[PaleoMammalogy • 2019] Nehalaennia devossi • A New Balaenopterid Whale (Cetacea, Mysticeti) from the late Miocene of the Southern North Sea Basin and the Evolution of Balaenopterid Diversity

Nehalaennia devossi 

Bisconti​, Munsterman & Post, 2019  

   DOI: 10.7717/peerj.6915

Abstract 

Background:

Balaenopterid mysticetes represent the most successful family-rank group of this clade. Their evolutionary history is characterized by a rich fossil record but the origin of the living genera is still largely not understood. Recent discoveries in the southern border of the North Sea revealed a number of well preserved fossil balaenopterid whales that may help resolving this problem. In particular, skull NMR 14035 shares morphological characters with the living humpback whale, Megaptera novaeangliae and, for this reason, its characteristics are investigated here.

Methods:

The comparative anatomical analysis of the new specimen formed the basis of a new phylogenetic analysis of the Mysticeti based on a matrix including 350 morphological character states scored for 82 Operational Taxonomic Units. The stratigraphic age of the specimen was determined based on the analysis of the dinocyst assemblage recovered in the associated sediment. We assessed clade diversity in Balaenopteridae by counting the numbers of clades in given time intervals and then plotted the results.

Results: 

Nehalaennia devossi n. gen. et sp. is described for the first time from the late Tortonian (8.7–8.1 Ma) of the Westerschelde (The Netherlands). This new taxon belongs to Balaenopteridae and shows a surprisingly high number of advanced characters in the skull morphology. Nehalaennia devossi is compared to a large sample of balaenopterid mysticetes and a phylogenetic analysis placed it as the sister group of a clade including the genus Archaebalaenoptera. The inclusion of this fossil allowed to propose a phylogenetic hypothesis for Balaenopteridae in which (1) Eschrichtiidae (gray whales) represents a family of its own, (2) Balaenopteridae + Eschrichtiidae form a monophyletic group (superfamily Balaenopteroidea), (3) Cetotheriidae is the sister group of Balaenopteroidea, (4) living Balaenoptera species form a monophyletic group and (5) living M. novaeangliae is the sister group of Balaenoptera. Our work reveals a complex phylogenetic history of Balaenopteridae and N. devossi informs us about the early morphological transformations in this family. Over time, this family experienced a number of diversity pulses suggesting that true evolutionary radiations had taken place. The paleoecological drivers of these pulses are then investigated.

 Figure 3: Dorsal view of the holotype skull of Nehalaennia devossi (NMR 999100014035). (A) Photographic representation. (B) Interpretation. Scale bar equals 10 cm.

Systematic Paleontology

Mammalia Linneaus, 1758
Cetartiodactyla Montgelard, Caatzeflis & Douzery, 1997

Cetacea Brisson, 1762
Neoceti Fordyce & De Muizon, 2001

Mysticeti Flower, 1864
Chaeomysticeti Mitchell, 1989
Thalassotherii Bisconti, Lambert & Bosselaers, 2013

Balaenopteridae Gray, 1864

Nehalaennia new genus

Nehalaennia devossi new species

Holotype: Specimen 999100014035 of the collection of the Natuurhistorisch Museum Rotterdam.

Etymology: The genus name is one of the spellings of the name of the Keltic pagan goddess of the sea which was also accepted by Romans when they conquered what is now the most southern province of The Netherlands. The species name is given to honor Dr. John de Vos for his lifelong contribution to Dutch paleontology and his leading role in creating the unique bond and trust between Dutch professional and amateur paleontologists.
 
Differential diagnosis: Nehalaennia devossi differs from Archaebalaenoptera castriarquati in having a rounded anterior border of the supraoccipital, anterior half of the supraoccipital not strongly compressed transversely, widely concave posterior border of the maxilla, shorter and wider ascending process of the maxilla, significantly shorter nasal bones and anterior border of the supraorbital process of the frontal anterolaterally concave. It differs from Plesiobalaenoptera quarantellii in showing a lower superior portion of the periotic, shorter and wider ascending process of the maxilla, more slender lateral process of the maxilla with deeper antorbital notch, posterior end of the posterior process of the periotic more robust and round. It differs from ‘Megaptera’ hubachi in having a ventrally concave glenoid fossa of the squamosal with the postglenoid process projecting ventrally and forming a c. 90° angle with the zygomatic process of the squamosal, in having a rounded anterior border of the supraoccipital, and in lacking exposure of the alisphenoid in the temporal fossa. It differs from ‘Balaenoptera’ bertae in having a wider and rounder anterior border of the supraoccipital, in having an anterolaterally concave anterior border of the supraorbital process of the frontal, in having a vertically-oriented postglenoid process of the squamosal making the glenoid fossa of the squamosal more concave in lateral view. It differs from Incakujira anillodefuego in having a rounder and wider anterior border of the supraoccipital, in having a comparatively shorter and slender supraorbital process of the frontal and a comparatively shorter zygomatic process of the squamosal, in having the premaxilla terminating anteriorly to the nasal. It differs from ‘Megaptera’ miocaena in having a narrower anterior border of the supraoccipital, comparatively longer ascending process of the maxilla with ‘primary dorsal infraorbital foramina’, more concave glenoid fossa of the squamosal. It differs from Fragilicetus velponi in lacking a squamosal bulging into the temporal fossa, in having a wider anterior border of the supraoccipital, in having a less strongly protruding posterolateral corner of the exoccipital, in having a rounded dorsal border of the periotic. It differs from Protororqualus cuvieri in having a wider and rounder anterior border of the supraoccipital, in having shorter zygomatic process of the squamosal, in having a wider space between the posterior border of the maxilla and the anterior border of the supraorbital process of the frontal, and in having an anterolaterally concave anterior border of the supraorbital process of the frontal. The same differences are observed when Nehalaennia devossi is compared against ‘Balaenoptera’ cortesi var. portisi. It differs from Parabalaenoptera baulinensis in having shorter and wider ascending process of the maxilla, rounded supraoccipital and shorter nasal bones.

Nehalaennia devossi differs from the genus Balaenoptera in having a rounded anterior border of the supraoccipital, rounded posterior end of the ascending process of the maxilla, anterolaterally concave anterior border of the supraorbital process of the frontal, alisphenoid not exposed in the temporal fossa. It differs from Megaptera novaeangliae in having zygomatic process of the squamosal less diverging from the longitudinal axis of the skull, anterior border of the pars cochlearis of the periotic not strongly protruded, and more concave glenoid fossa of the squamosal in lateral view.

 Artistic reconstruction of Nehalaennia devossi shows two individuals during feeding upon schooling fishes. The leatherback turtle is used as a reference to show the hypothesized size of the rorqual. Credits for the illustration: Remie Bakker, Manimal Works, Rotterdam, The Netherlands.

Michelangelo Bisconti​, Dirk K. Munsterman and Klaas Post. 2019. A New Balaenopterid Whale from the late Miocene of the Southern North Sea Basin and the Evolution of Balaenopterid Diversity (Cetacea, Mysticeti). PeerJ. 7:e6915  DOI: 10.7717/peerj.6915

[PaleoMammalogy • 2016] Fragilicetus velponi • A New Mysticete Genus and Species and Its Implications for the Origin of Balaenopteridae (Cetacea, Mysticeti)

Fragilicetus velponi 

Bisconti & Bosselaers, 2016

Figure 14. Artistic interpretation of possible interaction between a large shark and Fragilicetus velponi gen. et. sp. nov. as suggested by the shark bite marks on the skull of the holotype specimen. 

The human in the upper right corner serves as a size reference. Illustration by Mark Bosselaers.
  DOI: 10.1111/zoj.12370

Abstract

A new extinct genus, Fragilicetus gen. nov., is described here based on a partial skull of a baleen-bearing whale from the Early Pliocene of the North Sea. Its type species is Fragilicetus velponi sp. nov. This new whale shows a mix of morphological characters that is intermediate between those of Eschrichtiidae and those of Balaenopteridae. A phylogenetic analysis supported this view and provided insights into some of the morphological transformations that occurred in the process leading to the origin of Balaenopteridae. Balaenopterid whales show specialized feeding behaviour that allows them to catch enormous amounts of prey. This behaviour is possible because of the presence of specialized anatomical features in the supraorbital process of the frontal, temporal fossa, glenoid fossa of the squamosal, and dentary. Fragilicetus velponi gen. et sp. nov. shares the shape of the supraorbital process of the frontal and significant details of the temporal fossa with Balaenopteridae but maintains an eschrichtiid- and cetotheriid-like squamosal bulge and posteriorly protruded exoccipital. The character combination exhibited by this cetacean provides important information about the assembly of the specialized morphological features responsible for the highly efficient prey capture mechanics of Balaenopteridae.  

Keywords: Belgium; feeding behavior; Fragilicetus; phylogeny; Pliocene

Figure 14. Artistic interpretation of possible interaction between a large shark and Fragilicetus velponi gen. et. sp. nov. as suggested by the shark bite marks on the skull of the holotype specimen shown in Figure 3.

The human in the upper right corner serves as a size reference. Illustration by Mark Bosselaers. 

Systematic Palaeontology

Class Mammalia Linnaeus, 1758

Order Cetacea Brisson, 1762

Suborder Mysticeti Cope, 1891

Chaeomysticeti Mitchell, 1989

Balaenomorpha Geisler & Sanders, 2003

Superfamily Thalassotherii Bisconti, Lambert & Bosselaers, 2013

Epifamily Balaenopteroidea Flower, 1864

Family Balaenopteridae Gray, 1864

Fragilicetus gen. nov.

Diagnosis: The diagnosis of Fragilicetus includes the presence of eschrichtiid-like and balaenopterid-like features in the same individual. Fragilicetus is distinguished from the other nonbalaenopterid mysticete families based on the presence, in the same individual, of a squamosal bulging into the temporal fossa; posterior projection of the posterolateral corner of the exoccipital; anterior placement of the posterior apex of the lambdoidal crest; squamosal cleft present and v-shaped (turning ventrally at its lateral end); abruptly depressed and flat supraorbital process of the frontal; anterior portion of temporal crest transversely elongated and forming a dorsal roof to the anterior portion of the temporal fossa; very short intertemporal region; infraorbital region of the frontal exposed dorsally between the ascending processes of the maxillae; anterior end of the parietal located more anteriorly than the posterior ends of the ascending process of the maxilla; descending suprameatal surface from the central portion of the periotic to the superior rim of the internal acoustic meatus; endocranial opening of the facial canal separated from the internal acoustic meatus by a thick crista transversa but not prolonged into a groove; triangular anterior process of the periotic; anterior process of the periotic and central portion of periotic on the same plane; groove for VII cranial nerve in posterior process reduced; anteroposteriorly short and flattened posterior process of the periotic.

Etymology: Fragilis, Latin, fragile, in reference to the extreme fragility of the holotype skull. Cetus, Latin, whale.

Type species: Fragilicetus velponi sp. nov. This is currently the only included species.

Figure 5. Holotype skull of Fragilicetus velponi gen. et. sp. nov. in lateral view. A, photographic representation; B, interpretative representation.

Abbreviations: eam, external acoustic meatus; exo, exoccipital; fr, frontal; max, maxilla; opt, optic channel; par, parietal; pgl, postglenoid process of squamosal; ppp, posterior process of the periotic; pt, pterygoid; soc, supraoccipital; sop, supraorbital process of the frontal; sq, squamosal; sqc, squamosal cleft; vom, vomer; zyg, zygomatic process of the squamosal. Scale bar = 300 mm.

Fragilicetus velponi sp. nov.

Holotype: Item no. NMR 999100007727, housed at the Natuurhistorisch Museum Rotterdam, The Netherlands (hereinafter, NMR).

Type locality: The specimen was found along the south-west border of the Deurganckdock, approximately 12 km north-west of Antwerp city centre and 4 km north of the village of Kallo (Fig. 1). The Deurganckdock is an artificial excavation located on the left side of the Scheldt River. The geographical coordinates of the discovery site are 51°17′05″N, 4°15′30″E.

Etymology: Velpon is the brand of the glue used in the preparation of the holotype skull.

Figure 6. Skull of Fragilicetus velponi gen. et. sp. nov. in anterior view. A, photo; B, line drawing.

Abbreviations: bocc, basioccipital; bs, basisphenoid; desc sop, descending part of supraorbital process of the frontal; fm, foramen magnum; fr, frontal; max, maxilla; par, parietal; pt, pterygoid; soc, supraoccipital; sop, supraorbital process of the frontal; sq, squamosal; sqc, squamosal cleft; vom, vomer. Scale bar = 300 mm.

Figure 3. Localizations and orientations of shark bite marks on the holotype skull of Fragilicetus velponi gen. et. sp. nov. as seen from the anterior view. The shark bite marks are in solid black. The skull is in anterior view; only the right side of the skull is shown because it is that part that bears the shark bite marks.

 Abbreviations: ali, alisphenoid; fr, frontal; pal, palatine; par, parietal; pgl, postglenoid process of squamosal; pt, pterygoid; soc, supraoccipital; sq, squamosal; sqc, squamosal cleft; sq-par, squamosal–parietal suture; sq-pt, squamosal-pterygoid suture; tc, temporal crest; vom, vomer; zyg, zygomatic process of the squamosal. Scale bar = 100 mm.

Figure 14. Artistic interpretation of possible interaction between a large shark and Fragilicetus velponi gen. et. sp. nov. as suggested by the shark bite marks on the skull of the holotype specimen shown in Figure 3.

The human in the upper right corner serves as a size reference. Illustration by Mark Bosselaers.

Conclusions

The new genus Fragilicetus is established based on the new fossil species F. velponi. Fragilicetus is the sister group of later Balaenopteridae; Eschrichtiidae is the sister group of the Balaenopteridae clade. Fragilicetus velponi shares several characters with Eschrichtiidae, e.g. the prominent bulge of the squamosal into the temporal fossa and the posterior protrusion of the posterolateral corner of the exoccipital; it shares with later Balaenopteridae the shape of the abruptly depressed supraorbital process of the frontal, details of the articulation of the rostrum with the frontal, and the shape of the supraoccipital. A phylogenetic analysis revealed that F. velponi is closer to Balaenopteridae than to Eschrichtiidae. The phylogenetic analysis also revealed high levels of homoplasy in the Balaenopteroidea clade but these did not prevent the construction of a highly resolved strict consensus tree. However, the high levels of homoplasy prevent unambiguous diagnoses of the internal nodes of Balaenopteridae. The morphological characters observed in F. velponi showed that the appearance of a wide and flat, abruptly depressed supraorbital process of the frontal and an anteriorly constricted supraoccipital preceded the loss of the squamosal bulge and the loss of strong attachment sites for neck muscles in the evolutionary process leading to the balaenopterid lineage. These characters have functional implications respectively related to the anterior placement of the attachment for the temporalis muscle and to the preservation of a mobile head in the earliest phases of balaenopterid evolution.

Finally, the fossil described in the present paper is a demonstration of direct interaction between sharks and mysticetes in the Pliocene; an illustration of this interaction is shown in Figure 14.

Michelangelo Bisconti and Mark Bosselaers. 2016. Fragilicetus velponi: A New Mysticete Genus and Species and Its Implications for the Origin of Balaenopteridae (Mammalia, Cetacea, Mysticeti). Zoological Journal of the Linnean Society. 177(2); 450–474.  DOI: 10.1111/zoj.12370

[PaleoMammalogy • 2013] Balaenoptera bertae • A New Marine Vertebrate Assemblage from the Late Neogene Purisima Formation in Central California, part II: Pinnipeds and Cetaceans

Balaenoptera bertae Boessenecker 2013

The newly discovered Upper Miocene to Upper Pliocene San Gregorio assemblage of the Purisima Formation in Central California has yielded a diverse collection of 34 marine vertebrate taxa, including eight sharks, two bony fish, three marine birds (described in a previous study), and 21 marine mammals. Pinnipeds include the walrus Dusignathus sp., cf. D. seftoni, the fur seal Callorhinus sp., cf. C. gilmorei, and indeterminate otariid bones. Baleen whales include dwarf mysticetes (Herpetocetus bramblei Whitmore & Barnes, 2008, Herpetocetus sp.), two right whales (cf. Eubalaena sp. 1, cf. Eubalaena sp. 2), at least three balaenopterids (“Balaenoptera” cortesi “var.” portisi Sacco, 1890, cf. Balaenoptera, Balaenopteridae gen. et sp. indet.) and a new species of rorqual (Balaenoptera bertae n. sp.) that exhibits a number of derived features that place it within the genus Balaenoptera. This new species of Balaenoptera is relatively small (estimated 61 cm bizygomatic width) and exhibits a comparatively narrow vertex, an obliquely (but precipitously) sloping frontal adjacent to vertex, anteriorly directed and short zygomatic processes, and squamosal creases. Fossil odontocetes include the lipotid “river dolphin” Parapontoporia sternbergi (Gregory & Kellogg, 1927), four true porpoises including a bizarre new genus also known from other strata (Phocoenidae indet., Phocoenidae unnamed genera 1 and 2, and cf. Phocoena), an indeterminate delphinid (Delphinidae indet.) a pilot whale-like delphinid (cf. Globicephalinae indet.), an undetermined sperm whale (cf. Physeteroidea indet.), and an indeterminate odontocete. The new record of Parapontoporia sternbergi is noteworthy as it represents the first association of any earbones (petrosal, tympanic bulla, malleus, and incus) for the extinct genus. Discovery and description of a complete marine mammal assemblage permits faunal comparisons with other published Pliocene marine mammal assemblages from around the globe. The aggregate Pliocene marine mammal assemblage from eastern North Pacific (ENP) shares little in common with the modern fauna, and is mostly composed of extinct genera; notably, phocoenids and odobenids were more diverse than in the ENP today. This indicates that the modern fauna of the ENP did not emerge until after the end of the Pliocene, and probably sometime during the Early Pleistocene. The Pliocene ENP assemblage is similar to that of Japan, and the North Pacific in general shares little with south Pacific, Mediterranean, or North Atlantic marine mammal assemblages, indicating the North Pacific hosted a provincial marine mammal fauna that evolved in isolation from the modern marine mammal fauna, which had already appeared in the North Atlantic by the Early Pliocene.

Keywords: Marine mammals, Pinnipedia, Mysticeti, Odontoceti, Pliocene, Pleistocene, Purisima Formation, new species, Mammiferes marins, Pinnipedia, Mysticeti, Odontoceti, Pliocene, Pleistocene, Formation Purisima, espece nouvelle

Robert W. Boessenecker. 2013. A New Marine Vertebrate Assemblage from the Late Neogene Purisima Formation in Central California, part II: Pinnipeds and Cetaceans. Geodiversitas. 35(4): 815-940. doi: 10.5252/g2013n4a5

Balaenoptera bertae: New Fossil Whale Species Discovered

Robert Boessenecker, a PhD student with the University of Otago, has discovered an extinct species of whale that lived during Pliocene, 3.35 – 2.5 million years ago.

http://www.sci-news.com/paleontology/science-balaenoptera-bertae-new-fossil-whale-species-01750.html