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The triple origin of whales

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Workers presume the traditional whale clade, Cetacea, is monophyletic when they support a hypothesis of relationships for baleen whales (Mysticeti) rooted on stem members of the toothed whale clade (Odontoceti). Here a wider gamut phylogenetic analysis recovers Archaeoceti + Odontoceti far apart from Mysticeti and right whales apart from other mysticetes. The three whale clades had semi-aquatic ancestors with four limbs. The clade Odontoceti arises from a lineage that includes archaeocetids, pakicetids, tenrecs, elephant shrews and anagalids: all predators. The clade Mysticeti arises from a lineage that includes desmostylians, anthracobunids, cambaytheres, hippos and mesonychids: none predators. Right whales are derived from a sister to Desmostylus. Other mysticetes arise from a sister to the RBCM specimen attributed to Behemotops. Basal mysticetes include Caperea (for right whales) and Miocaperea (for all other mysticetes). Cetotheres are not related to aetiocetids. Whales and hippos are not related to artiodactyls. Rather the artiodactyl-type ankle found in basal archaeocetes is also found in the tenrec/odontocete clade. Former mesonychids, Sinonyx and Andrewsarchus, nest close to tenrecs. These are novel observations and hypotheses of mammal interrelationships based on morphology and a wide gamut taxon list that includes relevant taxa that prior studies ignored. Here some taxa are tested together for the first time, so they nest together for the first time.
The triple origin of whales
Independent Researcher
311 Collinsville Avenue, Collinsville, IL 62234, USA
July 13, 2018
Keywords: Cetacea, Mysticeti, Odontoceti, Phylogenetic analyis
ABSTRACT—Workers presume the traditional whale clade, Cetacea, is monophyletic
when they support a hypothesis of relationships for baleen whales (Mysticeti) rooted on
stem members of the toothed whale clade (Odontoceti). Here a wider gamut phylogenetic
analysis recovers Archaeoceti + Odontoceti far apart from Mysticeti and right whales
apart from other mysticetes. The three whale clades had semi-aquatic ancestors with four
limbs. The clade Odontoceti arises from a lineage that includes archaeocetids, pakicetids,
tenrecs, elephant shrews and anagalids: all predators. The clade Mysticeti arises from a
lineage that includes desmostylians, anthracobunids, cambaytheres, hippos and
mesonychids: none predators. Right whales are derived from a sister to Desmostylus.
Other mysticetes arise from a sister to the RBCM specimen attributed to Behemotops.
Basal mysticetes include Caperea (for right whales) and Miocaperea (for all other
mysticetes). Cetotheres are not related to aetiocetids. Whales and hippos are not related to
artiodactyls. Rather the artiodactyl-type ankle found in basal archaeocetes is also found in
the tenrec/odontocete clade. Former mesonychids, Sinonyx and Andrewsarchus, nest
close to tenrecs. These are novel observations and hypotheses of mammal
interrelationships based on morphology and a wide gamut taxon list that includes relevant
taxa that prior studies ignored. Here some taxa are tested together for the first time, so
they nest together for the first time.
Marx and Fordyce (2015) reported the genesis of the baleen whale clade
(Mysticeti) extended back to Zygorhiza, Physeter and other toothed whales (Archaeoceti
+ Odontoceti). Earlier Fitzgerald (2006) and Fordyce and Marx (2016) discussed ‘basal
mysticetes’ with teeth. Even earlier Emlong (1966) described the newly discovered
Aetiocetus, “If it were not for the presence of functional teeth on this mature specimen,
this cetacean could easily be placed in the suborder Mysticeti.” Marx et al. (2016)
proposed a scenario in which “the transition from raptorial to baleen-assisted filter
feeding was mediated by suction, thereby avoiding the problem of functional interference
between teeth and the baleen rack.” Lambert et al. (2017) echoed this conjecture in their
description of toothy Mystacodon, a taxon they considered the earliest known member of
the Mysticeti. Thewissen (1994), Thewissen et al. (2007), and Thewissen et al. (2009)
supported the traditional view that “whales with legs”, like Maiacetus, were basal to all
hydropedal whales. Unfortunately, these studies did not include elephant shrews, tenrecs,
anthracobunids and desmostylians, taxa omitted from, but relevant to phylogenetic
studies of whales according to the present analysis (Fig. 1).
The monophyly of the traditional clade, Cetacea, has been challenged only rarely.
Kükenthal (1891) concluded, “We are justified in maintaining that the toothed whales are
of much earlier origin than whalebone whales, and that the terrestrial ancestors of the two
divisions were not identical.” Miller (1923), Yablokov (1965) and Zhemkova (1965)
thought odontocetes and mysticetes arose from different and unidentified ancestors. In
1968 Van Valen listed 20 traits that differ in odontocetes and mysticetes, but considered
cetaceans monophyletic and toothed Aetiocetus a mysticete. This has remained the
traditional view to the present day.
Here the taxon list of whale ancestors is expanded to include untested candidates.
The results of this wide-gamut, online phylogenetic analysis, commonly known as the
large reptile tree (= LRT;; subset in Fig. 1),
challenge the monophyly of the traditional clade ‘Cetacea.’ The LRT includes 1165
tetrapod taxa. With this wider gamut of taxa the two universally recognized clades of
whales, Odontoceti and Mysticeti, arise in parallel from distinct terrestrial mammal
clades (Fig. 1). Members of the Odontoceti are derived from aquatic archaeocetids,
pakicetids, tenrecs, terrestrial elephant shrews and anagalids in order of increasing
distance. Members of the Mysticeti are derived from aquatic desmostylians (Fig. 3),
anthracobunids, cambaytheres, hippos and terrestrial mesonychids. Right whales arise
from different desmostylians than other mysticetes do. This novel hypothesis of
interrelationships is well supported by fossils and extant taxa that document a gradual
accumulation of odontocete and mysticete traits in separate but convergent lineages. This
finds analogy in sirenians they also converge on whales in the loss of hind limbs and the
development of tail flukes. Purported transitional taxa, Aetiocetus (Emlong, 1966) and
Mystacodon (Lambert et al., 2017), nest with odontocetes, far from mysticetes in the
LRT. Janjucetus and Mammalodon nest with Anthracobune in the lineage of
desmostylians and mysticetes, far from odontocetes. Cetotheres were considered basal
mysticetes (Lambert et al. 2017) due to their resemblance to aetiocetids. Here the small
mysticetes, Caperea and Miocaperea, nest as basal mysticetes based on traits shared with
DNA studies do not include older fossils. Frequently molecular studies recover
family tree topologies that do not match those of morphological studies (Fordyce and de
Muizon, 2001). Molecular studies do not provide a gradual accumulation of physical
traits in order to check that evolutionary changes are indeed tenable.
The evolutionary history of cetacean swimming from cursorial mesonychids
promoted by Thewissen and Fish (1997) used the dorsoventral undulation of otters as a
living analog for unknown transitional taxa. They assumed a monophyletic Cetacea and
made no reference to tenrecs or desmostylians. They also expressed some concern in the
transition from a cursorial mesonychid to a hydropedal whale, ironically without
invoking the cursorial and aquatic hippo. Fordyce and de Muizon (2001) discussed
similar issues without an adequate phylogenetic framework.
The problem with all prior studies has been taxon exclusion. No prior studies
correctly identified the ancestors of pakicetids and mysticetes. Here all potential
candidates for whale ancestry are tested going back to Devonian tetrapods. Here the 3x
convergent loss of hind limbs and acquisition of tail flukes in the three ‘whale’ clades is
based on a phylogenetic framework with very short ghost lineages. The present study also
reveals several overlooked transitional traits and vestiges.
The key advancement provided by the present tree topology (Fig. 1) is taxon
inclusion. Prior workers with first hand access to whale specimens omitted relevant taxa.
Over 1150 candidates for mysticete and odontocete ancestry were tested here using
published images for most of the data. Results indicate that the exclusion of relevant taxa
is a greater problem than lacking first hand access. The present list of 1165 taxa
minimizes bias and tradition in the process of selecting ingroup and outgroup taxa for
smaller, more focused studies because all major and many minor tetrapod clades are
tested here. That means that all derived clades, including every tested whale, have
outgroups extending back to Late Devonian tetrapods.
No characters used in the LRT are specific to the clades that include whales and
their proximal ancestors. Although some characters are similar to those from various
prior analyses, the present list (see Supporting Data; to be
completed when the ms. is accepted) was largely built from scratch. Traits specific to
turtles or pterosaurs would have been useless on whales and tree shrews and vice versa.
Generalized characters were chosen or invented for their ability to lump and split clades
and for their visibility in a majority of tetrapod taxa, many of which had never been
tested together. Up to this point, the 231 multi-state character set has proven sufficient to
lump and separate 1165 taxa, typically with high Bootstrap scores. In the past, certain
workers considered 231 characters too small for the number of tested taxa—when the
taxon list was a quarter of the size it is now. Others thought the characters themselves
were less than optimally fashioned. Not all opinions can be accommodated given the
constraints of a single lifetime. Complete resolution in the LRT tree and high Bootstrap
scores falsify any blackwashing levied against the present character list. For all of its
faults, real or imagined, the LRT continues to work well as more taxa are added every
week. All taxon subsets of the LRT (e.g. Fig. 1) raise the character/taxon ratio. All taxa in
the LRT are generic, specific or species-based. Chimaeras are not employed.
Taxa and characters were compiled in MacClade 4.08 (Maddison and Maddison,
1990), then imported into PAUP* 4.0b (Swofford, 2002) and analyzed using parsimony
analysis with the heuristic search algorithm. All characters were treated as unordered and
no character weighting was used. Bootstrap support figures were calculated for 100
replicates. The cladogram, character list and data matrix accompany the manuscript and
will be available in permanent repository here: xxxxx and (to be completed when the ms. is accepted).
Abbreviations: mya = million years ago; IVPP = Institute of Vertebrate
Paleontology, Peking; NMV = Museum Victoria, Melbourne, Australia; RBCM = Royal
British Columbia Museum, Victoria, Canada; UHR = University of Hokkaido
Registration, Sapporo, Japan.
The LRT nests all tested taxa in near-complete resolution (2 MPTs with loss of
resolution at the incomplete fossil of Maelestes). High Bootstrap scores are typically
recovered. The traditional clades, Odontoceti and Mysticeti, nest apart from one another
(Fig. 1). Both odontocetes and mysticetes had limbed precursors. Right whales descend
from different desmostylians than all other mysticetes do. The last common ancestor of
all whales is a sister to the tiny tree shrew-like taxo, Maelestes (Late Cretaceous; Wible,
et al., 2007a, b).
The tenrec/odontocete clade
In the LRT (subset Fig. 1) odontocetes are derived from a placental mammal
clade that had its origin with tiny, formerly arboreal insectivores, like Maelestes. Rabbit-
sized Anagale (early Oligocene; Simpson, 1931) and Leptictis (early Oligocene; Leidy,
1868; Rose, 2006) split off next followed by the long-legged elephant shrew,
Rhynchocyon (extant). The other extant elephant shrew, Macroscelides nested elsewhere.
Former mesonychids, wolf-sized Sinonyx (late Paleocene; Zhou et al., 1995) and giant
Andrewsarchus (middle Eocene; Osborn, 1924), split off next, followed by the tenrec
clade. Mesonyx and other mesonychids nested elsewhere (see below). Bajpai and
Gingerich (1998) associated Himalayacetus (early Eocene;) with the lineage of toothed
whales. It is known from an incomplete dentary comparable to that of Sinonyx and is not
tested here.
In the LRT the tenrec clade includes Hemicentetes (extant) and Tenrec (extant,
formerly Centetes, Fig. 2), plus cat-sized Leptictidium (Early Eocene; Tobien 1962) and
Indohyus (Eocene; Rao, 1971). The shrew-like ‘tenrecs’, Limnogale, Microgale,
Micropotamogale and Potamogale, nested elsewhere and apart from the hedgehog-like
‘tenrec,’ Echinops.
At succeeding nodes, wolf-sized Pakicetus (Eocene) was considered “one of the
oldest whales known anywhere” (Gingerich and Russell, 1981) upon its discovery.
Larger Maiacetus (Eocene, Gingerich, et al., 2009) was unable to locomote on land due
to a much longer torso, hydropedal forelimbs and vestigial hind limbs. It had a more
robust tail, yet retained a skull similar to that of Tenrec (Fig. 2). Larger still, Mystacodon
(Late Eocene; Lambert et al., 2017) and Zygorhiza (Late Eocene; True, 1908) was more
fully hydropedal based on its relatively smaller pelvis. Aetiocetus (Oligocene; Emlong,
1966), Chonecetus (Oligocene; Russell, 1968; = Fucacia Marx et al., 2015) and NMV
P252567 (Oligocene, Marx et al., 2016) were originally considered basal to the clade
Mysticeti due to their resemblance to cetotheres. With more taxa these three nest together
between Zygorhiza and the two tested extant odontocetes, Orcinus and Physeter. Many
dozen species of extant and extinct toothed whales are known and could have been added
to this taxon list, but the focus here is on basal taxa with terrestrial traits.
The mesonychid/mysticete clade
In the LRT mysticetes are derived from a mammal clade that had its origin with
terrestrial Mesonyx (Eocene; Cope, 1872; Van Valen, 1966), derived from Paleocene or
older ungulates (see below and Fig. 1). Phylogenetically Mesonyx is followed by semi-
terrestrial Ocepeia (Paleocene; Gheerbrant et al., 2001, 2014) and Hippopotamus
(extant). The next split produced Cambaytherium (Eocene; Rose et al., 2014) and
Cornwallius (Early Oligocene; Cornwall 1922; Hay 1923, Beatty 2006a, b). The next
split produced Anthracobune (Eocene; Pilgrim, 1940), Mammalodon (late Oligocene;
Pritchard, 1939) and Janujucetus (late Oligocene; Fitzgerald 2006) in a clade. The basal
desmostylians followed. These include Neoparadoxia (Barnes 2013; Miocene),
Paleoparadoxia (Miocene; Reinhart, 1959), Desmostylus (Oligocene; Marsh, 1888;
Inuzuka, 2009; Uno and Kimura, 2004) and the RBCM.EH2007.008.0001 specimen
attributed to Behemotops (Oligocene; Domning, Ray and McKenna, 1986; Beatty and
Cockburn, 2015). The two tested right whales (Eubalaena and Caperea) nested with
Desmostylus. The remaining mysticetes nested with the RBCM specimen attributed to
Regaining a monophyletic ‘Cetacea’
In order to attract members of the Mysticeti to the strongly convergent
Odontoceti, only two mysticete outgroup taxa need to be removed: Desmostylus and the
RBCM specimen attributed to Behemotops. When that happens the mysticetes nest with
Physeter and Orcinus. Conversely, in order to attract extant members of the Odontoceti
to the Mysticeti, four odontocete outgroup taxa need to be removed: Zygorhiza and the
three aetiocetids. When that happens Physeter and Orcinus nest with the mysticetes
leaving Maiacetus and Mystacodon nesting with the pakicetids and tenrecs.
The traditional clades ‘Cetacea’ (Brisson, 1762) and ‘Cetartiodactylia’ are no
longer monophyletic in the LRT. The traditional clade ‘Artiodactylia’ is found to be
polyphyletic in the LRT—unless Hippopotamus and all whales are omitted. The
traditional clade ‘Whippomorpha’ (Cetacea + Hippopotamidae; Waddell, Okada and
Hasegawa, 1999) is no longer monophyletic in the LRT. The clade Neoceti (Odontoceti +
Mysticeti, Uhen, 2008) is no longer monophyletic. The clade Pelagiceti (Basilosauridae +
Neoceti, Uhen, 2008) is likewise no longer monophyletic.
The traditional clade Mesonychia is here expanded to include Mesonyx, Sus, their
last common ancestor and all of their descendants. In the LRT that list includes hippos,
desmostylians, mysticeti, artiodactyls, elephants, sirenians, chalichotheres and
perissodactyls. These are all decendants of a sister to Mesonyx.
A more restricted new clade, Mesonyketos (“middle claw-sea monster”), is
proposed to include Mesonyx, Hippopotamus, their last common ancestor and all of their
descendants. That clade includes the Desmostylia and both mysticete clades (Fig. 1). The
Desmostylia is no longer an extinct clade.
The clade Mysticeti (Cope, 1891) traditionally includes all baleen whales. In
order to remain monophyletic it must also include tested members of the Desmostylia
(Reinhart, 1959). This makes ‘Desmostylia’ a junior synonym for Mysticeti unless
applied to just Neoparadoxia and Paleoparadoxia, among tested taxa.
A new clade Tenreketos (“tenrec-sea monster”; from French tanrec, from
Malagasy tàndraka, plus Greek ketos) is proposed for Maelestes, Tenrec, their last
common ancestor and all of its descendants. That clade includes the traditional clade
Archaeoceti, which now includes the smaller clade Odontoceti.
A new clade, Edafosia (“ground” in Greek), is proposed for Maelestes,
Phenacodus, their last common ancestor and all of its descendants. In the LRT this clade
includes mammals that plesiomorphically became ground dwellers. Edafosia is the
smallest clade that contains all ‘whales.’
Many fossil taxa document transitional phases in the evolution of so-called ‘land
whales’ to hydropedal taxa (Thewissen et al., 2007, 2009). That lineage traditionally
begins with Pakicetus, Indohyus and related taxa. Here the ancestors of Pakicetus and
Indohyus are recovered for the first time, and they are not artiodactyls.
In the LRT, Maelestes (skull 2 cm long) nests at the base of the Tenreketos (Fig.
1). Similar in size and shape to extant tree shrews, Maelestes was originally (Wible et al.
2007a, b) allied with Asioryctes far outside of the Placentalia, but deep within the
Eutheria in a cladogram with few taxa in common with the LRT. The Wible et al. (2007a,
b) cladograms excluded taxa nesting here with Maelestes: IVPP V2385 (Ting et al.,
2004) and Anagale. The LRT includes Asioryctes, which nests a few nodes outside the
last common ancestor of all placental mammals, as in Wible et al. (2007a, b). On Anagale
the primordial nuchal crest and elongate hoof-like unguals are traits retained by many
members of the Tenreketos.
Like the related elephant shrew, Rhynchotus, the torso and tail of Leptictis (skull 6
cm long) were short and the legs were long with digitigrade extremities and a semi-
circular astragalus analogous with those of basal artiodactyls. The long rostrum is
retained in descendant taxa.
The next clade includes much larger predatory taxa, Sinonyx (skull 28 cm) and
Andrewsarchus (skull 83 cm long). Both had larger canines and parietal crests, traits
retained by descendant taxa.
Two extant tenrecs split off next, Tenrec (Fig. 2) and Hemicentetes. Both have
tiny tails, a derived trait. Related extinct taxa, Indohyus and Lepticitidium retained and
further developed long tails. Both were the first aquatic taxa in the Tenreketos. Today
tenrecs are found in Madagascar. Indohyus fossils are found in Kashmir. These
landmasses and their occupants split apart 89–85 million years ago (McKenzie Sclater,
1971; de Wit, 2003), pushing this node back twenty million years before the Late
Cretaceous extinction event.
Hemicentetes is known to make short duration tongue clicks (5000–17,000 cps)
that aid in echolocation (Gould, 1965). Odontocetes echolocate by producing short
duration clicks using ‘phonic lips’ located in the melon along the nasal passage outside of
the skull (Cranford, 2000). Given the close phylogenetic relationship of tenrecs and
odontocetes in the LRT, echolocation in odontocetes likely originated with Late
Cretaceous tenrecs similar to Hemicentetes and Indohyus.
Tenrecs typically travel and feed in family/social groups of kinship litters (Gould
and Eisenberg, 1966). Many skeletons of Indohyus were washed together, buried in
freshwater stream sediments (Thewissen et al., 2009). Given their close phylogenetic
relationship in the LRT, these kinship litters may be retained as pods in living
In the semi-aquatic tenrec, Indohyus, the tail is long, but not longer than the hind
limb. The hind limb is slightly longer than the fore limb. The paddle-like pes is
substantially larger than the manus. The limbs are made of denser bone with less marrow,
making them better suited to wading and swimming in lake shallows (Thewissen, et al.
2009). Unique among terrestrial taxa, the middle ear has a thickened internal lip, as found
in cetaceans. Further evidence for an aquatic habitat comes from the tooth chemistry of
Indohyus (Thewissen, et al. 2009). Based on its artiodactyl-like ankle, Indohyus was
earlier assumed to be a small deer-like herbivorous artiodactyl (Thewiessen, et al. 2007).
Phylogenetically that is a problem for a piscivorous odontocete ancestor, but not if that
ancestor is an insectivorous-grading-to-piscivorous aquatic tenrec. Thewiessen et al.
(2009) reported, “This shape of the astragalus, with a proximal trochlea (hinge joint) as
well as distal trochlea, only occurs in even-toed ungulates (artiodactyls).” An overlooked
convergent shape is also found in Rhynchocyon and Leptictidium, which also have an
ungulate-like digitigrade pes with hooves and elongate metatarsals. The unguals of
Indohyus were described (Thewiessen et al. 2009) as hooves, elongate and tapering, but
with an expanded tip. Such hooves are also present in Anagale.
Leptictidium had smaller fore limbs, elongate hind limbs and a muscular tail much
longer than its hind limbs. The skull and dentition are close matches to Tenrec and
Maiacetus. Originally considered a bipedal omnivore and saltator, Leptictidium had a
‘loose’ sacroiliac joint, different from those in typical saltators, but similar to those found
in so-called ‘land whales.’ Phylogenetic bracketing, comparative morphology and the
fossil’s lacustrine matrix indicate Leptictidium was aquatic, like its sister Indohyus. It
could swim by paddling those long hind legs together, coordinated with dorsoventral
undulations of that long, muscular tail. This pattern of swimming is further refined in
toothed whales.
More derived toothed taxa linking Pakicetus to archaoecetes and odontocetes are
well documented (e.g. Thewissen, et al. 2009), but in short: the teeth become simple
pegs, the external naris and lacrimal migrate over the cranium, the tail enlarges and
develops flukes, the forelimbs become flippers, the pelvis is reduced and the hind limbs
become internal vestiges.
The origin of the traditional clade Mysticeti has been called ‘a baffling problem’
(Fitzgerald, 2006) largely because relevant taxa have not been included in prior whale
analyses (e.g. Marx and Fordyce, 2015). Here that problem is resolved with the addition
of previously omitted taxa.
Mesonyx (30 cm skull length) nests at the base of the Mesonyketos. In the LRT
mesonychids were derived from basal ungulates of similar size, like Phenacodus (late
Paleocene). Distinct from Phenacodus, Mesonyx had a larger skull with a higher parietal
crest and larger canines. The mandibular joint was lower. The medial digits were
vestigial. Phylogenetic bracketing nests Mesonyx in the midst of many large herbivores,
so those large canines were likely used for display and fighting, as in the related
Hippopotamus, than against prey. Exceptionally meat-eating is known in hippos (Dudley,
1996, 1998).
The first taxon to split off is the neotonous Ocepeia (9 cm skull). Retaining
juvenile traits into adulthood, it had lower skull crests and smaller canines. The orbits
were raised to the top of the skull, as in related hippos. The appearance of Ocepeia in the
Paleocene argues for an earlier appearance of mesonychids than the Eocene.
Hippopotamus (70 cm skull) splits off next. This extant, graviportal herbivore is
more at home in the water, but still able to run on land. The orbits are elevated above the
elongated and laterally expanded rostrum. The lower incisors are elongate and oriented
anteriorly. The ribcage is expanded ventrally and posteriorly, reducing the lumbar region.
The tail is a vestige. Hair is nearly absent, but a deep layer of fat is present. Nursing and
communication takes place underwater. Some of these traits are retained in descendant
Next in the lineage of mysticetes, Cambaytherium (35 cm skull), was originally
considered a basal perissodactyl close to anthracobunids (Rose et al., 2014). Fossils were
found on the marine coastline of western India, coeval with Pakicetus. Cambaytherium
did not have the dorsal orbits and elongate muzzle of Hippopotamus, but it had a large
retroarticular process on the dentary, a trait retained in succeeding taxa. The teeth were
all more similar in size, lacking giant canines. The putative desmostylian, Cornwallius
(Hay 1923, Cornwall 1922, Beatty 2006a, b; Early Oligocene) nests with Cambaytherium
in the LRT. Adults had a downturned rostrum, as in desmostylians. Juveniles did not, as
in anthracobunids. The procumbent incisors and canines of Cornwallius were separated
from the suborbital desmotylian-grade molars by a long diastema (the paraglossal crest).
This is the first step toward toothlessness in the lineage of mysticetes.
At the next split, Anthracobune (20 cm skull) was originally considered a
proboscidean (Pilgrim, 1940), then a perissodactyl (Cooper et al., 2014) before nesting in
the LRT between cambaytheres and desmostylians. Anthracobune also nests at the base
of a small clade that includes toothy Mammalodon and Janjucetus, taxa known
principally from skulls. Both were earlier restored as hydropedal stem mysticetes
(Fitzgerald, 2006; Fordyce and Marx, 2016), but phylogenetic bracketing in the LRT
indicates these taxa must have had robust limbs. Here the naris opens more dorsally than
anteriorly, as in desmostylians and mysticetes.
The basal desmostylian, Neoparadoxia (Barnes, 2013) splits off next. Prior
studies (Reinhart, 1959; Barnes, 2013) nested desmostylians with sirenians and elephants.
Like hippos the orbit of Neoparadoxia is elevated. The wide rostrum is downturned and
includes a long diastema. The tail is a vestige, but the manus and pes are broad enough to
swim with, powered by dorsoventral undulations of the spine and hind limbs (Gingerich,
2005). This behavior would be retained by mysticetes, analogous to the aquatic taxa in
the clade Tenreketos.
Closely related Paleoparadoxia (50 cm skull), splits off next. The skull is at least
twice as wide as tall, as in mysticetes. The orbits are not elevated.
Desmostylus (35 cm skull; Marsh, 1888; Domning, Ray and McKenna, 1986; Uno
and Kimura, 2004; Inuzuka 2009) nests at the base of the clade that includes the right
whales, Eubalaena and Caperea. Desmostylus has fewer and smaller teeth. The rostrum
is narrower than the mandibles, a trait exaggerated in right whales. The wider, flatter
cranial roof of Desmostylus is retained in descendant right whales. The neck is
compressed to less than half the skull length. As in Caperea, the lumbar region is reduced
to two or three vertebrae in Desmostylus. The seven preserved coccygeal and caudal
vertebrae are small and flat, extending not much further than the posterior ilium. The
metacarpals are flattened, as in mysticete flippers. Compared to Paleoparadoxia the
limbs in Desmostylus are relatively smaller relative to the torso and the hind limbs are
shorter than the fore limbs. Distinct from most mammals, the humerus and femur have a
sprawling configuration.
The extant pygmy right whale, Caperea marginata (Gray, 1846), was recently
considered ‘the last of the cetotheres’ (Fordyce and Marx, 2013), but only in the absence
of desmostylians. Here it returns to its traditional nesting with the much larger, extant
right whale, Eubaelana. Like Desmostylus, Caperea has seven robust post-sacral
vertebrae plus eight smaller caudals between the flukes. The great reduction of the pelvis
in Caperea changed the coccygeal vertebrae into caudal vertebrae, therby producing a
longer practical tail. Caperera is toothless, with baleen deeper than its open and unfused
mandibles. Like the right whales, it is a ram-feeder on calanoid copepods. The sternal
elements and manus were reduced to vestiges compared to Desmostylus. Bisconti (2012)
noted, “Given that C. marginata possesses a mix of balaenid and balaenopterid
characters, it is difficult to understand which features are the result of convergence and
which are those representing the proof of true phylogenetic relationships.” The LRT
resolves this issue by nesting plesiomorphic Caperea near the base of both major
mysticete clades.
In the giant right whale, Eubalaena, the jaws are permanently open for ram
feeding and giant lower lips rise to close off the sides of the mouth. The lacrimals and
frontals extend laterally, matching the wide mandibles. In Eubalaena the 16 caudals in
series are not longer than the lumbar series, now increased to nine vertebrae. Distinct
from all other members of the Mesonyketos, Caperea and Eubalaena redevelop a tiny
manual digit 1.
In the LRT, the taxon Behemotops (40 cm skull) is scored based on the narrow
skull of the RBCM.EH2007.008.0001 specimen (Fig. 3; Beatty and Cockburn, 2015),
which does not match several wide and toothy dentaries previously assigned to this genus
(Domning, Ray and McKenna, 1986). A better fit to the concave maxilla of the RBCM
specimen is found in the elongate dentary of the Sanjussen specimen cf. of Vanderhoofius
sp. (UHR32380, Fig. 3, Uno and Kimura, 2004; Chiba et al., 2015), which does not have
a strong medial symphysis, as in mysticetes. Behemotops was originally considered the
most primitive desmostylian based on wide toothy mandibles, but the RBCM specimen
nests between Desmostylus and all other (non-right whale) mysticetes in the LRT. The
post-crania of the RBCM specimen are poorly known: some dorsal vertebrae, a distal
scapula and a large humerus that could be semi-terrestrial or hydropedal.
Distinct from all prior whale studies, toothless Miocaperea (Bisconti 2012; late
Miocene, 7–8 Ma; 1m skull length) nests at the base of all tested non-right whale
mysticetes, not with Caperea (contra Bisconti 2012, who omitted desmostylians).
Miocaperea is known from a skull three times the length of the Behemotops skull, but
broadly similar is morphology. Short patches of baleen are preserved. Like hydropedal
mysticetes, the orbit migrates posteriorly. A vestige of the jugal appears on the anterior
tip of the squamosal. Compared to the RBCM specimen of Behemotops, maxillary tusks
are absent in Miocaperea. The frontals, lacrimals and squamosals are laterally expanded.
The parietals do not appear to be roofed over by the supraoccipital to the extent
illustrated by Bisoconti (2012). Rather the supraoccipitals appear to extend no further
than the anteriormost extent of the squamosal as in Isanacetus.
In the LRT taxa preceding the RBCM specimen of Behemotops and Miocaperea
have robust limbs with free fingers. Taxa succeeding these taxa have hydropedal
forelimbs without free fingers and vestigial hind limbs that do not emerge from the body
wall. The transition from one body type to the other occurred between these two taxa.
With Behemotops in the Early Oligocene and Miocaperea in the Late Miocene, about 20
million years is available for this transition, unless these two are late survivors of an
earlier radiation.
Higher tested mysticetes split between cetotheres (Cetotherium + (Tokarahia +
Yamatocetus) and Eschrichtius + rorquals + (Isanacetus + Balaeonoptera). Cetotherium
has 15-16 caudals and the series is no longer than the lumbar section of the torso.
Other than Miocaperea, Eschrichtius appears to be the most primitive member of
this clade. Relicts of procumbent desmostylian tusk alveoli are present at the anterior tips
of the dentaries and maxillae (Fig. 3). Blood vessels and nerves still pass through these
openings as they did when tusks were present. Eschrichtius is the only baleen whale that
still scoops up sediments from the sea floor, similar to behavior imagined for shovel-
tusked desmostylians and, by phylogenetic bracketing, Miocaperea. Eschrichtius has 28
caudals nearly equal to its entire thorax length.
Isanacetus laticephalus (Kimura and Ozawa, 2002) has a skull similar in size to
that of Miocaperea. The rostrum and frontals are wider. The orbit is stationed more
anteriorly. The naris and nasals are narrower. The posterior squamosal descends and the
parietal raises diagonal nuchal crests. In palatal view the maxillae are in contact medially.
Extinct cetotheres, like Yamatocetus (Early Oligocene), Tokaraharia (late
Oligocene) and Cetotherium (late Miocene) had relatively straight jaw rims with a high
cornoid process, distinct from the more plesiomorphic ventrally concave maxillae of
rorquals, right whales and Behemotops. Thus cetotheres were not ancestral to extant
mysticetes and do not nest as transitional taxa arising from the toothed archaeocete,
Aetiocetus (contra Emlong, 1966; Van Valen 1968; Geisler et al. 2011).
Embryological studies
In a study of embryo bowhead whales (genus: Balaena), Thewissen et al. 2017
noted the rack of baleen was “implanted more or less where the tooth rows would be, but
there is no trace of teeth.” By following the hypothesis of an archaeocete origin for
mysticetes, they came to realize, “The pattern of dental evolution in mysticetes is thus
counterintuitive, first the number of teeth increases in evolution but then teeth disappear
altogether suddenly.” By contrast, in the present hypothesis where mysticetes evolve
from desmostylian ancestors, adult teeth and tusks disappear in the jaws gradually and
leave traces of their departure.
Thewissen et al., 2017 reported 41 upper and 35 lower tooth ‘caps’ in each jaw of
an embryo bowhead whale. Where do such large numbers come from? In the LRT there
are no tetrapods in the lineage of whales with more than 30 teeth in the maxilla, until one
extends the search to the pre-tetrapods, Tiktaalik and Panderichthys.
Peredo et al. (2017) provided a comprehensive review of the literature on tooth
buds and concluded, “Based on the available range of evidence, the origin and evolution
of baleen in mysticetes defies simple explanations.” The Peredo team did not consider
desmostylians, but held to the archaeocete hypothesis of mysticete origins.
In the LRT, the origin of baleen in mysticete whales can be traced to the
narrowing of the rostrum, the widening of the mandibles, the disappearance of the
premolars (= appeance of the long diastema), the reduction of all teeth and tusks along
with the increasing lateral exposure of the palatal portion of the ventrally concave maxilla
in desmostylians. That’s the simple explanation that comes with taxon inclusion.
DNA and supermatrix studies
Gatesy (1997) used molecules to nest hippos with whales (Balaenopteridae +
(Delphinoidea + Physeteridae)). A long list of artiodactyls nested elsewhere. Their short
tree topology matches the LRT sans tenrecs. Tenrecs were not tested by Gatesy (1997).
Geisler et al., (2011) created a supermatrix of traits, but omitted tenrecs and
desmostylians from their taxon list. By default (due to taxon exclusion) the
anthracobunids, Janjucetus and Mammalodon, nested between the toothed whales,
Zygorhiza and (Chonecetus + Aetiocetus). These are wrongly considered the last taxa
with teeth in the Geisler et al. lineage of mysticetes. There is no demonstrated gradual
loss of teeth in the Geisler et al. lineage. Their basalmost mysticetes include the flat-
jawed and toothless cetotheres Eomysticetus, Micromysticetus, Diorocetus and Pelocetus
nesting prior to the extremely derived, bow-skulled right whale, Eubaleaena. Once again,
there is no gradual accumulation of traits between transitional taxa in the Geisler et al.
cladogram. The primitive gray whale, Eschrichtius, nests at the third derived node in their
Mysticeti. So, their cladogram essentially reverses the order of mysticetes recovered by
the LRT, putting derived taxa in basal nodes and vice versa.
The addition of relevant taxa nests odontocetes and mysticetes in two clades
derived from predatory and non-predatory limbed ancestors. This invalidates the results
of earlier, smaller studies that nested mysticetes with odontocetes, archaeocetes and
artiodactyls when relevant taxa were excluded. Here the Odontoceti arise from aquatic
echolocating tenrecs and their kin. Here the Mysticeti arise from increasingly toothless
desmostylians and their kin. This report documents the gradual accumulation of derived
traits that led to the Odontoceti and the largely convergent Mysticeti.
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Figure captions
Figure 1.
Subset of the large reptile tree, (= LRT;,
focusing on the two whale clades, Odontoceti and Mysticeti and their proximal
outgroups. See link above for complete taxon list.
Figure 2.
Comparative lateral views of two taxa in the tenrec/odontocete clade. Above: The extant
tenrec, Tenrec. Below: The much larger ‘whale with legs’, Maiacetus (Eocene). Inside
the jaws of Maicetus is the skull of Tenrec to scale.
Figure 3.
Three taxa in the mesonychid/mysticete clade at the transition from desmostylians to
mysticetes: Left column: The RBCM skull originally attributed to Behemotops in dorsal,
lateral and palatal views along with the Sanjussen mandible scaled down to fit the skull
(dark gray). Bottom left column: The Sanjussen mandible specimen to scale with the
RBCM skull. Middle column: The gray whale (genus: Echrichtius) in anterior view.
Arrows point to former tusk alveoli. Lateral view of Isanacetus and the RBCM specimen
to scale. Palatal view of same with baleen in dark gray. Right column: The small baleen
whale, Isanacetus, skull in dorsal, lateral and palatal views. More extensive nutrient
foramina here root baleen to the palate.
Figure 4.
The reduction/retreat of the canine tusk in Desmostylus is shown here (upper arrow).
Lower arrow points to dentary tusks. Compare to Eschrichtius in figure 3.
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Full-text available
In utero, baleen whales initiate the development of several dozens of teeth in upper and lower jaws. These tooth germs reach the bell stage and are sometimes mineralized, but toward the end of prenatal life they are resorbed and no trace remains after birth. Around the time that the germs disappear, the keratinous baleen plates start to form in the upper jaw, and these form the food-collecting mechanism. Baleen whale ancestors had two generations of teeth and never developed baleen, and the prenatal teeth of modern fetuses are usually interpreted as an evolutionary leftover. We investigated the development of teeth and baleen in bowhead whale fetuses using histological and immunohistochemical evidence. We found that upper and lower dentition initially follow similar developmental pathways. As development proceeds, upper and lower tooth germs diverge developmentally. Lower tooth germs differ along the length of the jaw, reminiscent of a heterodont dentition of cetacean ancestors, and lingual processes of the dental lamina represent initiation of tooth bud formation of replacement teeth. Upper tooth germs remain homodont and there is no evidence of a secondary dentition. After these germs disappear, the oral epithelium thickens to form the baleen plates, and the protein FGF-4 displays a signaling pattern reminiscent of baleen plates. In laboratory mammals, FGF-4 is not involved in the formation of hair or palatal rugae, but it is involved in tooth development. This leads us to propose that the signaling cascade that forms teeth in most mammals has been exapted to be involved in baleen plate ontogeny in mysticetes.
Full-text available
The origin of baleen, the key adaptation of modern whales (Mysticeti), marks a profound yet poorly understood transition in vertebrate evolution, triggering the rise of the largest animals on Earth. Baleen is thought to have appeared in archaic tooth-bearing mysticetes during a transitional phase that combined raptorial feeding with incipient bulk filtering. Here we show that tooth wear in a new Late Oligocene mysticete belonging to the putatively transitional family Aetiocetidae is inconsistent with the presence of baleen, and instead indicative of suction feeding. Our findings suggest that baleen arose much closer to the origin of toothless mysticete whales than previously thought. In addition, they suggest an entirely new evolutionary scenario in which the transition from raptorial to baleen-assisted filter feeding was mediated by suction, thereby avoiding the problem of functional interference between teeth and the baleen rack.
Full-text available
A small, toothed fossil cetacean from Hakataramea Valley (South Canterbury, New Zealand) represents a new Late Oligocene species, Mammalodon hakataramea. The new material is from the Kokoamu Greensand (Duntroonian Stage, about 27 Ma, early to middle Chattian) of the Canterbury Basin, and thus about 2 Ma older than the only other species included in this genus, Mammalodon colliveri (Late Oligocene, Victoria, Australia). The anterior pedicle of the tympanic bulla is not fused to the periotic and resembles that of Delphinidae in basic structure. The teeth show extreme attritional and/or abrasive wear, which has obliterated the crowns. Like Mammalodon colliveri, M. hakataramea was probably raptorial or a benthic suction feeder.
Although combined molecular and morphological analyses point to a late middle Eocene (38-39 million years ago) origin for the clade Neoceti (Odontoceti, echolocating toothed whales plus Mysticeti, baleen whales, and relatives), the oldest known mysticete fossil dates from the latest Eocene (about 34 million years ago) of Antarctica [1, 2]. Considering that the latter is not the most stemward mysticete in recent phylogenies and that Oligocene toothed mysticetes display a broad morphological disparity most likely corresponding to contrasted ecological niches, the origin of mysticetes from a basilosaurid ancestor and its drivers are currently poorly understood [1, 3-8]. Based on an articulated cetacean skeleton from the early late Eocene (Priabonian, around 36.4 million years ago) of the Pisco Basin, Peru, we describe a new archaic tooth-bearing mysticete, Mystacodon selenensis gen. et sp. nov. Being the geologically oldest neocete (crown group cetacean) and the earliest mysticete to branch off described so far, the new taxon is interpreted as morphologically intermediate between basilosaurids and later toothed mysticetes, providing thus crucial information about the anatomy of the skull, forelimb, and innominate at these critical initial stages of mysticete evolution. Major changes in the morphology of the oral apparatus (including tooth wear) and flipper compared to basilosaurids suggest that suction and possibly benthic feeding represented key, early ecological traits accompanying the emergence of modern filter-feeding baleen whales' ancestors.