Antaecetus

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Antaecetus
Temporal range: Bartonian
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Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Infraorder: Cetacea
Family: Basilosauridae
Subfamily: Pachycetinae
Genus: Antaecetus
Gingerich, Amane & Zhouri, 2022 [1]
Type species
Antaecetus aithai
(Gingerich & Zhouri, 2015) [2]

Antaecetus is an extinct genus of pachycetine basilosaurid from the middle Eocene Aridal Formation of Morocco as well as the Fayum, Egypt. Antaecetus, although known from fewer remains in total, is markedly more complete than the closely related Pachycetus , with one specimen preserving large parts of the vertebral column up to the lumbar vertebrae and a well preserved skull. Based on these remains Antaecetus appears to have been smaller than Pachycetus, with a proportionally smaller head and much more gracile teeth. Both genera however share a highly osteosclerotic and pachyostotic skeleton, greatly increasing their weight and possibly serving as additional ballast. In turn the elongation and thickening of the vertebrae severely impacts the animals movement, most likely causing it to have been much slower and far less mobile than other archaeocetes. It has been proposed that Antaecetus was a slow moving inhabitant of shallow coastal waters, where it would ambush fish and marine molluscs. Only a single species of Antaecetus is known, Antaecetus aithai, which was originally described as a species of Pachycetus.

Contents

History and naming

Initially thought to represent a species of Platyosphys (now Pachycetus ), Antaecetus was described in 2015 based on fossils from the Aridal Formation of the Gueran depression, southwestern Morocco. The fossil material, consisting of a partial skull (later thought to have been that of Eocetus [1] ), a tympanic bulla and various vertebrae was recovered in 2014 along with remains of various other basal whales, leading to the recognition of the Gueran depression as an important assemblage for research on early cetaceans. [2] Several years after this initial paper was published a second publication was released, dealing in part with the taxonomy of Platyosphys in an attempt to clarify its status. At the same time, following the discovery of an additional, much better preserved specimen from El Brije, P. aithai was reexamined and found to be sufficiently distinct to warrant it being placed in its own genus, which the authors dubbed Antaecetus. The principal specimen of said study consists of a cranium, the second to eleventh thoracic vertebrae and an additional ten lumbar vertebrae. The same team further established the clade Pachycetinae based on shared characteristics of the skeleton that distinguish Antaecetus and Pachycetus from other basilosaurids. Material of Antaecetus has also been recovered from Fayum, Egypt, but not yet described or catalogued. [1]

Antaecetus was named after Antaeus (also known as Antaois or Anti), a figure from Greek mythology, in combination with the word "cetus" meaning whale. In the myths Antaeus was the son of Gaia and the sea god Poseidon and was said to reside in western North Africa with his tomb being found in Morocco. The species name derives from Amer Ait Ha, an experienced fossil collector who was responsible for guiding researchers to the Gueran locality. [2]

Description

Antaecetus (top) compared to Saghacetus (bottom) Antaecetus Saghacetus comparisson.png
Antaecetus (top) compared to Saghacetus (bottom)

The skull of Antaecetus is noticeably smaller than that of Pachycetus relative to its body, measuring a total of 69.5 cm (27.4 in) from the tip of the premaxillae to the occipital condyles. At its widest point the skull measures 31.4 cm (12.4 in). Antaecetus possesses three incisors in either premaxilla, followed by a canine tooth, four premolars and two molars in either maxilla. This means the dental formula for the upper jaw is 3.1.4.2., which is the standard formula for basilosaurids. The connection between the rostrum and the basicranium is formed by a narrow yet robust intertemporal constriction, which consists of the pterygoid and palatine bones. The constriction is bordered by the temporal fossae, which in turn sit next to the slender jugal that connects the maxillae with the squamosals. In its overall form the skull of Antaecetus resembles the typical basilosaurid condition, featuring a narrow rostrum connected to a much wider basicranium. [1]

The teeth are much more gracile than those of Pachycetus and lack the crenulated tooth enamel, instead having a smooth surface. The three incisors are single-rooted and conical with laterally compressed crowns. Among them the second incisors stand out as being more caniniform than the others and also projecting much further out of the tooth socket than the other two. Overall the canine teeth are similar to the incisors and roughly the same size as the second canine. Philip Gingerich and colleagues note that canine size may be sexually dimorphic in archaeocetes, meaning that the El Brije specimen could be a female. Following the canines there is a diastema, a toothless section that separates these early teeth from the premolars and molars. The first premolar is regarded as being among the distinguishing features between Antaecetus and Pachycetus. In the former, the tooth has a single root much like the incisors and canines and different from the double-rooted structure displayed by the second premolar and the teeth further back. In Pachycetus on the other hand the first premolar is already double-rooted. Furthermore, the posterior premolars have three accessory cusps or denticles on either side of the central apical cusp, one less than what is seen in its relative. The molars only have two accessory cusps each and the second molar is much simpler than the first. Though the protocones of the molars have been lost, the teeth of Antaecetus still retain an expansion in this area hinting at where the structure was once located. Both molars have a weak lateral cingulum, but differ in that the first has a stronger medial cingulum while there is none in the second molar. [1]

The full skeleton of the El Brije Antaecetus specimen Antaecetus el brije.png
The full skeleton of the El Brije Antaecetus specimen

The anterior vertebrae of the thorax (torso) of Antaecetus show clear tapering, with the posterior (back) end being much wider relative to the beginning of the vertebrae. These early vertebrae show a clear transition in how they articulate with the ribs, as they have capitular facets for the reception of the heads of the ribs on both ends of the centrum. However, further down the spine these facets become restricted to the anterior end and resemble more of a pit than a true facet. These middle thoratic vertebrae furthermore display a developed diapophysis that articules with the tubercles of the ribs. The presence of such a diapophysis is particularly highlighted by Gingerich and Samir Zhouri, who used this feature to distinguish A. aithai from the two species of Pachycetus recognized by them. The diapophysis is eventually lost by the posterior vertebrae of the thorax and the rib articulation is altered even further, with the capitular facets becoming even more pit-like and now being situated atop the elevated surface of the parapophysis. It has been noted that this would suggest that the ribs didn't articulate through synovial joints like in other basilosaurids but through cartilage or ligaments like in early sirenians. The neural spines are described as slender and relatively short. The lumbar vertebrae strongly resemble those of Pachycetus, with both the centra themselves and the robust transverse processes being elongated, the latter stretching almost the entire length of their respective centra. [2] [1]

Another significant trait of the vertebrae is their density relative to those of other basilosaurids. The vertebrae are osteosclerotic and pachyostotic, meaning that they feature thickened rings of cortical bone that are denser than would be typical. Gingerich and Zhouri initially characterize the vertebrae as being unique in containing cones of cancellous bone instead of cylinders, [1] though this notion was later rejected by Henk Jan van Vliet as having been much more common than initially assumed. [3] The lower surface of the centra appears to have had a pockmarked-texture, caused by the presence of nutrient foramina that penetrate the cortical bone. [2] The ribs of Antaecetus are robust, but only the anterior ones are pachyostotic. [1]

Antaecetus is described as having been a medium-sized basilosaurid, significantly smaller than the related species Pachycetus paulsonii, slightly smaller than Pachycetus wardii and in a similar size range as Pappocetus , a contemporary protocetid. The lumbar vertebrae of large males are in the same size range as those of female Eocetus and the skull is similar in size to that of Saghacetus. [2] [1]

Phylogeny

A phylogenetic analysis conducted in 2023 confirmed the hypothesis that Antaecetus was a close relative of Pachycetus, the relationship between them being strongly supported by the results and validating the clade Pachycetinae that was erected to contain the two taxa. Within the group, Antaecetus was recovered as the most derived species and notably the sister taxon to Pachycetus wardii from the eastern United States, suggesting that Pachycetus may be paraphyletic. Although typically considered basilosaurids, the same study recovered pachycetines as being more closely related to the clade Neoceti, rendering Basilosauridae paraphyletic as well. [4]

Pelagiceti

Eocetus

Chrysocetus

Ocucajea

Tutcetus rayanensis

Zygorhiza

Saghacetus

Ancalecetus

Basilosaurus

Dorudon

Pontogeneus peruvianus

Supayacetus

Pachycetus paulsonii

Pachycetus wardii

Antaecetus aithai

Neoceti

Paleobiology

Live reconstruction Antaecetus aithai.png
Live reconstruction

The elongation of the vertebrae in Antaecetus resembles what is known from Basilosaurus , consequently supporting the idea that their locomotion could have been similar. This would suggest that Antaecetus swam by undulating the entire body. However, the length of the transverse processes would have restricted this movement somewhat. More specifically, the reduced space between the processes meant that the side-to-side flexibility was limited, limiting the animal to swimming through up- and downstrokes. The increase in bone density would have further impacted its lifestyle, with Gingerich and colleagues listing a variety of potential causes for such an adaptation. For instance, the corresponding increase in ballast and lung volume would have been useful for a slow-moving animal, possibly one feeding in shallow waters close to the ocean floor. The articulation of the ribs would also fall in line with this interpretation, as ligamentous or cartilaginous rib articulation allows for the thorax to expand during air intake while also allowing the animal to collapse the thorax at the ocean floor to minimize buoyancy. At the same time, the increased density took a further toll on the mobility of Antaecetus, likely making it much slower to accelerate in addition to its poor maneuverability. [1]

From this Gingerich, Ayoub Amane and Zhouri hypothesize that Antaecetus, like Pachycetus, was a slow-moving animal inhabiting shallow coastal waters. However, its precise ecology proved to be more elusive, with its small, gracile teeth unfit to deal with either vegetation or hard-shelled prey (ruling out lifestyles akin to those of manatees and sea otters). Feeding directly from the ocean floor is also dismissed, as contact with sediment would lead the teeth to undergo rapid abrasion. The poor maneuverability and slow speed of Antaecetus indicate that it was not built for pursuing prey either, leaving the possibility that Antaecetus was an ambush hunter preying on fish and invertebrates. [1]

The remains of Antaecetus are known from the Aridal Formation of Morocco, where the animal coexisted with at least 5 additional early whales of different sizes. Antaecetus was one of the larger whales of this assemblage, together with the large protocetid Pappocetus and the basilosaurid Eocetus schweinfurthii, which was the largest cetacean of the locality. The remaining three taxa were all smaller than Antaecetus, including two indeterminate protocetids and the small basilosaurid Chrysocetus fouadassaii , which in terms of size falls in between the ranges of the afforementioned protocetids. [2]

Related Research Articles

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Rodhocetus is an extinct genus of protocetid early whale known from the Lutetian of Pakistan. The best-known protocetid, Rodhocetus is known from two partial skeletons that taken together give a complete image of an Eocene whale that had short limbs with long hands and feet that were probably webbed and a sacrum that was immobile with four partially fused sacral vertebrae. It is one of several extinct whale genera that possess land mammal characteristics, thus demonstrating the evolutionary transition from land to sea.

<i>Ambulocetus</i> Genus of extinct mammals of the order Cetacea

Ambulocetus is a genus of early amphibious cetacean from the Kuldana Formation in Pakistan, roughly 48 or 47 million years ago during the Early Eocene (Lutetian). It contains one species, Ambulocetus natans, known solely from a near-complete skeleton. Ambulocetus is among the best-studied of Eocene cetaceans, and serves as an instrumental find in the study of cetacean evolution and their transition from land to sea, as it was the first cetacean discovered to preserve a suite of adaptations consistent with an amphibious lifestyle. Ambulocetus is classified in the group Archaeoceti—the ancient forerunners of modern cetaceans whose members span the transition from land to sea—and in the family Ambulocetidae, which includes Himalayacetus and Gandakasia.

<i>Basilosaurus</i> Prehistoric cetacean genus from the Late Eocene epoch

Basilosaurus is a genus of large, predatory, prehistoric archaeocete whale from the late Eocene, approximately 41.3 to 33.9 million years ago (mya). First described in 1834, it was the first archaeocete and prehistoric whale known to science. Fossils attributed to the type species B. cetoides were discovered in the United States. They were originally thought to be of a giant reptile, hence the suffix "-saurus", Ancient Greek for "lizard". The animal was later found to be an early marine mammal, which prompted attempts at renaming the creature, which failed as the rules of zoological nomenclature dictate using the original name given. Fossils were later found of the second species, B. isis, in 1904 in Egypt, Western Sahara, Morocco, Jordan, Tunisia, and Pakistan. Fossils have also been unearthed in the southeastern United States and Peru.

<span class="mw-page-title-main">Basilosauridae</span> Family of mammals

Basilosauridae is a family of extinct cetaceans. They lived during the middle to the early late Eocene and are known from all continents, including Antarctica. They were probably the first fully aquatic cetaceans. The group is noted to be a paraphyletic assemblage of stem group whales from which the monophyletic Neoceti are derived.

<i>Dorudon</i> Genus of ancient whales

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<span class="mw-page-title-main">Archaeoceti</span> Paraphyletic group of primitive cetaceans from Early Eocene to Late Oligocene

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Pachycetus is an extinct genus of pachycetine basilosaurid from Middle Eocene of the eastern United States and Europe. The best known remains generally suggest that Pachycetus lived during the Bartonian, 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 Lutetian and Early Priabonian. Pachycetus is primarily known from vertebrae and ribs and is characterized by its highly osteosclerotic and pachyostotic skeleton. This means the bones not only featured thickened rings of cortical bone surrounding the internal cancellous bone, but the cortical bone was furthermore much denser than in other basilosaurids. Two species of Pachycetus are recognized: Pachycetus paulsonii from Europe and Pachycetus wardii from the United States. A third species might be represented by "Zeuglodon" wanklyni.

<i>Ichthyolestes</i> Genus of mammals

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<i>Georgiacetus</i> Extinct genus of mammals

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Gaviacetus is an extinct archaeocete whale that lived approximately 45 million years ago. Gaviacetus was named for its characteristic narrow rostrum and the fast pursuit predation suggested by its unfused sacral vertebrae.

Basiloterus is an extinct genus of late-Eocene archaeocete whale from the Drazinda Formation in southwestern Punjab, Pakistan and possibly also the Barton Group of England. Known from two isolated lumbar vertebrae, the elongated nature of these elements has been taken as possible evidence that Basiloterus was a close relative of the better-known Basilosaurus. This was also the reasoning behind its name, which roughly translates to "another king". However, publications since then not only lead to some major changes of the internal relationships within Basilosauridae but have also called into question how diagnostic elongated vertebrae are for members of this group, as other early whales have developed similar anatomy independently. Though the identity of Basiloterus as a basilosaurid is generally maintained, its exact position within more recent interpretations of the family is unclear.

Chrysocetus is a genus of extinct early whale known from Late Eocene-aged fossils of the eastern United States and western Africa.

Protocetidae, the protocetids, form a diverse and heterogeneous group of extinct cetaceans known from Asia, Europe, Africa, South America, and North America.

<i>Aegyptocetus</i> Species of mammal

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Babiacetus is an extinct genus of early cetacean that lived during the late Lutetian middle Eocene of India . It was named after its type locality, the Harudi Formation in the Babia Hills, Kutch District, Gujarat, India.

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Eocetus is an extinct protocetid early whale known from the early-late Eocene Giushi Formation in Gebel Mokattam, outside Cairo, Egypt. The specimen was first named by Fraas as Mesocetus schweinfurthi. However, the name Mesocetus was previously used causing a change to the species name to Eocetus schweinfurthi. Since the genus was first described in the early 20th century, several other specimens, mostly isolated vertebrae, have been attributed to Eocetus, but the taxonomic status of these widely distributed specimens remain disputed.

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Pachycetinae is an extinct subfamily of basilosaurid cetaceans that lived during the middle Eocene. The best dated remains stem from Bartonian strata, but some finds suggest that they could have first appeared during the Lutetian and may have survived until the Priabonian. Fossils of pachycetines are chiefly known from the southern United States, Ukraine, Morocco and Germany, among others. They differ from other basilosaurids in having pachyostotic and osteosclerotic vertebrae 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 sirenians. Analysis of the teeth suggests that pachycetines had a varying diet, with the robust teeth of the larger Pachycetus indicating that it possibly fed on sharks, whereas the more gracile teeth of Antaecetus suggest a diet of smaller prey items. The clade currently only includes two genera, Antaecetus and Pachycetus, but a 2023 study suggests that the Peruvian Supayacetus may at least be a close relative.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 Gingerich, P.D.; Amane, A.; Zouhri, S. (2022). "Skull and partial skeleton of a new pachycetine genus (Cetacea, Basilosauridae) from the Aridal Formation, Bartonian middle Eocene, of southwestern Morocco". PLOS ONE. 17 (10): e0276110. Bibcode:2022PLoSO..1776110G. doi: 10.1371/journal.pone.0276110 . PMC   9604876 .
  2. 1 2 3 4 5 6 7 Gingerich, P.D.; Zouhri, S. (2015). ""New fauna of archaeocete whales (Mammalia, Cetacea) from the Bartonian middle Eocene of southern Morocco"" (PDF). Journal of African Earth Sciences. 111: 273–286. Bibcode:2015JAfES.111..273G. doi:10.1016/j.jafrearsci.2015.08.006.
  3. van Vliet, H.J.; Bosselaers, M.E.J.; Munsterman, D.K.; Dijkshoorn, M.L.; de Groen, J.J.; Post, K. (2024). "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". PeerJ. 12: e16541. doi: 10.7717/peerj.16541 .
  4. Antar, M.S.; Glaohar, A.S.; El-Desouky, H.; Seiffert, E.R.; El-Sayed, S.; Claxton, A.G.; Sallam, H.M. (2023). "A diminutive new basilosaurid whale reveals the trajectory of the cetacean life histories during the Eocene". Commun Biol. 6 (707): 707. doi: 10.1038/s42003-023-04986-w . PMC   10415296 . PMID   37563270.