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A CT-based Survey of Supramedullary
Diverticula in Extant BirdsDr. Jessie Atterholt1,2 & Dr. Mathew Wedel1
1Western University of Health Sciences, Pomona, CA, USA2Raymond M. Alf Museum of Paleontology, Claremont, CA, USA
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(Müller, 1908)
lungs & trachea air sacs & diverticula
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(Müller, 1908)
cervical air sacs
abdominal air sacs
lungs
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canalis intertransversarius (Müller, 1908)
diverticula supervertebralia
(Müller, 1908)
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Supramedullary Airways: • diverticula branching from lungs & air sacs• inside vertebral canal• above spinal cord
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vertebralcanal
spinal cord
supramedullaryairways
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vertebralcanal
spinal cord
supramedullaryairways
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vertebralcanal
spinal cord
supramedullaryairways
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Research on Supramedullary Airways
Cover (1953) — turkey
Müller (1908) — pigeon
Bezuidenhout et al. (1999) — ostrich O’Connor (2006) — general description
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Research on Supramedullary Airways
Cover (1953) — turkeyMüller (1908) — pigeon
Bezuidenhout et al. (1999) — ostrich O’Connor (2006) — general description
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Prior Phylogenetic Sampling
Struthioniformes
GalliiformesAnseriformes
Charadriiformes Rallidae
Pelecaniformes
Procellariiformes
Gruiformes
Columbiformes Pteroclidiformes
CuculiformesMusophagiformes
CaprimulgiformesApodiformes
Strigiformes CathartidaeAccipitridae
Falconiformes
Psittaciformes
ColiiformesTrogoniformes
PiciformesCoraciiformes
Passeriformes
2 major clades
2 taxa
Combined analysis tree(Atterholt & Matzke, 2011)
GaviiformesPodicipediformes
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Broad Phylogenetic Sampling
Struthioniformes
GalliiformesAnseriformes
Charadriiformes Rallidae
Pelecaniformes
Procellariiformes
Gruiformes
Columbiformes Pteroclidiformes
CuculiformesMusophagiformes
CaprimulgiformesApodiformes
Strigiformes CathartidaeAccipitridae
Falconiformes
Psittaciformes
ColiiformesTrogoniformes
PiciformesCoraciiformes
Passeriformes
18 major clades
26 genera
Combined analysis tree(Atterholt & Matzke, 2011)
GaviiformesPodicipediformes
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Anas platyrhynchos(Anseriformes)
Bubo virginianus(Strigiformes)
Calypte anna (Apodiformes)
Meleagris gallopavo (Galliformes)
Puffinus griseus (Procellariiformes)
Zenaida macroura (Columbiformes)
Supramedullary Airways Present
Struthio camelus (Struthioniformes)
Pyrrhura molinae Eclectus roratus(Psittaciformes)
Aphelocoma californicaMelozone crisallis Psaltriparus minimus Turdus migratorius(Passeriformes)
Musophaga violacea (Musophagiformes)
Buteo jamaicensis Cathartes aura(Accipitriformes)
Larus occidentalisUria aalge(Charadriiformes)
Nycticorax nycticoraxPelecanus occidentalisPhalacrocorax penicillatus(Pelicaniformes)
Falco sparverius (Falconiformes)
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Supramedullary Airways Absent
Aechmorphus occidentalis(Podicipediformes)
Dryobates nuttallii (Piciformes)
Melanerpes formicivorus (Piciformes)
Gavia pacifica (Gaviiformes)
Gavia immer (Gaviiformes)
Gavia adamsii (Gaviiformes)
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Supramedullary Airways Absent
Aechmorphus occidentalis(Podicipediformes)
Gavia pacifica (Gaviiformes)
diving taxa
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Supramedullary Airways Absent
Dryobates nuttallii (Piciformes)
Melanerpes formicivorus (Piciformes)
???
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Supramedullary Airways Absent
Aechmorphus occidentalis(Podicipediformes)
Gavia pacifica (Gaviiformes)
Diving Taxa
Phalacrocorax penicillatus (Suliformes)
Supramedullary Airways Present
Uria aalge (Charadriiformes)
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Presence and Absence of SMAs
Struthioniformes
GalliiformesAnseriformes
Charadriiformes Rallidae
Pelecaniformes
Procellariiformes
Gruiformes
Columbiformes Pteroclidiformes
CuculiformesMusophagiformes
CaprimulgiformesApodiformes
Strigiformes CathartidaeAccipitridae
Falconiformes
Psittaciformes
ColiiformesTrogoniformes
PiciformesCoraciiformes
Passeriformes
18 major clades
26 taxa
Combined analysis tree(Atterholt & Matzke, 2011)
GaviiformesPodicipediformes
Phylogenetic Signal
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Morphology of Supramedullary Airways
anteriorview
dorsal view
spinalcord
anterior posterior
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Morphology of Supramedullary Airways
canalis intertransversarius only(no SMAs)
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Morphology of Supramedullary Airways
A. supramedullary airwaysat intervertebral joints
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Morphology of Supramedullary Airways
B. supramedullary airwaysdiscontinuous throughvertebral canals
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Morphology of Supramedullary Airways
C. supramedullary airwayscontinuous throughvertebral canals
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Morphology of Supramedullary Airways
D. supramedullary airwaysmerge with diverticula supervertebralia
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Morphology of Supramedullary Airways
variation across vertebral column is universal
A B C D
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Morphology of Supramedullary Airways
ostrich
2 dorsal tubes
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Morphology of Supramedullary Airways
bushtit
2 lateral tubes
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Morphology of Supramedullary Airwaysostrich
3 dorsal tubes
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Morphology of Supramedullary Airwayspelican
spinal cord jacketed by SMAs
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ostrich
Variation with Body Size
pelican
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Variation with Body Size?
bushtit turkey vulture
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Variation Across Vertebral Regions
cervical vertebraepresent: 22 genera
absent: 0 genera
red-tailed hawk
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Variation Across Vertebral Regions
dorsal vertebraepresent: 13 generaabsent: 9 genera
green-cheeked conure
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Variation Across Vertebral Regions
synsacrum*present: 5 generaabsent: 17 genera
*in posterior sacrals only
great-horned owl
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Variation Across Vertebral Regions
caudal vertebraepresent: 3 generaabsent: 19 genera
pelican
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Conclusions: Variation in Supramedullary Airways
1. phylogenetic position 2. body size3. vertebral region
Dryobates nuttallii (Piciformes)
Melanerpes formicivorus (Piciformes)
Gavia pacifica (Gaviiformes)
absent
present
Strong phylogenetic
signal
Pyrrhura molinae Eclectus roratus(Psittaciformes)
Buteo jamaicensis Cathartes aura(Accipitriformes)
Larus occidentalisUria aalge(Charadriiformes)
Aphelocoma californicaMelozone crisallis Psaltriparus minimus Turdus migratorius(Passeriformes)
Nycticorax nycticoraxPelecanus occidentalis
Phalacrocorax penicillatus
(Pelicaniformes)
Gavia immer (Gaviiformes)
Gavia adamsii (Gaviiformes)
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1. phylogenetic position2. body size? 3. vertebral region
Conclusions: Variation in Supramedullary Airways
bushtit turkey vulture
Future studies: quantify volume of SMAs and spinal cord
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Conclusions: Variation in Supramedullary Airways
Function influences distribution of SMAs along vertebral column?more movement—> large SMAs
restricted movement —> small (absent) SMAs
% g
ener
a pr
esen
t
0
25
50
75
100
vertebral region
cervical dorsal sacral caudal
1. phylogenetic position2. (body size)3. vertebral region
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Conclusions: Defining “Supramedullary Airways”
• diverticula branching from lungs & air sacs• inside vertebral canal• dorsal to spinal cord
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Conclusions: Defining “Supramedullary Airways”
• diverticula branching from lungs & air sacs• inside vertebral canal or at intervertebral joints• dorsal to spinal cord
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Conclusions: Defining “Supramedullary Airways”
• diverticula branching from lungs & air sacs• inside vertebral canal or at intervertebral joints• dorsal to spinal cord or otherwise in contact
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Conclusions: Paramedullary Diverticula
• diverticula branching from lungs & air sacs• inside vertebral canal or at intervertebral joints• dorsal to spinal cord or otherwise in contact
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Osteological Correlates
Phoebastria nigripes LACM 115139
Cervical 9
length 20mm
PelecanuserythrorhynchusLACM86262Cervical13
length40mm
foramina sculpting/fossae
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pelican
foramen in floor of vertebral canal
Osteological Correlates
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Osteological Correlates
(Schwarz & Fritsch, 2006: Fig. 4)
foramina sculpting/fossae
Giraffatitan Alamosaursus
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Conclusions: Paramedullary Airways in Other Archosaurs?
Illustration by Brian Engh (dontmesswithdinosaurs.com)
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Conclusions: Paramedullary Airways in Other Archosaurs?
Photo: Dave Hone Pteranodon (LACM 51760)
foramina in floor of vertebral canal
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Illustration by Brian Engh (dontmesswithdinosaurs.com)
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