APPENDIX 1

Phylogenetic Analysis of Microleo attenboroughi: Results and Description of Characters

The phylogenetic relationships of Microleo attenboroughi within the Thylacoleonidae were assessed as well as the relationships of thylacoleonids within the Suborder Vombatiformes, using a branch-and-bound search in PAUP * 4.0b10 (Swofford, 2002) based on a modified data matrix of Black et al. (2012) employing a total of 93 characters. Thirty characters were designated ordered. All characters were equally weighted and parsimony-informative. Five phascolarctid taxa in that matrix were not included in this analysis because many had high levels of missing data and the phascolarctids that were included were deemed informative for that taxon. The Oligo-Miocene peramelid Galadi speciosus and dasyurid Barinya wangala were used as outgroups. Within the thylacoleonid ingroup taxa, Priscileo pitikantensis, Wakaleo alcootaensis and Thylacoleo hilli have relatively high levels of missing data because they are only known from relatively incomplete specimens. The dentary specimen of Thylacoleo sp. cf. T. hilli (AM F63584) was used to code characters for the lower dentition of T. hilli on the basis of the assumption that it is likely to be representative of that species. Sixty five of the 71 characters of the original data matrix (Black et al., 2012) analysis were utilised. Six characters were removed because they either related to phascolarctid taxa not included in the analysis or were uninformative. One character relating to P3 morphology (ch.14) was modified to incorporate states in newly included taxa. A heuristic search was performed using branch and bound analysis. New characters relating to dental morphology (ch. 5, 6, 7, 8, 9, 10, 11, 12, 13, 15, 17 18, 27, 28, 30, 38, 39, 40, 41, 42, 43, 44, 48, 50, 53, 57, 58, 59) were included in the data matrix. Bootstrap values were calculated using 1000 replicates.

Results

Our phylogenetic analysis recovered nine most-parsimonious trees (see consensus tree in Supplementary Figure 2: tree length = 272 steps, consistency index = 0.580, rescaled consistency index = 0.435, retention index = 0.750) in all of which Microleo attenboroughi is recovered as the sister taxon to all other known thylacoleonid taxa. Unequivocal thylacoleonid apomorphies (node 10) include bunodont molars and features of P3, including its length relative to M1 length (> 1.2), a longitudinal blade that has a long v-shaped (notched) occlusal edge and lack of a basal posterior cingulum. Autapomorphies distinguishing Microleo from all other thylacoleonids are features of the P3; its fusiform shape, medial cuspule, a posterolingual crest and steep posterobuccal crest at the rear of the crown. Bootstrap support was relatively strong for the thylacoleonid clade (91%) and for the Priscileo - Wakaleo - Thylacoleo clade (70%; see Supplementary Figure 2). Synapomorphies for the Priscileo - Wakaleo - Thylacoleo clade are: a posterior longitudinal blade on P3 that is gently-angled and M2 with confluent postprotocrista and premetaconulecrista. Priscileo roskellyae is recovered as the sister taxon to P. pitikantensis which in turn is recovered as the sister taxon to a Wakaleo/Thylacoleo clade. Unequivocal synapomorphies of the Wakaleo - Thylacoleo clade are increased P3 length relative to cheektooth row (> 0.4) and loss of M4/m4. Bootstrap support for this clade was moderate (83%). Species of Wakaleo form a clade (bootstrap support 62%) but their relationships are unresolved probably as a result of the limited data for both W. oldfieldi and W. alcootaensis. Synapomorphies for the Wakaleo clade include; reduced number of upper and lower premolars, development of an anterolingual cuspule and a moderately-developed posterobuccal crest on P3. Monophyly for species of Thylacoleo was well-supported (bootstrap frequency = 97%) and based on synapomorphies of P3 including: width greater anteriorly than posteriorly, a convex posterior longitudinal blade, a posterolingual crest, a significant increase in length, and reduction of the number of lower incisors to one.

Vombatiform Relationships. The strict consensus tree (Supplementary Figure 2) places thylacoleonids as the sister taxon to all other vombatiform families. This phylogeny differs from previous phylogenies (e.g., Aplin and Archer, 1987; Marshall et al., 1990; Gillespie, 2007; Black et al., 2012) which place thylacoleonids within the vombatomorphian clade. In our analysis, synapomorphies that united phascolarctomorphians and vombatomorphian families (excluding thylacoleonids, node 2) include presence of a single upper and lower premolar, presence of a lingual cusp on P3, a rectangular M2, lower incisors reduced to one, mandibular symphysis that extends to below m1 and a diastema that is < 30% the length of the horizontal ramus. Bootstrap support for this clade was weak (59%). Black et al. (2012) found phascolarctids were united by characters of the lower molars and were the sister group to all other vombatiform families. Bootstrap support for the phascolarctomorphian clade was strong (100%) but was weaker (63%) for the vombatomorphian clade (excluding thylacoleonids). With the exception of thylacoleonids and Marada arcanum, the relationships of all the other vombatomorphian families agree with those presented by Black et al. (2012). Our analysis was unable to resolve the position of Marada with respect to vombatids, diprotodontids and palorchestids.In previous analyses (Aplin and Archer, 1987; Black et al., 2012) thylacoleonids have been united with other vombatomorphian families predominantly on the basis of cranial characters and their repositioning in this analysis may be due to the large number of dental characters in the data matrix. The results of this analysis suggest that thylacoleonids possibly form a clade within Vombatiformes that has similar taxonomic status to phascolarctomorphians. However, in contrast to thylacoleonids, phascolarctomorphians show a greater diversity at the generic level and, at present, stronger support for the clade is required before erecting a possible new thylacoleonid infraorder.

Character Description

Description of 93 characters used in the phylogenetic analysis. Reference is given to characters used in previous analyses and whether they are modified.* Indicates ordered characters. Abbreviations for references: A84b, Archer (1984b); AD82, Archer and Dawson (1982); B07, Black (2007); B12, Black et al. (2012); BA97a, Black and Archer (1997a); BA97b, Black and Archer (1997b); G07, Gillespie (2007); H93, Hand et al. (1993); HSV03, Horovitz and Sanchez-Villagra (2003); HW82, Hope and Wilkinson (1982); L02, Luo et al. (2002); L04, Louys (2004); M86, Murray (1986); M90, Marshall et al. (1990); M94, Myers (1994); MA97, Myers and Archer (1997); R87, Reig et al. (1987), R98, Rougier et al. (1998); S97, Springer et al. (1997); SVW02, Sánchez-Villagra and Wible, (2002); T10, Travouillon et al. (2010); WD 87, Woodburne et al. (1987); W98, Wroe et al. (1998); W00, Wroe et al. (2000).

1.* Number of upper incisors: five (0); four (1); three (2); two (3); one (4). (S97, R98, W00, HSV03, B12)
2. Size of I1/ relative to I2-3/: equal/similar length (0); large, elongate (1). (B12)
3. Upper canine: present (0); absent/vestigial (1). (Modified: S97, HSV03, B12)
4.* Number of upper premolars: three (0); two (1); one (2). (B12, AD82, B12)
5. Number of primary cusps on longitudinal blade: one (0); two (1); three (2); four (3).
6. P3/ occlusal shape: wider posteriorly than anteriorly (0); anterior and posterior widths similar, broadest centrally (1); wider anteriorly than posteriorly (2). (G07)
7.* Absolute size of P3: small (< 5mm) (0); moderate (5-10mm); (1); large (10-20mm) (2); very large (20-30mm) (3); extremely large (>30mm) (4).
8.* Size of P/p3 relative to cheektooth row length: very short (0); short (1); long (2); very long (3). (G07)
9. Curvature of the longitudinal blade of P3: straight (0); buccally convex (1); lingually convex (2).
10.* Development of the anterolingual crest on P3: absent (0); weak, long crest (1); moderate, long crest (2); cuspule (3). (G07)
11. Slope of the posterior end of P3 longitudinal blade: steep (0); gently-angled (1); gently convex (2). (G07)
12. Long v-shaped longitudinal blade on P3: absent (0); present (1).
13. P3 Posterolingual crest: absent (0); present (1).
14. P3 Lingual cusp: absent (0); present (1). (Modified: MA97, B12)
15. P3 posterobuccal crest: absent (0); well developed (1); moderately developed (2); weakly developed (3). (G07)
16. P3 transverse parametacone crest: absent (0); present (1). (Modified: H93, BA97a, B12)
17. P3 medial cuspule: absent (0); present (1).
18. Posterior cingulum development on P3: buccal and lingual (0); lingual (1); absent (2).
19.* Size of P3/ relative to M1/: P3L/M1L ≤ 0.8 (0); 0.9 < P3L/M1L < 1.1 (1); 1.1< P3L/ M1L≤ 1.2 (2); 1.2 < P3L/M1L≤ 1.4 (3); 1.4< P3L/ M1L≤ 1.5 (4); 1.5 < P3L/ M1L≤ 1.6 (5) P3L/ M1L > 1.6 (6). (Modified: BA97a, B12)
20.* Molar morphology: tribosphenic (0); selenodont (1); semi-lophodont (2); lophodont but stylar cusps evident on lophs (3); fully lophodont (4); bunodont (5). (Modified: B12)
21. Anteriorly concave lower/ anteriorly convex upper molar lophs: absent (0); present (1). (B12)
22. Ever-growing unrooted cheekteeth: absent (0); present (1). (B12)
23. Enamel crenulations: small (0); strong, crest-like (1). (Modified: S97, B12)
24.* Stylar cusp development: strong (0); moderate (1); weak/absent (2). (Modified: BA97b, B12)
25.* Parastyle development on M1: absent/small (0); moderately developed with expansion of anterobuccal tooth corner (1); large, cuspate and pyramid-like (2). (BA97b, B12)
26. Separation of stylar cusps C and D: close together (0); separated by large trough (1). (MA97, B12)
27.* M1 occlusal outline: triangular (0); subtriangular/subsquare (1); square (2); rectangular (3). (G07)
28. M1 metacone: moderate (0); large (1).
29.* M1 paraconule: absent/weak (0); moderate/strong, linear (1); strong, crescentic (2). (Modified: BA9, B12)
30. M1 anteroposterior buccal gradient: paracone much shorter than metacone (0); paracone slightly shorter than metacone (1); paracone and metacone similar height (2); paracone slightly taller than metacone (3); paracone much taller than metacone (4). (G07)
31.* M1 neometaconule: absent (0); weak/small (1); moderate/well developed (2). (Modified: WD87, BA97b, S97)
32. Protostyle: absent (0); present (1). (WD87, BA97b, B12)
33. Paracone and metacone placement: medial (0); buccal (1). (S97, HSV03, B12)
34. Paracone buccal basin on M1 deep, enclosed: absent (0); present (1). (B12)
35. Posterolingual paracristae: absent/weak (0); strongly developed (1). (BA97b, B12)
36. Postprotocrista: present (0); absent (1). (A84b, B12)
37.* M2 occlusal shape: triangular (0); subtriangular (1); rectangular i.e., longer than wide (2); square i.e., width ≥ length (3). (Modified: S97, HSV03, B12)
38.* M2 buccal height vs lingual height: similar (0); slightly taller (1); moderately taller (2); much taller (3). (G07)
39.* M2 buccal inflation of crown below paracone: absent (0); present (1). (G07)
40.* M2 postprotocrista and premetaconule crista: confluent (0); not confluent, separated (1).
41. M2 postparacrista direction: posterobuccal (0); posterior (1).
42. M2 width of trigon basin relative to crown width: broad (0); narrow (1).
43. m3: present (0); absent (1). (AD82, G07)
44. M/m4: present (0); absent (1). (AD82, G07)
45. M4 metaconule: absent or significantly reduced and retracted towards posterior cingulum (0); distinct, cuspate (1). (B12)
46.* Number of lower incisors: three (0); two (1); one (2). (S97, R98, W00, L02, HSV03, B12)
47. Inclination angle of i1: high, ≥ 30 degrees (0); low < 30 degrees (1). (M90, B07, B12)
48. Number of lower premolars: three (0); two (1); one (2).
49. p3 morphology: bicuspid, simple (0); multicusped/bladed (1). (B07, B12)
50. p3 anterolingual crest: well-developed (0); weakly-developed: absent (1); absent (2).
51. Posterior cingulum on p3: absent/weak (0); present (1). (M94, B12)
52. Molar gradient: ratio m4/m1 < 1 (0); m4/m1 ≥ 1 (1). (B07, B12)
53. m2 talonid height relative to paraconid and trigonid: slightly shorter (0); similar (1); much shorter (2).
54. Development of paraconid and paracristid on m1: paraconid present and paracristid well-developed (0); paraconid weak or absent, paracristid present but low (1); paraconid and paracristid absent (2). (B0, B12)
55. Position of protoconid on m1: buccal half of trigonid (0); lingual third of trigonid (1). (BA97b, B12)
56.* Protostylid on m1: absent (0); small (1); moderate (2); large (3). (WD87, S97, BA97b, MA97, B0, B12)
57.* m1 talonid basin width between entocristid and hypocristid: broad (0); slightly narrowed (1); narrow (2). (G07)
58.* m1 talonid width relative to trigonid width: broader (0); slightly narrower (1); much narrower (2). (G07)
59.* m3 talonid basin: broad (0); narrow (1); absent/lost (2).
60.* Metastylid development: absent (0); present, cuspate (1); present, metastylid fold (2). (Modified: BA97b, B12)
61. Entostylid ridge on m1: absent (0); present (1). (BA97b, B12)
62. Cristid obliqua: well-developed, does not meet postprotocristid (0); well-developed, meets postprotocristid lingual to horizontal tooth midline (1); well-developed, meets postprotocristid at or buccal to horizontal tooth midline (2); weak/absent (3). (B12)
63. Internal ribs on conids of lower molars: absent (0); present, meet in longitudinal valley (1); present, do not meet (2). (B12)
64.* Nasal aperture retracted beyond incisor arcade: absent (0); retracted to above diastema (1); retracted to above cheek tooth row (2). (B12)
65.* Masseteric process: absent/weak- dorsal to molar row (0); at level of molar row (1); elongate- extends ventral to molar row (2). (W98, B12)
66. Lacrimal tuberosity: absent (0); present (1). (R98; HSV03, B12)
67. Infraorbital shelf: well-developed (0); weak (1). (B12)
68.* Posterior palatal vacuities: anteriorly extensive to opposite or anterior to M1 (0); extends anteriorly to opposite M2 (1); confined within palatine, opposite M3-4 (2); absent (3). (A84b, R87, R98, W00, HSV03, B12)
69. Frontal/squamosal contact: absent, alisphenoid-parietal contact (0); present (1). (S97, W98, HSV03, B12)
70. Infratemporal crests: weak/absent (0); well-developed (1). (B12)
71. Postglenoid constriction: absent (0); present (1). (B12)
72. Glenoid fossa: flat articular eminence, shallow mandibular fossa (0); flat articular eminence, deep mandibular fossa (1); prominent articular eminence, mandibular fossa absent (2). (B12)
73.* Postglenoid process: elongate (0); short (1); absent (2). (Modified: S97, HSV03, B12)
74. Medial glenoid process: absent (0); present (1). (M86, B12)
75. Well-developed postglenoid cavity: absent (0); present (1). (L04, B12)
76. Position of postglenoid foramen: posterior to PGP and bounded medially by petrosal (0); anteromedial to or in line with postglenoid process (1); posteromedial to postglenoid process within squamosal (2); within epitympanic fenestra, surrounded by bony septum (3). (Modified: S97, W98, HSV03, B12)
77. Tympanic cavity roof elements: alisphenoid and petrosal (0); alisphenoid and squamosal (1); squamosal (2). (Modified: T10, B12)
78. Tympanic floor elements: alisphenoid (0); alisphenoid and squamosal (1); squamosal (2). (B12)
79.* Alisphenoid tympanic wing: absent (0); short (1); moderate, extends under periotic (2); elongate, completely floors middle ear (3). (S97, W98, HSV03, B12)
80. Epitympanic fenestra: absent (0); present (1). (B12)
81. Non auditory sinuses: absent (0); present (1). (M86, B12)
82.* Posterior epitympanic fossa: absent (0); present, shallow (1); deep, perforating squamosal and mastoid (2) (M86, B12)
83. Rostral tympanic process of periotic: strong (0); absent/weak (1). (R98, SVW02, HSV03, B12)
84. Posterior parietal width: broad (0); narrow (1). (B12)
85. Interparietal: present (0); absent (1). (L04, B12)
86. Narrow mastoid strip on occiput: absent (0); present (1). (M86, B12)
87. Ventrolaterally flared mastoid process on occiput: absent (0); present (1). (M86, B12)
88. Angle of the anterior border of the ascending ramus: < 70 degrees (0); ≥ 70 degrees (1). (HW82, B07, B12)
89.* Posterior extent of mandibular symphysis: anterior to p3 (0); below p3 (1); below m1 (2); below m2-3 (3). (B07, B12)
90.* Diastema (between i1 and p3): absent (0); present, length < 30% of horizontal ramus length (1); present, length > 30% of horizontal ramus length (2). (B07, B12)
91. Tooth row length (p3-m4) relative to horizontal ramus length: > 60% (0); ≤ 60% (1). (B12)
92. Masseteric foramen: absent (0); present (1). (B07, B12)
93. Flared masseteric eminences: absent/weak (0); moderately to strongly flared (1). (M98, B12)

APPENDIX 2

Data Matrix

Data matrix used for phylogenetic analysis of Vombatiformes. Abbreviations: '?' signifies 'missing data'; '-' signifies 'inapplicable'. Polymorphic states indicated by: A, (0,1).

  1
0
2
0
3
0
4
0
5
0
6
0
7
0
Barinya wangala 1000000000 0000000010 00-00-0100 0000000101 010000?002 011000000- -20100?110
Galadi speciosus 0000000000 0000000000 -0-0001000 00000012-0 010000?002 0000000000 0200000000
Litokoala kutjamarpensis ??? 2201012 0011001101 -012202121 21011032-1 00000???1? 00?0130002 112?1?0100
Nimiokoala greystanesi 2002200011 2002200011 -012202121 21011031-1 0000020212 0000130001 1120000?00
Vombatus ursinus 4 -121010-0 -000000- 02 01-2003102 00100020-- - -00121202 00210000-0 02-1010211
Warendja wakefieldi 4 -12- 000-0 - 000000212 01-2003- 02 00100020-- - -00121202 00210000-0 02-?00020?
Kuterintja ngama ??? 22010 -3 0001000011 0000112101 00010021-1 0000121212 1021000000 01-?2?0???
Namilamadeta albivenator 2002302001 0011001022 00-0112102 00000021-0 0000121210 1021000000 02-1210111
Marada arcanum ???2?0?0?? ?????????2 00-??????2 ??????? ??? ??00?21202 1001000000 02-???????
Propalorchestes novaculacephalus ??12022?20 0011000003 10-1013102 0010012001 - -00121?0? 1102000000 02-2??1301
Ngapakaldia tedfordi 2112001000 0011010004 10-2013102 0010012001 1-00121202 1102000000 03-1111311
Nimbadon lavarackorum 2112002000 0011010014 10-2013102 0010012001 --0012120- 1101000000 03-0211311
Microleo attenboroughi ??? 0110111 0110101235 - 002002??? ?0?? 001001 0000 ?????? ???????? 0? ?? 0???????
Priscileo roskellyae 2100101122 1100300235 -002001003 0000001000 0000?10000 0011100000 02- 0000001
Priscileo pitikantensis ? 100? 021?? ????????? 5 -00 ????? ?? ????0002? 0 ?100?????? ?????? ?? ?? ?????? ?? ??
Wakaleo oldfieldi 210A102223 1100200235 -002000103 0000002110 1101-10201 0011102110 020 ???????
Wakaleo vanderleueri 210A102223 1100200235 -002000104 0000003110 1101-10202 0011102120 0200000101
Wakaleo alcootaensis ?10 2103223 1100200235 -002?00??? ?????? ?? ?? ??11-??2?? ??????2?-? ??0???????
Thylacoleo hilli ???? 123? 21 21100002 ?? -0???????? ?????? ?? ?? ?????20000 0???0????? ?????? ????
Thylacoleo crassidentatus 2100124321 2110000245 -002100104 000000???? ?111-20?00 00111022-0 020????0??
Thylacoleo carnifex 2100124321 2110000245 -002103104 000000???? ??11-20001 00111022 -0 0200010011
               
  8
0
9
0
9
3
       
Barinya wangala 0100000020 00000?0000 100        
Galadi speciosus 0100000020 0000100?00 100        
Litokoala kutjamarpensis 0100111020 0000000??? ???        
Nimiokoala greystanesi ????1?1??0 0?000??021 010        
Vombatus ursinus 1221122200 0110111132 111        
Warendja wakefieldi ???11?2200 0??01??022 111        
Kuterintja ngama ????? ?? ?? ?????? ?12? ???        
Namilamadeta albivenator 1001122120 0110111021 010        
Marada arcanum ?????? ?? ?? ?????? ?012 110        
Propalorchestes novaculacephal 11111?21?1 1211111122 111        
Ngapakaldia tedfordi 1111132201 1211110122 110        
Nimbadon lavarackorum 1111132111 1211110121 1A0        
Microleo attenboroughi ?????? ?? ?? ?????? ?? ?? ???        
Priscileo roskellyae 0001122120 0010100?10 010        
Priscileo pitikantensis ?????? ?? ?? ?????? ?? ?? ???        
Wakaleo oldfieldi ?????? ?? ?? ?????? ?010 010        
Wakaleo vanderleueri 0001122120 0001110010 010        
Wakaleo alcootaensis ?????? ?? ?? ?????? ?? 1? ???        
Thylacoleo hilli ?????? ?? ?? ?????? ?? ?0 ???        
Thylacoleo crassidentatus ?????? ?? ?? ?????? ?? 10 010        
Thylacoleo carnifex 0001022200 0011110010 010        

List of Synapomorphies

List of nodal apomorphies resulting from the phylogenetic analysis. Node numbers refer to Supplementary Figure 2.

Node Character Change
1 1 1→2
  10 0→1
  20 0→2
  24 0→2
  42 1→0
  46 0→1
  73 0→1
  75 0→1
  76 0→2
  77 0→2
  89 0→1
  91 1→0
  92 0→1
2 4 0→2
  14 0→1
  37 1→2
  46 1→2
  48 0→2
  89 1→2
  90 0→1
3 17 0→1
  18 0→1
  23 0→1
  25 1→2
  29 0→2
  31 0→2
  32 0→1
  34 0→1
  35 0→1
  37 2→3
  56 0→3
  60 0→1
  61 0→1
  62 2→1
  77 2→1
  85 1→0
4 45 0→1
  47 0→1
  51 0→1
  64 0→1
  66 0→1
  71 0→1
  82 0→1
  86 0→1
  87 0→1
5 24 2→0
  53 0→2
  65 1→2
6 3 0→1
  10 1→0
  27 2→3
  33 0→1
  38 1→0
  68 1→2
  73 0→1
  79 2→0
  90 1→2
  91 0→1
7 13 0→1
  20 2→3
  21 0→1
  36 0→1
  52 0→1
  67 0→1
  68 2→3
  80 0→1
  81 0→1
  82 1→2
  84 0→1
8 16 0→1
  20 3→4
  62 2→3
9 1 2→4
  14 1→0
  22 0→1
  51 1→0
  53 0→2
  78 1→2
  93 0→1
10 12 0→1
  18 0→2
  19 0→3
  20 2→5
11 11 0→1
  40 1→0
12 7 1→2
  37 1→0
  38 0→2
  42 0→1
13 8 1→2
  44 0→1
14 4 0→2
  10 2→3
  15 0→2
  48 0→2
15 6 0→2
  7 2→3
  11 1→2
  13 0→1
  46 1→2
16 7 3→4

APPENDIX 3

Body Mass Estimate

An estimate of the body mass of Microleo attenboroughi, Priscileo roskellyae and Wakaleo sp. nov. was made using a regression equation formulated by Myers (2001) from correlations of marsupial body mass with cranio-dental measurements. The equation used for the analysis was the highest possible ranked regression from the ‘Diprotodontians data set’ of Myers (2001, table 5, p.106) and employed the measurement for upper molar row length (UMRL, i.e., M1-M4). This equation (log y = -0.418 + 3.011 [log x], where x equals UMRL) was chosen because it was based on alveolar measurements rather than molar measurements; the posterior teeth are missing in both fossil taxa. The UMRL measurements (average of the left and right lengths) for M. attenboroughi (11.3 mm) P. roskellyae (QM F23453, 16.4 mm) and Wakaleo sp. nov. (QM F45200, 23.3 mm) resulted in body weight estimates of 590 g, 1813 g, and 5221 g, respectively. As recommended by Myers (2001), a smearing estimate (4.4%) was applied to the equation. Myers (2001) indicates that the regression equations are not appropriate for the larger species of marsupial lions (based on their unusual reduced molar numbers that result in large underestimates of body mass), hence, an estimate was not calculated for Wakaleo vanderleueri.