2952:
2881:
621:
1335:, stating that the sigmoid process of this bone was broad and plate-like with a transverse rim that was not well developed. A prominent keel emerges from the medial margin of the bulla and the involucrum is much steeper towards the front, both features setting the taxon aparat from most other basilosaurids. Gol'din and Zvonok further note that both the anterior process and the posterior (back) end of the bulla are rounded and blunt. Overall its size and shape resemble the ear bones of
2972:
2963:
1387:
155:
809:
1118:
4468:
2318:
135:
2329:
1252:
1227:, which is currently only known from the UK. However this species is only known from very fragmentary remains, making it unclear if it represents a distinct species or if it simply represents one of the other two species having lived in Britain. Fossils of a small species with less compact vertebrae have been recovered from the Lutetian to Bartonian of
3011:. This abrasion is seen across all types of teeth, but is least pronounced on the canines. The wear is described as uniform, leaving smooth margins and occurring perpendicular to the long axis of the tooth. In orcas, similar rapid abrasion is associated with the animals preying on sharks, causing the teeth to be worn down by their
780:, but failed to mention the material from Germany. It is thought by van Vliet that this was likely due to the fact that Kuhn had only identified the Helmstedt material as archaeocetes a year earlier, with Kellogg simply having been unaware of this research. Things went quiet for the following 70 years until 2001, when
2905:. This however came at the cost of making the animal much slower to accelerate and less maneuverable while swimming. The animal's maneuverability was also negatively affected by the anatomy of the transverse processes of the vertebrae. Since the transverse processes were much more elongated than those of
2929:
While acceleration and mobility were limited, there may have also been a variety of advantages in the denser bones, especially if coupled with an increase in lung volume. Slow swimmers, animals requiring large oxygen stores and shallow water inhabitants feeding close to the ocean floor are all listed
3835:
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
2359:
may have been a basal member of basilosauridae owning in part to its well developed innominate bones, which somewhat mirrors the position the taxon occupied in the work of Gol'din and Zvonok. This interpretation however is contrasted by a phylogenetic analysis published alongside the description of
1382:
preserving 12 thoratic vertebrae (though this could also be the ancestral condition) and six lumbars. The presence of a smaller number of longer vertebrae is similar to what is observed in modern whales and highlighted by Gol'din and Zvonok as differing greatly from
Basilosaurus, which despite its
3076:. During the time periods these formations were formed the sea was rather shallow, reaching a depth of only around 150 m (490 ft) at the time the Anneberg Formation was deposited and a depth between 50–100 m (160–330 ft) during the deposition of the Gehlberg Formation.
839:
consisted of a variety of bones including fragments of the skull, various vertebrae, ribs and parts of the innominate bone of the pelvic region. Additional specimens were referred to this species in 2001 by Uhen himself and in 2011 by Weems and colleagues, who expanded the range of
1405:
Though the individual vertebrae are elongated in a similar manner to those of basilosaurines, they differ in how much denser they are, which is achieved through the internal architecture of the bone. In basilosaurines, the vertebrae have been described as containing cylinders of
2364:. While this study also supported a paraphyletic Basilosauridae, pachycetines in this study appeared to be more derived rather than basal, appearing as a sister group to Neoceti with all other basilosaurids having split off before the appearance of this clade. Within the family
3598:
2994:
based on the patterns of tooth wear. Teeth from
Ukraine show a mixed pattern of latitudinal and cross-oriented scratches as well as deeper gouges that cover the surface of the teeth. Prior studies have linked coarse patterns like gouges to a carnivorous diet like in
3091:
are also known from shallow marine sediments. The Piney Point
Formation represents a shallow marine shelf that was deposited at a depth of around 60–120 ft (18–37 m). Molluscs from this formation suggest mild to warm temperatures and the terrestrial
924:
as well. However, as pointed out by
Gingerich and Zouhri in 2015, the availability of the holotype is not a prerequisite for the continued validity of a taxon. Instead the descriptions provided by Brandt, Fedorowskij and Kellogg were deemed sufficient for
804:
eventually came into question, with other authors highlighting that the material is very poorly preserved, casting doubt over the
Gritsenko's claims of it being distinct, and overall not diagnostic enough to be referred to any species or genus.
3444:
1427:
its name. However, Van Vliet and colleagues argue that the presence of cones is more widespread than presumed by
Gingerich and Zouhri and in fact common across basilosaurids. Regardless, the cortical bone within the vertebrae is
668:, which represented the first example of a fossil whale being found within the country. Later publications also make mention of another centra and a rib, which alongside the first centrum and some other whales bones were sent to
1354:, a size increase so extreme that even individual vertebrae already appear trapezoid in shape with a much larger posterior end. According to van Vliet the vertebrae are wider than they are tall, separating them from those of
3649:
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".
1167:
stages of the Eocene, while
Gingerich later stated that all well-dated material stems from the Bartonian. The possibility that some fossils were reworked into younger strata is mentioned by van Vliet, but deemed unlikely.
1414:
and other pachycetines meanwhile the cancellous bone is present in the form of two cones with their tips facing each other, meaning the central section of the vertebrae is surrounded by a much thicker and denser ring of
1279:
of the upper jaw is either double-rooted or had two fused roots. The premolars further back display four accessory cusps (sometimes also referred to as denticles) preceding and following the apex of tooth, whereas
1086:
was chosen by van
Beneden as a clear allusion to the thickened nature of the bones of the genus, with the name directly translating to "thick whale". Several other names have been proposed in the past, including
3043:
fauna was present in the
Americas, with the Piney Point Formation yielding the remains of 19 shark species and 9 species or rays. They therefore hypothesize that the animal was specialised in feeding on sharks.
1172:
is noted for having been larger than the American species, although Gingerich and colleagues do not list any other diagnostic features to separate the two species. The species was named after Otto Mikhaĭlovich
1486:
did not articulate at the sacrum, clearly setting it apart from the more basal protocetids. Furthermore, again like with other basilosaurids, the innominates retain features like the acetabular fossa and the
743:, another seven vertebrae were recovered by farmworkers from the region. The ten vertebrae, regarded as lumbar, saccral and caudal vertebrae by Fedorowskij, were described in 1912 and assigned to the species
2921:
would have had to rely on dorsoventral undulation, meaning it was limited to moving its body up and down. This limited mobility, combined with the overall increase in density, is at times compared to modern
558:
has a long and complex history, in part due to the often fragmentary and isolated nature of much of the material referred to this genus. The earliest recorded mention of material now assigned to
696:
based on some of the smaller fossils sent to him by Geinitz. Kuhn published a redescription of the two taxa in 1935, though he was less convinced of their distinctiveness. He rejected the name
2938:, though slow-moving, was a powerful swimmer regardless. Given that these adaptations are poorly suited for pursuing prey items, it has been hypothesized that pachycetines were ambush-hunter.
1362:. The lumbar vertebrae meanwhile are elongated in a fashion similar to those of basilosaurines, though not quite as extreme in their proportions. A major difference in the lumbar vertebrae of
2909:, there is less space between each process and those preceding and following it. This in turn means less space for muscle contraction, which leads to a decrease in lateral flexibility. While
2901:
and pachycetines in general is the great increase in bone density achieved via pachyostosis and osteosclerosis. This would have greatly increased the animal's weight and served as
1383:
elongated centra still retained a high vertebrae count. The surface of each vertebra is covered in small vascular openings, resulting in the bones having a pockmarked texture.
1219:
and is known from fewer remains, though said remains are more complete than any of the European finds. It was named in honour of Lauck "Buck" Ward who discovered the holotype.
464:
most likely represented a single taxon, which came to combine the generic name of the former and the species name of the latter. A species from the USA previously known as
456:, though the reasoning for the latter was regarded as unjustified and was not accepted in much of the subsequent literature. By the 2020s it was recognized that the German
436:
has a long and complex history, with its earliest recorded remains having been found in what is now Ukraine in the late 19th century. The fossils were initially dubbed
957:
While this seemingly settled the matter of validity, the nomenclature was still subject to change. In a paper published in 2020, van Vliet noted that the vertebrae of
828:
3735:"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"
720:). Kuhn furthermore described several additional fossils and correctly hypothesized that they were Eocene in age, while Van Beneden had assumed them to be Oligocene.
870:, some of which were featured a map by Evgenij Zvonok that figured archaeocete localities across Ukraine and Russia. Material discovered include teeth and ribs from
4533:
1545:
has a long history of research, its precise relationship with other basilosaurids has long remained unknown. In 2013 Gol'din and Zvonok found basilosaurids to be a
477:
Given the limited material known of this genus, its biology is only poorly understood. The dense skeletal structure is commonly compared to that seen in modern-day
1029:
on the other hand, described by van Beneden in 1883, did not receive the same treatment. Instead it was speculated that it could represent a whale distinct from
978:
3123:, formed at depths between 50–500 m (160–1,640 ft) and most common between 200–300 m (660–980 ft), which has been interpreted to mean that
3247:"Skull and partial skeleton of a new pachycetine genus (Cetacea, Basilosauridae) from the Aridal Formation, Bartonian middle Eocene, of southwestern Morocco"
1370:
these processes are almost as long as the centra of the corresponding vertebrae, which is not the case in basilosaurines. Both the pedicles (surrounding the
503:, a suggestion largely congruent with the German and American sediments it has been recovered from. Some fossils from the Ukraine do however suggest that
1529:. In the former the lumbar vertebrae, including the epiphysis, measure up to 266 mm (10.5 in) compared to only 171 mm (6.7 in) in
1436:, are pachyostotic as well. The ribs also contribute to the increased density of the skeleton, being pachyostotic themselves and having replaced the
4607:
4520:
1432:, meaning it is even denser than is typical for whales, with much fewer porosities. Like the centra, the pedicles of the vertebrae, which flank the
374:, however, fossil finds have also been recovered from sediments of less certain age that could suggest that it may have also lived during the Late
3131:
may have entered deeper waters while migrating, which would have been made possible by their powerful swimming musculature and its good ballast.
3498:
Uhen, M. D. (1999). "New species of protocetid archaeocete whale, Eocetus wardii (Mammalia: Cetacea) from the middle Eocene of North Carolina".
4592:
489:) and it has been suggested that it was a powerful if comparably inflexible swimmer that swam by moving its entire body up and down similar to
3399:
Gol'din, P.; Zvonok, E. (2013). "Basilotritus uheni, a New Cetacean (Cetacea, Basilosauridae) from the Late Middle Eocene of Eastern Europe".
3907:
4597:
1468:
is how the core of cancellous bone is situated in the middle of the ribs, while in other basilosaurids the core is slightly off-center.
531:. Some supporting evidence for this hypothesis may be found in the high number of sharks and rays found within the same sediments as
4602:
2934:. Gol'din and Zvonok come to a similar conclusion while further highlighting how the large vertebral processes would indicate that
2880:
3060:. During the Paleogene the ocean retreated and expanded repeatedly, creating a mix of terrestrial and marine conditions. Multiple
448:, a name that would eventually fade into obscurity as time passed on. The Ukrainian material would eventually come to be named
1211:
was found, has been suggested to be either late Lutetian or early Bartonian in age and a similar age has been assumed for the
3007:. Gol'din and colleagues especially highlight the strongly abraded tips of the teeth of the Nagornoye teeth they assigned to
866:
The 2000s and 2010s saw an increase in newly described basilosaurid material from Ukraine that may or may not have come from
613:
would be found. Paulson seemingly didn't object to the change, though later authors would occasionally refer to the taxon as
2951:
1331:. Gol'din and Zvonok describe the tympanic bulla, an element created by the fusion of the ear bones, in their diagnosis for
1040:, though one less drastic than the work of van Vliet. Following a summary of the genus' complex history, they conclude that
570:
1561:, was recovered as a basal off-shoot of the latter alongside MUSM 1443 from Peru. Their follow-up publication describing
1499:. Additional material has been described for the European species and Gol'din and Zvonok include in their diagnosis for
3056:
originated, was once located at the southern edge of the Paleogene North Sea where rivers from the continent filled an
2917:, undulating the entire body to move, it was much more restricted in its movement. Gingerich and colleagues argue that
601:, who published the description in his 1873 monograph on European fossil whales. Paulson initially dubbed the material
2984:
may have included sharks like the sand tiger shark (top), epaulette sharks (bottom left) and catsharks (bottom right).
1495:
and other basilosaurids as well. For instance, the obturator foramen, an opening in the innominate, is much larger in
3340:
3177:"Eocene cetaceans from the Helmstedt region, Germany, with some remarks on Platyosphys, Basilotritus and Pachycetus"
3015:. It is noted by the team that the same sediments yielding these teeth also preserve 35 species of sharks including
154:
2930:
as profiting from such anatomy, with Gingerich and colleagues interpreting pachycetines as slow inhabitants of the
871:
566:
3900:
669:
3610:
1378:
are also elongated. The elongation of the vertebrae is accompanied by a notable reduction in their number, with
3552:"An occurrence of the protocetid whale "Eocetus" wardii in the Middle Eocene Piney Point Formation of Virginia"
594:
3784:
Antar, M.S.; Glaohar, A.S.; El-Desouky, H.; Seiffert, E.R.; El-Sayed, S.; Claxton, A.G.; Sallam, H.M. (2023).
740:
724:
640:. The nearly complete skull discovered in 1872, which was damaged during collection, was described in 1876 by
3786:"A diminutive new basilosaurid whale reveals the trajectory of the cetacean life histories during the Eocene"
3700:"An archaeocete vertebra re-examined: indications for a small-sized species of Pachycetus from Spain, Europe"
1478:, the element formed by the fusion of the hip bones, with the best material of this bone being preserved for
527:, arguing that the high levels of abbrasion seen on some of the teeth was caused by contact with their preys
4587:
3445:""New fauna of archaeocete whales (Mammalia, Cetacea) from the Bartonian middle Eocene of southern Morocco""
1204:
781:
598:
3064:
within the region around Helmstedt trace their origins to this period, including the Lutetian to Bartonian
1461:
620:
4488:
3733:
van Vliet, H.J.; Bosselaers, M.E.J.; Munsterman, D.K.; Dijkshoorn, M.L.; de Groen, J.J.; Post, K. (2024).
3310:
2650:
1232:
3065:
1009:, but refrains from committing to this given how fragmentary the Helmstedt material is. In light of this
770:, primarily basing his work on the material previously described by Fedorowskij. He furthermore referred
628:
was coined by Pierre-Joseph van Beneden, though his work was largely overlooked in the following century.
4559:
3361:
Gritsenko, V. (2001). "New species Platiosphys einori (Archaeoceti) from Oligocenic deposits of Kyiv".
3061:
1212:
728:
4471:
3893:
3847:
3659:
3563:
3507:
3459:
3408:
3258:
3127:
at least occasionally left the shallows to venture into the open ocean. One hypothesis suggests that
1457:
3699:
2971:
2962:
2486:
653:
641:
820:
815:
was described based on assorted cranial and postcranial remains including parts of the innominate.
507:
at least occasionally entered deeper waters, which has been interpreted as potential evidence for
3871:
3622:
3579:
3523:
3069:
1240:
748:
657:
300:
149:
3035:
1386:
410:, but the cortical bone was furthermore much denser than in other basilosaurids. Two species of
4564:
875:
118:
Though the best-dated remains are Bartonian in age, some could suggest its presence during the
4546:
4538:
4031:
3863:
3817:
3766:
3675:
3614:
3315:
Mélanges biologiques tirés du Bulletin de l'Académie impériale des sciences de St. Pétersbourg
3286:
2791:
1488:
759:
736:
706:
4551:
3599:"A new protocetid whale (Cetacea, Archaeoceti) from the late middle Eocene of South Carolina"
3080:
would have shared this environment with a second, much smaller basilosaurid originally named
1339:
and Gol'din and Zvonok describe it as displaying a mix of features seen in basilosaurids and
3855:
3838:
3807:
3797:
3756:
3746:
3667:
3606:
3571:
3515:
3467:
3416:
3276:
3266:
1475:
1351:
1327:, although little of the cranium is known, with the most substantial material being that of
578:
508:
142:
1419:. This condition of having a thickened layer of lamellar cortical bone is what is known as
1117:
993:, the former takes priority over the latter, which means that the type species would still
808:
4068:
3040:
1407:
516:
407:
3851:
3812:
3785:
3761:
3734:
3663:
3567:
3511:
3463:
3412:
3262:
4384:
4285:
4088:
3281:
3246:
3012:
3004:
2193:
1437:
1429:
1296:. The molar, unlike the premolar, only has three accessory cusps. Overall the teeth of
1002:
909:
832:
610:
528:
395:
347:
229:
1533:, width some individual vertebrae reaching a length of over 280 mm (11 in).
1491:
which have been lost by today's whales. There are however notable differences between
1482:. Like in other members of its family and derived ceteaceans, the innominate bones of
1109:("broad loin" in reference to the wide transverse processes of the lumbar vertebrae).
852:
had fallen out of favour, with Geisler, Sanders and Luo (2005) suggesting it be named
4581:
4150:
4139:
4111:
4040:
4020:
3997:
3986:
3875:
3583:
3551:
3471:
3109:
2345:, with which it forms the subfamily Pachycetinae. Top shows the holotype specimen of
1730:
1416:
1200:
582:
403:
343:
74:
3626:
3527:
882:). These new finds eventually lead to another attempt at clarifying the taxonomy of
4418:
4393:
4317:
4294:
4193:
4179:
4170:
4125:
4118:
4079:
4047:
3101:
3023:
3017:
2931:
2609:
2317:
2066:
2042:
1788:
1696:
1546:
1504:
1433:
1420:
1371:
1340:
1305:
1272:
1093:
901:
888:
800:, he failed to provide an adequate diagnosis for this new species. The validity of
793:
665:
648:. Though the holotype has been lost since then, various other elements referred to
491:
399:
339:
336:
282:
242:
134:
3175:
van Vliet, H.; Bosselaers, M.; Vahldiek, B.W.; Paymans, T.; Verheijen, I. (2020).
1147:
is currently only known from European deposits, with fossils having been found in
933:
was questioned. The same publication went on to not only reaffirm the validity of
4511:
3271:
1553:(which includes today's whales) and what they refer to as "crown basilosaurids".
680:. Taking note of the robust nature of the bones and their similarity to those of
609:, reasoning that the name was better suited in case material outside of the then
4448:
4400:
4366:
4345:
4310:
4303:
4249:
4235:
4228:
4159:
4061:
4006:
3963:
3953:
3916:
2726:
2576:
2445:
2328:
2009:
1961:
1843:
1672:
1375:
1289:
1105:
1099:
633:
206:
49:
3885:
3859:
3802:
3671:
656:
published on a vertebral centrum recovered from the phosphate beds (likely the
4427:
4359:
4352:
4331:
4263:
4256:
4242:
4207:
4200:
4186:
4132:
4104:
4054:
4013:
3941:
3618:
3519:
3120:
3029:
2559:
1992:
1812:
1747:
1713:
1648:
1624:
1164:
1125:
are known from across Europe and the eastern United States, while the related
1046:
716:
470:
379:
123:
94:
59:
3679:
3176:
723:
Additional archaeocete material from eastern Europe was described in 1894 by
4441:
4373:
4270:
4221:
4214:
4097:
3979:
3972:
3113:
2923:
2528:
1937:
1607:
1583:
1445:
1251:
1236:
1160:
797:
776:
685:
681:
661:
478:
387:
371:
166:
99:
43:
37:
20:
3867:
3821:
3770:
3290:
1036:
Two years later Gingerich, Amane and Zhouri published another revision to
4505:
4338:
3935:
3363:
Visnyk Heolohila Kyivskyi Natsionalyi Universytet Imeni Tarasa Shevchenka
3097:
2469:
2170:
1441:
1301:
1276:
845:
586:
563:
524:
500:
482:
375:
351:
186:
119:
89:
84:
69:
64:
54:
3751:
2384:
from the Bartonian of South America was also recovered as a relative of
4525:
4407:
4324:
3057:
2902:
2834:
2633:
2407:
2208:
2083:
1771:
1550:
1508:
1156:
1148:
942:
789:
684:, Van Beneden described them as a new genus of what he thought to be a
677:
590:
486:
363:
359:
216:
104:
79:
3420:
886:
by Gol’din and Zvonok in 2013. In their work they name the new genus
444:. Simultaneously, fossils from the north of Germany were described as
3947:
3929:
3575:
2355:
Although no full analysis was performed, Van Vliet hypothesized that
1152:
1044:
was sufficiently distinct to warrant its own genus, which they named
762:
recognized the distinct nature of the taxon and coined the new genus
673:
355:
196:
176:
4482:
1366:
versus those of basilosaurines lies in the transverse processes. In
402:
skeleton. This means the bones not only featured thickened rings of
3698:
van Vliet, H.J.; Bosselaers, M.; Paijmans, T.; Calzada, S. (2023).
632:
Around the same time another archaeocete was discovered within the
3105:
3093:
3000:
2879:
1385:
1250:
1228:
1116:
807:
619:
520:
3116:, which indicate that the climate was tropical with dry winters.
874:(initially referred to Basilosauridae indet.) and vertebrae from
562:
dates back to 1871, when fossils of this whale where reported by
474:
these animals would become the basis of the family Pachycetinae.
3550:
Weems, R.E.; Edwards, L.E.; Osborne, J.E.; Alford, A.A. (2011).
856:
until further study could determine its generic identification.
574:
426:
from the United States. A third species might be represented by
4486:
3889:
1456:
indicate that they were similar. The densest ribs are the late
2913:
would still share a swimming style broadly similar to that of
752:
735:, and even more complete material was discovered in 1909 near
700:
and instead suggested that the fossils may have been those of
652:
have been described in subsequent years. In Germany meanwhile
391:
1410:
that fill up the anterior and posterior of each vertebra. In
965:
were identical to those from Germany originally described as
755:
are not known and the remains are currently considered lost.
3003:
while the scratches mirror those seen in modern piscivorous
1159:. Van Vliet and colleagues assign the German remains to the
908:
was named on the basis of various postcranial fossils and a
792:
south of Kyiv. Although Gritsenko took note of the advanced
3611:
10.1206/0003-0082(2005)480[0001:ANPWCA]2.0.CO;2
3311:"*Über bisher in Russland gefundene Reste von Zeuglodonten"
637:
577:. The material consisted of three complete and one partial
912:
while a variety of less complete remains were referred to
739:. In addition to the initial three vertebra discovered by
2888:
is at times compared to modern sirenians like the dugong.
848:. By that point however the assignment of the fossils to
511:. Tooth wear has been used as evidence for the fact that
3084:, though it is now believed to have been a dorudontine.
1565:
remains from Nagornoye recovered results much the same.
1033:, a sentiment later echoed by Gingerich and colleagues.
3087:
Similar to the German remains, the American fossils of
796:
of the material, likening it to what is seen in modern
3119:
Ukrainian material is known from sediments containing
1517:
is described as having been larger than the related
4495:
4417:
4383:
4293:
4284:
4169:
4149:
4087:
4078:
4030:
3996:
3962:
920:, ultimately keeping the species but placing it in
468:was also introduced to the genus and together with
1223:A potential third species might be represented by
3597:Geisler, J.H.; Sanders, A.E.; Luo, Z.-X. (2005).
3072:that is most likely to have yielded the bones of
1350:show a rapid increase in size as one follows the
1058:. The team also agrees with van Vielt in sinking
366:). The best known remains generally suggest that
3052:Helmstedt, where much of the German material of
2376:being successively diverging taxa leading up to
1203:, specifically North Carolina and Virginia. The
985:had priority. Simultaneously, since the species
829:United States National Museum of Natural History
3245:Gingerich, P.D.; Amane, A.; Zouhri, S. (2022).
1448:. Though the articulation is best described in
1316:have been described by Gol'din and colleagues.
747:before being introduced into the collection of
335:(meaning "thick whale") is an extinct genus of
2990:Some inferences have been made on the diet of
499:preferred shallow waters and fed close to the
3901:
1440:articulation between ribs and vertebrae with
1021:were all carried over as distinct species of
827:based on material from the collection of the
692:. He furthermore described a second species,
593:. These remains were eventually described by
8:
1284:only has three. The two also differ in that
727:, who discusses a vertebra found within the
1259:with the "pockmarks" highlighted by arrows.
973:was named in 1883, a whole 43 years before
4483:
4290:
4084:
3908:
3894:
3886:
1464:. One feature highlighted for the ribs of
916:. The two also wrote on the assignment of
133:
27:
3811:
3801:
3760:
3750:
3280:
3270:
3728:
3726:
3724:
3722:
3720:
3693:
3691:
3689:
3240:
3238:
3236:
3234:
3232:
3230:
3228:
3226:
3224:
3222:
3220:
3218:
3216:
3214:
3170:
3168:
3166:
3164:
2388:and the basalmost member of this clade.
941:based on fossil material collected from
3644:
3642:
3640:
3638:
3636:
3545:
3543:
3541:
3539:
3537:
3212:
3210:
3208:
3206:
3204:
3202:
3200:
3198:
3196:
3194:
3162:
3160:
3158:
3156:
3154:
3152:
3150:
3148:
3146:
3144:
3140:
1300:resemble those of other basilosaurids.
831:, which had been discovered in 1977 in
758:In his 1936 monograph on archaeocetes,
3438:
3436:
3434:
3432:
3430:
3394:
3392:
3390:
3388:
3386:
3384:
3382:
3380:
3378:
3376:
1549:clade giving rise to both the derived
1474:is among the basilosaurids with known
1323:had a proportionally larger head than
3493:
3491:
3489:
3487:
3485:
3483:
3481:
3334:
3332:
3330:
3328:
3304:
3302:
3300:
751:, though their whereabouts following
571:Third Russian Congress of Naturalists
495:. It has further been suggested that
307:
7:
3443:Gingerich, P.D.; Zouhri, S. (2015).
1319:Gingerich and colleagues argue that
1050:, and furthermore commit to lumping
929:to remain valid while the status of
440:, only to be immediately renamed to
2766:
2742:
2718:
2711:
2625:
2601:
2551:
2544:
2520:
2461:
2437:
2430:
2423:
2399:
2391:
2368:was found to be paraphyletic, with
2186:
2162:
2058:
2034:
1984:
1977:
1953:
1929:
1866:
1859:
1835:
1828:
1804:
1763:
1688:
1664:
1640:
1599:
1575:
1568:
1191:and later transferred to the genus
394:and is characterized by its highly
3348:Carnegie Institution of Washington
2897:One of the most notable traits of
937:, but also to name a new species:
342:from Middle Eocene of the eastern
14:
3452:Journal of African Earth Sciences
1187:Originally named as a species of
896:, reasoning that the loss of the
605:, but Brandt changed the name to
4467:
4466:
3472:10.1016/j.jafrearsci.2015.08.006
3068:and the Bartonian to Priabonian
2970:
2961:
2950:
2327:
2316:
153:
4608:Eocene mammals of North America
2349:, bottom a live reconstruction.
1503:that the section formed by the
1267:are more robust than those of
731:that he referred to the genus
1:
4593:Fossil taxa described in 1883
3341:"A Review of the Archaeoceti"
1525:being noticeably larger than
1239:off the Dutch-Belgian coast (
1091:("third king" in allusion to
900:should constitute it being a
3272:10.1371/journal.pone.0276110
1507:extends forward whereas the
1452:, the rough surface seen in
1308:that could have belonged to
997:. Van Vliet recognizes that
788:from material discovered in
567:Afanasii Semenovich Rogovich
515:might have fed primarily on
4598:Prehistoric cetacean genera
4624:
3860:10.1038/s41586-023-06381-1
3803:10.1038/s42003-023-04986-w
3672:10.1016/j.crpv.2013.11.002
3024:Hemiscyllium bruxelliensis
2884:Due to its dense skeleton
1346:The thoratic vertebrae of
1271:and possessing crenulated
1199:is exclusively known from
18:
4462:
3924:
3603:American Museum Novitates
3520:10.1017/s002233600002802x
2832:
2788:
2771:
2764:
2747:
2740:
2723:
2716:
2709:
2647:
2630:
2623:
2606:
2599:
2573:
2556:
2549:
2542:
2525:
2518:
2483:
2466:
2459:
2442:
2435:
2428:
2421:
2404:
2397:
2206:
2191:
2184:
2167:
2160:
2080:
2063:
2056:
2039:
2032:
2006:
1989:
1982:
1975:
1958:
1951:
1934:
1927:
1901:
1884:
1871:
1864:
1857:
1840:
1833:
1826:
1809:
1802:
1785:
1768:
1761:
1744:
1727:
1710:
1693:
1686:
1669:
1662:
1645:
1638:
1621:
1604:
1597:
1580:
1573:
670:Pierre-Joseph Van Beneden
595:Otto Mikhaĭlovich Paulson
406:surrounding the internal
306:
299:
272:
267:
150:Scientific classification
148:
141:
132:
30:
4603:Eocene mammals of Europe
2893:Locomotion and lifestyle
2337:The closest relative of
1288:lacks a swelling on the
1207:, where the holotype of
1070:now deemed a synonym of
741:Alexandre S. Fedorowskij
725:Leonid Ivanovich Lutugin
452:before being changed to
386:is primarily known from
19:Not to be confused with
3556:Journal of Paleontology
3500:Journal of Paleontology
3401:Journal of Paleontology
1129:inhabited North Africa.
835:. The type specimen of
599:Johann Friedrich Brandt
313:Goldin and Zvonok 2013
16:Extinct genus of whales
3652:Comptes Rendus Palevol
3039:. A similarly diverse
2889:
2651:Pontogeneus peruvianus
1402:
1260:
1255:A lumbar vertebrae of
1215:. It was smaller than
1205:Castle Hayne Formation
1130:
816:
782:Volodymyr P. Gritsenko
688:whale, which he named
629:
581:, discovered near the
4560:Paleobiology Database
3309:Brandt, J.F. (1873).
3096:supported members of
3062:geological formations
3030:Carcharias acutissimu
2883:
1557:, at the time called
1462:vertebrochondral ribs
1389:
1312:and were referred to
1254:
1213:Piney Point Formation
1120:
927:Platyosphys paulsonii
898:Platyosphys paulsonii
892:as a replacement for
811:
623:
3339:Kellogg, R. (1936).
2751:Pachycetus paulsonii
2370:Pachycetus paulsonii
1523:Pachycetus paulsonii
1458:vertebrosternal ribs
1390:Innominate bones of
1233:Folgarolas Formation
1225:"Zeuglodon" wanklyni
1217:Pachycetus paulsonii
1170:Pachycetus paulsonii
1143:The type species of
1136:Pachycetus paulsonii
1072:Pachycetus paulsonii
995:Pachycetus paulsonii
428:"Zeuglodon" wanklyni
417:Pachycetus paulsonii
3852:2023Natur.620..824B
3752:10.7717/peerj.16541
3664:2014CRPal..13..267G
3568:2011JPal...85..271W
3512:1999JPal...73..512U
3464:2015JAfES.111..273G
3413:2013JPal...87..254G
3263:2022PLoSO..1776110G
3066:Annenberg Formation
2487:Tutcetus rayanensis
1905:Basilotritus wardii
1511:is highly reduced.
1019:Basilotritus wardii
977:, the rules of the
768:Zeuglodon paulsonii
690:Pachycetus robustus
654:Hanns Bruno Geinitz
642:Harry Govier Seeley
607:Zeuglodon paulsonii
544:Early discoveries,
442:Zeuglodon paulsonii
3181:Cainozoic Research
3082:Pachycetus humilis
3070:Gehlberg Formation
2890:
1888:Basilotritus uheni
1501:Basilotritus uheni
1403:
1261:
1241:Maldegem Formation
1131:
1027:Pachycetus humilis
1015:Basilotritus uheni
1011:Platyosphys aithai
953:and recent studies
939:Platyosphys aithai
931:Basilotritus uheni
906:Basilotritus uheni
817:
786:Platyosphys einori
772:Zeuglodon wanklyni
749:Kharkiv University
745:Zeuglodon rossicus
729:Donets River basin
694:Pachycetus humilis
658:Gehlberg Formation
646:Zeuglodon wanklyni
630:
615:Zeuglodon rossicum
603:Zeuglodon rossicus
539:History and naming
509:migratory behavior
460:and the Ukrainian
438:Zeuglodon rossicus
4575:
4574:
4547:Open Tree of Life
4489:Taxon identifiers
4480:
4479:
4458:
4457:
4280:
4279:
4032:Remingtonocetidae
3846:(7975): 824–829.
3421:10.1666/12-080R.1
3089:Pachycetus wardii
3036:Clerolamna umovae
2872:
2871:
2863:
2862:
2854:
2853:
2845:
2844:
2821:
2820:
2812:
2811:
2803:
2802:
2792:Antaecetus aithai
2775:Pachycetus wardii
2698:
2697:
2689:
2688:
2680:
2679:
2671:
2670:
2662:
2661:
2588:
2587:
2507:
2506:
2498:
2497:
2374:Pachycetus wardii
2309:
2308:
2300:
2299:
2291:
2290:
2282:
2281:
2273:
2272:
2264:
2263:
2255:
2254:
2246:
2245:
2237:
2236:
2228:
2227:
2219:
2218:
2149:
2148:
2140:
2139:
2131:
2130:
2122:
2121:
2113:
2112:
2104:
2103:
2095:
2094:
2021:
2020:
1916:
1915:
1527:Pachycetus wardii
1489:obturator foramen
1480:Pachycetus wardii
1380:Pachycetus wardii
1329:Pachycetus wardii
1292:which is seen in
1180:Pachycetus wardii
1042:Pachycetus aithai
813:Pachycetus wardii
760:Remington Kellogg
717:Saghacetus osiris
707:Basilosaurus isis
597:at the behest of
423:Pachycetus wardii
370:lived during the
328:
327:
322:
314:
293:
280:
263:
262:van Beneden, 1883
127:
4615:
4568:
4567:
4555:
4554:
4542:
4541:
4529:
4528:
4516:
4515:
4514:
4484:
4470:
4469:
4291:
4085:
3919:genera by family
3910:
3903:
3896:
3887:
3880:
3879:
3832:
3826:
3825:
3815:
3805:
3781:
3775:
3774:
3764:
3754:
3730:
3715:
3714:
3704:
3695:
3684:
3683:
3646:
3631:
3630:
3594:
3588:
3587:
3576:10.1666/10-083.1
3547:
3532:
3531:
3495:
3476:
3475:
3449:
3440:
3425:
3424:
3396:
3371:
3370:
3358:
3352:
3351:
3345:
3336:
3323:
3322:
3306:
3295:
3294:
3284:
3274:
3257:(10): e0276110.
3242:
3189:
3188:
3172:
3048:Paleoenvironment
3018:Scyliorhinus sp.
2974:
2965:
2954:
2767:
2743:
2719:
2712:
2626:
2602:
2552:
2545:
2521:
2462:
2438:
2431:
2424:
2400:
2392:
2331:
2320:
2187:
2163:
2059:
2035:
1985:
1978:
1954:
1930:
1867:
1860:
1836:
1829:
1805:
1764:
1689:
1665:
1641:
1600:
1576:
1569:
1476:innominate bones
1352:vertebral column
918:"Eocetus" wardii
914:Basilotritus sp.
854:"Eocetus" wardii
712:Zeuglodon osiris
579:vertebral centra
466:"Eocetus" wardii
420:from Europe and
414:are recognized:
320:
312:
291:
278:
261:
254:
241:
228:
158:
157:
143:Life restoration
137:
117:
116:
46:
36:Temporal range:
28:
4623:
4622:
4618:
4617:
4616:
4614:
4613:
4612:
4578:
4577:
4576:
4571:
4563:
4558:
4550:
4545:
4537:
4532:
4524:
4519:
4510:
4509:
4504:
4491:
4481:
4476:
4454:
4413:
4379:
4276:
4165:
4145:
4074:
4069:Remingtonocetus
4026:
3992:
3958:
3920:
3914:
3884:
3883:
3834:
3833:
3829:
3783:
3782:
3778:
3732:
3731:
3718:
3702:
3697:
3696:
3687:
3648:
3647:
3634:
3596:
3595:
3591:
3549:
3548:
3535:
3497:
3496:
3479:
3447:
3442:
3441:
3428:
3398:
3397:
3374:
3360:
3359:
3355:
3343:
3338:
3337:
3326:
3308:
3307:
3298:
3244:
3243:
3192:
3174:
3173:
3142:
3137:
3050:
3041:chondrichthyian
2988:
2987:
2986:
2985:
2977:
2976:
2975:
2967:
2966:
2957:
2956:
2955:
2944:
2895:
2878:
2873:
2864:
2855:
2846:
2822:
2813:
2804:
2699:
2690:
2681:
2672:
2663:
2589:
2508:
2499:
2353:
2352:
2351:
2350:
2334:
2333:
2332:
2323:
2322:
2321:
2310:
2301:
2292:
2283:
2274:
2265:
2256:
2247:
2238:
2229:
2220:
2150:
2141:
2132:
2123:
2114:
2105:
2096:
2022:
1917:
1539:
1408:cancellous bone
1249:
1115:
1080:
955:
864:
553:
541:
517:chondrichthyans
408:cancellous bone
260:
252:
239:
226:
152:
128:
115:
114:
113:
112:
107:
102:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
42:41.2–37.8
41:
40:
34:
24:
17:
12:
11:
5:
4621:
4619:
4611:
4610:
4605:
4600:
4595:
4590:
4588:Basilosauridae
4580:
4579:
4573:
4572:
4570:
4569:
4556:
4543:
4530:
4517:
4501:
4499:
4493:
4492:
4487:
4478:
4477:
4475:
4474:
4463:
4460:
4459:
4456:
4455:
4453:
4452:
4445:
4438:
4431:
4423:
4421:
4415:
4414:
4412:
4411:
4404:
4397:
4389:
4387:
4385:Basilosaurinae
4381:
4380:
4378:
4377:
4370:
4363:
4356:
4349:
4342:
4335:
4328:
4321:
4314:
4307:
4299:
4297:
4288:
4286:Basilosauridae
4282:
4281:
4278:
4277:
4275:
4274:
4267:
4260:
4253:
4246:
4239:
4232:
4225:
4218:
4211:
4204:
4197:
4190:
4183:
4175:
4173:
4167:
4166:
4164:
4163:
4155:
4153:
4147:
4146:
4144:
4143:
4136:
4129:
4122:
4115:
4108:
4101:
4093:
4091:
4089:Georgiacetinae
4082:
4076:
4075:
4073:
4072:
4065:
4058:
4051:
4044:
4036:
4034:
4028:
4027:
4025:
4024:
4017:
4010:
4002:
4000:
3994:
3993:
3991:
3990:
3983:
3976:
3968:
3966:
3960:
3959:
3957:
3956:
3950:
3944:
3938:
3932:
3925:
3922:
3921:
3915:
3913:
3912:
3905:
3898:
3890:
3882:
3881:
3827:
3776:
3716:
3685:
3658:(4): 267–276.
3632:
3605:(3480): 1–68.
3589:
3562:(2): 271–278.
3533:
3506:(3): 512–528.
3477:
3426:
3407:(2): 254–268.
3372:
3353:
3324:
3296:
3190:
3139:
3138:
3136:
3133:
3049:
3046:
3013:placoid scales
3005:toothed whales
2979:
2978:
2969:
2968:
2960:
2959:
2958:
2949:
2948:
2947:
2946:
2945:
2943:
2940:
2894:
2891:
2877:
2874:
2870:
2869:
2866:
2865:
2861:
2860:
2857:
2856:
2852:
2851:
2848:
2847:
2843:
2842:
2839:
2838:
2831:
2828:
2827:
2824:
2823:
2819:
2818:
2815:
2814:
2810:
2809:
2806:
2805:
2801:
2800:
2797:
2796:
2787:
2784:
2783:
2780:
2779:
2770:
2765:
2763:
2760:
2759:
2756:
2755:
2746:
2741:
2739:
2736:
2735:
2732:
2731:
2722:
2717:
2715:
2710:
2708:
2705:
2704:
2701:
2700:
2696:
2695:
2692:
2691:
2687:
2686:
2683:
2682:
2678:
2677:
2674:
2673:
2669:
2668:
2665:
2664:
2660:
2659:
2656:
2655:
2646:
2643:
2642:
2639:
2638:
2629:
2624:
2622:
2619:
2618:
2615:
2614:
2605:
2600:
2598:
2595:
2594:
2591:
2590:
2586:
2585:
2582:
2581:
2572:
2569:
2568:
2565:
2564:
2555:
2550:
2548:
2543:
2541:
2538:
2537:
2534:
2533:
2524:
2519:
2517:
2514:
2513:
2510:
2509:
2505:
2504:
2501:
2500:
2496:
2495:
2492:
2491:
2482:
2479:
2478:
2475:
2474:
2465:
2460:
2458:
2455:
2454:
2451:
2450:
2441:
2436:
2434:
2429:
2427:
2422:
2420:
2417:
2416:
2413:
2412:
2403:
2398:
2396:
2390:
2336:
2335:
2326:
2325:
2324:
2315:
2314:
2313:
2312:
2311:
2307:
2306:
2303:
2302:
2298:
2297:
2294:
2293:
2289:
2288:
2285:
2284:
2280:
2279:
2276:
2275:
2271:
2270:
2267:
2266:
2262:
2261:
2258:
2257:
2253:
2252:
2249:
2248:
2244:
2243:
2240:
2239:
2235:
2234:
2231:
2230:
2226:
2225:
2222:
2221:
2217:
2216:
2213:
2212:
2205:
2202:
2201:
2198:
2197:
2194:Kekenodontidae
2190:
2185:
2183:
2180:
2179:
2176:
2175:
2166:
2161:
2159:
2156:
2155:
2152:
2151:
2147:
2146:
2143:
2142:
2138:
2137:
2134:
2133:
2129:
2128:
2125:
2124:
2120:
2119:
2116:
2115:
2111:
2110:
2107:
2106:
2102:
2101:
2098:
2097:
2093:
2092:
2089:
2088:
2079:
2076:
2075:
2072:
2071:
2062:
2057:
2055:
2052:
2051:
2048:
2047:
2038:
2033:
2031:
2028:
2027:
2024:
2023:
2019:
2018:
2015:
2014:
2005:
2002:
2001:
1998:
1997:
1988:
1983:
1981:
1976:
1974:
1971:
1970:
1967:
1966:
1957:
1952:
1950:
1947:
1946:
1943:
1942:
1933:
1928:
1926:
1923:
1922:
1919:
1918:
1914:
1913:
1910:
1909:
1900:
1897:
1896:
1893:
1892:
1883:
1880:
1879:
1876:
1875:
1870:
1865:
1863:
1858:
1856:
1853:
1852:
1849:
1848:
1839:
1834:
1832:
1827:
1825:
1822:
1821:
1818:
1817:
1808:
1803:
1801:
1798:
1797:
1794:
1793:
1784:
1781:
1780:
1777:
1776:
1767:
1762:
1760:
1757:
1756:
1753:
1752:
1743:
1740:
1739:
1736:
1735:
1726:
1723:
1722:
1719:
1718:
1709:
1706:
1705:
1702:
1701:
1692:
1687:
1685:
1682:
1681:
1678:
1677:
1668:
1663:
1661:
1658:
1657:
1654:
1653:
1644:
1639:
1637:
1634:
1633:
1630:
1629:
1620:
1617:
1616:
1613:
1612:
1603:
1598:
1596:
1593:
1592:
1589:
1588:
1579:
1574:
1572:
1567:
1538:
1535:
1460:and the early
1430:osteosclerotic
1257:Eocetus wardii
1248:
1245:
1221:
1220:
1184:
1183:
1175:
1174:
1140:
1139:
1114:
1111:
1079:
1076:
1003:junior synonym
981:dictated that
954:
947:
910:tympanic bulla
863:
858:
833:North Carolina
825:Eocetus wardii
702:Zeuglodon isis
611:Russian Empire
552:
542:
540:
537:
529:placoid scales
396:osteosclerotic
348:North Carolina
326:
325:
324:
323:
315:
304:
303:
297:
296:
295:
294:
286:
279:(Brandt, 1873)
270:
269:
265:
264:
250:
246:
245:
237:
233:
232:
230:Basilosauridae
224:
220:
219:
214:
210:
209:
204:
200:
199:
194:
190:
189:
184:
180:
179:
174:
170:
169:
164:
160:
159:
146:
145:
139:
138:
130:
129:
110:
109:
108:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
47:
35:
15:
13:
10:
9:
6:
4:
3:
2:
4620:
4609:
4606:
4604:
4601:
4599:
4596:
4594:
4591:
4589:
4586:
4585:
4583:
4566:
4561:
4557:
4553:
4548:
4544:
4540:
4535:
4531:
4527:
4522:
4518:
4513:
4507:
4503:
4502:
4500:
4498:
4494:
4490:
4485:
4473:
4465:
4464:
4461:
4451:
4450:
4446:
4444:
4443:
4439:
4437:
4436:
4432:
4430:
4429:
4425:
4424:
4422:
4420:
4416:
4410:
4409:
4405:
4403:
4402:
4398:
4396:
4395:
4391:
4390:
4388:
4386:
4382:
4376:
4375:
4371:
4369:
4368:
4364:
4362:
4361:
4357:
4355:
4354:
4350:
4348:
4347:
4343:
4341:
4340:
4336:
4334:
4333:
4329:
4327:
4326:
4322:
4320:
4319:
4315:
4313:
4312:
4308:
4306:
4305:
4301:
4300:
4298:
4296:
4292:
4289:
4287:
4283:
4273:
4272:
4268:
4266:
4265:
4261:
4259:
4258:
4254:
4252:
4251:
4247:
4245:
4244:
4240:
4238:
4237:
4233:
4231:
4230:
4226:
4224:
4223:
4219:
4217:
4216:
4212:
4210:
4209:
4205:
4203:
4202:
4198:
4196:
4195:
4191:
4189:
4188:
4184:
4182:
4181:
4177:
4176:
4174:
4172:
4168:
4162:
4161:
4157:
4156:
4154:
4152:
4151:Makaracetinae
4148:
4142:
4141:
4140:Pontobasileus
4137:
4135:
4134:
4130:
4128:
4127:
4123:
4121:
4120:
4116:
4114:
4113:
4112:Carolinacetus
4109:
4107:
4106:
4102:
4100:
4099:
4095:
4094:
4092:
4090:
4086:
4083:
4081:
4077:
4071:
4070:
4066:
4064:
4063:
4059:
4057:
4056:
4052:
4050:
4049:
4045:
4043:
4042:
4041:Andrewsiphius
4038:
4037:
4035:
4033:
4029:
4023:
4022:
4021:Himalayacetus
4018:
4016:
4015:
4011:
4009:
4008:
4004:
4003:
4001:
3999:
3998:Ambulocetidae
3995:
3989:
3988:
3987:Ichthyolestes
3984:
3982:
3981:
3977:
3975:
3974:
3970:
3969:
3967:
3965:
3961:
3955:
3951:
3949:
3945:
3943:
3939:
3937:
3933:
3931:
3927:
3926:
3923:
3918:
3911:
3906:
3904:
3899:
3897:
3892:
3891:
3888:
3877:
3873:
3869:
3865:
3861:
3857:
3853:
3849:
3845:
3841:
3840:
3836:morphology".
3831:
3828:
3823:
3819:
3814:
3809:
3804:
3799:
3795:
3791:
3787:
3780:
3777:
3772:
3768:
3763:
3758:
3753:
3748:
3744:
3740:
3736:
3729:
3727:
3725:
3723:
3721:
3717:
3712:
3708:
3701:
3694:
3692:
3690:
3686:
3681:
3677:
3673:
3669:
3665:
3661:
3657:
3653:
3645:
3643:
3641:
3639:
3637:
3633:
3628:
3624:
3620:
3616:
3612:
3608:
3604:
3600:
3593:
3590:
3585:
3581:
3577:
3573:
3569:
3565:
3561:
3557:
3553:
3546:
3544:
3542:
3540:
3538:
3534:
3529:
3525:
3521:
3517:
3513:
3509:
3505:
3501:
3494:
3492:
3490:
3488:
3486:
3484:
3482:
3478:
3473:
3469:
3465:
3461:
3457:
3453:
3446:
3439:
3437:
3435:
3433:
3431:
3427:
3422:
3418:
3414:
3410:
3406:
3402:
3395:
3393:
3391:
3389:
3387:
3385:
3383:
3381:
3379:
3377:
3373:
3368:
3364:
3357:
3354:
3350:(482): 1–366.
3349:
3342:
3335:
3333:
3331:
3329:
3325:
3320:
3316:
3312:
3305:
3303:
3301:
3297:
3292:
3288:
3283:
3278:
3273:
3268:
3264:
3260:
3256:
3252:
3248:
3241:
3239:
3237:
3235:
3233:
3231:
3229:
3227:
3225:
3223:
3221:
3219:
3217:
3215:
3213:
3211:
3209:
3207:
3205:
3203:
3201:
3199:
3197:
3195:
3191:
3187:(1): 121–148.
3186:
3182:
3178:
3171:
3169:
3167:
3165:
3163:
3161:
3159:
3157:
3155:
3153:
3151:
3149:
3147:
3145:
3141:
3134:
3132:
3130:
3126:
3122:
3117:
3115:
3111:
3107:
3103:
3099:
3095:
3090:
3085:
3083:
3079:
3075:
3071:
3067:
3063:
3059:
3055:
3047:
3045:
3042:
3038:
3037:
3032:
3031:
3026:
3025:
3020:
3019:
3014:
3010:
3006:
3002:
2998:
2993:
2983:
2973:
2964:
2953:
2941:
2939:
2937:
2933:
2927:
2925:
2920:
2916:
2912:
2908:
2904:
2900:
2892:
2887:
2882:
2875:
2868:
2867:
2859:
2858:
2850:
2849:
2841:
2840:
2837:
2836:
2830:
2829:
2826:
2825:
2817:
2816:
2808:
2807:
2799:
2798:
2795:
2794:
2793:
2786:
2785:
2782:
2781:
2778:
2777:
2776:
2769:
2768:
2762:
2761:
2758:
2757:
2754:
2753:
2752:
2745:
2744:
2738:
2737:
2734:
2733:
2730:
2729:
2728:
2721:
2720:
2714:
2713:
2707:
2706:
2703:
2702:
2694:
2693:
2685:
2684:
2676:
2675:
2667:
2666:
2658:
2657:
2654:
2653:
2652:
2645:
2644:
2641:
2640:
2637:
2636:
2635:
2628:
2627:
2621:
2620:
2617:
2616:
2613:
2612:
2611:
2604:
2603:
2597:
2596:
2593:
2592:
2584:
2583:
2580:
2579:
2578:
2571:
2570:
2567:
2566:
2563:
2562:
2561:
2554:
2553:
2547:
2546:
2540:
2539:
2536:
2535:
2532:
2531:
2530:
2523:
2522:
2516:
2515:
2512:
2511:
2503:
2502:
2494:
2493:
2490:
2489:
2488:
2481:
2480:
2477:
2476:
2473:
2472:
2471:
2464:
2463:
2457:
2456:
2453:
2452:
2449:
2448:
2447:
2440:
2439:
2433:
2432:
2426:
2425:
2419:
2418:
2415:
2414:
2411:
2410:
2409:
2402:
2401:
2394:
2393:
2389:
2387:
2383:
2379:
2375:
2371:
2367:
2363:
2358:
2348:
2344:
2340:
2330:
2319:
2305:
2304:
2296:
2295:
2287:
2286:
2278:
2277:
2269:
2268:
2260:
2259:
2251:
2250:
2242:
2241:
2233:
2232:
2224:
2223:
2215:
2214:
2211:
2210:
2204:
2203:
2200:
2199:
2196:
2195:
2189:
2188:
2182:
2181:
2178:
2177:
2174:
2173:
2172:
2165:
2164:
2158:
2157:
2154:
2153:
2145:
2144:
2136:
2135:
2127:
2126:
2118:
2117:
2109:
2108:
2100:
2099:
2091:
2090:
2087:
2086:
2085:
2078:
2077:
2074:
2073:
2070:
2069:
2068:
2061:
2060:
2054:
2053:
2050:
2049:
2046:
2045:
2044:
2037:
2036:
2030:
2029:
2026:
2025:
2017:
2016:
2013:
2012:
2011:
2004:
2003:
2000:
1999:
1996:
1995:
1994:
1987:
1986:
1980:
1979:
1973:
1972:
1969:
1968:
1965:
1964:
1963:
1956:
1955:
1949:
1948:
1945:
1944:
1941:
1940:
1939:
1932:
1931:
1925:
1924:
1921:
1920:
1912:
1911:
1908:
1907:
1906:
1899:
1898:
1895:
1894:
1891:
1890:
1889:
1882:
1881:
1878:
1877:
1874:
1869:
1868:
1862:
1861:
1855:
1854:
1851:
1850:
1847:
1846:
1845:
1838:
1837:
1831:
1830:
1824:
1823:
1820:
1819:
1816:
1815:
1814:
1807:
1806:
1800:
1799:
1796:
1795:
1792:
1791:
1790:
1783:
1782:
1779:
1778:
1775:
1774:
1773:
1766:
1765:
1759:
1758:
1755:
1754:
1751:
1750:
1749:
1742:
1741:
1738:
1737:
1734:
1733:
1732:
1731:Carolinacetus
1725:
1724:
1721:
1720:
1717:
1716:
1715:
1708:
1707:
1704:
1703:
1700:
1699:
1698:
1691:
1690:
1684:
1683:
1680:
1679:
1676:
1675:
1674:
1667:
1666:
1660:
1659:
1656:
1655:
1652:
1651:
1650:
1643:
1642:
1636:
1635:
1632:
1631:
1628:
1627:
1626:
1619:
1618:
1615:
1614:
1611:
1610:
1609:
1602:
1601:
1595:
1594:
1591:
1590:
1587:
1586:
1585:
1578:
1577:
1571:
1570:
1566:
1564:
1560:
1556:
1552:
1548:
1544:
1536:
1534:
1532:
1528:
1524:
1520:
1516:
1512:
1510:
1506:
1502:
1498:
1494:
1490:
1485:
1481:
1477:
1473:
1469:
1467:
1463:
1459:
1455:
1451:
1447:
1443:
1439:
1435:
1431:
1426:
1422:
1418:
1417:cortical bone
1413:
1409:
1401:
1397:
1393:
1388:
1384:
1381:
1377:
1373:
1369:
1365:
1361:
1357:
1353:
1349:
1344:
1342:
1338:
1334:
1330:
1326:
1322:
1317:
1315:
1311:
1307:
1303:
1299:
1295:
1291:
1287:
1283:
1278:
1274:
1270:
1266:
1263:The teeth of
1258:
1253:
1246:
1244:
1242:
1238:
1234:
1230:
1226:
1218:
1214:
1210:
1206:
1202:
1201:North America
1198:
1194:
1190:
1186:
1185:
1182:
1181:
1177:
1176:
1171:
1166:
1162:
1158:
1154:
1150:
1146:
1142:
1141:
1138:
1137:
1133:
1132:
1128:
1124:
1119:
1112:
1110:
1108:
1107:
1102:
1101:
1096:
1095:
1090:
1085:
1077:
1075:
1073:
1069:
1065:
1061:
1057:
1053:
1049:
1048:
1043:
1039:
1034:
1032:
1028:
1024:
1020:
1016:
1012:
1008:
1004:
1000:
996:
992:
988:
984:
980:
976:
972:
968:
964:
960:
952:
948:
946:
944:
940:
936:
932:
928:
923:
919:
915:
911:
907:
903:
899:
895:
891:
890:
885:
881:
880:"Eocetus" sp.
877:
873:
869:
862:
859:
857:
855:
851:
847:
843:
838:
834:
830:
826:
822:
814:
810:
806:
803:
799:
795:
791:
787:
783:
779:
778:
774:to the genus
773:
769:
765:
761:
756:
754:
750:
746:
742:
738:
734:
730:
726:
721:
719:
718:
713:
709:
708:
703:
699:
695:
691:
687:
683:
679:
675:
671:
667:
663:
659:
655:
651:
647:
643:
639:
635:
627:
622:
618:
616:
612:
608:
604:
600:
596:
592:
588:
584:
583:Tiasmyn River
580:
576:
572:
568:
565:
561:
557:
551:
547:
543:
538:
536:
534:
530:
526:
522:
518:
514:
510:
506:
502:
498:
494:
493:
488:
484:
480:
475:
473:
472:
467:
463:
459:
455:
451:
447:
443:
439:
435:
431:
429:
425:
424:
419:
418:
413:
409:
405:
404:cortical bone
401:
397:
393:
389:
385:
381:
377:
373:
369:
365:
361:
357:
353:
349:
345:
344:United States
341:
338:
334:
333:
321:Kellogg 1936
319:
316:
311:
308:
305:
302:
298:
290:
287:
284:
277:
274:
273:
271:
266:
259:
258:
251:
248:
247:
244:
238:
235:
234:
231:
225:
222:
221:
218:
215:
212:
211:
208:
205:
202:
201:
198:
195:
192:
191:
188:
185:
182:
181:
178:
175:
172:
171:
168:
165:
162:
161:
156:
151:
147:
144:
140:
136:
131:
125:
121:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
45:
39:
33:
29:
26:
22:
4496:
4447:
4440:
4434:
4433:
4426:
4419:Pachycetinae
4406:
4399:
4394:Basilosaurus
4392:
4372:
4365:
4358:
4351:
4344:
4337:
4330:
4323:
4318:Cynthiacetus
4316:
4309:
4302:
4295:Dorudontinae
4269:
4262:
4255:
4248:
4241:
4234:
4227:
4220:
4213:
4206:
4199:
4194:Crenatocetus
4192:
4185:
4180:Aegyptocetus
4178:
4171:Protocetinae
4158:
4138:
4131:
4126:Natchitochia
4124:
4119:Georgiacetus
4117:
4110:
4103:
4096:
4080:Protocetidae
4067:
4060:
4053:
4048:Attockicetus
4046:
4039:
4019:
4012:
4005:
3985:
3978:
3971:
3954:Artiodactyla
3843:
3837:
3830:
3796:(707): 707.
3793:
3789:
3779:
3742:
3738:
3710:
3706:
3655:
3651:
3602:
3592:
3559:
3555:
3503:
3499:
3455:
3451:
3404:
3400:
3366:
3362:
3356:
3347:
3318:
3314:
3254:
3250:
3184:
3180:
3128:
3124:
3118:
3102:Juglandaceae
3088:
3086:
3081:
3077:
3073:
3053:
3051:
3034:
3028:
3022:
3016:
3009:Basilotritus
3008:
2997:Basilosaurus
2996:
2991:
2989:
2981:
2980:The diet of
2935:
2932:neritic zone
2928:
2918:
2915:Basilosaurus
2914:
2910:
2907:Basilosaurus
2906:
2898:
2896:
2885:
2876:Paleobiology
2833:
2790:
2789:
2774:
2773:
2772:
2750:
2749:
2748:
2725:
2724:
2649:
2648:
2632:
2631:
2610:Basilosaurus
2608:
2607:
2575:
2574:
2558:
2557:
2527:
2526:
2485:
2484:
2468:
2467:
2444:
2443:
2406:
2405:
2385:
2381:
2377:
2373:
2369:
2365:
2361:
2356:
2354:
2346:
2342:
2338:
2207:
2192:
2169:
2168:
2082:
2081:
2067:Basilosaurus
2065:
2064:
2043:Cynthiacetus
2041:
2040:
2008:
2007:
1991:
1990:
1960:
1959:
1936:
1935:
1904:
1903:
1902:
1887:
1886:
1885:
1872:
1842:
1841:
1811:
1810:
1789:Georgiacetus
1787:
1786:
1770:
1769:
1746:
1745:
1729:
1728:
1712:
1711:
1697:Aegyptocetus
1695:
1694:
1671:
1670:
1647:
1646:
1623:
1622:
1606:
1605:
1582:
1581:
1563:Basilotritus
1562:
1559:Basilotritus
1558:
1554:
1547:paraphyletic
1542:
1540:
1530:
1526:
1522:
1518:
1514:
1513:
1509:ischiac part
1500:
1496:
1492:
1483:
1479:
1471:
1470:
1465:
1453:
1449:
1434:neural canal
1424:
1421:pachyostosis
1411:
1404:
1399:
1396:Basilosaurus
1395:
1391:
1379:
1372:neural canal
1367:
1363:
1359:
1356:Basilosaurus
1355:
1347:
1345:
1336:
1333:Basilotritus
1332:
1328:
1324:
1320:
1318:
1314:Basilotritus
1313:
1309:
1306:canine teeth
1297:
1293:
1285:
1281:
1275:. The first
1273:tooth enamel
1268:
1264:
1262:
1256:
1224:
1222:
1216:
1208:
1196:
1193:Basilotritus
1192:
1188:
1179:
1178:
1169:
1144:
1135:
1134:
1126:
1122:
1104:
1098:
1094:Basilosaurus
1092:
1089:Basilotritus
1088:
1083:
1081:
1071:
1067:
1063:
1060:Basilotritus
1059:
1056:P. paulsonii
1055:
1051:
1045:
1041:
1037:
1035:
1030:
1026:
1022:
1018:
1014:
1010:
1007:P. paulsonii
1006:
998:
994:
990:
987:P. paulsonii
986:
982:
974:
970:
966:
963:Basilotritus
962:
958:
956:
950:
938:
934:
930:
926:
922:Basilotritus
921:
917:
913:
905:
902:nomen dubium
897:
893:
889:Basilotritus
887:
883:
879:
867:
865:
861:Basilotritus
860:
853:
849:
841:
836:
824:
821:Mark D. Uhen
818:
812:
801:
794:pachyostosis
785:
775:
771:
767:
763:
757:
744:
732:
722:
715:
711:
705:
701:
697:
693:
689:
666:Lower Saxony
649:
645:
631:
625:
617:regardless.
614:
606:
602:
559:
555:
554:
549:
545:
532:
512:
504:
496:
492:Basilosaurus
490:
476:
469:
465:
461:
457:
454:Basilotritus
453:
449:
445:
441:
437:
433:
432:
427:
422:
421:
416:
415:
411:
400:pachyostotic
383:
367:
340:basilosaurid
331:
330:
329:
317:
310:Basilotritus
309:
292:(Uhen, 1999)
288:
275:
256:
255:
243:Pachycetinae
213:Infraorder:
207:Artiodactyla
31:
25:
4497:Platyosphys
4449:Platyosphys
4401:Basiloterus
4367:Supayacetus
4346:Pontogeneus
4311:Chrysocetus
4304:Ancalecetus
4250:Qaisracetus
4236:Phiomicetus
4229:Peregocetus
4160:Makaracetus
4062:Kutchicetus
4007:Ambulocetus
3964:Pakicetidae
3917:Archaeocete
3790:Commun Biol
3458:: 273–286.
2727:Supayacetus
2577:Ancalecetus
2446:Chrysocetus
2395:Pelagiceti
2382:Supayacetus
2010:Ancalecetus
1962:Chrysocetus
1844:Supayacetus
1673:Qaisracetus
1400:Qaisracetus
1376:neural arch
1341:protocetids
1247:Description
1121:Remains of
1106:Platyosphys
1100:Basiloterus
1052:P. robustus
1001:could be a
999:P. robustus
991:P. robustus
975:Platytholus
959:Platyosphys
949:Revival of
935:Platyosphys
894:Platyosphys
884:Platyosphys
764:Platyosphys
650:Z. wanklyni
634:Barton Clay
550:Platyosphys
462:Platyosphys
450:Platyosphys
337:pachycetine
318:Platyosphys
276:P. paulsoni
236:Subfamily:
4582:Categories
4435:Pachycetus
4428:Antaecetus
4360:Stromerius
4353:Saghacetus
4332:Masracetus
4264:Takracetus
4257:Rodhocetus
4243:Protocetus
4208:Gaviacetus
4201:Dhedacetus
4187:Artiocetus
4133:Pappocetus
4105:Babiacetus
4055:Dalanistes
4014:Gandakasia
3942:Vertebrata
3940:Subphylum
3745:: e16541.
3619:4630500767
3321:: 111–112.
3135:References
3129:Pachycetus
3125:Pachycetus
3121:glauconite
3078:Pachycetus
3074:Pachycetus
3054:Pachycetus
2992:Pachycetus
2982:Pachycetus
2936:Pachycetus
2919:Pachycetus
2911:Pachycetus
2899:Pachycetus
2886:Pachycetus
2560:Saghacetus
2386:Pachycetus
2378:Antaecetus
2366:Pachycetus
2357:Pachycetus
2347:Antaecetus
2343:Antaecetus
2339:Pachycetus
1993:Saghacetus
1873:MUSM 1443
1813:Babiacetus
1748:Gaviacetus
1714:Protocetus
1649:Artiocetus
1625:Rodhocetus
1555:Pachycetus
1543:Pachycetus
1519:Antaecetus
1515:Pachycetus
1497:Pachycetus
1493:Pachycetus
1484:Pachycetus
1472:Pachycetus
1466:Pachycetus
1454:Pachycetus
1450:Antaecetus
1425:Pachycetus
1423:and gives
1412:Pachycetus
1392:Pachycetus
1374:) and the
1368:Pachycetus
1364:Pachycetus
1348:Pachycetus
1325:Antaecetus
1321:Pachycetus
1310:Pachycetus
1298:Pachycetus
1294:Antaecetus
1286:Pachycetus
1282:Antaecetus
1269:Antaecetus
1265:Pachycetus
1235:) and the
1165:Priabonian
1145:Pachycetus
1127:Antaecetus
1123:Pachycetus
1084:Pachycetus
1064:Pachycetus
1047:Antaecetus
1038:Pachycetus
1031:Pachycetus
1023:Pachycetus
983:Pachycetus
971:Pachycetus
967:Pachycetus
951:Pachycetus
868:Pachycetus
823:described
784:described
698:Pachycetus
644:and named
626:Pachycetus
560:Pachycetus
556:Pachycetus
546:Pachycetus
533:Pachycetus
513:Pachycetus
505:Pachycetus
497:Pachycetus
471:Antaecetus
458:Pachycetus
446:Pachycetus
434:Pachycetus
412:Pachycetus
384:Pachycetus
380:Priabonian
378:and Early
368:Pachycetus
332:Pachycetus
257:Pachycetus
124:Priabonian
32:Pachycetus
4512:Q20817932
4442:Perucetus
4374:Zygorhiza
4271:Togocetus
4222:Maiacetus
4215:Indocetus
4098:Aegicetus
3980:Nalacetus
3973:Pakicetus
3876:260433513
3680:1631-0683
3584:129160656
2924:sirenians
2529:Zygorhiza
1938:Zygorhiza
1608:Maiacetus
1584:Pakicetus
1537:Phylogeny
1531:P. wardii
1446:cartilage
1442:ligaments
1337:Zygorhiza
1237:North Sea
1209:P. wardii
1197:P. wardii
1161:Bartonian
1082:The name
1078:Etymology
989:predates
876:Kurenevka
872:Nagornoye
842:E. wardii
837:E. wardii
802:P. einori
798:sirenians
777:Zygorhiza
733:Zeuglodon
686:mysticete
682:sirenians
662:Helmstedt
624:The name
585:south of
479:sirenians
388:vertebrae
372:Bartonian
358:(chiefly
289:P. wardii
173:Kingdom:
167:Eukaryota
38:Bartonian
21:Pakicetus
4506:Wikidata
4472:Category
4339:Ocucajea
3948:Mammalia
3936:Chordata
3930:Animalia
3928:Kingdom
3868:37532931
3822:37563270
3813:10415296
3771:38774542
3762:11107809
3627:54060094
3528:85977537
3369:: 17–20.
3291:36288346
3251:PLOS ONE
3098:Fagaceae
2470:Ocucajea
2362:Tutcetus
2171:Ocucajea
1438:synovial
1302:Incisors
1277:premolar
1173:Paulson.
1068:B. uheni
846:Virginia
819:In 1999
737:Koropove
587:Chyhyryn
564:botanist
501:seafloor
483:manatees
376:Lutetian
352:Virginia
301:Synonyms
268:Species
223:Family:
197:Mammalia
187:Chordata
183:Phylum:
177:Animalia
163:Domain:
120:Lutetian
4552:3612303
4539:1033898
4526:4832522
4408:Eocetus
4325:Dorudon
3934:Phylum
3848:Bibcode
3713:(1–16).
3707:Deinsea
3660:Bibcode
3564:Bibcode
3508:Bibcode
3460:Bibcode
3409:Bibcode
3282:9604876
3259:Bibcode
3110:tupelos
3058:estuary
2903:ballast
2835:Neoceti
2634:Dorudon
2408:Eocetus
2209:Neoceti
2084:Dorudon
1772:Eocetus
1551:Neoceti
1521:, with
1360:Eocetus
1189:Eocetus
1157:Germany
1149:Ukraine
1113:Species
1066:, with
943:Morocco
850:Eocetus
790:Pyrohiv
678:Belgium
636:of the
591:Ukraine
569:at the
487:dugongs
364:Ukraine
360:Germany
249:Genus:
217:Cetacea
203:Order:
193:Class:
111:↓
3952:Order
3946:Class
3874:
3866:
3839:Nature
3820:
3810:
3769:
3759:
3678:
3625:
3617:
3582:
3526:
3289:
3279:
1541:While
1290:molars
1153:Russia
1103:) and
710:) and
674:Leuven
521:sharks
356:Europe
354:) and
350:&
4565:36815
4534:IRMNG
3872:S2CID
3739:PeerJ
3703:(PDF)
3623:S2CID
3580:S2CID
3524:S2CID
3448:(PDF)
3344:(PDF)
3114:palms
3106:ferns
3094:flora
3001:orcas
1505:ilium
1229:Spain
1062:into
1054:into
969:. As
844:into
714:(now
704:(now
660:) of
519:like
4521:GBIF
3864:PMID
3818:PMID
3767:PMID
3676:ISSN
3615:OCLC
3287:PMID
3112:and
3033:and
2999:and
2942:Diet
2372:and
2341:was
1444:and
1398:and
1358:and
1304:and
1155:and
1097:and
1017:and
979:ICZN
961:and
766:for
575:Kyiv
548:and
525:rays
523:and
485:and
398:and
392:ribs
390:and
362:and
283:type
122:and
50:PreꞒ
3856:doi
3844:620
3808:PMC
3798:doi
3757:PMC
3747:doi
3668:doi
3607:doi
3572:doi
3516:doi
3468:doi
3456:111
3417:doi
3277:PMC
3267:doi
1243:).
1163:to
1005:of
753:WW2
672:in
573:in
4584::
4562::
4549::
4536::
4523::
4508::
3870:.
3862:.
3854:.
3842:.
3816:.
3806:.
3792:.
3788:.
3765:.
3755:.
3743:12
3741:.
3737:.
3719:^
3711:21
3709:.
3705:.
3688:^
3674:.
3666:.
3656:13
3654:.
3635:^
3621:.
3613:.
3601:.
3578:.
3570:.
3560:85
3558:.
3554:.
3536:^
3522:.
3514:.
3504:73
3502:.
3480:^
3466:.
3454:.
3450:.
3429:^
3415:.
3405:87
3403:.
3375:^
3367:20
3365:.
3346:.
3327:^
3317:.
3313:.
3299:^
3285:.
3275:.
3265:.
3255:17
3253:.
3249:.
3193:^
3185:20
3183:.
3179:.
3143:^
3108:,
3104:,
3100:,
3027:,
3021:,
2926:.
2380:.
1394:,
1343:.
1195:,
1151:,
1074:.
1025:.
1013:,
945:.
904:.
676:,
664:,
638:UK
589:,
535:.
430:.
382:.
100:Pg
44:Ma
3909:e
3902:t
3895:v
3878:.
3858::
3850::
3824:.
3800::
3794:6
3773:.
3749::
3682:.
3670::
3662::
3629:.
3609::
3586:.
3574::
3566::
3530:.
3518::
3510::
3474:.
3470::
3462::
3423:.
3419::
3411::
3319:9
3293:.
3269::
3261::
1231:(
878:(
481:(
346:(
285:)
281:(
253:†
240:†
227:†
126:.
105:N
95:K
90:J
85:T
80:P
75:C
70:D
65:S
60:O
55:Ꞓ
23:.
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