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Mancallinae
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Mancallinae is an of prehistoric flightless that lived on the coast of today's and from the late epoch to the early (ranging from at least 7.4 million to 470,000 years ago). They are sometimes collectively referred to as Lucas auks after the scientist who described the first , Frederic Augustus Lucas.


Description
They had along somewhat similar lines as the , their counterpart, but their decidedly stubbier wings were in some aspects more convergent with . Compared with the great auk, they were also smaller ( see also: Bergmann's Rule): Praemancalla species have been estimated to have weighed about 3 kg. Most Mancalla forms weighed somewhat less (about 2.4 kg), with M. milleri being a smaller (1.65 kg) and M. emlongi a much larger bird (3.8 kg) than the rest. The last species thus stood around 55–60 cm high in life. The largest species, Miomancalla howardi, was the largest charadriiforme of all time.Smith, N. 2015. Evolution of body mass in the Pan-Alcidae (Aves, Charadriiformes): the effects of combining neontological and paleontological data. Paleobiology. doi: 10.1017/pab.2015.24


Evolution and classification
In a 2011 paper, N. Adam Smith published a review and revision of the Mancallinae. Smith defined Mancallinae as the containing Mancalla, Miomancalla, their common ancestor and all its descendants. A summary of his results follows.

  • Pan- Smith, 2011
    • Mancallinae L.Miller, 1946
      • Lucas, 1901
        • Mancalla californiensis Lucas, 1901
        • Mancalla cedrosensis Howard, 1971
        • Mancalla lucasi Smith, 2011
        • Mancalla vegrandis Smith, 2011
      • Smith, 2011
        • Miomancalla howardi (Smith, 2011)
        • Miomancalla wetmorei (Howard, 1976)

The following species could not be identified as unquestionable mancallines, and have been assigned to Pan-Alcidae incertae sedis. They may be mancallines, or more closely related to true .

  • Alcodes ulnulus Howard, 1968
  • Mancalla diegensis Miller, 1937
  • Mancalla milleri Howard, 1970

The following species could not be differentiated from other recognized species of mancallines (see above), and have been assigned to Mancallinae incertae sedis.

  • Mancalla emlongi Olson, 1981
  • Praemancalla lagunensis Howard, 1966

The mancallines probably evolved from proto-, which must have been birds not dissimilar to the rhinoceros auklet. Accordingly, their status as a subfamily has been questioned as this would make the Alcinae (true auks) . However, the mancallines were a very distinct and unique evolutionary lineage and are thus usually retained as a subfamily. They must have diverged from flying ancestors during the mid-Miocene, roughly 15 mya.

Alcodes is known from a single found in Late Miocene (, 9–12 mya) deposits at , . While assignment of such a fragmentary fossil is always problematical, the ulna is a fairly distinctive bone and that of Alcodes is quite peculiar. However, it is more allied with the Mancallines as a matter of convenience; additional material would be needed to confirm this relationship. From the bone's measurements, it seems probable that this species was flightless and judging from its age, it either represents an earlier development parallelling Mancalla, or a third lineage of flightless auks.

Praemancalla is known from Clarendonian to Early remains. It is similar to Mancalla, but less extreme in its adaptations and it is quite possibly that the latter evolved from one of the two known species. Mancalla was a common species throughout the Pliocene, appearing in the stage of the Late Miocene (5–9 mya), and spreading in the Pliocene, with 4 species apparently coexisting at one time on the coast of southern California.

As with many marine birds, the mancalline auks were much affected by the extinction crisis in the late Pliocene oceans. This coincided with the diversification of , but may ultimately have been caused by increased activity in the vicinity of the . Despite their apparent awkwardness, they seem to have been quite well adapted for flightless birds, with the fossil record suggesting that the last remnants did not disappear until the Early Pleistocene, some time after the ecological changes had passed their peak.

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