Romaleodelphis
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''Romaleodelphis'' is a genus of extinct archaic
toothed whale The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales with teeth, such as beaked whales and the sperm whales. 73 species of toothed wha ...
from the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
Ebelsberg Formation of
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
. It had long and slender jaws and homodont teeth and roamed the marine waters of the Central
Paratethys The Paratethys sea, Paratethys ocean, Paratethys realm or just Paratethys (meaning "beside Tethys"), was a large shallow inland sea that covered much of mainland Europe and parts of western Asia during the middle to late Cenozoic, from the lat ...
that once covered much of Europe. Phylogenetic analysis has shown that ''Romaleodelphis'' is most closely related to other odotontocetes previously grouped together as the "'' Chilcacetus'' clade, now a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
grade basal to true dolphins and
beaked whale Beaked whales (systematic name Ziphiidae) are a Family (biology), family of cetaceans noted as being one of the least-known groups of mammals because of their deep-sea habitat, reclusive behavior and apparent low abundance. Only three or four of ...
s. The inner ear of ''Romaleodelphis'' shows that it was capable of hearing hearing narrow-band high frequency sounds, an ability it convergently evolved with several modern types of whales. It is possible that this was a useful tool in evading other predatory whales, though some recent research suggests that it may have also been an early step in the development of the cetacean
biosonar Echolocation, also called bio sonar, is a biological active sonar used by several animal groups, both in the air and underwater. Echolocating animals emit calls and listen to the echoes of those calls that return from various objects near them ...
.


History and naming

The holotype skull of ''Romaleodelphis'' was discovered in 1980 in the sediments of the Ebelsberg Formation during the construction of a hydroelectric powerplant near the town of Pucking,
Upper Austria Upper Austria ( ; ; ) is one of the nine States of Austria, states of Austria. Its capital is Linz. Upper Austria borders Germany and the Czech Republic, as well as the other Austrian states of Lower Austria, Styria, and Salzburg (state), Salzbur ...
. At the time, several amateur paleontologists, among them Jürgen Pollerspöck, removed fossil material from the site to prevent their destruction. The total material collected for ''Romaleodelphis'' consisted of a multitude of skull fragments that were put together by the collectors as well as several postcranial remains. Put together, the skull is mostly complete but lacks the tips of the rostrum and much of the basicranium. However, while the skull material was donated to the Staatliche Naturwissenschaftliche Sammlungen Bayerns by Pollerspöck, the postcranial material was kept by other collectors and their whereabouts are currently unknown. Despite being donated in 1992, it would take another 30 years for the skull to be described, with the paper being published by Catalina Sanchez-Posada and colleagues in 2024. The name ''Romaleodelphis'' derives from the Greek "'romaléos" meaning sturdy or robust, in reference to the strong zygomatic process of the squamosal bone, and the -suffix "delphis". This suffix is commonly employed for the scientific names of dolphins and traces its meaning to the Greek "dophis" for womb, initially used to highlight the fact that unlike fish, dolphins have a uterus. The species name on the other hand references Jürgen Pollerspöck, who discovered, saved and donated the holotype fossil.


Description

''Romaleodelphis'' was a medium sized toothed whale with long and slender jaws that make up about 71% of the total skull length. The animal was homodont, meaning that like most modern toothed whales and unlike several now extinct forms that coexisted with ''Romaleodelphis'' it possessed only a single type of tooth in its jaws.


Phylogeny

The
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
conducted in the type description of ''Romaleodelphis'' recovered 15 most parsimonious trees, the strict consensus of which showing that the animal was a derived toothed whale. Specifically, ''Romaleodelphis'' was found to clade in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
with '' Chilcacetus'', '' Argyrocetus bakersfieldensis'' and a larger grouping consisting of the extant members of
Delphinidae Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as the ...
(dolphins) and
Ziphiidae Beaked whales (systematic name Ziphiidae) are a family of cetaceans noted as being one of the least-known groups of mammals because of their deep-sea habitat, reclusive behavior and apparent low abundance. Only three or four of the 24 existing s ...
(beaked whales), their fossil relatives as well as a series of fossil whales previously grouped as the "''Chilcacetus'' clade". In the strict consensus tree, the "''Chilcacetus'' clade" forms an evolutionary grade leading up to dolphins and beaked whales and includes the name-giving ''Chilcacetus'', ''Romaleodelphis'', three species of '' Argyrocetus'' (none of which are each others closest relatives) and '' Macrodelphinus''. More broadly, this phylogeny finds that the "''Chilcacetus'' clade", dolphins and beaked whales are each others closest relatives and that the sister group to these three is formed by the river dolphins of the
Platanistoidea River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant rive ...
and their extinct relatives the Eurhinodelphinidae. Other taxa included in the study were ''Squalodon'' and ''Waipatia'', two much more basally branching toothed whales. The supplementary material further includes a 50% majority-rule consensus tree in which rather than being a grade, a slimmed down version of the "''Chilcacetus'' clade" (consisting of ''Romaleodelphis'', ''Argyrocetus bakersfieldensis'' and ''Chilcacetus'') actually forms a monophyletic group with ''Romaleodelphis'' as the earliest diverging member. It is therefore possible that the "''Chilcacetus'' clade" originated in the Paratethys before spreading across the Atlantic and colonizing the coasts of North and South America, were most members of this group are known from.


Paleobiology


Hearing abilities

Principal component analysis Principal component analysis (PCA) is a linear dimensionality reduction technique with applications in exploratory data analysis, visualization and data preprocessing. The data is linearly transformed onto a new coordinate system such that th ...
(PCA) has shown that the anatomy of ''Romaleodelphis earbones would indicate that it was capable of hearing narrow-band high frequency sounds. This makes ''Romaleodolphin'' the first known archaic homodont toothed whale with the ability to perceive the kinds of sounds made by modern toothed whales like
porpoises Porpoises () are small dolphin-like cetaceans classified under the family Phocoenidae. Although similar in appearance to dolphins, they are more closely related to narwhals and belugas than to the true dolphins. There are eight extant speci ...
and dwarf spermwhales, which acquired this ability convergently. The PCA also suggests that some early members of ''Platanistoidea'' fall into the same morphospace, likely representing yet another example of narrow-band high frequency hearing arising independently among toothed whales. Notably however, the shape of the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
differs between these early platanistoids and ''Romaleodelphis'', which may be related to them filling different niches in their respective
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s. Whatever the case, the hearing apparatus of ''Romaleodelphis'' provides additional evidence for the early and repeated evolution of such hearing, which is seen in Early Oligoccene platanistoids from New Zealand, ''Romaleodelphis'' from Early Miocene Austria and the Middle Miocene eurhinodelphinid ''Xiphiacetus'' (also from Europe). The function of narrow-band high frequency hearing is uncertain. One hypothesis has proposed that such hearing is advantageous when trying to avoid large predatory whales like
macroraptorial sperm whale Macroraptorial sperm whales were highly predatory whales of the sperm whale superfamily (Physeteroidea) of the Miocene epoch (geology), epoch that hunted large marine mammals, including other whales, using their large teeth. They consist of six ge ...
s and
orcas The orca (''Orcinus orca''), or killer whale, is a toothed whale and the largest member of the oceanic dolphin family. The only extant species in the genus '' Orcinus'', it is recognizable by its black-and-white-patterned body. A cosmopoli ...
, whereas other research has suggested that this form of hearing is simply an early stage in the evolution of echolocation. Whatever the case, the anatomy of the semicircular canals in ''Romaleodelphis'' suggests that the hearing was more sensitive to head movement than in ''Xiphiacetus'' and porpoises, as inferred based on comparison with certain baleen whales and the bottlenose dolphin.


Paleoenvironment

''Romaleodelphis'' was discovered in the sediments of the Ebelsberg Formation near the Upper Austrian town of Pucking. This
Lagerstätte A Fossil-Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that preserves an exceptionally high amount of palaeontological information. ''Konzentrat-Lagerstätten'' preserv ...
is known to represent a marine
neritic The neritic zone (or sublittoral zone) is the relatively shallow part of the ocean above the drop-off of the continental shelf, approximately in depth. From the point of view of marine biology it forms a relatively stable and well-illuminated ...
environment of the Central
Paratethys The Paratethys sea, Paratethys ocean, Paratethys realm or just Paratethys (meaning "beside Tethys"), was a large shallow inland sea that covered much of mainland Europe and parts of western Asia during the middle to late Cenozoic, from the lat ...
, which once covered much of Central Europe. In addition to the clear signs of a marine environment, not limited to the present fauna which includes the sunfish '' Austromola'', various sharks, at least one other dolphin and many marine fish, the Ebelsberg Formation also preserves evidence of some freshwater influence through nearby rivers that emptied into the Paratethys. Since narrow-band high frequency hearing is not associated with any biome in particular (dwarf sperm whales are
pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
animals that inhabit the open oceans, porpoises stay close to the coast and the
La Plata dolphin The La Plata dolphin, franciscana or toninha (''Pontoporia blainvillei'') is a species of river dolphin found in coastal Atlantic waters of southeastern South America. It is a member of the Inioidea group and the only one that lives in the ocean ...
even swims up estuaries), Sanchez-Posada and colleagues speculate that it is possible that ''Romaleodelphis'' could have theoretically been capable of traveling between fresh-, salt- and brackish water.


References

{{Taxonbar, from=Q130969797 Miocene cetaceans Miocene mammals of Europe Fossils of Austria Fossil taxa described in 2024