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Fixed cervical count and the origin of the mammalian diaphragm


  • Emily A. Buchholtz,

    Corresponding author
    • Department of Biological Sciences, Wellesley College, Wellesley, MA 02481, USA
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  • H. Grady Bailin,

  • Susan A. Laves,

  • Jennifer T. Yang,

  • Mei-Yee Chan,

  • Lucie E. Drozd

Author for correspondence (email:


Why is mammalian cervical count fixed across the historically long and ecologically broad mammalian radiation? Multiple lines of evidence, considered together, suggest a link between fixed cervical count and the muscularization of the diaphragm, a key innovation in mammalian history. We test this hypothesis by documenting the anteroposterior (AP) movement of the diaphragm, a lateral plate derivative, relative to that of the somitic thoracolumbar transition in unusually patterned mammals, by comparing the temporal occurrence of an osteological proxy for the diaphragm and fixed cervical counts in the fossil record, and by quantifying morphological differentiation within the mammalian cervical series. We then integrate these anatomical observations with details of diaphragm function and development to propose a sequence of innovations in mammalian evolution that could have led to fixed cervical count. We argue that the novel commitment of migratory muscle precursor cells (MMPs) of mid-cervical somites to a fate in the abaxial diaphragm defined these somites as a new and uniquely mammalian modular subunit. We further argue that the coordination of primaxial abaxial patterning constrained subsequent AP migration of the forelimb, thereby secondarily fixing cervical count.