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Numbers of Vertebrae in Hominoid Evolution

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Spinal Evolution

Abstract

Vertebral formulae, the combination of regional numbers of vertebrae making up the bony spine, vary across vertebrates and within hominoid primates. Reconstructing the ancestral vertebral formulae throughout hominoid evolution has proved a challenge due to limited fossil evidence and disagreement among researchers. Proposed “long-backed” and “short-backed” ancestors have implications for the evolution of bipedalism and human evolutionary history generally. Here, we analyze a large dataset of hominoid vertebral formulae, including previously unstudied species and subspecies. We find more variation within and between species than expected, particularly in hylobatids (gibbons or lesser apes) and in gorilla and chimpanzee subspecies. Our results suggest that combined thoracic and lumbar numbers of vertebrae are somewhat phylogenetically structured, with outgroup taxa (two species of Old World monkeys, or cercopithecoids) retaining the primitive number of 19 thoracolumbar vertebrae, hylobatids generally possessing 18 thoracolumbar vertebrae, and hominids (great apes and humans) having 17 or 16 thoracolumbar vertebrae. When compared to cercopithecoids, and to putative stem hominoids, extant hominoids show evidence for homeotic change at both the lumbosacral (e.g., decrease in lumbar vertebrae; increase in sacral segments) and in the position of the transitional vertebrae. Homeotic changes are probably also responsible for the differences between African apes and modern humans, with differences in the number of thoracic and lumbar within a 17-segment thoracolumbar framework.

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Acknowledgments

We thank Ella Been for her leadership in organizing this volume and for inviting us to contribute to it. Peer review improved the manuscript. We thank N. Duncan, G. Garcia, E. Hoeger, S. Ketelsen, A. Marcato, B. O’Toole, M. Surovy, and E. Westwig (American Museum of Natural History); M. Milella, M. Ponce de León, and C. Zollikofer (Anthropological Institute and Museum, University of Zurich); H. Taboada (Center for the Study of Human Origins, New York University); Y. Haile-Selassie and L. Jellema (Cleveland Museum of Natural History); D. Katz and T. Weaver (Department of Anthropology, U.C. Davis); B. Patterson, A. Goldman, M. Schulenberg, L. Smith, and W. Stanley (Field Museum of Natural History); C. McCaffery and D. Reed (Florida Museum of Natural History, University of Florida); S. McFarlin (the George Washington University); J. Ashby (Grant Museum of Zoology; University College London); J. Chupasko, J. Harrison, and M. Omura (Harvard Museum of Comparative Zoology); E. Gilissen and W. Wendelen (Musée Royal de l’Afrique Centrale); S. Jancke, N. Lange, and F. Mayer, (Musée für Naturkunde, Berlin); C. Lefèvre (Muséum national d’Histoire naturelle); C. Conroy (Museum of Vertebrate Zoology, U.C. Berkeley); N. Edmison, L. Gordon, K. Helgen, E. Langan, D. Lunde, J. Ososky, and R. Thorington (National Museum of Natural History, Smithsonian Institution); P. Jenkins, L. Tomsett, and R. Portela (Natural History Museum, London); J. Soderberg and M. Tappen (Neil C. Tappen Collection, University of Minnesota); M. Nowak-Kemp (Oxford University Museum of Natural History); S. Bruaux and G. Lenglet (Royal Belgian Institute of Natural Sciences); A. Zihlman, C. Underwood, and J. Hudson (University of California, Santa Cruz); R. Asher (University Museum of Zoology, University of Cambridge); B. Zipfel, S. Jirah, L. Berger, B. Billings, and J. Hemmingway (University of the Witwatersrand); and M. Hiermeier (Zoologische Staatssammlung München) for facilitating access to specimens in their care. SAW has been funded by the National Science Foundation (BCS-0925734), the Leakey Foundation, and the New York University Research Challenge Fund. Part of the data collection done by AGO was possible thanks to the support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by the European Community Research Infrastructure Action under the FP7 “Capacities” Program (projects BE-TAF-4132 and GB-TAF-3674). AGO has received support from the Spanish Ministerio de Ciencia y Tecnología (project: CGL-2015-65387-C3-2-P, MINECO/FEDER), the Spanish Ministerio de Ciencia, Innovación y Universidades (project PGC2018-093925-B-C33), Research Group IT1418-19 from the Eusko Jaurlaritza-Gobierno Vasco. We gratefully acknowledge the Rwandan government for permission to study skeletal remains curated by the Mountain Gorilla Skeletal Project (MGSP). The authors also thank the continuous efforts of researchers, staff, and students from the Rwanda Development Board’s Department of Tourism and Conservation, Gorilla Doctors, DFGFI, the George Washington University, New York University College of Dentistry, Institute of National Museums of Rwanda, and other universities in Rwanda and the USA.

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Authors and Affiliations

  1. Center for the Study of Human Origins, Department of Anthropology, New York University, New York, NY, USA

    Scott A. Williams

  2. New York Consortium in Evolutionary Primatology, New York, NY, USA

    Scott A. Williams

  3. Departamento de Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco/Euskal Herriko Unibertsitatea (UPV/EHU), Leioa, Spain

    Asier Gómez-Olivencia

  4. IKERBASQUE. Basque Foundation for Science, Bilbao, Spain

    Asier Gómez-Olivencia

  5. Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain

    Asier Gómez-Olivencia

  6. Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA

    David R. Pilbeam

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Correspondence to Scott A. Williams .

Editor information

Editors and Affiliations

  1. Department of Sports Therapy, Department of Anatomy and Anthropology, Faculty of Health Professions, Ono Academic College, Sackler Faculty of Medicine, Kiryat Ono, Tel Aviv, Israel

    Ella Been

  2. Departmento de Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco/Euskal Herriko Unibertsitatea (UPV/EHU), IKERBASQUE, Basque Foundation for Science, Bilbao, Spain, Centro Mixto UCM-ISCIII de Evolución y, Comportamiento Humanos Avda. Monforte de Lemos, Leioa, Madrid, Spain

    Asier Gómez-Olivencia

  3. Departments of Anthropology and Orthopaedics and Sports Medicine, University of Washington, Seattle, WA, USA

    Patricia Ann Kramer

Appendix

Appendix

Taxon

Cervical

Thoracic

Lumbar

Sacral

TV levela

TLb

CTLSc

Total N/TV N (top) and # form. (bottom), freq.d

H. sapiens

Mean

7.00

11.99

4.98

5.22

18.70

16.97

29.19

784/732

CTLS modee

7

12

5

5

–

17

29

63.2%

CTLS 2ndf

7

12

5

6

–

17

30

14.9%

CTLS+TV modeg

7

12

5

5

19

17

29

36.6%

CTLS+TV 2ndh

7

12

5

5

18

17

29

20.4%

CTLS+TV 3rd

7

12

5

6

19

17

30

10.9%

Range

(6–7.5)

(11–13)

(4–6)

(4–7)

(18–20)

(15–18)

(27–31)

39/75

P. paniscus

Mean

7.01

13.51

3.55

6.35

20.17

17.06

30.42

72/51

CTLS mode

7

13

4

6

–

17

30

18.2%

CTLS 2nd

7

14

3

7

–

17

31

10.9%

CTLS+TV mode

7

13

4

6

20

17

30

11.8%

Range

(6.5–8)

(12.5–14)

(2–4)

(5–8)

(19–21)

(16–18)

(28–32)

24/35

P. troglodytes

Mean

7.00

13.11

3.67

5.69

19.78

16.77

29.46

526/342

CTLS mode

7

13

4

6

–

17

30

28.5%

CTLS 2nd

7

13

4

5

–

17

29

21.0%

CTLS 3rd

7

13

3

6

–

16

29

13.6%

CTLS+TV mode

7

13

4

6

20

17

30

24.3%

CTLS+TV 2nd

7

13

4

5

20

17

29

16.4%

Range

(6–7.5)

(12–14)

(2–5)

(4–7)

(18–20)

(15–18)

(28–31)

45/60

P. t. schweinfurthii

Mean

7.01

13.15

3.83

5.84

19.81

16.99

29.84

52/44

CTLS mode

7

13

4

6

–

17

30

45.8%

CTLS 2nd

7

13

4

5

–

17

29

10.4%

CTLS+TV mode

7

13

4

6

20

17

30

36.4%

CTLS+TV 2nd

7

13

4

6

19

17

30

11.4%

Range

(7–7.5)

(12–14)

(2–5)

(4.5–7)

(19–21)

(16–18)

(28–31)

19/22

P. t. troglodytes

Mean

7.00

13.06

3.72

5.65

19.74

16.78

29.44

241/203

CTLS mode

7

13

4

6

–

17

30

29.5%

CTLS 2nd

7

13

4

5

–

17

29

20.5%

CTLS 3rd

7

13

3

6

–

16

29

12.0%

CTLS+TV mode

7

13

4

6

20

17

30

20.2%

CTLS+TV 2nd

7

13

4

5

20

17

29

14.8%

Range

(6.5–7.5)

(12–14)

(2–5)

(4–7)

(18–21.5)

(15–18)

(28–31)

35/50

P. t. verus

Mean

6.98

13.23

3.44

5.73

19.71

16.67

29.38

24/17

CTLS mode

7

13

3

6

–

16

29

8.5%

CTLS 2nd

7

13

4

5

–

17

29

16.7%

CTLS 3rd

7

13

4

6

–

17

30

16.7%

CTLS+TV mode

7

13

4

6

20

17

30

23.5%

CTLS+TV 2nd

7

13

4

5

20

17

29

17.6%

CTLS+TV 3rd

7

13

3

6

20

16

29

11.8%

CTLS+TV 4th

7

14

3

6

20

17

30

11.8%

Range

(6.5–7)

(13–14)

(3–4)

(5–7)

(18.5–20)

(16–18)

(28–31)

10/10

G. gorilla

Mean

6.99

13.03

3.53

5.59

20.01

16.57

29.16

375/319

CTLS mode

7

13

4

5

–

17

29

24.0%

CTLS 2nd

7

13

3

6

–

16

29

21.9%

CTLS 3rd

7

13

4

6

–

17

30

16.3%

CTLS+TV mode

7

13

4

5

20

17

29

17.9%

CTLS+TV 2nd

7

13

3

6

20

16

29

15.7%

Range

(6–7)

(12–15)

(2–5)

(4–8)

(18–23)

(16–18)

(27–31.5)

42/81

G. beringei

Mean

6.99

12.90

3.16

5.96

19.81

16.06

29.01

126/83

CTLS mode

7

13

3

6

–

16

29

71.4%

CTLS+TV mode

7

13

3

6

20

16

29

39.8%

CTLS+TV 2nd

7

13

3

6

19

16

29

19.3%

Range

(6–7)

(12–13)

(2–4)

(5–7)

(17–23)

(15–17)

(27–30)

14/24

G. b. beringei

Mean

6.99

12.86

3.19

5.92

19.95

16.06

28.96

71/41

CTLS mode

7

13

3

6

–

16

29

64.7%

CTLS 2nd

7

12

4

6

–

16

29

11.8%

CTLS+TV mode

7

13

3

6

20

16

29

39.0%

CTLS+TV 2nd

7

13

3

6

19

16

29

12.2%

Range

(6–7)

(12–13)

(3–4)

(5–7)

(18–23)

(15–17)

(28–30)

12/19

G. b. graueri

Mean

7.00

12.96

3.00

6.07

20.04

15.96

29.04

28/27

CTLS mode

7

13

3

6

–

16

29

82.1%

CTLS 2nd

7

13

3

7

–

16

30

10.7%

CTLS 3rd

7

13

3

6

20

16

29

44.4%

CTLS+TV mode

7

13

3

6

19

16

29

18.5%

CTLS+TV 2nd

7

13

3

6

22

16

29

11.1%

Range

–

(12–13)

(2–4)

(5–7)

(18–23)

(15–16)

(27–30)

4/9

Pongo

Mean

6.98

11.95

4.01

5.12

19.13

15.96

28.07

330/163

CTLS mode

7

12

4

5

–

16

28

39.0%

CTLS 2nd

7

12

4

6

–

16

29

14.0%

CTLS+TV mode

7

12

4

5

19

16

28

30.1%

Range

(6–7)

(11–13)

(3–5)

(4–7)

(18–21)

(14.5–17)

(26–30)

39/53

P. pygmaeus

Mean

6.98

11.90

4.02

5.24

19.07

15.92

28.15

148/83

CTLS mode

7

12

4

5

–

16

28

33.6%

CTLS 2nd

7

12

4

6

–

16

29

16.8%

CTLS+TV mode

7

12

4

5

19

16

28

27.7%

CTLS+TV 2nd

7.0

12.0

4.0

6.0

19

16

29

13.3%

Range

(6–7)

(11–13)

(3–5)

(4–7)

(18–20)

(14.5–17)

(27–30)

31/36

P. abelii

Mean

7.0

12.1

4.0

5.0

19.3

16.10

28.11

50/32

CTLS mode

7

12

4

5

–

16

28

47.8%

CTLS+TV mode

7

12

4

5

19

16

28

28.1%

CTLS+TV 2nd

7

12

4

4

19

16

27

12.5%

Range

(6.5–7)

(11–13)

(3–5)

(4–6)

(18–21)

(15–17)

(27–30)

12/17

S. syndactylus

Mean

7.00

13.11

4.53

4.70

19.83

17.64

29.34

87/45

CTLS mode

7

13

5

5

–

18

30

20.2%

CTLS 2nd

7

13

5

4

–

18

29

17.9%

CTLS 3rd

7

13

4

5

–

17

29

17.9%

CTLS+TV mode

7

13

5

5

20

18

30

20.0%

CTLS+TV 2nd

7

13

5

4

20

18

29

13.3%

CTLS+TV 3rd

7

13

4

5

20

17

29

11.1%

Range

–

(12–14)

(4–5)

(4–6)

(18–21)

(16–19)

(27–31)

21/22

H. hoolock

Mean

7

12.9

4.92

4.2

19.5

17.82

29.02

25/4

CTLS mode

7

13

5

4

–

18

29

52.0%

CTLS 2nd

7

13

5

5

–

18

30

12.0%

Range

–

(12–13.5)

(4–6)

(3–5)

(19–20)

(17–19)

(28–30)

10/4

H. lar

Mean

7.00

13.05

5.13

3.86

19.57

18.19

29.04

262/239

CTLS mode

7

13

5

4

–

18

29

47.7%

CTLS+TV mode

7

13

5

4

20

18

29

23.4%

CTLS+TV 2nd

7

13

5

4

19

18

29

23.4%

Range

(6–8)

(12–14)

(4–6)

(3–5)

(18.5–21)

(17–19)

(28–31)

37/50

H. agilis

Mean

7.00

13.00

4.87

4.30

20.00

17.87

29.16

34/20

CTLS mode

7

13

5

4

–

18

29

37.5%

CTLS 2nd

7

13

5

5

–

18

30

18.8%

CTLS+TV mode

7

13

5

4

20

18

29

30.0%

CTLS+TV 2nd

7

13

5

5

20

18

30

15.0%

CTLS+TV 3rd

7

13

5

4

19

18

29

10.0%

Range

–

(12–14)

(4–6)

(3.5–5)

(19–21)

(17–19)

(28–30)

15/12

H. moloch

Mean

7.00

13.06

4.77

4.66

19.50

17.83

29.49

33/6

CTLS mode

7

13

5

5

–

18

30

40.0%

CTLS 2nd

7

13

5

4

–

18

29

26.7%

Range

(7–8)

(12–14)

(4–5.5)

(3.5–6)

(19–20)

(17–19)

(28–30)

11/5

H. klossii

Mean

7.00

13.04

5.04

4.54

20.17

18.08

29.62

13/5

CTLS mode

7

13

5

5

–

18

30

50.0%

CTLS 2nd

7

13

5

4

–

18

29

16.7%

CTLS+TV mode

7

13

5

4

20

18

29

40.0%

CTLS+TV 2nd

7

13

5

5

20

18

30

40.0%

Range

–

(13–13.5)

(4–6)

(4–5)

(20–21)

(17–19)

(28–30)

6/3

H. muelleri

Mean

7.03

13.09

4.70

4.41

19.69

17.79

29.23

35/31

CTLS mode

7

13

5

4

–

18

29

25.7%

CTLS 2nd

7

13

5

5

–

18

30

17.1%

CTLS 3rd

7

13

5

4.5

–

18

30

11.4%

CTLS+TV mode

7

13

5

4

20

18

29

12.9%

CTLS+TV 2nd

7

13

5

5

20

18

30

12.9%

Range

(7–8)

(12–14)

(3.5–6)

(4–5)

(19–21)

(16–19)

(27–31)

18/20

H. pileatus

Mean

7.00

12.44

5.07

4.00

19.67

17.51

28.51

8/2

CTLS mode

7

12

5

4

–

17

28

42.9%

CTLS 2nd

7

13

5

4

–

18

29

28.6%

Range

–

(12–13)

(4.5–6)

(4–4)

19–20

(17–18)

(28–29)

4/2

N. concolor

Mean

7.00

13.12

4.92

4.58

20.21

18.04

29.62

25/6

CTLS mode

7

13

5

4

–

18

29

41.7%

CTLS 2nd

7

13

5

5

–

18

30

29.2%

CTLS+TV mode

7

13

5

5

19.5

18

30

33.3%

Range

–

(12–14.5)

(4–6)

(4–6)

(19–21)

(17–19)

(28–32)

8/5

N. gabrielle

Mean

7.00

13.93

4.89

4.96

21.00

18.82

30.79

21/2

CTLS mode

7

14

5

5

–

19

31

57.1%

CTLS 2nd

7

14

4

5

–

18

30

14.3%

Range

–

(13–14)

(4–6)

(4–5.5)

(21–21)

(18–20)

(30–32)

6/2

T. cristatus

Mean

7.00

12.02

7.00

2.90

17.02

19.02

28.93

50/50

CTLS mode

7

12

7

3

–

19

29

87.5%

CTLS+TV mode

7

12

7

3

17

19

29

87.5%

Range

–

(12–13)

(6.5–7.5)

(2–3)

(17–18)

(19–20)

(28–30)

6/6

P. cynocephalus

Mean

7.00

12.48

6.44

2.90

16.98

18.92

28.82

88/88

CTLS mode

7

13

6

3

–

19

29

42.0%

CTLS 2nd

7

12

7

3

–

19

29

32.0%

CTLS 3rd

7

12

7

2

–

19

28

10.0%

CTLS+TV mode

7

13

6

3

17

19

29

20.5%

CTLS+TV 2nd

7

12

7

3

17

19

29

18.2%

Range

–

(11.5–13)

(5.5–7)

(2–4)

(16–18)

(17–20)

(27–30)

9/12

  1. aTV level: the absolute vertebral level at which the transitional vertebra occurs
  2. bTL: Number of thoracic and lumbar (thoracolumbar) vertebrae
  3. cCTLS: Number of cervical, thoracic, lumbar, and sacral (precaudal) vertebrae
  4. dThe first row of each taxon contains the total N and N of transitional vertebra data (shown as N/N); the last row contains the number of vertebral formulae sampled for the total N and N of transitional vertebra data (shown as N/N); the rows in between the first and last show the frequency of vertebral formulae sampled at ≥10%
  5. eCTLS mode: The modal vertebral formula sampled in each taxon (not including the position of the transitional vertebra)
  6. fCTLS 2nd, etc.: The second (and third, etc.) most common vertebral formula sampled (sans the TV)
  7. gCTLS+TV mode: The modal vertebral formula (with position of the transitional vertebra) sampled in each taxon
  8. hCTLS+TV 2nd, etc.: The second (and third, etc.) most common vertebral formula (with position of the transitional vertebra) sampled in each taxon

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Williams, S.A., Gómez-Olivencia, A., Pilbeam, D.R. (2019). Numbers of Vertebrae in Hominoid Evolution. In: Been, E., Gómez-Olivencia, A., Ann Kramer, P. (eds) Spinal Evolution. Springer, Cham. https://doi.org/10.1007/978-3-030-19349-2_6

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