Current Biology

Minireview

The Ecological Rise of Whales Chronicled

by the Fossil Record
Nicholas D. Pyenson
Department of Paleobiology, MRC-121, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA
Correspondence: pyensonn@si.edu
http://dx.doi.org/10.1016/j.cub.2017.05.001

The evolution of cetaceans is one of the best examples of macroevolution documented from the fossil record.
While ecological transitions dominate each phase of cetacean history, this context is rarely stated explicitly.
The first major ecological phase involves a transition from riverine and deltaic environments to marine ones,
concomitant with dramatic evolutionary transformations documented in their early fossil record. The second
major phase involves ecological shifts associated with evolutionary innovations: echolocation (facilitating
hunting prey at depth) and filter-feeding (enhancing foraging efficiency on small prey). This latter phase
involves body size shifts, attributable to changes in foraging depth and environmental forcing, as well as
re-invasions of freshwater systems on continental basins by multiple lineages. Modern phenomena driving
cetacean ecology, such as trophic dynamics and arms races, have an evolutionary basis that remains mostly
unexamined. The fossil record of cetaceans provides an historical basis for understanding current ecological
mechanisms and consequences, especially as global climate change rapidly alters ocean and river ecosys-
tems at rates and scales comparable to those over geologic time.

Introduction ecosystems, showing that these changes were not unidirectional

Cetacean evolution receives wide recognition as a preeminent
example of macroevolutionary change [1]. This recognition
derives primarily from an excellent fossil record of their early
evolution, mostly from the middle to late Eocene epoch (from
55–34 Ma, or million years ago). The early evolution of whales
documents a suite of evolutionary transformations to anatomical
systems (and inferred life history and behavioral changes) that
occurred concomitant with a major ecological transition from
terrestrial ancestry to obligate aquatic life [2–4]. In the subse-
quent 34 million years, cetacean diversification is associated
with evolutionary innovations that are unique among mammals
(i.e., filter-feeding and underwater echolocation) [5–6], and the
attainment of body sizes that are unrivaled by any other verte-
brate, living or extinct [7].
Today, there are over 80 cetacean species divided into
two clades: mysticetes, or baleen whales; and odontocetes, or
toothed whales. Mysticetes comprise right whales, bowhead
whales, pygmy right whales, gray whales, and rorquals, the
most species-rich mysticete group. Odontocetes comprise
pygmy and dwarf sperm whales, sperm whales, beaked whales,
separate ‘river dolphin’ lineages, belugas, narwhals, true por-
poises, and true oceanic dolphins, the most species-rich odon-
tocete clade (Figure 1). Many mysticetes and odontocetes have
extinct relatives that are closely related to living species, while
other fossil lineages are unique, without descendants [5–6].
The evolutionary relationships among living and fossil cetaceans
remain works in progress.
The distinction between evolutionary transformation and
ecological transition in cetacean history is important for several
reasons. First, evolutionary and ecological processes have man-
ifestly different mechanisms, scales, modes, rates, and conse-
quences [8]. Second, multiple ecological transitions happened
in cetacean history, across terrestrial, aquatic, and marine
[9]. During these transitions, evolutionary transformations may
lag, accelerate, or change independently with respect to habitat
changes. Lastly, most fossil cetacean studies have focused on
evolutionary change as read from their skeletal record, with little
ecological context. Reframing cetacean evolution in the context
of their changing ecological role illuminates key areas of investi-
gation for resolving how cetaceans underwent an ecological rise
from initial absence to their current dominance as top consumers
in ocean food webs (Figure 1).
At the Water’s Edge
Over the past 250 million years, dozens of tetrapod lineages
have adapted to aquatic life from terrestrial ancestry. Cetaceans
represent just one of the more recent episodes of these inva-
sions. Prior to the Cenozoic era (66 Ma to today), over a dozen
diapsid reptile lineages underwent similar transitions in the
Mesozoic era (252–66 Ma), resulting in lineages adapted to a
range of semi-aquatic to pelagic lifestyles, such as ichthyosaurs
and mosasaurs, whose bauplans mirror different stages in ceta-
cean evolution [10]. The early cetacean fossil record, in this light,
is unusual because a comparable record for any other tetrapod
group early in their history, undergoing similar ecological transi-
tions, is sparse.
Basal branching cetacean lineages, sometimes called ‘ar-
chaeocetes’ (or stem cetaceans, a term used here; Box 1)
represent lineages ranging from quadrupedal forms to obligate
aquatic ones with vestigial hind limbs (Figure 1). The ecomorpho-
logic diversity of quadrupedal stem cetaceans is relatively broad
[2–4]. Feeding ecomorphologies ranged from longirostral, gav-
ialid crocodilian-like taxa (Remingtonocetus) [4] to potentially
molluscivorous taxa with ventrally deflected rostra (Makarace-
tus) [11]. Locomotory modes among stem cetaceans also
ranged from likely cursorial taxa (e.g., Pakicetus) to lineages

R558 Current Biology 27, R558–R564, June 5, 2017 Published by Elsevier Ltd.
Current Biology

Minireview

Figure 1. The ecological rise of whales from the Eocene to today.

(A) Reconstructed assemblage of stem cetaceans from the middle to late Eocene that did not live together, nor in similar environments (see [5]), including
(clockwise from the top), a stranded Dorudon, Ambulocetus, Pakicetus, Kutchicetus, and Rodhocetus (art by Carl Buell). (B) Blue whale (Balaenoptera musculus)
feeding on a prey patch off the coast of California. (Figures modified from [5] and [24], the latter by the BBC under J. Calambokidis NMFS permit 16111.)

that were semi-aquatic (e.g., Ambulocetus, Rodhocetus, Maia-
cetus) and moved in the water using some combination of
fore- and hindlimb propulsion [3,4,12]. Investigations into the
bony structures of the inner and middle ear of stem cetaceans
have revealed several traits already present in these quadru-
pedal taxa that presaged underwater, directional hearing in later
obligate marine taxa [13]. Although quadrupedal bauplans domi-
nated the incipient stages of cetacean evolution from 53–41
Ma, these bauplans disappeared as obligate marine stem ceta-
ceans such as Basilosaurus and Dorudon evolved in the middle
to late Eocene (43–34 Ma) [2–4,14] (Figure 2).
With the exception of a handful of genera represented by
dense monotaxic bonebeds or near-complete individual skele-
tons [15,16], most early stem cetaceans are known from far
less complete skeletons than many artistic interpretations would
indicate (Figure 1). The incompleteness of these fossil taxa,
especially for early quadrupedal forms such as Pakicetus, under-
scores the challenges of reconstructing their ecology. Although
the early stages of cetacean evolution have been commonly
described as a ‘land to sea’ transition, it is clear that the earliest
stem cetaceans were mostly semi-aquatic in habit and feeding,
as opposed to being characterized purely as terrestrial [17].
More specifically, the land to sea shorthand pertains more to
the environmental setting of these stem cetaceans, rather than
their ecomorphology. Semi-aquatic stem cetacean lineages
were likely ecological analogs to today’s estuarine and coastal
tetrapods of similar body size (e.g., crocodylians, carnivorans),
based on isotopic characterizations of their habitat use and
diet from analyses of their bones and teeth [18]. Well-sampled
stratigraphic sections that contain an abundance of stem ceta-
ceans, such as those in the Wadi Hitan of Egypt [19], can provide
a fine-scale control on environmental change through time,
underpinning more detailed analyses of changing diets or life
history.
Oceans Onwards
The second major phase in the ecological history of cetaceans
relates to their diversification in ocean ecosystems. Some line-
ages of semi-aquatic stem cetaceans, such as protocetids,
achieved a broad geographic distribution by the middle Eocene,
ranging to North America, Europe, Africa, and Asia [2,4]. The
dispersal of these lineages, however, can be mostly explained
via coastal habitats and island archipelagos that connected
much of Laurasia and Africa in the Paleogene. Protocetids

Box 1. Glossary of terms

Stem cetaceans: Extinct cetacean lineages that evolved prior to the origin of crown Cetacea. This paraphyletic group includes all
extinct lineages more closely related to living whales than to Hippopotamus, their closest living relative. The phylogenetic grouping
of living whales and Hippopotamus has been termed Whippomorpha, although there were likely extinct lineages more closely
related to whales than hippos, such as Raoellidae, which are Eocene age artiodactyls from South Asia [15]. The oldest stem ceta-
cean with robust stratigraphic constraint is Himalayacetus subathuensis at about 53.2 Ma (see [31] for more details).
Crown cetaceans: A clade defined by the most recent common ancestor between all living baleen and toothed whales, and all of
their descendants, extinct or extant. Some authors have also called this clade Neoceti [31]. The oldest crown cetacean is the stem
mysticete Mystacodon selenensis, whose estimated age is about 36.4 Ma [33].
Mysticeti: A clade that includes all living baleen whales and stem relatives that are closer to them than the lineage leading to odon-
tocetes. All living mysticetes are filter-feeders [23] and lack adult, mineralized teeth [27], although many stem mysticetes
possessed adult dentition [24,25].
Odontoceti: A clade that includes all living toothed whales and extinct relatives that are closer to them than the lineage leading to
mysticetes. All living odontocetes echolocate. The oldest odontocete is Simocetus rayi from the early Oligocene of Oregon, USA,
as much as 32 Ma (see [31]).

Current Biology 27, R558–R564, June 5, 2017 R559

 Current Biology

Minireview

Eocene Oligocene Miocene Pli Q (Box 1 and Figure 2). Two major evolutionary innovations that
56.0 33.9 23.0 5.3
2.62.6 0 Ma characterize crown cetaceans reflect ecological modes without
Hippopotamus
parallel in mammals: filter-feeding in mysticetes; and underwater
Pakicetidae
Ambulocetidae
echolocation in odontocetes. The evolutionary origins of both in-
Origin of underwater hearing novations are not straightforward, although they both represent
Remingtonocetidae Stem cetaceans
or key innovations that fueled the explosive radiation of crown ce-
“Protocetidae” “archaeocetes”
Semi-aquatic
taceans during the Oligocene [22]. Living mysticetes filter-feed
using keratinous plates, called baleen, which hang from the
Obligate aquatic
roof of the mouth and enhance foraging efficiency on fish or krill
Basilosauridae
Mystacodon
Mystacodon selenensis suspended in the water column [23]. However, many stem mys-
Toothed stem mysticetes
“toothed” Stem ticetes possessed mineralized adult teeth [5,24,25], and the
Llanocetus
Llanocetus dentricrenatus
dentricrenatus
mysticetes disappearance of teeth in some crownward stem mysticetes
?
has been presumably linked to the appearance of baleen,
Eomysticetidae
Eomysticetidae

Crown
Crown Mysticeti
Mysticeti
Origin of filter-feeding
Crown Cetacea
Origin of echolocation
Echovenator sandersi
Echovenator sandersi
Other
other Xenorophidae
Simocetus rayi
Simocetus rayi
Other stem Odontoceti
Crown Odontoceti
Odontoceti
Mixed
Freshwater
Marine
Crown clades
Figure 2. Whale phylogeny and evolution.
Generalized view of cetacean evolution over geologic time, showing major
lineages, innovations and ecological changes. Phylogenetic relationships
follow morphological studies for fossil (stem) lineages [2,5,6,14,25,29,33], and
molecular studies for living taxa [22,32]. Higher taxonomic groups among
crown cetaceans show stratigraphic ranges that include both stem and crown
lineages. Origin times and phylogenetic placements of key innovations
are based on [13] for underwater hearing, [29] for echolocation, and [27] for
filter-feeding. Abbreviations: Ma, millions of years ago; Pli, Pliocene; and
Q, Quaternary.
were smaller in body size than obligate marine basilosaurids,
which achieved a global distribution by the late Eocene
[2–4,16]. Basilosaurids have been reported from every continent
and ranged in body size from 5 to 18 m-long taxa [2,14]. Based
on rare stomach contents, basilosaurids were piscivorous [14],
and some species likely consumed other stem cetaceans [20],
in a hypercarnivorous mode reminiscent of mosasaurs — Creta-
ceous diapsids with a convergent bauplan [10]. Fossil assem-
blages of stem cetaceans including both semi-aquatic and
obligate marine forms are best represented by the fossil-rich
Eocene age lagoonal deposits of the Wadi Hitan [2,3,19].
There is evidence that some pelagic stem cetaceans survived
through the Eocene–Oligocene boundary (34 Ma) [21], implying
a co-existence with the first crown cetaceans, a group that in-
cludes lineages leading to today’s baleen and toothed whales
Balaenidae
although there is no direct evidence for baleen in any mysticetes
Neobalaenidae older than the middle Miocene [26]. Thus, the exact antiquity of
Eschrichtiidae baleen-enabled filter-feeding remains unclear [27].
Balaenopteridae
Echolocation is a form of biological sonar that requires sound
generation, sound reception, and signal processing using fat
bodies, air sinuses and muscles of the face, along with fat bodies
that connect the jaw to the ear bones [28]. Echolocation has
Stem
evolved several times among tetrapods, although odontocetes
odontocetes are the only ones that echolocate underwater. Evidence for
high frequency hearing, a trait for all living odontocetes, has
Physeteroidea been documented in the stem odontocete Echovenator sand-
Ziphiidae ersi, pointing to an Oligocene origin for echolocation (27–24
Platanista
Ma) [29]. More work on the timing and mode of the other two
Lipotes
Inioidea
components in echolocation (sound generation and signal pro-
cessing) may be difficult to infer without clear bony correlates
Delphinoidea, of their soft tissue bases. The co-occurrence of stem and crown
including:
Monodontidae, cetaceans in the Oligocene signals non-analogous complexities
Phocoenidae,
and to cetacean community structure that require more study [21].
Delphinidae
Fordyce [30] originally pointed to global ocean changes at
Current Biology the Eo–Oligocene boundary as a potential driver for the origin
of crown cetaceans, with the establishment of the Antarctic
Circumpolar Current (ACC) system and the large-scale changes
to oceanic productivity that accompanied it. While intuitively
appealing, there are several problems with invoking this explana-
tion, in terms of timing and agency. First, an increasingly better
reported fossil record of Oligocene age crown cetaceans points
to the start of this epoch as the primary time for their diversifica-
tion [31], but molecular estimates of divergence times consis-
tently point to an older origin for crown Cetacea, specifically
in the late Eocene [32]. For decades, the only crown cetacean
known from this age was a stem mysticete, Llanocetus dentri-
crenatus, from late Eocene sediments of Seymour Island,
Antarctica, at approximately 34.2 Ma [32]. Lambert et al. [33]
recently reported an older crown cetacean, Mystacodon sele-
nensis, from the late Eocene of Peru, which is geochronologically
older than Llanocetus at an estimated age of 36.4 Ma. There is no
record of odontocetes older than the early Oligocene (32 Ma),
yet the antiquity of Mystacodon implies a ghost lineage for odon-
tocetes extending to the late Eocene as well (Figure 2).
Second, there is a lack of clarity about specific ecological
mechanisms that explain how Eo-Oligocene ocean changes
promoted the diversification of crown cetaceans. Berger [34]
emphasized the presence of silicate in deep water upwelling
from the ACC as a resource opportunity for both mysti-
cetes and odontocetes. However, opportunities are necessary
but not sufficient to explain the selection pressures for bulk

R560 Current Biology 27, R558–R564, June 5, 2017

Current Biology
Minireview
A B
Total length (meters)
0 5 10 15 20
0
First right whales
10
First rorquals
Ma
20
Crown Mysticeti
Origin of filter-feeding
30
Llanocetus dentricrenatus
40
filter-feeding and biosonar-assisted hunting (see [35,36]), espe-
cially if both traits did not evolve until 4–10 million years after
the establishment of the ACC (Figures 2, 3). There is a need
for testable models that explain the ecological context of ceta-
cean evolution with known mechanisms and responses [37],
especially for relevant metrics that can be observed in the fossil
record; some approaches to meet these challenges are dis-
cussed below.
Escalation, Predation and Coevolution
Today’s cetaceans are major ecological consumers with large
body sizes (even the smallest cetaceans are large compared
to most mammal species), they possess high metabolic rates,
and they occupy high trophic levels in ocean food webs [37].
This ecological position is the result of evolutionary pro-
cesses, yet there are few studies that have explicitly consid-
ered their trophic interactions as a consequence of long-
term generative processes, such as arms races [38]. For
example, living sperm whales (Physeter macrocephalus) are
the largest odontocetes of all-time, and they pursue some of
the largest cephalopods known (e.g., giant and colossal
squids) as single prey items at depth. The long fossil record
Figure 4. Killer whale attack on a beaked whale.
A killer whale (Orcinus orca) pursues and bites a beaked whale (Mesoplodon sp.) off Western Australia. Images reproduced from Figure 6 in [42].
Figure 3. Baleen whale body size evolution
18
O over geologic time.
25 0 2 4 6 (A) Mean body lengths for living and fossil
mysticetes over geologic time. Shaded areas
Q show quantiles of Brownian motion simulations on
Ice Ages mysticete phylogeny to show that the range of
Pli
modern body size lacks small species (less than
5 m) and is over-represented by large ones (more
Miocene
than 10 m), relative to the fossil record. Circles,
baleen-bearing mysticetes; triangles; toothed
mysticetes. (B) A smooth-spline fitted to oxygen
Start of
isotope curve as a proxy for modelling tempera-
Antarctic ture-dependent body size evolution. From [7].
Olig
Circumpolar
Current
Eo
of sperm whales supports this escala-
Current Biology
tion in body size, but the diversity of
fossil relatives points to hypercarnivo-
rous feeding modes during the Neogene without analog for
today’s sperm whales [39].
A plurality of co-evolutionary interactions also likely shaped
cetacean evolution. Parasitic or commensal relationships with
whale barnacles have been proposed as co-opted anti-predatory
responses (i.e., armor) for some mysticetes, in response to killer
whale (Orcinus orca) predation [40]. Intra-guild (and cross-clade)
predation by Orcinus (e.g., [41]; Figure 4) is among the few exam-
ples for this feeding mode among living marine tetrapods [10], but
its prevalence in the geological history of cetaceans is unknown.
The few studies on the fossil record of coevolutionary factors
(e.g., stratigraphic and geographic co-occurrences between
barnacles and cetaceans) have already yielded insight into the
antiquity of migration in humpback whales [42]. The selection
advantages conferred by these interactions are compelling,
and they range across many taxa and habitats. For example,
the deposition of whale carcasses on the seafloor creates unique
communities that persist for decades, depending on water depth.
Numerous fossil whale examples demonstrate that these com-
munities had different components over geologic time, including
linkages with other deep-sea habitats such as woodfalls, cold
seeps, and hydrothermal vents [31].
Current Biology 27, R558–R564, June 5, 2017 R561

Figure 5. Whale mass stranding from the Miocene.
High dynamic range image of orthogonal three-dimensional point clouds
showing several over-lapping adult and juvenile mysticetes from Cerro Bal-
lena, a Miocene mass stranding fossil locality. Small scale bars 20 cm, large
scale bars 30 cm. See [45] for more details.
Morphological Parallelism — Signs of Trophic
Convergence?
The diversity of living cetaceans shows clear examples of
morphological and ecological parallelism. For example, delphi-
nid species belonging to Cephalorhynchus have radiated
into small, protected coastal habitats of the Southern Hemi-
sphere [43], and have converged in their small size, external
morphology, and even biosonar frequency range with coastal
phocoenids (or true porpoises). There are examples of this
type of parallelism (i.e., convergence within clades) in the fossil
record of cetaceans: Australodelphis, a fossil delphinid from
Pliocene age rocks of the Sørsdal Formation, East Antarctica,
exhibits a cranial morphology most similar to living beaked
whales (Ziphiidae) [31]. Beyond parallelism within clades, the
cetacean fossil record also shows a compelling example of
cross-clade convergence with Odobenocetops, a fossil odonto-
cete from Mio-Pliocene rocks of South America, and living wal-
ruses (Odobenus), known only from the Northern Hemisphere,
based on its heavily foreshortened rostrum, vaulted palate, and
large tusks [31]. Analyses using taxonomically broad datasets
from the diet of marine tetrapods show that such patterns of
similarities in feeding morphology are tied to trophic conver-
gences, which cut across deep divergences in over 250 million
years of tetrapod phylogeny [44]. There are likely common
causes for these similarities, but they remain unexplored.
Abundance and Body Size as Semaphores of Ecological
Power
Traditionally, paleontologists have relied on taxonomic richness
(i.e., counts of taxa within time bins) as a metric for ecological
diversity through geologic time. Hyperabundant fossil cetacean
localities in Egypt, California, Peru, and Chile [19,31,45] provide
an additional avenue for answering paleoecological questions
using abundance-based metrics. In these scenarios, strati-
graphic and taphonomic bias control remain paramount, but
the ability to count individuals and specimens provides a direct
means to understand the ecological structure in a given fossil
assemblage, sometimes with statistical strength. For example,
R562 Current Biology 27, R558–R564, June 5, 2017
Current Biology
Minireview
the high relative abundance of large fossil mysticetes at
Cerro Ballena (Figure 5), in the Atacama of Chile [45], along
with other taphonomic traits, mirror similar patterns in toxin-
mediated mass strandings today [46], strongly pointing to
harmful algal blooms as the mechanism responsible for this
Miocene death assemblage.
Body size is also a direct means to evaluate ecological power
because organismal size correlates strongly with many meta-
bolic functions and life history traits. While odontocete body
size is likely strongly related to foraging and dive depth [47],
the extremely large body size among mysticetes such as blue,
fin, bowhead and right whales have independent origins [7]
(Figure 3). These body size changes are also linked to the
appearance, at the start of the Ice Ages, of seasonal, dense,
but patchy prey aggregations on which the largest mysticetes
most efficiently feed [24].
Past and Future Ecological Contexts
Geologic-scale changes to marine habitats, through eustatic
sea-level changes, tectonic uplift, and ocean portal openings
and closures, have likely had a major impact on cetacean evolu-
tion [31]. For example, large-scale sea-level fluctuations in
the Pleistocene repeatedly eliminated seafloor foraging habitats
and altered migration routes [42,48], yet many lineages persisted
through these changes, showing ecological flexibility. In other
cases, flooding of continental basins (e.g., Amazonia) provided
ecological opportunities for odontocete invasions of freshwater
ecosystems [49]. The implications of future sea-level rise, in
terms of cetacean habitat creation and/or elimination, along
with changes to prey availability, remain poorly understood.
Humans have hunted cetaceans for thousands of years, but
technological innovations in the past few hundred years have
accelerated and increased the rate and scale of impact [37,50].
Pelagic whaling, for example, decimated the abundances of
baleen whales, which has had downstream effects on other ma-
rine mammals via trophic cascades [37,50]. Other detrimental
impacts, such as by-catch or wholesale habitat modification
(e.g., dam construction) threaten marginal cetacean species
(e.g., river dolphins) with extinction. Anthropogenic climate
change, especially via shrinking sea ice and altered ocean
thermal gradients at polar latitudes, will lead to range shifts
for many cetacean species, precipitating new trophic interac-
tions and/or colonization opportunities [10,50]. These changes
may be abrupt, episodic, and difficult to predict [8]. Like other
historical data, the fossil record provides a crucible for the
ecological limits and capabilities of cetaceans, and should be a
source of continued data for predicting future states in the
Anthropocene.
ACKNOWLEDGEMENTS
I thank A.T. Boersma, J.A. Goldbogen, A.H. Fleming, A.S. Friedlaender, N.P.
Kelley, M.S. Leslie, M.R. McCurry, M.F. McKenna, A. O’Dea, J.F. Parham,
C.M. Peredo, J. Ve
lez-Juarbe, G.J. Vermeij and M. Yamato for helpful conver-
sations about these topics over many years.
REFERENCES
1. Zimmer, C. (2011). Evolution: The Triumph of an Idea (New York:
HarperCollins Publishers, Inc).
Current Biology
Minireview
2. Uhen, M.D. (2010). The origin (s) of whales. Annu. Rev. Eart. Planet. Sci. 38,
189–219.
3. Gingerich, P.D. (2003). Land-to-sea transition in early whales: evolu-
tion of Eocene Archaeoceti (Cetacea) in relation to skeletal propor-
tions and locomotion of living semiaquatic mammals. Paleobiology
29, 429–454.
4. Thewissen, J.G., and Williams, E.M. (2002). The early radiations of Ceta-
cea (Mammalia): evolutionary pattern and developmental correlations.
Annu. Rev. Ecol. Syst. 33, 73–90.
5. Deme
re

, T.A., McGowen, M.R., Berta, A., and Gatesy, J. (2008). Morpho-
logical and molecular evidence for a stepwise evolutionary transition from
teeth to baleen in mysticete whales. Syst. Biol. 57, 15–37.
6. Geisler, J.H., Colbert, M.W., and Carew, J.L. (2014). A new fossil species
supports an early origin for toothed whale echolocation. Nature 508,
383–386.
7. Slater, G.J., Goldbogen, J.A., and Pyenson, N.D. (2017). Independent evo-
lution of baleen whale gigantism linked to Plio-Pleistocene ocean dy-
namics. Proc. R. Soc. B 284, 20170546.
8. Jackson, J.B., and Erwin, D.H. (2006). What can we learn about ecology
and evolution from the fossil record? Trends Ecol. Evol. 21, 322–328.
9. Vermeij, G.J., and Dudley, R. (2000). Why are there so few evolutionary
transitions between aquatic and terrestrial ecosystems? Biol. J. Linn.
Soc. 70, 541–554.
10. Kelley, N.P., and Pyenson, N.D. (2015). Evolutionary innovation and ecol-
ogy in marine tetrapods from the Triassic to the Anthropocene. Science
348, aaa3716.
11. Gingerich, P.D., Zalmout, I.S., Ul-Haq, M., and Bhatti, M.A. (2005).
Makaracetus bidens, a new protocetid archaeocete (Mammalia, Ceta-
cea) from the early middle Eocene of Balochistan (Pakistan). Univer-
sity of Michigan Contributions from the Museum of Paleontology 31,
197–210.
12. Thewissen, J.G.M., and Fish, F.E. (1997). Locomotor evolution in the
earliest cetaceans: functional model, modern analogues, and paleonto-
logical evidence. Paleobiology 23, 482–490.
13. Nummela, S., Thewissen, J.G.M., Bajpai, S., Hussain, S.T., and Kumar, K.
(2004). Eocene evolution of whale hearing. Nature 430, 776–778.
14. Uhen, M.D. (2004). Form, function, and anatomy of Dorudon atrox (Mam-
malia, Cetacea): an archaeocete from the middle to late Eocene of Egypt.
Univ. Michigan Papers Paleont 34, 1–222.
15. Thewissen, J.G., Cooper, L.N., Clementz, M.T., Bajpai, S., and Tiwari, B.N.
(2007). Whales originated from aquatic artiodactyls in the Eocene epoch of
India. Nature 450, 1190–1194.
16. Gingerich, P.D., von Koenigswald, W., Sanders, W.J., Smith, B.H., and
Zalmout, I.S. (2009). New protocetid whale from the middle Eocene of
Pakistan: birth on land, precocial development, and sexual dimorphism.
PLoS One 4, e4366.
17. Thewissen, J.G.M., Sensor, J.D., Clementz, M.T., and Bajpai, S. (2011).
Evolution of dental wear and diet during the origin of whales. Paleobiology
37, 655–669.
18. Clementz, M.T., Goswami, A., Gingerich, P.D., and Koch, P.L. (2006). Iso-
topic records from early whales and sea cows: contrasting patterns of
ecological transition. J. Vert. Paleont 26, 355–370.
19. Peters, S.E., Antar, M.S.M., Zalmout, I.S., and Gingerich, P.D. (2009).
Sequence stratigraphic control on preservation of late Eocene whales
and other vertebrates at Wadi Al-Hitan, Egypt. Palaios 24, 290–302.
20. Fahlke, J.M. (2012). Bite marks revisited—evidence for middle-to-late
Eocene Basilosaurus isis predation on Dorudon atrox (both Cetacea, Basi-
losauridae). Palaeont. Electronica 15, 32A.
21. Clementz, M.T., Fordyce, R.E., Peek, S.L., and Fox, D.L. (2014). Ancient
marine isoscapes and isotopic evidence of bulk-feeding by Oligocene ce-
taceans. Palaeo. Palaeo. Palaeo. 400, 28–40.
22. Slater, G.J., Price, S.A., Santini, F., and Alfaro, M.E. (2010). Diversity
versus disparity and the radiation of modern cetaceans. Proc. R. Soc. B
277, 3097–3104.
23. Goldbogen, J.A., Cade, D.E., Calambokidis, J., Friedlaender, A.S., Pot-
vin, J., Segre, P.S., and Werth, A.J. (2017). How baleen whales feed:
the biomechanics of engulfment and filtration. Annu. Rev. Mar. Sci. 9,
367–386.
24. Fitzgerald, E.M. (2012). Archaeocete-like jaws in a baleen whale. Biol. Lett.
8, 94–96.
25. Marx, F.G., Tsai, C.H., and Fordyce, R.E. (2015). A new Early Oligocene
toothed ‘baleen’whale (Mysticeti: Aetiocetidae) from western North
America: one of the oldest and the smallest. R. Soc. Open Sci. 2,
150476.
26. Gioncada, A., Collareta, A., Gariboldi, K., Lambert, O., Di Celma, C., Bo-
naccorsi, E., Urbina, M., and Bianucci, G. (2016). Inside baleen: Excep-
tional microstructure preservation in a late Miocene whale skeleton from
Peru. Geol. 44, 839–842.
27. Peredo, C.M., Pyenson, N.D., and Boersma, A.T. (2017). Decoupling tooth
loss from the evolution of baleen in whales. Front. Mar. Sci. 4, 67.
28. Au, W.W.L. (1993). The Sonar of Dolphins (Springer-Verlag).
29. Churchill, M., Martinez-Caceres, M., de Muizon, C., Mnieckowski, J., and
Geisler, J.H. (2016). The origin of high-frequency hearing in whales. Curr.
Biol. 26, 2144–2149.
30. Fordyce, R.E. (1977). The development of the Circum-Antarctic Current
and the evolution of the Mysticeti (Mammalia: Cetacea). Palaeo. Palaeo.
Palaeo. 21, 265–271.
31. Marx, F.G., Lambert, O., and Uhen, M.D. (2016). Cetacean Paleobiology
(Chichester: John Wiley & Sons).
32. McGowen, M.R., Spaulding, M., and Gatesy, J. (2009). Divergence date
estimation and a comprehensive molecular tree of extant cetaceans.
Mol. Phylo. Evol. 53, 891–906.
33. Lambert, O., Martı́nez-Cáceres, M., Bianucci, G., Di Celma, C., Salas-Gis-
mondi, R., Steurbaut, E., Urbina, M., and de Muizon, C. (2017). Earliest
mysticete from the late Eocene of Peru sheds new light on the origin of
baleen whales. Curr. Biol. 27, 1535–1541.
34. Berger, W.H. (2007). Cenozoic cooling, Antarctic nutrient pump, and the
evolution of whales. Deep Sea Res. Part II 54, 2399–2421.
35. Marx, F.G., and Uhen, M.D. (2010). Climate, critters, and cetaceans:
Cenozoic drivers of the evolution of modern whales. Science 327,
993–996.
36. Pyenson, N.D., Irmis, R.B., and Lipps, J.H. (2010). Comment on ‘‘Climate,
Critters, and Cetaceans: Cenozoic Drivers of the Evolution of Modern
Whales’’. Science 330, 178–178.
37. Estes, J.A., Demaster, D.P., Doak, D.F., Williams, T.M., and Brownell, R.L.,
Jr. (2006). Whales, Whaling, and Ocean Ecosystems (Oakland: University
of California Press).
38. Lindberg, D.R., and Pyenson, N.D. (2007). Things that go bump in the
night: evolutionary interactions between cephalopods and cetaceans in
the tertiary. Lethaia 40, 335–343.
39. Boersma, A.T., and Pyenson, N.D. (2015). Albicetus oxymycterus, a new
generic name and redescription of a basal physeteroid (Mammalia, Ceta-
cea) from the Miocene of California, and the evolution of body size in
sperm whales. PLoS One 10, e0135551.
40. Ford, J.K., and Reeves, R.R. (2008). Fight or flight: antipredator strategies
of baleen whales. Mamm. Rev. 38, 50–86.
41. Wellard, R., Lightbody, K., Fouda, L., Blewitt, M., Riggs, D., and Erbe, C.
(2016). Killer whale (Orcinus orca) predation on beaked whales (Mesoplo-
don spp.) in the Bremer Sub-Basin, Western Australia. PLoS One 11,
e0166670.
42. Bianucci, G., Landini, W., and Buckeridge, J. (2006). Whale barnacles and
Neogene cetacean migration routes. NZ J. Geol. Geophys. 49, 115–120.
Current Biology 27, R558–R564, June 5, 2017 R563

43. Pichler, F.B., Robineau, D., Goodall, R.N.P., Meyer, M.A., Olivarria, C., and
Baker, C.S. (2001). Origin and radiation of Southern Hemisphere coastal
dolphins (genus Cephalorhynchus). Mol. Ecol. 10, 2215–2223.
44. Kelley, N.P., and Motani, R. (2015). Trophic convergence drives morpho-
logical convergence in marine tetrapods. Biol. Lett. 11, 20140709.
45. Pyenson, N.D., Gutstein, C.S., Parham, J.F., Le Roux, J.P., Chavarrı́a,
C.C., Little, H., Metallo, A., Rossi, V., Valenzuela-Toro, A.M., Velez-Juarbe,
J., et al. (2014). Repeated mass strandings of Miocene marine mammals
from Atacama Region of Chile point to sudden death at sea. Proc. R.
Soc. B. 281, 20133316.
46. Ha€ussermann, V., Gutstein, C.S., Beddington, M., Cassis, D., Olavarria,
C., Dale, A.C., Valenzuela-Toro, A.M., Perez-Alvarez, M., Sepúlveda,
H.H., McConnell, K.M., Horwitz, F.E., and Försterra, G. (2017). Largest
baleen whale mass mortality during strong El Niño event is likely related
to harmful toxic algal bloom. PeerJ Preprints 5, e2707v1.
R564 Current Biology 27, R558–R564, June 5, 2017
Current Biology
Minireview
47. Mirceta, S., Signore, A.V., Burns, J.M., Cossins, A.R., Campbell, K.L., and
Berenbrink, M. (2013). Evolution of mammalian diving capacity traced by
myoglobin net surface charge. Science 340, 1234192.
48. Pyenson, N.D., and Lindberg, D.R. (2011). What happened to gray
whales during the Pleistocene? The ecological impact of sea-level
change on benthic feeding areas in the North Pacific Ocean. PLoS One
6, e21295.
49. Pyenson, N.D., Ve
lez-Juarbe, J., Gutstein, C.S., Little, H., Vigil, D., and
O’Dea, A. (2015). Isthminia panamensis, a new fossil inioid (Mammalia, Ce-
tacea) from the Chagres Formation of Panama and the evolution of ‘river
dolphins’ in the Americas. PeerJ 3, e1227.
50. McCauley, D.J., Pinsky, M.L., Palumbi, S.R., Estes, J.A., Joyce, F.H., and
Warner, R.R. (2015). Marine defaunation: Animal loss in the global ocean.
Science 347, 1255641.