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Volume 27.1
January–April 2024
Full table of contents
ISSN: 1094-8074, web version;
1935-3952, print version
Recent Research Articles
See all articles in 27.1 January-April 2024
See all articles in 26.3 September-December 2023
See all articles in 26.2 May-August 2023
See all articles in 26.1 January-April 2023
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
Ukhaatherium
Maelestes
Asioryctes
Zhelestidae
Zalambdalestidae
Cimolestes magnus
Batodon
Gypsonictops
Deccanolestes
Afrodon/Bustylus
Fordonia
Leptictidium
Prodiacodon/Palaeictops
Leptictis
Aphronorus
Coriphagus
Palaeosinopa
Escavadodon
Miacis/Vulpavus
Viverravus
Protungulatum
Loxolophus
Arctocyon
Pseudictops
Gomphos
Rhombomylus
Macrocranion
Apheliscus
Haplomylus
Paschatherium
Rhynchocyon
Soricidae
Echinosorex
Erinaceus
Onychonycteris
Icaronycteris
Rhinolophus
Leptacodon
Cryptotopos
Tupaia
Ptilocercus
Mixodectes
Eudaemonema
Elpidophorus
Plagiomene
Cynocephalus
Purgatorius
Micromomyidae
Ignacius
Plesiadapis
Cantius/Notharctus
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
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
APPENDIX FIGURE 1. Illustrations of left P 5 in crown view for character 20 states 0-8 and their parsimonious transformations.
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.
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.
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 Appendix 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.
47. (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.
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.
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.
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).
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
FIGURE 16. Scattergram of length against proximal width of first phalanges of Nyctitheriidae.
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;
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.
Jerry J. Hooker
Department of Earth Sciences
Natural History Museum
Cromwell Road
London SW7 5BD
UK
j.hooker@nhm.ac.uk
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.
New postcranial bones of the extinct mammalian family Nyctitheriidae (Paleogene, UK): primitive euarchontans with scansorial locomotion
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
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Review: The Princeton Field Guide to Mesozoic Sea Reptiles
The Princeton Field Guide to Mesozoic Sea Reptiles
Article number: 26.1.1R
April 2023