@ARTICLE{TreeBASE2Ref16892,
author = {Maureen A. O'Leary and Jonathan H. Geisler},
title = {The Position of Cetacea Within Mammalia: Phylogenetic Analysis of Morphological Data from Extinct and Extant Taxa.},
year = {1999},
keywords = {Artiodactyla; astragalus; Cetacea; fossils; homoplasy; Mesonychia; morphology; phylogeny},
doi = {10.1080/106351599260102},
url = {},
pmid = {},
journal = {Systematic Biology},
volume = {48},
number = {3},
pages = {455--490},
abstract = {Knowledge of the phylogenetic position of the order Cetacea (whales, dolphins, and porpoises) within Mammalia is of central importance to evolutionary biologists studying the transformations of biological form and function that accompanied the shift from fully terrestrial to fully aquatic life in this clade. Phylogenies based on molecular data and those based on morphological data both place cetaceans among ungulates but are incongruent in other respects. Morphologists argue that cetaceans are most closely related to mesonychians, an extinct group of terrestrial ungulates. They have disagreed, however, as to whether Perissodactyla (odd-toed ungulates) or Artiodactyla (even-toed ungulates) is the extant clade most closely related to Cetacea, and have long maintained that each of these orders is monophyletic. The great majority of molecule-based phylogenies show, by contrast, not only that artiodactyls are the closest extant relatives of Cetacea, but also, that Artiodactyla is paraphyletic unless cetaceans are nested within it, often as the sister group of hippopotamids. We tested morphological evidence for several hypotheses concerning the sister taxon relationships of Cetacea in a maximum parsimony analysis of 123 morphological characters from 10 extant and 30 extinct taxa. We advocate treating certain multistate characters as ordered because such a procedure incorporates information about hierarchical morphological transformation. In all most-parsimonious trees, whether multistate characters are ordered or unordered, Artiodactyla is the extant sister taxon of Cetacea. With certain multistate characters ordered, the extinct clade Mesonychia (Mesonychidae + Hapalodectidae) is the sister taxon of Cetacea, and Artiodactyla is monophyletic. When all fossils are removed from the analysis, Artiodactyla is paraphyletic with Cetacea nested inside, indicating that inclusion of mesonychians and other extinct stem taxa in a phylogenetic analysis of the ungulate clade is integral to the recovery of artiodactyl monophyly. Phylogenies derived from molecular data alone may risk recovering inconsistent branches because of an inability to sample extinct clades, which by a conservative estimate amounts to 89% of the ingroup. Addition of data from recently described cetacean astragali attributed to cetaceans does not overturn artiodactyl monophyly. [Artiodactyla; astragalus; Cetacea; fossils; homoplasy; Mesonychia; morphology; phylogeny.]}
}
Matrix 2174 of Study 589

Citation title:
"The Position of Cetacea Within Mammalia: Phylogenetic Analysis of Morphological Data from Extinct and Extant Taxa.".

This study was previously identified under the legacy study ID S387
(Status: Published).
Matrices
Title: Fig. 4
Description: Legacy TreeBASE Matrix ID = M540
Rows
|
Taxon Label |
Row Segments |
Characters 1?–30 |
| Didelphis |
(none)
|
00001100000?????0??00101?10??0 |
| Equus |
(none)
|
010011000110?20010200011?3100? |
| Hippopotamus |
(none)
|
110?111031100031102100{01}1?30 |
| Hexaprotodon |
(none)
|
11????1031100031??210{01}11?2? |
| Sus |
(none)
|
000111100110020010210021?20??0 |
| Camelus |
(none)
|
000111002110010111100001?00??0 |
| Ovis |
(none)
|
1001010?2110013010200001?10??0 |
| Tragulus |
(none)
|
0?01000031100{12}011?100001?00 |
| Balaenoptera |
(none)
|
121311112111103011011121?00??0 |
| Tursiops |
(none)
|
121311110111123011001121??0??0 |
Columns
| Column |
Character Description |
|
1
|
Subarcuate Fossa
|
|
2
|
shape Tegmen Tympani
|
|
3
|
Anterior Proc Petr
|
|
4
|
Tensor Tympani Fossa
|
|
5
|
transpromontorial sulcusICA
|
|
6
|
Stapedial Groove
|
|
7
|
Mastoid process
|
|
8
|
Stylomastoid Foramen
|
|
9
|
Facial Nerve Sulcus
|
|
10
|
Pars cochlearis/Basioccpital
|
|
11
|
Ectotympanic Bulla
|
|
12
|
Pachyo Bulla
|
|
13
|
Lat furrow Bulla
|
|
14
|
Med furrow Bulla
|
|
15
|
Art Bulla to Squamosal
|
|
16
|
ECtoTymp/Exocc contact
|
|
17
|
Sigmoid Process
|
|
18
|
Sigmoid process shape
|
|
19
|
Ectotympanic meatal floor
|
|
20
|
External Auditory Meatus squamo
|
|
21
|
Basioccipital crests (falcate p
|
|
22
|
Internal Carotid Foramen
|
|
23
|
Post-Glenoid Foramen
|
|
24
|
PRESENCE For ram sup.
|
|
25
|
POSITION for ram sup
|
|
26
|
Posit For Ovale
|
|
27
|
Alisphenoid Canal
|
|
28
|
Ant Open Alar Canal
|
|
29
|
Post Open Alar Canal
|
|
30
|
Foramen Rotundum
|
|
31
|
Mastoid Foramen
|
|
32
|
Posttemporal canal
|
|
33
|
Preglenoid process
|
|
34
|
Supraorbital process
|
|
35
|
posorbital bar
|
|
36
|
Contact Front/Max
|
|
37
|
Lacrimal Tubercle
|
|
38
|
palatine fissure
|
|
39
|
Post margin nares
|
|
40
|
Mandibular Foramen
|
|
41
|
condyle
|
|
42
|
Dentary angle
|
|
43
|
Upp Inc 1-2
|
|
44
|
Incisor Alignment
|
|
45
|
Embrasure pits
|
|
46
|
P1 presence
|
|
47
|
P3 roots
|
|
48
|
P4 protocone
|
|
49
|
P4 paracone height
|
|
50
|
P4 metacone
|
|
51
|
stylar shelf
|
|
52
|
M2 metacone
|
|
53
|
M1 parastyle
|
|
54
|
ling cing M2
|
|
55
|
Ectocingula
|
|
56
|
Hypocone M1M2
|
|
57
|
Trigon basin
|
|
58
|
Paraconule M2/
|
|
59
|
Metaconule M2/
|
|
60
|
M3
|
|
61
|
lower canines
|
|
62
|
lower incisors
|
|
63
|
lower Canine shape
|
|
64
|
p1
|
|
65
|
p3 metaconid
|
|
66
|
p4 metaconid
|
|
67
|
lin cing molars
|
|
68
|
m1 parac
|
|
69
|
m1 parac/crist posit
|
|
70
|
m2 parac
|
|
71
|
m2 parac/crist pos
|
|
72
|
m3 parac
|
|
73
|
m3 parac/cris posit
|
|
74
|
m1 metaconid
|
|
75
|
m2 metaconid
|
|
76
|
m3 metaconid
|
|
77
|
m1 m2 hypoconulid
|
|
78
|
molar protoconid
|
|
79
|
protoloph
|
|
80
|
metaloph
|
|
81
|
ant low lophs
|
|
82
|
post low lophs
|
|
83
|
Reen groove
|
|
84
|
Talonid basin molar
|
|
85
|
M3 Hypoconulid
|
|
86
|
dp4
|
|
87
|
Elongate shear facets on molars
|
|
88
|
Cerv vert length
|
|
89
|
Art Sacral/Illium
|
|
90
|
Number Sacrals
|
|
91
|
Scapular Spine
|
|
92
|
Entepicondyle
|
|
93
|
Entepi foramen
|
|
94
|
Olecranon process
|
|
95
|
Olec. fossa
|
|
96
|
Proximal radius
|
|
97
|
Distal radius
|
|
98
|
Centrale
|
|
99
|
Manus
|
|
100
|
Greater troch
|
|
101
|
3rd troch
|
|
102
|
Distal Femur
|
|
103
|
Astragalar Canal
|
|
104
|
Navicular Facet
|
|
105
|
Sustentacular facet
|
|
106
|
Cuboid facet
|
|
107
|
Lateral process
|
|
108
|
Pes
|
|
109
|
Proxi astrag
|
|
110
|
Nav/Cub fusion
|
|
111
|
m3/m4 fusion
|
|
112
|
Metatarsal 1
|
|
113
|
Metatarsal 2
|
|
114
|
Metatarsal 5
|
|
115
|
Unguventral
|
|
116
|
Ungdorsal
|
|
117
|
Stomach epithelium
|
|
118
|
Lumen
|
|
119
|
Lumen diverticulation
|
|
120
|
Penile Cavernous Tissue
|
|
121
|
Hair
|
|
122
|
sebac glands
|
|
123
|
Primary Bronchi
|