Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends
Megalospheric specimens of Nummulitidae from eight localities in western and central Cuba were morphometrically investigated using test characters described by 11 growth-independent and growth-invariant attributes that provide a complete geometric reconstruction of nummulitid equatorial morphology. The species Nummulites striatoreticulatus, Palaeonummulites trinitatensis, Operculinoides floridensis and O. soldadensis were classified by an agglomerative cluster analysis. Discriminant analysis yielded significant morphological separators between the species such as the backbend angle, marginal radius increase, perimeter ratio and first chamber length. The transition of tightness to laxity of the spiral was an important morphological separator at the generic level, representing a clear general trend coupled with the change in palaeodepth. Based on further discriminant analysis, an increase in proloculus size was detected in Nummulites striatoreticulatus from the middle Eocene to early late Eocene, supporting this important evolutionary pattern in many lineages of Nummulites. Operculinid forms showed an opposite and more weakly pronounced time-dependent trend in the size decrease of the proloculus. In the Cuban localities, Nummulites striatoreticulatus occurs from the Lutetian to Priabonian, while Palaeonummulites trinitatensis is restricted to the Bartonian to Priabonian. The moderately to loosely coiled operculinid taxa O. floridensis and O. soldadensis have longer stratigraphical ranges from the middle Eocene to probably the early Oligocene. Operculinoides floridensis and O. soldadensis show a broader variability in marginal radius increase, and thus probably occupied wider niches than N. striatoreticulatus. The latter seems to be restricted to the shelf edge and to the shallowest parts of the upper slope. A possible phylogenetic connection between Heterostegina and Operculinoides is suggested by the closest equatorial morphology of Heterostegina sp. indet. to tightly coiled forms of Operculinoides floridensis. Discriminant analysis documents the strongest similarities in perimeter ratio, backbend angle, initial marginal radius and proloculus mean diameter.