Character list and data matrix:
[1] Premaxillary teeth present (0), absent (1) (irreversible)
[2] Parapophyses located in the cranial (0) or central (1) part of the bodies of dorsal vertebrae
[3] Fibula does (0) not (1) reach calcaneum
[4] Humeral head cranially concave and caudally convex (0) or globose (1)
[5] Scapula and coracoid sutured or fused (0), forming a flat joint (1), a joint coracoid in scapula (2), a shallow joint scapula in coracoid (3) or a deep joint scapula in coracoid (4) (see stepmatrix)
[6] Supracoracoid nerve foramen not (0) opening into an elongate furrow medially and separated from the medial margin of the coracoid by a thick, bony bar (1)
[7] Broad, deep fossa on the dorsal side of the coracoid: absent (0), present (1)
[8] Supracoracoid nerve foramen opens above the coracoidal fossa (0) or inside it (1)
[9] Prominent scapular acromion (0) absent, costolaterally not (1) wider than deep (2) (ordered)
[10] Costal surface of scapular blade not (0) with a prominent, longitudinal
furrow (1)
[11] Dorsal and ventral margins of the furcula: subequal in width (0), ventral margin clearly wider than the dorsal margin (1)
[12] Hypocleideum: absent or poorly developed (0) or well developed (1)
[13] Ossified sternal keel: absent (0), not (1) reaching the cranial border of the sternum or beyond it (2) (ordered)
[14] Manual digit I reaches beyond mc II (0), +- reaches its end (1) or is markedly shorter (2) (ordered)
[15] Manual digit III: 4 separate phalanges (0), 3 (1), 2 (2), 1 (3) (irreversible)
[16] Coracoid without (0), with a tiny (1) or with a cylindrical part longer than 1/3 of the coracoid (2)
(ordered)
[17] Dorsal margin of the humeral head not (0) concave in its central portion, rising ventrally and dorsally
(1)
[18] Prominent bicipital crest of humerus, cranioventrally projecting: absent (0), present
(1)
[19] Ventral face of the humeral bicipital crest not (0) with a small fossa for muscular attachment
(1)
[20] Distal end of humerus not (0) very compressed craniocaudally (1)
[21] Dorsal cotyla of ulna not (0) separated from the olecranon by a groove
(1)
[22] Shaft of radius not (0) with a long longitudinal groove on its caudoventral surface
(1)
[23] Pygostyle: absent (0), middle (1), long (2), short (3) (see
stepmatrix)
[24] Mc III not (0) projecting distally more than mc II (1)
[25] Manual digit II: proximal phalanx shorter (0) or longer (1) than intermediate
phalanx
[26] Pubic foot: present (0), absent (1)
[27] Femoral posterior trochanter: absent or weakly developed (0), hypertrophied
(1)
[28] Caudal projection of the lateral border of the distal end of the femur: absent (0), present
(1)
[29] Very narrow, deep intercondylar sulcus on tibiotarsus that proximally undercuts condyles: absent (0), present
(1)
[30] Trochlea of mt II not (0) much broader than the trochlea of mt III
(1)
[31] Mt IV not (0) significantly thinner than mt II & III (1)
[32] Medial rim of the trochlea of mt III with a strong plantar projection: absent (0), present
(1)
[33] Mt I not (0) distinctly J-shaped in medial aspect (1)
[34] Mt V present (0), absent (1) (irreversible)
[35] Strong transverse convexity of the dorsal surface of the midshaft of
mt III: absent (0), present
(1)
[36] Distal end of mt II not (0) strongly curved medially (1)
[37] Sacral vertebrae: 5 (0), 6 (1), 7 (2), 8 (3), 9 (4), 10 or more (5)
(ordered)
[38] Pedal digit II without the claw not (0) significantly shorter than IV
(1)
[39] Largest pedal claw on digit III (0), II (1), I (2) (unordered)
[40] Pedal digit II not (0) hyperextensible (1)
[41] Distal fusion between mc II and III: absent (0), present (1)
[42] Distal fusion between mt II, III and IV: absent (0), ontogenetically after (1) or before proximal fusion (2)
(unordered)
[43] Patella(r groove on femur): absent (0), present (1)
[44] Hypotarsus: absent (0), present (1)
[45] Proximal end of mt III not (0) broadened (1), displaced plantarly (2) or narrowed (subarctometatarsal condition) (3) (see stepmatrix)
[46] Middle tail vertebrae not (0) at least twice as long as the proximal ones
(1)
[47] Extensor process on mc I: absent (0), present (1)
[48] Sterna sutured (0) or fused (1)
[49] Procoracoid process: absent (0), present (1)
[50] Pubic symphysis more (0), less than 1/3 of pubic length (1) or absent (2)
(irreversible)
[51] Femoral neck: present (0), absent (1)
[52] More (0) or fewer (1) than 10 free caudal vertebrae (irreversible)
[53] Preacetabular process of ilium less than (0) twice as long as postacetabular process
(1)
[54] Bicipital scar on ulna not (0) developed as a tubercle (1)
[55] Ventral tubercle of humerus: absent (0), present (1)
[56] Coracoid less (0), more (1) than 1.75 times as long (glenoid to sternum) as broad (maximal distance craniomedial to caudolateral
margin)
[57] Articular pneumaticity: absent (0), present (1)
[58] Intercotylar eminence: absent (0), present (1)
[59] Fossa for mt I on distal plantar surface of mt II: absent (0), present
(1)
[60] Flange on phalanx II-1 (for attachment of remiges): absent (0), present
(1)
[61] Ilia: close together over entire length (0), far apart and vertical over entire length (1) or sigmoid, close together cranially to the acetabulum and far apart caudally (2)
(unordered)
[62] Pubic peduncle of ilium about 1 (0) or 3 (1) times as long craniocaudally as the ischiadic one (measured at the base = dorsal
end)
[63] Transition point near the middle (1), base (0) or end of the tail (2)
(ordered)
[64] Tail distal to transition point has: short, broad prz (or none) (0), short,
narrow prz (1), long prz and poz (2) (unordered)
[65] Dorsal vertebrae with connective tissue bridging transverse processes:
absent (0), present (1)
[66] Lateral process of coracoid: absent (0), present (1)
[67] Toes (0) shorter, (1) longer than metatarsus
5 |
10 |
15 |
20 |
25 |
30 |
35 |
40 |
45 |
50 |
55 |
60 |
65 |
67 | |
allzero outgroup |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00000 |
00100 |
00 |
Scansoriopterygidae |
??0?0 |
????? |
00?00 |
000?0 |
???10 |
0???0 |
0?000 |
00010 |
00??{03} |
10?0? |
100?? |
0?000 |
??020 |
00 |
Archaeopteryx |
00000 |
00?10 |
00000 |
00000 |
00000 |
00000 |
00000 |
0010? |
00003 |
10000 |
1010? |
(01)?000 |
11010 |
01 |
Rahonavis ostromi |
?01?2 |
???1? |
????? |
????? |
00??? |
0??00 |
000?0 |
01111 |
?0000 |
1??00 |
1010? |
??00? |
?1010 |
?1 |
Shenzhouraptor sinensis |
1?1?{123} |
0001? |
00010 |
00??0 |
??001 |
0??00 |
0?000 |
011{01}? |
00000 |
10000 |
101?? |
10001 |
?1010 |
01 |
| Jixiangornis orientalis | 0???? | ????? | ?0100 | 2???0 | ??000 | ????0 | 0???? | 0211? | 00??{01} | 10100 | ?0??? | 0???? | 1?010 | 01 |
Hulsanpes perlei |
????? |
????? |
????? |
????? |
????? |
????0 |
00??? |
0???1 |
?0?01 |
????? |
????? |
??0?? |
????? |
?? |
Yandangornis longicaudus |
1?1?? |
????? |
??0?{01} |
????0 |
???0? |
00?00 |
0??00 |
0?110 |
00001 |
??1?{01} |
00??0 |
??00? |
????? |
?0 |
Sapeornis chaoyangensis |
0?000 |
???00 |
00?02 |
01000 |
00101 |
0?000 |
0??00 |
02000 |
00000 |
00?01 |
11(01)0? |
0?0?1 |
?1200 |
01 |
| Omnivoropteryx sinousaorum | 0?0?? | ????? | ????1? | ????0 | ???00 | 0???0 | 0???? | 1?10? | ?0??? | ????1 | ????? | ??0?1 | ????0 | ?1 |
Confuciusornis sanctus |
10100 |
00?00 |
00(01)00 |
10001 |
00200 |
0?000 |
00000 |
12110 |
00001 |
00101 |
01010 |
10001 |
10200 |
01 |
Jibeinia luanhera |
0?10? |
???{12}? |
??011 |
2?1?? |
??201 |
0???? |
0??1? |
031{01}0 |
00??1 |
00101 |
11??1 |
1?0?? |
??{12}0? |
?1 |
Protopteryx fengningensis |
01101 |
?0??? |
?1102 |
21101 |
?0210 |
0???1 |
0000? |
0201? |
00001 |
00111 |
11??? |
1?0?1 |
??200 |
0? |
Longipteryx chaoyangensis |
0?102 |
?0??? |
01102 |
2?1?1 |
??211 |
0???? |
0??1? |
03??? |
0000{01} |
00101 |
?10?1 |
1?0?1 |
1?{12}?0 |
0? |
Eocathayornis walkeri |
01?02 |
?11{12}0 |
??112 |
21111 |
?1{12}11 |
????? |
????? |
????? |
0???? |
?010? |
????1 |
1???1 |
????0 |
0? |
Iberomesornis romerali |
?010? |
????? |
?11?? |
20001 |
??2?? |
?1??1 |
0??1? |
03100 |
?00?1 |
0?10? |
?10?? |
1?0?? |
{12}0200 |
00 |
Noguerornis gonzalezi |
???0? |
????? |
11??? |
?10?1 |
?0?1? |
????? |
????? |
????? |
0???? |
?0??? |
????? |
????? |
????? |
?? |
Halimornis thompsoni |
?1??2 |
???21 |
????? |
?111? |
??2?? |
??0?? |
????? |
????? |
????? |
0???? |
????? |
????? |
??{01}00 |
?? |
Cuspirostrisornis houi |
0?1?? |
???{12}? |
??22? |
2?01? |
??21? |
0???? |
1??1? |
?{234}?2? |
00?0? |
001?1 |
0???? |
1?0?? |
??{12}?? |
01 |
Concornis lacustris |
?1102 |
10?2? |
1112? |
21111 |
1???1 |
1?1?1 |
1?011 |
0??20 |
000?1 |
?0101 |
0??11 |
1?0?1 |
????? |
00 |
Sinornis santensis |
01102 |
10010 |
?1123 |
21111 |
?1211 |
01011 |
1101? |
0310? |
00001 |
00101 |
01011 |
1?0?1 |
?0200 |
01 |
| Longirostravis hani | 0?1?? | ???1? | 111?? | 21??? | ??11? | ???10 | 0???? | 12??? | 00??1 | 001?? | 010?? | 1?0?? | ??100 | 0? |
| Vescornis hebeiensis | 0???{123} | ????1 | 1112{23} | 21111 | ??211 | ??1?1 | 0?1?0 | 13010 | ?1??1 | 0010? | 1???1 | 1?001 | ??20? | 01 |
Gobipteryx minuta |
1?1?2 |
?001? |
11??? |
2???? |
?111? |
1??01 |
010?0 |
1{45}??? |
?0?0? |
0??01 |
????? |
100?? |
????? |
0? |
Lectavis bretincola |
??1?? |
????? |
????? |
????? |
????? |
???0? |
1???0 |
????? |
?{01}?11 |
????? |
????? |
??0?? |
????? |
?? |
Soroavisaurus australis |
??1?? |
????? |
????? |
????? |
????? |
???11 |
111?1 |
1??2? |
?0?01 |
????? |
????? |
??0?? |
????? |
?? |
Yungavolucris brevipedalis |
????? |
????? |
????? |
????? |
????? |
????1 |
10??0 |
0???1 |
?0?01 |
????? |
????? |
??0?? |
????? |
?? |
Avisaurus gloriae |
????? |
????? |
????? |
????? |
????? |
????1 |
11??1 |
1???? |
?1?01 |
????? |
????? |
??0?? |
????? |
?? |
Enantiornis leali |
???02 |
10021 |
????? |
21110 |
11?1? |
????? |
????? |
????? |
00??? |
?0?0? |
???01 |
1???? |
????? |
0? |
Neuquenornis volans |
?1??2 |
10111 |
102?? |
2???1 |
?1?1? |
?11?1 |
111?1 |
0??2? |
000?? |
?0101 |
????1 |
1??01 |
????? |
0? |
Boluochia zhengi |
1?1?? |
????? |
??1?? |
????? |
??2?? |
0??11 |
00?1? |
0???? |
?0??1 |
001?? |
0???? |
??0?? |
??20? |
?1 |
Otogornis genghisi |
???0{34} |
1???? |
???2? |
211?1 |
???0? |
????? |
????? |
????? |
0???? |
???0? |
????1 |
1???? |
????? |
0? |
Alexornis antecedens |
????2 |
???{12}? |
????? |
2???? |
1???? |
????? |
????? |
????? |
??0?? |
???0? |
????? |
1???? |
????? |
?? |
Eoalulavis hoyasi |
???02 |
1011? |
11123 |
21111 |
?1?11 |
?1??? |
????? |
????? |
0???? |
?010? |
??001 |
1???1 |
2???0 |
0? |
Eoenantiornis buhleri |
0?10? |
?0??? |
11?1{23} |
{12}???? |
?1211 |
01??1 |
0??1? |
?{123}??? |
00??1 |
0010? |
?1??? |
1???1 |
???0? |
0? |
unnamed Lebanon |
????? |
10?20 |
?1??? |
2???? |
????? |
????1 |
0???? |
0??2? |
?0??1 |
??10? |
????? |
1?0?? |
????? |
01 |
Liaoningornis longidigitis |
??1?? |
????? |
??2?? |
????? |
????? |
0???0 |
0??1? |
1?00? |
?1??1 |
??1?? |
0???? |
??0?? |
????? |
01 |
Apsaravis ukhaana |
?0113 |
101{12}? |
??2?? |
20111 |
003?? |
1??10 |
0???0 |
150?0 |
?2112 |
01102 |
01011 |
1?0?1 |
2?{12}{01}? |
0? |
Ambiortus dementjevi |
???1{34} |
0???? |
002?? |
200?? |
????1 |
????? |
????? |
????? |
????? |
??11? |
????? |
1???1 |
????? |
?? |
Yanornis martini |
0?1?{34} |
?0??? |
?0203 |
2?0?0 |
??301 |
1???0 |
0?0?0 |
0410? |
12??2 |
00111 |
????? |
1???1 |
2??{01}? |
01 |
Yixianornis grabaui |
0?1?{34} |
?0??? |
?0213 |
20??0 |
??301 |
1???0 |
0?0?0 |
0410? |
12?02 |
01111 |
??0?? |
1?0?1 |
???{01}? |
01 |
Patagopteryx deferrariisi |
?0103 |
00?21 |
????? |
200?1 |
00?00 |
1?010 |
000?0 |
04110 |
12011 |
0??02 |
0?01? |
10?00 |
20{12}?0 |
?1 |
Vorona berivotrensis |
??1?? |
????? |
????? |
????? |
????? |
?0110 |
00?00 |
0???0 |
?2001 |
????? |
0???? |
??00? |
????? |
?? |
Songlingornis linghensis |
0???{34} |
01??? |
002?? |
2???? |
????? |
????? |
????? |
????? |
?{12}??? |
??11? |
????? |
1???? |
????? |
0? |
Gansus yumenensis |
??1?? |
????? |
????? |
????? |
????? |
???00 |
00??0 |
0?110 |
?2?12 |
????? |
????? |
??10? |
?????? |
?1 |
Hesperornithes |
101?3 |
01?0? |
000?3 |
10000 |
??30? |
10000 |
00010 |
05100 |
?2112 |
0?112 |
010?0 |
0(01)11? |
20200 |
01 |
Ichthyornis dispar |
10114 |
0100? |
00223 |
20010 |
00301 |
10000 |
00010 |
15??? |
12112 |
01112 |
01011 |
11111 |
20201 |
1? |
Anas platyrhynchos |
10114 |
01?1? |
00223 |
20000 |
?0301 |
10000 |
00010 |
05000 |
12112 |
01112 |
01011 |
11111 |
20201 |
11 |
Microraptor |
00000 |
???1? |
00000 |
0?0?0 |
??000 |
00?00 |
0??00 |
00111 |
00003 |
10(01)00 |
100?1 |
??0?1 |
11020 |
00 |
Caudipteryx |
0?000 |
???0? |
00002 |
0???0 |
??{01}00 |
0??00 |
0?000 |
00?00 |
00003 |
00000 |
000?? |
0?0?0 |
?0000 |
00 |
basal Troodontidae |
00000 |
???{12}? |
0?000 |
000?0 |
?0?00 |
00000 |
001?0 |
10111 |
00003 |
10?00 |
?000? |
??000 |
1(01)000 |
00 |
{} = uncertainty (one of the character states occurs)
() = polymorphism (all character states occur)
Taxon comments:
Scansoriopterygidae: Composite of Scansoriopteryx heilmanni and Epidendrosaurus ningchengensis
– very similar, identical apart from "?" in this matrix.
Archaeopteryx: A. lithographica and A. bavarica together. Only difference except for position of
"?" :
A. lithographica has 56(0), A. bavarica has
(1). Therefore polymorphism.
Iberomesornis romerali: Character 67 after measurements by Sereno
2000. Consistent with longish mt; impossible to see where bone ends and horn
sheath continues alone in illustrations.
Hesperornithes: Composite of Hesperornis regalis
and Baptornis advenus. Differences in missing data and inapplicable
codings, and perhaps in character 57.
Microraptor: Composite of Microraptor zhaoianus and Microraptor gui
– identical apart from "?" in this
matrix.
Caudipteryx: Species-level taxonomy (C. zoui, C. dongi, C. sp.) uncertain and ignored. All differences would be in the positions of
"?"
.
basal Troodontidae: Coded after Sinovenator and Sinornithoides. Only difference except for position of
"?" :
-venator has 62(1), -ornithoides appears to have (0). Therefore
coded as polymorphism.
Character comments:
8: In case of 65(0) the theoretical position of the coracoid fossa was
guessed.
13: In absence of ossified sternum coded as 0.
52: If not irreversible, Yandangornis likes to cluster with Confuciusornis. That said, it likes to cluster with
C. anyway.
53: Preacetabular process measured from the cranialmost point of the ilium to the cranial border of the acetabulum. Postacetabular process measured from the caudalmost point of the ilium to the caudal border of the
ischiadic peduncle of the ilium.
61: In the absence of visible ilia, coded from the length of the sacral ribs.
63: Ordered in this way – plesiomorphy = (1) – to spare me a stepmatrix.
Occasionally the transition point is in the pygostyle.
64: Should perhaps be
ordered like 63... Pygostyles coded as (0) because of Halimornis, Sinornis,
Boluochia? etc..
Stepmatrices:
character 5 character
23 character 45
0 1 2 3 4 0 1 2 3
0 1 2 3
- 1 1 2 3 - 1 2
2 - 1 2 1
1 - 1 1 2 1 - 1
1 1 - 1 2
1 1 - 2 3 2 1 -
1 2 1 - 1
2 1 2 - 1 1 1 2
- 1 2 2 -
3 2 3 1 -
Cladograms:
Yeah, er. Coming.
Strict consensus when all taxa are included and when Alexornis, Hulsanpes, Lectavis and Yungavolucris are deleted: Grass.
Questions? Including requests for the NEXUS file?
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