Indices Academiatrum, or, a Dialogue on an AIG Webcast with Jonathan Sarfati, on Avian Phylogeny
Webcast Source:
http://www.gospelcom.net/cgi-bin/ram...thUntruths1.rm
1. Introduction
In an interview featured on the website of Answers in Genesis, the Australian creationist organization, Jonathan Sarfati is featured in a discussion concerning the marvelous science of avian phylogenetics. The intent of the Dialogue presented herein is to address multiple aspects of the Sarfati interview and elucidate with all brevity possible, the data, which renders several conclusions offered by Sarfati untenable. Specifically, the authors will focus on the following areas:
a)Theropod aerobic capacity, with particular emphasis on Maniraptoriformes
b)Integumentary structure of Maniraptoriformes
c)Integumentary derivates and exaptation thereof
d)Phylogenetic placement of Mononykus olecranus, Protarchaeopteryx robusta, and Caudipteryx zhoui
e)Comparative body proportions in Theropoda, with particular emphasis on Maniraptoriformes
f)Temporal discontinuity
g)The plesiomorphic osteology of Archaeopteryx lithographica
h)Digital homology
i)Support for theropod origin
2. Aerobic Capacity in Theropoda, with Particular Emphasis on Maniraptoriformes
The principal arguments advanced by Sarfati in this regard during the AIG interview are based on the research of John Ruben of the University of Oregon and his colleagues. Said arguments conclude that Theropoda possessed a pelvovisceral piston in which diaphragmatic musculature anchored to the procumbent pubes operates upon the hepatic capsule, ventilating the lungs (Ruben et al, 1996, 1997, 1998, 1999).
Pelvovisceral systems are further characterized by an airtight post-pulmonary septum inferior to the liver, which preserves a pressure differential between the cranial and caudal aspects of the thorax (Gans & Clark 1976, Perry 1988, 1990). Osteological characters associated with such a system include (following Mook 1921, Gans & Clark 1976, Duncker 1978, Perry 1988, 1990, Ruben et al 1997, Paul 2002):
a) Thoracic ribs articulate with the dorsal vertebrae via hyper-elongated transverse processes and single proximal heads
b) Gastralia do not meet medially, and are instead set in a cartilaginous sheet
c) Pubes mobile, procumbent
d) Lumbar region formed by reduction of the caudal thoracic ribs
e) Sternal ribs doubled
f) Diverticula largely absent, pneumatic excavation of bones inconsistent or absent, most especially caudal to the post-pulmonary septum
Characters a, b, c, and d, are biomechanical necessities of a pelvovisceral pump. The modified rib articulation with the dorsal vertebrae and failure of the gastralia to meet medially permits the cranial aspect of the thorax (superior to the post-pulmonary septum) to present a smooth surface both dorsally and ventrally in which the viscera can readily move in concert with the action of the diaphragmatic musculature. Similarly, such musculature requires mobility and a procumbent orientation of the pubes (and most likely a high breadth ratio). The presence of a lumbar region is consistently observed in all taxa in which a pelvovisceral pump is found, and is therefore most logically considered to be an essential aspect of the system as a whole. The doubling of the sternal ribs assists in regulation of the pelvovisceral mechanism. The absence or inconsistent presence of pneumatic excavation of the bones caudal to the post-pulmonary septum is particularly indicative of such an aerobic system, as the septum maintains a pressure differential between the fore and aft thoracic compartments.
Ruben et al presented their conclusions on the aerobic system present in Theropoda following the 1996 description of the type Sinosauropteryx prima (NGMC 2124) (Ji & Ji 1996) from the Lower Cretaceous Yixian lagerstatten of China, in which a concentration of carbonized material is preserved in the caudal region of the thorax. Ruben and his colleagues identified the material as the liver, and argued that the cranial aspect of this region was convex, demonstrating the presence of a post-pulmonary septum, and thus by inference, a pelvovisceral pump. Ruben et al further asserted that muscle fibers preserved in the type Sinosauropteryx were indicative of the requisite diaphragmatic musculature, inserting on the distal pubes (Ruben et al 1997, Feduccia 1999). Ruben and his colleagues have further maintained that the type Scipionyx samniticus (Sasso & Signore 1998) recovered from Cretaceous strata of Italy validates said claims (Ruben et al 1999).
Ruben et al have relied exclusively on the data available from preserved soft-anatomy, minimizing osteological factors in their analyses (Ruben 1995, Ruben & Jones 2000). The soft anatomy of the type Sinosauropteryx displays an overall poor state of preservation, and thus data derived from this material is largely ambiguous until such time as soft-tissues are found preserved with greater fidelity in similar theropods (Martill et al 2000, Paul 2002).
Due to haphazard and unsupervised preparation of NGMC 2124 by amateurs, the slab was shattered symmetrically into at least twelve pieces (Ji & Ji 1996, Paul 2002), and subsequent restorative measures have largely obscured the original extent and form of the carbonized region (Paul 2002). Breakage and infilling have destroyed the original cranial aspect of the carbonized region, such that its form and the degree to which it extended cranially cannot be ascertained with certainty (Paul 2002). The dorsal, central, and ventral margins identified by Ruben et al (1997) as displaying a convex arc, in fact delineate a pseudo-margin resulting from the inadequate preparation of the specimen. Thus, the initial conclusion that the carbonized material displays a convex arc in cranial aspect is not unequivocally substantiated at this time.
Moreover, the liver of both birds and crocodiles, the two extant nodes of Archosauria, display convex cranial arcs, and the presence of such an arc in and of itself is not indicative of a pelvovisceral system (Duncker 1979, Brackenbury 1987, Paul 1988, Paul 2002). Ruben et al infer a post-pulmonary septum, in concordance with their argument that the carbonized material displays a convex arc in cranial aspect, however, the reason why such a septum must be present given a convex cranial arc, is not clear. Furthermore, the height of the liver (assuming the material represents as much) is not a reliable correlate of a pelvovisceral system. Liver anatomy is highly variable throughout the ontogeny of any given taxon and individual animal, and more germane, there is no consistent difference in the size and position of the liver in crocodiles, and birds (Siwe 1937, Brackenbury 1987, Secor & Diamond 1995, Paul 2002). Curiously, a post-pulmonary septum is observed in none of the remaining Sinosauropteryx material (Ackerman 1998, Chen et al 1998, Paul 2002).
Ruben et al (1997) attempted to mitigate these ambiguities with the soft-anatomy of Sinosauropteryx by arguing that preserved muscle fibers in the type Scipionyx were oriented in a position to be expected of diaphragmatic musculature. However, these fibers—if they are indeed muscle fibers—have subsequently been identified as components of M. obliquus and M rectus, muscles of the abdomini, by virtue of their configuration (Paul 2002).
Further doubt is cast on the conclusions pertaining to the aerobic capacity in Sinosauropteryx made by Ruben and his colleagues, by the soft anatomy of Scipionyx samniticus. In the type Scipionyx, the intestinal tract is preferentially preserved at the expense of other viscera, and a structure immediately caudal to the sternum, distal ends of the humeri and proximal blade of the scapula, is most logically associated with the liver (Sasso & Signore 1998, Ruben et al 1999, Paul 2002). Ironically, the cranial orientation of the liver in Scipionyx is entirely incongruent with the position of the alleged liver in Sinosauropteryx. The position of the intestines in Scipionyx and the carbonized matter in Sinosauropteryx are identical, and given these data, the most logical conclusion is that the latter represents the fossilized post-mortem decay of the intestinal tract (Paul 2002).
Thus, even offering Ruben’s research the benefit of the doubt (i.e., assuming they have identified the carbonized material in Sinosauropteryx correctly), the data presented in Ruben et al (1997, 1999), is not sine qua non verification of a pelvovisceral pump in Theropoda, in that the soft-anatomy is sufficiently ambiguous to render definitive conclusions based on such, wholly speculative at this time.
Given these limitations, the osteological data is currently the most reliable in determining the aerobic faculties of Theropoda. While Ruben (1995) and Ruben & Jones (2000) have argued that osteological factors are largely inconsequential to the operation of aerobic systems, there is no apparent reason why this should be so, and indeed the suite of characters which are biomechanical necessities for the respective systems of aerobic regulation, seem to directly refute this assertion.
Theropoda lack all the osteological modifications permitting the operation of a pelvovisceral pump, and in particular, lack the modification of the proximal heads of the ribs and the communication of the gastralia seen in crocodiles. In Theropoda, the thoracic ribs articulate with moderate or reduced transverse processes of the dorsal vertebrae via bifurcated proximal heads, and thus the dorsal surface of the rib cage in theropods is sharply corrugated (Ostrom 1969, Paul 1988, Britt 1997, Paul 2002). The gastralia meet medially in Theropoda, and were not imbedded within a cartilaginous sheet (Ostrom 1969, Paul 1988, Dodson et al 1990, Paul 2002). These characters would have precluded the formation of a sub-cylindrical, smooth-walled tube in which the viscera could be acted upon by diaphragmatic muscles (Paul 2002), and these data alone, demonstrate that the presence of a pelvovisceral system in Theropoda, was a biomechanical impossibility.
The absence of a lumbar region, inadequacy of the distal pubes to support diaphragmatic musculature and immobility of the pubes (especially in opisthopubic taxa, e.g. Eumaniraptora) further argue against the presence of such a hepatic piston, as is seen in crocodiles (contra Ruben et al).
To preclude the assertion of negative evidence, review of the osteological characters associated with avian or paravian aerobic capacities, which are, or are not present in Theropoda, is germane. Characters associated with the avian aerobic system include (after Zimmer 1935, Bellairs & Jenkin 1960, King 1966, Schmidt-Nielsen 1972, Olson 1973, Perry 1983, Beale 1985, Paul 1988, Perry 1989, McLelland 1989, Bramble & Jenkins 1998, Paul 2002):
a)Postcrania consistently pneumatic, infiltrated by pulmonary diverticula
b)Proximal thoracic rib heads bifurcated, forming a corrugated dorsal rib cage
c)Caudal ribs not reduced
d)Sternocostal hinge present
e)Sternal ribs are singular, not double, and usually ossified
f)Ossified uncinate processes
g)Gastralia meet medially
In Theropoda the postcrania are consistently pneumatized, displaying invasion of pulmonary diverticula. Pneumaticity of postcranial elements extends to the sacrum and caudal vertebrae in some Theropoda (Russell & Dong 1993, Britt 1997, Paul 2002), in contrast to the pattern expected had an airtight post-pulmonary septum existed in life. The proximal heads of the thoracic ribs, in even the most basal Theropoda, are bifurcated and articulate with the dorsal vertebrae via moderate transverse processes, as detailed above. The caudal ribs in Theropoda are not reduced, and consequently, a lumbar region is absent, as detailed above. The sterna and coracoids of avepectoran theropods (i.e., Maniraptoriformes), articulate via a hinge joint permitting sternal kinesis (Paul 1987, 1988, Norell & Makovicky 1999, Burnham et al 2000, Paul 2002). Contrary to Ostrom (1969), the sternal ribs of Maniraptoriformes are singular, and ossified (Barsbold 1983, Paul 1988, Ji et al 1998, Norell & Makovicky 1999, Xu, Wang & Wu 1999, Paul 2002). Ossified uncinate processes are present in derived maniraptoran taxa (Paul 1988, Xu, Wang & Wu, 1999, Norell & Makovicky 1999, Xu et al 2000, Burnham et al 2000, Paul 2002). Lastly, the gastralia of Theropoda meet medially, and were not set in a cartilaginous sheet, as in crocodiles, as detailed elsewhere.
It is thus concluded, that the osteology of Theropoda is inconsistent with the presence of a pelvovisceral pump, and at least in two regards, the operation of such a pump would have been a biomechanical impossibility. On the contrary, all the modifications of the thoracic and pectoral skeleton, as well as the vertebrae seen in Theropoda, are congruent with those observed in Avialae, arguing the presence of a similar aerobic system in at least some theropods. Considering the ambiguity of the soft-anatomy at hand, current analyses must rely on the osteological data until such time as better-preserved soft-tissues become available. These data directly contradict Sarfati’s comments in the interview in question.
2. Integumentary Structure of Maniraptoriformes
Evidence of feathery integument in Theropoda is solely restricted to the clade Maniraptoriformes. The first specimen to provide empirical data suggesting the presence of such integument in this taxon, was the type Sinosauropteryx prima (NGMC 2124) (Ji & Ji 1996), in which tubular integumentary structures are observed from the crown and nape, down the back, and onto the dorsal and ventral aspects of the tail. The integumentary structures also appear to be present on parts of the skull and along the antebrachial margin of the forelimb (Chen et al 1998, Feduccia 1999, Paul 2002). Multiple authors have asserted that these structures represent integumentary fibers, which are precursors to feathers sensu stricto (Ji & Ji 1996, 1997, Chen et al 1998, Griffiths 1998/2000, Paul 2002).
Although Geist et al (1997), Feduccia (1999), and Ruben & Jones (2000) have argued that the integumentary structures are collagenous, there has been no data to substantiate said conclusion presented. The inferred aquatic habitus of compsognathid coelurosaurs (Feduccia 1999) is entirely speculative and directly refuted by the osteology of known compsognathids. Furthermore, the suggestion that sub-dermal collagen fibers can account for the integumentary structures observed in Sinosauropteryx is irreconcilable with the examples of varanid lizards shown by Feduccia (1999, 378). The integumentary structures seen in Sinosauropteryx are most parsimoniously interpreted as keritanaceous (Ji & Ji 1996, 1997, Chen et al 1998, Paul 2002).
The emphasis placed upon the integument of Sinosauropteryx is curious, in that evidence of feathery integument is by no means limited to this theropod. Far more unambiguous are the structures preserved with high fidelity in Sinornithosaurus millenii, Beipiaosaurus inexpectus, Microraptor gui, Caudipteryx zhoui, and Protarchaeopteryx robusta (Ji & Ji 1997, Ackerman 1998, Ji et al 1998, Padian & Chiappe 1998, Xu, Tang & Wang 1999, Sloan 1999, Xu, Wang & Wu 1999, Xu et al 2000, Hoffman 2000, Zhou & Wang 2000, Xu et al 2001, Paul 2002, Currie 2003). The presence of feathery integument is inferred from the presence of ulnar quill knobs in both Rahonavis ostromi, and Unenlagia comahuensis (Novas 1997, Forster, Sampson, Chiappe & Krause, 1998). There has to date been no data presented to refute the presence of feathery integument in these taxa.
Therefore, Sarfati’s comments that there are no feathered Theropoda are lacking in substantiation.
3. Integumentary Derivatives and Exaptation Thereof
Arguments concerning the origin of feathery integument that do not take account of the developmental and molecular data, which has accrued in the past decade, are obsolete. Modern hypotheses advanced for modeling the origin and early development of feathers synthesize data gleaned from developmental processes and the pleiotropic interaction of regulatory genes, and proteins (Brush 1993, 1996, 2000, Ji et al 1998, Padian & Chiappe 1998, Prum 1999, Xu, Tang & Wang 1999, Xu, Wang & Wu 1999, Maderson & Alibardi 2000, Brush 2000, Harris, Fallon & Prum 2002, Prum & Brush 2002, 2003). Research into the developmental stages by which feathers are derived in modern birds had yielded a coherent, incremental developmental model by which the origin and early evolution of feathers can be quantified, and predicted (Prum & Brush 2002, 2003).
Specifically, the influence of regulatory gene Sonic Hedgehog (Shh) and Bone Morphogenetic Protein 2 (Bmp2) at specific times and in specific regions of the epidermal placodes are responsible for the helical growth patterns of the follicle collar and its subsequent differentiation into the rachis ridge, barbule plates, and barb ridges. These data have shown that contrary to classical arguments, feathers can be derived at their most basal level without direct aerodynamic impetus. In other words, the developmental model of feather origins is completely compatible with the exaptation of basal feathers to aerodynamic function (Brush 1993, 1996, 2000, Prum 1999, Brush 200, Harris, Fallon & Prum 2002, Prum & Brush 2002, 2003).
Considering that the most basal feathers are, following a developmental model, predicted to be hollow cylinders arising from epidermal placodes, the presence of such structures on multiple theropod taxa is astounding corroboration of said model (Paul 2002, Prum & Brush 2002, 2003).
Data can, and has, been produced to falsify the simplistic if cherished notion of feathers being “scales gone frayed,” and consequently this postulate is no longer at the forefront of research into the origin of feathers, and has not been seriously advanced since Regal’s study in 1975. The developmental model is also interesting in that it falsifies the objections that feathery integument should not be observed in theropods requiring a thermoregulatory coat, by demonstrating that the earliest derivatives of feathers would have been quite capable of insulatory function. Perhaps even more pertinent, is the ability of the developmental model to substantiate arguments that the integumentary structures of some of the most paravian maniraptorans are congruent with the pattern, which should be observed in neoflightless post-urvogels (Paul 2002).
Contrary to the assertions made by Sarfati in the AIG interview, there is no data to suggest that exaptation is not a viable explanation for the development of flight feathers.
Continuation presented in next post.
Vindex Urvogel