Table 1.
Dimensions (in millimeters) of selected bones of Anhinga pannonica from the Hammerschmiede clay pit in comparison to fossil and extant Anhingidae (of Giganhinga kikuyensis and Meganhinga chilensis no comparable measurements were published, but these species are much larger than A. pannonica).
Fig 1.
Section of the excavation plan Hammerschmiede level HAM 4 (excavation year 2019).
Black dots represent vertebrate fossil specimens (black stripes denote the orientation of elongated objects). Excavated bird bones are shown with green triangles, and Anhinga pannonica bones, most probably belonging to the same individual, are highlighted with red stars (specimen GPIT/AV/00215 is a surface find from this area without coordinates). Associated bones of A. pannonica are arranged over a distance of nine meters parallel to the reconstructed flow direction (SSW-NNE) of the river.
Fig 2.
5th or 6th cervical vertebra of Anhinga pannonica from the late Miocene (MN 8) of the Hammerschmiede clay pit near Pforzen, Germany (A, C, D, F) in comparison to the holotype of Anhinga pannonica (B, E, G; from [25], original labeling removed) and the 4th (H–M) and 6th (N–S) cervical vertebrae of extant A. anhinga (SMF 9967) and A. rufa (SMF 9106). A, B, J, K, P, Q: dorsal view; C, right lateral view; D, E, H, I, N, O: left lateral view; F, G, L, M, R, S: ventral view. Abbreviations: car, sulcus caroticus; cir, circular expansion of vertebral corpus; fac, facies articularis; fcd, facies articularis caudalis; fcr, facies articularis cranialis; for, foramen delimited by the ridge-like shelf along the ventrolateral margin of the vertebral corpus; pcs, processus costalis; rdg, ridge bordering sulcus caroticus; slt, slit separating zygapophyses caudales; zcd, zygapophysis caudalis. The scale bar equals 10 mm (the size of the A. pannonica holotype is based on the measurements in [25]).
Fig 3.
Various postcranial bones of Anhinga pannonica from the late Miocene (MN 8) of the Hammerschmiede clay pit near Pforzen, Germany.
A–C, thoracic vertebra (GPIT/AV/00215) in cranial (A), left lateral (B), and caudal (C) view. D–H, extremitas omalis of left coracoid (GPIT/AV/00145) in dorsal (D), lateral (E), medial (F), ventromedial (G), and ventral (H) view. I–K, distal end of right humerus (GPIT/AV/00217) in cranial (I), caudal (J), and distal (K) view. L–N, proximal end of right ulna (GPIT/AV/00127) in caudodorsal (L), cranial (M), and cranioventral (N) view. O, Anhinga anhinga (SMF 9967), proximal end of right ulna in cranioventral view. P, Phalacrocorax carbo (Phalacrocoracidae; SMF 2939), proximal end of right ulna in cranioventral view. Q, R, distal end of right ulna (GPIT/AV/00127) in dorsal (Q) and ventral (R) view. S, A. anhinga (SMF 9967), distal end of right ulna in ventral view. T, P. carbo (SMF 2939), distal end of right ulna in ventral view. Abbreviations: cdd, condylus dorsalis; cdv, condylus ventralis; csc, cotyla scapularis; ctd, cotyla dorsalis; ctv, cotyla ventralis; fac, facies articularis clavicularis; fcd, facies articularis caudalis; fcr, facies articularis cranialis; for, foramen in vertebral corpus; olc, olecranon; ppc, processus procoracoideus; prg, papilla remigalis; tbc, tuberculum carpale. The scale bars equal 10 mm.
Fig 4.
Major limb and pectoral girdle elements of Anhinga pannonica from the late Miocene (MN 8) of the Hammerschmiede clay pit near Pforzen, Germany.
A, B, partial right coracoid (GPIT/AV/00223) in dorsal (A) and ventral (B) view. C, right coracoid of Anhinga anhinga (SMF 9967) in dorsal view. D–F, extremitas omalis of right coracoid GPIT/AV/00223 in dorsomedial (D), lateral (E) and ventromedial (F) view. G–I, partial right carpometacarpus (GPIT/AV/00216). in dorsal (G), ventral (H), and distal (I) view. J, right carpometacarpus of A. rufa (SMF 9106) in ventral view. K, L, right carpometacarpus of A. anhinga (SMF 9967) in ventral (K) and distal (L) view. M, N, right humerus (GPIT/AV/00217) in cranial (M) and caudal (N) view. O, right humerus of A. rufa (SMF 9106) in caudal view. P, right humerus of A. anhinga (SMF 9967) in caudal view. Q, R, proximal portion of right femur (GPIT/AV/00220) in cranial (Q) and caudal (R) view. S, T, left femur lacking distal end (GPIT/AV/00264) in caudal (S) and cranial (T) view. U, left femur (cranial view) of A. melanogaster (SMF 19890). V, left femur (cranial view) of A. anhinga (SMF 9967). W, left femur (cranial view) of Phalacrocorax carbo (Phalacrocoracidae; SMF 2939). X–Z, distal end of right tibiotarsus (GPIT/AV/00198) in caudal (X), cranial (Y), and distal (Z) view. AA, distal end of right tibiotarsus of A. rufa (SMF 9106) in cranial view. BB, distal end of right tibiotarsus of P. carbo (SMF 2939) in cranial view. Abbreviations: cbp, crita bicipitalis; cdf, crus dorsale fossae; cdl, condylus lateralis; cdm, condylus medialis; cdp, crista deltopectoralis; ext, sulcus extensorius; fib, fibula; flg, ridge-like flange on distal end of os metacarpale majus; flx, processus flexorius. The scale bars equal 10 mm; same scale bar for all images except I, L, and Z.
Fig 5.
Fossils of Anhinga pannonica from the middle Miocene (MN 5) of Regensburg-Dechbetten near Regensburg, Germany.
A–E, Left coracoid (BSPG 2008 LI 671; holotype of Phalacrocorax praecarbo von Ammon, 1918) in dorsal (A), dorsolateral (B), medial (C), ventromedial (D), and ventral (E) view. F, Left coracoid of the extant Anhinga anhinga (SMF 9967) in medial view. G, Partial left carpometacarpus (BSPG 2008 LI 607; holotype of Ardea brunhuberi von Ammon, 1918) in cranioventral view; H, the former specimen as it was figured by von Ammon [47]. I, referred left carpometacarpus of A. pannonica from the late Miocene of Brusturi in Romania (mirrored to ease comparisons and original labeling removed; from [25]). J–P, cervical vertebra (BSPG 2008 LI 676; holotype of Botaurites avitus von Ammon, 1918) in different views (J: cranial; K: ventral [cranial portion]/right lateral [caudal portion]; M: dorsal [cranial portion]/left lateral [caudal portion]; N: left lateral [cranial portion]/ventral [caudal portion]; O: right lateral [cranial portion]/dorsal [caudal portion]); L and P are from [47] and show the original condition of the bone in ventral (L) and right lateral (P) view; the arrows in K and M–O indicate the line of breakage separating the incorrectly reassembled cranial and distal portions of the vertebra. Abbreviations: car, canalis caroticus; fac, facies articularis clavicularis; fcd, facies articularis caudalis; ppc, processus procoracoideus. The scale bar equals 10 mm (the size of H, I, L, and P is based on measurements in [25] and [47].
Fig 6.
Humeri of extant and fossil Anhingidae and Phalacrocoracidae.
A, B, Anhinga pannonica from the Hammerschmiede clay pit (GPIT/AV/00217), C, D, Phalacrocorax intermedius from the early Miocene of France; from ([52]: pl. 43), E, F, extant Phalacrocorax carbo (SMF 2939), and G, H, extant Anhinga anhinga (SMF 9967); the bones are shown in caudal (A, C, E, G) and cranial (B, D, F, H) view. The arrows indicate the distal ends of the crista bicipitalis and crista deltopectoralis. Note the long crista deltopectoralis and poorly developed crus dorsale fossae of Phalacrocorax intermedius. Abbreviations: cbp, crista bicipitalis; cdf, crus dorsale fossae; cdp, crista deltopectoralis. The scale bars equal 10 mm; the size of P. intermedius was inferred from ([52]: pl. 43), which depicts the fossil in natural size).
Fig 7.
Geographic and stratigraphic distribution of Anhinga pannonica in Europe and Africa.
Early middle Miocene (yellow stars): 1 –Hambach (Germany), 2 –Regensburg-Dechbetten (Germany). Late middle Miocene (green star): 3 –Ngorora Formation (Kenya). Early late Miocene (red stars): 4 –Hammerschmiede (Germany), 5 –Götzendorf (Austria), 6 –Brusturi/Tataros (Romania), 7 –Beglia Formation (Tunisia). Late late Miocene (blue stars): 8 –Sahabi Formation (Libya), 9 –Toros-Menalla (Chad).