Supporting online information to:
Anatomy of a new cetotheriid genus and species from the Miocene of Herenthals, Belgium,
and the phylogenetic and paleobiogeographic relationships of Cetotheriidae s.s.
(Mammalia, Cetacea, Mysticeti)
Michelangelo Bisconti*a
a
San Diego Natural History Museum, 1588 El Prado, 92101, San Diego, California.
Index
1. Taxonomic dataset (p. 2).
2. Morphological dataset (p. 7).
3. Matrix for phylogenetic analysis (p. 19).
4. Table S1. Ages and areas of occurrence of Cetotheriidae s.s. taxa and Diorocetus and their
translations into Operational Geographic Units as used in the paleobiogeographic analysis
(p. 25).
5. Table S2. Matrix generated for the Secondary BPA and used in the paleobiogeographic
analysis (p. 26).
6. References (p. 27).
*
Corresponding author. E-mail: michelangelobisconti@gmail.com
1. Taxonomic dataset
Several specimens were examined in order to get a complete coverage of the morphological
diversity of the mysticete skeleton. Four archaeocetes are also included in the dataset: i.e.,
Protocetus atavus (Fraas 1904), Georgiacetus goedertorum (Hulbert et al. 1996), Dorudon atrox
(Kellogg 1936; Uhen 1998, 2004), and Zygorhiza kochii (Kellogg 1936; Uhen 1998) as they are
used as outgroup taxa. All the remaining taxa included in the dataset form the ingroup and are
assigned to the suborder Mysticeti.
The taxa and the specimens examined are reported in below.
The mysticetes included in the database are represented by both toothed and baleen-bearing taxa.
The toothed mysticetes include Mammalodon colliveri (Fitzgerald 2010) and Aetiocetus weltoni
(Deméré & Berta 2008). The baleen-bearing mysticetes include representatives from all the
mysticete radiations. The Late Oligocene Eomysticetoidea are represented by Eomysticetus
whitmorei and Micromysticetus rothauseni (Sanders & Barnes 2002a, b). Balaenidae are
represented by the living Balaena mysticetus (Reeves & Leatherwood 1985) and Eubalaena
glacialis (Cummings 1985a), and by the fossil Balaenella brachyrhynus (Bisconti 2005) and
Balaenula astensis (Bisconti 2000, 2003). Caperea marginata is the only included neobalaenid
species (Beddard 1901; Baker 1985; Bisconti 2013). Almost all the living balaenopterid species are
included in the dataset; they are Megaptera novaeangliae, Balaenoptera physalus, B. musculus, B.
acutorostrata, B. edeni, B. borealis together with several fossil species such as Plesiobalaenoptera
quarantellii (Bisconti 2010), Protororqualus cuvieri (Bisconti 2007a), Archaebalaenoptera
castriarquati (Bisconti 2007b), Diunatans luctoretemergo (Bosselaers & Post 2010), “Megaptera”
hubachi (Dathe 1983; Bisconti 2011), “Balaenoptera” siberi (Pilleri, 1989), and Parabalaenoptera
baulinensis (Zeigler et al. 1997). Eschrichtiidae is represented by the living Eschrichtius robustus
(Wolman 1985) and the extinct Eschrichtioides gastaldii (Bisconti 2008) and Archaeschrichtius
ruggieroi (Bisconti & Varola 2000, 2006).
2
Many fossil mysticete taxa were assigned to Cetotheriidae, a family that, over the years, has been
regarded as a paraphyletic taxonomic basket. Recently, Cetotheriidae has been re-defined as to
include only a group of species characterized by shared morphological characters observed in the
rostrum-frontal interface and in the ear bones (Bouetel & De Muizon 2006; Whitmore & Barnes
2008; Bisconti 2008). The cetotheriid taxa used here are: Cetotherium rathkei (Brandt 1873; Pilleri
1986), Piscobalaena nana (as redescribed by Bouetel & De Muizon 2006), Metopocetus durinasus
(Kellogg 1968a), Nannocetus eremus (Kellogg 1926), Herpetocetus transatlanticus, H. scaldiensis,
H. bramblei (Whitmore & Barnes 2008), a Japanese Herpetocetus sp. (Oishi et al. 1985), and
Mixocetus elysius (Kellogg 1934c). The latter has been considered a problematic taxon by some
authors as its skull is heavily reconstructed and it appears difficult to understand which parts are
original and which are not. However, in the pictures published by Kellogg (1934c), the
reconstructed portions are clearly highlighted thus allowing the correct identification of the real
morphological features. For this reason, Mixocetus is included here.
Several additional taxa previously assigned to Cetotheriidae are included in the present dataset as
they represent an important portion of the past diversity of the baleen-bearing mysticetes. These
taxa lack the diagnostic characters of Cetotheriidae and are considered variously by different
authors as Cetotheriidae sensu lato, Cetotheriidae incertae sedis or Cetotheriidae sedis mutabilis.
Bisconti et al. (2013) proposed to call them basal thalassotherians. The lack of the morphological
characters defining the family Cetotheriidae is crucial and implies that these taxa are not part of
Cetotheriidae. They are: Diorocetus hiatus (Kellogg 1968b), “Aglaocetus” patulus (Kellogg
1968c), Parietobalaena palmeri (Kellogg 1924, 1968d), Pelocetus calvertensis (Kellogg 1965),
Uranocetus gramensis (Steeman 2009), Cophocetus oregonensis (Kellogg 1934a), Aglaocetus
moreni (Kellogg 1934b), Titanocetus sammarinensis (Capellini 1900; Bisconti 2006), Isanacetus
laticephalus (Kimura & Ozawa 2002), Joumocetus shimitzui (Kimura & Hasegawa 2010). The
Early Miocene specimen USNM 187416 represents a new mysticete taxon still to be described but
that is included here because it exhibits very primitive character states and seems the sister group of
3
a large mysticete radiation including Cetotheriidae, Eschrichtiidae, Balaenopteridae and all the
other non-balaenid taxa.
The specimens included in the analysis are the following:
Protocetus atavus: SMNS 11084 (holotype); Fraas (1904); Middle Eocene.
Georgiacetus vogtlensis: Hulbert et al. (1996); Hulbert (1998); Middle Eocene.
Dorudon atrox: Uhen (2004); Late Eocene.
Zygorhiza kochii: USNM 4748, 16638, 449538; Kellogg (1936), Uhen (1998); Late Eocene.
Mammalodon colliveri: Fitzgerald (2010); Late Oligocene.
Aetiocetus weltoni: Barnes et al. (1995), Deméré et al. (2008); Late Oligocene.
Micromysticetus rothauseni: ChM PV4844 (holotype), Sanders & Barnes (2002a); Late Oligocene.
Eomysticetus whitmorei: ChM PV4253 (holotype), Sanders & Barnes (2002b); Late Oligocene.
Caperea marginata: AMNH AMO 36692; IRSN 1536; ISAM ZM 41126, ZM 14407, ZM 19944,
ZM 40626, 03/06 on display; Bisconti (2012), Baker (1985), Beddard (1901); Recent.
Balaenella brachyrhynus: NMB 42001; Bisconti (2005); Early Pliocene.
Balaena mysticetus: USNM 257513; NML 1680, 3997, 2563, 2001; Bisconti (2003), Reeves &
Leatherwood (1985); Recent.
Balaena montalionis: MSNT MC CF 31 (holotype); Bisconti (2000, 2003); Early Pliocene.
Balaenula astensis: MSNT MC CF 35 (holotype); Bisconti (2000); Early Pliocene.
Eubalaena glacialis: AMNH 42752, 256803, 90241; MSNT 264; USNM 267612, 3339990, 23077,
301637; Bisconti (2003), Cummings (1985a), True (1904); Recent.
USNM 187416: unpublished pers. obs; Early Miocene.
Pelocetus calvertensis: USNM 11976 (holotype); Kellogg (1965); Middle Miocene.
Uranocetus gramensis: Steeman (2009); Late Miocene.
Metopocetus durinasus: USNM 60460 (holotype); Kellogg (1968a); Late Miocene.
Diorocetus hiatus: USNM 16783 (holotype), 205990; Kellogg (1968b); Middle Miocene.
4
“Aglaocetus” patulus: USNM 23690 (holotype); Kellogg (1968c); Middle Miocene.
Parietobalaena palmeri: AMNH 128885; USNM 10677, 16570, 24883, 10909; Kellogg (1968d);
Middle Miocene.
Joumocetus shimitzui: Kimura & Hasegawa (2010); Late Miocene.
Isanacetus laticephalus: Kimura & Ozawa (2002); Early Miocene.
Cetotherium rathkei: Pilleri (1986); Middle Miocene.
Cophocetus oregonensis: Kellogg (1934a); Early Miocene.
Aglaocetus moreni: Kellogg (1934b); Early Miocene.
Mixocetus elysius: Kellogg (1934c); Late Miocene.
Piscobalaena nana: Bouetel & De Muizon (2006); Late Miocene-to-Early Pliocene.
Herentalia nigra: NML ZMA 5069 (holotype); this work; Late Miocene.
Nannocetus eremus: Kellogg (1929), Whitmore & Barnes (2008); Late Miocene.
Herpetocetus scaldiensis: Van Beneden (1872), Whitmore & Barnes (2008); Late Miocene.
Herpetocetus transatlanticus: Whitmore & Barnes (2008); Early Pliocene.
Herpetocetus bramblei: Whitmore & Barnes (2008); Late Miocene.
Herpetocetus sp. (Japan): Oishi et al. (1985); Early Pliocene.
Eschrichtius robustus: AMNH 181374, 34260, 1750 (‘Eschrichtius cephalum’), A; NMB 42001;
USNM 364969, 364580, 571931, 364969, 364977, 364970, 364973, 504305; NML St20350,
St13130, 630 (‘Eschrichtius gibbosus’); Wolman (1985), True (1904); Pleistocene to Recent.
Eschrichtioides gastaldii: MGPT 13802 (holotype); Portis (1885), Bisconti (2008); Early Pliocene.
Titanocetus sammarinensis: MGB 9073 1CMC172 (1-6) (holotype); Capellini (1900), Bisconti
(2006); Middle Miocene.
Protororqualus cuvieri: Cortesi (1819), Cuvier (1823), Van Beneden (1875), Strobel (1881),
Bisconti (2007a); Middle Pliocene.
Archaebalaenoptera castriarquati: MCA 240536 (inventory of the Soprintendenza per i Beni
Archeologici dell’Emilia Romagna) (holotype); Bisconti (2007b); Middle Pliocene.
5
Plesiobalaenoptera quarantellii: MPST 240505 (inventory of the Soprintendenza per i Beni
Archeologici dell’Emilia Romagna) (holotype); Bisconti (2010); Late Miocene.
Parabalaenoptera baulinensis: Zeigler et al. (1997); Late Miocene.
Megaptera hubachi: MNBe MB.Ma 28570 (holotype); Dathe (1983), Bisconti (2011); Middle
Pliocene.
Megaptera novaeangliae: AMNH 24679; MSNT 263; USNM 269982, 486175 (1-2), 13656/16252,
21492; ZMA 14964, 14953 (1-2), 14952 (1-2), 14965, 14966, 14967; Clapham & Mead (1999),
Winn & Reichley (1985); Recent.
“Balaenoptera” siberi: Pilleri (1989); Early Pliocene.
Diunatans luctoretemergo: Bosselaers & Post (2010); Early Pliocene.
Balaenoptera acutorostrata: AMNH 181411, 35680; IRSN 1537; MSNT 260, 261; ZMA 12873;
Stewart & Leatherwood (1985), True (1904); Recent.
Balaenoptera borealis: USNM 504699, 504698, 504701, 504244, 486174; Gambell (1985a);
Recent.
Balaenoptera physalus: AMNH 35026, 256796; MSNT 251, 252, 253, 258, 255, 257; ZMA 14950
(1-2), 14927 (1-2), 14935 (1-2), 23353, 14947; Gambell (1985b); Recent.
Balaenoptera musculus: AMNH 234949, 256797, 256798; MSNT 250; ZMA 23356, 23354, 23355,
14946, 14942, 14961; Yochem & Leatherwood (1985), True (1904); Recent.
Balaenoptera edeni: USNM 504692, 236680 (1-3); Cummings (1985b), Junge (1950); Recent.
Balaenoptera omurai: Wada et al. (2003); Recent.
6
3. Morphological dataset
Maxilla
1.
Rostrum length: 0, rostrum forms around half of the length of the skull; 1, rostrum forms less than half of the
skull; 2, rostrum forms more than half of the length of the skull.
2.
Rostrum width at base (between bases of lateral processes of the maxillae): 0, rostrum wide; 1, rostrum narrow.
3.
Lateral borders of maxilla posteriorly (in dorsal view) (between bases of lateral processes of the maxillae): 0,
gently diverging from the longitudinal axis; 1, parallel to longitudinal axis.
4.
Lateral borders of maxilla in dorsal view: 0, straight; 1, converging.
5.
Rostrum straight in lateral view: 0, yes; 1, no (rostrum anteroposteriorly arched).
6.
Rostrum arc: 0, moderate; 1, rostrum highly arched.
7.
Rostrum arc: 0, continuous; 1, discontinuous.
8.
Lateral process of maxilla: 0, absent; 1, present.
9.
Lateral process of maxilla: 0, long; 1, very short to absent; 2, very long.
10.
Antorbital groove: 0, absent; 1, present.
11.
Antorbital notch: 0, absent; 1, present.
12.
Grooves on the palate for vasculature of baleen epithelium: 0, absent; 1, present.
13.
Baleen: 0, absent; 1, present.
14.
Baleen: 0, long; 1, short.
15.
Ascending process of maxilla: 0, absent; 1, present.
16.
Ascending process of maxilla: 0, short; 1, long.
17.
Ascending process of maxilla: 0, wide at base; 1, narrow at base.
18.
Ascending process of maxilla: 0, posteriorly rounded; 1, posteriorly pointed; 2, posteriorly squared.
19.
Ascending process of maxilla: 0, located anterior to antorbital corner of frontal; 1, posterior.
20.
Ascending process of maxilla: 0, lateral border forms a distinctive angle with posterior border of maxilla; 1,
lateral border forms a wide curve with posterior border of maxilla; 2, posterior border forms a straight line
inclined towards the longitudinal axis of the skull and posteriorly.
21.
Ascending process of maxilla: 0, the distinctive angle is obtuse; 1, the distinctive angle is around 90°.
22.
Ascending process of maxilla: 0, terminates at posterior border of nasal; 1, terminates posteriorly to the nasal; 2,
terminates anteriorly to the nasal.
7
23.
Ascending processes of maxillae: 0, do not meet posteriorly along midline; 1, meet posteriorly to the nasal
along midline.
24.
Ascending processes of maxillae: 0, do not meet supraoccipital posteriorly; 1, meet supraoccipital posteriorly.
25.
Dorsal exposure of ascending process of the maxilla: 0, wide; 1, narrow because they are exposed laterally on
the depression of the supraorbital process of the frontal.
26.
Lateral surface of maxilla: 0, vertical; 1, flat and exposed dorsally.
27.
Lateral border of maxilla: 0, thick; 1, thin.
28.
Posterior border of maxilla: 0, tightly joined with frontal; 1, loosely joined with frontal or no joined at all.
29.
Infraorbital foramen: 0, bilateral foramens; 1, distintegrated into a series of foramens.
30.
Medial border of maxilla crest-like in the posterior half: 0, no; 1, yes.
31.
Infraorbital foramen distribution: 0, scattered in the posterior surface of the maxilla; 1, allineated closely to
medial border of maxilla.
Premaxilla
32.
Premaxillary foramen: 0, present; 1, absent.
33.
Anterior expansion in premaxilla: 0, absent; 1, present.
34.
Posterior termination of premaxilla: 0, anterior to posterior end of maxilla; 1, at the same level of maxilla and
nasal.
35.
Posterior termination of premaxilla if anterior to maxilla: 0, lateral to the nasal; 1, anterior to the nasal.
36.
Teeth in maxilla and premaxilla: 0, present; 1, absent.
37.
Mesorostral groove: 0, absent; 1, present.
Nasal
38.
Anterior end of nasal: 0, close to anterior end of rostrum; 1, around mid-rostrum; 2, slightly anterior to anterior
border of supraorbital process of frontal; 3, at the same level of anterior border of supraorbital process of
frontal; 4, posterior to anterior border of frontal.
39.
Nasal proportions: 0, nasal width much lower than nasal length; 1, nasal width close to nasal length; 2, nasal
width higher than nasal length.
40.
Width of posterior portion of nasal: 0, rather wide; 1, very thin.
41.
Lateral borders of nasal: 0, parallel to longitudinal axis of the skull; 1, anteriorly diverging; 2, anteriorly
converging; 3 transversely constricted.
8
42.
Anterior border of nasal: 0, concave; 1, transversely flat; 2, convex; 3, lateral corner more anterior than medial
corner.
43.
Nasal length relative to condylobasal length: 0, long; 1, moderate; 2, short.
44.
Nasal dorsal surface: 0, flat; 1, sagittal keel anterior half.
Lacrimal
45.
Lacrimal exposition in dorsal view: 0, lacrimal absent in dorsal view; 1, lacrimal well exposed in dorsal view.
Frontal
46.
Supraorbital process of frontal: 0, horizontal and continuous with interorbital region of frontal; 1, gently
descending from interorbital region; 2, abruptly depressed from interorbital region.
47.
Supraorbital process of frontal: 0, emergence anteroposteriorly narrow; 1, emergence anteroposteriorly
elongated.
48.
Supraorbital process of frontal: 0, very short; 1, short; 2, long.
49.
Posterior border of supraorbital process of frontal: 0, concave; 1, straight.
50.
Orientation of anterior border of supraorbital process of frontal: 0, lateral; 1, anterior; 2, posterior.
51.
Ascending temporal crest: 0, very evident; 1, evident; 2, reduced to a line.
52.
Ascending temporal crest: 0, located on posterodorsal edge of supraorbital process of frontal; 1, developed from
postorbital corner of sop to the anteromedial corner of sop; 2, posteriorly concave with lateral end terminating at
postorbital corner; 3, located at anterior border of supraorbital process of frontal.
53.
Postorbital corner of supraorbital process of frontal shows a clearly visible articular facet for zygomatic process
of squamosal: 0, no; 1, yes.
54.
Postorbital corner of supraorbital process of frontal far from zygomatic process of squamosal: 0, yes; 1, no, they
are close.
55.
Coronal (frontoparietal) suture in dorsal view: 0, transversely concave; 1, straight; 2, transversely convex
because parietal is superimposed on interorbital region of frontal posteriorly; 3, transversely concave because
posteromedial elements of rostrum are interposed between parietal squamae.
56.
Posterior contact of interorbital region of frontal in dorsal view: 0, parietal; 1, supraoccipital.
57.
Position of nasofrontal suture: 0, within interorbital region of frontal; 1, anterior border of interorbital region of
frontal.
9
58.
Anterior exposure of interorbital region of frontal: 0, long; 1, short because posteromedial elements of rostrum
are located on it; 2, short because supraoccipital superimposes on it; 3, short because parietal superimposes on
it.
59.
Paraxial length of interorbital region of frontal: 0, long; 1, very long; 2, short.
60.
Interorbital region of frontal forms subtle sheet of bone surrounding ascending process of maxilla: 0, absent; 1,
present.
61.
Transverse width of interorbital region of the frontal between the depressions of the supraorbital processes of
the frontal: 0, narrow; 1, wide.
Jugal
62.
Jugal shape: 0, long and mainly straight; 1, short and ventrally rounded.
63.
Zygomatic arch lateral to the plane on which the posterior edge of the exoccipital lies: 0, yes; 1, no.
Parietal
64.
Exposure of parietal at vertex: 0, long; 1, short; 2, very short; 3, absent.
65.
Sagittal crest formed by parietal at vertex: 0, present; 1, absent.
66.
Sagittal crest formed by parietal at vertex: 0, acute; 1, transversely flat and formed by two opposite concavities
on both sides of longitudinal axis of the skull.
67.
Width between lateral concavities formed by parietal at vertex: 0, short; 1, wide; 2, very wide.
68.
Parietal anteriorly overwhelmed by temporal crest: 0, no; 1, yes.
69.
Parietal posteriorly overwhelmed by temporal crest: 0, no; 1, yes.
70.
Transverse width of postorbital constriction: 0, very narrow; 1, narrow; 2, wide.
71.
Elongation of postorbital constriction: 0, very long; 1, long; 2, short.
72.
Post-parietal foramen: 0, present; 1, absent.
73.
Lambdoid crest in dorsal view: 0, wide and rounded; 1, triangular.
74.
Posterior apex of lambdoid crest in dorsal view: 0, anterior or same level of occipital condyles; 1, posterior to
occipital condyles.
75.
Parietal and squamosal bulging in temporal fossa: 0, absent; 1, present.
76.
Strong concavity in temporal fossa posterior to emergence of supraorbital process of frontal: 0, absent; 1,
present.
10
77.
Anterior end of parietal: 0, posterior to the posteromedial elements of rostrum; 1, anterior to the posteromedial
elements of rostrum (parietal interdigitated with maxilla).
78.
Anterolateral portion of parietal squama: 0, superimposed on the emergence of supraorbital process of frontal; 1,
not.
Interparietal
79.
Interparietal exposition at vertex: 0, absent; 1, present.
80.
Anterior border of interparietal: 0, in contact with parietal; 1, in contact with interorbital region of frontal.
Squamosal
81.
Squamosal cleft: 0, absent; 1, present.
82.
Squamosal cleft shape: 0, straight; 1, lateral part forms a distinctive angle with medial part.
83.
Squamosal cleft emergence at adulthood: 0, from squamosal-alisphenoid suture; 1, from parietal-squamosal
suture; 2, from squamosal-pterygoid suture.
84.
Squamosal cleft emergence during later ontogeny: 0, from parietal-alisphenoid suture; 1, from parietalsquamosal suture; 2, from squamosal-pterygoid suture.
85.
Cranio-mandibular joint: 0, close articulation; 1, loose articulation.
86.
Cranio-mandibular joint: 0, posterior to orbit and elongated anteroposteriorly; 1, approximately under the orbit
and elongated dorsoventrally.
87.
Dorsoventral elongation of squamosal: 0, absent; 1, present.
88.
Supramastoid crest: 0, absent; 1, present.
89.
Secondary squamosal crest: 0, absent; 1, present.
90.
Zygomatic process of squamosal in lateral view: 0, long; 1, short; 2, very short.
91.
Zygomatic process of squamosal in lateral view: 0, projecting anteriorly; 1, projecting anteriorly and ventrally.
92.
Zygomatic process of squamosal in lateral view, lateral fossa: 0 absent; 1, present.
93.
Zygomatic process of squamosal in dorsal view (lateral border): 0, projecting medially; 1, projecting anteriorly;
2, projecting anterolaterally; 3, strongly diverging from anteroposterior axis of skull.
94.
Apex of zygomatic process of squamosal twisted medially: 0, no; 1, yes.
95.
Postglenoid process of squamosal: 0, projecting ventrally much more than zygomatic process; 1, projecting
ventrally close to zygomatic process.
96.
Transversely compressed postglenoid process of squamosal: 0, no; 1, yes.
11
97.
Glenoid fossa of squamosal in lateral view: 0, right angle; 1, highly concave (crescent-shaped); 2, straight and
inclined ventroposteriorly; 3, straight and oriented anteroposteriorly; 4, slightly concave.
Alisphenoid
98.
Alisphenoid exposure in temporal fossa: 0, present; 1, absent.
99.
Alisphenoid contacts in temporal fossa: 0, alisphenoid between frontal, parietal, squamosal, and pterygoid; 1,
alisphenoid between parietal, squamosal, and pterygoid; 2, alisphenoid between parietal and pterygoid; 3,
alisphenoid between parietal and squamosal.
Supraoccipital
100.
Position of anterior border of supraoccipital: 0, posterior to anterior end of zygomatic process of squamosal; 1,
anterior to anterior end of zygomatic process of squamosal.
101.
Attach sites for neck muscles on lateral surfaces of supraoccipital: 0, low to absent; 1, present.
102.
Attach sites for neck muscles on lateral surfaces of supraoccipital: 0, well developed; 1, strong.
103.
Nuchal plane: 0, concave; 1, flat; 2, convex.
104.
External occipital protuberance: 0, concave; 1, convex; 2, flat.
105.
Sagittal crest on supraoccipital: 0, absent; 1, present.
106.
Massive elongation of supraoccipital: 0, absent; 1, present.
107.
Anterior border of supraoccipital: 0, rounded; 1, borders converging and forming a sort of triangle in dorsal
view.
108.
Anterior border of supraoccipital in dorsal view: 0, narrow and rounded; 1, pointed; 2, narrow and squared; 3,
wide and squared.
109.
Lateral borders of supraoccipital: 0, uniformly convex; 1, uniformly concave; 2, straight; 3, sinusoidal.
110.
Transverse constriction of supraoccipital: 0, moderate; 1, strong.
111.
Lateral border of supraoccipital anterior to the strong constriction: 0, concave; 1, straight.
Exoccipital
112.
Protrusion of posterolateral corner of exoccipital in ventral view: 0, absent; 1, strong and reaching a point
posterior to occipital condyles; 2, strong and reaching a point anterior to occipital condyles.
113.
Protrusion of posterolateral corner of exoccipital in ventral view: 0, largely medial to postglenoid process of
squamosal; 1, posterior to postglenoid process of squamosal.
12
114.
Occipital condyles: 0, show highly convex surfaces both dorsoventrally and transversely; 1, show convex
surfaces only dorsoventrally and nearly flat surfaces transversely.
115.
Occipital condyles: 0, with laterally concave neck; 1, with very short-to-absent, scarcely concave neck.
Basicranium
116.
Infraorbital plate of maxilla: 0, absent; 1, present.
117.
Location of optical channel: 0, under middle part of supraorbital region of frontal; 1, in posterior part of
supraorbital process of frontal.
118.
Basioccipital-basisphenoid suture obliterated by posterior termination of vomer: 0, no; 1, yes.
119.
Palatine elongation: 0, palatine does not reach posterior border of skull; 1, palatine reaches posterior border of
skull in ventral view.
120.
Ventral lamina of pterygoid: 0, absent; 1, present.
121.
Foramen pseudo-ovale: 0, absent; 1, present.
122.
Foramen pseudo-ovale: 0, infundibulum incomplete; 1, infundibulum complete.
123.
Location of foramen pseudo-ovale: 0, between squamosal and pterygoid; 1, within squamosal but with fissure
towards pterygoid; 2, within pterygoid.
124.
Posterior lacerate foramen: 0, absent; 1, present.
125.
Posterior lacerate foramen located near to the posterior border of the skull: 0, no; 1, yes.
126.
Anterior condyloid foramen: 0, present; 1, absent.
Periotic
127.
Parabullary ridge in medial view: 0, highly convex dorsally; 1, mildly convex; 2, flat.
128.
Maximum convexity of parabullary ridge located located over internal acoustic meatus in medial view: 0, yes;
1, no, it is more anterior.
129.
Strong bulging of dorsal part of periotic: 0, absent; 1, present.
130.
Endocranial opening of facial canal (adulthood): 0, located within internal acoustic meatus; 1, separated from
internal acoustic meatus.
131.
Endocranial opening of facial canal (later ontogeny): 0, located within internal acoustic meatus; 1, separated
from internal acoustic meatus.
132.
Endocranial opening of facial canal connected by a groove to the anterior rim of internal acoustic meatus: 0, no;
1, yes.
13
133.
Endocranial opening of facial canal with fissure (adulthood): 0, no; 1, yes.
134.
Endocranial opening of facial canal with fissure (later ontogeny): 0, no; 1, yes.
135.
Suprameatal area gently descending towards internal acoustic meatus rim: 0, no, it is vertical; 1, strongly
concave; 2, yes; 3, no, it protrudes.
136.
Transverse elongation of pars cochlearis: 0, absent; 1, present.
137.
Anteroposterior elongation of pars cochlearis: 0, absent; 1, present.
138.
Anterior inflation of pars cochlearis: 0, present; 1, absent.
139.
Median promontorial groove: 0, absent; 1, present.
140.
Tensor tympani groove: 0, present; 1, absent.
141.
Round window and perilymphatic duct confluent: 0, no; 1, yes.
142.
Dorsal tuberosity of periotic cranially rounded: 0, no; 1, yes.
143.
Anterior edge of pars cochlearis protruding cranially and forming a dorsoventral crest: 0, no; 1, yes.
144.
Long and thick roof of stylomastoid fossa: 0, absent; 1, present.
145.
Stylomastoid fossa: 0, broad and barely concave; 1, highly concave and forming a sort of notch at the
emergence of posterior process.
146.
Fossa for stapedial muscle well separated from round window by a thin crest: 0, yes; 1, no.
147.
Anterior process of periotic: 0, absent; 1, present.
148.
Anterior process of periotic: 0, long; 1, very short-to-nearly absent.
149.
Anterior process of periotic in medial view: 0, transversely compressed (blade-like); 1, thick.
150.
Anterior process of periotic in dorsal view: 0, roughly round; 1, squared; 2, triangular.
151.
Anterior process of periotic in dorsal view, triangular apex: 0, rounded; 1, pointed.
152.
Anterior process of periotic in dorsal view, triangular and pointed apex: 0, medial border straight-to-slightly
convex; 1, medial border concave.
153.
Anterior process of periotic abruptly depressed from strong bulging of central part: 0, absent; 1, present.
154.
Lateral process of anterior process of periotic: 0, absent; 1, present.
155.
Lateral process of anterior process of periotic: 0, located at the emergence of the anterior process; 1, formed by
the whole lateral edge of anterior process.
156.
Lateral process of anterior process of periotic: 0, rounded; 1, triangular and short; 2, triangular and long.
157.
Compound posterior process: 0, very short; 1, long.
158.
Compound posterior process: 0, prismatic; 1, flattened.
159.
Articulation between compound posterior process and squamosal: 0, tight; 1, weak.
14
160.
Groove for facial nerve on posterior process: 0, long; 1, short.
161.
Groove for facial nerve on posterior process: 0, wide; 1, narrow-to-tube-like.
162.
Relative size of petrotympanic complex: 0, large; 1, very small; 2, small.
Tympanic bulla
163.
Inner posterior prominence in dorsal view: 0, projects posteriorly; 1, does not project posteriorly.
164.
Outer posterior prominence in dorsal view: 0, projects posteriorly; 1, does not project.
165.
Posterior surface of tympanic bulla: 0, fissurated; 1, not fissurated.
166.
Eustachian opening: 0, low; 1, high.
167.
Dorsoventral height of tympanic cavity: 0, high; 1, low.
168.
Interprominential notch in posterior view: 0, present; 1, absent.
169.
Ventral surface of tympanic bulla: 0, highly concave; 1, convex; 2, keeled.
170.
Anterolateral portion of tympanic bulla: 0, not expanded (pyruliform bulla sensu Bisconti, 2010); 1, expanded.
171.
Anterolateral expansion: 0, short; 1, long.
172.
Anterolateral expansion: 0, continuous with anteromedial corner; 1, anteriorly separated from anteromedial
corner by a lateral concavity visible in dorsal view.
173.
Epitympanic hiatus: 0, small; 1, wide because of massive reduction of the height of conical process.
174.
Dorsolateral border of involucrum in dorsal view: 0, straight; 1, concave; 2, convex.
175.
Dorsal border of involucrum in medial view: 0, descending anteriorly; 1, projecting dorsally at anterior end.
Dentary
176.
Teeth on dentary: 0, present; 1, absent.
177.
Mental symphysis: 0, present; 1, absent.
178.
Groove for mental ligament: 0, absent; 1, present.
179.
Coronoid process: 0, high; 1, low; 2, absent.
180.
Coronoid crest: 0, short; 1, long.
181.
Position of coronoid process: 0, close to the condyle; 1, far from the condyle.
182.
Postcoronoid crest and fossa: 0, absent; 1, present.
183.
Postcoronoid fossa: 0, wide; 1, vestigial.
184.
Satellite process: 0, absent; 1, present.
185.
Satellite process: 0, high; 1, low.
15
186.
Pterygoid fovea: 0, absent; 1, present.
187.
Pterygoid fovea: 0, located medially; 1, located posteriorly.
188.
Pterygoid fovea located posteriorly: 0, highly inclined; 1, scarcely inclined.
189.
Protrusion of angular process: 0, absent; 1, ventral; 2, posterior.
190.
Proportional size of angular process relative to the condyle in the dorsoventral plane (in lateral view): 0, angle
larger than condyle; 1, angle half the size of the condyle; 2, angle severly reduced.
191.
Condyle: 0, oriented dorsally and posteriorly; 1, oriented dorsally; 2, oriented posteriorly.
192.
Articular surface of condyle in lateral view: 0, highly convex; 1, posteriorly flat.
193.
Anterior torsion: 0, slight-to-absent; 1, strong.
194.
Attach for mylohyoidal muscle: 0, apparently not developed; 1, sharp groove; 2, concavity; 3, crest.
195.
Ventral border of dentary shows a lateral groove: 0, no; 1, yes.
196.
Mandibular foramen: 0, wide; 1, small.
197.
Mandibular foramen: 0, concave in medial view; 1, triangular.
198.
Mental foramen: 0, bilateral foramens; 1, disintegrated into a series of foramens.
199.
Gingival foramens and grooves: 0, absent; 1, present.
200.
Dorsoventral arc of dentary ramus: 0, absent; 1, present in anterior-most portion; 2, present and continuous
along the dentary.
201.
Lateral arc of dentary ramus: 0, absent; 1, slight; 2, strong.
Vertebral column
202.
Cervical vertebrae: 0, elongated; 1, very shortened.
203.
Cervical vertebrae: 0, free; 1, fused.
204.
Dorsal process in cervical vertebra No. 3: 0, present; 1, absent.
205.
Dorsal process in cervical vertebra No. 4: 0, present; 1, absent.
206.
Dorsal process in cervical vertebra No. 5: 0, present; 1, absent.
207.
Dorsal process in cervical vertebra No. 6: 0, present; 1, absent.
208.
Dorsal process in cervical vertebra No. 7: 0, present; 1, absent.
209.
Ventral process in cervical vertebra No. 3: 0, present; 1, absent.
210.
Ventral process in cervical vertebra No. 4: 0, present; 1, absent.
211.
Ventral process in cervical vertebra No. 5: 0, present; 1, absent; 2, tubercle-like.
212.
Ventral process in cervical vertebra No. 6: 0, present; 1, absent; 2, tubercle-like.
16
213.
Ventral process in cervical vertebra No. 7: 0, present; 1, absent; 2, tubercle-like.
214.
Foramen transversarium complete in cervical vertebra No. 3: 0, yes; 1, no.
215.
Foramen transversarium complete in cervical vertebra No. 4: 0, yes; 1, no.
216.
Foramen transversarium complete in cervical vertebra No. 5: 0, yes; 1, no.
217.
Foramen transversarium complete in cervical vertebra No. 6: 0, yes; 1, no.
218.
Foramen transversarium complete in cervical vertebra No. 7: 0, yes; 1, no.
219.
Neural processes in cervical vertebrae: 0, free; 1, first-to-seventh cervical vertebrae have fused neural processes;
3, second-to-seventh vertebrae have fused neural processes.
Scapula
220.
Inclination of cranial border: 0, scarce; 1, elevated;
221.
Inclination of caudal border: 0, scarce; 1, elevated.
222.
Supra-spinous fossa in lateral view: 0, present and wide; 1, present but reduced; 2, absent.
223.
Infra-spinous fossa in lateral view: 0, concave; 1, flat.
224.
Acromion process: 0, present; 1, absent.
225.
Coracoid: 0, present; 1, absent.
226.
Scapular spine: 0, present; 1, strongly reduced to absent.
Humerus
227.
Forelimb proportions: 0, humerus has same length as radius and ulna; 1, humerus is shorter than radius and
ulna; 2, humerus is longer than radius and ulna.
228.
Medial tuberculum (deltopectoral crest): 0, present; 1, absent.
Radius
229.
Radius and ulna can rotate at humeral articulation: 0, yes; 1, no.
Ulna
230.
Olecranon process: 0, present; 1, absent.
Manus
231.
Carpal bones in articulation: 0, yes; 1, no.
17
232.
First digit: 0, present; 1, absent.
233.
Fifth digit: 0, present; 1, absent.
234.
Trapezium: 0, present; 1, absent.
235.
Hyperphalangy: 0, no; 1, yes.
Hindlimb
236.
Pelvis articulated with vertebral column: 0, yes; 1, no.
237.
Functional hindlimb in adult: 0, yes; 1, no.
Ribs
238.
Number of ribs that are articulated with sternum: 0, more than one pair; 1, one pair.
239.
Rib 2: 0, bifid head; 1, single headed.
240.
Rib 3: 0 bifid head; 1, single headed.
241.
Rib 4: 0 bifid head; 1, single headed.
242.
Rib 5: 0 bifid head; 1, single headed.
243.
Rib 6: 0 bifid head; 1, single headed.
244.
Rib 7: 0 bifid head; 1, single headed.
245.
Rib 8: 0 bifid head; 1, single headed.
246.
Rib 9: 0 bifid head; 1, single headed.
247.
Rib 10: 0 bifid head; 1, single headed.
248.
Rib 11: 0 bifid head; 1, single headed.
Sternum
249.
Sternum formed by several sternebra: 0, yes; 1, no, it is formed by manubrium only.
18
4. Matrix for phylogenetic analysis
Protocetus atavus Fraas, 1904
0-000--0- 0000-0---- ------0000 0000000000 0000000000 0000000000 0-?0000-00
00100000-0 -0---00010 00000000?? 00-00000-0 --00000010 01?0101??? ??????000?
?0???1???? ??????0000 0-00000000 ----10???? ?????????? ?????????? ??0000?000
0000000000 00???????? ?????????? ??????????
Georgiacetus vogtlensis Hulbert et al., 1996
0-000--0- 0000-0---- ------0000 0010000010 0000000000 0000000000 0-00000-00
00100000-0 -0---00000 0000000000 00-00000-0 --00000000 0110100??? ??????000?
?0?????1?? ???????000 ?-00000000 0---100000 -00-0-0--0 0000000010 0000?00?0?
00?0?00?00 ?????????? ??????00?0 000000000?
Dorudon atrox Andrews, 1906
0-000--0- 0000-0---- ------0000 0010000010 0000000000 0000000000 0-00000-00
00000000-0 -0---00010 0000000000 00-00100-0 --00000000 0110101000 0000000001 000000100 ?--00--000 1-00000000 0--0000000 -00-0-0--0 0000000010 0000000000
0000000000 0000000000 0000001000 0000000110
Zygorhyza kochii (USNM 11962) Reichenbach in Carus, 1947
0-000--0- 0000-10111 0110000000 0010000010 0330010000 2000010000 0-00000-00
0000001010 -0---00010 0000000000 00-01000-0 --00000010 0110100000 0?00000000
0000000101 ???????000 ??00000000 0--0000000 000-0-0--0 0000000010 00000??0?0
??000??000 0000000000 0?????11?0 0000001110
Mammalodon colliveri Pritchard, 1939
100?0--11 1000-10101 1-0000?011 00??010?20 00?2000000 0000?00000 0-?0000-00
00100000-0 -0---??0?0 ?????????? 00-01100-0 --0?0010?? ?110100000 0?00001000
0000000101 1--00--??? ??00000000 0--0100000 -00-0-0--0 2000000010 00?0??????
?????????? ?????????? ?????????? ?????????0
Aetiocetus weltoni Barnes et al., 1995
00000--11 1011111111 0100001101 00001-0120 0001010000 0000100010 0-00000-00
1110001010 -0---10010 0100010100 0?????0??? ??00001000 0111101??? ??????????
00???0?1?1 ???????1?0 0000000000 0--0100110 -00-0-0--0 1000000010 00????????
?????????? ?????????? ?????????? ??????????
Micromysticetus rothauseni Sanders & Barnes, 2002a
????????? ?????????? ?????????? ?????????? ?????????? ??????0??? ????100-00
1110000010 -0---10011 000200?1?? 0100110100 --1011??10 0110101000 0?00010000
000000?100 1--00--000 101??????? ?????????? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
Eomysticetus whitmorei Sanders & Barnes, 2002b
1???0--?? ??1110---- ------11?? 00101-1110 00010?1010 0000000012 0-?0000-00
0010000010 -0---10001 00020001?? 0100110112 --1111?010 0110101000 1?00010000
000000?100 1--00--001 1010000000 0--0001110 -00-0-0--0 1000000011 001000?000
0?00???110 1??001?001 0????????0 000000111?
Caperea marginata Gray, 1846
000011010 101100---- ------1111 00101-1120 0022002011 2210121122 0-103---10
2210101010 -102211110 2001000402 10-21110-0 --11111010 1112111100 1?00020001
1001000100 2??????100 0001110110 1001201112 --0-0-10-0 1100101011 2011000001
1111111111 1120000111 0101111110 0000011111
19
Balaenula astensis Trevisan, 1941
211011010 10?100---- ------0111 00101-1120 00020?1021 0110121122 0-?03---00
2210000000 -0---11110 20010004?? 10-21010-0 --20111011 100011020? ?????000??
0?0010?101 1--011?101 0001111110 1001201112 --0-0-10-0 1101101011 20????????
?????????? ?????????? ?????????? ??????????
Eubalaena glacialis Müller, 1776
211011112 101100---- ------0111 00101-1120 0002001021 0110121122 0-103---11
2210000000 -0---10110 2001010301 10-21010-0 --20111011 1010111000 0000031001
0001101101 1--0111101 0001111110 1001201112 0-0-0-10-0 1101101011 2211000000
0000111112 0011011111 1100111110 0000000011
Balena mysticetus Linnaeus, 1758
211011012 101100---- ------0111 00101-1120 0002001021 2210121122 0-103---11
2210000010 -0---10110 2102010301 10-2101123 --20111011 1100111000 0000000001
0000100101 1--0102100 0001210110 1001201112 0-0-0-10-0 1101101011 2211000000
0012111112 0010011111 1100111111 0000000011
Balaenella brachyrhynus Bisconti, 2005
000011012 101100---- ------0111 00101-1121 0222001021 2210121122 0-?03---11
2210000010 -0---10110 2002010301 10-1101123 --201110?0 1100111??? ??????0001
00?010?101 1--0112100 0001110110 100120???? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
Pelocetus calvertensis Kellogg, 1965
20110--10 101110---- ------1111 00101-1120 11?2001020 1130120132 0-?0100-00
1110100000 -0---10011 0002000002 10-0000112 --10111010 0111101000 0?00000001
0000010101 0--0120100 0001110011 1100101??1 01100-10-1 1200000011 021000?000
0?0011?110 1000100211 ????????10 000000011?
“Aglaocetus” patulus Kellogg, 1968c
20110--10 101110---- ------1111 00101-1120 11?2001020 0210120132 0-?0100-00
1110000000 -0---10010 0002000002 10-0000112 --20111010 0111101000 0?00?00001
0000000101 0--0120100 0001110011 110010???? ?????-???? ?????????? ??100?0??0
?0??1?1??0 ?????????? ?????????? ??????????
Isanacetus laticephalus Kimura & Ozawa, 2002
20110--10 1011110001 ?-----1111 00101-1120 11?2011020 1210120132 0-?0100-00
1110100010 -0---10010 000200021- 10-0010112 --20111010 0110101000 0?00?00001
000000?101 20-0100100 0001010011 110010111? ?????-???? ????00??11 ??100??0?0
??1?1??1?0 ?????????? ?????????? ??????????
Uranocetus gramensis Steeman, 2009
20110--10 101110---- ------1111 00111-1120 1112001020 011??20132 0-?010?-00
1110100010 -0---1001? 00?20002?? 10-0?10113 --20111010 0???101??? 0?00??0001
00?000010? 0--0?????? ?001110011 1100101??1 ????0-???1 1200000011 0?1000???0
0???00???0 0000000111 0???????10 ?????????1
Parietobalaena palmeri Kellogg, 1924
20110--11 101110---- ------1111 00111-1120 1112001020 1110120132 0-?0100-00
1110000010 -0---10010 0012000402 1100110113 0011110010 0111101000 1?10?00001
0000010110 0--0100101 0001110011 0--0001111 01100-10-1 1200001011 01100000?0
000??????0 ?????????? ?????????? ??????????
Aglaocetus moreni Kellogg, 1934b
20110--11 1011110101 0000001111 000111-112 0032001020 0110130112 0-?0101000
1110100010 -0---10010 0002000401 10-0000113 --11111010 0111101??? ??????000?
?????????? ???????101 00???????? ??????1111 01100-10-1 1200001011 02????????
?????????? ?????????? ?????????? ??????????
20
Titanocetus sammarinensis Bisconti, 2006
20110--11 0011110101 0022000111 0010001120 0332001020 1130130112 0-?0101000
2210110010 -0---10010 0002000?01 00-00000-0 --20111010 0???101??? ??????????
?????????? ???????101 ?????????? ??????1111 011?0-0--0 1210001111 02100????0
?????????0 ?????????? ?????????? ??????????
Cophocetus oregonensis Kellogg, 1934a
20110--10 1111110001 1-00001111 00111-1120 0002001020 1130110112 0-?0101000
1210100010 -0---10010 0001000?01 10-0000113 --10111010 0111101??? 0?00?00001
00?0??0101 ???????101 0??1110011 1100001??1 01100-10-1 1200000111 0110??00?0
0001?????0 0010000111 0100?111?? ??????????
Diorocetus hiatus Kellogg, 1968b
21110--10 1011110001 1-00001111 00111-1120 1002001020 1110120112 0-?0100-00
1210100010 -0---10010 0012000402 1100010113 0020111010 0111101010 1?10?00001
00000?0101 20-00--101 0001110011 0--0101111 01100-10-2 1200001011 0110???00?
??11?????0 11??000??1 0????????? ??????????
Joumocetus shimitzui Kimura & Hasegawa, 2010
20?10--?? ??1110---- ------1111 00111-1120 11320?10?? 021??20132 0-?0100-00
12100?0010 -0---??0?? ??????0??? ?0-0010113 001?11101? ????101??? ??????0110
?0?0???100 ???????000 ??0111??11 0--?10111? ?????????2 121000??11 01????????
?????????? ?????????? ?????????? ??????????
Cetotherium rathkii Brandt, 1873
?11?0--11 1111111001 1-10001111 00??001?20 1112001010 111013-1-2 1-?0201100
121010001? ?????10010 00010002?? 0100110101 0011111010 011?101??? ??????????
?????????? ?????????? ?101110011 0--0?01??? ?????????? ???000??11 01????????
?????????? ?????????? ?????????? ??????????
Piscobalaena nana Pilleri & Siber, 1989
21100--11 1111111001 1-11001111 0011001130 1112001010 111013-100 1-10101100
1210100110 -0---10010 001200041- 00-0010100 0011111110 0111101110 1110000001
0010010101 0--0100001 0101210011 0--0101111 01100-10-2 1000001011 0210000000
0001001110 1110000111 01????1110 0000011111
Mixocetus elysius Kellogg, 1934c
21100--11 1111111001 1-00001111 00111-1130 0102001010 111013-0-2 1-?0201100
121011011? ?????10010 00?10?0?01 ?????????? ??11111110 0111101??? ??????000?
??????1??? ???????100 0????????? ??????1111 11000-???? ?200001?11 ??10000000
00000????0 1110000??? ?????????? ??????????
Metopocetus durinasus Cope, 1886
????????? ?????1100? 1-1100???? ????01???0 11???????? ?????3-?-2 1-?0201100
1210?101?0 -0---??010 ??????0??? ?100010103 00??????1? ???????110 1?10?00001
00100??101 ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
Herentalia nigra this work
????????? ?????1100? 1-1100???? ????01??20 11?20?10?? ?12??3-0-2 1-??201110
12100101?1 110?????1? ?????????? ?100010000 01?101??1? 0110101??0 ??????00??
???????100 ???0100111 0111?????? ?????????? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
Herpetocetus transatlanticus Whitmore & Barnes, 2008
????????? ?????110?1 1-1100???? ?????????0 11????1010 ?12??3-0-2 1-?1?????0
1201100010 -0---10010 00020114?? 0?????0??? ??1111??1? 0110101100 1111300001
0000000100 0--0100101 0111210011 0--000???? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
21
Herpetocetus bramblei Whitmore & Barnes, 2008
????????? ?????????? ?????????? ?????????? ??????1?10 ???01????? ???1????00
1200010??? ?0---10010 10110114?? ?????????3 002111???? 0110101100 1?11?00001
00010?0100 0--0100101 01112????? ????????1? ?????????? ??????00?1 01????????
?????????? ?????????? ?????????? ??????????
Herpetocetus scaldiensis Van Beneden, 1872
????????? ?????????? ?????????? ?????????? ?????????? ?????????? ???1??????
?????????? ?0---10010 00110114?? ?????????? ??21?????? ????10?100 1?11?00001
00010?0100 ???0???101 011??????? ???????111 10110-10-2 1000001011 01????????
?????????? ?????????? ?????????? ??????????
Herpetocetus sp. (Japan; Oishi et al., 1985
21100--11 0111111000 1-11001111 0010001120 1112001010 012003-0-2 1-?1201100
12?001010? ?10??10010 10110114?? 0100?1010? ??1???1?1? ????1????? ??????????
?????????? ?????????? ?????????? ???????111 10110-1012 10?00???11 0110??????
?????????? ?111000111 10100?1111 ??????????
Nannocetus eremus Kellogg, 1929
????????? ?????1???? ??11?0???? ?????????? ??????1010 012003-0-2 1-?12???00
12?0010??? ?0---10010 10110112?? 0?????0000 002111?01? 0110101100 1?11?00010
0000???100 0--0100101 0112111011 0--010???? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????????
USNM 187416 (pers. obs.)
21100--10 101110---- ------1111 00101-1120 1102001010 1110120132 0-?0100-00
1210010010 -0---10010 00010004?? 00-0000113 0011111110 0110101??? ??????????
?????????? ???0???101 000111??11 0--???1111 01100-10-0 1200001011 01????????
?????????? ?????????? ?????????? ??????????
Eschrichtius robustus (Lilljeborg, 1861)
211010011 1111111111 0100011111 00101-1120 0012002011 2220030010 1010101100
1210011110 -100?10010 1001000201 0112100023 0011111110 0110101010 0?00?11101
010100?101 20-00--111 0001111011 1100201112 -1--1110-2 1100201011 2110000000
0000111110 0020000111 0101111111 0000011111
Eschrichtioides gastaldii Bisconti, 2008
2???0--?? ??11111101 0100001111 001?001120 0012002011 012?130010 1010101100
1211011010 -101?10010 10020102?? 01112000-0 --11111?10 0110101??? ??????????
?????????? ???????110 ??01111011 1100001111 011-1110-1 1100201011 2210??????
?????????? ???????111 0?????11?? ??????????
Archaeschrichtius ruggieroi Bisconti & Varola, 2006
????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ?????????? ?????????? ?????????? ??????????
?????????? ?????????? ?????????? ??????1110 011010???? ???000??11 22????????
?????????? ?????????? ?????????? ??????????
Archaebalaenoptera castriarquati Bisconti, 2007a
20100--10 1011111101 0100011111 0011001120 0001002121 2231130011 10?0201210
2211101101 0????10010 00030103?? 1100110103 101???1??? ??????1??? ??????????
?????????? ???????110 ??0??????? ??????111? ?????????? ??0?????11 01????????
?????????? ?????????? ?????????? ??????????
Parabalaenoptera baulinensis Zeigler et al., 1997
20??0--?? ??11111111 0100011111 00101-1120 1031002120 2230131011 10?0301211
2210001100 -????10010 00010002?? 10-1?00122 --01111?10 0110101??? ??????????
?????????? ???????110 ??0111101? ??????1111 11010-11?0 ?2100?1011 02100????0
????0????0 ????????11 ?????????? ??????????
22
Megaptera novaeangliae Brisson, 1762
20110--12 1011111100 0100001111 0010001141 0002102121 2231130011 1110201211
2211001100 -112110000 100300001- 10-1210103 0000111110 0110101110 0?00011100
0001011101 2000101111 1001110011 1100201112 -10?0-1110 2210011011 0210000000
0021111110 1120111111 0101111111 0011111111
Balaenoptera physalus Linnaeus, 1758
20110--10 1011111121 0100001111 00101-1141 0002002121 2231130012 1110201211
2211001101 0112110000 0101010101 10-1000133 0020111110 0110101110 1100121100
1000001101 2100101111 1001111012 1110211111 01110-1110 2210011011 0210000000
0002000110 1110000111 0101111110 0111111111
Balaenoptera acutorostrata Lacépède, 1804
01110--10 1011111121 0100001111 00101-1130 0122002121 2231130011 1110201211
2211001111 1112100000 0101010101 10-1200133 0000111110 0110111210 0100111111
0000011101 2110101111 1001111012 1110211111 01110-1110 2210011011 0210000000
0021001110 1120000111 0101111111 0000111111
Balaenoptera edeni Anderson, 1879
20110--10 1011111121 0100001111 00101-1130 0112002121 2011130011 1110201211
2211001110 -112?10000 0101010101 10-1000133 0000111110 01101012-0 0100131100
1001011101 2100101111 1001111012 1110211111 01110-1110 2210011111 021000000?
?????????? 1120000111 010111111? ?????????1
Balaenoptera borealis Lesson, 1828
20110--10 1011111121 0100001111 00101-1130 0112002121 2231130011 1110201211
2211001101 1112?10000 0101010102 10-1000133 0000111110 01101012-0 0100131110
1101001101 2110101111 1001111012 1110211111 01110-1110 2210011011 021000000?
?????????? 1120000111 010111111? ?????????1
Balaenoptera musculus Linnaeus, 1758
20110--10 1011111121 0100101111 00101-1130 0012002121 2231131011 11103---11
2210001100 -110210000 0103010101 1101200133 0000111110 0111101??1 ??????1100
01??00?101 2101101111 1001111012 1110211111 11110-1110 2210011011 0210000000
0001000110 1110000111 0101111110 0000111111
“Balaenoptera” siberi Pilleri, 1989
20110--10 1011111121 0100101111 00101-1141 0002102121 2230131011 1110011210
2210001110 -????10010 00030000?? 10-01000-0 --10111?1? ?1????1??? ??????????
?????????? ?????????? ?????????? ??????111? ?????????? ???0????11 0210??????
?????????? 1110000111 010111111? ?????????1
Protororqualus cuvieri Bisconti, 2007b
20110--11 0111111101 0100011111 10101-1130 0112002120 2230130011 10?0201211
1111001100 -????10010 0??30????? 1100010113 1110111110 0110101??? ??????????
?????????? ?????????? ???111?012 ????2?1111 01100-???0 22100?1011 01100????0
?????????0 1120000111 0????1111? 0011111111
Plesiobalaenoptera quarantellii Bisconti, 2010
20110--11 101111110? 01?0011111 0011??11?? ??????212? 2??????0?1 ?0??????11
?????0???1 ?????1???? ?????????? ?0-0000122 ????111??? ??????1011 1?10?11111
0000000101 200110011? 1001111012 1100211111 01000-1110 2210200011 021000???0
0???00???0 ?????????? ?????????? ?????????1
“Megaptera” hubachi Dathe, 1983
20110--?? ??11111101 0100001111 00111-1140 0100102121 2231130011 10??201211
221100111? ?110?10010 00030000?? 10-0000123 0-10111110 0111101??? 1?????1100
?1?????101 210??????? ??0??????? ??????1111 01100-1110 1210011011 0210000000
0001111110 1110000111 0101111110 0000111111
23
Diunatans luctoretemergo Bosselaers & Post, 2010
????????? ??11111121 ??0000??1? ????1-1141 00021021?? ?????30011 11??211211
22110011?? ?111?10010 1001000100 10-0100133 0-1011??10 01101012-0 1?00?11101
0000?10101 210-101110 ??01111012 ????11???? ?????????? ?????????? ??1000???0
0????????? ?????????? ?????????? ??????????
24
5. Table S1
Ages and areas of occurrence of Cetotheriidae s.s. taxa and Diorocetus and their translations into
Operational Geographic Units as used in the paleobiogeographic analysis.
Taxon
AREA(S) OF
OPERATIONAL
OCCURRENCE
GEOGRAPHIC UNIT
AGE
Diorocetus hiatus
Langhian
Maryland
North Atlantic
Diorocetus shobarensis
Burdigalian
Japan
North Pacific
Diorocetus (Germany)
Serravallian
Germany
Parathetys
Mixocetus elysius
Tortonian
California
North Pacific
Cetotherium rathkii
Serravallian
Ukraina
Parathetys
Peru
South Pacific
Tortonian/Messinian-toPiscobalaena nana
Zanclian
Metopocetus durinasus
Langhian
Virginia, Portugal
North Atlantic
Herpetocetus (Japan)
Zanclian
Japan
North Pacific
Herpetocetus scaldiensis
Tortonian-to-Zanclian
Belgium
North Sea
Herpetocetus transatlanticus
Zanclian
North Carolina
North Atlantic
Herpetocetus bramblei
Messinian
California
North Pacific
Nannocetus eremus
Tortonian/Messinian
California, Mexico
North Pacific
Herentalia nigra
Tortonian
Belgium
North Sea
25
6. Table S2.
Matrix generated for the Secondary Brooks Parsimony Analysis.
Parathetys 1
100000000 0000001000 00
North Pacific1
100000000 0000001000 00
North Atlantic 1
100000000 0000001000 00
North Pacific 2
010000000 0000001100 00
Parathetys 2
001000000 0000001110 00
South Pacific 1
000100000 0000101111 00
North Atlantic 2
000010000 0000101111 00
North Sea 1
000001000 0001011111 10
North Pacific 3
000000100 0001011111 10
North Pacific 4
000000010 0000011111 10
North Pacific 5
000000001 0010001111 11
North Sea 2
000000000 1001000111 11
North Atlantic 3
000000000 0100001111 11
26
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