A new carnivorous dinosaur from the late Jurassic Solnhofen archipelago
Ursula B. Göhlich1 and Luis M. Chiappe2
1Dept. for Geo- and Environmental Sciences, Section Paleontology, University of Munich, Richard-Wagner-Strasse 10, D-80333 Munich, Germany, email: .
2 The Dinosaur Institute, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA90007, USA, email:
Supplementary Figure 1. Location of the type locality Schamhaupten (Southern Germany, Bavaria, Franconian Alb). (Modified from Meyer & Schmidt-Kaler, 1989).
Supplementary Figure 2. Interpretive drawing of the bow-shaped zygapophyses of the mid-caudal (C18 to C21) vertebrae of Juravenator starki. Abbreviations: C18-C21, caudal vertebrae 18th to 21st; ha, haemal arches; ncs, neurocentral sutures; zy, zygapophyses.
Table 1. Selected measurements of the bones of Juravenator starki (JME Sch 200).
length / lengthskull / 82 mm
scapula (left)
humerus (left)
ulna (left)
radius (left)
manus (with claws) (left) / 42 mm
27 mm
19.5 mm
~19.3 mm
41 mm / ilium (right)
femur (right)
tibia (left/right)
pes (with claws) (left) / 40 mm
52 mm
58,1 mm
63 mm
metacarpal I (left)
metacarpal II (left)
metacarpal III (left)
phalanx I-1 (left)
phalanx II-1 (left)
phalanx II-2 (left)
phalanx III-1 (left)
phalanx III-2 (left)
phalanx III-3 (left)
claw I (left)
claw II (left/right)
claw III (left/right) / 4.5 mm
11.5 mm
9 mm
10.5 mm
8 mm
10 mm
4 mm
4.5 mm
5.5 mm
~10 mm
9 mm / 10 mm
5.5 mm / 7 mm / metatarsal I (left/right)
metatarsal II (left)
metatarsal III (left)
metatarsal IV (left/right)
metatarsal V (left/right)
phalanx I-1 (left/right)
phalanx II-1 (left)
phalanx II-2 (left/right)
phalanx III-1 (left/right)
phalanx III-2 (left/right)
phalanx III-3 (left)
phalanx IV-1 (left/right)
phalanx IV-2 (left/right)
phalanx IV-3 (right)
phalanx IV-4 (left/right)
claw I (left/right)
claw II (left/right)
claw III (left/right)
claw IV (left/right) / 4.6 mm / 4.5 mm
26.5 mm
34.0 mm
29.6 mm / 29.8 mm
8 mm / 6,8 mm
5.8 mm / 6 mm
10.4 mm
9.0 mm / 8 mm
11.9 mm / 11.5 mm
8.1 mm / 8 mm
7.4 mm
7.4 mm / 7 mm
5.5 mm / 6.5 mm
4.5 mm
4.2 mm / 4 mm
6.0 mm / 3.5 mm
10.7 mm / 11.5 mm
7.4 mm / 6.6 mm
7.2 mm / 5.8 mm
Cladistic Analysis. Morphological data of Juravenator starki was included in a matrix of 189 characters (see Appendix 1). The 35 terminal taxa included Juravenator starki, 30 coelurosaur species (Microraptor zhaoianus, Sinornithosaurus millenii, Velociraptor mongoliensis, Shuuvuia deserti, Mononykus olecranus, Patagonykus puertai, Confuciusornis sanctus, Archeopteryx lithographica, Caudipteryx zoui, Conchoraptor gracilis, Citipati osmolskae, Shenzhousaurus orientalis, Ornithomimus edmonticus, Gallimimus bullatus, Troodon formosus, Saurornithoides mongoliensis, Sinovenator changii, Alxasaurus elesitaiensis, Erlikosaurus andrewsi, Albertosaurus sarcophagus, Tyrannosaurus rex, Ornitholestes hermanni, Huaxiagnathus orientalis, Sinosauropteryx prima, Compsognathus longipes, Mirischia asymmetrica, Scipionyx samniticus, Nqwebasaurus thwazi, Coelurus fragilis, Nedcolbertia justinhoffmanni), two basal tetanurans (Allosaurus fragilis and Sinraptor dongi), and two non-tetanuran theropods (Coelophysis bauri and Dilophosaurus wetherilli). An initial analysis of all these taxa resulted in a strict consensus cladogram (107 trees of 534 steps) with limited resolution at the base of the tree. In this tree, Juravenator was nested with a number of other basal coelurosaurs in a large polichotomy (Supplementary Figure 3). A second analysis omitting seven taxa, either less relevant (i.e., Alxasaurus elesitaiensis, Erlikosaurus andrewsi) or largely incomplete (i.e., Mirischia asymmetrica, Scipionyx samniticus, Nqwebasaurus thwazi, Coelurus fragilis, Nedcolbertia justinhoffmanni), resulted in the cladogram discussed and figured in the text (Figure 4 of text). Both analyses wereconducted using 5 replications of random addition sequences of the taxa creating Wagner trees, each followed by TBR (Tree Bisection Reconnection) branch swapping and 100 iterations of jackknife ratchet (Nixon, 1999), collapsing the trees on TBR rearrangements. This search strategy was implemented using Goloboff’s (1993) program NONA. Given the size of the present data, this strategy is almost certain to produce an entirely accurate consensus tree (see Goloboff and Farris, 2001). All cladograms used Coelophysis bauri as the root.
Supplementary Figure 3. Strict consensus tree (534 steps) resultant from the cladistic analysis of the data matrix listed in the Appendix 2.
Appendix 1. Character list for cladistic analysis based on Hwang et al. (2004). Of the 222 characters used by Hwang et al. (2004), we have omitted 43 characters (i.e., Hwang et al.’s characters 6, 30, 32, 37, 45, 49, 50, 55, 92, 95, 99, 100, 103, 107, 109, 119, 122, 134, 135, 141, 143, 156, 163, 166, 168, 169, 170, 174, 177, 182, 189, 191, 198, 199, 203, 204, 210, 211, 214, 215, 216, 219, 220). We have also added 10 characters to the remaining list and modified (description and/or states) a number of Hwang et al.’s (2004) original characters (see below). The states of multistate characters 40, 44, 70, 77, 89, 90, 110, 113, 120, 121, 148, 159, 167, 168, 172, and 186 were treated as ‘ordered’ (additive characters) (see below).
Integument
1. Vaned feathers on forelimb: symmetric (0) or asymmetric (1).
Skull
2. Orbit in lateral or dorsolateral view: round (0) or dorsoventrally elongate (1).
3. Anterior process of postorbital: projects into orbit (0) or does not project into orbit (1).
4. Frontal (anterior) process of postorbital in lateral view: straight (0) or anterodorsally curved with concave dorsal border (1).
5. Postorbital bar: parallels quadrate, lower temporal fenestra rectangular in shape (0) or jugal and postorbital approach or contact quadratojugal to constrict lower temporal fenestra (1).
6. Crista interfenestralis: confluent with lateral surface of prootic and opisthotic (0) or distinctly depressed within middle ear opening (1).
7. Subotic recess (pneumatic fossa ventral to fenestra ovalis): absent (0) or present (1).
8. Basisphenoid recess: located between basisphenoid and basioccipital (0), entirely within basisphenoid (1), or absent (2).
9. Posterior opening of basisphenoid recess: single (0) or divided into two small, circular foramina by a thin bar of bone (1).
10. Base of cultriform process (parasphenoid rostrum): not highly pneumatized (0) or expanded and pneumatic (parasphenoid bulla) (1).
11. Basipterygoid processes: projecting ventrally or anteroventrally (0) or lateroventrally (1).
12. Basipterygoid processes: well developed, extending as a distinct process from the base of the basisphenoid (0) or abbreviated or absent (1).
13. Basipterygoid processes: solid (0) or hollow (1).
14. Basipterygoid recesses on dorsolateral surface of basipterygoid processes: absent (0) or present (1).
15. Depression for pneumatic recess on prootic: absent (0) or present as dorsally open fossa on prootic/opisthotic (1) or present as deep, posterolaterally directed concavity (2).
16. Accessory tympanic recess dorsal to crista interfenestralis: absent (0), small pocket present (1), or extensive with indirect pneumatization (2).
17. Caudal (posterior) tympanic recess: absent (0), present as opening on anterior surface of paroccipital process (1), or extends into opisthotic posterodorsal to fenestra ovalis, confluent with this fenestra (2).
18. Exits of cranial nerves X-XII: flush with surface of exoccipital (0) or cranial nerve exits located together in a bowl-like basisphenoid depression (1).
19. Maxillary process of premaxilla: contacts nasal to form posterior border of external nares (0), maxillary process reduced so that maxilla participates broadly in external naris (1), or maxillary process extends posteriorly to separate maxilla from nasal posterior to nares (2).
20. Internarial bar: rounded (0) or flat (1).
21. Premaxillary subnarial depth: main body as long or longer rostrocaudally than high dorsoventrally (0) or very high, much shorter rostrocaudally than dorsoventrally (1). In birds, the main body of the premaxilla is not entirely ventral to the external nares—the latter has migrated caudally in more or less degree. For this reason, we have scored Archaeopteryx and Confuciusornis as non-comparable. (added)
22. Crenulate margin on buccal edge of premaxilla: absent (0) or present (1).
23. Caudal margin of naris: farther rostral than (0), or nearly reaching or overlapping (1), the rostral border of the antorbital fossa.
24. Premaxillary symphysis: acute, V-shaped (0) or rounded, U-shaped (1).
25. Secondary palate: formed by premaxilla only (0) or by premaxilla, maxilla, and vomer (1).
26. Palatal shelf of maxilla: flat (0) or with midline ventral ‘tooth-like’ projection (1).
27. Pronounced, round accessory antorbital fenestra (maxillary fenestra): absent (0) or present (1).
28. Accessory antorbital fenestra (maxillary fenestra): situated at rostral border of antorbital fossa (0) or situated posterior to rostral border of fossa (1).
29. Tertiary antorbital fenestra (promaxillary fenestra): absent (0) or present (1).
30. Maxillary rostral margin: convex (0), concave, rostral ramus equal or shorter than higher (1), concave, rostral ramus longer than higher (2). (added)
31. Nasal participation in antorbital fossa: minimal or absent (0), present (1). (added)
32. Narial region: apneumatic or poorly pneumatized (0) or with extensive pneumatic fossae, especially along posterodorsal rim of fossa (1).
33. Jugal tall beneath lower temporal fenestra, twice or more as tall dorsoventrally as it is wide transversely (0) or rod-like (1). We coded the very short quadratojugal process of the jugal of Archaeopteryx (Elzanowski and Wellnhofer, 1996) as non-comparable.
34. Jugal pneumatic recess in posteroventral corner of antorbital fossa: present (0) or absent (1).
35. Medial jugal foramen: present on medial surface ventral to postorbital bar (0) or absent (1).
36. Quadratojugal: without horizontal process posterior to ascending process (reversed “L” shape) (0) or with process (i.e., inverted ‘T’ or ‘Y’ shape) (1).
37. Supraorbital crests on lacrimal in adult individuals: absent (0) or dorsal crest above orbit (1) or lateral expansion anterior and dorsal to orbit (2).
38. Enlarged foramen or foramina opening laterally at the angle of the lacrimal: absent (0) or present (1).
39. Lacrimal posterodorsal process: absent (inverted ‘L’ shaped) (0) or lacrimal ‘T’ shaped in lateral view (1) or anterodorsal process much longer than posterior process (2).
40. Prefrontal dorsal exposure: large, similar to or larger than that of lacrimal (0) or greatly reduced in size (1) or absent (2). (ordered)
41. Anterior end of frontals: narrow, forming a wedge between nasals (0) or ending abruptly anteriorly, suture with nasal transversely oriented (1).
42. Anterior emargination of supratemporal fossa on frontal: straight or slightly curved (0) or strongly sinusoidal and reaching onto postorbital process (1) (see Currie 1995).
43. Postorbital process of frontal in dorsal view: smooth transition from orbital margin (0) or sharply demarcated from orbital margin (1) (see Currie 1995).
44. Dorsal surface of parietals: flat, lateral ridge borders supratemporal fenestra (0), parietals dorsally convex with very low sagittal crest along midline (1), or dorsally convex with well developed sagittal crest (2). (ordered)
45. Parietals: separate (0) or fused (1).
46. Descending process of squamosal: parallel (0) or nearly perpendicular (1) to quadrate shaft.
47. Dorsal process of quadrate: single headed (0) or with two distinct heads, a lateral one contacting the squamosal and a medial head contacting the braincase (1).
48. Quadrate: vertical (0) or strongly inclined anteroventrally so that distal end lies far forward of proximal end (1).
49. Quadrate: solid (0) or hollow, with depression on posterior surface (1).
50. Quadrate head in lateral view: covered by squamosal (0) or exposed, quadrate cotyle of squamosal open laterally (1).
51. Lateral border of quadrate shaft: straight in caudal view (0) or with lateral tab that touches squamosal and quadratojugal above an enlarged quadrate foramen (1), when these bones are in contact to each other. Although the squamosal and quadratojugal of Shuvuuia do not contact each other (Chiappe et al., 2002), the lateral border of the quadrate of this taxon is laterally notched and not straight—thus, we have still coded as “1” despite the absence of contact between the squamosal and quadratojugal. (states modified)
52. Occipital condyle: without constricted neck (0) or subspherical with constricted neck (1).
53. Paroccipital process: elongate and slender, with dorsal and ventral edges nearly parallel (0) or short and deep, with convex distal end (1).
54. Paroccipital process: straight, projects laterally or posterolaterally (0) or distal end curves ventrally, pendant (1).
55. Dorsal edge of paroccipital process: straight (0) or twisted rostrolaterally at distal end (1) (see Currie 1995).
56. Ectopterygoid: with constricted opening into fossa (0) or with open ventral fossa in the main body of the element (1).
57. Ventral recess on ectopterygoid: absent (0) or present (1).
58. Flange of pterygoid: well developed (0) or reduced in size or absent (1).
59. Palatine and ectopterygoid: separated by pterygoid (0) or contact each other (1) (see Currie 1995).
60. Palatine: tetraradiate, with jugal process (0) or triradiate, jugal process absent (1).
61. Suborbital fenestra: similar in length to orbit (0) or reduced in size (less than one quarter orbital length) or absent (1).
62. Depression (possibly pneumatic) on ventral surface of postorbital process of laterosphenoid: absent (0) or present (1).
63. Basal tubera: set far apart, level with or beyond lateral edge of occipital condyle and/or foramen magnum (may connected by a web of bone or separated by a large notch) (0) or tubera small, directly below condyle and foramen magnum, and separated by a narrow notch (1).
Mandible
64. Symphyseal region of dentary in occlusal view: almost straight (symphyseal surface aligned with medial margin of mandible) (0) or medially recurved (1). (states modified; see Rauhut, 2003)
65. Dentary symphyseal region: in line with main part of buccal edge (0) or symphyseal end downturned (1).
66. Mandibular coronoid prominence: absent (0) or present (1).
67. Posterior end of dentary: straight or only slightly concave (minimal or no development of process dorsal to mandibular fenestra, if present) (0) or strongly forked (with strong process above mandibular fenestra, if present) (1) (see Rauhut, 2003). (states modified)
68. Labial face of dentary: flat (0) or with lateral ridge and inset tooth row (1).
69. Shape of dentary in lateral view: subtriangular (0) or with subparallel dorsal and ventral edges (1) (see Currie 1995).
70. Dentary: fully toothed (0) or only with teeth rostrally (1) or edentulous (2). (ordered)
71. External mandibular fenestra: large (0), reduced or absent (1). (added)
72. Nutrient foramina on external surface of dentary: superficial (0) or lie within deep groove (1).
73. Large external mandibular fenestra: oval (0) or subdivided by a spinous rostral process of the surangular (1). (states modified)
74. Internal mandibular fenestra: absent or small and slit-like (0) or large and rounded (1) (see Currie 1995). (states modified)
75. Foramen in lateral surface of surangular rostral to mandibular articulation, absent (0) or present (1).
76. Lateral exposure of splenial: not widely exposed (0) or exposed as a broad triangle between dentary and angular (1).
77. Coronoid ossification: large (0), or only a thin splint (1), or absent (2). (ordered)
78. Laterally inclined flange along dorsal edge of surangular for articulation with lateral process of lateral quadrate condyle: absent (0) or present (1).
79. Angular exposed almost to end of mandible in lateral view: reaches or almost reaches articular (0) or excluded from posterior end angular suture turns ventrally and meets ventral border of mandible rostral to glenoid (1).
80. Articular: without elongate, slender medial, posteromedial, or mediodorsal process from retroarticular process (0) or with process (1).
81. Retroarticular process: short, stout (0) or elongate and slender (1).
82. Mandibular articulation surface: as long as distal end of quadrate (0) or twice or more as long as quadrate surface, allowing anteroposterior movement of mandible (1).
83. Jaws occlusion: throughout their length (0) or jaws diverge rostrally due to kink and downward deflection in dentary buccal margin (1).
Dentition
84. Premaxilla teeth: present (0) or absent (1).
85. Premaxillary teeth: all or some serrated (0), without serrations (1). (added)
86. Second premaxillary tooth: approximately equivalent in size to other premaxillary teeth (0) or markedly larger than third and fourth premaxillary teeth (1) (see Currie 1995).
87. Maxillary teeth: present (0) or absent (1).
88.Caudal extension of upper tooth row: to within the level of orbit (0), to rostral margin of, or anterior to the orbit (1). (added)
89. Maxillary and dentary teeth: serrated (0) or some without serrations anteriorly (except at base in Saurornithoides mongoliensis) (1) or all without serrations (2). (ordered)
90. Dentary and maxillary teeth: large, less than 25 in dentary (0) or moderate number of small teeth (25-30 in dentary) (1) or teeth relatively small, and numerous (more than 30 in dentary) (2). (ordered)
91. Serration denticles: large (0) or small (1).
92. Serrations: simple, denticles convex (0) or distal and often mesial edges of teeth with large, hooked denticles that point toward the tip of the crown (1).
93. Teeth constriction between root and crown: present (0) or absent (1).
94. Dentary teeth: evenly spaced (0) or anterior dentary teeth smaller, more numerous, and more closely appressed than those in middle of tooth row (1).
95. Dentary interdental plates: absent (0) or present medially between teeth (1).
96. Premaxillary tooth crowns in cross section: sub-oval to sub-circular (0) or asymmetrical (D-shaped in cross section) with flat lingual surface (1).
Axial skeleton
97. Axial epipophyses: absent or poorly developed, not extending past posterior rim of postzygopophyses (0) or large and posteriorly directed, extend beyond postzygapophyses (1).
98. Axial neural spine: flared transversely (0) or compressed mediolaterally (1).
99. Anterior cervical centra: level with or shorter than posterior extent of neural arch (0) or extending beyond posterior limit of neural arch (1).
100. Carotid process on posterior cervical vertebrae: absent (0) or present (1).
101. Cervical vertebrae pneumaticity: two pneumatic foramina arranged horizontally (0) or a single pneumatic foramen (1). (added)
102. Anterior cervical centra: subcircular or square in anterior view (0) or distinctly wider than high, kidney shaped (1).
103. Cervical and anterior dorsal vertebrae: amphiplatyan (0) or opisthocoelous (1).
104. Anterior dorsal vertebrae: without prominent hypapophyses (0) or with large hypapophyses (1).
105. Hyposphene-hypantrum articulations in dorsal vertebrae: absent (0) or present (1).
106. Zygapophyses of dorsal vertebrae: abutting one another above neural canal, opposite hyposphenes meet to form lamina (0), or zygapohyses placed lateral to neural canal and separated by groove for interspinuous ligaments, hyposphenes separated (1).
107. Presacral vertebrae pneumaticity: cervical vertebrae but not dorsal vertebrae pneumatic (0) or all presacral vertebrae pneumatic (1).
108. Apices of neural spines of dorsal vertebrae: not expanded (0) or expanded transversally to form a ‘spine table’ (1).
109. Neural spines on posterior dorsal vertebrae in lateral view: rectangular or square (0) or anteroposteriorly expanded distally, fan-shaped (1).
110. Number of sacral vertebrae: 4 or 5 (0), or 6 or 7 (1), or 8 or more (2). (states modified; ordered) Our coding for Gallimimus follows Osmolska et al. (1972) and differs from that given by Hwang et al. (2004). (states modified)
111. Sacral vertebrae: with unfused zygapophyses (0) or with fused zygapophyses forming a sinuous ridge in dorsal view (1).
112. Ventral surface of posterior sacral centra: convex and gently rounded (0), or ventrally flattened, sometimes with a shallow sulcus (1), or centrum strongly constricted transversely, ventral surface keeled (2).
113. Pleurocoels on sacral vertebrae: absent (0), or present on anterior sacrals only (1), or present on all sacrals (2). (ordered)
114. Last sacral centrum: with flat posterior articulation surface (0) or convex articulation surface (1).
115. Caudal vertebrae: with distinct transition point, from shorter centra with long transverse processes proximally to longer centra with small or no transverse processes distally (0) or vertebrae homogeneous in shape, without transition point (1).
116. Transition point in caudal series: begins distal to the 10th caudal (0) or at or proximal to the 10th caudal vertebra (1).
117. Anterior caudal centra: tall, oval in cross section (0), with box-like centra in caudals I-V (1), or anterior caudal centra laterally compressed with ventral keel (2).
118. Neural spines of caudal vertebrae: simple, undivided (0) or separated into anterior and posterior alae throughout much of caudal sequence (1).
119. Prezygapophyses of distal caudal vertebrae: between 1/3 and whole centrum length (0), or with extremely long extensions of the prezygapophyses (up to 10 vertebral segments long in some taxa) (1), or strongly reduced as in Archaeopteryx lithographica (2).