.40 6.30 19.60 13.50 8.80

Depth of distal shaft versus 

radius length

0.22 0.18 0.20 0.25 0.16 0.24 0.22 0.26

Ulna

Maximum ulna length (mm) 67.8 — — 60.5 51.2 115.1 — 50.7

Ulna length versus humerus 

length

0.88 — — 0.97 1.06 0.95 — 1.14

Maximum length of olecranon 

(mm)

18.1 — — 20.7 9.3 23.4 — 13.8

Length of olecranon versus ulna 

length

0.27 — — 0.34 0.18 0.20 — 0.27

Maximum depth of distal end 

of shaft (mm)

8.5 — — 7.7 4.5 12.7 — 7.4

Depth of distal end of shaft 

versus ulna length

0.13 — — 0.13 0.09 0.11 — 0.15

Length of styloid process (mm) 2.8 — — 3.0 3.5 — — —

Length of styloid process versus 

ulna length

0.04 — — 0.05 0.07 — — —
n u m b e r 9 8 • 9 7

TABLE A11. Measurements of the scapholunar in Patriomanis americana and related living pangolin

Scapholunar measuremen

Patriomanis 

americana†, 

USNM-P 2999

Manis 

pentadactyla, 

USNM 3087

Phataginus tricuspis, 

USNM 5377

Smutsia gigantea, 

AMNH 5385

Maximum transverse width (mm) 14.5 15.3 13.1 28.5

Maximum dorsoventral depth (mm) 9.9 11.8 6.5 19.5

Depth versus width 0.68 0.77 0.5 0.68

†Extinct.

TABLE A12. Measurements of the metacarpals in Patriomanis americana and related living and extinct (†) pangolins. A dash (—

indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-

299960

Patriomanis 

americana†,

USNM-

494439

Cryptomanis 

gobiensis, 

AMNH

26140

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea,

AMNH

53858

Eomanis 

waldi†, 

Pohl 

specimen

Greatest length of metacarpals

Metacarpal I (mm) 13.3 14.8 25.5 9.6 6.5 11.5 —

Metacarpal II (mm) 19.2 — ~26 13.6 11.1 19.1 —

Metacarpal III (mm) 21.9 — 28.3 14.6 1.8 20.9 13.7

Length of metacarpal III versus maximum 

length of humerus

0.28 — 0.30 0.23 0.24 0.19 0.28

Metacarpal IV (mm) 19.3 21.4 28.9 12.4 11.0 17.8 10.2

Metacarpal V (mm) 12.6 — — 8.6 10.2 17.2 —

Midshaft width of metacarpals

Metacarpal I (mm) 2.6 2.7 3.6 1.6 4.1 2.3 —

Metacarpal II (mm) 3.8 — 6.0 2.2 1.8 4.6 —

Metacarpal III (mm) 4.8 — 10.0 5.8 3.8 9.3 —

Metacarpal IV (mm) 4.3 4.7 8.1 3.8 1.7 5.8 —

Metacarpal V (mm) 3.6 — 5.3 2.3 1.7 3.2 —

Midshaft depth of metacarpals

Metacarpal I (mm) 2.6 2.7 4.0 2.3 1.6 4.1 —

Metacarpal II (mm) 3.7 — 4.5 3.9 2.1 4.9 —

Metacarpal III (mm) 4.7 — 5.5 6.4 3.7 10.1 6.0

Metacarpal IV (mm) 4.0 4.0 5.5 4.1 2.6 7.4 5.0

Metacarpal V (mm) 2.6 — 3.2 3.5 1.8 4.5 —

Ratio of metacarpal width to length

Metacarpal I 0.20 0.18 0.14 0.14 0.18 0.2 —

Metacarpal II 0.20 — 0.23 0.16 0.16 0.24 —

Metacarpal III 0.22 — 0.35 0.4 0.32 0.44 —

Metacarpal IV 0.22 0.22 0.28 0.31 0.15 0.32 —

Metacarpal V 0.29 — — 0.27 0.17 0.19 —
9 8 • 

TABLE A13. Measurements of the manual phalanges in Patriomanis americana and related living and extinct (†) pangolins. A dash (—

indicates data are not available.

Measurement

Patriomanis 

americana†,

USNM-

299960

Patriomanis 

americana†, 

USNM-

494439

Patriomanis 

americana†,

F:AM 78999 

(type)

Manis 

pentadactyla, 

USN

308733

Phataginus 

tricuspis, 

USN

537785

Smutsia 

gigantea, 

AMNH

53858

Cryptomanis 

gobiensis†,

AMNH

26140

Eomanis 

waldi†, 

Pohl 

specimen

Greatest length of proximal phalanges

Digit I (mm) 11.1 12.8 — 6.2 5.8 11.6 >13.5 —

Digit II (mm) 11.1 13.3 12.7 7.4 5.9 11.6 9.2 —

Digit III (mm) 9.6 9.7 10.0 6.4 8.7 14.3 7.2 5.5

Digit IV (mm) 8.7 12.0 10.2 5.2 6.1 11.8 9.8 —

Digit V (mm) 7.6 9.0 — 3.3 5.6 6.7 11.1 —

Maximum proximal width of proximal phalanges (Ratio of width/length)

Digit I (mm) 4.7 (0.4) 4.5 (0.35) — 2.5 (0.4) 1.4 (0.2) 4.3 (0.4) 6.3 (<0.5) —

Digit II (mm) 6.4 (0.6) 8.3 (0.6) 7.3 (0.6) 3.7 (0.5) 2.9 (0.5) 8.4 (0.7) 9.4 (1.0) —

Digit III (mm) 7.9 (0.8) 8.1 (0.8) 8.2 (0.8) 6.9 (1.1) 5.2 (0.6) 12.2 (0.85) 10.2 (1.4) —

Digit IV (mm) 6.6 (0.8) 6.2 (0.5) 6.9 (0.7) 5.4 (1.0) 3.0 (0.5) 10.5 (0.9) 9.8 (1.0) —

Digit V (mm) 5.6 (0.7) 5.6 (0.7) — 3.5 (1.1) 2.7 (0.5) 5.6 (0.8) 7.9 (0.7) —

Greatest length of intermediate phalanges (Ratio of intermediate/proximal phalanx length)

Digit II (mm) 11.2 (1.0) 12.6 (1.0) — 8.8 (1.2) 7.1 (1.2) 16.9 (1.5) — —

Digit III (mm) 10.6 (1.1) 13.0 (1.0) — 12.5 (2.0) 11.9 (1.4) 18.3 (1.3) 13.1 (1.8) 4.8 (0.9)

Digit IV (mm) 10.1 (1.2) 12.6 (1.1) — 9.2 (1.8) 8.7 (1.4) 17.2 (1.5) — —

Digit V (mm) 7.6 (1.0) 9.4 (1.0) — 5.6 (1.7) 7.7 (1.4) 9.9 (1.5) 10.6 (1.0) —

Greatest length of ungual phalanges

Digit I (mm) 9.1 10.3 10.1 8.0 4.9 13.7 — —

Digit II (mm) 12.2 15.8 — 16.5 11.6 32.3 — 15.8

Digit III (mm) 13.4 — — 33.3 19.6 52.5 — 20.7

Digit IV (mm) 11.0 13.6 — 23.4 13.7 43.1 — 13.5

Digit V (mm) 9.2 — — 11.6 10.8 19.0 — —
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