TABLE 1. Muscles attached to the basihyal and thyrohyal.
Muscles | Muscle attachments on the basihyal + thyrohyal |
Other side of the muscle attachments |
Function | Reference of function |
Hyoglossus muscle | Anterior suraface of the anterior process and lateral portion of the body |
It ends left and right sides of the tongue |
The hyoglossus pulls the tongue ventrally | Werth and Ito, 2017: rorquals |
Geniohyoid muscle | The anterior surface of the body | The mandibular symphysis | The geniohyoid muscle protracts the tongue | Werth and Ito, 2017: rorquals |
Basio-keratic muscle | The anterolateral border of the lateral portion | The stylohyal | The basio-keratic muscle flexes the thyrohyal posteriorly and decreases the angle forme by the thyrohyal and ceratohyal | Kienle et al., 2015: gray whales |
Sternohyoid muscle | The medial part of the posterior border |
The sternum | The sternohyoid and thyrohyoid similaryly retract and depress the tongue root | Werth and Ito, 2017: rorquals |
Thyrohyoid muscle | The posterior process | The tyroid cartilage | The sternohyoid and thyrohyoid similaryly retract and depress the tongue root | Werth and Ito, 2017: rorquals |
TABLE 2. Specimens used for analysis in this study with variation in prey capture tactics and prey types among extant mysticetes. See cited references in Appendix 2.
Family | ID | Scientific name |
Abbreviation | Prey capture tactics |
Prey types | Specimen number or ID |
Reference |
Balaenidae | 0 | Balaena mysticetus | MYS | Skim | Small prey | - | Nishiwaki and Kasuya, 1970 |
1 | Eubalaena glacialis | EUG | Skim | Small prey | Plate XV, 1 | Omura, 1964 | |
2 | Eubalaena glacialis | EUG | Skim | Small prey | Plate XV, 3 | Omura, 1964 | |
3 | Eubalaena glacialis | EUG | Skim | Small prey | Plate XV, 4 | Omura, 1964 | |
4 | Eubalaena glacialis | EUG | Skim | Small prey | Plate XV, 5 | Omura, 1964 | |
5 | Eubalaena japonica | EUJ | Skim | Small prey | 61A | Omura et al., 1969 | |
6 | Eubalaena japonica | EUJ | Skim | Small prey | 61B | Omura et al., 1969 | |
7 | Eubalaena japonica | EUJ | Skim | Small prey | - | Omura, 1958 | |
Neobalaenidae | 8 | Caperea marginata | CAP | Skim | Small prey | NMV C40064 | Evans EvoMorph Lab. 2020 |
Eschrichtiidae | 9 | Eschrichtius robustus | ESC | Multiple | Small prey | M-804A | Nakamura and Kato, 2014 |
10 | Eschrichtius robustus | ESC | Multiple | Small prey | M-804B | Nakamura and Kato, 2014 | |
11 | Eschrichtius robustus | ESC | Multiple | Small prey | AMNH 34260 | Andrews, 1914 | |
12 | Eschrichtius robustus | ESC | Multiple | Small prey | - | Kienle et al 2015 | |
Balaenopteridae | 13 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | USNM 21492 | Cope, 1871 |
14 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 1 | Omura, 1964 | |
15 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 2 | Omura, 1964 | |
16 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 3 | Omura, 1964 | |
17 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 4 | Omura, 1964 | |
18 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 5 | Omura, 1964 | |
19 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 6 | Omura, 1964 | |
20 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | Plate XIV, 10 | Omura, 1964 | |
21 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | NMNS M 24369 | - | |
22 | Megaptera novaeangliae | MEG | Lunge | Both large and small prey | OMNH-M 3042 | - | |
23 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 1 | Omura, 1964 | |
24 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 2 | Omura, 1964 | |
25 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 3 | Omura, 1964 | |
26 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 4 | Omura, 1964 | |
27 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 6 | Omura, 1964 | |
28 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 7 | Omura, 1964 | |
29 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 8 | Omura, 1964 | |
30 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 9 | Omura, 1964 | |
31 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 10 | Omura, 1964 | |
32 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | Plate XIII, 11 | Omura, 1964 | |
33 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | - | Arnold et al., 1987 | |
34 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | NMNS M 24347 | - | |
35 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | NMNS M 24355 | - | |
36 | Balaenoptera acutorostrata | ACU | Lunge | Both large and small prey | NMNS M 24357 | - | |
37 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 4 | Satake and Omura, 1974 | |
38 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 5 | Satake and Omura, 1974 | |
39 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 6 | Satake and Omura, 1974 | |
40 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 7 | Satake and Omura, 1974 | |
41 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 8 | Satake and Omura, 1974 | |
42 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 9 | Satake and Omura, 1974 | |
43 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate I, 11 | Satake and Omura, 1974 | |
44 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate II, 15 | Satake and Omura, 1974 | |
45 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate II, 17 | Satake and Omura, 1974 | |
46 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate II, 24 | Satake and Omura, 1974 | |
47 | Balaenoptera bonaerensis | BON | Lunge | Small prey | Plate II, 25 | Satake and Omura, 1974 | |
48 | Balaenoptera borealis | BOL | Multiple | Both large and small prey | Plate VI, 3 | Omura, 1964 | |
49 | Balaenoptera borealis | BOL | Multiple | Both large and small prey | Plate VI, 8 | Omura, 1964 | |
50 | Balaenoptera borealis | BOL | Multiple | Both large and small prey | Plate VIII, 1 | Omura, 1964 | |
51 | Balaenoptera borealis | BOL | Multiple | Both large and small prey | Plate VIII, 10 | Omura, 1964 | |
52 | Balaenoptera borealis | BOL | Multiple | Both large and small prey | - | Nishiwaki and Kasuya, 1971 | |
53 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate X, 6 | Omura, 1964 | |
54 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate X, 10 | Omura, 1964 | |
55 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate X, 11 | Omura, 1964 | |
56 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate XI, 1 | Omura, 1964 | |
57 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate XI, 5 | Omura, 1964 | |
58 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate XI, 7 | Omura, 1964 | |
59 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate XI, 9 | Omura, 1964 | |
60 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | Plate XI, 10 | Omura, 1964 | |
61 | Balaenoptera edeni | EDE | Lunge | Large prey (fish prymarily) | 77N62, Plate 1 | Omura et al., 1981 | |
62 | Balaenoptera ricei | RIC | Unknown | Unknown | USNM 594665 | Rosel et al., 2021 | |
63 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 1 | Omura, 1964 | |
64 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 2 | Omura, 1964 | |
65 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 3 | Omura, 1964 | |
66 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 5 | Omura, 1964 | |
67 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 6 | Omura, 1964 | |
68 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | Plate I, 8 | Omura, 1964 | |
69 | Balaenoptera musculus brevicauda | MUS | Lunge | Small prey | - | Omura et al., 1970 | |
70 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate II, 3 | Omura, 1964 | |
71 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate II, 4 | Omura, 1964 | |
72 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate II, 5 | Omura, 1964 | |
73 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate II, 7 | Omura, 1964 | |
74 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate II, 9 | Omura, 1964 | |
75 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate IV, 8 | Omura, 1964 | |
76 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate IV, 10 | Omura, 1964 | |
77 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate IV, 12 | Omura, 1964 | |
78 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | Plate IV, 13 | Omura, 1964 | |
79 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | OMNH-M 1000 | - | |
80 | Balaenoptera physalus | PHY | Lunge | Both large and small prey | OMNH in the 3rd exhibition room | - | |
81 | Balaenoptera omurai | OMU | Lunge | Both large and small prey | NMNS | Shinohara, 2011 | |
Extinct taxa | |||||||
Balaenopteridae | 82 | "Megaptera" hubachi | HUB | Unknown | Unknown | MPST 240505 | Bisconti, 2010 |
Cetotheriidae | 83 | Piscobalaena nana | PIS | Unknown | Unknown | MNHN SAS 1617 | Bouetel and de Muizon, 2006 |
Eomysticetidae | 84 | Yamatocetus canaliculatus | YAM | Unknown | Unknown | KMNH VP 000,017 | Okazaki, 2012 |
- | 85 | Mysticeti indet. | NFL | Unknown | Unknown | NFL 2634 | Shinohara, 2011 |
TABLE 3. P-values for pairwise Hotelling’s tests. Significant values (<.005) are in bold.
Invertebrates | Both | Fish | |
Invertebrates | 8.26E-07 | 5.23E-10 | |
Both | 8.26E-07 | 2.36E-09 | |
Fish | 5.23E-10 | 2.36E-09 | |
Skim | Multiple | Lunge | |
Skim | 0.056496 | 7.05E-05 | |
Multiple | 0.056496 | 0.035062 | |
Lunge | 7.05E-05 | 0.035062 |
TABLE 4. Qualitative morphological comparison. The articular process is categorized by the ratio of the width and length of the process (Very short [Over 3.0], Short [2.9 to 1.2], Long [1], Very long [less than 1]). The notch between the articular process is categorized by the ratio of the width and length of the notch (Narrow [less than 0.3], Wide [over 0.5]).
Scientific name | articular process | notch between the articular processes |
processes at the posterior margin |
orientation of lateral portion |
lateral portion anteroposerior expansion |
Extant taxa | |||||
Balaena mysticetus | short | wide | absent | posterolaterally | absent |
Eubalaena glacialis | short | wide | small | posterolaterally | absent |
Eubalaena japonica | short | wide | small | posterolaterally | absent |
Caperea marginata | short | wide | small | posterolaterally | absent |
Eschrichtius robustus | short | wide | small or absent | posterolaterally | absent |
Megaptera novaeangliae | short | wide | small or absent | laterally / posterolaterally | absent |
Balaenoptera omurai | short | wide | large and pointed | laterally | absent |
Balaenoptera bonaerensis | short | wide | small or absent | laterally | absent |
Balaenoptera acutorostrata | short | wide | small or large | laterally / posterolaterally | absent |
Balaenoptera physalus | long to short | wide to narrow | large and pointed | posterolaterally | absent |
Balaenoptera musculus | short | wide | large and pointed | posterolaterally | present |
Balaenoptera ricei | short | narrow | large and pointed | laterally | absent |
Balaenoptera borealis | long to short | wide to narrow | large and wide | anterolaterally | absent |
Balaenoptera edeni | very long | narrow | large and wide | anterolaterally | absent |
Extinct taxa | |||||
"Megaptera" hubachi | short | wide | absent | laterally | absent |
Pelocetus calvertensis | short | wide | absent | laterally | absent |
Piscobalaena nana | short | wide | absent | posterolaterally | absent |
NFL 2634 | short | wide | absent | posterolaterally | absent |
Yamatocetus canaliculatus | very short | wide | absent | more posteriorly | absent |
Fucaia buelli | very short | wide | - | - | absent |
TABLE 5. Estimated feeding strategies for extinct taxa and Balaenoptera ricei.
Family | ID | Scientific name | Estimated prey types | Specimen number |
Balaenopteridae | 62 | Balaenoptera ricei | Both large and small prey | USNM 594665 |
Balaenopteridae | 82 | "Megaptera" hubachi | Both large and small prey, or small prey | MPST 240505 |
Cetotheriidae | 83 | Piscobalaena nana | Both large and small prey | MNHN SAS 1617 |
Eomysticetidae | 84 | Yamatocetus canaliculatus | Small size prey | KMNH VP 000,017 |
- | 85 | Mysticeti indet. | Both large and small prey, or small prey | NFL 2634 |