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Eggs

Evolutionary theories and controversies

Phylogenetic affinities

Family: Balaenicipitidae

Genus: Balaeniceps

Order: Pelecaniformes 

Controversies about shoebill's order

Shoebill was initially classified under the order Ciconiiformes due to its similar behavior to the other members of the Ciconiidae families (Buxton et al., 1978, cited in Muir & King 2012) like its bill-clattering display which resembles the Up-down greeting display by storks (Slikas 1998).

However, after several molecular studies, shoebill is now classified in the Pelecaniformes order.

 

Related studies:

1.  Mikhailov (1995) found that shoebill's eggshell microstructure is similar to that of Scopus and most Pelecaniformes birds.

2. Tuinen et al. (2001) analyzed the DNA-DNA hybridization data, nuclear and mitochondrial DNA sequences and found that the taxon [Scopidae + (Balaenicipitidae + Pelecanidae)] is monophyletic, meaning they are more closely related to each other than to other bird families.

3.  Another study revealed that the closest relative of shoebill is hamerkop (Scopus umbretta) from the Scopidae family and Pelecaniformes order. It was also found that shoebill and hamerkop have very similar skeletal structure, including a hooked premaxilla (Mayr 2003). 

4. Mayr (2008) noted that Shoebills have two distinct anatomical features that are exclusively shared by Pelecanidae and Scopidae

  • The presence of os palatinum and pars choanalis located deep in dorsoventral direction, with short ossa pterygoidea;

  • A small or completely absent musculus ambiens

 

These molecular studies collectively led to shoebill's reclassification within the Pelecaniformes order.

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Hamerkops, the closest relative to shoebill (Image credit to: Molenaar, Wild South Africa Kruger National Park)

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Eggshell structure in radial section showing (A) General pattern of eggshells, (B) Parallelism of increasing microglobular material concentration in Pelecaniformes and Ardeidae (Image credit to: Mikhailov, Canadian Journal of Zoology)

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Skull in comparison. A, Balaeniceps rex (Balaenicipitidae); B, Scopus umbretta (Scopidae). (Image credit to: Mayr,

 Journal für Ornithologie)

Evolutionary Theories

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Endo et. al theory

Endo et. al (2013) proposed that shoebills evolved to have large bill and enlarged pharynx to pass large food into its alimentary tract.

Additionally, they suggested that since shoebill's tongue is not functionally important in their feeding, it has degenerated over the course of shoebill's evolutionary history. This degenerate tongue creates more space in the oral cavity, aiding in the process of swallowing large food items.
 

Cracraft's theory

Cracraft (1985) theorized that the physical similarities between shoebill and Pelecaniformes were not due to a common evolutionary ancestry, but convergence evolution. He suggested that these resemblances in physical characteristics emerged as adaptations to shared feeding behavior.

Yet, Cracarft's theory does not apply to Scopus which, despite having different living and feeding habits from those of Balaeniceps and Pelecaniformes, still exhibit similar eggshell structure (del Hoyo et al. 1992)
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