Forelimb Joint Mobility and the Evolution of Wing-Propelled Diving in Birds

Raikow, Robert J.; Bicanovsky, Lesley; Bledsoe, Anthony H.

doi:10.1093/auk/105.3.446

Cited by 56 publications

(54 citation statements)

References 7 publications

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“…Among Pan-Alcidae volant species have lower forelimb RBT values (A. torda forelimb RBT 5 26%) and higher values of wing flexion (see Storer, 1960, Fig. 4;Raikow et al, 1988;Smith, 2011b, Fig. 16).…”

Section: Osteohistological Correlates Of Flightlessness In Pan-alcidaementioning

confidence: 99%

“…It is seen in pan-alcids (Charadriiformes), penguins (Sphenisciformes), diving petrels and some shearwaters (Procellariiformes) and dippers (Passeriformes). Studies of osteological morphology, gross myology, neuroanatomy and biomechanics have all documented anatomical modifications associated with wing-propelled diving (Stettenheim, 1959;Storer, 1960;Spring, 1971;Schreiweis, 1982;Pennycuick, 1987;Baldwin, 1988;Livezey, 1988;Raikow et al, 1988;Bannasch, 1994;Habib and Ruff, 2008;Habib, 2010;Smith, 2011a;Smith and Clarke, 2012). However, histological investigations of the derived locomotor strategies of wing-propelled diving birds are quite limited.…”

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confidence: 99%

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Osteological Histology of the Pan‐Alcidae (Aves, Charadriiformes): Correlates of Wing‐Propelled Diving and Flightlessness

Smith

Clarke

2013

The Anatomical Record

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Although studies of osteological morphology, gross myology, myological histology, neuroanatomy, and wing-scaling have all documented anatomical modifications associated with wing-propelled diving, the osteohistological study of this highly derived method of locomotion has been limited to penguins. Herein we present the first osteohistological study of the derived forelimbs and hind limbs of wing-propelled diving Pan-Alcidae (Aves, Charadriiformes). In addition to detailing differences between wing-propelled diving charadriiforms and nondiving charadriiforms, microstructural modifications to the humeri, ulnae and femora of extinct flightless pan-alcids are contrasted with those of volant alcids. Histological thin-sections of four species of pan-alcids (Alca torda, †Alca grandis, †Pinguinus impennis, †Mancalla cedrosensis) and one outgroup charadriiform (Stercorarius longicaudus) were compared. The forelimb bones of wing-propelled diving charadriiforms were found to have significantly thicker ( 22%) cortical bone walls. Additionally, as in penguins, the forelimbs of flightless pan-alcids are found to be osteosclerotic. However, unlike the pattern documented in penguins that display thickened cortices in both forelimbs and hind limbs, the forelimb and hind limb elements of pan-alcids display contrasting microstructural morphologies with thickened forelimb cortices and relatively thinner femoral cortices. Additionally, the identification of medullary bone in the sampled †Pinguinus impennis specimen suggests that further osteohistological investigation could provide an answer to longstanding questions regarding sexual dimorphism of Great Auks. Finally, these results suggest that it is possible to discern volant from flightless wing-propelled divers from fragmentary fossil remains.

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Section: Osteohistological Correlates Of Flightlessness In Pan-alcidaementioning

confidence: 99%

mentioning

confidence: 99%

Osteological Histology of the Pan‐Alcidae (Aves, Charadriiformes): Correlates of Wing‐Propelled Diving and Flightlessness

Smith

Clarke

2013

The Anatomical Record

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“…2g. Forelimb swimmers (Cerorhinca monocerata, Uria aalge, Spheniscus magellanicus): Forelimb-propelled diving has evolved at least five times in birds, specifically in the penguins (Spheniscidae), auks (Alcidae), diving-petrels (Pelecanoididae), dippers (Cinclidae) and the extinct Plotopteridae (Raikow, Bicanovsky, & Bledsoe, 1988). Of these, those represented here are two auks (the rhinoceros auklet and common murre) and the magellanic penguin.…”

Section: F Marine Soaring Specialists (Diomedea Exulans Puffinus Gmentioning

confidence: 99%

The effects of locomotion on the structural characteristics of avian limb bones

Habib¹,

Ruff

2008

Zoological Journal of the Linnean Society

113

142

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Despite the wide range of locomotor adaptations in birds, little detailed attention has been given to the relationships between the quantitative structural characteristics of avian limb bones and bird behaviour. Possible differences in forelimb relative to hindlimb strength across species have been especially neglected. We generated cross-sectional, geometric data from peripheral quantitative computed tomography scans of the humerus and femur of 127 avian skeletons, representing 15 species of extant birds in 13 families. The sample includes terrestrial runners, arboreal perchers, hindlimb-propelled divers, forelimb-propelled divers and dynamic soarers. The hindlimb-propelled diving class includes a recently flightless island form. Our results demonstrate that locomotor dynamics can be differentiated in most cases based on cross-sectional properties, and that structural proportions are often more informative than bone length proportions for determining behaviour and locomotion. Recently flightless forms, for example, are more easily distinguished using structural ratios than using length ratios. A proper phylogenetic context is important for correctly interpreting structural characteristics, especially for recently flightless forms. Some of the most extreme adaptations to mechanical loading are seen in aquatic forms. Penguins have forelimbs adapted to very high loads. Aquatic species differ from non-aquatic species on the basis of relative cortical thickness. The combination of bone structural strength and relative cortical area of the humerus successfully differentiates all of our locomotor groups. The methods used in this study are highly applicable to fossil taxa, for which morphology is known but behaviour is not. The use of bone structural characteristics is particularly useful in palaeontology not only because it generates strong signals for many locomotor guilds, but also because analysing such traits does not require knowledge of body mass, which can be difficult to estimate reliably for fossil taxa.

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“…Penguins generate thrust during both the downstroke and upstroke (Clark and Bemis, 1979). Raikow et al (1988) measured joint mobility in the wings of wingpropelled divers as well as non-diving birds and found great reduction in the range of motion of the intrinsic wing joints of penguins compared with other groups, resulting in a degree of stiffening of the wing incompatible with aerial flight. The ulnar condyle in extant penguins is reduced to a nearly flat articular surface and the humerus is tightly bounded to the ulna and radius by a set of ligaments.…”

Section: Evolution Of the Penguin Flippermentioning

confidence: 99%

The phylogeny of the living and fossil Sphenisciformes (penguins)

Ksepka¹,

Bertelli²,

Giannini

2006

Cladistics

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We present the first phylogenetic analysis of the Sphenisciformes that extensively samples fossil taxa. Combined analysis of 181 morphological characters and sequence fragments from mitochondrial and nuclear genes (12S, 16S, COI, cytochrome b, RAG-1) yields a largely resolved tree. Two species of the New Zealand Waimanu form a trichotomy with all other penguins in our result. The much discussed giant penguins Anthropornis and Pachydyptes are placed in two clades near the base of the tree. Stratigraphic and phylogenetic evidence suggest that some lineages of penguins attained very large body size rapidly and early in the clade's evolutionary history. The only fossil taxa that fall inside the crown clade Spheniscidae are fossil species assigned to the genus Spheniscus. Thus, extant penguin diversity is more accurately viewed as the product of a successful radiation of derived taxa than as an assemblage of survivors belonging to numerous lineages. The success of the Spheniscidae may be due to novel feeding adaptations and a more derived flipper apparatus. We offer a biogeographical scenario for penguins that incorporates fossil distributions and paleogeographic reconstructions of the Southern continent's positions. Our results do not support an expansion of the Spheniscidae from a cooling Continental Antarctica, but instead suggest those species that currently breed in that area are the descendants of colonizers from the Subantarctic. Many important divergence events in the clade Spheniscidae can instead be explained by dispersal along the paths of major ocean currents and the emergence of new islands due to tectonic events.

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Forelimb Joint Mobility and the Evolution of Wing-Propelled Diving in Birds

Cited by 56 publications

References 7 publications

Osteological Histology of the Pan‐Alcidae (Aves, Charadriiformes): Correlates of Wing‐Propelled Diving and Flightlessness

Osteological Histology of the Pan‐Alcidae (Aves, Charadriiformes): Correlates of Wing‐Propelled Diving and Flightlessness

The effects of locomotion on the structural characteristics of avian limb bones

The phylogeny of the living and fossil Sphenisciformes (penguins)

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