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Additionally, the lead author will host a formatted, down-loadable version of the cladistic character matrix on The Field Museum public access server


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Additionally, the lead author will host a formatted, down-loadable version of the cladistic character matrix on The Field Museum public access server.

Supplementary information to MS 2005-05-05206:




The earliest dromaeosaurid theropod from South America.

Peter J. Makovicky1, Sebastian Apesteguía2,3, & Federico L. Agnolín4



1 Department of Geology, The Field Museum, 1400 S Lake Drive, Chicago IL 60605, USA.

2 Sección de Paleontología de Vertebrados, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Av. Ángel Gallardo 470, Buenos Aires (1405), Argentina.

3 Fundación de Historia Natural "Félix de Azara",CAECNA, Universidad Maimónides, V. Virasoro 732, Buenos Aires (1405), Argentina.

4 Laboratorio de Anatomía Comparada, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Av. Ángel Gallardo 470, Buenos Aires (1405), Argentina.




  1. Character list

  2. Data matrix

  3. Analysis protocol and results

  4. Clade diagnoses and definition of Unenlagiinae

  5. Supplementary Discussion

  6. Select measurements of Buitreraptor

  7. Comparative measurements of basal paravian limb elements

  8. References

1. Character list modified and expanded from (Xu and Norell, 2004). A number of novel characters were added in order to test the relationships of Unenlagiinae. These include the pedal flange character introduced by (Novas and Pol, 2005) [char. 229], as well as iliac features described by (Novas, 2004) [char. 226], Senter et al. (2004) [char. 231] or recognized here [char. 227], and pedal characteristics [char. 228], all relevant to paravian interrelationships. Other characters or states related to the tail [char. 116, state 2], ischium [chars. 230, 232], foot [char. 235] and forelimb length [char. 236] were added in order to more rigorously test the phylogenetic position of Rahonavis found in this analysis. A few previously employed characters such as the arctometatarsal foot [char. 203], were redefined to better capture phylogenetically informative variation following recent descriptions. A single character (character 30 [antorbital fossa emargination] of Xu and Norell [2004]) was eliminated, as it is too vaguely defined to score in a non-arbitrary manner. Its deletion had no topological effect. Matrix and characters are downloadable from the following web-page http://fm1.fieldmuseum.org/aa/staff_page.cgi?staff=pmakovicky in NONA format).





  1. Vaned feathers on forelimb symmetric (0) or asymmetric (1). The barbs on opposite sides of the rachis differ in length; in extant birds, the barbs on the leading edge of flight feathers are shorter than those on the trailing edge.



Skull


  1. Orbit round in lateral or dorsolateral view (0) or dorsoventrally elongate (1).

  2. Anterior process of postorbital projects into orbit (0) or does not project into orbit (1).

  3. Postorbital in lateral view with straight anterior (frontal) process (0) or frontal process curves anterodorsally and dorsal border of temporal bar is dorsally concave (1).

  4. 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).

  5. Otosphenoidal crest vertical on basisphenoid and prootic, and does not border an enlarged pneumatic recess (0) or well developed, crescent shaped, thin crest forms anterior edge of enlarged pneumatic recess (1). This structure forms the anterior, and most distinct, border of the “lateral depression” of the middle ear region (Currie, 1985; Currie and Zhao, 1993) of troodontids and some extant avians.

  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 present between basisphenoid and basioccipital (0) or 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 not highly pneumatized (0) or base of cultriform process (parasphenoid rostrum) expanded and pneumatic (parasphenoid bulla present) (1).

  11. Basipterygoid processes ventral or anteroventrally projecting (0) or lateroventrally projecting (1).

  12. Basipterygoid processes well developed, extending as a distinct process from the base of the basisphenoid (0) or processes abbreviated or absent (1).

  13. Basipterygoid processes solid (0) or processes hollow (1).

  14. Basipterygoid recesses on dorsolateral surfaces 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). The dorsal tympanic recess referred to here is the depression anterodorsal to the middle ear on the opisthotic, not the recess dorsal to the crista interfenestralis within the middle ear as seen in Archaeopteryx lithographica, Shuuvuia deserti and Aves.

  16. Accessory tympanic recess dorsal to crista interfenestralis absent (0) small pocket present (1) or extensive with indirect pneumatization (2). According to (Witmer, 1990), this structure may be an extension from the caudal tympanic recess, although it has been interpreted as the main part of the caudal tympanic recess by some previous authors.

  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 C. N. X-XII flush with surface of exoccipital (0) or cranial nerve exits located together in a bowl-like depression (1).

  19. Maxillary process of premaxilla contacts nasal to form posterior border of nares (0) or maxillary process reduced so that maxilla participates broadly in external naris (1) or maxillary process of premaxilla extends posteriorly to separate maxilla from nasal posterior to nares (2).

  20. Internarial bar rounded (0) or flat (1).

  21. Crenulate margin on buccal edge of premaxilla absent (0) or present (1).

  22. Caudal margin of naris farther rostral than (0), or nearly reaching or overlapping (1), the rostral border of the antorbital fossa (Chiappe et al., 1998).

  23. Premaxillary symphysis acute, V-shaped (0) or rounded, U-shaped (1).

  24. Secondary palate short (0) or long, with extensive palatal shelves on maxilla (1) REDEFINED.

  25. Palatal shelf of maxilla flat (0) or with midline ventral ‘tooth-like’ projection (1)

  26. Pronounced, round accessory antorbital fenestra absent (0) or present (1). A small fenestra, variously termed the accessory antorbital fenestra or maxillary fenestra, penetrates the medial wall of the antorbital fossa anterior to the antorbital fenestra in a variety of coelurosaurs and other theropods.

  27. Accessory antorbital fossa situated at rostral border of antorbital fossa (0) or situated posterior to rostral border of fossa (1).

  28. Tertiary antorbital fenestra (fenestra promaxillaris) absent (0) or present (1).

  29. Narial region apneumatic or poorly pneumatized (0) or with extensive pneumatic fossae, especially along posterodorsal rim of naris (1).

  30. Jugal and postorbital contribute equally to postorbital bar (0) or ascending process of jugal reduced and descending process of postorbital ventrally elongate (1).

  31. Jugal tall beneath lower temporal fenestra, twice or more as tall dorsoventrally as it is wide transversely (0) or rod-like (1).

  32. Jugal pneumatic recess in posteroventral corner of antorbital fossa present (0) or absent (1).

  33. Medial jugal foramen present on medial surface ventral to postorbital bar (0) or absent (1).

  34. Quadratojugal without horizontal process posterior to ascending process (reversed L shape) (0) or with process (i.e., inverted T or Y shape) (1).

  35. Jugal and quadratojugal separate (0) or quadratojugal and jugal fused and not distinguishable from one another (1).

  36. Supraorbital crests on lacrimal in adult individuals absent (0) or dorsal crest above orbit (1) or lateral expansion anterior and dorsal to orbit (2).

  37. Enlarged foramen or foramina opening laterally at the angle of the lacrimal above antorbital fenestra absent (0) or present (1).

  38. Lacrimal anterodorsal process absent (inverted L shaped) (0) or lacrimal T shaped in lateral view (1) or anterodorsal process much longer than posterior process (2). .

  39. Prefrontal large, dorsal exposure similar to that of lacrimal (0) or greatly reduced in size (1) or absent (2). .

  40. Frontals narrow anteriorly as a wedge between nasals (0) or end abruptly anteriorly, suture with nasal transversely oriented (1).

  41. Anterior emargination of supratemporal fossa on frontal straight or slightly curved (0) or strongly sinusoidal and reaching onto postorbital process (1) (Currie, 1995).

  42. Frontal postorbital process (dorsal view): smooth transition from orbital margin (0) or sharply demarcated from orbital margin (1) (Currie, 1995).

  43. Frontal edge smooth in region of lacrimal suture (0) or edge notched (1) (Currie, 1995).

  44. Dorsal surface of parietals flat, lateral ridge borders supratemporal fenestra (0) or parietals dorsally convex with very low sagittal crest along midline (1) or dorsally convex with well developed sagittal crest (2).

  45. Parietals separate (0) or fused (1).

  46. Descending process of squamosal parallels quadrate shaft (0) or nearly perpendicular to quadrate shaft (1).

  47. Descending process of squamosal contacts quadratojugal (0) or does not contact quadratojugal (1).

  48. Posterolateral shelf on squamosal overhanging quadrate head absent (0) or present (1) (Currie, 1995).

  49. 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).

  50. Quadrate vertical (0) or strongly inclined anteroventrally so that distal end lies far forward of proximal end (1).

  51. Quadrate solid (0) or hollow, with foramen on posterior surface (1).

  52. Lateral border of quadrate shaft straight (0) or with broad, triangular process along lateral edge of shaft contacting squamosal and quadratojugal above an enlarged quadrate foramen (1)(Currie, 1995).

  53. Foramen magnum subcircular, slightly wider than tall (0) or oval, taller than wide (1) (Makovicky and Sues, 1998).

  54. Occipital condyle without constricted neck (0) or subspherical with constricted neck (1).

  55. Paroccipital process elongate and slender, with dorsal and ventral edges nearly parallel (0) or process short, deep with convex distal end (1).

  56. Paroccipital process straight, projects laterally or posterolaterally (0) or distal end curves ventrally, pendant (1).

  57. Paroccipital process with straight dorsal edge (0) or with dorsal edge twisted rostrolaterally at distal end (1) (Currie 1995).

  58. Ectopterygoid with constricted opening into fossa (0) or with open ventral fossa in the main body of the element (1).

  59. Dorsal recess on ectopterygoid absent (0) or present (1).

  60. Flange of pterygoid well developed (0) or reduced in size or absent (1).

  61. Palatine and ectopterygoid separated by pterygoid (0) or contact (1) (Currie 1995).

  62. Palatine tetraradiate, with jugal process (0) or palatine triradiate, jugal process absent (1)(Elzanowski and Wellnhofer, 1996).

  63. Suborbital fenestra similar in length to orbit (0) or reduced in size (less than one quarter orbital length) or absent (1)(Clark et al., 1994).



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