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Systematics and Phylogeny of Paleocene-Eocene Nyctitheriidae (Mammalia, Eulipotyphla?) with Description of a new Species from the Late Paleocene of the Clarks Fork Basin, Wyoming, USA

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Abstract

Nyctitheriidae is a diverse group of small, insectivorous mammals from the Paleogene of Asia, North America, and Europe that have alternately been linked to Eulipotyphla (shrews, moles, hedgehogs, solenodons), Euarchonta (primates, tree shrews, dermopterans), or Chiroptera (bats). Even intrafamilial relationships are poorly understood, resulting in ambiguity regarding morphological character polarity critical for evaluating supraordinal relationships and paleobiogeographic patterns. To help address this issue, we performed a cladistic analysis of 51 North American, European, and Asian nyctitheriid species, including a new nyctitheriid, Plagioctenodon thewisseni sp. nov. from the late Paleocene of Wyoming, using 66 characters derived from dental morphology. Although the oldest nyctitheriids are found in North America, the resulting most-parsimonious cladograms support an Asian origin of the family with dispersal into North America by the early Paleocene. Among North American and European groups, the subfamilies Nyctitheriinae and Amphidozotheriinae, and the genera Leptacodon and Saturninia are not monophyletic and require future study and revision. The multi-species genera Nyctitherium, Plagioctenodon (including P. thewisseni), Plagioctenoides, Cryptotopos, and Euronyctia are found to be monophyletic, whereas Wyonycteris is paraphyletic, having Pontifactor bestiola nested within it. The earliest known European nyctitheriids (Leptacodon nascimentoi, Placentidens lotus, Plagioctenodon dormaalensis, Wyonycteris richardi) appear in the early Eocene and are each found in an otherwise strictly North American clade consisting of either solely Paleocene or a combination of Paleocene and Eocene taxa, suggesting at least four earliest Eocene dispersals between North America and Europe.

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Notes

  1. The codings for the M1 and M2 of Leptacodon nascimentoi were based on Fig. 10 (SV3-300) and Fig. 9 (SV2-12), respectively, which is opposite to how they are identified in Estravis (1996). See the reasoning in the Body Mass Calculations subsection of the Materials and Methods section.

  2. Leptacodon proserpinae was taken out of the final analysis. It is only mentioned in one publication with a single tooth, the P4, illustrated. Only a small fraction of the characters in this analysis could be coded (8 of 66), which caused a great deal of instability in the placement of the taxon and resulted in a large polytomy in the strict consensus tree.

  3. The holotype specimen number in Setoguchi (1973) is given as TTM 1219 but based on the figures, the specimen (broken P4, M1-M3) is the same as the holotype for Nyctitherium christopheri in Krishtalka and Setoguchi (1977), although that specimen number is reported as CM 16996.

  4. Posterior premolars are referred to as penultimate and ultimate to avoid confusion because Maelestes gobiensis has five premolars and nyctitheriids have four. It is understood that the third premolar is eventually lost in eutherians when the count goes down to four, so that the P4 and P5 of Maelestes are homologous to the P3 and P4 of nyctitheriids.

  5. For characters that just specify “lower molar” or “upper molar,” all taxa were coded using the second molar, if possible. If that position was unknown, the first molar was used.

  6. sensu Menu and Sigé 1971: The entoconid and hypoconulid are closely appressed but separated by a fissure and the hypoconid and hypoconulid are connected by a crest.

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Acknowledgments

We thank Richard Hulbert, Kristen MacKenzie, and Jason Bourque (UF) for help in cataloguing and preparation of the UF specimens. We thank Philip Gingerich (UM) and Gregg Gunnell (now of Duke Lemur Center) for access to specimens and comparative casts and for help with loans at UM, as well as William Sanders (UM) for help with preparation of specimens. We thank Peter Houde (New Mexico State University) for access to limestones containing invaluable fossils. We thank Stephen Chester (now of Brooklyn College) and Josh Van Houten (Department of Internal Medicine, Yale University) for micro-CT imaging done at the Yale University Core Center for Musculoskeletal Disorders microCT facility; and Doug Boyer, Jimmy Thostenson, and Judit Marigo (Duke University) for access to and assistance with the microCT facilities at the Shared Materials Instrumentation Facility at Duke University. We thank Jason Bourque, Stephen Chester, Jerry Hooker, Paul Morse, Ken Rose, and Aaron Wood for helpful discussions. We thank John Wible and an anonymous reviewer for providing helpful comments that improved the manuscript. Research was in part funded by the Miss Lucy Dickinson Fellowship, the Geological Sciences Graduate Award, and a Geological Society of America Student Research Grant to C.L.M. The version of TNT used in the cladistic analysis was produced with support by the Willi Hennig Society. This is University of Florida Contribution to Paleobiology 679.

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Authors and Affiliations

  1. Department of Geological Sciences, University of Florida, Gainesville, FL, 32611, USA

    Carly L. Manz

  2. Florida Museum of Natural History, University of Florida, Gainesville, FL, 32611, USA

    Carly L. Manz & Jonathan I. Bloch

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  1. Carly L. Manz
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Correspondence to Carly L. Manz.

Appendices

Appendix 1. Taxa Selected for Analysis and Sources of Data

Outgroup

Maelestes gobiensis Wible et al., 2007—Wible et al. (2007, 2009)

Erinaceomorpha

Adunator minutus (Jepsen, 1930)—Jepsen (1930); Gingerich (1983); Secord (2008); YPM-PU 19463 (cast)

Macrocranion junnei Smith et al., 2002—Smith et al. (2002); Rose et al. (2012); UM 93378 (cast)

Nyctitheriidae

Plagioctenodon krausae Bown, 1979—Bown (1979); Bown and Schankler (1982)

Plagioctenodon savagei Bown and Schankler, 1982—Bown and Schankler (1982)

Plagioctenodon dormaalensis (Quinet, 1964)—Smith (1996) as Leptacodon dormaalensis

Plagioctenodon rosei (Gingerich, 1987)—Rose (1982) as cf. Leptacodon packi; Rose and Gingerich (1987) as cf. Leptacodon packi; Gingerich (1987) as Leptacodon rosei; UF 303728; UF 303729; UF 303730; UM 71650 (cast); UM 76895 (cast); UM 77032 (cast)

Plagioctenodon thewisseni this publication— UM 39873; UM 76906; UM 76920; UM 82576; UM 83931; UM 86725; UF 289746; UF 289747; UF 294696

Leptacodon tener Matthew and Granger, 1921—Simpson (1935); McKenna (1968); Krishtalka (1976); Scott (2003)

Leptacodon munusculum Simpson, 1935—Krishtalka (1976); Gingerich et al. (1983); Krause and Gingerich (1983); Scott (2003)

Leptacodon catulus Krishtalka, 1976—Krishtalka (1976); Gheerbrant and Hartenberger (1999)

Leptacodon packi Jepsen, 1930—Jepsen (1930); Krishtalka (1976); Secord (2008)

Leptacodon nascimentoi Estravis, 1996—Estravis(1996)Footnote 1

Leptacodon acherontus Secord, 2008—Secord (2008); YPM-PU 19957 (cast)

Leptacodon donkroni Rose et al., 2012—Rose et al. (2012)

Leptacodon proserpinae Footnote 2 Van Valen, 1978—Van Valen (1978)

Leptacodon choristus Secord, 2008—Secord (2008)

Nyctitherium velox Marsh, 1872—Robinson (1968); YPM 13510 (cast)

Nyctitherium serotinum (Marsh, 1872)—Robinson (1968)

Nyctitherium krishtalkai Christiansen and Stucky, 2013—Christiansen and Stucky (2013)

Nyctitherium christopheri Krishtalka and Setoguchi, 1977—Setoguchi (1973)as Nyctitherium robinsoni Footnote 3; Krishtalka and Setoguchi (1977)

Acrodentis rosenorum Christiansen and Stucky, 2013—Christiansen and Stucky (2013)

Ceutholestes dolosus Rose and Gingerich, 1987—Rose and Gingerich (1987); UM 82503 (cast)

Limaconyssus habrus Gingerich, 1987—Gingerich (1987); UM 86724 (cast)

Pontifactor bestiola West, 1974—West (1974); Bown (1979)

Wyonycteris chalix Gingerich, 1987—Gingerich (1987)

Wyonycteris richardi Smith, 1995—Smith (1995)

Wyonycteris primitivus Beard and Dawson, 2009—Beard and Dawson (2009)

Wyonycteris galensis Secord, 2008—Secord (2008); YPM-PU 14138 (cast)

Wyonycteris microtis Secord, 2008—Secord (2008); YPM-PU 19479 (cast)

Plagioctenoides microlestes Bown, 1979—Bown (1979); Rose et al. (2012)

Plagioctenoides tombowni Rose et al., 2012—Rose et al. (2012)

Amphidozotherium cayluxi Filhol, 1877—Sigé (1976)

Saturninia gracilis Stehlin, 1940—Crochet (1974); Sigé (1976)

Saturninia mamertensis Sigé, 1976— Sigé (1976)

Saturninia grandis Sigé, 1976— Sigé (1976)

Saturninia rigasii Hooker and Weidman, 2000—Hooker and Weidmann (2000)

Saturninia ceciliensis Storch and Haubold, 1989—Storch and Haubold (1989)

Saturninia intermedia Sigé, 1976— Sigé (1976)

Saturninia pirenaica Gibert Clols and Agustí Ballester, 1979—Gibert Clols and Agustí Ballester (1979); Sigé (1997); Ziegler (2007)

Saturninia pelissiei Sigé, 1997—Sigé (1997); Ziegler (2007)

Saturninia carbonum Sigé and Storch, 2001—Sigé (2001)

Cryptotopos beatus Crochet, 1974—Crochet (1974); Sigé (1976) as Saturninia beata; Hooker and Weidmann (2000); Ziegler (2007)

Cryptotopos woodi (Cray, 1973)—Cray (1973); Ziegler (2007)

Cryptotopos hartenbergi (Sigé, 1976)— Sigé (1976); Ziegler (2007)

Scraeva hatherwoodensis Cray, 1973—Cray 1973

Placentidens lotus Russell et al.1973—Russell et al. (1973)

Euronyctia montana Sigé, 1997— Sigé (1997)

Euronyctia grisollensis (Sigé, 1976)—Sigé (1976); Sigé (1997)

Paradoxonycteris soricodon Revilliod, 1922—Hooker and Weidman (2000)

Asionyctia guoi Missiaen and Smith, 2005—Missiaen and Smith (2005)

Bumbanius rarus Russell and Dashzeveg, 1986—Russell and Dashzeveg (1986); Lopatin (2006)

Eosoricodon terrigena Lopatin, 2005—Lopatin (2005)

Oedolius perexiguus Russell and Dashzeveg, 1986—Russell and Dashzeveg (1986); Lopatin (2006)

Edzenius lus Lopatin, 2006—Lopatin (2006)

Appendix 2. Unambiguous Synapomorphies for Key Nodes

Node A

52 (1) upper molar stylocone absent

61 (1) upper molar hypocone present

Node B: Adunator minutus + Macrocranion junnei

22 (2) upper ultimate premolar postcingulum absent

25 (2) M1 trigonid less wide than talonid

26 (2) M2 trigonid less wide than talonid

50 (1) M1 parastylar lobe anterolabially projecting

63 (1) M3 parastylar lobe small

Node C: Nyctitheriidae

14 (1) lower ultimate premolar paraconid lingual to midline

34 (1) lower molar paraconid anteriorly projecting

37 (2) lower molar protoconid larger than metaconid

46 (2) lower molar entoconid smaller than hypoconid

66 (0) posteriormost mental foramen below penultimate premolar or more anterior

Node D

15 (2) lower ultimate premolar metaconid present

48 (1) upper molar length between 0.7-0.95 of the width

Node E: North American and European nyctitheriids

17 (1) lower ultimate premolar talonid basin present

18 (1 or 2) lower ultimate premolar talonid has two or three cusps

26 (1) M2 trigonid width subequal to talonid

44 (0) lower molar hypoconulid positioned on midline

46 (0) lower molar entoconid subequal in height to hypoconid

58 (2) upper molar paraconule lingual to metaconule and closer to protocone

Node F

14 (0) lower ultimate paraconid on midline

21 (0) upper ultimate premolar precingulum present

33 (0) lower molar paraconid cuspate in shape

37 (1) lower molar protoconid and metaconid subequal in size

Node G

24 (1) upper ultimate premolar metacone present

49 (0) M1 ectoflexus present

54 (1) upper molar centrocrista strong and rectilinear

55 (1) upper molar precingulum strong, reaching past paraconule

Node H

11 (3) lower ultimate premolar width greater than 0.8 of the length

14 (1) lower ultimate premolar paraconid lingual to midline

29 (1) M1 length greater than that of M2

40 (0) lower molar cristid obliqua meets the protocristid lingual to notch

47 (1) lower molar postcingulid present

Node I: Placentidens lotus + Ceutholestes dolosus

6 (1) lower penultimate premolar metaconid present

37 (0) lower molar metaconid larger than protoconid

46 (1) lower molar entoconid taller than hypoconid

Node J: Nyctitherium

20 (1) anterodorsal shift in upper ultimate premolar visible from labial view

23 (1) upper ultimate premolar hypocone present

56 (0) upper molar paraconule small, not strongly winged

60 (1) upper molar postcingulum greatly expanded into shelf

Node K

7 (1) lower penultimate premolar anteriorly canted

Node L

8 (1) lower penultimate premolar smaller than surrounding premolars

Node M: Plagioctenodon

12 (2) lower ultimate premolar paraconid present and high on anterior of trigonid

59 (1) upper molar paracone and metacone height subequal

Node N

33 (1) lower molar paraconid crestiform in shape

40 (2) lower molar cristid obliqua ascends the protocristid to the tip of the metaconid

55 (2) upper molar precingulum strong with pericone

Node O

11 (1) lower ultimate premolar width between 0.5-0.65 of the length

43 (1) cristid obliqua is deeply concave occlusally

44 (2) lower molar hypoconulid positioned lingually in nyctalodont condition

46 (1) lower molar entoconid taller than hypoconid

66 (1) posteriormost mental foramen below ultimate premolar or more posterior

Node P: Plagioctenoides

18 (0) lower ultimate premolar has one talonid cusp

Node Q: Wyonycteris + Pontifactor

27 (0) lower molar trigonid anteroposteriorly compressed

Node R: W. galensis + W. chalix + W. richardi + P. bestiola

16 (0) lower ultimate premolar talonid narrower than trigonid or absent

31 (1) lower molar metaconid twinned

33 (0) lower molar paraconid cuspate in shape

53 (1) upper molar mesostyle present

54 (2) upper molar centrocrista dilambdodont

Pontifactor bestiola autapomorphies

55 (1) upper molar precingulum strong, reaching past paraconule

58 (1) upper molar paraconule lingual to metaconule and halfway to protocone

59 (1) upper molar paracone and metacone subequal

60 (1) upper molar postcingulum greatly expanded into shelf

Node S

22 (1) upper ultimate premolar postcingulum greatly expanded into shelf

23 (1) upper ultimate premolar hypocone present

60 (1) upper molar postcingulum greatly expanded into shelf

64 (1) dentary has one mental foramen

Node T: Cryptotopos?

11 (3) lower ultimate premolar width greater than 0.8 of the length

33 (0) lower molar paraconid cuspate in shape

42 (1) lower molar cristid obliqua has notch halfway along the length

Node U: Cryptotopos?

27 (0) lower molar trigonid anteroposteriorly compressed

43 (2) cristid obliqua is convex occlusally

54 (0) upper molar centrocrista absent or weak

Node V

55 (0) upper molar precingulum absent or weak

56 (0) upper molar paraconule small, not strongly winged

57 (0) upper molar metaconule small, not strongly winged

Node W

47 (1) lower molar postcingulid present

Node X: Paradoxonycteris + Euronyctia

54 (2) upper molar centrocrista dilambdodont

56 (2) upper molar paraconule large, not strongly winged

57 (1) upper molar metaconule small, strongly winged

Node Y

25 (0) M1 trigonid wider than talonid

26 (0) M2 trigonid wider than talonid

33 (0) lower molar paraconid cuspate in shape

37 (2) lower molar protoconid larger than metaconid

Node Z: Scraeva + Limaconyssus

13 (1) lower ultimate premolar paraconid is large cusp

28 (0) lower molar trigonid height twice that of the talonid

Appendix 3. Characters Used in Cladistic Analysis

1. Lower incisor shape: conical (0), spatulate (1), or pectinate (2)

2. Second lower incisor number of cusps: one (0), two (1), three (2), four (3), or five (4)

3. Lower canine posterior shelf or basal cusp: absent (0) or present (1)

4. Second lower premolar number of roots: two (0) or one (1)

5. Lower penultimate premolarFootnote 4 number of roots: two (0) or one (1)

6. Lower penultimate premolar metaconid: absent (0) or present (1)

7. Lower penultimate premolar protoconid orientation: dorsoventral (0) or anteriorly canted (1)

8. Lower penultimate premolar size relative to other premolars: similarly sized to large (0) or small (1)

9. Upper penultimate premolar protocone: present (0) or absent (1)

10. Upper penultimate premolar metacone: absent (0) or present (1)

11. Lower ultimate premolar shape (width/length): less than 0.5 (0), 0.5-0.65 (1), 0.65-0.8 (2), or more than 0.8 (3)

12. Lower ultimate premolar paraconid position: absent (0), present and low on trigonid (1), or present and high on trigonid (2)

13. Lower ultimate premolar paraconid size: absent or very small cuspule on low anterior cingulid (0) or large cusp (1)

14. Lower ultimate premolar paraconid or anterior-most projection of tooth position: on midline or buccal to midline (0) or lingual to midline (1)

15. Lower ultimate premolar metaconid: absent (0), swelling (1), or present (2)

16. Lower ultimate premolar width of talonid: narrower than trigonid (0) or as wide as or wider than trigonid (1)

17. Lower ultimate premolar talonid basin: absent (0) or present (1)

18. Lower ultimate premolar talonid number of cusps: one (0), two (1), or three (2)

19. Upper ultimate premolar highly waisted: present (0) or absent (1)

20. Upper ultimate premolar anterodorsal shift (from labial view): absent (0) or present (1)

21. Upper ultimate premolar precingulum: present (0) or absent (1)

22. Upper ultimate premolar postcingulum size: small (0), greatly expanded (1), or absent (2)

23. Upper ultimate hypocone: absent (0) or present (1)

24. Upper ultimate premolar metacone: absent (0) or present (1)

25. M1 trigonid width: wider than talonid (0), equal to talonid (1), or narrower than talonid (2)

26. M2 trigonid width: wider than talonid (0), equal to talonid (1), or narrower than talonid (2)

27. Lower molarFootnote 5 trigonid length: anteroposteriorly compressed (0) or about equal to talonid or longer (1)

28. Lower molar trigonid height: nearly twice the height of the talonid (0), less than twice the height of the talonid (1), or nearly the same height as the talonid (2)

29. M1 and M2 length: subequal, within 5% (0), M1 longer than M2 (1), or M2 longer than M1 (2)

30. M2 and M3 length: M3 longer than M2 (0), subequal, within 5% (1), or M2 longer than M3 (2)

31. Lower molar (more visible on the M1) metaconid “twinned,” or a groove runs alongside the cristid obliqua up the posterior surface of the metaconid: absent (0) or present (1)

32. Lower molar precingulid: strong, anteriorly projecting (0) or reduced, not projecting (1)

33. Lower molar paraconid shape: cuspate (0) or crestiform (1)

34. Lower molar paraconid anteriorly projecting: absent (0) or present (1)

35. Lower molar paraconid position: on midline (0) or lingually positioned (1)

36. Lower molar protoconid position relative to metaconid: protoconid anterior to metaconid (0) or the cusps are transverse (1)

37. Lower molar protoconid and metaconid size: metaconid is larger (0), subequal (1), or protoconid is larger (2)

38. Lower molar protoconid and metaconid height: subequal (0), protoconid is taller (1), or metaconid is taller (2)

39. Lower molar labial cingulid: absent (0) or present along length of tooth (1)

40. Lower molar cristid obliqua position: ends lingual to notch in protocristid (0), ends labial to notch in protocristid (1), or ascends to apex of metaconid (2)

41. Lower molar mesoconid on cristid obliqua: absent (0) or present (1)

42. Lower molar notch in cristid obliqua: absent (0), present, located halfway along cristid obliqua (1), or present, located on distal third of cristid obliqua (2)

43. Lower molar cristid obliqua shape in labial view: straight or slightly concave occlusally (0), deeply concave occlusally (1), or convex occlusally (2)

44. Lower molar hypoconulid position: on midline (0), lingual to midline (1), close to entoconid in nyctalodont conditionFootnote 6 (2), or incipient on postcingulid to absent (3)

45. Lower molar entoconid position: anterolingual to hypoconulid (0) or on posterolingual margin of tooth and even posteriorly with hypoconulid (1)

46. Lower molar entoconid height: subequal to hypoconid (0) or taller than hypoconid (1)

47. Lower molar postcingulid: absent (0) or present (1)

48. Upper molar shape (length/width): less than 0.7 (0), 0.7-0.95 (1), or greater than 0.95 (2)

49. M1 ectoflexus: present (0) or absent (1)

50. M1 parastylar lobe orientation: anteriorly projecting (0) or anterolabially projecting (1)

51. M2 ectoflexus: present, relatively deep (0) or absent to very shallow (1)

52. Upper molar stylocone (stylar cusp B): present (0) or absent (1)

53. Upper molar mesostyle: absent (0) or present (1)

54. Upper molar centrocrista (between protocone and metacone): absent or weak (0), strong, rectilinear (1), or strong, dilambdodont (2)

55. Upper molar precingulum: unlobed and small (0), unlobed and large, reaching labially past paraconule (1), or with pericone (2)

56. Upper molar paraconule: small and not strongly winged (0), large and strongly winged (1), or large and not strongly winged (2)

57. Upper molar metaconule: small and not strongly winged (0), large and strongly winged (1), or large and not strongly winged (2)

58. Upper molar paraconule and metaconule positions: level with each other transversely, closer to paracone and metacone than protocone (0), paraconule more lingual than metaconule and located halfway between paracone and protocone, metaconule closer to metacone than protocone (1), or paraconule more lingual than metaconule and located closer to protocone than paracone, metaconule halfway between metacone and protocone (2)

59. Upper molar paracone and metacone height: paracone taller than metacone (0) or subequal (1)

60. Upper molar postcingulum size: absent or small (0) or expanded into a large shelf (1)

61. Upper molar hypocone: absent (0) or present (1)

62. Upper molar postcingulum shape: flat posteriorly (0) or rounded posteriorly (1)

63. M3 parastylar lobe size: large, expanded anterolabially (0) or small (1)

64. Number of mental foramina: two (0) or one (1)

65. Location of anteriormost mental foramen: below P1 or anterior (0), below P2 (1), or below penultimate premolar or posterior

66. Location of posteriormost mental foramen (this character is coded if only one mental foramen): below penultimate premolar or more anterior (0), below ultimate premolar or more posterior (1)

Appendix 4. Data Matrix Used in Cladistic Analysis

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Manz, C.L., Bloch, J.I. Systematics and Phylogeny of Paleocene-Eocene Nyctitheriidae (Mammalia, Eulipotyphla?) with Description of a new Species from the Late Paleocene of the Clarks Fork Basin, Wyoming, USA. J Mammal Evol 22, 307–342 (2015). https://doi.org/10.1007/s10914-014-9284-3

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