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New postcranial bones of the extinct mammalian family Nyctitheriidae (Paleogene, UK): primitive euarchontans with scansorial locomotion

Jerry J. Hooker

Plain Language Abstract

Nyctitheres were mouse-sized insectivorous mammals that became extinct about 30 million years ago. Long known from shrew-like teeth, more recently ankle and heel bones, located by screenwashing sediments, suggested a relationship closer to tree shrews and a life mainly in the trees. Here new nyctithere limb and foot bones are described that more fully substantiate their tree-climbing abilities and, through phylogenetic analysis, show their relationship with the group combining primates, tree shrews and colugos, called the Euarchonta.

Resumen en Español

Nuevos huesos postcraneales de la extinta familia de mamíferos Nyctitheriidae (Paleógeno, Reino Unido): euarcontos primitivos con locomoción arborícola

Nuevos huesos postcraneales del Eoceno tardío y del Oligoceno temprano de la Cuenca Hampshire, Reino Unido, son identificados como pertenecientes a la familia extinta Nyctitheriidae. Anteriormente, los astrágalos y los calcáneos eran los únicos elementos adecuados para ser reconocidos de manera inequívoca, aparte de los elementos dentales y de las mandíbulas. Ahora, se han recolectado húmeros, radios, un metacarpiano, fémures, tibias distales, naviculares, cuboides, un ectocuneiforme, metatarsos y falanges, junto con astrágalos y calcáneos adicionales. Estos huesos se muestran como pertenecientes a una diversidad de taxones de Nyctitheriidae previamente nombrados a partir de restos dentales. El análisis funcional muestra que los Nyctitheriidae tenían articulaciones del hombro y de la cadera móviles, podrían realizar movimientos de pronación y supinación del radio, podrían revertir parcialmente el pie a nivel de las articulaciones astrágalo-calcáneo y del tobillo superior utilizando potentes músculos flexores, todo lo cual es indicativo de un estilo de vida trepador y de la posibilidad de realizar descensos de cabeza en superficies verticales. La escalada parece haber sido dominada por la flexión de los antebrazos y los pies. El análisis cladístico empleando una variedad de mamíferos euterios primitivos muestra que los Nyctitheriidae son euarcontos troncales, en lugar de corresponder a los Lipotyphla, grupo con el que habían sido previamente relacionados sobre la base de características dentales. No se confirman las primeras ideas que sostenían que estaban relacionados con los extintos Adapisoriculidae, recientemente considerados euterios troncales.

Palabras clave: Eutheria; Lipotyphla; placentario; Scandentia; musarañas; Soricidae

Traducción: Enrique Peñalver

Résumé en Français

Nouveaux os postcrâniens de la famille éteinte de mammifères Nyctitheriidae (Paléogène, Royaume-Uni): euarchontiens primitifs avec locomotion arboricol

De nouveaux os postcrâniens de l'Eocène supérieur et de l'Oligocène inferieur du bassin Hampshire, au Royaume-Uni, sont identifiés comme appartenant à la famille éteinte Nyctitheriidae. Auparavant, les astragales et calcanéums étaient les seuls éléments en dehors des dents et des mâchoires à être reconnu sans équivoque. Maintenant, des humérus, des radius, un métacarpien, des fémurs, des tibias distaux, des naviculaires, des cuboïdes, un ecto-cunéiforme, des métatarses et des phalanges, avec des astragales et calcanéums supplémentaires, ont été recueillies. Ceux-ci sont présentés comme appartenant à une diversité de taxons nyctithères précédemment nommé à partir de restes dentaires. L'analyse fonctionnelle montre que les nyctithères avaient les épaules et les hanches mobiles, le radius pouvait être en pronation et en supination, le pied pouvait être partiellement inversé au niveau des joints astragale-calcanéum et de la cheville supérieure à l'aide de puissants muscles fléchisseurs, tous signe d'un mode de vie arboricole et permettant une descente la tête la première sur des surfaces verticales. L'escalade semble avoir été dominée par la flexion des avant-bras et des pieds. Notre analyse cladistique utilisant une gamme de mammifères primitifs euthériens montre que les nyctithères sont des euarchontiens souches, plutôt que des lipotyphliens, avec qui ils avaient déjà été classés à partir de caractères dentaires. Les conceptions antérieures de relations avec les Adapisoriculidae éteint, récemment considéré comme eutheriens souches, ne sont pas confirmées.

Mots-clés: Eutheria; Lipotyphla; placentaire; Scandentia ; musaraignes; Soricidae

Translator: Kenny J. Travouillon

Deutsche Zusammenfassung

Neue postcraniale Knochen der ausgestorbenen Säugertierfamilie Nyctitheriidae (Paläogen, vereinigtes Königreich): primitive Euarchonte mit kletternder Lebensweise

Neue postcraniale Knochen aus dem späten Eozän und frühesten Oligozän des Hampshire Beckens des Vereinigten Königreichs gehören zu der ausgestorbenen Familie Nyctitheriidae. Bisher waren außer Zähnen und Kiefern nur die Astragali und Calcanea die einzig eindeutig bestimmten Elemente. Nun wurden Humeri, Radii, ein Metacarpale, Femora, distale Tibiae, Naviculare, Cuboide, ein Ectocuneiforme, Metatarsale and Phalangen zusammen mit zusätzlichen Astragali und Calcanea gesammelt. Es wird gezeigt, dass diese zu einer Diversität an nyctitheren Taxa gehören, die bisher über Zahnüberreste benannt wurde. Funktionsanalysen zeigen, dass Nyctitheren mobile Schulter-und Becken-Artikluationen hatten, den Radius pronieren und supinieren konnten und teilweise den Fuß am astragalocalcanealen und oberen Fußgelenk mit kräftigen Flexormuskeln invertierten. Dies alles weist auf eine kletternde Lebensweise hin und erlaubt kopfüber Bewegungen an vertikalen Flächen. Das Klettern scheint von der Flexion der Unterarme und Füße dominiert worden zu sein. Kladistische Analysen mit einer Reihe von primitiven eutherischen Säugetieren zeigen, dass Nyctiheren Stamm-Euarchonten sind, eher als Lipotyphle mit denen sie bisher auf der Basis von dentalen Merkmalen zusammengefasst waren. Frühere Vorschläge von Verwandtschaften mit den ausgestorbenen Adapisoriculidae, kürzlich als Stamm-Euarchonten bezeichnet, können nicht aufrecht erhalten werden.

Schlüsselwörter: Eutheria; Lipotyphla; placental; Scandentia; Spitzmäuse; Soricidae

Translator: Eva Gebauer

Arabic

in progress

Translator: Ashraf M.T. Elewa

 

 

APPENDIX 2.

Character-taxon matrix

Note that characters of the astragalus and navicular of Fordonia are from as yet undescribed material. Some characters unknown for Palaeosinopa and Cryptotopos have been substituted from Pantolestes and Scraeva respectively.

Outgroup

00000 00000 10000 00100 00000 00000 10110 00000 00100 10100 00011 10110
00010 00010 01000 30000 00000 01010 00000 00000 10001 00000 01000 00000
00020 00011 00000 00001 00000 00300 00000 10010 00100 00110 10000 00000
00000 00100 00000 00000 00000 00000 00010 00000 01011 10000 10110 00000
00000 00000 00000 00100 00000 00000 10000 00000 00000 10003 10002 10000
00000 00100                    

Ukhaatherium

00001 11000 20100 00100 00000 00110 20120 00000 00130 20000 01122 10110
00011 10010 0100? ?02?1 0??00 110?0 000?0 11010 100?1 10200 11?01 ?0??0
??000 10?2? 0001? 00000 ??000 10112 10200 10210 ?1000 0??0? 1?001 00211
?0001 00110 ?0011 ??31? ????? ??000 01010 01020 010?? ?1??? ?01?? 0?001
?11?? 0??01 100?0 0???0 00210 00000 0?000 10000 00010 10203 1?002 10000
????? ?0100                    

Maelestes

??201 10110 20000 00001 00110 00000 20000 10011 01021 12100 01022 10110
00112 11010 01101 31210 00201 ?101? ??0?? ??000 ??112 10200 31101 10000
01?0? ??020 ???1? 00110 ???10 ?0010 00200 10010 30100 00000 ??001 00011
20021 00?0? 00?11 12210 0012? ??000 ?1111 11020 01100 11001 0?01? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Asioryctes

00000 01000 20100 00100 00000 00100 20110 10000 00030 20000 01122 10110
00011 10010 01001 31111 10000 11010 ?0000 01010 100?1 10200 110?1 00000
0??00 10021 0001? 00000 12000 10112 1?200 10210 01000 00?01 ??001 00211
10011 001?0 000?? ??311 00110 ????? ????? ????? ????? ????? ????0 00???
????? ????? ????? 0010? ??2?0 ????? 0???? ??010 0?010 10002 1?002 ?0120
????0 00?00                    

Zhelestidae

4?001 10100 100?1 10031 00100 00000 20121 10021 01121 21201 10121 10110
01122 01011 12102 40000 10000 11010 ?0?00 1???? ????? 1?2?0 ?0??? ?????
?100? ???10 ????? ????? ????? ????? ????? 10?1? 00000 00??? ?0?00 00?00
0???? ????? ??0?1 022?? ????? ????? ????? ??001 10100 100?? ????? ????2
11121 11000 1000? ????? ????? ????? ????? ????1 10043 01013 10021 20100
????? ?????                    

Zalambdalestidae

411?0 01100 20100 00030 00000 00000 11122 00111 01120 20200 01131 10100
00114 01011 01102 20001 41101 11010 00000 00000 00021 10100 01101 20010
10020 20010 00010 00010 12000 ?0112 10200 10210 11000 00000 1?001 00211
20000 00110 200?1 ??111 00021 ?011? ?0111 11001 11100 2000? 0??20 10002
01022 01010 00001 30400 00023 00110 21200 01002 10153 01030 10021 10100
?0021 00?00                    

Cimolestes magnus

???01 11?0? 20000 10200 00011 10000 21000 10001 00121 21200 01121 10210
00002 11000 02002 300?1 10?01 2???? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Batodon

????? 11?10 ?0110 10000 00011 01000 11001 10011 01121 22200 11111 10110
00012 11010 01000 4???? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Gypsonictops

???0? 10000 21202 00022 00110 00010 41123 30111 01221 22201 11122 10100
00113 11010 11102 401?1 1010? 2?0?? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Deccanolestes

????? 11?0? 2010? ????? ????? ?0??? 1000? 10101 01110 20200 01110 20110
00012 11010 0100? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ??0?0 12012 110?? ????? ?????
????? ????? ????? ????? 00211 00000 10101 11002 10120 11003 10001 10120
????? ?????                    

Afrodon/Bustylus

????? ?1?1? ????0 20013 00010 00000 10211 40101 01110 20200 00111 20110
00012 11010 01010 5???1 ???0? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ??010 00020 12012 1?0?? ????? ?????
??0?1 10110 00??? ????? 00211 00000 10101 1?00? 10120 11003 10001 ?0120
????? ?????                    

Fordonia

????? ?1??? ?1??2 1?022 10110 00110 31122 40011 01221 22201 21121 10000
00013 00010 0110? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? 11123 00110 ?0110 041?? ????? ????? ?????
??01? ?????                    

Leptictidium

30300 21000 20??2 10022 00110 01010 01112 30111 21211 23211 21120 20100
01023 11010 03014 52021 40101 2112? 0???1 ?1??? ????? ???00 ?1??? ?????
???2? ????? ??1?? ????? ????0 ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ??1?? 01?1? ?011? 11?3? 1???? ????? ????? ??01?
?0??2 01020 001?1 11210 ????? ????? ????? ????2 101?? 01021 ???10 ?0111
0???1 00100                    

Prodiacodon/Palaeictops

31301 21110 22302 20002 10110 00113 21222 31101 01121 12201 11122 10100
01113 11010 01102 41111 00101 21220 010?1 ?1??? ????1 0??00 ?1??? ?????
???0? ????? 00?1? ???0? ???0? ????? ????? ???1? ????? ?0??? ????? ?????
????? ????? ????? ????? ????? ?210? ?0010 01030 02000 1?00? 02?11 0???1
01002 01040 10111 23310 00123 00110 21410 03003 10153 010?1 12021 00111
?1101 12?00                    

Leptictis

31301 21010 21102 10002 00110 00003 11322 20022 01321 15001 21112 10100
00113 11010 02101 21111 40201 21221 01011 10010 10001 11100 11110 00000
01001 01011 10011 11101 21110 20011 00110 10010 11000 00?00 ?1001 00210
01110 00010 21011 113?1 11??? 02111 10021 1103? ?11?? ????? ?2020 11012
10012 01110 10111 33310 ?0221 ??11? ????? ??00? 10??? 110?2 ????? 1??20
??1?1 12?0?                    

Aphronorus

??201 11220 20111 22002 10110 01000 01001 30012 01101 23200 21122 10111
01203 01012 01010 50011 30101 ??01? 01011 ????? ??0?0 10?00 ????? ?????
????? ????? 0?01? ????? ???0? ?00?? ????? ??01? ????? ????? ????? ?????
????? ????? ?1??? ????? ????? ????? ???2? 0??0? ????? ????? ?0??? ?????
????? ????? ????? 2?3?? ????3 ??1?? ??4?? ????? ????? ????? ????? ?????
????? ?????                    

Coriphagus

???0? 11?10 20010 12?02 00011 00000 11100 40010 01111 23200 11130 20120
00?13 00011 02022 40??1 00001 1???? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Palaeosinopa

1?301 11110 10110 00000 00010 00010 11010 00021 01111 23201 11231 20210
00113 11011 02002 51111 40201 22221 01001 10010 ????? ???00 ?0??? 0???0
1??00 ???22 010?? ?1??? ????1 ?00?1 ?02?? 100?0 ????1 00011 1?0?? 0??1?
?1010 ????? ???0? ???01 1?1?0 01110 11020 00030 1?000 0?0?? 10?2? 00?11
100?0 01140 01011 03310 00121 00010 11310 030?2 10143 00012 10?21 20110
??10? 12000                    

Escavadodon

????1 11?1? 22202 00000 00010 00110 21222 30??1 01300 20201 31021 10110
00013 10011 01002 3???1 ?0??? ????? ????? ????? ????? ????? ????? ?????
????? ????? 0?0?? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ?1121 00030 02011 0000? 02?21 ????1
1??10 0?120 11?11 23?10 00221 00110 2?410 0301? ????? ??0?? ?2021 2012?
?1001 12?00                    

Miacis/Vulpavus

10201 11110 20010 00300 00011 10010 11010 00001 00111 11100 01111 20210
00002 10000 02001 21011 30101 11111 01001 11010 10012 00301 100?1 00000
11030 ?0110 00111 11002 21200 ?0000 11200 00000 ?1200 10100 0?000 00010
01000 2???0 2?0?? ????? ????? ?11?0 01111 01031 12011 1110? 01401 00??1
00001 11020 00001 03200 10221 00001 20300 12003 10130 01213 12021 10110
?1001 10?00                    

Viverravus

???01 11000 10010 00300 00011 11003 00010 20001 00131 16000 01110 20220
00013 00000 24201 20??1 ?0?01 ??00? ????? ??01? ??01? ???01 ?1??? ?????
???1? ????? 0?01? ????? ??100 ?00?? 11??? 0?012 ????? 00??? ????? 00??0
?1110 2?2?0 ????? ???01 1???? ?111? ?1011 11020 02000 1110? 0041? ??011
00011 11110 100?1 ????0 11122 00001 20411 130?? ????? ????? ????? ?????
???0? ?????                    

Protungulatum

???01 11?1? 20100 10001 00100 00110 10221 00101 01301 24201 11230 10110
01012 02011 22121 32211 30201 2???? ????? ????0 ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? 00010 010?? 0?201 ??001 00210
1???? ????? 2?0?1 023?? ????? ????? ????? ??010 01010 010?? ????? ?????
????? ????? ????? ????? 00121 00000 20311 03001 10140 00013 11?10 20120
????? ?????                    

Loxolophus

1??01 11110 20100 00001 00110 00000 10110 11111 01322 17101 21121 10010
02213 02011 12022 10021 30101 ?1020 ?1011 11??? ??0?1 0??10 01??? ?????
??01? ???20 0?11? ????? ???00 200?? ????? ???1? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ???10 0?000 1?000 0?0?? ????? ???10
000?? 1?14? ?0??? 0?10? 0?121 ??000 ????1 ??0?1 1014? 00111 ???20 ?00??
?300? ?0?00                    

Arctocyon

10201 11110 20100 01001 00000 00000 10010 00132 01321 17201 31230 20111
12213 02011 32121 30101 30201 10010 01011 10010 20010 10310 00??? 0??00
1?01? 0?020 0011? ????2 ?1200 2000? ?1??? ??010 1???? ?0??? ??0?? 00??0
?111? 1???? 2???? ???01 11100 ??111 01121 01030 11000 10000 12110 01?11
00010 11110 10001 02200 01121 01001 20411 13011 10133 00111 10020 10130
?1011 0?000                    

Pseudictops

81701 11100 21301 10046 00100 00202 30303 30031 023?2 06002 41122 00001
00113 00120 02102 1???1 ?0??? ??0?? ????? ????? ??0?? ??000 ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ???1?
????2 ????0 1?111 03200 00122 20010 20110 02002 10131 00030 11023 20140
??001 1??00                    

Gomphos

90814 51223 21310 21107 00010 00002 51002 30132 02410 17102 31221 00000
11212 11122 02113 42021 30200 21001 0000? ?1011 ?10?2 00301 401?0 10??0
??00? ?0?1? ??01? 11001 ?1?01 11100 0210? 003?? ????? ?0??? ????? ?????
?1??0 ?0210 200?? ????1 11??? ?0110 ?0??? ??0?? ??1?? ????? ???2? ???11
?0??2 01130 ??111 03210 02122 00110 2?411 03012 10123 11121 12023 20131
?0101 1?00?                    

Rhombomylus

90814 51224 21311 22147 00010 00111 51213 30032 013?0 07022 41131 00001
11103 10122 02115 42211 30200 21002 00001 11011 11002 00301 40110 10110
10030 10122 00011 10100 21101 21100 02200 10311 11210 10120 32?01 00200
01020 20211 20001 013?? ?1??1 ?11?0 10011 00011 11100 01010 1111? ???10
10012 01120 10111 03210 12122 00110 1?411 14012 10121 00111 12013 20121
?0101 1??00                    

Macrocranion

31301 21112 00000 21003 00110 01000 01101 01111 01111 23201 21200 01111
01213 02011 01011 42211 30201 12011 ???01 100?? ????? 0??00 41??? ?????
??00? ???1? 0?0?? ????? ???0? 201?0 ????? ?101? ????? ?0??? 1?0?? 00?1?
?0?11 ?00?0 ?0??? ????? ????? 01100 1011? 0?02? ????? ????? ??01? 0?01?
????2 01?4? 1??01 3?200 21121 20110 20011 03111 10141 00112 10013 10111
0?001 ?0100                    

Apheliscus

???0? ?1??0 100?0 22043 00010 01000 21200 21122 01321 16102 31121 01101
01214 02010 2202? 32011 51001 ??01? ????? ????? ????? ????0 ????? ?????
???0? ????? 0?0?? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ??011 10010 1?01? 02?2? ????0
10001 01140 10110 33300 01122 20110 2?411 04112 10143 00021 12023 2012?
????0 1????                    

Haplomylus

???0? ?1??? ????1 10031 00110 00220 10221 30121 01321 28202 31220 10111
02225 02010 22013 4???1 ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ?1100 11011 10110 100?? ????? ????0
10002 01040 1011? 31300 01222 20110 0?411 03112 10153 00021 12023 1012?
????0 1????                    

Paschatherium

??30? 21?00 21201 11012 00110 00110 20002 40121 01211 13101 21102 00011
01213 02010 02012 523?1 30111 2???? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? 01123 20110 21410 14111 10131 01001 12021 20111
????? ?????                    

Rhynchocyon

91700 41000 21212 02014 03010 00003 10322 30022 01320 19221 41110 10011
02225 02022 04212 42001 41211 21121 01101 01010 20000 01010 11000 01000
01020 11101 00010 11002 21000 21201 01111 ?1100 21011 00120 11?0? ??10?
3?100 00111 21111 12301 11110 01111 11111 11001 10100 20000 12021 00012
10002 01140 10110 33201 21223 20110 10410 01112 10153 00121 01033 10141
03001 11000                    

Soricidae

61602 51123 00010 20004 01010 01120 11000 11112 31120 19121 11120 20120
01334 02010 33115 51301 30111 22221 11011 1111? ?0020 10100 63?00 ?1???
?1002 ?0021 00011 11001 2?010 203?1 11200 01010 21200 00100 01000 00211
31010 00100 20100 01301 11120 ?2100 11010 00000 12000 00011 00320 21211
00112 01042 00001 30300 02221 10101 00410 00002 10114 20011 12010 00120
02020 10000                    

Echinosorex

31300 11112 20210 11044 01000 00000 10001 01132 01122 19102 21120 20021
00324 02010 23025 31201 40211 23112 11111 11110 00020 10001 51111 10111
00001 01010 01011 11001 21100 20001 01111 02010 11210 00000 11001 00010
?1100 20000 10011 02101 11110 12110 11110 11001 11100 20000 11020 11211
00012 01010 10111 33301 12123 21000 00410 01002 10114 10012 12011 00110
02120 10000                    

Erinaceus

41403 31223 20210 22104 00010 01000 00321 01132 001?2 09101 31220 20121
00324 02010 24223 42221 40110 23112 11011 10111 20100 11101 51110 10111
00000 00012 01011 11001 21000 20111 00111 01010 11011 00110 11001 00011
11010 20000 10011 0?201 11110 12110 11110 01111 01000 21010 12010 21211
10300 11131 11001 33401 22121 20000 00110 01012 10123 10020 10020 10110
02020 00000                    

Onychonycteris

2?201 31222 10010 2000? 0001? 01000 00000 01101 110?? ????1 01?21 ???10
00?23 12010 3300? ?20?? 30101 ????? ????? ????? ????? ????0 ?1??? ?????
????? ????? 0???? ????? ????? ????? ????? ????? ????? ?0??? ????? ?????
????? ????? ????? ????? ????? 02?1? ?010? ????? ?20?? ????? ????? ???1?
????? ??00? ?0??1 1???0 ????? ????? ????? ????0 10??? 1?0?2 ???2? ??1??
1???? ???11                    

Icaronycteris

21701 11222 10110 20003 00010 00000 00221 20101 11011 22200 01121 20110
01123 02010 33014 00201 20101 2?11? ????? ????? ????? ???00 ?1??? ?????
????? ????? ????? ???0? ???0? ?0??? ????? ???1? ????? ?1??? ????? ?????
????? ?00?? ????? ????? ????? 0??1? ?110? 0??0? ????? ????? ????? 2?21?
?1??0 ???0? ????1 0???0 ????? ????? ????? ????? ????? ????? ????? ?????
????? ???11                    

Rhinolophus

21701 11224 00010 22003 00000 01000 01000 00102 21031 12001 11210 20120
01332 02011 33014 00101 20101 1422? 1???? ?101? 200?2 00000 51??0 ?1???
?0001 ?102? 0111? ?1002 ??000 20000 0?2?0 00010 ?1000 01000 ?2101 00200
01110 101?0 2?011 01?01 11110 01010 01101 00100 02022 10000 10210 21211
01200 11002 10001 12200 02222 20110 20410 03000 10123 20033 11023 20110
11000 10211                    

Leptacodon

????? ?1010 10011 10001 00110 00010 11121 31011 01111 22200 11110 10110
00122 12010 11010 3010? 00001 ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Cryptotopos

6?602 31110 20211 20012 01110 01010 11222 31011 01111 23211 11112 10110
00111 12010 11015 31100 00101 2?11? ????? ?101? ????? ???00 ?1??? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ?0010 00021 12021 11101 1020? ????1
10101 11041 10??? 1320? 00122 21100 20301 12013 10120 10202 01021 10111
?3001 10100                    

Tupaia

51502 11221 20010 20003 00010 00100 00111 01112 31000 13101 10110 20010
01233 02012 02010 21001 40201 10010 01011 00000 10100 10101 02101 01000
01140 10012 01011 11103 11100 20001 11110 00112 21211 00101 11110 00200
00110 01010 21011 12301 11110 01110 00110 01011 11100 00000 11210 20010
00102 11140 00101 03000 22221 11101 00201 12012 10130 21313 01033 10141
03001 10100                    

Ptilocercus

51503 31221 20010 20003 00010 01000 01211 30122 01131 17011 11121 20110
01224 12010 32020 11000 40201 13100 01011 11110 10001 10001 22111 00100
11040 10010 11001 11002 21100 20011 01110 00112 11211 00101 11100 00100
00110 01010 21011 12301 11110 01110 00110 01031 12042 10101 11300 10111
00100 10121 10001 02000 20220 01111 10201 11002 10112 21303 01022 20140
03000 10100                    

Mixodectes

4?4?4 51222 ?0010 00001 00110 01000 01300 10121 21121 23201 21110 10010
02224 02011 01010 4???1 ?0?0? 2???? ????? ????? ????? ???00 ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ???1? ????? ???0? ?2022 1?100 0040? ?????
1?0?1 11110 1000? ????? 20?20 0?000 20??? ??0?2 11122 11203 10131 02140
?0?0? ????0                    

Eudaemonema

??4?? 21?12 ?0111 00021 10110 01000 11212 30111 21221 23202 21111 10111
01222 12012 21012 4???? ????? ????? ????? ????? ????? ????0 ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Elpidophorus

80??2 ?1110 22201 10018 10110 01110 51322 31101 21112 26201 11121 10101
01233 12010 31010 4?3?1 ?0?0? 1?0?? ????? ????? ????? ????0 ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Plagiomene

??702 31110 22302 10128 10110 00222 41323 51111 30112 16201 21221 20200
02323 12010 31011 22311 ?0101 2?01? ????? ????? ????? ????1 ????? ?????
????? ????? ???1? ??1?3 ???0? ????1 ???11 ??11? ??101 ?0??? ????? 0????
?0?10 ?02?0 ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Cynocephalus

81703 41224 22002 02018 10010 00122 40123 50112 31020 10102 11100 21011
02234 12012 02010 50301 50201 10000 01001 11010 20041 11000 01011 00000
21130 00110 11111 11102 22000 00400 01200 10310 21110 00200 11011 00?20
01111 10211 21011 01301 11110 02110 00101 01031 10122 00000 12010 21111
01110 11112 00001 02100 21120 02111 10110 01003 01122 21212 00133 12101
03011 10200                    

Purgatorius

705?? ?1?10 ????1 10003 00011 00100 21300 41021 01111 22201 11220 10110
01122 02010 11111 3???? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ????? ?????
????? ?????                    

Micromomyidae

704?3 31221 20201 00001 10110 01100 00300 40122 01221 24201 21221 20110
01103 02010 10010 41111 00101 2?00? ????? ????? ????? ????0 ????? ?????
???0? ????? ????? ????? ????? ????? ????? ??01? ????? ????? ????? ?0???
????? ????1 ????? ????? ????? ????? ??010 00?3? ????2 ????? ????0 00111
00111 10110 10001 1?200 ????? ????? ????? ????? ????? ????? ????? ?????
????? ?0100                    

Ignacius

704?2 51223 11011 02004 01010 01200 10300 40112 01221 28121 21020 20100
12123 02012 00023 51001 40101 12000 10001 1001? 00000 10011 50?1? ?111?
?100? ???22 00011 ?0000 ?1000 2001? ????? ??11? ????? ?0??? ????? ?1???
?0110 000?1 ?1??? 0???1 ????? ?1??? ?0000 00030 12022 10101 0040? 00011
00011 10110 10001 03200 21221 01?00 00001 120?3 10132 11203 12?32 11121
0???? ?0200                    

Plesiadapis

70404 51223 20202 01013 00110 00200 50300 40122 21221 28201 21231 20010
11015 02010 10120 51021 30101 23202 01001 01110 10001 11301 0010? 00110
1?000 ?0122 0001? 101?0 11010 20001 01100 02010 11210 001?? 32111 11200
31110 01001 200?? ??311 1???1 ?11?? ?0110 00030 12032 1110? 01400 00011
10011 10120 00001 03200 21221 01000 20201 12003 10130 10303 00032 10142
???01 0?200                    

Cantius/Notharctus

30301 11110 20210 02023 00110 00000 10212 10122 01221 28202 21220 20010
11105 02011 10121 22021 00201 22011 11001 11111 10001 01300 401?0 00??0
??040 ?0?22 0000? 10000 ?1010 ?0000 01100 02112 11211 00101 ?1?11 11200
31110 01001 200?? ?2311 111?? 01??0 ?0110 01031 02042 11101 00100 00010
00010 10111 00011 01200 21221 01100 20201 10003 10122 21321 00032 21142
?1011 10200                    
 

APPENDIX 3
Character states at numbered nodes

The following numbered characters and their states relate to the numbered nodes in Figure 18. Bold type indicates unequivocal synapomorphy; plain type indicates homoplasy. Square brackets enclose character states found only by ACCTRAN optimization; parentheses enclose character states found only by DELTRAN optimization.

Node 1

7.1, 11.2, 13.1, 46.2, 52.1, 53.1, 65.1, 66.1, 78.1, 79.1, 80.1, 86.1, 115.1, 134.1, 141.1, 142.1, 146.1, 153.2, 169.0, 175.1, 178.2, 179.1, 194.1, 195.1, 198.3, 200.1, 203.1, 204.1, 212.1, 219.2, 227.1, 240.1, 264.1, 284.1 [5.1, 6.1, 36.1, 44.1, 54.2, 75.1, 77.1, 96.1, 97.1, 99.1, 104.1, 106.1, 108.2, 122.1, 124.0, 126.1, 129.2, 140.0, 142.2, 148.0, 149.1, 162.1, 163.0, 180.1, 189.1, 197.2, 199.1, 217.1, 242.1, 244.1, 245.1, 248.1, 251.1, 263.2, 276.1, 298.1] (108.3, 124.1, 263.1)

Node 2

28.1, 31.2, 44.3, 48.0, 55.2, 111.1, 148.1, 150.2, 151.1, 158.2, 185.1, 271.0 [170.1, 181.1, 243.1, 250.1] (54.2, 97.1, 99.1, 106.1, 108.2, 124.0, 126.1, 129.2, 140.0, 149.1, 162.1, 163.0, 180.1, 199.1, 263.2)

Ukhaatherium

29.1, 34.2, 78.2, 288.2 [36.0, 77.0, 298.0] (5.1, 6.1, 96.1, 189.1, 217.1, 242.1, 243.1, 250.1, 251.1, 276.1)

Asioryctes

43.0, 81.1, 184.1, 279.1, 290.2, 299.2 [5.0, 6.0, 96.0] (75.1, 77.1, 142.2, 170.1, 181.1)

Node 3

18.0, 20.1, 24.1, 35.1, 40.1, 42.1, 48.2, 65.2, 67.1, 221.1, 246.1, 249.1, 265.1, 277.1, 281.1, 283.1, 284.2, 287.1, 295.1, [1.1, 3.2, 8.1, 9.1, 16.1, 81.3, 83.1, 85.1, 113.1, 121.1, 130.0, 146.2, 161.1, 191.2, 208.1, 209.1, 233.0, 255.1, 258.2, 275.1, 280.2, 305.1] (6.1, 36.1, 245.1, 280.1, 298.1)

Node 5

39.1, 45.1, 47.1, 247.1, 264.2, 271.2, 273.2, 286.0, 289.1, 294.2 [47.2, 63.1, 213.1, 215.1, 225.0, 263.1, 276.0, 284.4] (1.1, 5.1, 8.1, 83.1, 85.1, 104.1, 113.1, 122.1, 130.0, 148.0, 197.2, 255.1, 258.2, 284.3, 305.1)

Node 6

44.2, 73.1, 139.1, 168.0, 198.2, 204.2, 216.1, 223.1, 224.0, 231.0 [1.4, 2.1, 7.0, 13.0, 16.0, 23.1, 31.2, 80.0, 97.0, 99.0, 101.0, 116.1, 126.2, 162.0, 181.2, 205.1, 230.1, 236.1, 240.2, 244.2, 248.0, 256.3, 258.4, 263.0, 265.3, 268.1, 269.1, 272.1, 275.0, 285.3, 304.2] (54.2, 63.1, 75.1, 106.1, 108.2, 140.0, 149.1, 199.1, 213.1, 215.1, 217.1, 225.0)

Maelestes

33.0, 34.0, 35.0, 43.0, 46.1, 48.1, 53.0, 55.2, 78.2, 83.2, 103,1, 105.2, 111.3, 138.1, 144.1, 161.3, 178.0, 184.2, 185.1, 196.1, 200.0, 221.0 [81.0, 121.0, 163.1, 189.0, 191.0, 208.0, 209.0] (3.2, 7.0, 9.1, 13.0, 23.1, 31.2, 47.2, 77.1, 80.0, 116.1, 124.0, 129.2, 180.1, 181.2, 230.1, 233.0)

Node 7

19.3, 24.0, 34.2, 66.0, 70.1, 75.2, 78.0, 79.0, 219.0, 220.1, 227.0 [3.0, 9.0, 47.1, 77.0, 104.2, 116.2, 119.2, 129.1, 148.1, 150.2, 151.1, 158.2, 203.0, 212.0, 234.2] (1.4, 163.0, 240.2, 242.1, 244.2, 284.4, 285.3)

Zhelestidae

11.1, 16.1, 39.2, 50.1, 51.1, 52.0, 62.1, 64.2, 71.1, 72.2, 76.4, 81.1, 83.0, 85.0, 112.0, 175.0, 179.0, 222.0, 241.1, 243.1, 249.0, 283.0, 296.2 [161.0, 180.0, 181.0, 280.1] (7.0, 13.0, 15.1, 23.1, 31.2, 80.0, 96.1, 124.0, 251.1)

Zalambdalestidae

3.1, 5.0, 6.0, 20.0, 32.1, 35.2, 36.0, 38.1, 45.0, 47.0, 54.3, 59.0, 65.4, 76.2, 81.4, 82.1, 108.1, 122.0, 124.2, 162.1, 198.1, 229.2, 245.2, 246.0, 284.5, 289.3, 290.0 [7.1, 13.1, 23.0, 31.1, 80.1, 96.0, 251.0] (2.1, 101.0, 104.2, 116.2, 119.1, 121.1, 126.2, 142.2, 148.1, 150.2, 151.1, 158.2, 161.1, 180.1, 181.2, 189.1, 191.2, 203.0, 205.1, 208.1, 209.1, 212.0, 234.2, 236.1, 256.3, 258.4, 263.0, 265.3, 268.1, 269.1, 272.1, 280.2, 304.2)

Node 8 - Placentalia

51.1, 92.1, 95.1, 114.1, 135.1, 136.1, 137.1, 141.2, 152.1, 156.0, 172.1, 183.1, 184.1, 202.1, 207.1, 239.1, 257.3, 273.3, 299.1, 302.1 [50.1, 55.0, 62.1, 67.2, 71.1, 74.1, 86.2, 106.0, 108.3, 124.1, 126.0, 140.2, 142.1, 149.0, 180.0, 182.1, 188.0, 192.1, 199.0, 242.0, 249.2, 306.1] (3.2, 9.1, 16.1, 47.2, 81.3, 96.1, 99.1, 146.2, 162.1, 191.2, 208.1, 209.1, 251.1, 275.1)

Node 9

14.1, 32.1, 128.1, 163.2, 222.2, 228.1, 233.2, 276.1, 280.3, 288.2 [54.1, 75.1, 129.1, 213.0, 220.1, 225.1, 234.0, 261.1, 277.2, 284.3] (97.1, 121.1, 306.1)

Node 10

20.0, 25.1, 33.0 [34.0, 44.2, 50.0, 55.1, 62.0, 63.0, 67.1, 71.0, 74.0, 77.0, 78.0, 90.1, 105.2, 110.1, 111.1, 113.0, 143.1, 151.1, 171.0, 175.0, 178.0, 186.2, 188.2, 231.0, 233.4, 235.1, 270.1, 292.2]

Node 11

26.1, 13.0, 18.2, 35.0, 39.0, 42.0, 51.0, 58.2, 69.0, 72.2, 75.1 [9.0, 47.1, 64.0] (78.0)

Cimolestes magnus

14.0, 31.2, 54.2, 75.2, 81.1 (9.0, 34.0, 44.2, 47.1, 64.0, 86.2)

Node 12 - Carnivora

16.0, 18.3, 46.1, 48.1, 56.2, 67.0, 76.2 [34.1, 44.1, 86.1] (110.1, 151.1, 182.1, 186.2, 215.1, 217.1, 231.0, 233.4, 261.1, 270.1)

Miacis/Vulpavus

29.1, 36.0, 77.1, 88.1, 112.0, 124.3, 133.1, 143.2, 159.0, 166.1, 183.0, 184.0, 213.1, 219.3, 232.1, 251.0, 263.2, 275.0 [9.1, 244.0, 248.0] (47.1, 64.0, 90.1, 105.2, 111.1, 113.0, 140.2, 171.0, 175.0, 178.0, 220.1, 234.0, 235.1, 249.2, 277.2, 292.2)

Viverravus

8.0, 11.1, 27.1, 30.3, 31.0, 32.0, 36.2, 44.3, 47.6, 48.0, 55.0, 59.2, 65.3, 66.0, 71.2, 72.4, 73.2, 89.0, 160.2, 216.1, 221.0, 224.0, 225.0, 262.1, 265.2, 273.4, 274.1, 277.3 [64.1, 220.0, 234.1, 249.1] (9.0, 143.1, 188.2, 244.1, 248.1)

Batodon

27.1, 76.4 (34.0, 44.2)

Node 13 - Euarchonta

1.4, 5.2, 15.1, 33.2, 64.2, 212.0, 224.2, 267.1, 284.2, 286.1, 300.1 [3.4, 6.3, 36.3, 79.0, 104.0, 106.0, 108.0, 140.3, 150.1, 153.1, 158.1, 164.1, 165.1, 170.1, 173.1, 174.1, 179.0, 182.0, 215.0, 217.0, 238.1, 243.1, 250.1, 268.1, 289.0, 291.0, 302.3] (6.2, 36.4, 50.1, 63.1, 67.2, 71.1, 74.1)

Node 14

1.6, 37.1 [80.0, 81.0, 88.1, 230.1, 241.1. 244.0, 248.0, 249.4, 256.1, 261.0, 265.2, 266.2, 279.1, 287.0, 290.2, 292.1]

Node 15 - Nyctitheriidae

23.1, 29.1, 34.2 [3.6, 62.0] (36.3, 79.0, 81.0)

Leptacodon

8.0, 11.1, 13.0, 33.1, 50.0, 83.0 [77.0] (55.0)

Cryptotopos

13.2, 16.2, 19.1, 20.2, 22.1, 27.1, 35.2, 47.3, 49.1, 55.2, 64.1, 65.1, 75.5 (3.6, 6.3, 77.1, 80.0, 86.2, 88.1, 220.1, 230.1, 234.0, 241.1, 243.1, 249.4, 250.1, 256.1, 265.2, 266.2, 268.1, 277.2, 279.1, 287.0, 289.0, 290.2, 291.0, 292.1, 302.3)

Purgatorius

1.7, 3.5, 20.3, 25.1, 28.1, 31.2, 33.3, 34.0, 35.0, 39.2, 53.2, 54.2, 66.0, 73.1, 75.1 [36.4] (55.0, 62.1)

Node 16

8.2, 10.1, 38.1, 41.2, 63.2, 225.2, 261.2, 273.2, 285.2, 294.3, 299.4 [9.2, 47.3, 65.3, 70.1, 76.4, 89.0, 219.3, 227.0, 249.1, 262.1, 263.2, 265.0, 297.2] (62.1, 77.1, 106.1, 161.1, 164.1, 174.1, 179.0, 248.1)

Node 17

46.1, 56.2, 87.0, 124.3, 132.1, 213.1, 217.1, 232.1, 258.1, 270.1, 271.1, 286.2, 295.3 [2.1, 3.2, 13.0, 28.1, 40.2, 41.3, 48.1, 64.3, 72.2, 83.2, 86.1, 123.1, 131.1, 138.1, 154.1, 161.2, 222.1, 223.1, 226.0, 228.0, 234.1, 236.2, 245.0, 251.0, 257.2, 269.1, 277.1, 289.1] (65.1, 108.0, 140.2, 189.1, 192.1, 219.3, 220.1, 227.0, 236.1, 243.1, 268.1, 291.0, 302.3)

Node 18 - Scandentia

1.5, 3.5, 15.0, 16.2, 20.3, 31.0, 76.2, 78.0, 81.4, 94.1, 110.1, 112.2, 124.4, 126.1, 128.0, 143.1, 160.2, 175.0, 187.1, 196.1, 249.2, 258.0, 280.2, 288.3, 292.1 [44.0, 244.0, 297.0] (2.1, 9.2, 13.0, 40.2, 47.3, 48.1, 72.2, 79.0, 83.2, 104.0, 150.1, 153.1, 154.1, 158.1, 165.1, 170.1, 173.1, 188.0, 263.2)

Tupaia

6.1, 32.0, 33.1, 36.0, 37.1, 43.0, 45.0, 52.0, 58.0, 66.0, 70.2, 71.0, 96.0, 97.0, 99.0, 103.1, 105.0, 108.1, 114.0, 117.1, 121.0, 130.2, 141.1, 151.1, 219.1, 224.0, 225.0, 240.0, 245.2, 249.4, 253.1, 262.2, 266.1, 271.0, 277.2, 279.1, 284.3, 285.0 [89.1, 131.0, 238.0, 250.0, 257.3, 265.1, 269.0] (28.1, 41.3, 44.0, 64.3, 123.1, 138.1, 140.3, 161.2, 222.1, 223.1, 226.0, 228.0, 236.2, 251.0)

Ptilocercus

5.3, 27.1, 39.2, 41.0, 44.3, 47.7, 48.0, 49.1, 54.2, 65.4, 71.3, 74.2, 76.1, 80.0, 87.3, 88.1, 98.1, 111.2, 118.1, 134.0, 149.1, 174.0, 178.1, 224.4, 230.1, 233.3, 234.0, 247.0, 284.1, 289.0, 294.2, 295.2, 296.2, 300.0, 305.0 [28.0, 55.1, 64.2, 70.0, 123.0, 138.0, 140.2, 161.1, 222.2, 223.0, 226.1, 228.1, 236.1, 251.1, 262.0] (6.3, 36.3, 89.0, 131.1, 238.1, 245.0, 250.1, 257.2, 265.0, 269.1)

Node 19 - Dermoptera

1.8, 3.7, 12.2, 14.0, 19.1, 20.8, 21.1, 34.2, 78.3, 163.1, 188.2 [15.2, 29.2, 30.2, 31.4, 35.3, 36.5, 47.2, 60.1, 62.2, 81.5, 101.2, 104.4, 107.1, 113.0, 121.2, 133.1, 142.2, 146.0, 148.4, 153.2, 156.1, 168.2, 170.0, 173.0, 179.2, 186.1, 197.1, 207.2, 214.0, 215.1, 222.0, 232.2, 233.0, 237.1, 242.1, 250.2, 263.1, 267.2, 273.1, 274.1, 275.0, 276.0, 281.0, 282.1, 290.2, 293.1, 299.0, 304.1, 308.2] (28.1, 29.1, 31.5, 35.2, 36.3)

Node 20 - Plagiomenidae

8.1, 10.0, 13.2, 23.1, 33.3, 37.1, 45.2, 47.6, 54.2, 59.0, 71.3 [2.0, 9.1, 40.1, 48.2, 55.1, 70.0, 72.1, 77.2, 79.1, 83.1, 89.1, 155.1, 164.0]

Elpidophorus

27.1, 39.0, 46.2, 56.1, [15.1, 29.1, 30.0, 31.5, 35.2, 36.3, 41.2, 62.1] (60.1, 64.3, 76.4)

Plagiomene

13.3, 18.1, 19.2, 28.2, 42.0, 51.2, 53.2, 58.2, 63.3, 75.1, 76.2, 86.2, 110.1 [60.0, 64.2] (6.3, 15.2, 29.2, 30.2, 31.4, 35.3, 36.5, 41.3, 62.2, 77.2, 140.3, 150.1, 154.1, 155.1, 158.1, 164.0, 165.1)

Cynocephalus

5.3, 6.4, 10.4, 16.0, 17.2, 32.0, 33.1, 43.0, 44.2, 45.0, 47.0, 50.2, 54.0, 57.1, 58.0, 65.4, 70.2, 71.0, 76.5, 77.0, 158.3, 182.1, 185.1, 190.1 [140.2, 150.0, 154.0, 165.0] (2.1, 9.2, 13.0, 15.2, 29.2, 30.2, 31.4, 35.3, 36.5, 40.2, 41.3, 48.1, 55.0, 60.1, 62.2, 64.3, 72.2, 79.0, 81.5, 83.2, 89.0, 101.2, 104.4, 107.1, 113.0, 121.2, 123.1, 131.1, 133.1, 142.2, 146.0, 148.4, 156.1, 161.2, 168.2, 179.2, 186.1, 197.1, 207.2, 214.0, 215.1, 222.0, 223.1, 226.0, 228.0, 232.2, 233.0, 236.2, 237.1, 238.1, 242.1, 244.1, 245.0, 250.2, 251.0, 257.2, 262.1, 265.0, 267.2, 269.1, 273.1, 274.1, 275.0, 276.0, 281.0, 282.1, 290.2, 293.1, 297.2, 299.0, 304.1, 308.2)

Node 21

10.2, 16.0, 23.1, 27.1, 44.2, 51.2 [6.2, 43.2, 79.1, 87.2, 91.1, 111.4, 112.0, 115.0, 118.1, 119.1, 124.0, 129.2, 130.2, 137.0, 140.0, 141.1, 157.2, 171.3, 176.1, 181.3, 189.0, 190.1, 199.1, 205.1, 220.0, 231.0, 233.4, 243.0, 250.0, 268.0, 291.1, 302.1] (3.4, 70.1, 76.4)

Node 22

6.5, 33.3, 34.0, 35.0, 39.2, 66.0 [31.0, 36.4] (9.2, 65.3, 86.2, 231.0, 233.4, 234.0, 289.0)

Mixodectes

5.4, 13.0, 15.0, 36.1, 58.0, 62.2, 65.4, 71.0, 219.0, 241.1, 280.2, 282.1, 293.1, 296.0, 300.0, 302.0 [43.1, 262.0] (31.0, 47.3, 55.0, 265.0, 297.2)

Node 23

1.7, 28.1, 32.0, 40.2, 41.0, 47.4, 54.2, 56.2, 63.1, 72.0, 247.0 [13.2, 53.2, 64.1, 230.1, 265.1, 284.3, 295.2, 297.1] (43.2, 89.0, 124.0, 190.1, 219.3, 244.1)

Micromomyidae

5.3, 6.3, 10.1, 14.0, 21.1, 64.0, 70.0, 81.0, 243.1, 256.1 [55.1, 158.0] (13.2, 31.0, 53.2, 238.1)

Node 24

20.3, 47.8, 61.1, 74.2, 78.0, 177.1, 308.2 [10.3, 17.2, 28.2, 31.1, 76.5, 110.1, 238.0] (17.1, 87.2, 104.0, 112.0, 118.1, 119.1, 129.2, 130.2, 137.0, 140.0, 188.0, 230.1, 262.1, 263.2, 295.2)

Ignacius

11.1, 12.1, 13.0, 20.4, 22.1, 23.0, 39.1, 48.1, 49.2, 53.0, 59.0, 62.2, 70.2, 71.0, 75.3, 79.0, 81.4, 86.1, 92.0, 97.0, 101.0, 105.0, 109.1, 111.5, 117.1, 149.1, 214.0, 271.0, 273.0, 292.2, 299.2 [64.2] (10.3, 17.2, 28.2, 55.0, 76.5, 91.1, 108.0, 110.1, 158.1, 192.1, 227.0, 277.2, 284.3, 297.1)

Node 25

19.1, 27.0, 58.0, 65.5, 73.1, 79.2, 90.1, 98.1, 107.1, 144.1, 182.1, 187.1, 213.1, 224.3, 251.0, 288.3, 291.0, 300.2 [108.3, 114.0, 192.0, 227.1] (13.2, 53.2, 64.1, 153.1, 157.2, 176.1, 181.3, 199.1)

Plesiadapis

5.4, 14.0, 15.2, 31.5, 41.2, 54.3, 63.0, 70.0, 87.3, 88.2, 90.2, 96.0, 138.1, 172.2, 232.1, 241.1, 249.2, 285.0, 287.0, 306.0 [17.1, 55.1, 91.0, 111.0, 158.0, 165.0, 297.0] (10.3, 28.2, 76.5, 110.1, 114.0, 141.1, 150.1, 171.3, 173.1, 205.1, 277.2, 284.3)

Cantius/Notharctus

1.3, 3.3, 5.1, 6.1, 8.1, 9.1, 10.0, 15.0, 19.2, 28.0, 33.2, 34.1, 35.2, 36.1, 50.2, 64.0, 75.1, 76.2, 77.2, 81.0, 83.2, 89.1, 100.1, 106.0, 124.4, 134.0, 160.2, 217.1, 220.1, 221.0, 224.4, 233.1, 240.0, 245.0, 250.1, 254.1, 257.1, 268.1, 277.0, 286.2, 289.2, 290.1, 296.2, 304.1 [110.0, 150.0, 284.2] (17.2, 55.0, 91.1, 111.4, 115.0, 158.1, 165.1, 170.1, 297.1)

Eudaemonema

19.2, 21.1, 35.2, 50.2, 60.1, 70.2, 71.2, 75.2 [9.1, 55.1, 65.2] (36.3, 43.2, 47.3)

Node 26

38.1, 66.0, 70.1, 72.2, 274.1, 277.3, 287.0 [36.0, 43.3, 47.4, 74.2, 94.1, 101.2, 115.0, 127.1, 154.1, 219.0, 226.0, 237.1, 245.0, 280.1, 295.0, 308.0] (50.1, 54.2, 62.1, 67.2, 74.1, 75.1, 163.0, 225.0, 284.4)

Node 27

63.2, 65.3, 224.0, 227.0, 273.4, 279.1, 285.3, 290.2 [35.0, 47.5, 51.2, 86.1, 197.1, 262.1] (47.3, 161.1, 182.1, 213.1, 244.1)

Node 29

2.1, 76.4, 90.1, 108.0, 124.0, 130.1, 211.1, 246.0, 256.2, 268.1, 269.1 [14.1, 32.1, 34.0, 43.1, 47.3, 71.0, 74.1, 87.2, 121.0, 132.1, 140.1, 217.0, 222.2, 275.0, 295.1] (115.0, 197.1, 233.0, 249.2, 306.1)

Node 36

67.1, 214.2, 258.3 [36.4, 38.0, 55.1, 62.0, 75.2, 101.1, 104.0, 111.1, 143.1, 150.1, 152.0, 156.1, 169.1, 196.1, 234.2, 235.1, 237.0, 241.1, 252.1, 254.1, 259.1, 262.0, 272.1, 280.2, 296.2, 307.2] (32.1, 192.1)

Node 38

3.3, 29.1, 63.1, 66.1, 74.0, 86.2, 89.2, 219.3 [16.0, 20.0, 34.1, 88.2, 94.0, 107.1] (62.0, 75.2, 77.1, 150.1, 156.1, 234.2, 241.1, 248.1, 254.1, 259.1, 275.0, 280.2, 296.2, 307.2)

Node 39

12.1, 13.2, 15.2, 31.2, 34.2, 35.2, 36.3, 43.2, 232.2, 292.2 [1.3, 8.0, 14.0, 28.1, 33.2, 38.1, 47.2, 72.1, 87.1, 129.1, 132.0, 153.1, 210.1, 221.0, 231.0, 253.1, 287.1, 289.2, 290.1, 300.1] (51.2, 215.1, 235.1)

Node 40

20.2, 23.1, 59.0, 70.0, 212.0, 217.1, 245.2 [6.2, 16.1, 44.2, 73.1, 214.1, 216.1, 222.1, 223.1, 226.1, 252.0, 256.3, 265.2] (1.3, 104.0, 153.1, 189.1, 253.1, 287.1)

Node 42

9.0, 266.1, 278.1 [5.0, 19.2, 33.1, 88.0, 124.2, 126.1, 128.1, 147.1, 148.1, 152.1, 164.1, 169.2, 179.0, 184.0, 188.1, 190.1, 220.1, 221.1, 229.1, 231.1, 240.0, 256.1, 258.2, 262.1, 266.1, 272.0, 284.3, 285.1, 307.0] (8.0, 19.1)

Node 43

31.3 [6.1, 77.0, 81.1, 267.1, 269.2, 270.3, 271.0, 273.1, 277.0, 280.0, 281.0, 282.4, 304.1] (19.2, 44.2, 47.2, 72.1, 73.1)

Gypsonictops

7.0, 16.0, 31.4, 35.3, 51.1, 55.2, 71.1 [28.0] (77.0, 81.1)

Fordonia

21.1, 36.4, 38.0, 58.0, 63.0, 66.0, 67.0 (28.1, 267.1, 269.2, 270.3, 271.0, 273.1, 277.0, 280.0, 281.0, 282.4, 304.1)

Node 44

34.1, 62.1, 74.1, 77.2, [44.1, 47.3, 72.2, 73.0, 76.5, 78.0, 266.2, 277.4] (6.2, 97.1, 214.1, 290.1, 300.1)

Leptictidium

2.0, 12.0, 27.1, 31.0, 41.2, 49.1, 55.0, 56.2, 63.0, 64.2, 72.3, 75.4, 79.2, 81.4, 88.1, 133.1, 248.0, 251.0, 257.1, 294.1, 295.0, 306.0 [28.0, 308.1] (5.0, 19.2, 76.5, 78.0, 124.2, 216.1, 256.1, 258.2, 289.2, 307.0)

Node 45

15.1, 17.1, 34.0, 39.2, 46.1,48.1, 56.0, 60.1, 66.0 [1.9, 19.1, 32.0, 58.0, 63.2, 67.2, 89.1, 219.1, 259.0, 307.1] (262.1, 265.2, 266.2, 277.4)

Node 46

29.0, 33.2, 43.3, 50.2, 51.3, 287.0, 295.3, 299.2 [3.7, 14.1, 19.4, 23.0, 40.2, 47.6, 76.4, 83.2, 275.1] (1.9, 3.8, 19.0, 47.7, 126.1, 128.1, 147.1, 164.1, 169.2, 179.0, 219.1, 220.1, 289.2, 308.0)

Node 47 - Anagalida

8.1, 13.3, 20.6, 39.3, 68.1, 69.2, 73.1, 256.0, 278.0 [5.1, 30.2, 35.3, 38.0, 42.2, 46.0, 48.0, 51.4, 63.1, 67.0, 85.0, 89.0, 92.0, 100.1, 102.1, 105.2, 107.0, 111.4, 112.0, 116.1, 118.1, 119.1, 121.1, 122.0, 127.0, 130.2, 145.1, 150.0, 152.2, 154.0, 158.3, 160.1, 163.2, 166.1, 171.3, 172.2, 183.0, 164.2, 186.2, 188.2, 192.0, 194.0, 196.0, 205.1, 210.0, 213.0, 216.0, 217.0, 226.0, 227.1, 232.1, 233.1, 302.0] (30.1, 58.0, 258.2, 284.3)

Pseudictops

1.8, 6.1, 17.0, 24.0, 28.2, 31.3, 33.3, 45.2, 55.2, 62.0, 74.0, 76.1, 262.0, 268.0, 273.1, 277.2, 279.0, 289.3, 290.0, 292.1, 299.4, 300.0 [14.0, 23.1, 40.1, 275.0] (3.7, 19.4, 30.2, 32.0, 35.3, 38.0, 42.2, 46.0, 47.6, 48.0, 51.4, 67.0, 259.0, 285.1)

Node 48 - Glires

2.0, 4.1, 5.4, 6.5, 8.2, 9.2, 10.3, 16.2, 18.1, 20.7, 31.5, 45.0, 61.1, 66.1, 70.2, 75.5, 108.3, 110.1, 262.2, 266.0, 284.2, 288.1, 303.1 [3.8, 32.1, 47.7, 259.1] (14.1, 23.0, 40.2, 83.2, 85.0, 89.0, 92.0, 100.1, 102.1, 105.2, 111.4, 112.0, 116.1, 145.1, 148.1, 150.0, 152.2, 158.3, 188.2, 192.0, 223.1, 275.1, 302.0)

Gomphos

15.0, 33.0, 43.4, 53.2, 60.0, 63.2, 65.2, 75.3, 79.2, 146.1, 156.0, 207.0, 249.3, 277.3, 286.1, 287.1 299.3 [19.0, 28.0, 35.2, 38.1, 46.1, 48.1, 51.3, 67.1, 124.0, 190.0, 240.1, 285.3] (30.2, 42.2, 78.0)

Rhombomylus

10.4, 17.2, 29.1, 34.1, 49.2, 54.3, 64.0, 78.2, 90.2, 124.3, 129.2, 137.0, 138.1, 140.0, 153.2, 234.1, 261.1, 271.1, 276.1, 289.1, 294.1 [30.1, 42.1] (19.4, 28.1, 35.3, 38.0, 46.0, 48.0, 51.4, 67.0, 118.1, 119.1, 121.1, 122.0, 130.2, 143.1, 160.1, 163.2, 166.1, 171.3, 172.2, 183.0, 184.2, 186.2, 190.1, 194.0, 205.1, 213.0, 217.0, 226.0, 227.1, 229.1, 232.1, 233.1, 240.0, 285.1)

Node 49 - Macroscelidea

44.2, 65.4, 82.1, 222.0, 244.0, 249.4, 255.0, 256.3 [6.4, 17.2, 20.3, 28.0, 35.1, 58.1, 71.2, 93.1, 96.0, 101.2, 105.0, 109.1, 113.0, 114.0, 117.1, 129.0, 135.0, 140.2, 143.0, 148.2, 155.1, 157.1, 158.1, 159.0, 161.2, 165.1, 178.1, 181.3, 193.1, 197.2, 212.1, 224.1, 258.3, 284.4, 285.3, 302.3, 305.0] (20.1, 63.2, 67.2, 78.0, 259.0)

Apheliscus

11.1, 12.0, 13.0, 15.0, 16.2, 27.1, 35.0, 36.2, 37.1, 57.1, 74.2, 76.3, 81.5, 83.0, 231.0, 245.1 [32.1, 124.0, 223.0] (17.2, 19.4, 20.3, 23.0, 40.2, 47.6, 71.2, 89.1, 224.1, 229.1, 240.0, 275.1, 305.0)

Node 50

31.1, 34.2, 48.2, 55.0, 56.1, 59.1, 62.2, 64.2, 65.5, 263.2, 271.1, 277.3, 284.5, 296.1 [19.0, 79.0, 81.4, 84.1, 88.1, 89.2, 229.0] (32.0)

Haplomylus

17.0, 19.3, 28.2, 29.2, 46.2, 47.8, 53.2, 75.3, 214.0, 215.0, 248.0, 257.1, 271.0 [20.1, 23.1, 40.1] (35.1, 71.2, 224.1, 240.0, 275.1, 305.0)

Rhynchocyon

15.2, 16.0, 19.1, 20.4, 22.3, 30.3, 33.3, 38.0, 45.0, 47.9, 49.2, 50.1, 51.4, 54.1, 58.0, 69.2, 70.2, 72.4, 73.2, 219.0, 226.2, 240.2, 258.2, 260.1, 261.2, 265.3, 277.1, 288.1, 291.0, 292.1, 294.3, 299.4 [35.2, 71.0, 224.0, 275.0, 305.1] (3.7, 5.0, 6.4, 14.1, 17.2, 23.0, 40.2, 79.0, 81.4, 83.2, 84.1, 88.1, 93.1, 96.0, 101.2, 105.0, 107.1, 109.1, 111.1, 113.0, 114.0, 117.1, 124.2, 127.1, 129.0, 135.0, 140.2, 148.2, 154.1, 155.1, 157.1, 158.1, 159.0, 161.2, 165.1, 178.1, 181.3, 184.0, 190.1, 193.1, 196.1, 197.2, 210.1, 302.3, 307.1)

Paschatherium

33.0, 36.4, 54.0, 55.2, 58.0, 59.1, 78.3, 84.1, 265.3, 272.1, 276.1, 280.1, 289.0 (28.1, 32.0, 63.2, 67.2, 76.5, 284.3, 285.1)

Node 41 - Leptictidae

30.3, 46.1, 55.2, 207.2, 213.0, 279.0, 296.1, 303.1 [94.1, 108.1, 125.1, 138.1, 149.1, 169.0, 237.1, 280.3, 284.5] (6.2, 33.2, 44.2, 73.1, 88.2)

Prodiacodon/Palaeictops

12.2, 13.3, 16.2, 21.1, 37.1, 39.0, 43.1, 51.1, 62.1, 81.0, 90.0, 209.0, 215.0, 222.2, 234.1, 241.0, 242.1, 244.0, 248.0, 249.4, 265.3, 296.0 [8.1, 216.0, 223.0, 256.2] (28.1, 47.2, 72.1, 97.1, 214.1, 221.0, 231.0, 272.1, 289.3, 284.5, 290.1, 300.1)

Leptictis

13.1, 29.0, 31.1, 33.3, 36.2, 38.0, 39.2, 40.2, 43.3, 47.5, 48.0, 54.1, 75.1, 76.2, 81.4, 83.2, 106.1, 131.1, 144.1, 235.0, 236.1, 240.2, 249.1, 263.2, 286.1, 299.2 [28.0, 72.2, 97.0, 214.2, 265.1, 290.2, 300.0] (8.0, 94.1, 107.1, 108.1, 111.1, 125.1, 127.1, 138.1, 143.1, 149.1, 152.0, 154.1, 188.0, 196.1, 210.1, 216.1, 223.1, 237.1, 256.3)

Escavadodon

12.2, 43.3, 44.0, 45.0, 47.0, 51.3, 53.0, 63.0, 67.0, 76.3, 224.1, 225.1, 263.2, 299.2 (16.0, 20.0, 28.1, 33.2, 72.1, 221.0, 222.2, 226.0, 231.0, 245.0, 252.1)

Palaeosinopa

11.1, 33.0, 36.0, 39.2, 53.2, 54.3, 56.2, 58.2, 76.5, 81.4, 83.2, 112.0, 121.1, 130.2, 145.1, 165.1, 168.0, 170.1, 183.0, 194.0, 213.0, 249.4, 251.0, 256.0, 268.0, 271.1, 273.3, 303.1 [51.1, 97.0, 215.0] (14.1, 16.0, 20.0, 35.0, 38.0, 87.2, 88.2, 129.2, 132.1, 152.0, 169.1, 226.0, 245.0, 252.1, 272.1, 308.0)

Node 37 - Pentacodontidae

17.2, 20.2, 50.0 [8.2, 77.0, 78.0, 105.0, 106.1, 265.3] (14.1, 34.0, 36.4, 38.0)

Aphronorus

9.2, 15.1, 16.2, 21.1, 23.1, 27.1, 31.0, 33.0, 36.3, 40.2, 44.0, 55.2, 60.1, 64.0, 70.2, 72.1, 75.0, 76.5 [35.1, 62.1] (8.2, 51.2, 78.0, 94.1, 105.0, 106.1, 219.0, 265.3)

Coriphagus

13.0, 25.1, 40.0, 54.3, 55.0, 56.2, 59.2, 67.0, 74.2, 81.0, 83.0 [51.1] (35.0, 62.0, 75.2)

Node 30

6.2, 10.2, 11.1, 13.0, 16.2, 20.3, 27.1, 37.1, 111.4, 256.3, 266.2, 295.3 [31.0, 78.2, 91.1, 104.2, 116.1, 117.1, 118.1, 119.1, 120.1, 122.0, 125.1, 157.1, 189.0, 192.0, 226.1, 245.2, 248.0, 250.2, 294.1, 308.1] (34.0, 36.0, 77.1, 87.2, 262.1)

Macrocranion

1.3, 3.3, 11.0, 17.1, 23.1, 53.2, 54.0, 56.0, 57.1, 60.1, 72.1, 77.2, 83.2, 148.1, 182.0, 185.1, 209.0, 212.0, 249.4, 261.2, 273.0, 278.1, 285.1, 288.1, 300.1 [14.0, 35.1, 94.0, 97.0, 129.1, 219.2, 275.1] (31.0, 32.1, 51.2, 55.0, 78.2, 157.1, 188.0, 245.2, 294.1)

Node 31

9.2, 33.0, 56.2, 64.2, 70.0, 72.3, 75.5, 79.0, 88.1, 111.5, 236.2, 238.2, 262.2, 279.0, 284.2, 286.1, 305.0 [32.0, 41.1, 46.1, 48.1, 51.1, 63.3, 71.3, 86.2, 207.2, 263.2, 292.1] (14.1, 71.1, 91.1, 97.1, 219.0, 237.1, 250.1, 275.0)

Node 32 - Lipotyphla

20.4, 40.2, 44.2, 47.9, 59.2, 65.4, 84.1, 98.1, 105.0, 106.1, 150.1, 181.1, 258.3, 269.0, 271.0, 280.2, 295.0, 302.2, 304.2, 308.0 [1.4, 3.4, 5.2, 10.3, 22.1, 31.1, 73.1, 108.1, 127.0, 163.2, 180.1, 183.0, 206.1, 215.0, 229.1, 234.2, 277.0, 284.1, 285.4, 290.1, 292.2, 296.0] (46.1, 48.1, 55.0, 63.3, 78.2, 86.2, 94.1, 157.1, 207.2)

Soricidae

1.6, 3.6, 6.5, 11.0, 28.1, 29.2, 32.1, 36.1, 41.3, 45.0, 49.2, 64.3, 76.5, 78.3, 88.2, 89.2, 111.6, 112.3, 114.0, 125.2, 144.1, 148.3, 151.1, 161.2, 171.0, 175.0, 181.3, 193.1, 194.0, 195.0, 204.2, 209.0, 213.0, 226.0, 230.1, 231.0, 233.3, 243.1, 249.4, 251.0, 257.0, 266.1, 270.1, 286.2, 299.2 [122.1, 132.0, 154.0, 188.1] (5.2, 10.3, 22.1, 35.0, 71.3, 73.1, 104.2, 108.1, 117.1, 129.2, 163.2, 180.1, 183.0, 222.2, 229.1, 234.2, 245.2, 250.2, 263.2, 277.0, 284.1, 285.4, 290.1, 292.2, 294.1, 296.0)

Node 33 - Erinaceidae

11.2, 13.2, 17.1, 39.3, 45.2, 51.2, 60.1, 62.0, 71.2, 74.2, 81.4, 87.3, 90.2, 110.1, 153.1, 155.1, 164.1, 178.0, 186.2, 191.1, 217.1, 220.1, 226.2, 232.1, 260.1, 261.1, 268.0, 277.1 [5.3, 35.1, 41.0, 117.0, 129.1, 197.2, 198.1, 222.1, 250.1, 263.1] (5.0, 116.1, 118.1, 119.1, 120.1, 122.0, 132.1, 154.1, 188.0, 206.1)

Echinosorex

1.3, 3.3, 6.1, 9.1, 16.1, 19.4, 24.0, 27.0, 50.2, 58.0, 76.3, 83.2, 93.1, 101.0, 115.1, 127.1, 130.0, 143.1, 157.2, 168.0, 184.0, 216.1, 223.1, 236.1, 249.1, 250.0, 253.1, 254.1, 265.3, 267.1, 295.1, 303.1 [5.0, 10.2, 73.0, 108.0, 180.0, 183.1, 229.0, 290.2] (22.1, 104.2, 125.1, 163.2, 197.2, 198.1, 234.2, 245.2, 284.1, 285.4, 292.2, 294.1, 296.0)

Erinaceus

6.3, 8.2, 17.2, 18.1, 31.0, 33.3, 34.2, 42.0, 46.0, 51.3, 53.2, 72.4, 73.2, 75.3, 77.2, 79.2, 85.0, 97.0, 100.1, 103.1, 104.0, 107.1, 130.2, 148.1, 149.1, 152.0, 165.1, 169.1, 198.2, 218.1, 219.1, 221.0, 227.1, 232.2, 241.1, 243.3, 244.0, 245.0, 246.1, 248.1, 249.3, 252.1, 258.4, 261.2, 273.1, 279.1, 289.2, 290.0, 306.0 [22.0, 125.0, 163.0, 234.1, 284.2, 285.3, 292.0, 294.2, 296.1] (1.4, 3.4, 5.3, 10.3, 101.2, 108.1, 180.1, 183.0, 229.1)

Node 34 - Chiroptera

1.2, 8.2, 39.0, 43.0, 214.0, 249.0, 256.1, 280.0, 301.1, 309.1, 310.1 [47.2, 51.0, 55.1, 75.4, 76.0, 78.1, 87.4, 105.2, 133.1, 140.2, 157.0, 168.0, 172.2, 173.1, 179.0, 186.1, 208.0, 211.0, 218.1, 221.0, 224.2, 225.2, 233.2, 242.1, 243.2, 244.0, 245.0, 246.1, 257.2, 265.2, 289.3, 294.2, 296.2, 308.2] (31.0, 41.1, 71.3, 222.2)

Onychonycteris

6.3, 62.0, 66.1, 74.0, 77.2, 78.0, 212.0 (35.0, 51.0, 207.2)

Node 35

3.7, 6.1, 37.0, 77.0, 81.2, 167.1 [207.1, 286.2, 290.3] (47.2, 75.4, 76.0, 242.1, 245.0)

Icaronycteris

13.1, 27.0, 33.2, 34.2, 36.2, 50.0, 63.1, 78.2, 256.0 [35.1, 46.2, 48.2] (51.0, 86.2, 188.0)

Rhinolophus

10.4, 11.0, 17.2, 24.0, 32.1, 40.2, 41.2, 44.3, 48.0, 53.2, 54.1, 55.0, 59.2, 64.3, 65.2, 70.1, 88.2, 89.2 [51.1, 86.1, 188.1] (35.0, 46.1, 63.3, 87.4, 101.2, 105.2, 117.1, 122.0, 125.1, 127.1, 129.2, 132.1, 133.1, 140.2, 168.0, 172.2, 173.1, 179.0, 186.1, 208.0, 211.0, 215.1, 218.1, 221.0, 224.2, 225.2, 233.2, 243.2, 244.0, 246.1, 250.2, 257.2, 263.2, 265.2, 286.2, 289.3, 290.3, 292.1, 296.2, 308.2)

Node 28

16.0, 44.2, 46.1, 47.7, 62.2, 109.1, 133.1, 288.1, 290.1 [77.0, 143.2, 186.1, 204.0, 210.1, 232.2, 233.1, 257.2, 267.1, 276.1, 299.3, 306.0] (35.0, 43.3, 51.2, 71.1, 74.2, 94.1, 129.2, 248.1, 295.0)

Loxolophus

23.1, 37.1, 45.2, 48.1, 58.0, 75.2, 76.1, 78.0, 79.2, 89.2, 240.0, 249.4, 258.1, 298.0, 302.3 [36.1, 55.1, 215.0] (97.1, 219.0, 226.0)

Arctocyon

17.1, 24.0, 33.0, 39.3, 40.2, 51.3, 53.2, 54.3, 56.2, 60.1, 61.1, 71.3, 73.1, 79.0, 83.2, 87.0, 105.0, 106.1, 112.0, 214.2, 219.3, 270.1, 284.3, 304.1 [97.0, 226.1, 249.1] (36.0, 55.0, 101.2, 121.1, 140.2, 143.2, 186.1, 204.0, 210.1, 215.1, 217.1, 232.2, 237.1, 245.0, 257.2, 262.1, 267.1, 276.1, 299.3, 308.0)

Protungulatum

23.1, 24.0, 28.1, 29.1, 33.2, 34.2, 39.0, 44.0, 53.2, 54.3, 71.2, 73.1, 77.2, 78.2, 83.2, 168.2, 170.1, 181.1, 219.1, 221.0, 292.1, 294.1, 296.2, 299.2 [63.0, 161.0] (36.0, 43.3, 47.4, 55.0, 74.2, 86.2, 226.0, 295.0)

Node 4 - Adapisoriculidae

38.1, 56.2, 222.2, 225.2, 273.1, 299.2 [16.2, 19.1, 20.3, 54.1, 75.0, 76.5, 217.0, 251.0] (263.2, 275.1, 276.1)

Deccanolestes

33.0, 34.0, 55.0 [9.0] (280.2)

Afrodon/Bustylus

33.2, 36.4, 52.0, 74.1 (9.1, 16.2, 19.1, 20.3, 76.5, 248.1)

 
 

APPENDIX 1. List of characters

Note that the zero state is primitive unless otherwise stated. Multistate characters are ordered unless stated to be unordered or a stepmatrix.

Teeth

1. Upper incisors: more than 3, simple structure, similar-sized (0); 3, simple structure, similar sized (1); 2, simple structure, similar-sized (2); 3 or 2, similar-sized, spatulate (3); 3 or 2, the first enlarged without heel (posterocone) (4); 2, both enlarged, simple (5); 3, the first enlarged with heel (posterocone) but only one terminal cusp (6); 3 or 2, the first enlarged, with heel (posterocone) and 2 or 3 terminal cusps (7); 3 or 2, similar-sized, serrated/denticulate (8); 1 or none (9). Primitive state based on Eomaia and Vincelestes. STEPMATRIX.

  0 1 2 3 4 5 6 7 8 9
0 - 1 2 2 2 3 3 4 2 3
1 1 - 1 1 1 2 2 3 1 2
2 2 1 - 2 2 1 3 4 2 1
3 2 1 2 - 2 3 3 4 2 1
4 2 1 2 2 - 1 1 2 2 3
5 3 2 1 3 1 - 2 3 3 2
6 3 2 3 3 1 2 - 1 3 4
7 4 3 4 4 2 3 1 - 4 5
8 2 1 2 2 2 3 3 4 - 1
9 3 2 1 1 3 2 4 5 1 -

2. Mesial gap between mesialmost upper teeth: absent (0); present (1). Primitive state based on Eomaia and Vincelestes. Modified from Wible et al. (2009) character 9.

3. Lower incisors: 4, simple structure, similar-sized (0); 4, first enlarged (1); 3, simple structure, similar-sized (2); 3 or 2, similar-sized, spatulate (3); 3 or 2, first enlarged, simple (4); 3, first two enlarged, simple (5); first incisor of 3 or 2, or sole if one, enlarged and serrated (6); 3 or 2, similar-sized, serrated/denticulate (7); absent (8). Primitive state based on Eomaia and Prokennalestes. STEPMATRIX.

  0 1 2 3 4 5 6 7 8
0 - 1 1 2 2 2 3 2 3
1 1 - 2 3 1 2 2 3 4
2 1 2 - 1 1 1 2 1 2
3 2 3 1 - 2 2 3 2 1
4 2 1 1 2 - 1 1 2 3
5 2 2 1 2 1 - 2 2 3
6 3 2 2 3 1 2 - 1 2
7 2 3 1 2 2 2 1 - 1
8 3 4 2 1 3 3 2 1 -

4. DI2 /2 simple, small, fully enamelled, rooted (if unknown, inferred from state of I2 /2 or form of the dentary), or absent (0); hypselodont, chisel-shaped with buccally restricted enamel (1). Primitive state based on ingroup commonality.

5. Upper canine: large, two-rooted (0); large, single-rooted (1); small, single rooted (2); small, two-rooted (3); absent (4). Primitive state based on Vincelestes. Modified from Wible et al. (2009) characters 23-24. STEPMATRIX.

  0 1 2 3 4
0 - 1 2 1 2
1 1 - 1 2 2
2 2 1 - 1 1
3 1 2 1 - 1
4 2 2 1 1 -

6. Lower canine: large, two-rooted (0); large, single-rooted (1); small, single rooted, without talonid cusp (2); small, single rooted, with talonid cusp (3); small, two-rooted (4); absent (5). Primitive state based on Prokennalestes. Modified from Wible et al. (2009) characters 25-26. STEPMATRIX.

  0 1 2 3 4 5
0 - 1 2 2 1 2
1 1 - 1 2 2 2
2 2 1 - 1 1 1
3 2 2 1 - 1 1
4 1 2 1 1 - 1
5 2 2 1 1 1 -

7. P3 /3 present (0); absent (1). Primitive state based on Prokennalestes.

8. P1 root number: 2 (0); 1 (1); tooth absent (2). Primitive state based on Juramaia. Modified from Wible et al. (2009) character 33.

9. P1 root number: 2 (0); 1 (1); tooth absent (2). Primitive state based on Juramaia. Modified from Wible et al. (2009) character 48.

10. P2 root number: 2 or more (0); 2 upper, one lower (1); 1 (2); 2 or one upper, P2 absent (3); P2 absent (4). Primitive state based on Juramaia. Those only known from P2 are assumed to have same P2 root number.

11. P4 root number: 1 (0); 2 (1) PRIMITIVE; 3 (2). Primitive state based on Juramaia.

12. P4 metacone absent (0); small, close to paracone (1); large, distant from paracone (2); no cusp structure (3). Primitive state based on Juramaia and Prokennalestes. Modified from Hooker and Russell (2012) character 2. STEPMATRIX

  0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 1
3 1 1 1 -

13. P4 protocone lobe essentially absent (0); small, short with small protocone (1); relatively long with small protocone (2); relatively long with large protocone (3). Primitive state based on Juramaia and Prokennalestes. Modified from Hooker and Russell (2012) character 3. STEPMATRIX

  0 1 2 3
0 - 1 2 3
1 1 - 1 2
2 2 1 - 1
3 3 2 1 -

14. P4 parastyle mesially projecting (0); poorly developed (1). Primitive state based on Juramaia and Prokennalestes. Modified from Hooker and Russell (2012) character 4.

15. P5 metacone small to absent (0); prominent but smaller than paracone (1); as large as paracone (2). Primitive state based on Juramaia and Prokennalestes. Modified from Hooker and Russell (2012) character 5.

16. P5 preparacrista: strong (0); weak (1); absent (2). Primitive state based on Juramaia and Prokennalestes. Modified from Hooker and Russell (2012) character 6.

17. P5 parastyle large, mesially projecting (0); medium (1); small (2). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 7.

18. P5 metastylar wing (homologue of postmetacrista or, when metacone is absent, of postparacrista): continuous (0); cleft (1) PRIMITIVE; deeply notched, short (2); deeply notched, long (3). Primitive state based on Prokennalestes.

19. P5 preprotocrista-paracingulum complex without paraconule (0); paraconule present on preprotocrista-paracingulum complex (1); paraconule present with postparaconule crista (2); preprotocrista not linked to parastyle and without paraconule or postparaconule crista, precingulum (if present) joining paracingulum (3); preprotocrista linked to paracone and without paraconule or postparaconule crista, precingulum joining paracingulum (4). Primitive state based on Prokennalestes and Juramaia. Modified from Hooker and Russell (2012) character 9. STEPMATRIX.

  0 1 2 3 4
0 - 1 2 1 1
1 1 - 1 2 2
2 2 1 - 3 3
3 1 2 3 - 1
4 1 2 3 1 -

20. (see s figure19Appendix Figure 1). P5 with: undifferentiated postprotocrista + metacingulum from protocone to metastyle (0); undifferentiated postprotocrista + metacingulum, with weak, short postcingulum (1); confluent postprotocrista + metacingulum, orientated distobuccally, postcingulum strong, long, with incipient to small hypocone (2); postprotocrista orientated nearly distally, joining long distal cingulum (3); postprotocrista short, distally orientated, joining prehypocrista, with hypocone (4); postprotocrista orientated buccally to distobuccally towards metacone or (where metacone absent) towards postparacrista, independent of distal cingulum (composed of postcingulum + metacingulum), with postflexus (5); as (5), but with no postflexus (6); as (6), but with lingual end of distal cingulum joind to protocone as postprotocingulum (7); postprotocrista deflected distally with buccal part weak to absent, distal cingulum present or absent (8). Primitive state based on Prokennalestes and Juramaia. Modified from Hooker and Russell (2012) character 10. STEPMATRIX.

  0 1 2 3 4 5 6 7 8
0 - 1 2 2 3 3 4 5 4
1 1 - 1 1 2 2 3 4 3
2 2 1 - 2 3 1 2 3 2
3 2 1 2 - 1 3 4 5 4
4 3 2 3 1 - 4 5 6 5
5 3 2 1 3 4 - 1 2 1
6 4 3 2 4 5 1 - 1 2
7 5 4 3 5 6 2 1 - 3
8 4 3 2 4 5 1 2 3 -

21. P5 metaconule +/- premetaconule crista: absent (0); present (1). Primitive state based on Prokennalestes.

22. P5 : without talon shelf (0); with talon shelf (1). Primitive state based on Prokennalestes and Juramaia.

23. P5 : without precingulum (0); with precingulum (1). Primitive state based on Prokennalestes and Juramaia. Wible et al. (2009) character 43.

24. P5 distinct ectoflexus: present (0); absent (1). Primitive state based on Prokennalestes.

25. P5 buccal protocone crest (central crista of Archibald et al., 2011) absent (0); present (1). Primitive state based on Prokennalestes.

26. P5 protocone lobe transversely orientated (0); twisted mesially, making protocone tilt distally (1). Primitive state based on Prokennalestes.

27. P4 /4 subequal in size to P5 /5 (0); much smaller (1). Primitive state based on Prokennalestes and Juramaia.

28. P4 paraconid height: low (0); nearly half the height of the protoconid (1); more than half the height of the protoconid (2). Primitive state based on Prokennalestes.

29. P4 paraconid strength: weak (0); medium (1); strong (2). Primitive state based on Prokennalestes.

30. P4 metaconid: absent (0); about half the height of the protoconid, lingually placed (1); of subequal height to the protoconid, part of widely open trigonid (2); about half the height of the protoconid, distally placed (3). Primitive state based on Prokennalestes. STEPMATRIX.

0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 2
3 1 1 2 -

31. P4 talonid: single-cusped, no basin (0); single-cusped, tiny basin (1) PRIMITIVE; prominent hypoconid (the single cusp), small basin (2); basined with hypoconid and hypoconulid (3); basined with hypoconid, hypoconulid and entoconid (4); basined with hypoconid and entoconid (5). Primitive state based on Prokennalestes. STEPMATRIX.

  0 1 2 3 4 5
0 - 1 2 3 4 3
1 1 - 1 2 3 2
2 2 1 - 1 2 1
3 3 2 1 - 1 2
4 4 3 2 1 - 1
5 3 2 1 2 1 -

32. P5 : not exodaenodont (0); exodaenodont (1). Primitive state based on Prokennalestes.

33. P5 paraconid height: low (0); about a third of the height of the protoconid (1) PRIMITIVE; about half the height of the protoconid (2); more than half the height of the protoconid (3). Primitive state based on Prokennalestes.

34. P5 paraconid strength: weak (0); medium (1) PRIMITIVE; strong (2). Primitive state based on Prokennalestes.

35. P5 metaconid: absent or negligeably developed (0); about half the height of the protoconid, lingually placed (1); of subequal height to the protoconid, part of widely open trigonid (2); of subequal height to the protoconid, part of fully formed short trigonid (3). Primitive state based on Prokennalestes.

36. P5 talonid: single-cusped, basin tiny or absent (0); prominent hypoconid (the single cusp), small basin (1); basined with hypoconid and hypoconulid (2); basined with hypoconid, hypoconulid and entoconid (3); basined with hypoconid and entoconid (4); basined with hypoconid, entoconid and doubled entoconid (also lower molars) (5). Primitive state based on Prokennalestes. STEPMATRIX.

  0 1 2 3 4 5
0 - 1 2 3 2 4
1 1 - 1 2 1 3
2 2 1 - 1 2 2
3 3 2 1 - 1 1
4 2 1 2 1 - 2
5 4 3 2 1 2 -

37. M1 transverse axis: buccolingual (0); slightly distobuccal-mesiolingual (1). Primitive state based on Prokennalestes. Hooker and Russell (2012) character 28.

38. M1 metacone: smaller than paracone (0); equal sized or slightly larger (1). Primitive state based on Prokennalestes. Hooker and Russell (2012) character 29.

39. M1 preparacrista: nearly transverse, joining stylocone (0); directed mesiobuccally, joining parastyle (possibly including a reduced, mesially shifted stylocone) (1); distinct, directed mesially (2); indistinct, directed mesially (3). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 30.

40. M1 parastyle: massive and shelf-like (0); large, cuspate (1); small (2). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 31.

41. Preultimate upper molar centrocrista: straight or slightly lingually deflected (0); buccally deflected, but not reaching buccal margin (1); buccally deflected, meeting buccal margin at single mesostyle (2); buccally deflected, meeting buccal margin at double mesostyle (3). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 33.

42. Preultimate upper molar postmetacrista: strong, cleft or notched (0); strong, continuous (1); weak, cleft (2). Primitive state based on Prokennalestes. UNORDERED.

43. M1 postmetacrista: distinct, long, nearly buccally directed (0); distinct, long, curved or angled distobuccally then buccally (1) PRIMITIVE; distinct, short distobuccally directed (2); distinct, short, distally directed (3); indistinct, short, distally directed (4). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 34.

44. M1 : paraconule and metaconule situated close to protocone (0); paraconule closer to protocone than is metacone (1); paraconule and metaconule equally more distant from protocone (2); paraconule close to protocone, metaconule absent (3). Primitive state based on Prokennalestes. Modified from Hooker and Russell (2012) character 35. STEPMATRIX.

  0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 2
3 1 1 2 -

45. Preultimate upper molar(s): preprotocrista +/- paracingulum (incorporating preparaconule crista) without precingulum (0); Preultimate upper molar(s): preprotocrista + paracingulum with precingulum (1); precingulum and paracingulum joined, from which preprotocrista (including preparaconule crista) is isolated and which may join paracone (2). Primitive state based on Prokennalestes.

46. Preultimate upper molar paraconule and postparaconule crista absent (0); paraconule present, postparaconule crista absent (1) PRIMITIVE; paraconule and postparaconule crista present (2). Primitive state based on Prokennalestes.

s figure2047. (see Appendix Figure 2). Preultimate upper molar: postprotocrista +/- metacingulum present, postcingulum absent (0); postprotocrista + metacingulum present, postcingulum present (1); postprotocrista + metacingulum present, postcingulum expanded into a short hypocone shelf (2); postprotocrista +/- metacingulum present, short hypocone shelf bearing hypocone (3); postprotocrista + metacingulum present, postcingulum expanded into a long hypocone shelf (4); postprotocrista + metacingulum present, long hypocone shelf bearing hypocone (5); postcingulum confluent with metacingulum, bypassing postprotocrista (6); postcingulum confluent with metacingulum, bypassing postprotocrista and bearing hypocone (7); as (6) but with postprotocingulum (Nannopithex fold) joining or nearly joining hypocone shelf (8); as (7) but with prehypocrista (erinaceid crest) (9). Primitive state based on Prokennalestes. STEPMATRIX.

  0 1 2 3 4 5 6 7 8 9
0 - 1 2 3 3 4 4 5 4 6
1 1 - 1 2 2 3 3 4 3 5
2 2 1 - 1 1 2 2 3 2 4
3 3 2 1 - 2 1 3 2 3 3
4 3 2 1 2 - 1 1 2 1 3
5 4 3 2 1 1 - 2 1 2 2
6 4 3 2 3 1 2 - 1 1 2
7 5 4 3 2 2 1 1 - 2 1
8 4 3 2 3 1 2 1 2 - 3
9 6 5 4 3 3 2 2 1 3 -

48. Preultimate upper molar: metaconule and premetaconule crista absent (0); metaconule present, premetaconule crista absent (1) PRIMITIVE; metaconule and premetaconule crista present (2). Primitive state based on Prokennalestes.

49. Preultimate upper molar inflated talon shelf: absent (0); present, restricted lingually (1); present, extending buccally (2). Primitive state based on Prokennalestes.

50. Preultimate upper molar ectoflexus: deep on M2 , shallow on M1 (0); shallow on M2 and shallow or absent on M1 (1); absent (2). Primitive state based on Prokennalestes.

51. Upper molar buccal margin with: stylar shelf nearly a third of the tooth width (0); stylar shelf narrow (1); no stylar shelf, strong ectocingulum (2); no stylar shelf, ectocingulum weak or incomplete (3); stylar shelf and ectocingulum absent (4). Primitive state based on Prokennalestes.

52. Preultimate upper molar stylar cusp D present (0); absent (1). Primitive state based on Prokennalestes.

53. Upper molar protocone tilt: lingual (0); more or less vertically orientated from crown base (1); buccal (2). Primitive state based on Prokennalestes.

54. M1 paraconid height from lingual side: about a third the height of the protoconid (0); about half the height of the protoconid (1) PRIMITIVE; about two thirds the height of the protoconid (2); nearly as tall as protoconid (3). Primitive state based on Prokennalestes.

55. Transverse position of M1 paraconid tip with respect to that of metaconid: lingual (0); sublingual (1) PRIMITIVE; median (2). Primitive state based on Prokennalestes.

56. M1 paraconid strength: weak (0); medium (1) PRIMITIVE; strong (2). Primitive state based on Prokennalestes.

57. M1 paraconid shape: cuspate (0); crestiform (1). Primitive state based on Prokennalestes.

58. Lower molar paracristid notch: absent (0); present as cleft or angle (1) PRIMITIVE; present as deep (carnassial) notch (2). Primitive state based on Prokennalestes.

59. Shape of M1 trigonid: very narrow mesiodistally (0); approximately an equilateral triangle (1) PRIMITIVE; elongate mesially (2). Primitive state based on Prokennalestes.

60. M2 paraconid: cuspate (0); crestiform (1). Primitive state based on Prokennalestes.

61. M2 paraconid: separate from metaconid (0); the two cusps connate (1). Primitive state based on Prokennalestes.

62. M1 hypoconid height: half or less than half height of protoconid (0); about two thirds the height of protoconid (1); subequal (2). Primitive state based on Prokennalestes.

63. M1 entoconid: less than half height of metaconid (0); about half height of metaconid (1); about two thirds height of metaconid (2); the two cusps subequal (3). Primitive state based on Prokennalestes.

64. Lower molar hypoconulid buccal of midline (0); central (1) PRIMITIVE; lingual of midline (2); at or near lingual margin, nyctalodont condition (3). Primitive state based on Prokennalestes.

65. Lower molar cristid obliqua meets trigonid: lingually, rising up distolingual edge of metaconid (0); at about midpoint, rising obliquely up the back of the metaconid (1); at about midpoint, not or scarcely rising up back of trigonid (2); slightly buccal of midpoint, not or scarcely rising up back of trigonid (3); nearly at buccal margin, not or scarcely rising up back of trigonid (4); from a buccally convex trajectory, rising up the buccal side of the metaconid (5). Primitive state based on Prokennalestes.

66. Lower molar cristid obliqua: unnotched (0); notched (1). Primitive state based on Prokennalestes.

67. M1 talonid: narrower than trigonid (0); equal in width to trigonid (1); wider than trigonid (2). Primitive state based on Prokennalestes.

68. Lower molar posthypocristid: complete or only narrowly notched (0); broken (1). Primitive state based on Prokennalestes.

69. Lower molar protocristid: fissured with carnassial notch (0); angled/cleft (1) PRIMITIVE; a smooth continuous crest (2). Primitive state based on Prokennalestes.

70. Lower molar precingulid: strong (0); weak (1); absent (2). Primitive state based on Prokennalestes.

71. Preultimate lower molar postcingulid: absent (0); present basally (1); rising to join hypoconulid (2); present basally, extended mesially to join precingulid, i.e., forming ectocingulid (3). Primitive state based on Prokennalestes. STEPMATRIX.

  0 1 2 3
0 - 1 2 2
1 1 - 1 1
2 2 1 - 2
3 2 1 2 -

72. M3 hypoconulid: distally protruding, inflated to enclose a basin (0); distally protruding, a distinct cusp (1) PRIMITIVE; contracted, not protruding significantly from talonid, entoconid present (2); contracted, not protruding significantly from talonid, entoconid absent (3); absent together with talonid or entire tooth (4). Primitive state based on Prokennalestes.

73. M3 hypoconulid: overhangs distal tooth margin (0); distally extensive basally beyond tip (1); absent together with talonid or entire tooth (2). Primitive state based on Prokennalestes.

74. Relative height of molar cusps judged from height of M1/2 protoconid as % of tooth length: more than 95% (0); between 75 and 95% (1); less than 75% (2). Primitive state based on Prokennalestes.

Dentary

75. Anterior mental foramen below P2 (0); between P1 and P2 (1); below P1 (2); between P4 and P5 (3); at extreme anterior position (4); absent (5). Primitive state based on Juramaia. Modified from Wible et al. (2009) character 129. STEPMATRIX.

  0 1 2 3 4 5
0 - 1 2 1 1 1
1 1 - 1 2 2 1
2 2 1 - 2 1 1
3 1 2 2 - 2 1
4 1 2 1 2 - 1
5 1 1 1 1 1 -

76. Posterior mental foramen: below P2 (0); between P2 and P4 (1); below P4 (2); between P4 and P5 (3) PRIMITIVE; below P5 (4); between P5 and M1 or below M1 (5). Primitive state based on Juramaia. Modified from Wible et al. (2009) character 130.

77. Height of condyle above alveolar line: one molar length (0); two molar lengths (1); three molar lengths (2); four molar lengths or more (3). Primitive state based on Juramaia. Modified from Wible et al. (2009) character 150.

78. Tilting of coronoid process; angle of anterior border to alveolar line: 120-130 degrees (0); 110 degrees (1); 100 degrees (2); 90 degrees (3). Primitive state based on Juramaia and Prokennalestes. Modified from Wible et al. (2009) character 135.

79. Height of coronoid process above alveolar line: two to three molar lengths (0); four molar lengths (1); five molar lengths or more (2). Primitive state based on Juramaia and Prokennalestes. Modified from Wible et al. (2009) character 133.

80. Lateral mandibular foramen present (0); absent (1). Primitive state based on Eomaia, Juramaia and Prokennalestes. Modified from Wible et al. (2009) character 139: polarity reversed.

81. Angular process directed posteroventrally (0); posteroventromedially (1); posteroventrolaterally (2); posteriorly (3); dorsally to dorsomedially inflected (4); in the form of a rounded posteroventral corner (5); rounded with dorsal hook (6). Primitive state based on Eomaia, Juramaia and Prokennalestes. Modified from Wible et al. (2009) character 143. STEPMATRIX.

  0 1 2 3 4 5 6
0 - 1 1 1 2 2 3
1 1 - 2 1 2 2 3
2 1 2 - 1 2 2 3
3 1 1 1 - 1 1 2
4 2 2 2 1 - 2 1
5 2 2 2 1 2 - 1
6 3 3 3 2 1 1 -

82. Depression in ventral dentary margin anterior to angle present (0); absent (1). Primitive state based on Eomaia, Juramaia and Prokennalestes.

83. Angular process projects posteriorly less than dentary condyle (0); about equal (1); farther (2). Primitive state based on Eomaia, Juramaia and Prokennalestes. Wible et al. (2009) character 147.

84. Angular process vertical position: at posteroventral border (0); posterodorsal, at or near alveolar border (1). Primitive state based on Eomaia, Juramaia and Prokennalestes. Wible et al. (2009) character 146.

85. Dentary condyle with posteriorly directed peduncle (0); without (1). Primitive state based on Eomaia, Juramaia and Prokennalestes. Wible et al. (2009) character 148. Cimolestes magnus recoded 1 (Lillegraven 1969, figure 37.3); Erinaceus recoded 0 (personal obs.).

86. Vertical position of mandibular foramen: near ventral margin at root of angle (0); recessed dorsally from ventral margin, but below alveolar plane (1); recessed dorsally from ventral margin, at or above alveolar plane (2). Primitive state based on Eomaia and Prokennalestes. Wible et al. (2009) character 157.

Rostrum

87. Premaxilla-maxilla suture on face: vertically orientated, contacting nasals (0); projecting posterodorsally no further than canine, but without contacting frontal (1) PRIMITIVE; projecting posterodorsally further than canine, but without contacting frontal (2); projecting posterodorsally and contacting frontal (3); premaxilla lacking facial process (4). Primitive state based on Vincelestes. Modified from Wible et al. (2009) characters 160-162. STEPMATRIX.

  0 1 2 3 4
0 - 1 2 2 1
1 1 - 1 2 1
2 2 1 - 1 1
3 2 2 1 - 1
4 1 1 1 1 -

88. Infraorbital foramen dorsal to: P4 or more anterior (0); dorsal to P5 (1); Dorsal to M1 or more posterior (2). Primitive state based on Eomaia and Prokennalestes. Wible et al. (2009) character 165, with changed state numbering.

89. Vertical position of anterior margin of orbit with respect to tooth loci. Above P5 or more anterior (0); M1 (1) PRIMITIVE; M2 or more posterior (2). Primitive state based on Eomaia. This, together with character 15, approximates length of infraorbital canal (Wible et al., 2009, character 166).

90. Nasal widest posteriorly (0); with sides subparallel (1); widest anteriorly (2). Primitive state based on Eomaia. Wible et al. (2009) character 168.

91. Nasal overhangs external nasal aperture (0); does not overhang (1). Primitive state based on Vincelestes. Wible et al. (2009) character 169.

92. Nasofrontal suture with medial process of frontals wedged between nasals (0); not wedged between nasals (1). Primitive state based on Eomaia. Wible et al. (2009) character 170.

93. Posterior margin of nasals posterior to or even with anterior orbital rim (0); anterior to anterior orbital rim (1). Primitive state based on Eomaia. Modified from Wible et al. (2009) character 171.

94. Anterior process of frontal weak/absent (0); elongate/thin (1). Primitive state based on Eomaia. Modified from Wible et al. (2009) character 174.

95. No frontal maxillary contact on rostrum (0); contact (1). Primitive state based on Eomaia. Wible et al. (2009) character 173.

96. Facial process of lacrymal large, triangular and pointed anteriorly (0); small and rectangular or crescentic (1). Primitive state based on Eomaia. Wible et al. (2009) character 177. Erinaceus rescored 1 as, although fused to maxilla in adult, its extent can be judged from juveniles (Butler, 1948).

97. Lacrymal tubercle present (0); absent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 178.

98. Lacrymal foramen not exposed on face (0); exposed on face (1). Primitive state based on Eomaia. Wible et al. (2009) character 179, but with reversed state numbering.

99. Number of lacrymal foramina: two (0); one (1). Primitive state based on Vincelestes. Wible et al. (2009) character 180.

100. Lacrymal foramen within lacrymal (0); with maxillary contribution (1); with jugal contribution (2). UNORDERED. Primitive state based on Vincelestes. Wible et al. (2009) character 181.

Palate

101. Premaxillary-maxillary suture on palate wedge-shaped, pointing anteriorly (0); transverse (1) PRIMITIVE; wedge-shaped, pointing posteriorly (2). Primitive state based on Vincelestes. Wible et al. (2009) character 184, with states 0 and 1 reversed.

102. Incisive foramina small, length of one or two incisors (0); length of three or four incisors (1). Primitive state based on Vincelestes. Wible et al. (2009) character 185.

103. Palatal vacuities absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 187.

104. Major (anterior) palatal foramen single within palatine (0); single between palatine and maxilla (1); single within maxilla (2). Primitive state based on Vincelestes. Wible et al. (2009) character 188.

105. Palatal expansion posterior to last molar (0); even with last molar (1) PRIMITIVE; anterior to last molar (2). Primitive state based on Vincelestes. Wible et al. (2009) character 190, with states 0 and 1 reversed.

106. Postpalatine torus absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 191.

107. Posterior nasal spine weak or absent (0); prominent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 192. Ptilocercus rescored as 0 (pers. obs.).

108. Minor (posterior) palatal foramen small, single or multiple within palatine or between maxilla and palatine (0); large with thin posterior bony bridge, within palatine or between maxilla and palatine (1); large, with thin posterior bony bridge between palatine and pterygoid (2); absent (3). Primitive state based on Vincelestes. Combined from Wible et al. (2009) characters 193 and 194. Ptilocercus rescored as 0 (personal obs.). STEPMATRIX.

  0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 1
3 1 1 1 -

109. Maxilla without large shelf-like expansion posterior to last molar (0); with (1). Primitive state based on Vincelestes. Wible et al. (2009) character 195.

Zygoma

110. Posterior edge of anterior zygomatic root aligned with last molar (0); with anterior molars (1). Primitive state based on Eomaia. Wible et al. (2009) character 196.

111. Zygomatic process of maxilla: not bifurcated, jugal contribution to anteroventral orbit and zygoma, and jugal-lacrymal contact (0); bifurcated, jugal contribution to anteroventral orbit and zygoma, and jugal-lacrymal contact (1); bifurcated, jugal contribution to anteroventral orbit and zygoma, but no jugal-lacrymal contact (2); absent/vestigial, jugal contribution to anteroventral orbit and zygoma, and jugal-lacrymal contact (3); not bifurcated, jugal contribution to anteroventral orbit and zygoma, but no jugal-lacrymal contact (4); not bifurcated, jugal contribution only to zygoma and no jugal-lacrymal contact (5); present but no jugal (6). Primitive state based on Vincelestes. Wible et al. (2009) characters 197-201. STEPMATRIX.

  0 1 2 3 4 5 6
0 - 1 2 1 1 2 3
1 1 - 1 2 2 3 4
2 2 1 - 3 1 2 3
3 1 2 3 - 2 3 4
4 1 2 1 2 - 1 2
5 2 3 2 3 1 - 1
6 3 4 3 4 2 1 -

112. Zygomatic arch: stout, without malar foramen (0); delicate, without malar foramen (1) PRIMITIVE; delicate, with malar foramen (2); absent (3). Primitive state based on Eomaia. Modified from Wible et al. (2009) character 202. STEPMATRIX.

  0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 1
3 1 1 1 -

Orbit

113. Palatine reaches infraorbital canal (0); does not reach (1). Primitive state based on Vincelestes. Wible et al. (2009) character 204.

114. Lacrymal contributes to maxillary foramen (0); does not contribute (1). Primitive state based on Vincelestes. Wible et al. (2009) character 205.

115. No groove connecting maxillary and sphenopalatine foramina (0); groove present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 206.

116. Sphenopalatine (orbitonasal) foramen within palatine (0); between palatine and maxilla (1); between palatine, maxilla and frontal (2); within maxilla (3). Primitive state based on Vincelestes. Wible et al. (2009) character 207. STEPMATRIX.

  0 1 2 3
0 - 1 1 2
1 1 - 1 1
2 1 1 - 1
3 2 1 1 -

117. Sphenopalatine (orbitonasal) foramen not proximal to maxillary foramen (0); proximal (1). Primitive state based on Vincelestes. Wible et al. (2009) character 208.

118. Maxilla excluded from medial orbital wall (0); not excluded (1). Primitive state based on Vincelestes. Wible et al. (2009) character 209.

119. No frontal-maxilla contact in medial orbital wall (0); contact (1). Primitive state based on Vincelestes. Wible et al. (2009) character 210.

120. Orbital process of palatine present (0); absent or with thin sliver in ventromedial wall (1). Primitive state based on Vincelestes. Wible et al. (2009) character 211.

121. Ethmoidal foramen between frontal and orbitosphenoid (0); within frontal (1); absent (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 213. UNORDERED.

122. Foramen for frontal diploic vein absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 214.

123. Frontal foramen on cranial roof absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 215.

124. Postorbital process and dorsal process of jugal absent (0); postorbital process weak, dorsal process of jugal absent (1); postorbital process strong, dorsal process of jugal absent (2) PRIMITIVE; postorbital process strong, dorsal process of jugal present (3); postorbital bar composed of postorbital process and dorsal process of jugal (4). Primitive state based on Vincelestes. Wible et al. (2009) characters 216, 218, 219.

125. Optic foramen narrowly separated from sphenorbital fissure (0); broadly separated (1); not visible in lateral view (2). Polarity problematic as optic foramen absent or its state unknown in potential outgroups. Wible et al. (2009) character 221. UNORDERED.

126. Orbitosphenoid expanded anteriorly from optic foramen (or with anterior process) (0); expanded dorsally from optic foramen (or with dorsal process) (1). Primitive state based on Vincelestes. Wible et al. (2009) character 222, with changed state numbering.

127. Suboptic foramen absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 223. N.B., presence in Leptictis shown by Butler (1956) but stated by Novacek (1986), quoting Butler (1956, p. 473) to be absent anterior to the sphenorbital fissure, but not necessarily absent per se (see Butler 1956, figure 7), thus contra Wible et al. (2009). N.B., absence in Erinaceus is shown by Butler (1948, figure 7) contra Wible et al. (2009).

128. Orbitotemporal canal (sinus canal, cranio-orbital foramen) present (0); absent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 224.

Braincase

129. Frontal length on midline more than 50% longer than parietal (0); subequal to slightly shorter than parietal (1) PRIMITIVE; less than half length of parietal (2). Primitive state based on Eomaia. Wible et al. (2009) character 226, with state numbering changed.

130. Fronto-parietal suture: with anterior process of parietal off the midline (0); transverse (1) PRIMITIVE; with anterior process on the midline (2). Primitive state based on Vincelestes. Wible et al. (2009) character 227.

131. Temporal lines (crista frontalis externa) meet on midline to form sagittal crest (0); do not meet (1). Primitive state based on Eomaia. Wible et al. (2009) character 228.

132. Interparietal absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 229.

133. Nuchal crest level with or anterior to foramen magnum (0); posterior to foramen magnum (1). Primitive state based on Eomaia. Wible et al. (2009) character 230.

Mesocranium

134. Choanae as wide as posterior palate (0); narrower (1). Primitive state based on Vincelestes. Wible et al. (2009) character 234.

135. Vomer does not contact pterygoid (0); contacts pterygoid (1). Primitive state based on Vincelestes. Wible et al. (2009) character 235.

136. Pterygoids contact on midline (0); do not contact (1). Primitive state based on Vincelestes. Wible et al. (2009) character 236.

137. Midline crest in basipharyngeal canal present (0); absent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 238, with state numbering reversed.

138. Entopterygoid process ends at anterior basisphenoid (0); approaches ear region (1). Modified from Wible et al. (2009) character 239.

139. Midline rod-shaped eminence on basisphenoid absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 240.

140. Ectopterygoid process of alisphenoid: approaches ear region as long crest (0); ends at anterior basisphenoid as long crest (1) PRIMITIVE; ends at anterior basisphenoid as narrow process (2); approaches ear region as narrow process (3); absent (4). Wible et al. (2009) characters 241-242. STEPMATRIX.

  0 1 2 3
0 - 1 2 1
1 1 - 1 2
2 2 1 - 1
3 1 2 1 -

141. Exit for maxillary nerve (cranial nerve V2) relative to alisphenoid: behind (0); within (1); in front (2). Primitive state based on Vincelestes. Wible et al. (2009) character 244.

142. Foramen ovale in petrosal (anterior lamina) (0); in alisphenoid (1); between alisphenoid and squamosal (2). Primitive state based on Vincelestes. Wible et al. (2009) character 246, with changed states.

143. Alisphenoid canal absent (0); present, posterior opening separate from foramen ovale (1); present, posterior opening in common depression with foramen ovale (2). Primitive state based on Vincelestes. Wible et al. (2009) characters 248-249. UNORDERED.

Basicranium

144. Glenoid fossa on zygoma (0); partly on braincase (1). Primitive state based on Vincelestes. Wible et al. (2009) character 251.

145. Glenoid fossa even dorsoventrally with sphenoid on midline cranial base (0); higher (1). Primitive state based on Vincelestes. Wible et al. (2009) character 253.

146. Jugal contributes to glenoid with articular facet (0); contributes to glenoid without articular facet (1); does not contribute to glenoid (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 254. State 2 for Ptilocercus (Wible, 2011) contra Wible et al. (2009) (0).

147. Postglenoid process present (0); absent (1). Primitive state based on Eomaia. Wible et al. (2009) character 256.

148. Postglenoid foramen within squamosal, behind postglenoid process (0); within squamosal, medial or anterior to postglenoid process or on lateral aspect of braincase (1); behind squamosal, behind postglenoid process (2); absent (3) PRIMITIVE. Primitive state based on Vincelestes. Wible et al. (2009) characters 257-259. UNORDERED.

149. Suprameatal foramen absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 260.

150. Entoglenoid process of squamosal absent (0); present, separated from postglenoid process (1); present, continuous with postglenoid process (2). Primitive state based on Eomaia. Wible et al. (2009) character 261. UNORDERED.

151. Posttympanic crest of squamosal absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 262.

152. Carotid foramen within basisphenoid (0); between basisphenoid and petrosal (1); absent (2). Primitive state based on Vincelestes. Wible et al. (2009) character 263. UNORDERED.

153. Cavum epiptericum floor composition: petrosal (0); petrosal and alisphenoid (1); primarily open as pyriform (piriform) fenestra (2). Primitive state based on Vincelestes. Wible et al. (2009) character 264. UNORDERED.

154. Alisphenoid tympanic process absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 265. Recoded 1 in Ptilocercus as shown present by Cartmill and MacPhee (1980) and Wible (2011), contra Wible et al. (2009), where coded 0.

155. Basisphenoid tympanic process absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 266.

156. Medial flange of petrosal absent (0); present (1). Polarity is equivocal as it is absent in Vincelestes, but present in Prokennalestes. 1 treated here as PRIMITIVE. Wible et al. (2009) character 268.

157. Rostral tympanic process of petrosal on posteromedial aspect of promontorium: absent or low ridge (0); moderate ridge, contributing to posterodorsomedial bulla (1); tall ridge, contributing to ventral bulla (2). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 269. UNORDERED.

158. Course of internal carotid artery (ICA) and of stapedial artery: ICA lateral (transpromontorial), both in sulcus (0); ICA lateral (transpromontorial), both in intratympanic vascular canal (1); medial (perbullar or extrabullar), both in sulcus (2); course indication absent (3). Primitive state based on Vincelestes and Prokennalestes. Modified from Wible et al. (2009) characters 270-271, 273-274 and Bloch et al. (2007) character 87. STEPMATRIX.

  0 1 2 3
0 - 1 1 1
1 1 - 2 2
2 1 2 - 1
4 1 2 1 -

159. Promontorium shape: flat (0); globose (1). Polarity is equivocal as it is flat in Vincelestes, but globose in Prokennalestes. 1 treated here as PRIMITIVE. Wible et al. (2009) character 278.

160. Intratympanic course of facial nerve: open in sulcus (0); open anteriorly, canal posteriorly (1); in canal (2). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 280.

161. Aperture of hiatus Fallopii: in tympanic roof through petrosal (0); at anterior edge of petrosal (1); through cerebral surface, via fenestra semilunaris if present (2); absent (3). Primitive state based on Vincelestes and Prokennalestes. Modified from Wible et al. (2009) character 281. STEPMATRIX.

  0 1 2 3
0 - 1 2 1
1 1 - 1 1
2 2 1 - 1
3 1 1 1 -

162. Prootic canal present (0); absent (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 282.

163. Length of bony shelf lateral to promontorium (lateral trough or tegmen tympani): extended anteriorly as far as promontorium (0); confined posterolaterally (1) PRIMITIVE; prolonged anterior to promontorium (2). Polarity is equivocal as Vincelestes is 0, but Prokennalestes is 1. 1 treated here as PRIMITIVE. Wible et al. (2009) character 285.

164. Width of bony shelf lateral to promontorium (lateral trough or tegmen tympani): uniform (0); expanded anteriorly (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 286.

165. Stapedial canal on bony shelf lateral to promontorium (lateral trough or tegmen tympani): absent (0); present (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 288.

166. Tensor tympani fossa on petrosal: shallow to absent (0); deep circular pit (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 289.

167. Cochlear width less than 20% of skull length (0); greater than 20%. Primitive state based on Vincelestes. Novacek (1987).

168. Epitympanic recess to fossa incudis relative size: subequal (0); epitympanic recess larger (1); no visible depression for epitympanic recess (2). Polarity is equivocal as Vincelestes is 0, but Prokennalestes is 1. 1 treated here as PRIMITIVE. Wible et al. (2009) character 292.

169. Epitympanic recess lateral wall: with extensive contribution to lateral wall by squamosal (0); with small contribution to posterolateral wall by squamosal (1) PRIMITIVE; no squamosal contribution (2). Polarity is equivocal as Vincelestes is 1, but Prokennalestes is 0. 1 treated here as PRIMITIVE. Wible et al. (2009) character 293, with states 0 and 1 reversed.

170. Fossa incudis continuous with epitympanic recess (0); separated from it (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 294.

171. Foramen for ramus superior of stapedial artery: posterior or lateral to fenestra vestibuli (ovalis) (0); anterior to fenestra vestibuli (1); absent (2). Polarity is equivocal as Vincelestes is 0, but Prokennalestes is 1. 1 treated here as PRIMITIVE. Wible et al. (2009) character 298. UNORDERED.

172. Ascending canal (for arteria diploëtica magna): intramural (0); intracranial (1); absent (2). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 299. UNORDERED.

173. Stapedius fossa twice the size of fenestra vestibuli (ovalis) (0); small and shallow (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 300.

174. Cochlear canaliculus not visible in middle ear space (0); visible (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 301.

175. Postpromontorial tympanic sinus: dorsal to cochlear fossula (0); at same level as cochlear fossula (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 302.

176. Fenestra cochleae (rotunda) position to fenestra vestibuli (ovalis): posteromedial (0); posterior (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 303.

177. Posterior septum shielding fenestra cochleae (rotunda) absent (0); present (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 304.

178. Paroccipital process of petrosal: vertical (0); slanted, projecting as flange towards back of promontorium (1); indistinct to absent (2). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 305. UNORDERED.

179. Caudal tympanic process of petrosal: not notched (0); notched (1); absent (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 306. UNORDERED.

180. Crista interfenestralis and caudal tympanic process: not connected by curved ridge (0); connected by curved ridge (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 307.

181. ‘Tympanic process’ of Kielan-Jaworowska: absent (0); present, low, composed of petrosal (1); present, low, composed of petrosal and exoccipital (2); present, high, composed of petrosal (3). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) characters 308-309. STEPMATRIX.

  0 1 2 3
0 - 1 1 2
1 1 - 1 1
2 1 1 - 2
3 2 1 2 -

182. Jugular (posterior lacerate) foramen size relative to fenestra cochleae (rotunda): subequal (0); larger (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 312.

183. Jugular (posterior lacerate) foramen: confluent with opening for inferior petrosal sinus (0); separated from inferior petrosal sinus (1). Primitive state based on Vincelestes. Wible et al. (2009) character 313.

184. Hypoglossal foramen number: two or more (0); one (1); variable (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 314. UNORDERED.

185. Hypoglossal foramen not housed in opening larger than jugular (posterior lacerate) foramen (0); housed in opening larger than jugular foramen (1). Primitive state based on Vincelestes. Wible et al. (2009) character 315.

186. Paracondylar (paroccipital) process of exoccipital: weak or absent (0); prominent, vertical (1); prominent, posteriorly directed (2). Primitive state based on Vincelestes. Wible et al. (2009) character 316. UNORDERED.

187. Ectotympanic: phaneric or visible in ventral view (0); aphaneric or hidden by auditory bulla (1). Wible et al. (2009) character 317.

188. Ectotympanic: ring-like (0); fusiform (1) PRIMITIVE; expanded (2). Wible et al. (2009) character 318, but recoded (0) for Plesiadapis (Bloch and Silcox 2001, figure 7).

189. Anterior crus of ectotympanic: does not broadly contact facet on squamosal (0); does broadly contact facet on squamosal (1). Primitive state based on Vincelestes. Wible et al. (2009) character 319.

190. Elongate ossified external acoustic canal: absent (0); present (1). Primitive state based on ingroup commonality. Wible et al. (2009) character 320.

191. Roof of external acoustic meatus: petrosal (0); petrosal and squamosal (1); squamosal (2). Polarity equivocal, as Vincelestes is 0, Prokennalestes is 1. 0 treated here as PRIMITIVE. Modified from Wible et al. (2009) character 321. UNORDERED.

192. Entotympanic absent (0); present (1). Primitive state based on Vincelestes Wible et al. (2009) character 322.

193. Hyoid arch does not contribute to bulla (0); does contribute (1). Primitive state based on Vincelestes and Prokennalestes. Wible et al. (2009) character 324.

194. Dorsum sellae tall (0); low (1). Primitive state based on Vincelestes. Wible et al. (2009) character 325.

195. Wall separating cavum supracochleare from cavum epiptericum: incomplete, with fenestra semilunaris (0); complete (1). Polarity equivocal, as Vincelestes is 0, Prokennalestes is 1. 0 treated here as PRIMITIVE. Modified from Wible et al. (2009) character 328.

196. Crista petrosa: vestigial or absent (0); a tall, thin crest (1). Primitive state based on Vincelestes. Wible et al. (2009) character 329.

197. Anterior (superior) semicircular canal: forms lateral wall of unconstricted subarcuate fossa aperture (0); forms lateral wall of constricted subarcuate fossa aperture (1); does not form lateral wall of constricted subarcuate fossa aperture (2). Primitive state based on Vincelestes. Wible et al. (2009) characters 330-331. UNORDERED.

198. Posttemporal canal on occiput: large, between petrosal and squamosal (0); small, between petrosal and squamosal (1); small, within petrosal (2); absent (3). Primitive state based on Vincelestes. Wible et al. (2009) characters 333-335. STEPMATRIX.

  0 1 2 3
0 - 1 2 2
1 1 - 1 1
2 2 1 - 1
3 2 1 1 -

Vertebrae

199. Atlantal foramen present (0); absent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 339.

200. Atlas neural hemiarches unfused (0); fused (1). Primitive state based on Vincelestes. Wible et al. (2009) character 340.

201. Atlas neural arch and intercentrum unfused (0); fused (1). Primitive state based on Vincelestes. Wible et al. (2009) character 341.

202. Axis with suture between atlantal and axial parts (0); without (1). Wible et al. (2009) character 342.

203. Axis with extra pair of transverse processes on ventral surface (0); without (1). Primitive state based on Vincelestes. Wible et al. (2009) character 343.

204. Axis prezygapophyses extend ventral to dens (0); linked (1); not linked to dens (2). Primitive state based on Vincelestes. Wible et al. (2009) character 344.

205. Inferior lamellae present on posterior cervical vertebrae (0); absent (1). Primitive state based on Vincelestes. Wible et al. (2009) character 345.

206. Number of thoracic vertebrae 14 or fewer (0); 15 or more (1). Primitive state based on Eomaia. Modified from Wible et al. (2009) character 347.

207. Number of sacral vertebrae 2 (0); 3 (1); 4 or more (2). Primitive state based on Vincelestes. Wible et al. (2009) character 350.

Scapula

208. Infraspinous fossa position with respect to supraspinous fossa: different planes (in part, medial to) (0); coplanar (1). Primitive state based on Vincelestes. Wible et al. (2009) character 352.

209. Suprascapular incisure absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 353.

210. Acromion reaches distal to glenoid articulation (0); is proximal (1). Primitive state based on Vincelestes. Wible et al. (2009) character 354.

211. Metacromion weak or absent (0); a well developed process (1). Primitive state based on Eomaia. Wible et al. (2009) character 355.

Humerus

212. Greater and lesser tuberosities lower than or at same level as head (0); Greater tuberosity higher than head (1). Primitive state based on Vincelestes. Wible et al. (2009) character 356.

213. Proximal third of shaft curved posteriorly (0); ~ straight (1). Primitive state based on Vincelestes.

214. Deltopectoral crest extending less than one third the length from the proximal end (0); between one third and one half (1) PRIMITIVE; more than a half (2). Primitive state based on Vincelestes.

215. Deltopectoral crest laterally displaced (0); median (1). Primitive state based on Vincelestes.

216. Deltopectoral crest strong (0); weak (1). Primitive state based on Vincelestes.

217. Tuberosity for teres major muscle a crestiform or tubercular structure (0); absent or represented only by rugose surface (1). Primitive state based on Vincelestes.

218. Entepicondylar foramen present (0); absent (1). Primitive state based on Vincelestes, Henkelotherium and Juramaia. Wible et al. (2009) character 360).

219. Supinator crest weak and short (0); weak and long (1); strong, short, extending scarcely proximal of the flaring of the medial epicondyle (2); strong extending more proximally (3). Primitive state based on Vincelestes. STEPMATRIX

  0 1 2 3
0 - 1 1 2
1 1 - 2 1
2 1 2 - 1
3 2 1 1 -

220. Medial epicondyle extends from medial trochlear ridge more than shaft width (0); less (1). Primitive state based on Vincelestes.

221. Medial epicondyle projects slightly distomedially (0); medially to slightly proximomedially (1). Primitive state based on Vincelestes and Juramaia.

222. Olecranon fossa very deep (0); moderately deep (1) (PRIMITIVE); shallow (2). Primitive state based on Vincelestes. Modified from Goswami et al. (2011) character 413.

223. Olecranon fossa imperforate (0); perforate (1). Primitive state based on Vincelestes. Reworded from Wible et al. (2009) character 361.

224. Trochlea concave with sharp medial keel, projecting further distally than capitulum (0); convex, without clear lateral crest, projecting farther distally than capitulum (1) PRIMITIVE; convex, without lateral crest, not projecting further distally than capitulum (2); convex, narrow, with lateral crest (3); semicylindrical, wide, with lateral crest (4). Primitive state based on Vincelestes.

225. Capitulum semi-cylindrical, confluent with trochlea (0); semi-spherical, separated from trochlea by intercondylar groove (1) PRIMITIVE; spherical, separated from trochlea by intercondylar groove (2). Primitive state based on Vincelestes.

226. Capitulum with no distinct lateral flange (0); with distinct lateral flange confluent with rest of capitulum (1) PRIMITIVE; lateral flange separated from rest of capitulum by groove (2). Primitive state based on Vincelestes.

227. Dorsoepitrochlear fossa absent (0); present (1). Primitive state based on Vincelestes.

228. Radial fossa present, but coronoid fossa absent (0); both radial and coronoid fossae present (1). Primitive state based on Vincelestes.

Radius

229. Capitular eminence: weak to absent (0); strong (1). Primitive state based on Vincelestes.

230. Ulnar facet proximodistally shallow (0); deep (1). Primitive state based on Vincelestes.

231. Facet for humeral trochlea absent (0); present (1) PRIMITIVE. Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 364.

232. Shaft strongly curved proximally with asymmetrical head (0); weakly curved with asymmetrical head (1); gently curved to straight with symmetrical head (2). Primitive state based on Vincelestes.

233. Bicipital tuberosity weak to absent (0); present, simple, confluent with proximal articulation, without lateral crest (1) PRIMITIVE; confluent with proximal articulation, with lateral crest (2); separate from proximal articulation, without lateral crest (3); separate from proximal articulation, with lateral crest (4). Primitive state based on Vincelestes.STEPMATRIX.

  0 1 2 3 4
0 - 1 1 2 2
1 1 - 1 1 2
2 1 1 - 2 1
3 2 1 2 - 1
4 2 2 1 1 -

234. Ulnar facet strongly rounded (0); gently rounded (1) PRIMITIVE; flat or nearly so (2). Primitive state based on Vincelestes.

235. Radial articulation with carpals: a single fossa (0); two fossae (1). Primitive state based on Vincelestes. Wible et al. (2009) character 367.

Carpals

236. Scaphoid and lunar separate (0); partially fused, retaining surface division between the bones (1); fully fused (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 368.

237. Centrale separate (0); fused or lost (1). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 369.

Pelvis

238. Pubic symphysis extensive (0); narrow (1); separate (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 370.

239. Epipubic bones present (0); absent (1). Primitive state based on Vincelestes and Eomaia. Wible et al. (2009) character 371.

Femur

240. Head spherical, not extending onto neck (0); head spherical, extending onto neck proximally (1); head more cylindrical (2). Primitive state based on Henkelotherium, Eomaia and Purbeck cladotherian femur NHMUK.PV.OR48250.

241. Head extending far round neck distally (0); not extending far round distally (1). Primitive state based on stem cladotherian NHMUK.PV.OR48250.

242. Neck distinct (0); poorly indented proximally (1). Primitive state based on Eomaia and stem cladotherian NHMUK.PV.OR48250.

243. Head and neck project medially (0); anteromedially (1); proximomedially (2); anteroproximomedially (3); UNORDERED. Primitive state based on stem cladotherian NHMUK.PV.OR48250.

244. Teres fovea central (0); posterior (1); absent (2). UNORDERED. Primitive state based on Henkelotherium. Modified from Wible et al. (2009) character 373.

245. Greater trochanter extends proximally less than head (0); to about level of head (1); farther (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 374.

246. Lesser trochanter posteromedial (0); medial (1). Primitive state based on Henkelotherium and stem cladotherian M13126.

247. Lesser trochanter projects as far as head or beyond (0); less than head (1). Primitive state based on Henkelotherium and stem cladotherians NHMUK.PV.OR48250, M13126. Modified from Wible et al. (2009) character 375.

248. Lesser trochanter near head (0); more distal (1). Primitive state based on Vincelestes and stem cladotherian M13126.

249. Third trochanter absent (0); small less than a third of the distance from proximal end (1); small c. one third distance from proximal end (2); small, more than a third distance from proximal end (3); large c. one third distance from proximal end (4). Primitive state based on Vincelestes, Henkelotherium and stem cladotherians NHMUK.PV.OR48250, M13126. Modified from Wible et al. (2009) character 376. STEPMATRIX.

  0 1 2 3 4
0 - 1 2 3 3
1 1 - 1 2 2
2 2 1 - 1 1
3 3 2 1 - 2
4 3 2 1 2 -

250. Trochanteric fossa deep (0); shallow (1); essentially absent (2). Primitive state based on stem cladotherian M13126.

251. Intertrochanteric crest restricted to lateral position (0); reaches or nearly reaches the base of the lesser trochanter (1). Primitive state based on stem cladotherian M13126.

252. Shaft essentially round in cross section (0); anteroposteriorly compressed (1). Primitive state based on Henkelotherium and stem cladotherian M13126.

253. Distal femur similar in size in anteroposterior and mediolateral dimensions (0); longer anteroposteriorly (1). Primitive state based on Vincelestes. Wible et al. (2009) character 378.

254. Patellar groove broad and shallow (0); narrow and elevated (1). Primitive state based on Vincelestes. Wible et al. (2009) character 379.

255. Articulation between femur and fibula (0); no articulation (1). Primitive state based on Eomaia. Wible et al. (2009) character 381, but polarity coding reversed.

Tibia

256. Tibia separate from fibula distally (0); in contact along part of the shaft (1); co-ossified distally (2); co-ossified along much of the distal half (3). Primitive state based on Vincelestes. STEPMATRIX

  0 1 2 3
0 - 1 1 2
1 1 - 2 1
2 1 2 - 1
3 2 1 1 -

257. Shaft smooth (0); with groove distally for flexor digitorum fibularis tendon (1); with groove distally for flexor digitorum tibialis and tibialis posterior tendons (2); with grooves for flexor digitorum fibularis tendon and for flexor digitorum tibialis and tibialis posterior tendons (3). UNORDERED. Primitive state based on Vincelestes.

258. Medial malleolus distally long (i.e., more than half as long as the distal width of the tibia) with articular surface for sustentaculum of calcaneum (0); Medial malleolus distally long (i.e., more than half as long as the distal width of the tibia) without articular surface (1) PRIMITIVE; length about one third of the distal width of the tibia (2); short (3); absent (4). Primitive state based on Vincelestes.

259. Posterior process absent (0); present (1). Primitive state based on Vincelestes.

260. No tibia-fibula fusion proximally (0); fusion (1). Primitive state based on Vincelestes. Wible et al. (2009) character 382.

Astragalus

261. Body broader (measured proximally) than long (measured laterally) (0); approximately equidimensional (1); longer than broad (2). Primitive state based on Ukhaatherium, which, although belonging to the ingroup, is broadly primitive. Hooker (2001) character 1.

262. Lateral trochlear ridge taller than medial one (0); trochlear ridges of approximately equal height (1); medial trochlear ridge taller than lateral one (2). Primitive state based on Ukhaatherium. Hooker (2001) character 2.

263. Neck not distinct from body medially as navicular facet extends as far as body medially (0); free neck, navicular facet extending c. half way up medial side (1); free neck, navicular facet extending less than half way up medial side (2). Part of Hooker 2001, character 15. Polarity based on Vincelestes, which lacks a neck, so the more metatherian-like pattern in zalambdalestids (Szalay and Sargis, 2006) is likely to be primitive.

264. Medial and lateral walls of body (i.e., for articulation with the fibula and the medial malleolus of the tibia) converge dorsally at more than 60 degrees (0); at 60 degrees (1); essentially parallel (2). From Horovitz 2000, character 14 and description of Vincelestes.

265. Lateral tibial facet not excavated (0); gently excavated (1); deeply excavated, U-shaped (2); deeply excavated, V-shaped (3). Primitive state (0) as ‘ancestral therian’, Horovitz (2000). N.B., the deep V-shape of zalambdalestids separates two dorsally-facing facets. These were labelled ‘medial and lateral tibial facets by Szalay and Sargis (2006, figures 16-17), presumably as there is no medially facing facet on the body. However, it seems more likely that a medial tibial facet is missing, along with the medial malleolus of the tibia and that the ‘medial’ and ‘lateral’ facets are together homologous with the lateral facet of other therians. Modified from Horovitz (2000) character 11 and Hooker (2001) character 16.

266. Lateral tibial facet follows the curvature of the trochlear ridges (0); gently indented distally, into which cavity, convexity of tibial facet or anterior process of tibia fits (1); deeply indented distally for anterior process of tibia (2). Primitive state based on Ukhaatherium.

267. No extension of trochlea onto neck (0); extension of medial ridge (1); extension between ridges (2). UNORDERED. Primitive state based on Ukhaatherium. Modified from Hooker (2001) character 3.

268. Proximal foramen of astragalar canal present (0); absent (1). Primitive state based on Vincelestes. Horovitz (2000) character 17; Hooker (2001) character 4. This structure labelled as present in the dermopteran Galeopterus variegatus by Smith et al. (2010, figure 1h) is in fact the dorsal edge of the groove for the flexor digitorum fibularis tendon (personal obs.; Beard, 1993, figure 10.12).

269. Ventral limit of trochlea on proximal surface of body at midpoint (0); more ventral (1). Primitive state based on Vincelestes. Hooker (2001) character 5.

270. Groove for flexor digitorum fibularis tendon shallow (0); deep (1). Primitive state based on ingroup commonality. Hooker (2001) character 6.

271. Medial plantar tuberosity large (0); medium (1) PRIMITIVE; small (2). Modified from Horovitz 2000, character 15; Hooker 2001, character 7. Primitive state based on Eomaia (Luo et al., 2003, figure 3D). Deccanolestes scored incorrectly by Horovitz 2000 as having large state, probably because of distorted cast figured by Prasad and Godinot (1994), see Hooker (2001).

272. Attachment area on lateral body wall for fibuloastragalar ligament proximal (0); central (1). Primitive state based on ingroup commonality. Hooker (2001) character 8.

273. Sustentacular facet extensive, confluent with navicular facet (0); restricted distally to lateral of astragalocalcaneal ligament area, confluent with navicular facet (1); restricted distally to medial of astragalocalcaneal ligament area, confluent with navicular facet (2); extensive mediolaterally, restricted to proximal of astragalocalcaneal ligament area, not confluent with navicular facet (3); subcircular, isolated (4). Primitive state based on Eomaia (Luo et al., 2003, figure 3D). Modified from Hooker (2001) characters 9 (in part) and 10. STEPMATRIX.

  0 1 2 3 4
0 - 1 1 2 2
1 1 - 2 2 1
2 1 2 - 1 2
3 2 2 1 - 1
4 2 1 2 1 -

274. Sustentacular facet strongly convex (0); flat or nearly so (1). Polarity based on Ukhaatherium being primitive. Modified from Hooker (2001) character 9 (in part).

275. Navicular facet not projecting over lateral side of neck (0); projecting over lateral side of neck (1). Primitive state based on Eomaia (Luo et al., 2003, figure 3D).

276. Head irregular or rounded in only one plane (0); evenly convex (1). Primitive state based on Vincelestes. Hooker (2001) character 11.

277. Navicular facet in distal view: nearly circular, equidimensional (0); elongate oval, long axis 1.4-1.5 times short axis, horizontally aligned (1); elongate oval, long axis 1.6 times short axis or more, horizontally aligned (2); elongate oval, long axis 1.4-1.5 times short axis, dipping medially (3); elongate oval, long axis 1.6 times short axis or more, dipping medially (4). Primitive state based on Ukhaatherium. Modified from Hooker (2001) characters 12, 13, 14. STEPMATRIX.

  0 1 2 3 4
0 - 1 2 1 2
1 1 - 1 1 2
2 2 1 - 2 1
3 1 1 2 - 1
4 2 2 1 1 -

278. Cotylar fossa absent (0); present (1). Primitive state based on Vincelestes. Wible et al. (2009) character 388.

279. Head lacks cuboid facet (0); bears cuboid facet (1). Primitive state based on Vincelestes. Wible et al. (2009) character 394.

Calcaneum

280. Tuber curved/twisted ventromedially and dorsoventrally compressed (0); straight, cylindrical to 1.25 times higher dorsoventrally than wide mediolaterally (1); straight, 1.3-1.6 times higher than wide (2); 1.7 or more times higher than wide (3). Primitive state based on Vincelestes and supported by more primitive mammals, fide Horovitz (2000), although Vincelestes is only medially deflected and not dorsoventrally compressed (Rougier, 1993). Modified from Horovitz (2000) characters 2 and 3; Hooker (2001) character 18.

281. Lateral edge of tuber continuous with peroneal process (0); no lateral edge (1). Polarity based on Vincelestes. Modified from Horovitz (2000) character 10.

282. Groove on lateral surface of tuber for peroneus brevis tendon absent (0); present (1). Primitive state based on ingroup commonality. Hooker (2001) character 24, from Szalay and Lucas (1996).

283. Ectal facet medial of midline passing through tuber (0); essentially median or more lateral (1). Primitive state based on Vincelestes.

284. Ectal facet orientation: distal end more medial than proximal end (0); proximodistal (1); proximomedial-distolateral 5-15 degrees (2); proximomedial-distolateral 25-30 degrees (3); proximomedial-distolateral 40-60 degrees (4); mediolateral with or without wrap around (5). Primitive state based on Vincelestes, backed up by Morganucodon, but no eutherian has primitive state. Modified from Horovitz (2000) character 7; Hooker (2001) character 20.

285. Ectal facet strongly evenly convex (0); gently evenly convex (1); helical (2); unevenly convex (3); sigmoidal (4). UNORDERED. Primitive state based on Vincelestes. Modified from Hooker (2001) character 19.

286. Ectal facet nearer to distal than proximal end (i.e., tuber relatively long) (0); at about midpoint along length (1) PRIMITIVE; nearer to proximal than distal end (i.e., tuber short) (2). Primitive state based on Vincelestes. Polarity changed from Hooker (2001) character 17.

287. Fibular facet present (0); absent (1). Primitive state based on Vincelestes. Horovitz (2000) character 4; Hooker (2001) character 21.

288. Peroneal process fully distal, its distal edge reaching level of cuboid facet (0); apex of peroneal process positioned at 10-15% of calcaneal length from the distal extremity (1); 17-25% from distal extremity (2); 35-55% from distal extremity (3). Primitive state based on Vincelestes. Horovitz (2000) character 5; modified from Hooker (2001) character 26.

289. Peroneal process lateral projection as percentage of calcaneal length: 15-30% (0); 9-13% (1); 5-7% (2); no lateral projection (3). Primitive state based on Vincelestes. Polarity reversed from Hooker (2001) character 27.

290. Peroneal process restricted distally (0); extending for 15-20% of calcaneal length (1); 21-27% (2); 32-46% (3) PRIMITIVE. Primitive state based on Eomaia. Hooker (2001) character 28.

291. Ectal and sustentacular facets do not overlap on long axis (0); do overlap (1) PRIMITIVE. Primitive state based on Vincelestes. Polarity changed from Hooker (2001) character 22; Wible et al. (2009) character 399.

292. Sustentacular facet reaches edge of cuboid facet (0); divided into a sustentacular facet sensu stricto and a distal sustentacular facet (1); single, but distinctly separated from edge of cuboid facet (2). UNORDERED. Primitive state based on Vincelestes. Modified from Hooker (2001) character 23, see Horovitz (2000). N.B. the 0 state is often treated as derived and an archontan synapomorphy, e.g., Hooker (2001). This change in polarity has important implications for phylogeny. The character is represented (and is interdependent) in the astragalus by a level of connection between the sustentacular facet and the navicular facet and is not treated as a separate character. However, the exact nature of this connection is a distinct character (q.v.).

293. Navicular facet absent (0); present (1). Primitive state based on Vincelestes. Hooker (2001) character 25.

294. Cuboid facet faces distoventrally (0); faces distomedially at c.55-60 degrees to long axis of calcaneum (1); at c.70-75 degrees to long axis of calcaneum (2); faces essentially distally (3). STEPMATRIX. Primitive state based on Vincelestes. Horovitz (2000) character 8; modified from Hooker (2001) character 31.

  0 1 2 3
0 - 1 1 1
1 1 - 1 2
2 1 1 - 1
3 1 2 1 -

295. Cuboid facet dorsoventrally concave (0); shallowly evenly concave (1); deeply evenly concave (2) PRIMITIVE; sellar (3). UNORDERED. Primitive state based on Vincelestes. Polarity and ordering changed from Hooker (2001) character 29.

296. Cuboid facet deeper dorsoventrally than wide mediolaterally (0); subequal (1) PRIMITIVE; wider mediolaterally than deep dorsoventrally (2). Primitive state based on Vincelestes.Wible et al. (2009) character 407.

297. Cuboid facet without plantar pit (0); with plantar pit allowing rotation (1); with grooved plantar pit inhibiting rotation (2). UNORDERED. Primitive state based on Vincelestes. Hooker (2001) character 30.

298. Plantar groove present (0); absent (1). Primitive state based on Vincelestes and Eomaia (Luo et al. 2003, figure 3J). Modified from Horovitz (2000) character 6.

299. Distal plantar tubercle absent (0); present distal, medial (1); present distal central (2); proximal of distal edge, medial (3); proximal of distal edge, central (4). UNORDERED. Primitive state based on Vincelestes. Modified from Hooker (2001) characters 33-34.

300. Groove for tendon of flexor digitorum fibularis absent (0); shallow (1); deep (2). Primitive state based on Vincelestes. Modified from Wible et al. (2009) character 408.

301. Calcar absent (0); present (1). Primitive state based on ingroup commonality.

Navicular

302. Ectocuneiform and mesocuneiform facets not distinguished (0); the two facets of equal length (1); ectocuneiform facet longer dorsoventrally than mesocuneiform facet (2); mesocuneiform facet longer dorsoventrally than ectocuneiform facet (3). STEPMATRIX. Primitive state based on simplest form and 0 state in Zalambdalestidae).

  0 1 2 3
0 - 1 2 2
1 1 - 1 1
2 2 1 - 2
3 2 1 2 -

303. Plantar process scarcely or not projecting distally (0); projecting distinctly distally (1). Primitive state based on Eomaia (Luo et al., 2003).

304. Tuber tibialis absent (0); short (1); long (2). Primitive state based on Eomaia (Luo et al., 2003).

Cuboid

305. Edge of calcaneal facet overhangs dorsal wall, making it concave (0); not overhanging, dorsal wall essentially straight (1). Primitive state based on Asioryctes.

306. Plantar process strong (Kielan-Jaworowska, 1979, plate 9, figures 1a,1b) (0); weak (1). Primitive state based on Asioryctes (Kielan-Jaworowska, 1977, plate 18, figure 2b).

Metapodials

307. Tubercles for insertion of m. extensor carpi radialis on metacarpals absent (0); present on M/C III (1); present on M/C II-III (2). UNORDERED. Primitive state based on Henkelotherium. See Rose (1999), Rose and Lucas (2000), Rose and Koenigswald (2005).

Phalanges

308. Attachment on phalanx 1 for annular ligament absent or weak (0); present as tubercles (1) PRIMITIVE; present as long ridges (2). Primitive state based on Henkelotherium and Eomaia.

309. Manual phalanges relatively short (0); those of digits II-V greatly elongate (1). Primitive state based on ingroup commonality.

310. Unguals well developed (0); reduced to absent on manual digits III-V (1). Primitive state based on ingroup commonality.

 

FIGURE 1. Scanning electron micrographs of right humerus (reversed) type 1 (Scraeva hatherwoodensis), M61009, Mammal Bed Hordle, Hampshire, in anterior (1), posterior (2) and medial (3) views. Scale bar equals 1 mm.

figure1 

FIGURE 2. Scanning electron micrographs of humeri of Nyctitheriidae. 1, right humerus (reversed) type 1 (Scraeva hatherwoodensis), M61009; 2, distal left humerus type 2 (Euronyctia grisollensis), M60810; 3, distal right humerus (reversed) type 4 (Paradoxonycteris sp. 1?), M61422; 4, distal right humerus (reversed) type 3 (Cryptotopos woodi), M95898. Views are proximal (1), anterior (2.1, 3, 4.4), distal (2.2, 4.2), posterior (2.3, 4.5), medial (2.4, 4.3) and lateral (2.5, 4.1). 3 is from the How Ledge Limestone, Headon Hill, Isle of Wight, the rest are from the Mammal Bed, Hordle, Hampshire. Scale bar equals 1 mm.

 figure2

FIGURE 3. Scanning electron micrographs of right radii (reversed) of Nyctitheriidae. 1, Nearly complete radius lacking end, M95899; 2, proximal fragment, M60817; 3, proximal fragment, HZM.324.28063. 1-2 are attributed to Scraeva hatherwoodensis, 3 to Paradoxonycteris sp. 1? The extent of the ulnar facet (uf) in 3.1 is shown by arrows. Views are: anterior (1.1, 2.1, 3.2), lateral (1.2, 2.2, 3.3), posterior (2.3, 3.4), medial (2.4, 3.5) and proximal (3.1). 1-2 are from the Mammal Bed, Hordle, Hampshire; 3 is from the Rodent Bed, Hordle. Scale bar equals 1 mm.

 figure3

FIGURE 4. Scanning electron micrographs of right femora (reversed) of Nyctitheriidae. 1, nearly complete femur type 1 (Scraeva hatherwoodensis ?), lacking distal end, M60701; 2, proximal fragment of femur type 2 (Cryptotopos woodi ?), M95900; 3, left metacarpal II, HZM.266.27528. Views are: anterior (1.1, 3.1), posterior (1.2, 2) and proximal (3.2). All from the Mammal Bed, Hordle, Hampshire. Scale equals 1 mm.

figure4 

FIGURE 5. Scanning electron micrographs of right femora (reversed) of Nyctitheriidae. 1, nearly complete femur type 1 (Scraeva hatherwoodensis ?), lacking distal end, M60701; 2, proximal fragment of femur type 2 (Cryptotopos woodi ?), M95900. Views are: medial (1.1), proximal (1.2, 2) and lateral (1.3). Both from the Mammal Bed, Hordle, Hampshire. Scale equals 1 mm.

 figure5

FIGURE 6. Scanning electron micrographs of distal tibiae, shown as from left side, of Nyctitheriidae. 1.1, right tibia (reversed) type 1 (Cryptotopos woodi), M60823; 1.2, left tibia type 1 (Cryptotopos woodi), M60822; 2, left tibia type 2 (Scraeva hatherwoodensis), M61087; 3, right tibia (reversed) type 3 (Paradoxonycteris sp. 1), M95949; 4, left tibia type 4 (Euronyctia grisollensis), M95951. Views are: distal (1.1, 2.1, 3.1, 4.1) and posterior (1.2, 2.2, 3.2, 4.2). 1-2 from the Mammal Bed, Hordle, Hampshire; 3-4 from the How Ledge Limestone, Headon Hill, Isle of Wight. Scale equals 1 mm.

 figure6

FIGURE 7. Scanning electron micrographs of right distal tibia (reversed) type 1 (Cryptotopos woodi), M95901, Mammal Bed, Hordle, Hampshire. Views are: anterior (1), medial (2) and lateral (3). Scale equals 1 mm.

figure7 

FIGURE 8. Scanning electron micrographs of right distal tibia (reversed) type 3 (Paradoxonycteris sp. 1), M95949, How Ledge Limestone, Headon Hill, Isle of Wight. Views are: anterior (1), medial (2) and lateral (3). Scale equals 1 mm.

 figure8

FIGURE 9. Scanning electron micrographs of left astragali of Nyctitheriidae. 1, type 2 (Paradoxonycteris sp. 1), M95952; 2, type 3 (Scraeva hatherwoodensis), M60101; 3, type 4 (Euronyctia grisollensis), HZM.108.25595. Views are: dorsal (1.1, 2.1, 3.1), ventral (1.4, 2.2, 3.2), lateral (1.2), medial (1.3), proximal (1.5) and distal (1.6). 3 is from the Rodent Bed, Hordle, Hampshire; the others from the Mammal Bed, Hordle. Scale equals 1 mm.

 figure9

FIGURE 10. Logarithmic linear regressions of tooth versus astragalar and calcaneal dimensions in millimetres, following the methodology of Cifelli (1983) and Coillot et al. (2013). 1, M 2 area versus astragalar length of a range of mammals of known tooth and tarsal association; measurements of the numbered taxa are from Table 2; regression slope 1.4264, intercept -0.18646. 2, M 2 area versus calcaneal length as for 1; regression slope1.4686, intercept -0.52107. 3, M 2 area of named nyctitheres versus lengths of astragalar (A) and calcaneal (C) numbered types, using the regression lines from 10.1 and 10.2; M 2 areas of Saturninia gracilis and Euronyctia grisollensis (How Ledge Limestone, Headon Hill) are nearly identical; those of Cryptotopos woodi and Paradoxonycteris sp. 1, although similar, are from the Mammal Bed, Hordle, and How Ledge Limestone, Headon Hill, respectively, removing doubt of attribution of type1 and 2 tarsals from the same locations; calcaneal plots are closer to the regression line than are those of the astragali. 4, M 1 distal width versus lengths of astragalar (A) and calcaneal (C) numbered types; here the substantial difference in M 1 distal width between E. grisollensis and S. gracilis supports the referral of calcaneum type 5 to the latter; Scraeva hatherwoodensis is from the Mammal Bed, Hordle. Measurements in 10.3 and 10.4 are derived from Table 3.

 figure10

FIGURE 11. Scanning electron micrographs of calcanea, shown as left, of Nyctitheriidae. 1, left calcaneum type 2 (Paradoxonycteris sp. 1), M95960; 2, right calcaneum (reversed) type 3 (Scraeva hatherwoodensis), M95911; 3, left calcaneum type 4 (Euronyctia grisollensis), M95959; 4, left calcaneum type 5 (Saturninia gracilis ?), M95965. Views are: dorsal (1.1, 2.1, 3.1, 4.1), ventral (1.2, 2.2, 3.2, 4.2) and lateral (1.3, 2.3, 3.3, 4.3) . 1-2 are from the Mammal Bed, Hordle, Hampshire; 3-4 are from the How Ledge Limestone, Headon Hill, Isle of Wight. Scale equals 1 mm.

figure11 

FIGURE 12. Scanning electron micrographs of calcanea, shown as left, of Nyctitheriidae. 1, left calcaneum type 2 (Paradoxonycteris sp. 1), M95960; 2, right calcaneum (reversed) type 3 (Scraeva hatherwoodensis), M95911; 3, left calcaneum type 4 (Euronyctia grisollensis), M95959; 4, left calcaneum type 5 (Saturninia gracilis ?), M95965. Views are: medial (1.1, 2.1, 3.1, 4.1) and distal (1.2, 2.2, 3.2, 4.2) . 1-2 are from the Mammal Bed, Hordle, Hampshire; 3-4 are from the How Ledge Limestone, Headon Hill, Isle of Wight. Scale equals 1 mm.

figure12 

FIGURE 13. Scanning electron micrographs of naviculars, cuboids and ectocuneiform, shown as left, of Nyctitheriidae. 1, left navicular type 1 (Cryptotopos woodi), HZM.204.27205; 2, right navicular (reversed) type 2 ( Paradoxonycteris sp.1?), M95967; 3, right cuboid (reversed) type 1 (Cryptotopos woodi), M60970; 4, right cuboid (reversed) type 2 (Scraeva hatherwoodensis), M95918; 5, right cuboid (reversed) type 3 (Euronyctia grisollensis), M95969; 6, left ectocuneiform (Scraeva hatherwoodensis ?), M95919. Views are: dorsal (1.1, 3.1, 4, 5.1, 6.1), distal (1.2, 2.1, 3.5, 6.5), proximal (1.3, 2.2, 3.6, 5.4, 6.4), ventral (1.4, 3.3, 5.3), medial (1.5, 3.4, 6.3), lateral (1.6, 2.3, 3.2, 5.2, 6.2) . 1, 3, 4, 6 are from the Mammal Bed, Hordle, Hampshire; 2, 5 are from the How Ledge Limestone, Headon Hill, Isle of Wight. Scale equals 1 mm.

figure13 

FIGURE 14. Scanning electron micrographs of composite reconstructed right ankle region (reversed), inverted at astragalo-calcaneal joint, of Cryptotopos woodi (type 1), Mammal Bed, Hordle, Hampshire. Distal tibia, M95901; astragalus, M95905; calcaneum, M95910; navicular, M95913; cuboid, M60970. Tibia is omitted from 6-7. Ankle dorsiflexed (1, 4), plantarflexed (2, 3, 5) . Views are: medial (1-2), posterior (3), anterior (4-5), dorsal (6), ventral (7) . Scale equals 2 mm.

 figure14

FIGURE 15. Scanning electron micrographs of metatarsals and phalanges of Nyctitheriidae. 1, right metatarsal IV (reversed), M95976; 2, left metatarsal III, M95970; 3, right metatarsal II (reversed), M95975; 4-5, 7-8, first phalanges; 6, second phalanx. 5 is manual?, 4, 7-8 are pedal?. 4, M60843; 5, M61114; 6, M60989; 7, HZM.588.30474; 8, M60790. Views are: proximal (1.1, 2.1, 3.1, 4.5, 5.3, 6.1, 8.1), distal (1.2, 4.4), dorsal (1.3, 2.2, 3.2, 4.1, 5.1, 6.2, 7.1, 8.2), medial (1.4), ventral (4.3, 6.4) and side (4.2, 5.2, 6.3, 7.2, 8.3) . 2, Paradoxonycteris sp. 1?; 4-6, Cryptotopos woodi ?; 7, Euronyctia grisollensis or Saturninia gracilis ; 8, Scraeva hatherwoodensis ? 1, lower Hamstead Member, Bouldnor, Isle of Wight; 2, How Ledge Limestone, Headon Hill, Isle of Wight; 3, Fishbourne Member, Woodside, Isle of Wight; 7, Rodent Bed, Hordle, Hampshire; 4-6, 8, Mammal Bed, Hordle. Scale equals 1 mm.

 figure15

FIGURE 16. Scattergram of length against proximal width of first phalanges of Nyctitheriidae.

 figure16

FIGURE 17. Strict consensus of four maximum parsimony trees generated by PAUP 4.0b10 from the character-taxon matrix in Appendix 2. Node numbers are Bremer decay indices. Groups shown as monophyletic are labelled. Abbreviations: Anagal, Anagalida; Carniv, Carnivora; Chiropt, Chiroptera; Dermopt, Dermoptera; Erin, Erinaceidae; Lept, Leptictidae; Lipo, Lipotyphla; Macro, Macroscelidea; Pholid, Pholidota; Plag, Plagiomenidae; Prim, Primates; Scand, Scandentia;

 figure17

FIGURE 18. One of four maximum parsimony trees of 2704 steps generated by PAUP 4.0b10 from the character-taxon matrix in Appendix 2. Character states at numbered nodes are listed in Appendix 3.

figure18

 

 

TABLE 1. Occurrences of nyctitheriid and equivalent-sized non-nyctitheriid taxa, mostly known dentally, in the levels and localities from which postcranials are described herein. The Hamstead Member (earliest Oligocene) belongs to the Bouldnor Formation, the rest to the Headon Hill Formation (Late Eocene). The levels at Hordle and Headon Hill are in the Totland Bay Member. These records update Hooker (1987, 2010) and Hooker et al. (2005), where details of the stratigraphy can be found. 1, C. beata Crochet, 1974, previously recorded, is indistinguishable from C. woodi . 2, Based tentatively only on an ulna.

 

Mammal Bed, Hordle

Rodent Bed, Hordle

How Ledge Limestone, Headon Hill

Fishbourne Member, Woodside

L. Hamstead Member, Bouldnor

Nyctitheriidae

 

 

 

 

 

Cryptotopos woodi 1

X

X

X

 

 

Scraeva hatherwoodensis

X

 

 

X

 

Paradoxonycteris tobieni

 

 

 

 

X

Paradoxonycteris sp. 1

 

X

X

 

 

Paradoxonycteris sp. 2

 

 

 

 

X

Amphidozotherium spp.

 

 

 

X

X

Euronyctia grisollensis

X

 

X

 

 

Saturninia gracilis

X

X

X

 

X

Other nyctithere-sized taxa

 

 

 

 

 

Suevosciurus bosmae

X

X

X

 

 

Treposciurus gardneri

X

 

X

 

 

Glamys priscus

X

X

X

 

 

G. fordi

 

 

 

 

X

Gliravus daamsi and aff.

X

 

X

 

 

Bransatoglis bahloi

 

 

X

 

 

B. planus

 

 

 

 

X

Miniglis minor

 

X

X

 

 

Eotalpa anglica

X

 

X

 

X2

Heterohyus nanus

 

 

X

 

 

Pseudoloris parvulus

X

X

X

 

 

Amphiperatherium spp.

X

X

X

X

X

Peratherium cuvieri

 

 

X

 

 

TABLE 2. Tooth and tarsal measurements in millimetres of taxa numbered in Figure 10.1 and 10.2. All are means or single measurements. Sorex minutus , Ptilocercus lowii , Tupaia , Cynocephalus variegatus and Echinosorex gymnura are modern. Numbered sources: 1, Coillot et al. (2013); 2, Tobien (1962), Storch (1993); 3, Kondrashov and Lucas (2012).

 

Length X width
M 2

Length Astragalus

Length Calcaneum

Source

1. Arctocyon primaevus

144.64

28.5

52.5

1

2. Arfia shoshoniensis

47.24

19.95

34.95

1

3. Cynocephalus variegatus

11.75

12.6

15.5

herein

4. Echinosorex gymnura

22.46

9.28

17.14

herein

5. Gazinocyon vulpeculus

39.4

20.33

32.9

1

6. Macrocranion tenerum

3.52

3.14

6.71

2

7. Miacis exiguus

12.76

14.2

-

1

8. Notoparamys costilloi

20.52

20

28.4

1

9. Plesiadapis cookei

35.14

13.02

18.8

1

10. Plesiadapis rex

22.3

7.94

-

1

11. Prototomus martis

34.78

14.1

-

1

12. Ptilocercus lowii

5.15

4.35

5.15

herein

13. Rhombomylus turpanensis

16.85

8.38

13.07

1

14. Sorex minutus

0.77

1.27

2.18

herein

15. Tetraclaenodon puercensis

54.27

16.4

30.4

3

16. Tupaia sp.

7.54

5.9

8.15

herein

TABLE 3. Tooth (named) and tarsal (numbered types) measurements in millimetres of nyctitheriid taxa used in Figure 10.3 and 10.4. All are means or single measurements.

 

Length X width
M 2

Distal width
M 1

Type

Length
Astragalus

Length Calcaneum

Cryptotopos woodi

2.36

2.46

1

2.7

4.09

Paradoxonycteris sp. 1

2.22

2.48

2

2.41

3.51

Scraeva hatherwoodensis

1.71

2.15

3

2.04

2.97

Euronyctia grisollensis

1.09

1.54

4

1.53

2.24

Saturninia gracilis

1.07

1.73

5

-

2.65

 

APPENDIX FIGURE 1. Illustrations of left P 5 in crown view for character 20 states 0-8 and their parsimonious transformations.

figure19

APPENDIX FIGURE 2, Illustrations of left preultimate upper molars in crown view for character 47 states 0-9 and their parsimonious transformations.
Note that the zero state is primitive unless otherwise stated. Multistate characters are ordered unless stated to be unordered or a stepmatrix.

 figure20

 

portraitJerry J. Hooker
Department of Earth Sciences
Natural History Museum
Cromwell Road
London SW7 5BD
UK
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Jerry Hooker is Scientific Associate in the Department of Earth Sciences, Natural History Museum, London, following retirement from a long career at this institution. He graduated at London University and gained his PhD, entitled ‘Mammals from the Bartonian (Middle/Late Eocene) of the Hampshire Basin, southern England’, from University College London, both while employed. He has published monographs and numerous papers on mammals, mainly from the Paleogene, but also a few from the Mesozoic. He applies his studies to phylogeny, biostratigraphy and palaeoenvironmental reconstruction. He has an active programme of field collecting in the UK.