Felix G. Marx

Climate, Critters, and Cetaceans: Cenozoic Drivers of the Evolution of Modern Whales

From Big Fish to Big Whales Whales are the largest animals today, and many feed on the abundant plankton, particularly diatoms, in the oceans. Whales arose and diversified in the Cenozoic, about 30 to 40 million years ago (see the Perspective by Cavin). Marx and Uhen (p. 993) show that their diversity parallels the diversity of diatoms and changes in ocean temperature. Whether there were large predators of plankton before whales has been enigmatic, because the fossil record during the Mesozoic (245 to 65 million years ago) is sparse. Friedman et al. (p. 990) now show that a group of large fish filled this role for nearly 100 million years in the Mesozoic. Although not as large as whales, these globally distributed fish were still several meters long. Their extinction at the Cretaceous-Paleogene boundary 65.5 million years ago may have cleared the seas for the evolution of whales. The diversity of whales during the Cenozoic varied in response to the diversity of diatoms and ocean temperatures. Modern cetaceans, a poster child of evolution, play an important role in the ocean ecosystem as apex predators and nutrient distributors, as well as evolutionary “stepping stones” for the deep sea biota. Recent discussions on the impact of climate change and marine exploitation on current cetacean populations may benefit from insights into what factors have influenced cetacean diversity in the past. Previous studies suggested that the rise of diatoms as dominant marine primary producers and global temperature change were key factors in the evolution of modern whales. Based on a comprehensive diversity data set, we show that much of observed cetacean paleodiversity can indeed be explained by diatom diversity in conjunction with variations in climate as indicated by oxygen stable isotope records (δ18O).

A behavioural framework for the evolution of feeding in predatory aquatic mammals

Extant aquatic mammals are a key component of aquatic ecosystems. Their morphology, ecological role and behaviour are, to a large extent, shaped by their feeding ecology. Nevertheless, the nature of this crucial aspect of their biology is often oversimplified and, consequently, misinterpreted. Here, we introduce a new framework that categorizes the feeding cycle of predatory aquatic mammals into four distinct functional stages (prey capture, manipulation and processing, water removal and swallowing), and details the feeding behaviours that can be employed at each stage. Based on this comprehensive scheme, we propose that the feeding strategies of living aquatic mammals form an evolutionary sequence that recalls the land-to-water transition of their ancestors. Our new conception helps to explain and predict the origin of particular feeding styles, such as baleen-assisted filter feeding in whales and raptorial ‘pierce’ feeding in pinnipeds, and informs the structure of present and past ecosystems.

How whales used to filter: exceptionally preserved baleen in a Miocene cetotheriid

Baleen is a comb‐like structure that enables mysticete whales to bulk feed on vast quantities of small prey, and ultimately allowed them to become the largest animals on Earth. Because baleen rarely fossilises, extremely little is known about its evolution, structure and function outside the living families. Here we describe, for the first time, the exceptionally preserved baleen apparatus of an entirely extinct mysticete morphotype: the Late Miocene cetotheriid, Piscobalaena nana, from the Pisco Formation of Peru. The baleen plates of P. nana are closely spaced and built around relatively dense, fine tubules, as in the enigmatic pygmy right whale, Caperea marginata. Phosphatisation of the intertubular horn, but not the tubules themselves, suggests in vivo intertubular calcification. The size of the rack matches the distribution of nutrient foramina on the palate, and implies the presence of an unusually large subrostral gap. Overall, the baleen morphology of Piscobalaena likely reflects the interacting effects of size, function and phylogeny, and reveals a previously unknown degree of complexity in modern mysticete feeding evolution.

A new Miocene baleen whale from the Peruvian desert.

The Pisco-Ica and Sacaco basins of southern Peru are renowned for their abundance of exceptionally preserved fossil cetaceans, several of which retain traces of soft tissue and occasionally even stomach contents. Previous work has mostly focused on odontocetes, with baleen whales currently being restricted to just three described taxa. Here, we report a new Late Miocene rorqual (family Balaenopteridae), Incakujira anillodefuego gen. et sp. nov., based on two exceptionally preserved specimens from the Pisco Formation exposed at Aguada de Lomas, Sacaco Basin, southern Peru. Incakujira overall closely resembles modern balaenopterids, but stands out for its unusually gracile ascending process of the maxilla, as well as a markedly twisted postglenoid process of the squamosal. The latter likely impeded lateral (omega) rotation of the mandible, in stark contrast with the highly flexible craniomandibular joint of extant lunge-feeding rorquals. Overall, Incakujira expands the still meagre Miocene record of balaenopterids and reveals a previously underappreciated degree of complexity in the evolution of their iconic lunge-feeding strategy.

A new Miocene baleen whale from Peru deciphers the dawn of cetotheriids

Cetotheriidae are an iconic, nearly extinct family of baleen whales (Mysticeti) with a highly distinct cranial morphology. Their origins remain a mystery, with even the most archaic species showing a variety of characteristic features. Here, we describe a new species of archaic cetotheriid, Tiucetus rosae, from the Miocene of Peru. The new material represents the first mysticete from the poorly explored lowest portion of the highly fossiliferous Pisco Formation (allomember P0), and appears to form part of a more archaic assemblage than observed at the well-known localities of Cerro Colorado, Cerro los Quesos, Sud-Sacaco and Aguada de Lomas. Tiucetus resembles basal plicogulans (crown Mysticeti excluding right whales), such as Diorocetus and Parietobalaena, but shares with cetotheriids a distinct morphology of the auditory region, including the presence of an enlarged paroccipital concavity. The distinctive morphology of Tiucetus firmly places Cetotheriidae in the context of the poorly understood ‘cetotheres’ sensu lato, and helps to resolve basal relationships within crown Mysticeti.

Reply to comment by Kienle et al. 2017

Kienle et al . [1] suggest amendments to our framework for feeding in predatory aquatic mammals [2]. Below we reply to their suggestions and demonstrate that they are fundamentally flawed from both a mechanical (feeding cycle, strategies) and an evolutionary perspective. They do, however, inspire an important addition to the range and structuring of capture behaviours encoded in our framework. Feeding cycle . Our framework groups feeding behaviours with similar functions, such as capture and processing, and thus clarifies how different species perform similar tasks during feeding. Kienle et al . [1] suggest that these groupings should be broken up, with capture, ‘external’ processing and manipulation behaviours instead being clustered into a single ‘ingestion’ stage. We question the biological justification for lumping behaviours as disparate as chasing, killing and dismembering. Capturing prey is unrelated to, and need not be followed by, processing. By contrast, ‘external’ and intraoral processing behaviours are functionally akin, with both aiming to dismember prey to start the digestive process. Lumping capture and processing furthermore deviates from the most recent conceptualization of the tetrapod feeding cycle by Schwenk & Rubega [3], which, contra [1], both explicitly includes a separate capture/subjugation stage (p. 12) and specifically associates ‘external’ with intraoral processing (p. 21). Besides the inclusion of water removal, our model differs from [3] only in not recognizing a separate ‘ingestion’ stage. This is because we view ingestion as a moment in time—namely, when food enters the mouth [4]—that can occur during multiple stages, and be achieved and reversed several times during feeding. By contrast, capture, manipulation and processing reflect periods of time over which specific behaviours are performed. Feeding strategies . Kienle et al. [1] criticize our use of a semi-aquatic …

Clawed forelimbs allow northern seals to eat like their ancient ancestors

Streamlined flippers are often considered the defining feature of seals and sea lions, whose very name ‘pinniped’ comes from the Latin pinna and pedis, meaning ‘fin-footed’. Yet not all pinniped limbs are alike. Whereas otariids (fur seals and sea lions) possess stiff streamlined forelimb flippers, phocine seals (northern true seals) have retained a webbed yet mobile paw bearing sharp claws. Here, we show that captive and wild phocines routinely use these claws to secure prey during processing, enabling seals to tear large fish by stretching them between their teeth and forelimbs. ‘Hold and tear’ processing relies on the primitive forelimb anatomy displayed by phocines, which is also found in the early fossil pinniped Enaliarctos. Phocine forelimb anatomy and behaviour therefore provide a glimpse into how the earliest seals likely fed, and indicate what behaviours may have assisted pinnipeds along their journey from terrestrial to aquatic feeding.

Enigmatic Humerus of an Archaic Oligocene—Miocene Neocete from Miyazaki Prefecture, Kyushu, Japan

Abstract. The late Palaeogene represents a crucial time in cetacean evolution that witnessed the origin of modern baleen and toothed whales (Neoceti) from their “archaeocete” ancestors. So far, this fundamental transition has been discussed mainly in terms of cranial morphology, whereas descriptions of postcranial material remain rare. Here, we report a small cetacean humerus from the Nichinan Group (lower Oligocene to lower Miocene), Kushima City, Miyazaki Prefecture, southern Kyushu, Japan. Our specimen resembles archaeocete humeri in being proximodistally elongate and in retaining a distinct deltoid ridge, but shares with neocetes the defining feature of an immobilised elbow joint. It resembles most Oligocene odontocetes in its small size and in lacking a notch marking the position of the distal epiphysis, and is furthermore similar to the enigmatic Microzeuglodon in having a transversely compressed shaft. A morphometric analysis based on five linear measurements, however, fails to cluster our specimen with any other known group of cetaceans, indicating that it is not easily referable to either basal mysticetes or odontocetes. Therefore, we here classify it as Neoceti incertae sedis.