Cheng-Hsiu Tsai

A new Early Oligocene toothed ‘baleen’ whale (Mysticeti: Aetiocetidae) from western North America: one of the oldest and the smallest

Archaic toothed mysticetes represent the evolutionary transition from raptorial to bulk filter feeding in baleen whales. Aetiocetids, in particular, preserve an intermediate morphological stage in which teeth functioned alongside a precursor of baleen, the hallmark of all modern mysticetes. To date, however, aetiocetids are almost exclusively Late Oligocene and coeval with both other toothed mysticetes and fully fledged filter feeders. By contrast, reports of cetaceans from the Early Oligocene remain rare, leaving the origins of aetiocetids, and thus of baleen, largely in the dark. Here, we report a new aetiocetid, Fucaia buelli, from the earliest Oligocene (ca 33–31 Ma) of western North America. The new material narrows the temporal gap between aetiocetids and the oldest known mysticete, Llanocetus (ca 34 Ma). The specimen preserves abundant morphological detail relating to the phylogenetically informative ear bones (otherwise poorly documented in this family), the hyoid apparatus and much of the (heterodont) dentition. Fucaia comprises some of the smallest known mysticetes, comparable in size with the smallest odontocetes. Based on their phylogenetic relationships and dental and mandibular morphology, including tooth wear patterns, we propose that aetiocetids were suction-assisted raptorial feeders and interpret this strategy as a crucial, intermediary step, enabling the transition from raptorial to filter feeding. Following this line of argument, a combination of raptorial and suction feeding would have been ancestral to all toothed mysticetes, and possibly even baleen whales as a whole.

Disparate Heterochronic Processes in Baleen Whale Evolution

Skulls of living baleen whales show distinctive patterns of heterochronic ontogenetic change with implications for mysticete evolution. Here, three baleen whale species are analysed and considered in a heterochronic context. Landmarks show that, during ontogeny, skull morphology changes significantly in the rorqual Balaenoptera borealis and humpback Megaptera novaeangliae (both Balaenopteridae), while the pygmy right whale Caperea marginata (Cetotheriidae: Neobalaeninae) retains an overall juvenile morphology from foetus to adult. Geometric morphometric analyses show that foetal and adult C. marginata are similar, whereas the balaenopterids are more disparate: foetal M. novaeangliae and B. borealis appear in one group, and adult M. novaeangliae and B. borealis are grouped closely. Heterochrony involves paedomorphosis for Caperea, and peramorphosis for the balaenopterids. Heterochrony might cause limited or released developmental constraints, leading to low taxonomic diversity in the single surviving species of neobalaenine, and higher diversity amongst balaenopterids.

Juvenile morphology in baleen whale phylogeny

Phylogenetic reconstructions are sensitive to the influence of ontogeny on morphology. Here, we use foetal/neonatal specimens of known species of living baleen whales (Cetacea: Mysticeti) to show how juvenile morphology of extant species affects phylogenetic placement of the species. In one clade (sei whale, Balaenopteridae), the juvenile is distant from the usual phylogenetic position of adults, but in the other clade (pygmy right whale, Cetotheriidae), the juvenile is close to the adult. Different heterochronic processes at work in the studied species have different influences on juvenile morphology and on phylogenetic placement. This study helps to understand the relationship between evolutionary processes and phylogenetic patterns in baleen whale evolution and, more in general, between phylogeny and ontogeny; likewise, this study provides a proxy how to interpret the phylogeny when fossils that are immature individuals are included. Juvenile individuals in the peramorphic acceleration clades would produce misleading phylogenies, whereas juvenile individuals in the paedomorphic neoteny clades should still provide reliable phylogenetic signals.

Ancestor-descendant relationships in evolution: origin of the extant pygmy right whale, Caperea marginata.

Ancestor–descendant relationships (ADRs), involving descent with modification, are the fundamental concept in evolution, but are usually difficult to recognize. We examined the cladistic relationship between the only reported fossil pygmy right whale, †Miocaperea pulchra, and its sole living relative, the enigmatic pygmy right whale Caperea marginata, the latter represented by both adult and juvenile specimens. †Miocaperea is phylogenetically bracketed between juvenile and adult Caperea marginata in morphologically based analyses, thus suggesting a possible ADR—the first so far identified within baleen whales (Cetacea: Mysticeti). The †Miocaperea–Caperea lineage may show long-term morphological stasis and, in turn, punctuated equilibrium.

Archaic baleen whale from the Kokoamu Greensand: earbones distinguish a new late Oligocene mysticete (Cetacea: Mysticeti) from New Zealand

ABSTRACT †Whakakai waipata is a new genus and species of Mysticeti from the Kokoamu Greensand (Chattian, Late Oligocene, 25–27 Ma) of ‘The Earthquakes’, North Otago. The holotype of †Whakakai waipata is a partial skull, periotics, tympanic bulla, vertebrae and scapula. The earbone morphology of †Whakakai waipata shows a unique combination of diagnostic features including: massive periotic; flange of posterior process and caudal tympanic process flooring stapedial muscle fossa; and an elongated stylomastoid fossa that invades the robust posterior process of the periotic. The tympanoperiotic features in †Whakakai waipata suggest a different acoustic niche from other Oligocene mysticetes, and by inference different feeding habits and habitat. http://zoobank.org/NomenclaturalActs/9D04EC04-1425-4631-9B96-3C279C51E928 http://zoobank.org/urn:lsid:zoobank.org:act:9D04EC04-1425-4631-9B96-3C279C51E928

Quaternary Fossil Gray Whales from Taiwan

Abstract. Two specimens of fossil juvenile gray whale from the sea bottom between Taiwan and the Penghu Islands are Quaternary in age, and probably early Holocene, no older than 11–12 ka. Both specimens preserve the posterior portion of the skull from the occipital condyles to the broken frontals; the earbones are missing. A key diagnostic feature of eschrichtiids, paired tuberosities on the supraoccipital, occurs in both specimens. Because of incompleteness and differences with the living gray whale, the fossils are designated as Eschrichtius sp. rather than Eschrichtius robustus. This report of fossil gray whales is the first from Taiwan. The fossils expand the known range of Quaternary gray whales, and this occurrence of juveniles is consistent with a possible paleo-breeding- or nursery range.

An Early Pleistocene gray whale (Cetacea: Eschrichtiidae) from the Rio Dell Formation of northern California

Abstract. The earliest fossil gray whale (Eschrichtius) from the eastern North Pacific is reported from the Lower Pleistocene Rio Dell Formation of Humboldt County, Northern California. This specimen, a tympanic bulla and posterior process, is identical in morphology to extant Eschrichtius robustus and differs from Pliocene Eschrichtius sp. from the western North Pacific (Japan). Thus, it suggests that the modern bulla morphology of the gray whale had been acquired by the Early Pleistocene. The absence of fossil Eschrichtius in the Pliocene of the eastern North Pacific may indicate that the extant gray whale lineage originated in the western North Pacific during the Pliocene before invading the eastern North Pacific during the Early Pleistocene. Further discoveries of Plio-Pleistocene gray whale fossils will help test this hypothesis and properly interpret the evolutionary history of eschrichtiid clade.

Northern pygmy right whales highlight Quaternary marine mammal interchange

The pygmy right whale, Caperea marginata, is the most enigmatic living whale. Little is known about its ecology and behaviour, but unusual specialisations of visual pigments [1], mitochondrial tRNAs [2], and postcranial anatomy [3] suggest a lifestyle different from that of other extant whales. Geographically, Caperea represents the only major baleen whale lineage entirely restricted to the Southern Ocean. Caperea-like fossils, the oldest of which date to the Late Miocene, are exceedingly rare and likewise limited to the Southern Hemisphere [4], despite a more substantial history of fossil sampling north of the equator. Two new Pleistocene fossils now provide unexpected evidence of a brief and relatively recent period in geological history when Caperea occurred in the Northern Hemisphere (Figure 1A,B).

A Miocene breeding ground of an extinct baleen whale (Cetacea: Mysticeti)

Locating breeding sites is definitely a key to understanding the ecological requirements and maintaining the sustainability of populations/species. Here I re-examined published specimens of an extinct baleen whale, Parietobalaena yamaokai, from the lower part of Itahashi Formation (16.1–15.6 Ma, Middle Miocene) in Shobara, Hiroshima, Japan. A critical and previously unnoticed feature, the open suture between the supraoccipital and exoccipital, in one specimen indicates the preservation of a very young individual–under six months old and even close to a new-born calf. Given the occurrence of a new-born whale and relatively abundant assemblage of Parietobalaena yamaokai, I propose a previously hidden and unknown breeding ground for the extinct baleen whale, P. yamaokai, in the Middle Miocene of Shobara (16.1–15.6 Ma), Hiroshima. Discovery of paleo-breeding sites of extinct populations/species should further help us to understand biological extinctions from a long-term perspective as conservation paleobiology aims to offer new insights into policy making for conserving endangered populations/species.

A new archaic baleen whale Toipahautea waitaki (early Late Oligocene, New Zealand) and the origins of crown Mysticeti

A new genus and species of extinct baleen whale †Toipahautea waitaki (Late Oligocene, New Zealand) is based on a skull and associated bones, from the lower Kokoamu Greensand, about 27.5 Ma (local upper Whaingaroan Stage, early Chattian). The upper jaw includes a thin, elongate and apparently toothless maxilla, with evidence of arterial supply for baleen. Open sutures with the premaxilla suggest a flexible (kinetic) upper jaw. The blowhole is well forward. The mandible is bowed laterally and slightly dorsally; unlike the Eomysticetidae, there are no mandibular alveoli, and the coronoid process is tapered and curved laterally. Jaw structure is consistent with baleen-assisted gulp-feeding. The age of early Chattian makes †Toipahautea a very early, if not the oldest named, toothless and baleen-bearing mysticete, suggesting that the full transition from toothed to baleen-bearing probably occurred in the Early Oligocene. Late Oligocene mysticetes vary considerably in jaw form and kinesis, tooth form and function, and development of baleen, implying a wide range of raptorial, suctorial and filter-feeding behaviour. More study may elucidate the function of jaws, teeth and baleen in terms of opportunist/generalist feeding, as in modern gray whales, versus specialized feeding. We here propose that early mysticetes, when transitioned from toothed to baleen-bearing, were generalists and opportunists instead of specializing in any forms of feeding strategies. In addition, two different phylogenetic analyses placed †Toipahautea either in a polytomy including crown Mysticeti, or immediately basal to the crown, and above †Eomysticetidae in both cases. Because the †Toipahautea waitaki holotype is an immature individual, it may plot more basally in phylogeny than its true position.