Keuppia is in the family Palaeoctopodidae, and one of the earliest representatives of the order Octopoda. These ancient marine beauties are in the class Cephalopoda making them relatives of our modern octopus, squid and cuttlefish.
There are two species of Keuppia, Keuppia hyperbolaris and Keuppia levante, both of which we find as fossils. We find their remains, along with those of the genus Styletoctopus, in Cretaceous-age Hâqel and Hjoula localities in Lebanon.
For many years, Palaeoctopus newboldi (Woodward, 1896) from the Santonian limestones at Sâhel Aalma, Lebanon, was the only known pre‐Cenozoic coleoid cephalopod believed to have an unambiguous stem‐lineage representative of Octobrachia Fioroni.
With the unearthing of some extraordinary specimens with exquisite soft‐part preservation in the Lebanon limestones, our understanding of ancient octopus morphology has blossomed. The specimens are from the sub‐lithographical limestones of Hâqel and Hâdjoula, in north‐west Lebanon. These localities are about 15 km apart, 45 km away from Beirut and 15 km away from the coastal city of Jbail. Fuchs et al. put a nice little map in their 2009 paper that I've included and referenced here.
Palaeoctopus newboldi had a spherical mantle sac, a head‐mantle fusion, eight equal arms armed with suckers, an ink sac, a medially isolated shell vestige, and a pair of (sub‐) terminal fins. The bipartite shell vestige suggests that Palaeoctopus belongs to the octopod stem‐lineage, as the sister taxon of the Octopoda, the Cirroctopoda, is characterized by an unpaired clasp‐like shell vestige (Engeser 1988; Haas 2002; Bizikov 2004).
Again from the Santonian–Campanian of Canada and Japan, Tanabe et al. (2008) reported on at least four different jaw morphotypes. Two of them (Paleocirroteuthis haggarti Tanabe et al. , 2008 and Paleocirroteuthis pacifica Tanabe et al ., 2008) have been interpreted as being of cirroctopod type, one of octopod type, and one of uncertain octobrachiate type.
Interestingly Fuchs et al. have gone on to describe the second species of Palaeoctopus, the Turonian Palaeoctopus pelagicus from limestones at Vallecillo, Mexico. While more of this fauna will likely be recovered in time, their work is based solely on a medially isolated shell vestige.
Five new specimens have been found in the well-known Upper Cenomanian limestones at Hâqel and Hâdjoula in Lebanon that can be reliably placed within the Octopoda. Fuchs et al. described these exceptionally well‐preserved specimens and to discuss their morphology in the context of phylogeny and evolution in their 2008 paper (2009 publishing) in the Palaeontology Association Journal, Volume 51, Issue 1.
The presence of a gladius vestige in this genus shows a transition from squid to octopus in which the inner shell has divided in two in early forms to eventually be reduced to lateralized stylets, as can be seen in Styletoctopus.
The adorable fellow you see here with his remarkable soft-bodied preservation and inks sack and beak clearly visible is Keuppia levante. He hails from Late Cretaceous (Upper Cenomanian) limestone deposits near Hâdjoula, northwestern Lebanon. The vampyropod coleoid, Glyphiteuthis abisaadiorum n. sp., is also found at this locality. This specimen is in the collection of the deeply awesome David Appleton.
Fuchs, D.; Bracchi, G.; Weis, R. (2009). "New octopods (Cephalopoda: Coleoidea) from the Late Cretaceous (Upper Cenomanian) of Hâkel and Hâdjoula, Lebanon". Palaeontology. 52: 65–81. doi:10.1111/j.1475-4983.2008.00828.x.
Photo credit: David Appleton. Figure Two: Topographic map of north‐western Lebanon with the outcrop area in the upper right-hand corner. Fuchs et al, 2009.