Showing posts with label River Dolphin. Show all posts
Showing posts with label River Dolphin. Show all posts

Thursday, April 19, 2018

[PaleoMammalogy • 2018] Macrosqualodelphis ukupachai • A New Large Squalodelphinid (Cetacea, Odontoceti) from Peru Sheds Light on the Early Miocene Platanistoid Disparity and Ecology


Macrosqualodelphis ukupachai  
Bianucci, Bosio, Malinverno, de Muizon, Villa, Urbina & Lambert, 2018

   DOI: 10.1098/rsos.172302 

Abstract
The South Asian river dolphin (Platanista gangetica) is the only extant survivor of the large clade Platanistoidea, having a well-diversified fossil record from the Late Oligocene to the Middle Miocene. Based on a partial skeleton collected from the Chilcatay Formation (Chilcatay Fm; southern coast of Peru), we report here a new squalodelphinid genus and species, Macrosqualodelphis ukupachai. A volcanic ash layer, sampled near the fossil, yielded the 40Ar/39Ar age of 18.78 ± 0.08 Ma (Burdigalian, Early Miocene). The phylogenetic analysis places Macrosqualodelphis as the earliest branching squalodelphinid. Combined with several cranial and dental features, the large body size (estimated body length of 3.5 m) of this odontocete suggests that it consumed larger prey than the other members of its family. Together with Huaridelphis raimondii and Notocetus vanbenedeni, both also found in the Chilcatay Fm, this new squalodelphinid further demonstrates the peculiar local diversity of the family along the southeastern Pacific coast, possibly related to their partition into different dietary niches. At a wider geographical scale, the morphological and ecological diversity of squalodelphinids confirms the major role played by platanistoids during the Early Miocene radiation of crown odontocetes.

KEYWORDS: Odontoceti, Squalodelphinidae, Early Miocene, Peru, phylogeny, palaeoecology


Systematic palaeontology
Cetacea Brisson, 1762
Neoceti Fordyce and Muizon, 2001

Odontoceti Flower, 1867
Platanistoidea Gray, 1863
Squalodelphinidae Dal Piaz, 1917

Type genus. Squalodelphis Dal Piaz, 1917

Other genera included. Huaridelphis, Medocinia, Notocetus, Phocageneus.


Macrosqualodelphis, gen. nov.

Etymology. From ‘Macro’, large, and ‘Squalodelphis’ the type genus of the family. Gender masculine.


Macrosqualodelphis ukupachai, sp. nov.

Holotype and only referred specimen. MUSM 2545 consists of a skull lacking the anterior portion of the rostrum, the ear bones, both mandibles and the hyoid bones. The ventralmost portion of the rostrum and of the basicranium is worn along a plane slightly anterodorsally sloping with respect to the horizontal plane of the skull (erupted portion of maxillary teeth, basioccipital crests, ventral part of exoccipitals and postglenoid processes of squamosals missing). MUSM 2545 also preserves three detached anterior teeth; the atlas, two thoracic, two lumbar and eight caudal vertebrae; the left humerus, radius and incomplete ulna; one phalanx and one metacarpal; and two small fragments of ribs.

Type locality. About 3 km south of the fossiliferous Cerro Colorado locality, Western Ica Valley, Ica Region, southern Peru. 710 m above sea level. The holotype was discovered and collected by one of the authors (M.U.).

Etymology. From ‘Uku Pacha’ (Uku = withininsidePacha = Earth), the Inca lower world, located below the Earth's surface, in reference to the discovery of the specimen buried in sediment.


Figure 16. Skeletal remains and inferred body outline of the squalodelphinids from the early Burdigalian of the Chilcatay Fm (Pisco Basin, Peru) and skeletal and body outline of the extant P. gangetica. Body lengths based on the Pyenson & Sponberg [2011] equation for the fossils and on Jefferson et al. [2008] for the extant P. gangetica.


Cranium of the holotype (MUSM 2545) of Macrosqualodelphis ukupachai, from the early Burdigalian of the Chilcatay Fm (Pisco Basin, Peru).

Figure 3. (a) Dorsal view; (b) corresponding explanatory line drawing;   Linear hatching indicates major breaks and cross-hatching areas covered by the sediment.
Figure 4. (a) Ventral view; (b) corresponding explanatory line drawing;  Linear hatching indicates major breaks, cross-hatching areas covered by the sediment and dark shading worn surface.
Figure 5. (a) Right lateral view; (b) corresponding explanatory line drawing; (c) left lateral view. Cross-hatching indicates supporting frame.

Conclusion: 
Macrosqualodelphis ukupachai is a new species of the extinct platanistoid family Squalodelphinidae based on a well-preserved partial skeleton collected from the Early Miocene (ca 19–18 Ma) fossiliferous beds of the Chilcatay Fm outcropping in the Western Ica Valley (southern coast of Peru). The age of this skeleton is further constrained via 40Ar/39Ar dating of a local volcanic ash layer to 18.78 ± 0.08 Ma (early Burdigalian).

Our phylogenetic analysis supports the referral of M. ukupachai to the monophyletic family Squalodelphinidae, of which it constitutes the earliest diverging lineage.

The main distinctive character of M. ukupachai is its large size: its estimated TBL is approximately 3.5 m, significantly larger than all other known squalodelphinids, including N. vanbenedeni (2.5 m) and H. raimondii (2.0 m), both also found in the Chilcatay Fm. Combined with cranial and dental features (robust rostrum less tapered than in other squalodelphinids, large temporal fossa, prominent nuchal and temporal crests, and more robust teeth), the large body size of M. ukupachai suggests that this squalodelphinid was able to prey upon larger prey items. Consequently, M. ukupachai would have been positioned higher along the local trophic chain than the roughly contemporaneous N. vanbenedeni and H. raimondii. Therefore, it is suggested that the squalodelphinid diversity, both locally and worldwide, could be related to their partition into different dietary niches, as is observed in the extant delphinids.

This new record further illustrates the first, Early Miocene, broad radiation of crown odontocetes in marine environments, with a major contribution of homodont platanistoids. This Early Miocene morphological and ecological diversification of platanistoids (including squalodelphinids) was followed by the radiation of delphinidans (porpoises, true dolphins and relatives) during the Middle–Late Miocene. The only extant survivor of the platanistoid ‘golden age’ is the endangered South Asian river dolphin P. gangetica, confined in freshwater ecosystems of the Ganges, Indus and Brahmaputra river basins.


Giovanni Bianucci, Giulia Bosio, Elisa Malinverno, Christian de Muizon, Igor M. Villa, Mario Urbina and Olivier Lambert. 2018. A New Large Squalodelphinid (Cetacea, Odontoceti) from Peru Sheds Light on the Early Miocene Platanistoid Disparity and Ecology. Royal Society Open Science. 5(4)  DOI: 10.1098/rsos.172302

Monday, December 25, 2017

[PaleoMammalogy • 2017] Urkudelphis chawpipacha • A New Tropical Oligocene Dolphin from Montañita/Olón, Santa Elena, Ecuador


Urkudelphis chawpipacha
 Tanaka, Abella, Aguirre-Fernández, Gregori & Fordyce, 2017  


Abstract
A new small probable Oligocene dolphin from Ecuador represents a new genus and speciesUrkudelphis chawpipacha. The new taxon is known from a single juvenile skull and earbones; it differs from other archaic dolphins in features including widely exposed frontals at the vertex, a dorsally wide open vomer at the mesorostral groove, and a strongly projected and pointed lateral tuberosity of the periotic. Phylogenetic analysis places it toward the base of the largely-extinct clade Platanistoidea. The fossil is one of a few records of tropical fossil dolphins.


Systematic paleontology
CETACEA Brisson, 1762
NEOCETI Fordyce & de Muizon, 2001

ODONTOCETI Flower, 1867
PLATANISTOIDEA Gray, 1863 sensu Fordyce, 1994

Comment: Urkudelphis chawpipacha shows these synapomorphies of the Platanistoidea (sensu Fordyce, 1994), as recognised previously by Tanaka and Fordyce [2017]: periotic with C-shaped parabullary sulcus; small articular rim, which forms a ridge anterolateral to posterior process of periotic and separated from it by a sulcus. Two phylogenies place Urkudelphis near the base of the Platanistoidea (sensu lato; including Platanistidae, Squalodelphis, Notocetus, Phocageneus, Otekaikea, Waipatiidae, Awamokoa and Squalodontidae [2017]).

Fig 2. Skull, Urkudelphis chawpipacha MO-1 (holotype) in dorsal view. Left, photo, right, line art.

Fig 4. Skull, Urkudelphis chawpipacha MO-1 (holotype) in right lateral view. Upper, photo, lower, line art.

Urkudelphis gen. nov.

Type species: Urkudelphis chawpipacha sp. nov.

Urkudelphis chawpipacha sp. nov.

Diagnosis: Urkudelphis chawpipacha is a small archaic odontocete with the following autapomorphic combination of characters: shallow antorbital notch (character 10); anteromedially oriented anterior edge of the supraorbital process (character 37); weakly dorsally convex nuchal crest in dorsoposterior view (character 119); approximately same sized apertures of the vestibular aqueduct and cochlear aqueduct (character 186); dorsoventrally thin pars cochlearis on the periotic (character 192); inner posterior prominence of the tympanic bulla is anterior to the outer posterior prominence (character 218); very strongly projecting and pointed lateral tuberosity; and an anteroposteriorly long accessory ossicle of the periotic. Urkudelphis chawpipacha differs from early branching odontocetes, including Agorophius, Ashleycetus, Simocetus, Mirocetus and Xenorophus in having the frontals on the vertex at a level behind the postorbital process; anteroposteriorly shorter and transversely wider frontals (approaching a square-shape rather than narrow and elongate); and parallel-sided posterior part of the ascending process of each maxilla forming a narrow elongate face. Urkudelphis differs from Early Miocene Papahu taitapu, Chilcacetus cavirhinus, Arktocara yakataga, Allodelphis pratti and Ninjadelphis ujiharai, having the frontals on the vertex flat and longer than the taxa above, which have more nodular and shorter frontals. Urkudelphis differs from Chilcacetus and Papahu in having a narrow premaxillary sac fossa. Urkudelphis chawpipacha also notably shows: frontals at the vertex invaded posteriorly by the interparietal; and long anteromedial projection of the palatine on the palate. Other diagnostic features of U. chawpipacha are shared with more-crownward Waipatiidae: a shallow suprameatal pit of the squamosal (character 152); an abruptly ventrally deflected anterior process of the periotic (character 172); and a nearly flat dorsal surface of the periotic in lateral view (character 181). In addition, Urkudelphis chawpipacha shares several characters with more-crownward Platanistoidea: a periotic with C-shaped parabullary sulcus (character 175); and a small articular rim, which forms a ridge anterolateral to the posterior process of the periotic and separated from it by a sulcus (character 195).

Holotype: MO-1, an incomplete skull (premaxilla, maxilla, vomer, pterygoid, frontal, parietal, interparietal, alisphenoid, squamosal and supraoccipital), including the right periotic, right tympanic bulla and right malleus.

Etymology: The generic name, Urkudelphis originates from Kichwa “urku” meaning mountain, referring to the type locality of Montañita, and Greek “delphis” for dolphin, which has been used widely as a suffix for dolphin generic names. Chawpipacha results from the combination of chawpi, meaning "half" or “middle” and pacha, meaning "the world" representing the equator, and thus Ecuador in Kichwa.

Type locality: MO-1 was found by one of the authors (JA) and several UPSE students in August 2015 in a boulder that collapsed from a cliff at the coastal locality here named Montañita/Olón (latitude 1°48'50.64" S, longitude 80°45'24.18" W). The Montañita/Olón (MO) locality (Fig 1) lies midway between the towns of Montañita and Olón (Santa Elena Province, Ecuador) and can only be accessed during low tides.
Conclusion: 
A new small dolphin from probable Oligocene (Chattian?) strata in Santa Elena, Ecuador is described as a new species and genus, Urkudelphis chawpipacha. The new taxon is characterized by: an anteromedially oriented anterior edge of the supraorbital process; weakly convex nuchal crest in dorsoposterior view; approximately same sized apertures of vestibular aqueduct and cochlear aqueduct; dorsoventrally thin pars cochlearis on periotic; and inner posterior prominence placed anterior to the outer posterior prominence. Urkudelphis chawpipacha differs from other Oligocene dolphins in the combination of: frontals on the vertex at a level posterior to the postorbital process; shorter and wider frontals; and parallel-sided posterior part of the ascending process of the maxilla. Phylogenetic analysis places it near the base of the largely-extinct clade Platanistoidea. The fossil is one of few fossil Neoceti reported from the equator, and is a reminder that Oligocene cetaceans may have ranged widely in tropical waters.


Yoshihiro Tanaka, Juan Abella, Gabriel Aguirre-Fernández, Maria Gregori and R. Ewan Fordyce. 2017. A New Tropical Oligocene Dolphin from Montañita/Olón, Santa Elena, Ecuador. PLoS ONE. 12(12); e0188380.  DOI: 10.1371/journal.pone.0188380

New ancient dolphin species Urkudelphis chawpipacha discovered in Ecuador http://phy.so/432980137 via @physorg_com


Wednesday, June 7, 2017

[PaleoMammalogy • 2017] Dilophodelphis fordycei • A New Fossil Dolphin Provides Insight Into the Evolution of Supraorbital Crests in Platanistoidea (Mammalia, Cetacea)


Dilophodelphis fordycei
Boersma, McCurry & Pyenson, 2017 

  DOI: 10.1098/rsos.170022 
Image: A. Boersma  @Boersma_Alex 

Abstract

Many odontocete groups have developed enlarged facial crests, although these crests differ in topography, composition and function. The most elaborate crests occur in the South Asian river dolphin (Platanista gangetica), in which they rise dorsally as delicate, pneumatized wings anterior of the facial bones. Their position wrapping around the melon suggests their involvement in sound propagation for echolocation. To better understand the origin of crests in this lineage, we examined facial crests among fossil and living Platanistoidea, including a new taxonDilophodelphis fordyceinov. gen. and sp., described herein, from the Early Miocene Astoria Formation of Oregon, USA. We measured the physical extent and thickness of platanistoid crests, categorized their relative position and used computed tomography scans to examine their internal morphology and relative bone density. Integrating these traits in a phylogenetic context, we determined that the onset of crest elaboration or enlargement and the evolution of crest pneumatization among the platanistoids were separate events, with crest enlargement beginning in the Oligocene. However, we find no evidence for pneumatization until possibly the Early Miocene, although certainly by the Middle Miocene. Such an evolutionary context, including data from the fossil record, should inform modelling efforts that seek to understand the diversity of sound generation morphology in Odontoceti.

KEYWORDS: cetacean, Platanistoidea, river dolphins, Miocene, pneumatization, computed tomography

Systematic palaeontology

Cetacea Brisson, 1762 
Odontoceti Flower, 1867 sensu Fordyce and Muizon, 2001 

Platanistoidea sensu Boersma and Pyenson 2016 
Platanistidae Gray, 1846  sensu Boersma and Pyenson 2016 

Dilophodelphis, gen. nov. 

Type and only included species. Dilophodelphis fordycei, sp. nov.

Etymology. From the Greek words di (double), lophos (crest) and delphis (dolphin), referring to the enlarged supraorbital crests on the dorsal surface of the skull, resembling twin mountain crests. This construction also evokes the dinosaur Dilophosaurus wetherilli Welles 1954, a double-crested theropod recovered from Early Jurassic sequences of the Kayenta Formation in Arizona, USA.

Dilophodelphis fordycei, sp. nov. 

Figure 1. Skull of Dilophodelphis (USNM 214911) in dorsal view.
(a) Illustrated skull with low opacity mask, interpretive line art and labels for skull elements. Dotted lines indicate uncertainty of sutures, and dashed lines highlight fossae. Hatched pattern indicates areas where sediment is obscuring the fossil. (b) Photograph of skull in dorsal view, photography by James Di Loreto, Smithsonian Institution. fr., frontals; max., maxilla; n., nasal; pmx., premaxilla; pmx. sac fossa, premaxillary sac fossa. 

Figure 2. Skull of Dilophodelphis (USNM 214911) in ventral view.
(a) Illustrated skull with low opacity mask, interpretive line art and labels for skull elements. Dotted lines indicate uncertainty of sutures, and dashed lines highlight fossae. Hatched pattern indicates areas where sediment is obscuring the fossil. (b) Photograph of skull in ventral view, photography by James Di Loreto, Smithsonian Institution. fr., frontal; max., maxilla; p., pterygoid; v., vomer. 

The endangered South Asian river dolphin, Platanista gangetica (middle), swimming alongside two of its fossil relatives: the longirostral Pomatodelphis inaequalis (bottom) and the new species Dilophodelphis fordycei (top). None of the species lived together at the same time. Glow-throughs to the skulls highlight the diversity in supraorbital crest shape and size among the members of this family.
Image: Alex Boersma  @Boersma_Alex 


Figure 3. Skull of Dilophodelphis (USNM 214911) in right and left lateral views.
 (a) Illustrated skull in right lateral view and (b) left lateral view with low opacity mask, interpretive line art and labels for skull elements. Dotted lines indicate uncertainty of sutures, and dashed lines highlight fossae. Hatched pattern indicates areas where sediment is obscuring the fossil. (c) Photograph of skull in right lateral view and (d) left lateral view, photography by James Di Loreto, Smithsonian Institution. exocc., exoccipital; m., maxilla; par., parietal; p., pterygoid; sq., squamosal; temp. fossa, temporal fossa; zyg. process, zygomatic process. 


Etymology: The species epithet honours Prof. R. Ewan Fordyce, FRSNZ, native New Zealander and prominent vertebrate palaeontologist. The epithet recognizes his extensive and long-lasting contributions to the field of marine mammal palaeontology, including his commitment to mentoring future scientists, especially in shaping the career paths of the authors herein. The epithet also honours his long-standing interest in the fossil marine mammal record of Oregon, which has yielded pivotal specimens for over a century, including Simocetus rayi Fordyce 2002, which he described.



Alexandra T. Boersma, Matthew R. McCurry and Nicholas D. Pyenson. 2017. A New Fossil Dolphin Dilophodelphis fordycei Provides Insight Into the Evolution of Supraorbital Crests in Platanistoidea (Mammalia, Cetacea). Royal Society Open Science.  DOI: 10.1098/rsos.170022


New, ancient dolphin species had weird skull crests australiangeographic.com.au/news/2017/06/new,-ancient-dolphin-species-had-weird-skull-crests   @ausgeo - Australian Geographic 



Sunday, August 28, 2016

[PaleoMammalogy • 2016] Arktocara yakataga • A New Fossil Odontocete (Mammalia, Cetacea) from the Oligocene of Alaska and the Antiquity of Platanistoidea


Arktocara yakataga 
Boersma & Pyenson, 2016

Artistic reconstruction of a pod of Arktocara yakataga, swimming offshore of Alaska during the Oligocene, about 25 million years ago, with early mountains of Southeast Alaska in the background. The authors speculate that Arktocara may have socialized in pods, like today's oceanic dolphins, while possessing a much longer snout, like its closest living relative in the freshwater rivers of South Asia.
Linocut print art by Alexandra Boersma

Abstract

The diversification of crown cetacean lineages (i.e., crown Odontoceti and crown Mysticeti) occurred throughout the Oligocene, but it remains an ongoing challenge to resolve the phylogenetic pattern of their origins, especially with respect to stem lineages. One extant monotypic lineage, Platanista gangetica (the Ganges and Indus river dolphin), is the sole surviving member of the broader group Platanistoidea, with many fossil relatives that range from Oligocene to Miocene in age. Curiously, the highly threatened Platanista is restricted today to freshwater river systems of South Asia, yet nearly all fossil platanistoids are known globally from marine rocks, suggesting a marine ancestry for this group. In recent years, studies on the phylogenetic relationships in Platanistoidea have reached a general consensus about the membership of different sub-clades and putative extinct groups, although the position of some platanistoid groups (e.g., Waipatiidae) has been contested. Here we describe a new genus and species of fossil platanistoid, Arktocara yakataga, gen. et sp. nov. from the Oligocene of Alaska, USA. The type and only known specimen was collected from the marine Poul Creek Formation, a unit known to include Oligocene strata, exposed in the Yakutat City and Borough of Southeast Alaska. In our phylogenetic analysis of stem and node-based Platanistoidea, Arktocara falls within the node-based sub-clade Allodelphinidae as the sister taxon to Allodelphis pratti. With a geochronologic age between ∼29–24 million years old, Arktocara is among the oldest crown Odontoceti, reinforcing the long-standing view that the diversification for crown lineages must have occurred no later than the early Oligocene.

Systematic paleontology

Cetacea Brisson, 1762
Odontoceti Flower, 1867 sensu Fordyce & Muizon, 2001
Platanistoidea (CCN) (node-based version of Fordyce, 1994)
Allodelphinidae (CCN) (node-based version of Barnes, 2006)

Arktocara, gen. nov. 
LSID: urn:lsid:zoobank.org:act:EE11B95B-8338-496B-97F4-1673ED90E709

The skull of Arktocara yakataga on an 1875 ethnographic map of Alaska drawn by William Healey Dall, a broadly trained naturalist who worked for several US government agencies, including the Smithsonian, and honored with several species of living mammals, including Dall's porpoise (Phocoenoides dalli). Near the skull of Arktocara is a cetacean tooth, likely belonging to a killer whale (Orcinus orca), collected by Aleš Hrdlička, a Smithsonian anthropologist who worked extensively in Alaska, and an Oligocene whale tooth collected by Donald Miller, a geologist who worked for the US Geological Survey, and collected the type specimen of Arktocara. Donald Orth's dictionary of Alaskan place names, published by the USGS, bookends the image.
photo: James Di Loreto, Smithsonian 


Definitions. Crown group Platanista refers to the crown clade arising from the last common ancestor of all lineages descending from Platanista, including two subspecies of Platanista gangetica (P. g. gangetica (Lebeck, 1801) and P. g. minor Owen, 1853), as recognized by The Society for Marine Mammology’ Committee on Taxonomy (2015).

Type and only included species: Arktocara yakataga, sp. nov.

Etymology. The name Arktocara derives from the combination of arktos from Greek and cara from Latin, which together signify “the face of the North.” The only preserved material of the type specimen, USNM 214830 consists of the cranium, or its face, and its type locality is the furthest north that a platanistoid has ever been found.

Age. Same as that of the species.
Diagnosis. Same as that of the species.

Arktocara yakataga, sp. nov. (Figs. 2–10 and Table 1)

The skull of Akrtocara yakataga rests on an 1875 ethnographic map of Alaska drawn by William Healey Dall, a broadly trained naturalist who worked for several US government agencies, including the Smithsonian, and honored with several species of living mammals, including Dall's porpoise (Phocoenoides dalli). Near the skull of Arktocara is a cetacean tooth, likely belonging to a killer whale (Orcinus orca), collected by Aleš Hrdlička, a Smithsonian anthropologist who worked extensively in Alaska, and an Oligocene whale tooth collected by Donald Miller, a geologist who worked for the US Geological Survey, and collected the type specimen of Arktocara. Donald Orth's dictionary of Alaskan place names, published by the USGS, bookends the image.
photo: James Di Loreto, Smithsonian  

Holotype. USNM 214830, consisting of an incomplete skull lacking the rostrum anterior of the antorbital notches, tympanoperiotics, dentition and mandibles (see Fig. 2).

Type locality. The precise geographic coordinates for the type locality of Arktocara yakataga are unknown. The type specimen (USNM 214830) was discovered and collected in 1951 by United States Geological Survey (USGS) geologist Donald J. Miller (1919–1961), who was mapping what was then the Yakataga District of Alaska (now the Yakutat City and Borough) between 1944 and 1963. Archival notes housed with the specimen at USNM state that Miller found the specimen in the Poul Creek Formation within the then-Yakataga District (see Age, below). Therefore, we delimit the area for the type’s provenance to exposures of the Poul Creek Formation in the Yakutat City and Borough, Alaska, USA, in a grid ranging approximately from 60°22′N, 142°30′W to 60°00′N, 143°22′W (see Fig. 1). While the formation has been named from its exposures along Poul Creek, it has been suggested that the most abundant macrofossils from this unit have been collected from outcrops along Hamilton Creek, White River, and Big River near Reare Glacier (Taliaferro, 1932). It is possible that Miller collected USNM 214830 from one of these exposures.

Formation. Poul Creek Formation.

Age. Archival documentation accessioned in the Department of Paleobiology with USNM 214830 indicate that the type specimen was collected from an unknown locality exposed about 400–500 m below the top of the Poul Creek Formation, which has a total stratigraphic thickness of around 1.9 km (Plafker, 1987). The Yakutat terrane of Southeast Alaska consists of the Kulthieth, Poul Creek, and Yakataga Formations (Perry, Garver & Ridgway, 2009; Plafker, Moore & Winkler, 1994; Miller, 1971). The Kulthieth Formation consists of mostly organic-rich sandstones deposited in nonmarine alluvial, deltaic, barrier beach and shallow marine environments, and is Early Eocene to Early Oligocene (∼54–33 Ma) in age based on the fossil assemblages present (Perry, Garver & Ridgway, 2009). The Upper Eocene to possibly Lower Miocene (∼40–20 Ma) Poul Creek Formation conformably overlies the Kulthieth Formation (Plafker, 1987; Miller, 1971). It is estimated to be approximately 1.9 km thick, and is composed of siltstones and organic-rich sandstones, in part glauconitic recording a marine transgression, interrupted by deposits of the Cenotaph Volcanics (Plafker, 1987). Finally, unconformably overlying the Poul Creek Formation is the Miocene to Pliocene Yakataga Formation (Miller, 1971). It is composed mainly of tillite and marine strata (Perry, Garver & Ridgway, 2009).

The Poul Creek Formation itself is broadly constrained to approximately 40–20 million years in age, from the latest Eocene to possibly early Miocene in age (Plafker, 1987; Miller, 1971). The depositional age of the unit has been further constrained to ∼24 to ∼29 Ma, or a mid to late Oligocene age, based on detrital zircon fission-track analyses of young grain-age populations (Perry, Garver & Ridgway, 2009). Using the broadest time duration for the formation (∼20 million years) and the coarse stratigraphic thickness of the sediments within it (∼2 km), a constant rate of sedimentation would suggest that the stratigraphic position of USNM 214830 at 500 m below the top of the formation would be roughly equivalent to an geochronologic age of ∼25 million years, an estimate that is consistent to the detrital zircon analyses. Overall, we propose a late Oligocene, or Chattian age for Arktocara, although we cannot exclude a Rupelian antiquity.

Diagnosis. Arktocara is a small to medium sized platanistoid odontocete (approximately 2.26 m in total length), which belongs, based on one equivocal synapomorphy, to the node-based Platanistoidea: width: width of the premaxillae >50% of the width of the rostrum at the antorbital notch (character 51[1]). More convincingly, Arktocara belongs to Platanistoidea based on its affinities to other members of the Allodelphinidae that possess unequivocal synapomorphies of the Platanistoidea (see ‘Discussion’ for further comments on the relationship of Allodelphinidae within the Platanistoidea). We also note that, for the purposes of this diagnosis, we use a broad definition of Waipatiidae that includes Otekaikea spp. (see Tanaka & Fordyce (2015a)), and Squalodelphinidae sensu Lambert, Bianucci & Urbina (2014). See ‘Discussion’ for further comments on systematics of these groups.

.........


Etymology. The species epithet ‘yakataga’ derives from the Tlingit name for the point of land along the southeast coast of Alaska between modern day Kayak Island and Ice Bay. This point, currently called Cape Yakataga, is located directly southwest of Watson Peak and represents the southeastern boundary of a floodplain drained by the Bering Glacier. The name Yakataga was first published by Tebenkov (1852: map 7), who was a cartographer and hydrographer of the Imperial Russian Navy, as “M[ys] Yaktaga” on an 1849 map of Alaska. The geographic place name has been alternatively spelled Cape Iaktag, Cape Yakaio, Cape Yakatag, and Yokataga Reef (Orth, 1967). According to the Geographic Names Information System (GNIS, 2016), developed by USGS in cooperation with the United States Board of Geographic Names (BGN), the name “Yakataga” means “canoe road,” referring to two reefs that form a canoe passage to the shore of the village.

Figure 12: Distribution map of fossil Allodelphinidae.
Mapped of fossil localities of allodelphinids, projected on a truncated Winkel Tripel map and centered on 25°N and 170°W. Occurrences for fossil data derive from location of type and referred localities for each taxon, are listed alphabetically by region, and are represented by orange dots.

Platanistoids first appear in the fossil record in the late Oligocene, and reach peak richness in the early Miocene (Kimura & Barnes, 2016; Tanaka & Fordyce, 2015a). The oldest platanistoids with solid age constraints are the waipatiids, all found in the Oligocene-Miocene Otekaike Limestone of New Zealand (Graham et al., 2000; Benham, 1935; Fordyce, 1994; Tanaka & Fordyce, 2014; Tanaka & Fordyce, 2015a). Based on both the lithology and the presence of age-diagnostic planktic foraminifera and ostracod species, Waipatia hectori (Benham, 1935) is the oldest reported waipatiid, from the uppermost Duntroonian Stage of the Otekaike Limestone, approximately 25.2 Ma (Tanaka & Fordyce, 2015b). Arktocara is possibly very similar in age to Waipatia hectori, constrained to the Chattian Stage of the upper Oligocene in the Poul Creek Formation, approximately ∼24–29 Ma (Perry, Garver & Ridgway, 2009). Unfortunately, the lack of robust locality data for either Waipatia hectori or Arktocara makes impossible to determine which is the oldest.

Arktocara is, however, very clearly the oldest known allodelphinid, expanding the previously reported age range of Allodelphinidae by as much as 9 million years (Kimura & Barnes, 2016). Other allodelphinids span temporally from the early to middle Miocene, which largely matches the stratigraphic range of other platanistoid lineages (Fig. 11). Interestingly, Arktocara is among the oldest crown Odontoceti, reinforcing the long-standing view that the timing for the diversification for crown lineages must have occurred no later than the early Oligocene.

Lastly, Allodelphinidae appear uniquely limited, in terms of geography, to marine rocks of the North Pacific Ocean, with occurrences in Japan, Alaska, Washington State, Oregon, and California (see Fig. 12; Kimura & Barnes, 2016). Arktocara expands this geographic range to sub-Arctic latitudes. At approximately 60°N in the Yakutat City and Borough, Arktocara is the most northern platanistoid yet reported. The next most northern platanistoid reported is an incomplete and unnamed specimen from the late Chattian marine Vejle Fjord Formation in northern Denmark, approximately 56.7°N, 9.0°E (Hoch, 2000).



Alexandra T. Boersma​ and Nicholas D. Pyenson. 2016. Arktocara yakataga, a new fossil odontocete (Mammalia, Cetacea) from the Oligocene of Alaska and the antiquity of Platanistoidea.  PeerJ. 4:e2321. DOI: 10.7717/peerj.2321

New species of extinct river dolphin discovered in Smithsonian collection bit.ly/2aGgRS8 via @EurekAlertAAAS

Wednesday, September 2, 2015

[PaleoMammalogy • 2015] Isthminia panamensis • A New Fossil inioid (Mammalia, Cetacea) from the Chagres Formation of Panama and the Evolution of ‘River Dolphins’ in the Americas


Isthminia panamensis
Pyenson, Vélez-Juarbe, Gutstein, Little, Vigil & O’Dea, 2015
Life reconstruction of Isthminia panamensis, feeding on a flatfish, which would have been abundant in the neritic zone of the late Miocene equatorial seas of Panama. 
DOI: 10.7717/peerj.1227

Abstract

In contrast to dominant mode of ecological transition in the evolution of marine mammals, different lineages of toothed whales (Odontoceti) have repeatedly invaded freshwater ecosystems during the Cenozoic era. The so-called ‘river dolphins’ are now recognized as independent lineages that converged on similar morphological specializations (e.g., longirostry). In South America, the two endemic ‘river dolphin’ lineages form a clade (Inioidea), with closely related fossil inioids from marine rock units in the South Pacific and North Atlantic oceans. Here we describe a new genus and species of fossil inioid, Isthminia panamensis, gen. et sp. nov. from the late Miocene of Panama. The type and only known specimen consists of a partial skull, mandibles, isolated teeth, a right scapula, and carpal elements recovered from the Piña Facies of the Chagres Formation, along the Caribbean coast of Panama. Sedimentological and associated fauna from the Piña Facies point to fully marine conditions with high planktonic productivity about 6.1–5.8 million years ago (Messinian), pre-dating the final closure of the Isthmus of Panama. Along with ecomorphological data, we propose that Isthminia was primarily a marine inhabitant, similar to modern oceanic delphinoids. Phylogenetic analysis of fossil and living inioids, including new codings for Ischyrorhynchus, an enigmatic taxon from the late Miocene of Argentina, places Isthminia as the sister taxon to Inia, in a broader clade that includes Ischyrorhynchus and Meherrinia, a North American fossil inioid. This phylogenetic hypothesis complicates the possible scenarios for the freshwater invasion of the Amazon River system by stem relatives of Inia, but it remains consistent with a broader marine ancestry for Inioidea. Based on the fossil record of this group, along with Isthminia, we propose that a marine ancestor of Inia invaded Amazonia during late Miocene eustatic sea-level highs.

Keywords: Ecology, Evolutionary Studies, Marine Biology, Paleontology, Zoology


Figure 1: Map of fossil and living Inioidea.
Global map of living and fossil inioids, projected onto an orthographic globe, centered on 15°N, 45°W. Extant distributions of Inia geoffrensis (teal and black waterways) and Pontoporia blainvillei (dark gray), follow data from the IUCN (International Union for Conservation of Nature) (2013) and Secchi, Ott & Danilewicz (2003), respectively. Occurrences for fossil data derive from location of type localities for each taxon, except for reports for the Northern Europe by Pyenson & Hoch (2007), Western South America by Gutstein et al. (2015), and Amazonia and Eastern South America by Cozzuol (2010). Major fossil localites for enumerated inioids identified at least to the generic level, are listed alphabetically by region, and represented by teal or blue dots, for freshwater and marine deposits, respectively.
Base map generated by Indiemapper (http://indiemapper.com).
DOI: 10.7717/peerj.1227




Systematic paleontology

Cetacea Brisson, 1762
Odontoceti Flower, 1867
Delphinida Muizon, 1988a

Inioidea Gray, 1846 sensu Muizon, 1988a

Pan-Inia (NCN) (panstem-based version of Inia (Blainville, 1817))

Isthminia, gen. nov. 
LSID: urn:lsid:zoobank.org:act:83F6A9B4-289D-45DE-A3D1-C361DAAAF973.


Definitions ‘Pan-Inia’ refers to the panstem that includes crown Inia (CCN), and all other lineages closer to Inia than to Pontoporia, such as Isthminia and Ischyrorhynchus. Subjective synonymies of Pan-Inia include: Iniidae Gray, 1846; Iniinae Flower, 1867; Saurocetidae Ameghino, 1891; Iniidae Muizon, 1984; Ischyrorhynchinae (Cozzuol, 1996); Iniidae Cozzuol, 2010; Iniidae Gutstein, Cozzuol & Pyenson, 2014b. Crown group Inia refers to the crown clade arising from the last common ancestor of all named species of Inia, including Inia boliviensis d’Orbigny, 1834 and Inia araguaiaensis Hrbek et al., 2014. Although we follow the suggestions of the Society for Marine Mammalogy’s Committee on Taxonomy (2014) in provisionally recognizing two sub-species of Inia geoffrensis (I. g. geoffrensis and I.g. humboldtiana Pilleri & Gihr, 1977), the phylogenetic definition of Inia can accommodate a plurality of species and subspecies.




Type and only known species. Isthminia panamensis, sp. nov.

Etymology. Isthm- reflects the type specimen’s provenance from the Isthmus of Panama and the crucial role that the formation of this isthmus played in Earth history and evolution of the biota of the Americas. This epithet follows in the tradition of another fossil cetacean from the Chagres Formation, Nanokogia isthmiaVelez-Juarbe et al., 2015. The feminine generic epithet Inia reflects its similarities to the living Amazon River dolphin (Inia geoffrensis). Pronunciation: ‘Ist-min-ee-a,’ with the emphasis on the second syllable.

Isthminia panamensis sp. nov.

Type locality. STRI locality 650009 (9°16′55.4880″N, 80°02′49.9200″W), less than 100 m northeast of the main road in the town of Piña, along the Caribbean Sea coastline of the Republic of Panama (Fig. 2).

Formation. Piña Facies of the Chagres Formation.

Etymology. The species epithet recognizes the Republic of Panama, its people, and the many generations of scientists who have studied its geological and biological histories.



Figure 14: Reconstruction of Isthminia.
Life reconstruction of Isthminia panamensis, feeding on a flatfish, which would have been abundant in the neritic zone of the late Miocene equatorial seas of Panama.
Art by Julia Molnar. DOI: 10.7717/peerj.1227

Figure 15: Stratigraphically calibrated phylogenetic tree of Inioidea.
Time calibrated phylogenetic tree of select Delphinida, pruned from our consensus cladogram in Fig. 13, including Isthminia, with Delphinoidea collapsed. Stratigraphic range data derives from published accounts for each taxon, including global ranges. Geologic time scale based on Cohen et al. (2013). Calibration for major nodes depths follow mean divergence date estimates by McGowen, Spaulding & Gatesy (2009: table 3) for the following clades: a, Delphinida (24.75 Ma); b, Inioidea + Lipotes (22.15 Ma); c, Delphinoidea (18.66 Ma); and Inioidea (in open white circle, 16.68 Ma). All minor node depths are graphical heuristics, and not intended to reflect actual divergence dates. Arc indicates stem-based clade, Pan-Inia. Ecological habitat preference is based on depositional environment or extant habitat.
Abbreviations: Aquitan., Aquitanian; H., Holocene; Langh., Langhian; Mess., Messinian; P., Piacenzian; Ple., Pleistocene; Plioc., Pliocene; Serra., Serravallian; Zan., Zanclean.  DOI: 10.7717/peerj.1227


Isthminia panamensis
Pyenson, Vélez-Juarbe, Gutstein, Little, Vigil & O’Dea, 2015
Life reconstruction of Isthminia panamensis, feeding on a flatfish, which would have been abundant in the neritic zone of the late Miocene equatorial seas of Panama. 



Nicholas D. Pyenson, Jorge Vélez-Juarbe, Carolina S. Gutstein, Holly Little, Dioselina Vigil and Aaron O’Dea. 2015. Isthminia panamensis, A New Fossil inioid (Mammalia, Cetacea) from the Chagres Formation of Panama and the Evolution of ‘River Dolphins’ in the Americas. PeerJ. 3:e1227 DOI: 10.7717/peerj.1227

Wednesday, September 10, 2014

[PaleoMammalogy • 2014] Huaridelphis raimondii • A New early Miocene Squalodelphinidae (Cetacea, Odontoceti) from the Chilcatay Formation, Peru


 skull and partial mandible of the new squalodelphinid species Huaridelphis raimondii, with an outline of the head and potential fish prey.
photo: G. Bianucci | DOI: 10.1080/02724634.2014.858050

ABSTRACT
The fossil record of odontocetes (toothed cetaceans) is relatively scarce during the Oligocene and early Miocene compared with later in the Miocene and Pliocene; most of the odontocete families from these epochs are known by a limited number of species and specimens. Among those, Squalodelphinidae is a family of small- to medium-sized platanistoids with single-rooted teeth, which until now has included only four genera based on diagnostic material, from the early Miocene of Europe, Argentina, and North America. Recent field work in the Pisco-Ica desert, southern coast of Peru, has resulted in the discovery of several marine vertebrate-rich localities in various levels of the late Oligocene–early Miocene Chilcatay Formation. Based on three specimens from Ullujaya and Zamaca, including two well-preserved skulls with periotics, we describe a new squalodelphinid genus and species, Huaridelphis raimondii. This new species increases the early Miocene diversity of the family and is also its smallest known member. It further differs from other squalodelphinids by its thin antorbital process of the frontal, abruptly tapering rostrum, and higher tooth count. A more fragmentary skull, from Zamaca, is referred to Squalodelphinidae aff. H. raimondii. This skull provides information on the morphology of the tympanic, malleus, and incus, currently unknown in H. raimondii. Focusing on platanistoids with single-rooted teeth, our phylogenetic analysis suggests that Squalodelphinidae are monophyletic and confirms the sister-group relationship between the latter and Platanistidae. The relationships within Squalodelphinidae are not fully resolved, but H. raimondii might be one of the earliest diverging taxa.


 the skull of the new squalodelphinid species Huaridelphis raimondii in dorsal and lateral view.
photo: G. Bianucci 

Etymology— From ‘Huari,’ ancient culture of the south-central Andes and coastal area of Peru (500–1000 AD), and from ‘delphis,’ the Latin word for dolphin. Gender masculine. The species name honors Antonio Raimondi (1826–1890), an Italian scientist who first documented fossil whales from Peru (Bianucci, 2010).


The examination of fossil cetacean remains in the locality of Ullujaya, Pisco Basin, Peru.
photo: G. Bianucci 



CONCLUSIONS
Based on three specimens, including two well-preserved skulls, from early Miocene localities of the Chilcatay Formation (Ullujaya and Zamaca), Pisco-Ica desert, southern coast of Peru, we describe a new genus and species of Squalodelphinidae, Huaridelphis raimondii. In addition to periotic characters, H. raimondii differs from other known squalodelphinids in, among others, its smaller size, the thin antorbital process of the frontal, the more abrupt tapering of the rostrum, and the higher tooth count. Another fragmentary skull from the Chilcatay Formation in Zamaca is referred to Squalodelphinidae aff. Huaridelphis raimondii. It brings additional information on the morphology of the tympanic, malleus, and incus, not yet known in H. raimondii. Our phylogenetic analysis of platanistoids with single-rooted teeth suggests that the family Squalodelphinidae is monophyletic; the analysis also confirms the sister-group relationship between the latter and Platanistidae. The relationships within Squalodelphinidae are not fully resolved, but H. raimondii might be one of the first diverging taxa of the family.


Olivier Lambert, Giovanni Bianucci & Mario Urbina. 2014. Huaridelphis raimondii, A New early Miocene Squalodelphinidae (Cetacea, Odontoceti) from the Chilcatay Formation, Peru
 Journal of Vertebrate Paleontology. 34(5); 987-1004.
DOI: 10.1080/02724634.2014.858050 

New species of extinct dolphin sheds light on river dolphin history

Friday, April 5, 2013

[Cetology • 2013] Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow, Acoustically Complex Habitat


A rare sight of the fast and shy
Ganges River Dolphin 
Platanista g. gangetica

(Photo by Rubaiyat Mansur, Whale and Dolphin Conservation Society)

Abstract
Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPapp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPapp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.

____________________

An Ancient Biosonar Sheds New Light On the Evolution of Echolocation in Toothed Whales

— Some 30 million years ago, Ganges river dolphins diverged from other toothed whales, making them one of the oldest species of aquatic mammals that use echolocation, or biosonar, to navigate and find food. This also makes them ideal subjects for scientists working to understand the evolution of echolocation among toothed whales.


New research, led by Frants Havmand Jensen, a Danish Council for Independent Research / Natural Sciences postdoctoral fellow at Woods Hole Oceanographic Institution, shows that freshwater dolphins produce echolocation signals at very low sound intensities compared to marine dolphins, and that Ganges river dolphins echolocate at surprisingly low sound frequencies. The study, "Clicking in shallow rivers," was published in the journal PLOS ONE.

"Ganges River dolphins are one of the most ancient evolutionary branches of toothed whales," says Jensen. "We believe our findings help explain the differences in echolocation between freshwater and marine dolphins. Our findings imply that the sound intensity and frequency of Ganges river dolphin may have been closer to the 'starting point' from which marine dolphins gradually evolved their high-frequent, powerful biosonar."

The scientists believe these differences evolved due to differences in freshwater and marine environments and the location and distribution of prey in those environments.

A complex, underwater environment
To sustain themselves, river dolphins must find their food, often small fish or crustaceans, in highly turbid water where visibility seldom exceeds a few inches. Like their marine relatives, they manage this using echolocation: They continuously emit sound pulses into the environment and listen for the faint echoes reflected off obstacles while paying special attention to the small details in the echoes that might signify a possible meal.
The environment that freshwater dolphins operate in poses very different challenges to a biosonar than the vast expanses of the sea where most dolphins later evolved. "Dolphins that range through the open ocean often feed on patchily distributed prey, such as schools of fish," Jensen says. "They have had a large advantage from evolving an intense biosonar that would help them detect prey over long distances, but we have little idea of how the complex river habitats of freshwater dolphins shape their biosonar signals."

Shy study animals with a surprisingly deep voice
To answer that question, the researchers recorded the echolocation signals of two species of toothed whales inhabiting the same mangrove forest in the southern part of Bangladesh: The Ganges river dolphin, an exclusively riverine species that is actually not part of the dolphin family but rather the Platanistidae family, and the Irrawaddy, a freshwater toothed whale from the dolphin family that lives in both coastal and riverine habitats.
Surprisingly, the echolocation signals turned out to be much less intense than those employed by marine dolphins of similar size and it seemed that the freshwater dolphins were looking for prey at much shorter distances. From this, the researchers surmise that both the dolphin species and the river dolphin were echolocating at short range due to the complex and circuitous river system that they were foraging in.
While both Irawaddy and Ganges river dolphin produced lower intensity biosonar, the Ganges river dolphin had an unexpectedly low frequency biosonar, nearly half as high as expected if this species had been a marine dolphin.
"It is very surprising to see these animals produce such low-frequent biosonar sounds. We are talking about a small toothed whale the size of a porpoise producing sounds that would be more typical for a killer whale or a large pilot whale," says Professor Peter Teglberg Madsen from Aarhus University in Denmark, an expert on toothed whale biosonar and co-author of the study.

A new perspective on the evolution of biosonar
The study suggests that echolocation in toothed whales initially evolved as a short, broadband and low-frequent click. As dolphins and other toothed whales evolved in the open ocean, the need to detect schools of fish or other prey items quickly favored a long-distance biosonar system. As animals gradually evolved to produce and to hear higher sound frequencies, the biosonar beam became more focused and the toothed whales were able to detect prey further away.
However, the Ganges river dolphin separated from other toothed whales early throughout this evolutionary process, adapting to a life in shallow, winding river systems where a high-frequency, long-distance sonar system may have been less important than other factors such as high maneuverability or the flexible neck that helps these animals capture prey at close range or hiding within mangrove roots or similar obstructions.

Improved tools for counting animals
Freshwater dolphins are among the most endangered animal species. Only around a thousand Ganges river dolphins are thought to remain, and they inhabit some of the most polluted and overfished river systems on Earth. The results of this study will help provide local collaborators with a new tool in their struggle to conserve these highly threatened freshwater cetaceans. Using acoustic monitoring devices to identify the local species may help researchers estimate how many animals remain, and to identify what areas are most important to them.


Frants H. Jensen, Alice Rocco, Rubaiyat M. Mansur, Brian D. Smith, Vincent M. Janik, Peter T. Madsen. 2013. Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow, Acoustically Complex Habitat. PLoS ONE. 8 (4): e59284 DOI: 10.1371/journal.pone.0059284