447:
1747:
swam by undulating their entire bodies. However, since the transverse processes of pachycetines are nearly as elongated as the vertebral centra themselves, they would be a lot more limited in the movements they could perform. Specifically, the smaller space between the adjacent processes meant there was less space for muscles to contract, which effectively hindered the ability of pachycetines to move their bodies side to side. Because of this, the mobility of these whales would be much lower than that of other basilosaurids and likely restricted to undulating their spine dorsoventrally (up-and-down), leading to researchers comparing pachycetines to manatees. At the same time, research also suggests that pachycetines would have been powerful swimmers, despite their low mobility.
1739:
1876:
1772:
1069:
999:
163:
900:
871:
782:
853:
3253:
1065:, which follow those of the thorax, are highly elongated, a trait shared by basilosaurines. Unlike basilosaurines however, the transverse processes that emerge from the sides of the vertebrae are nearly as elongated as the vertebral centra themselves. The surface of the bones is littered with numerous vascular canals, which give them a distinct pattern described as "pitted" or "pockmarked" by researchers.
1725:. The brief period of cooler temperatures between these may be another point of origin for pachycetines, as it was during this time that the sea levels began to rise, leading to the conditions seen during the Bartonian. This interpretation is supported by the divergence time calculated by Antar and colleagues, who suggest that pachycetines split from other archaeocetes some 43 to 42 million years ago.
120:
137:
809:
746:
758:
990:
821:
834:
2620:
Bianucci, G.; Lambert, O.; Urbina, M.; Merella, M.; Collareta, A.; Bennion, R.; Salas-Gismondi, R.; Benites-Palomino, A.; Post, K.; de Muizon, C.; Bosio, G.; Di Celma, C.; Malinverno, E.; Pierantoni, P.P.; Villa, I.M.; Amson, E. (2023). "A heavyweight early whale pushes the boundaries of vertebrate
1904:
of
Virginia, which represents shallow marine sediments deposited at a depth of approximately 60–120 ft (18–37 m). Molluscs from this formation further suggest mild to warm water temperatures whereas the terrestrial climate is interpreted as having been tropical with dry winters. There may
1750:
The pachyostosis, osteosclerosis and rib articulations also inform the lifestyle suggested for these animals. Gingerich and colleagues highlight how these traits may be advantageous for a multitude of purposes which may go hand in hand with each other. The increased density could serve as a form of
1033:
being restricted to two cones, with the narrowest points facing each other. The fact that the cancellous bone took on a cone shape was originally noted to be distinct to pachycetines, but later studies argued that this condition is much more widespread across basilosaurids and not diagnostic. While
1746:
The combination of a dense skeletal structure coupled with elongated vertebrae has been used to infer various aspects of the locomotion and lifestyle of pachycetines. For instance, the elongation of the lumbar vertebrae highly resembles what is seen in basilosaurines and suggests that these whales
1799:
had much more gracile teeth that would have been more prone to abrasion and wear, which
Gingerich and colleagues argue rules out the possibilities that it fed on hard-shelled molluscs or obtained food directly from the seafloor (as the ingested sand would contribute to tooth wear). Herbivory as
1135:
Historically, pachycetines were typically regarded as basilosaurids, with some older publications using the antiquated term zeuglodonts. Though this broader relationship is maintained across studies, few make any more definitive statements of how pachycetines may relate to other contemporary
1191:
with many of the core basilosaurids forming their own monophyletic group within this grade. The key difference lies in the position of pachycetines relative to other basilosaurids. Unlike previous publications, Antar and colleagues recovered pachycetines as being more derived than other
2232:
564:
While this would have provided a more accessible basis for future referrals, the proposition was not unanimously accepted by other researchers, with some later publications highlighting the fact that a missing holotype alone is not enough reason to disregard the use of
2408:
1883:
The idea that pachycetines were shallow-water animals that has been derived from the skeletal anatomy finds additional support in the types of environments that fossils of these animals were found in. For instance, the region around
387:
Pachycetines have a long history of research on a genus level, even though the family itself was only named in the early 2020s. This is primarily due to the taxonomic confusion and nomenclatural changes surrounding the genera
1708:
Most pachycetines with well-dated fossil remains suggest that the family was most prominent during the
Bartonian, however, some localities that have yielded pachycetines might extend their range. For instance, the fossils of
2350:
Gol’din, P.; Zvonok, E.; Rekovets, L.; Kovalchuk, A.; Krakhmalnaya, T. (2014). "Basilotritus (Cetacea: Pelagiceti) from the Eocene of
Nagornoye (Ukraine): New data on anatomy, ontogeny and feeding of early basilosaurids".
569:. Supporters of this line of thinking argue that the illustrations provided by Brandt and later authors are sufficient in diagnosing and comparing the material to more recent finds, therefore maintaining the validity of
446:
1913:, with the animals entering deeper waters while travelling. Such behaviour would profit from the increase in ballast and the powerful swimming musculature that is suggested by the anatomy of the vertebrae.
1759:, allowing for a greater intake of air at the surface and the option to expel air to reduce buoyancy after diving. In combination, these could indicate that pachycetines spent a lot of time feeding on the
1148:) as a comparably basal basilosaurid. In their study Basilosauridae, though broadly paraphyletic, still contained a monophyletic branch they referred to as "crown basilosaurids" containing taxa such as
348:
and ribs, making them denser and heavier by comparison. Based on this it has been suggested that these whales lived in shallow waters and that these thickened bones act as a buoyancy control as seen in
1050:. The density of the skeleton is at least somewhat increased by the articulation between the spine and the ribs as well. In pachycetines, the ribs are thought to connect to the vertebrae not through
1909:, a mineral formed between depths of 50–500 m (160–1,640 ft) and most common between 200–300 m (660–980 ft). Gol'din and Zvonok have suggested that this could hint at potential
1717:
remains may be as young as the
Priabonian. Regardless of the first confirmed appearance, it is likely that the origin of Pachycetinae is tied to the global warming that occurred during the
1038:
means that the cortical bone is much denser compared to the same structure in other whales, and contains fewer porosities. Overall these skeletal features have been compared to members of
1029:
forming thickened layers, which are prominent in the vertebrae and ribs of pachycetines. In the ribs in particular, the pachyostosis occurs around the middle of the bone, with
1755:, which is commonly associated with a greater lung capacity, a hypothesis further supported by the rib articulation that would allow the animal to expand and collapse its
1061:
rapidly increase in size, with individual bones appearing trapezoid in shape due to how much larger the posterior surface is relative to the beginning of the element. The
632:
could have been entirely synonymous was entertained, the limited material available for the latter effectively rendered a thorough comparison impossible. For this reason,
1200:
was found to not merely be a basilosaurid more basal than pachycetines but to clade with them, possibly being the basalmost pachycetine. This study also suggests that
517:, archaeocetes even more basal than basilosaurids, although subsequent research repeatedly disagreed on this matter, arguing that the fossils could not be assigned to
2520:"A vertebra of a small species of Pachycetus from the North Sea and its inner structure and vascularity compared with other basilosaurid vertebrae from the same site"
1017:
The most distinct feature of pachycetines, and the one they derive their name from, is the density of their skeleton. The vertebrae and ribs of pachycetines are both
952:
but may represent an additional species according to
Gingerich, Amane and Zhouri. Alternatively, it may not be distinct and instead could simply expand the range of
1823:
at least occasionally fed on sharks and may have even been specialised in them. Following this hypothesis, the teeth were worn down by coming in contact with the
1905:
however be some evidence that pachycetines were not restricted to shallow waters at all times. Ukrainian remains have been recovered from sediments that contain
1091:
No clear size estimates for the various pachycetines have been published, but several papers mention their size relative to other basilosaurids. Both species of
919:
was based on highly fragmentary remains, which were later determined to have been too poorly preserved to allow assignment beyond a family level. Subsequently,
2048:"Skull and partial skeleton of a new pachycetine genus (Cetacea, Basilosauridae) from the Aridal Formation, Bartonian middle Eocene, of southwestern Morocco"
592:
eventually returned to prominence in 2020, when Henk Jan van Vliet and his colleagues noticed the similarities between the German remains and those of
561:
on the other hand was dismissed as being too fragmentary and undiagnostic to be referred to anything more specific than the family
Basilosauridae.
396:, both of which trace their origins to fossils discovered during the late 19th century. The earliest finds were recovered from Ukraine and named
2134:
Uhen, M. D. (1999). "New species of protocetid archaeocete whale, Eocetus wardii (Mammalia: Cetacea) from the middle Eocene of North
Carolina".
2310:
Gol'din, P.; Zvonok, E. (2013). "Basilotritus uheni, a New
Cetacean (Cetacea, Basilosauridae) from the Late Middle Eocene of Eastern Europe".
2692:
848:
is known from across Europe but is not well understood due to the often isolated nature of the fossil finds. It is the largest pachycetine.
652:, whose validity had already come to be questioned by Gol'din and Zvonok as well as Gingerich and Zhouri, was not mentioned in their work.
1738:
667:, meaning that all material from continental Europe was now placed within a single species. The North African remains, previously named
1888:
that has yielded much of the German remains once formed the southern edge of the
Paleogene North Sea, with the environment featuring
3338:
1183:
Following the establishment of Pachycetinae as a recognized clade, the group was recovered in a notably different position in Antar
435:. Recognizing at least two morphotypes, van Beneden suggested that they belonged to two distinct but related species which he named
1800:
displayed by sirenians is also ruled out. Instead, a more generalized diet of fish and marine invertebrates has been proposed for
1742:
The skeletal structure of pachycetines is similar to what is seen in modern sirenians and interpreted as indicating slow movement.
1928:, a minimum of five other species are known from there, together representing a broad range in size. The smallest forms are two
655:
The most recent changes made to the taxonomy of pachycetines came in 2022 with another paper authored by Gingerich. In it, both
573:. Two of the researchers remarking on the matter, Phillip D. Gingerich and Samir Zhouri, subsequently question the validity of
486:
remains from Ukraine. In this publication, he argued for the fossil material to be distinct on a genus level, coining the name
624:, combining the older species' name with the older genus name following their respective priority. While the idea that both
1924:
has been discovered stands out primarily due to the large amount of cetaceans recovered from its sediments. In addition to
907:
Across the long research history, a variety of other species have been named, but few are still regarded as valid species.
671:, were split off and assigned to their own genus based on the discovery of additional remains. With this new genus, dubbed
1168:. A somewhat similar conclusion was reached by van Vliet in 2020 based in part on the well-developed innominate bone of
162:
3343:
2106:
2464:"Eocene cetaceans from the Helmstedt region, Germany, with some remarks on Platyosphys, Basilotritus and Pachycetus"
1875:
1722:
1187:
2023. Similar to Gol'din and Zvonok's prior work basilosaurids were recovered as a paraphyletic group leading up to
2685:
420:
2244:
1084:, which appears to show a much more basal condition than that of other basilosaurids and features a much larger
2186:"An occurrence of the protocetid whale "Eocetus" wardii in the Middle Eocene Piney Point Formation of Virginia"
1160:
they erected formed a clade of their own alongside specimen MUSM 1443, which was found to be more derived than
852:
781:
2569:
Antar, M.S.; Glaohar, A.S.; El-Desouky, H.; Seiffert, E.R.; El-Sayed, S.; Claxton, A.G.; Sallam, H.M. (2023).
1718:
2571:"A diminutive new basilosaurid whale reveals the trajectory of the cetacean life histories during the Eocene"
2487:"An archaeocete vertebra re-examined: indications for a small-sized species of Pachycetus from Spain, Europe"
1819:. These patterns of tooth wear have been compared to those seen in modern whales and seemingly indicate that
976:, though their fragmentary nature means that it is uncertain whether or not they represent distinct species.
541:
given the lack of material to compare more recent finds with. To compensate for this, they erected the genus
490:. Things largely went quiet around both the German and the Ukrainian remains in the following 70 years, with
3333:
2409:""New fauna of archaeocete whales (Mammalia, Cetacea) from the Bartonian middle Eocene of southern Morocco""
401:
353:. Analysis of the teeth suggests that pachycetines had a varying diet, with the robust teeth of the larger
3273:
2518:
van Vliet, H.J.; Bosselaers, M.E.J.; Munsterman, D.K.; Dijkshoorn, M.L.; de Groen, J.J.; Post, K. (2024).
1977:
1478:
777:
in having an overall smaller body size, more gracile teeth and a smaller skull in proportion to the body.
1893:
3305:
2274:
Gritsenko, V. (2001). "New species Platiosphys einori (Archaeoceti) from Oligocenic deposits of Kyiv".
1901:
1864:
1173:
1051:
889:
is known exclusively from the eastern United States. It was originally thought to be part of the genus
3256:
2678:
2632:
2423:
2360:
2319:
2197:
2143:
2059:
1763:
in relatively shallow water, that it was a slow swimmer and/or that it required large oxygen stores.
1578:
683:
to house the two genera. This led to the establishment of Pachycetinae. Finally, Seeley's remains of
2486:
1771:
1314:
501:
Research somewhat picked up again towards the end of the 20th century and the early 2000s. In 1999
405:
502:
2656:
2256:
2213:
2159:
1900:
was deposited at a depth of only around 50–100 m (160–330 ft). The same applies to the
1897:
1058:
409:
157:
3310:
1855:
1787:
Owing to their dense skeleton and the inferred slow speed, it is thought that pachycetines were
1068:
103:
Though the best-dated remains are Bartonian in age, some could suggest its presence during the
2816:
2648:
2602:
2551:
2376:
2248:
2087:
1982:
Mélanges biologiques tirés du Bulletin de l'Académie impériale des sciences de St. Pétersbourg
1811:
has been studied in greater detail. Several types of abrasion have been found on the teeth of
1619:
1602:
1085:
475:
2233:"A new protocetid whale (Cetacea, Archaeoceti) from the late middle Eocene of South Carolina"
2640:
2623:
2592:
2582:
2541:
2531:
2431:
2368:
2327:
2240:
2205:
2151:
2077:
2067:
1917:
1910:
1792:
1062:
596:, lumping the two genera into a single taxon. This effectively left them with five species:
998:
899:
529:
was reexamined in a more critical light. With the holotype presumably lost sometime during
454:
The German remains were eventually recognized as having belonged to archaeocetes. However,
2853:
1835:
fossils. Some Ukrainian localities for instance yield up to 35 species of shark including
1788:
1077:
1030:
648:
were carried over as per the works of Gol'din and Zvonok as well as Gingerich and Zhouri.
458:
rejected the idea that they represented a distinct genus, instead drawing comparisons to
2636:
2597:
2570:
2546:
2519:
2427:
2364:
2323:
2201:
2147:
2063:
3169:
3070:
2873:
2082:
2047:
1828:
1824:
1035:
1022:
925:
342:
299:
237:
3327:
2935:
2924:
2896:
2825:
2805:
2782:
2771:
2660:
2435:
2217:
2185:
1026:
876:
510:
413:
322:
59:
2260:
2163:
363:
suggest a diet of smaller prey items. The clade currently only includes two genera,
3296:
3178:
3102:
3079:
2978:
2964:
2955:
2910:
2903:
2864:
2832:
1929:
1843:
1837:
1437:
1193:
1137:
1018:
621:
538:
338:
119:
2462:
van Vliet, H.; Bosselaers, M.; Vahldiek, B.W.; Paymans, T.; Verheijen, I. (2020).
136:
2072:
1827:
of sharks and other cartilaginous fish. This notion finds support in the diverse
3233:
3185:
3151:
3130:
3095:
3088:
3034:
3020:
3013:
2944:
2846:
2791:
2748:
2738:
2701:
1934:
1554:
1404:
1273:
973:
375:
283:
214:
34:
2670:
2644:
2587:
2372:
3219:
3212:
3144:
3137:
3116:
3048:
3041:
3027:
2992:
2985:
2971:
2917:
2889:
2839:
2798:
2726:
2252:
2155:
1940:
1906:
1849:
1387:
1123:
1117:
791:
729:
514:
455:
369:
359:
318:
313:
strata, but some finds suggest that they could have first appeared during the
276:
269:
108:
79:
44:
2463:
2380:
1713:
stem from formations that span the Lutetian-Bartonian boundary, whereas some
450:
Johann Friedrich Brandt was among the first to describe pachycetine material.
357:
indicating that it possibly fed on sharks, whereas the more gracile teeth of
3226:
3158:
3055:
3006:
2999:
2882:
2764:
2757:
1889:
1885:
1795:. However, the specific diet likely differed significantly between species.
1356:
989:
965:
680:
521:. In 2001 more material from Europe was described, this time under the name
479:
432:
350:
345:
310:
174:
84:
2652:
2606:
2555:
2091:
1896:
seems to suggest a depth of 150 m (490 ft), while the overlying
3290:
3123:
2720:
2276:
Visnyk Heolohila Kyivskyi Natsionalyi Universytet Imeni Tarasa Shevchenka
1760:
1297:
1176:
was performed in this study, nor was one conducted in the description of
1047:
549:
material previously discovered. At the same time, they noted that Uhen's
314:
302:
194:
104:
74:
69:
54:
49:
39:
2536:
3192:
3109:
1950:
1756:
1752:
1662:
1461:
1235:
1188:
1039:
969:
826:
814:
751:
428:
334:
330:
326:
224:
89:
64:
2331:
2732:
2714:
2209:
1867:
preserving 19 species of sharks and an additional 9 species of rays.
1043:
839:
687:
were hypothesized to either represent an additional third species of
306:
204:
184:
3267:
2485:
van Vliet, H.J.; Bosselaers, M.; Paijmans, T.; Calzada, S. (2023).
1874:
1770:
1737:
1067:
961:
898:
763:
445:
2184:
Weems, R.E.; Edwards, L.E.; Osborne, J.E.; Alford, A.A. (2011).
1892:
fed by the rivers that ran into the sea from the continent. The
1879:
During the Lutetian much of Europe was covered by a shallow sea.
3271:
2674:
1136:
archaeocetes within this family, which is commonly found to be
1111:
has been described as "medium-sized" and slightly smaller than
337:, among others. They differ from other basilosaurids in having
1196:
to Neoceti. A further difference to the prior studies is that
636:
was retained, if only for a single vertebra and rib fragment.
530:
321:. Fossils of pachycetines are chiefly known from the southern
1007:
Fossil vertebrae (left) and innominate fragments (right) of
769:
The best preserved and most recently described pachycetine,
2245:
10.1206/0003-0082(2005)480[0001:ANPWCA]2.0.CO;2
1978:"*Über bisher in Russland gefundene Reste von Zeuglodonten"
1115:. It's said to be similar in size to the large protocetid
1034:
pachyostosis describes a thickening of the cortical bone,
903:
A map of localities that have yielded pachycetine fossils.
525:. With more material coming out of Ukraine, the status of
509:
based on cranial and postcranial remains from the eastern
1938:. The largest protocetid from the Aridal Formation is
915:
species, is now thought to be a distinct archaeocete.
1944:
which is thought to be within the same size range as
1057:
Another notable feature of the vertebrae is how the
513:, believing that the material represented a type of
3280:
3202:
3168:
3078:
3069:
2954:
2934:
2872:
2863:
2815:
2781:
2747:
1779:may have been a principal component of the diet of
620:became a new combination and was recognized as the
1140:. In their 2013 paper, Gol'din and Zvonok recover
960:are also known from isolated remains collected in
2231:Geisler, J.H.; Sanders, A.E.; Luo, Z.-X. (2005).
1859:. The American localities that were once home to
1156:itself. Within this grouping, the two species of
679:proper, the team further went on to create a new
553:was remarkably similar, referring the species to
2046:Gingerich, P.D.; Amane, A.; Zouhri, S. (2022).
1815:, which were notably more robust than those of
948:, which has never been formally included under
533:, Pavel Gol'din and Evgenij Zvonok argued that
1121:with a skull that is the same size as that of
1072:The innominate bones of various basilosaurids.
1025:. Pachoystosis describes the condition of the
373:, but a 2023 study suggests that the Peruvian
2686:
1192:basilosaurids, being placed as the immediate
956:to also include the UK. Smaller specimens of
8:
1204:itself may be paraphyletic, as the American
675:, clearly distinct from yet also similar to
1948:. The largest cetacean of the formation is
3268:
3075:
2869:
2693:
2679:
2671:
851:
780:
135:
118:
20:
2596:
2586:
2545:
2535:
2081:
2071:
923:is now simply regarded as Basilosauridae
2513:
2511:
2509:
2507:
2457:
2455:
2453:
2451:
2449:
2447:
2445:
2041:
2039:
2037:
2035:
2033:
2031:
2029:
2027:
2025:
2023:
2021:
2019:
2017:
2015:
937:) is regarded as a potential synonym of
702:
577:(alongside that of the poorly preserved
408:named poorly preserved remains from the
2402:
2400:
2398:
2396:
2394:
2392:
2390:
2179:
2177:
2175:
2173:
2013:
2011:
2009:
2007:
2005:
2003:
2001:
1999:
1997:
1995:
1963:
581:) while naming a species of their own:
2345:
2343:
2341:
2305:
2303:
2301:
2299:
2297:
2295:
2293:
2291:
2289:
482:on fossil archaeocetes, including the
2129:
2127:
2125:
2123:
1971:
1969:
1967:
498:likewise receiving little attention.
255:Gingerich, Amane, & Zouhri, 2022
7:
2407:Gingerich, P.D.; Zouhri, S. (2015).
1080:known from a pachycetine is that of
1054:but through ligaments or cartilage.
1831:fauna of the localities that yield
1594:
1570:
1546:
1539:
1453:
1429:
1379:
1372:
1348:
1289:
1265:
1258:
1251:
1227:
1219:
1042:, a group that includes the modern
309:. The best-dated remains stem from
2114:Carnegie Institution of Washington
944:Things are slightly different for
379:may at least be a close relative.
14:
2416:Journal of African Earth Sciences
911:, the second of the two original
3252:
3251:
2436:10.1016/j.jafrearsci.2015.08.006
997:
988:
869:
832:
819:
807:
756:
744:
317:and may have survived until the
161:
1775:Sharks like those of the genus
474:). Only a year later, in 1936,
431:which he thought belonged to a
1723:middle Eocene climatic optimum
691:or simply another instance of
1:
2107:"A Review of the Archaeoceti"
1095:are described as larger than
305:that lived during the middle
2073:10.1371/journal.pone.0276110
494:slipping into obscurity and
663:were fully integrated into
298:is an extinct subfamily of
3360:
2645:10.1038/s41586-023-06381-1
2588:10.1038/s42003-023-04986-w
2373:10.1016/j.crpv.2013.11.002
1844:Hemiscyllium bruxelliensis
1103:being notably larger than
3247:
2709:
2237:American Museum Novitates
2156:10.1017/s002233600002802x
1660:
1616:
1599:
1592:
1575:
1568:
1551:
1544:
1537:
1475:
1458:
1451:
1434:
1427:
1401:
1384:
1377:
1370:
1353:
1346:
1311:
1294:
1287:
1270:
1263:
1256:
1249:
1232:
1225:
850:
788:
421:Pierre-Joseph Van Beneden
265:
260:
158:Scientific classification
156:
144:A live reconstruction of
143:
134:
126:
117:
23:
3339:Eocene first appearances
1734:Locomotion and lifestyle
1719:Lutetian thermal maximum
1144:(by them referred to as
427:for fossil remains from
404:. Around the same time,
2312:Journal of Paleontology
2190:Journal of Paleontology
2136:Journal of Paleontology
1935:Chrysocetus fouadassaii
1863:were similar, with the
1212:than with the European
545:to include much of the
537:should be considered a
478:published an extensive
402:Johann Friedrich Brandt
2353:Comptes Rendus Palevol
1880:
1784:
1743:
1479:Pontogeneus peruvianus
1073:
904:
451:
3306:Paleobiology Database
1976:Brandt, J.F. (1873).
1932:and the basilosaurid
1902:Piney Point Formation
1878:
1865:Piney Point Formation
1850:Carcharias acutissimu
1774:
1741:
1174:phylogenetic analysis
1071:
1052:synovial articulation
933:(the type species of
902:
881:A smaller species of
449:
127:Skeletal material of
2105:Kellogg, R. (1936).
1715:Pachycetus paulsonii
1704:Evolutionary history
1579:Pachycetus paulsonii
1214:Pachycetus paulsonii
1164:yet more basal than
954:Pachycetus paulsonii
946:"Zeuglodon" wanklyni
939:Pachycetus paulsonii
846:Pachycetus paulsonii
799:Pachycetus paulsonii
685:"Zeuglodon" wanklyni
665:Pachycetus paulsonii
618:Pachycetus paulsonii
383:History of discovery
150:Pachycetus paulsonii
16:Subfamily of mammals
2637:2023Natur.620..824B
2537:10.7717/peerj.16541
2428:2015JAfES.111..273G
2365:2014CRPal..13..267G
2324:2013JPal...87..254G
2202:2011JPal...85..271W
2148:1999JPal...73..512U
2064:2022PLoSO..1776110G
1894:Annenberg Formation
1315:Tutcetus rayanensis
1208:claded closer with
1172:. However, no full
704:
657:Pachycetus robustus
437:Pachycetus robustus
406:Harry Govier Seeley
398:Zeuglodon paulsonii
3344:Eocene extinctions
2468:Cainozoic Research
1911:migrating behavior
1898:Gehlberg Formation
1881:
1785:
1744:
1074:
1059:thoracic vertebrae
921:Platyosphys einori
917:Platyosphys einori
909:Pachycetus humilis
905:
703:
583:Platyosphys aithai
559:Platyosphys einori
523:Platyosphys einori
452:
441:Pachycetus humilis
417:Zeuglodon wanklyni
146:Basilotritus uheni
3321:
3320:
3274:Taxon identifiers
3265:
3264:
3243:
3242:
3065:
3064:
2817:Remingtonocetidae
2631:(7975): 824–829.
2332:10.1666/12-080R.1
1861:Pachycetus wardii
1856:Clerolamna umovae
1793:pursuit predators
1711:Pachycetus wardii
1700:
1699:
1691:
1690:
1682:
1681:
1673:
1672:
1649:
1648:
1640:
1639:
1631:
1630:
1620:Antaecetus aithai
1603:Pachycetus wardii
1526:
1525:
1517:
1516:
1508:
1507:
1499:
1498:
1490:
1489:
1416:
1415:
1335:
1334:
1326:
1325:
1206:Pachycetus wardii
1086:obturator foramen
1082:Pachycetus wardii
1009:Pachycetus wardii
968:off the coast of
897:
896:
861:Pachycetus wardii
736:Antaecetus aithai
669:Pachycetus aithai
476:Remington Kellogg
293:
292:
256:
129:Antaecetus aithai
112:
3351:
3314:
3313:
3301:
3300:
3299:
3269:
3255:
3254:
3076:
2870:
2704:genera by family
2695:
2688:
2681:
2672:
2665:
2664:
2617:
2611:
2610:
2600:
2590:
2566:
2560:
2559:
2549:
2539:
2515:
2502:
2501:
2491:
2482:
2476:
2475:
2459:
2440:
2439:
2413:
2404:
2385:
2384:
2347:
2336:
2335:
2307:
2284:
2283:
2271:
2265:
2264:
2228:
2222:
2221:
2210:10.1666/10-083.1
2181:
2168:
2167:
2131:
2118:
2117:
2111:
2102:
2096:
2095:
2085:
2075:
2058:(10): e0276110.
2043:
1990:
1989:
1973:
1918:Aridal Formation
1871:Paleoenvironment
1838:Scyliorhinus sp.
1789:ambush predators
1595:
1571:
1547:
1540:
1454:
1430:
1380:
1373:
1349:
1290:
1266:
1259:
1252:
1228:
1220:
1063:lumbar vertebrae
1001:
992:
931:Pachycetus uheni
875:
873:
872:
855:
838:
836:
835:
825:
823:
822:
813:
811:
810:
784:
762:
760:
759:
750:
748:
747:
705:
661:Pachycetus uheni
551:"Eocetus" wardii
468:Zeuglodon osiris
423:coined the name
254:
249:
236:
166:
165:
139:
122:
102:
101:
31:
27:Temporal range:
21:
3359:
3358:
3354:
3353:
3352:
3350:
3349:
3348:
3324:
3323:
3322:
3317:
3309:
3304:
3295:
3294:
3289:
3276:
3266:
3261:
3239:
3198:
3164:
3061:
2950:
2930:
2859:
2854:Remingtonocetus
2811:
2777:
2743:
2705:
2699:
2669:
2668:
2619:
2618:
2614:
2568:
2567:
2563:
2517:
2516:
2505:
2489:
2484:
2483:
2479:
2461:
2460:
2443:
2411:
2406:
2405:
2388:
2349:
2348:
2339:
2309:
2308:
2287:
2273:
2272:
2268:
2230:
2229:
2225:
2183:
2182:
2171:
2133:
2132:
2121:
2109:
2104:
2103:
2099:
2045:
2044:
1993:
1975:
1974:
1965:
1960:
1873:
1769:
1736:
1731:
1706:
1701:
1692:
1683:
1674:
1650:
1641:
1632:
1527:
1518:
1509:
1500:
1491:
1417:
1336:
1327:
1133:
1078:innominate bone
1031:cancellous bone
1015:
1014:
1013:
1012:
1004:
1003:
1002:
994:
993:
982:
870:
868:
833:
831:
820:
818:
808:
806:
757:
755:
745:
743:
701:
385:
253:
247:
234:
160:
113:
100:
99:
98:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
37:
29:
28:
25:
17:
12:
11:
5:
3357:
3355:
3347:
3346:
3341:
3336:
3334:Basilosauridae
3326:
3325:
3319:
3318:
3316:
3315:
3302:
3286:
3284:
3278:
3277:
3272:
3263:
3262:
3260:
3259:
3248:
3245:
3244:
3241:
3240:
3238:
3237:
3230:
3223:
3216:
3208:
3206:
3200:
3199:
3197:
3196:
3189:
3182:
3174:
3172:
3170:Basilosaurinae
3166:
3165:
3163:
3162:
3155:
3148:
3141:
3134:
3127:
3120:
3113:
3106:
3099:
3092:
3084:
3082:
3073:
3071:Basilosauridae
3067:
3066:
3063:
3062:
3060:
3059:
3052:
3045:
3038:
3031:
3024:
3017:
3010:
3003:
2996:
2989:
2982:
2975:
2968:
2960:
2958:
2952:
2951:
2949:
2948:
2940:
2938:
2932:
2931:
2929:
2928:
2921:
2914:
2907:
2900:
2893:
2886:
2878:
2876:
2874:Georgiacetinae
2867:
2861:
2860:
2858:
2857:
2850:
2843:
2836:
2829:
2821:
2819:
2813:
2812:
2810:
2809:
2802:
2795:
2787:
2785:
2779:
2778:
2776:
2775:
2768:
2761:
2753:
2751:
2745:
2744:
2742:
2741:
2735:
2729:
2723:
2717:
2710:
2707:
2706:
2700:
2698:
2697:
2690:
2683:
2675:
2667:
2666:
2612:
2561:
2503:
2477:
2441:
2386:
2359:(4): 267–276.
2337:
2318:(2): 254–268.
2285:
2266:
2239:(3480): 1–68.
2223:
2196:(2): 271–278.
2169:
2142:(3): 512–528.
2119:
2097:
1991:
1962:
1961:
1959:
1956:
1872:
1869:
1829:chondrichthyan
1825:placoid scales
1768:
1765:
1735:
1732:
1730:
1727:
1705:
1702:
1698:
1697:
1694:
1693:
1689:
1688:
1685:
1684:
1680:
1679:
1676:
1675:
1671:
1670:
1667:
1666:
1659:
1656:
1655:
1652:
1651:
1647:
1646:
1643:
1642:
1638:
1637:
1634:
1633:
1629:
1628:
1625:
1624:
1615:
1612:
1611:
1608:
1607:
1598:
1593:
1591:
1588:
1587:
1584:
1583:
1574:
1569:
1567:
1564:
1563:
1560:
1559:
1550:
1545:
1543:
1538:
1536:
1533:
1532:
1529:
1528:
1524:
1523:
1520:
1519:
1515:
1514:
1511:
1510:
1506:
1505:
1502:
1501:
1497:
1496:
1493:
1492:
1488:
1487:
1484:
1483:
1474:
1471:
1470:
1467:
1466:
1457:
1452:
1450:
1447:
1446:
1443:
1442:
1433:
1428:
1426:
1423:
1422:
1419:
1418:
1414:
1413:
1410:
1409:
1400:
1397:
1396:
1393:
1392:
1383:
1378:
1376:
1371:
1369:
1366:
1365:
1362:
1361:
1352:
1347:
1345:
1342:
1341:
1338:
1337:
1333:
1332:
1329:
1328:
1324:
1323:
1320:
1319:
1310:
1307:
1306:
1303:
1302:
1293:
1288:
1286:
1283:
1282:
1279:
1278:
1269:
1264:
1262:
1257:
1255:
1250:
1248:
1245:
1244:
1241:
1240:
1231:
1226:
1224:
1218:
1132:
1129:
1036:osteosclerosis
1023:osteosclerotic
1006:
1005:
996:
995:
987:
986:
985:
984:
983:
981:
978:
926:incertae sedis
895:
894:
879:
866:
863:
857:
856:
849:
843:
804:
801:
796:
786:
785:
778:
767:
741:
738:
733:
725:
724:
721:
718:
715:
712:
709:
700:
697:
507:Eocetus wardii
460:Zeuglodon isis
384:
381:
343:osteosclerotic
291:
290:
289:
288:
280:
273:
263:
262:
258:
257:
245:
241:
240:
238:Basilosauridae
232:
228:
227:
222:
218:
217:
212:
208:
207:
202:
198:
197:
192:
188:
187:
182:
178:
177:
172:
168:
167:
154:
153:
141:
140:
132:
131:
124:
123:
115:
114:
95:
94:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
33:
32:
26:
15:
13:
10:
9:
6:
4:
3:
2:
3356:
3345:
3342:
3340:
3337:
3335:
3332:
3331:
3329:
3312:
3307:
3303:
3298:
3292:
3288:
3287:
3285:
3283:
3279:
3275:
3270:
3258:
3250:
3249:
3246:
3236:
3235:
3231:
3229:
3228:
3224:
3222:
3221:
3217:
3215:
3214:
3210:
3209:
3207:
3205:
3201:
3195:
3194:
3190:
3188:
3187:
3183:
3181:
3180:
3176:
3175:
3173:
3171:
3167:
3161:
3160:
3156:
3154:
3153:
3149:
3147:
3146:
3142:
3140:
3139:
3135:
3133:
3132:
3128:
3126:
3125:
3121:
3119:
3118:
3114:
3112:
3111:
3107:
3105:
3104:
3100:
3098:
3097:
3093:
3091:
3090:
3086:
3085:
3083:
3081:
3077:
3074:
3072:
3068:
3058:
3057:
3053:
3051:
3050:
3046:
3044:
3043:
3039:
3037:
3036:
3032:
3030:
3029:
3025:
3023:
3022:
3018:
3016:
3015:
3011:
3009:
3008:
3004:
3002:
3001:
2997:
2995:
2994:
2990:
2988:
2987:
2983:
2981:
2980:
2976:
2974:
2973:
2969:
2967:
2966:
2962:
2961:
2959:
2957:
2953:
2947:
2946:
2942:
2941:
2939:
2937:
2936:Makaracetinae
2933:
2927:
2926:
2925:Pontobasileus
2922:
2920:
2919:
2915:
2913:
2912:
2908:
2906:
2905:
2901:
2899:
2898:
2897:Carolinacetus
2894:
2892:
2891:
2887:
2885:
2884:
2880:
2879:
2877:
2875:
2871:
2868:
2866:
2862:
2856:
2855:
2851:
2849:
2848:
2844:
2842:
2841:
2837:
2835:
2834:
2830:
2828:
2827:
2826:Andrewsiphius
2823:
2822:
2820:
2818:
2814:
2808:
2807:
2806:Himalayacetus
2803:
2801:
2800:
2796:
2794:
2793:
2789:
2788:
2786:
2784:
2783:Ambulocetidae
2780:
2774:
2773:
2772:Ichthyolestes
2769:
2767:
2766:
2762:
2760:
2759:
2755:
2754:
2752:
2750:
2746:
2740:
2736:
2734:
2730:
2728:
2724:
2722:
2718:
2716:
2712:
2711:
2708:
2703:
2696:
2691:
2689:
2684:
2682:
2677:
2676:
2673:
2662:
2658:
2654:
2650:
2646:
2642:
2638:
2634:
2630:
2626:
2625:
2621:morphology".
2616:
2613:
2608:
2604:
2599:
2594:
2589:
2584:
2580:
2576:
2572:
2565:
2562:
2557:
2553:
2548:
2543:
2538:
2533:
2529:
2525:
2521:
2514:
2512:
2510:
2508:
2504:
2499:
2495:
2488:
2481:
2478:
2474:(1): 121–148.
2473:
2469:
2465:
2458:
2456:
2454:
2452:
2450:
2448:
2446:
2442:
2437:
2433:
2429:
2425:
2421:
2417:
2410:
2403:
2401:
2399:
2397:
2395:
2393:
2391:
2387:
2382:
2378:
2374:
2370:
2366:
2362:
2358:
2354:
2346:
2344:
2342:
2338:
2333:
2329:
2325:
2321:
2317:
2313:
2306:
2304:
2302:
2300:
2298:
2296:
2294:
2292:
2290:
2286:
2281:
2277:
2270:
2267:
2262:
2258:
2254:
2250:
2246:
2242:
2238:
2234:
2227:
2224:
2219:
2215:
2211:
2207:
2203:
2199:
2195:
2191:
2187:
2180:
2178:
2176:
2174:
2170:
2165:
2161:
2157:
2153:
2149:
2145:
2141:
2137:
2130:
2128:
2126:
2124:
2120:
2116:(482): 1–366.
2115:
2108:
2101:
2098:
2093:
2089:
2084:
2079:
2074:
2069:
2065:
2061:
2057:
2053:
2049:
2042:
2040:
2038:
2036:
2034:
2032:
2030:
2028:
2026:
2024:
2022:
2020:
2018:
2016:
2014:
2012:
2010:
2008:
2006:
2004:
2002:
2000:
1998:
1996:
1992:
1987:
1983:
1979:
1972:
1970:
1968:
1964:
1957:
1955:
1953:
1952:
1947:
1943:
1942:
1937:
1936:
1931:
1927:
1923:
1919:
1914:
1912:
1908:
1903:
1899:
1895:
1891:
1887:
1877:
1870:
1868:
1866:
1862:
1858:
1857:
1852:
1851:
1846:
1845:
1840:
1839:
1834:
1830:
1826:
1822:
1818:
1814:
1810:
1805:
1803:
1798:
1794:
1790:
1782:
1778:
1773:
1766:
1764:
1762:
1758:
1754:
1748:
1740:
1733:
1728:
1726:
1724:
1720:
1716:
1712:
1703:
1696:
1695:
1687:
1686:
1678:
1677:
1669:
1668:
1665:
1664:
1658:
1657:
1654:
1653:
1645:
1644:
1636:
1635:
1627:
1626:
1623:
1622:
1621:
1614:
1613:
1610:
1609:
1606:
1605:
1604:
1597:
1596:
1590:
1589:
1586:
1585:
1582:
1581:
1580:
1573:
1572:
1566:
1565:
1562:
1561:
1558:
1557:
1556:
1549:
1548:
1542:
1541:
1535:
1534:
1531:
1530:
1522:
1521:
1513:
1512:
1504:
1503:
1495:
1494:
1486:
1485:
1482:
1481:
1480:
1473:
1472:
1469:
1468:
1465:
1464:
1463:
1456:
1455:
1449:
1448:
1445:
1444:
1441:
1440:
1439:
1432:
1431:
1425:
1424:
1421:
1420:
1412:
1411:
1408:
1407:
1406:
1399:
1398:
1395:
1394:
1391:
1390:
1389:
1382:
1381:
1375:
1374:
1368:
1367:
1364:
1363:
1360:
1359:
1358:
1351:
1350:
1344:
1343:
1340:
1339:
1331:
1330:
1322:
1321:
1318:
1317:
1316:
1309:
1308:
1305:
1304:
1301:
1300:
1299:
1292:
1291:
1285:
1284:
1281:
1280:
1277:
1276:
1275:
1268:
1267:
1261:
1260:
1254:
1253:
1247:
1246:
1243:
1242:
1239:
1238:
1237:
1230:
1229:
1222:
1221:
1217:
1215:
1211:
1207:
1203:
1199:
1195:
1190:
1186:
1181:
1179:
1175:
1171:
1167:
1163:
1159:
1155:
1151:
1147:
1143:
1139:
1130:
1128:
1126:
1125:
1120:
1119:
1114:
1110:
1106:
1102:
1098:
1094:
1089:
1087:
1083:
1079:
1070:
1066:
1064:
1060:
1055:
1053:
1049:
1045:
1041:
1037:
1032:
1028:
1027:cortical bone
1024:
1020:
1010:
1000:
991:
979:
977:
975:
971:
967:
963:
959:
955:
951:
947:
942:
940:
936:
932:
928:
927:
922:
918:
914:
910:
901:
892:
888:
884:
880:
878:
867:
864:
862:
859:
858:
854:
847:
844:
842:
841:
829:
828:
816:
805:
802:
800:
797:
795:
794:
793:
787:
783:
779:
776:
773:differs from
772:
768:
766:
765:
753:
742:
739:
737:
734:
732:
731:
727:
726:
722:
719:
716:
713:
710:
707:
706:
698:
696:
694:
690:
686:
682:
678:
674:
670:
666:
662:
658:
653:
651:
647:
643:
639:
635:
631:
627:
623:
619:
615:
611:
607:
603:
599:
595:
591:
586:
584:
580:
576:
572:
568:
562:
560:
556:
552:
548:
544:
540:
536:
532:
528:
524:
520:
516:
512:
511:United States
508:
504:
499:
497:
493:
489:
485:
481:
477:
473:
469:
465:
461:
457:
448:
444:
442:
438:
434:
430:
426:
422:
418:
415:
414:Great Britain
411:
407:
403:
399:
395:
391:
382:
380:
378:
377:
372:
371:
366:
362:
361:
356:
352:
347:
344:
340:
336:
332:
328:
324:
323:United States
320:
316:
312:
308:
304:
301:
297:
286:
285:
281:
279:
278:
274:
272:
271:
267:
266:
264:
259:
252:
246:
243:
242:
239:
233:
230:
229:
226:
223:
220:
219:
216:
213:
210:
209:
206:
203:
200:
199:
196:
193:
190:
189:
186:
183:
180:
179:
176:
173:
170:
169:
164:
159:
155:
151:
147:
142:
138:
133:
130:
125:
121:
116:
110:
106:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
41:
36:
22:
19:
3282:Pachycetinae
3281:
3232:
3225:
3218:
3211:
3204:Pachycetinae
3203:
3191:
3184:
3179:Basilosaurus
3177:
3157:
3150:
3143:
3136:
3129:
3122:
3115:
3108:
3103:Cynthiacetus
3101:
3094:
3087:
3080:Dorudontinae
3054:
3047:
3040:
3033:
3026:
3019:
3012:
3005:
2998:
2991:
2984:
2979:Crenatocetus
2977:
2970:
2965:Aegyptocetus
2963:
2956:Protocetinae
2943:
2923:
2916:
2911:Natchitochia
2909:
2904:Georgiacetus
2902:
2895:
2888:
2881:
2865:Protocetidae
2852:
2845:
2838:
2833:Attockicetus
2831:
2824:
2804:
2797:
2790:
2770:
2763:
2756:
2739:Artiodactyla
2628:
2622:
2615:
2581:(707): 707.
2578:
2574:
2564:
2527:
2523:
2497:
2493:
2480:
2471:
2467:
2419:
2415:
2356:
2352:
2315:
2311:
2279:
2275:
2269:
2236:
2226:
2193:
2189:
2139:
2135:
2113:
2100:
2055:
2051:
1985:
1981:
1949:
1945:
1939:
1933:
1925:
1921:
1915:
1882:
1860:
1854:
1848:
1842:
1836:
1832:
1820:
1816:
1812:
1808:
1807:The diet of
1806:
1801:
1796:
1791:rather than
1786:
1780:
1777:Hemiscyllium
1776:
1749:
1745:
1729:Paleobiology
1714:
1710:
1707:
1661:
1618:
1617:
1601:
1600:
1577:
1576:
1553:
1552:
1477:
1476:
1460:
1459:
1438:Basilosaurus
1436:
1435:
1403:
1402:
1386:
1385:
1355:
1354:
1313:
1312:
1296:
1295:
1272:
1271:
1234:
1233:
1213:
1209:
1205:
1201:
1197:
1194:sister group
1184:
1182:
1177:
1169:
1165:
1161:
1158:Basilotritus
1157:
1154:Basilosaurus
1153:
1149:
1146:Basilotritus
1145:
1141:
1138:paraphyletic
1134:
1122:
1116:
1112:
1108:
1104:
1101:P. paulsonii
1100:
1096:
1092:
1090:
1081:
1075:
1056:
1019:pachyostotic
1016:
1008:
957:
953:
949:
945:
943:
938:
935:Basilotritus
934:
930:
924:
920:
916:
912:
908:
906:
890:
886:
882:
860:
845:
830:
817:
798:
790:
789:
774:
770:
754:
735:
728:
693:P. paulsonii
692:
688:
684:
676:
672:
668:
664:
660:
656:
654:
649:
645:
641:
637:
633:
629:
626:P. paulsonii
625:
622:type species
617:
613:
609:
605:
601:
598:P. paulsonii
597:
593:
589:
587:
582:
578:
574:
570:
566:
563:
558:
555:Basilotritus
554:
550:
546:
543:Basilotritus
542:
539:nomen dubium
534:
526:
522:
518:
506:
503:Mark D. Uhen
500:
495:
491:
487:
483:
471:
467:
464:Basilosaurus
463:
459:
453:
440:
436:
424:
416:
397:
393:
389:
386:
374:
368:
364:
358:
354:
339:pachyostotic
300:basilosaurid
296:Pachycetinae
295:
294:
282:
275:
268:
251:Pachycetinae
250:
221:Infraorder:
215:Artiodactyla
149:
145:
128:
24:Pachycetinae
18:
3234:Platyosphys
3186:Basiloterus
3152:Supayacetus
3131:Pontogeneus
3096:Chrysocetus
3089:Ancalecetus
3035:Qaisracetus
3021:Phiomicetus
3014:Peregocetus
2945:Makaracetus
2847:Kutchicetus
2792:Ambulocetus
2749:Pakicetidae
2702:Archaeocete
2575:Commun Biol
2422:: 273–286.
1930:protocetids
1555:Supayacetus
1405:Ancalecetus
1274:Chrysocetus
1223:Pelagiceti
1198:Supayacetus
1162:Supayacetus
980:Description
974:Netherlands
634:P. robustus
630:P. robustus
602:P. robustus
594:Platyosphys
571:Platyosphys
567:Platyosphys
547:Platyosphys
535:Platyosphys
527:Platyosphys
496:Platyosphys
488:Platyosphys
484:"Zeuglodon"
410:Barton Clay
390:Platyosphys
376:Supayacetus
284:Supayacetus
244:Subfamily:
3328:Categories
3297:Q121338969
3220:Pachycetus
3213:Antaecetus
3145:Stromerius
3138:Saghacetus
3117:Masracetus
3049:Takracetus
3042:Rodhocetus
3028:Protocetus
2993:Gaviacetus
2986:Dhedacetus
2972:Artiocetus
2918:Pappocetus
2890:Babiacetus
2840:Dalanistes
2799:Gandakasia
2727:Vertebrata
2725:Subphylum
2530:: e16541.
2253:4630500767
1988:: 111–112.
1958:References
1946:Antaecetus
1941:Pappocetus
1926:Antaecetus
1922:Antaecetus
1907:glauconite
1833:Pachycetus
1821:Pachycetus
1817:Antaecetus
1813:Pachycetus
1809:Pachycetus
1802:Antaecetus
1797:Antaecetus
1781:Pachycetus
1388:Saghacetus
1210:Antaecetus
1202:Pachycetus
1178:Antaecetus
1142:Pachycetus
1124:Saghacetus
1118:Pappocetus
1109:Antaecetus
1097:Antaecetus
1093:Pachycetus
958:Pachycetus
950:Pachycetus
913:Pachycetus
883:Pachycetus
865:Bartonian
803:Bartonian
792:Pachycetus
775:Pachycetus
771:Antaecetus
740:Bartonian
730:Antaecetus
689:Pachycetus
677:Pachycetus
673:Antaecetus
590:Pachycetus
515:protocetid
505:described
492:Pachycetus
472:Saghacetus
456:Oskar Kuhn
425:Pachycetus
394:Pachycetus
370:Pachycetus
365:Antaecetus
360:Antaecetus
355:Pachycetus
319:Priabonian
277:Pachycetus
270:Antaecetus
109:Priabonian
3227:Perucetus
3159:Zygorhiza
3056:Togocetus
3007:Maiacetus
3000:Indocetus
2883:Aegicetus
2765:Nalacetus
2758:Pakicetus
2661:260433513
2381:1631-0683
2218:129160656
1890:estuaries
1886:Helmstedt
1357:Zygorhiza
1170:P. wardii
1166:Zygorhiza
1131:Phylogeny
1113:P. wardii
1105:P. wardii
1076:The only
966:North Sea
887:P. wardii
717:Location
681:subfamily
650:P. einori
646:P. wardii
642:P. aithai
614:P. aithai
606:P. wardii
588:The name
579:P. einori
557:as well.
480:monograph
433:mysticete
351:sirenians
346:vertebrae
311:Bartonian
303:cetaceans
181:Kingdom:
175:Eukaryota
30:Bartonian
3291:Wikidata
3257:Category
3124:Ocucajea
2733:Mammalia
2721:Chordata
2715:Animalia
2713:Kingdom
2653:37532931
2607:37563270
2598:10415296
2556:38774542
2547:11107809
2282:: 17–20.
2261:54060094
2164:85977537
2092:36288346
2052:PLOS ONE
1761:seafloor
1721:and the
1298:Ocucajea
1048:manatees
972:and the
964:and the
711:Species
638:P. uheni
610:P. uheni
575:P. uheni
315:Lutetian
231:Family:
205:Mammalia
195:Chordata
191:Phylum:
185:Animalia
171:Domain:
105:Lutetian
3193:Eocetus
3110:Dorudon
2719:Phylum
2633:Bibcode
2500:(1–16).
2494:Deinsea
2424:Bibcode
2361:Bibcode
2320:Bibcode
2198:Bibcode
2144:Bibcode
2083:9604876
2060:Bibcode
1951:Eocetus
1757:ribcage
1753:ballast
1663:Neoceti
1462:Dorudon
1236:Eocetus
1189:Neoceti
1150:Dorudon
1099:, with
1040:Sirenia
970:Belgium
891:Eocetus
827:Germany
815:Ukraine
752:Morocco
699:Species
519:Eocetus
429:Germany
335:Germany
331:Morocco
327:Ukraine
261:Genera
225:Cetacea
211:Order:
201:Class:
96:↓
3311:462731
2737:Order
2731:Class
2659:
2651:
2624:Nature
2605:
2595:
2554:
2544:
2379:
2259:
2251:
2216:
2162:
2090:
2080:
1920:where
1185:et al.
1044:dugong
874:
840:Russia
837:
824:
812:
761:
749:
723:Image
720:Notes
708:Genus
466:) and
419:while
307:Eocene
2657:S2CID
2524:PeerJ
2490:(PDF)
2412:(PDF)
2257:S2CID
2214:S2CID
2160:S2CID
2110:(PDF)
962:Spain
764:Egypt
148:(now
2649:PMID
2603:PMID
2552:PMID
2377:ISSN
2249:OCLC
2088:PMID
1916:The
1853:and
1767:Diet
1152:and
1046:and
1021:and
714:Age
659:and
644:and
628:and
612:and
439:and
392:and
367:and
341:and
333:and
107:and
35:PreꞒ
2641:doi
2629:620
2593:PMC
2583:doi
2542:PMC
2532:doi
2432:doi
2420:111
2369:doi
2328:doi
2241:doi
2206:doi
2152:doi
2078:PMC
2068:doi
877:USA
531:WW2
412:of
400:by
152:).
3330::
3308::
3293::
2655:.
2647:.
2639:.
2627:.
2601:.
2591:.
2577:.
2573:.
2550:.
2540:.
2528:12
2526:.
2522:.
2506:^
2498:21
2496:.
2492:.
2472:20
2470:.
2466:.
2444:^
2430:.
2418:.
2414:.
2389:^
2375:.
2367:.
2357:13
2355:.
2340:^
2326:.
2316:87
2314:.
2288:^
2280:20
2278:.
2255:.
2247:.
2235:.
2212:.
2204:.
2194:85
2192:.
2188:.
2172:^
2158:.
2150:.
2140:73
2138:.
2122:^
2112:.
2086:.
2076:.
2066:.
2056:17
2054:.
2050:.
1994:^
1984:.
1980:.
1966:^
1954:.
1847:,
1841:,
1804:.
1216:.
1180:.
1127:.
1107:.
1088:.
941:.
929:.
893:.
885:,
695:.
640:,
616:.
608:,
604:,
600:,
585:.
443:.
329:,
325:,
85:Pg
2694:e
2687:t
2680:v
2663:.
2643::
2635::
2609:.
2585::
2579:6
2558:.
2534::
2438:.
2434::
2426::
2383:.
2371::
2363::
2334:.
2330::
2322::
2263:.
2243::
2220:.
2208::
2200::
2166:.
2154::
2146::
2094:.
2070::
2062::
1986:9
1783:.
1011:.
470:(
462:(
287:?
248:†
235:†
111:.
90:N
80:K
75:J
70:T
65:P
60:C
55:D
50:S
45:O
40:Ꞓ
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