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A new skeleton of the late Oligocene “Enspel cormorant”—from Oligocorax to Borvocarbo, and back again

A new specimen of the cormorant species from the late Oligocene (MP 28) fossil site Enspel in Germany—the third to be discovered at this site—is described. Previously unknown or controversial features bearing on the taxonomic and phylogenetic affinities of this bird are identified, which do not support the assignment of the Enspel cormorant to the taxon Borvocarbo. The species is therefore here, tentatively, transferred back to Oligocorax, in which it was first placed. The new osteological data confirm that ?Oligocorax stoeffelensis is a stem group representative of Phalacrocoracidae, and the species exhibits a previously unrecognised derived morphology of the pectoral girdle bones in that the furcula is fused with the apex carinae of the sternum, a feature absent in modern cormorants. A phylogenetic analysis revealed two alternative positions for ?O. stoeffelensis relative to other stem group Phalacrocoracidae, supporting a sister group relationship either to the clade including Nambashag, Nectornis and crown group Phalacrocoracidae or to a clade including ?O. stoeffelensis and crown group Phalacrocoracidae to the exclusion of Nambashag and Nectornis.

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I thank M. Wuttke for enabling study of the fossil and M. Poschmann (both Generaldirektion Kulturelles Erbe Rheinland-Pfalz, Mainz, Germany) for its excellent preparation. I am further indebted to T. Worthy and V. De Pietri (both Flinders University, Adelaide) for providing photographs of the humerus, coracoid, and tarsometatarsus of Nectornis miocaenus. S. Tränkner (SMF) took the photographs, and reviews by T. Worthy and U. Göhlich (Naturhistorisches Museum Wien) improved the manuscript.

1. Ornithological Section, Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325, Frankfurt am Main, Germany

   Gerald Mayr

Correspondence to Gerald Mayr.

This article is a contribution to the special issue “The Fossil-Lagerstaette Enspel—reconstructing the palaeoenvironment with new data on fossils and geology”

1. 1.

   Upper beak, praemaxilla with sharply hooked tip: no (0), yes (1). For ?O. stoeffelensis, this feature has been inferred from the dorsal curvature of the tip of the beak (see description).

2. 2.

   Upper beak, marked furrows distal of nasal openings: absent (0), present (1).

3. 3.

   Skull, processus paroccipitales very prominent and strongly protruding in caudal direction: no (0), yes (1).

4. 4.

   Skull, septum interorbitale greatly reduced, large fonticulus orbitocranialis in caudal wall of orbit: no (0); yes (1).

5. 5.

   Skull, processus styloideus: absent (0), present (1).

6. 6.

   Skull, additional crest caudal of crista nuchalis transversa: absent (0), present (1).

7. 7.

   Mandible with marked tubercle at insertion area of musculus adductor mandibulae externus, pars profunda: no (0), yes (1).

8. 8.

   8th-11th cervical vertebrae: processus carotici ankylosed along midline, forming an osseous canal: no (0), yes (1).

9. 9.

   Caudalmost thoracic vertebrae markedly opisthocoelous, i.e. cranial articular surface convex: no (0), yes (1).

10. 10.

    Furcula, processus acromialis measuring more than width of facies articularis acrocoracoidea: no (0), yes (1).

11. 11.

    Coracoid, processus procoracoideus well-developed, medial prominence as far as to medial margin of processus acrocoracoideus: no (0), yes (1).

12. 12.

    Coracoid, facies articularis humeralis with laterally projecting omal portion: no (0), yes (1).

13. 13.

    Coracoid, extremitas sternalis, linea intermuscularis, distal section: straight (0), markedly medially curved (1).

14. 14.

    Humerus, impressio coracobrachialis very marked and ventroproximally bordered by a marked edge formed by intumescentia humeri: no (0), yes (1).

15. 15.

    Humerus, crista bicipitalis very long, reaching as far distally as distal end of crista deltopectoralis: no (0), yes (1).

16. 16.

    Ulna, processus cotylaris dorsalis forming a distinct distal hook: no (0), yes (1). The condition in Nectornis miocaenus has been assessed based on the figures in Göhlich and Mourer-Chauviré (2010).

17. 17.

    Phalanx digiti minoris, measuring less than half of length of phalanx proximalis digiti majoris: yes (0), no (1).

18. 18.

    Pelvis, spina dorsolateralis ilii: greatly elongated and narrow: no (0), yes (1).

19. 19.

    Femur, short and stocky, with ratio length: proximal width being significantly less than 4.0: no (0), yes (1).

20. 20.

    Patella greatly enlarged and massive: no (0), yes (1).

21. 21.

    Tibiotarsus, proximal end, lateral portion of crista patellaris with proximal projection (Mayr 2007: fig. 6I): no (0), yes (1).

22. 22.

    Tibiotarsus, distal end, condylus medialis projecting much farther distally than condylus lateralis: no (0), yes (1).

23. 23.

    Tarsometatarsus short and stocky, ratio length: distal width less than 4.0, length of bone distinctly less than carpometacarpus length: no (0), yes (1).

24. 24.

    Tarsometatarsus, distinct furrow for tendon of musculus extensor hallucis longus on medial surface of shaft: absent (0), present, as wide or wider than shaft width (1), present, narrow, measuring less than the shaft width (2) (Worthy 2011: character 101).

25. 25.

    Tarsometatarsus, hypotarsus with tendon of musculus flexor hallucis longus enclosed in bony canal: yes (0), no (1). The tendon of musculus flexor hallucis longus is also enclosed in Pelecanidae, Balaenicipitidae and Scopidae, and this condition is likely to be plesiomorphic for Suloidea.

26. 26.

    Hypotarsus, crista medialis strongly plantarly projected, dorsoplantar depth of proximal end of tarsometatarsus greater than mediolateral width: no (0), yes (1).

27. 27.

    Hypotarsus, crista medialis forming distally directed hook (Mayr 2001): no (0), yes (1).

28. 28.

    Tarsometatarsus, trochlea metatarsi II distinctly longer than trochlea metatarsi IV, reaching as far distally as trochlea metatarsi III: yes (0), no (1).

29. 29.

    Fourth toe much longer than third: no (0), yes (1).

30. 30.

    Ungual phalanx of fourth toe very narrow: no (0), yes (1).

31. 31.

    Ungual phalanges with far distally situated tuberculum flexorium: no (0), yes (1).

Mayr, G. A new skeleton of the late Oligocene “Enspel cormorant”—from Oligocorax to Borvocarbo, and back again. Palaeobio Palaeoenv 95, 87–101 (2015). https://doi.org/10.1007/s12549-014-0167-7

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• Received: 21 October 2013

• Revised: 17 December 2013

• Accepted: 28 April 2014

• Published: 03 July 2014

• Issue Date: March 2015

• DOI: https://doi.org/10.1007/s12549-014-0167-7

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