Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends Author Torres-Silva, Ana. I. Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria; Author Eder, Wolfgang Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria; Author Hohenegger, Johann Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria; Author Briguglio, Antonino Dipartimento di Scienze della, Terra dell’Ambiente e della Vita, Universita ́ degli Studi di Genova, Corso Europa, 26, I- 16132 Genova, Italy text Journal of Systematic Palaeontology 2018 2018-04-13 17 7 557 595 http://dx.doi.org/10.1080/14772019.2018.1446462 journal article 10.1080/14772019.2018.1446462 1478-0941 PMC6474738 31057335 10883523 Operculinoides soldadensis Vaughan & Cole, 1941 ( Fig.17A–G ) 1941 Operculinoides soldadensis Vaughan & Cole : 18, pl. 9, figs 5–8, pl. 10, figs 1, 2. 1947 Nummulites ( Operculinoides ) floridensis (Vaughan & Cole) ; de Cizancourt: 517, pl. 25, figs 8–10, 13. 1961 Nummulites floridensis Heilprin ; Butterlin: 12, figs 5–6. 1975 Operculinoides soldadensis Vaughan & Cole ; Caudri: 537, pl. 1, fig. 11, pl. 8, figs 5–8, 10. 1996 Operculinoides suteri Caudri ; Caudri: 1189, pl. 10, fig. 9. Material. Twenty-five megalospheric specimens in equatorial section, comprising five from Loma Candelaria (98LC-1), four from Loma El Santo (CA-215), one from Loma Jabaco (LM-52) and 15 from Norona ( NOR-UN ). Description. External features. Test planispiral, flattened, last whorl fragile and laterally compressed, involute in the nepionic stage, becoming evolute in the last whorl. The prominent central umbo is surrounded by slightly raised septal sutures. Internal features. Megalospheric generation with spherical proloculus with a mean diameter of 0.09 mm, followed by reniform deuteroloculus and a loosely coiled spiral with commonly two to three whorls. Rapid increase in height of the last spiral with chamber height roughly 4– 5 times higher than chamber width. Primary operculine septa with strong backbend angle gently tapered towards inner ends and with septal undulations. A diagnostic characteristic are the numerous and narrow chambers. Characters and attributes (means and standard deviations) for Operculinoides soldadensis and comparisons with Nummulites striatoreticulatus , Palaeonummulites trinitatensis , Operculinoides floridensis (tightly coiled) and O. floridensis (loosely coiled) are given in Table 9 . Occurrences. Middle late Eocene to late Eocene, NP 16/ NP17, upper part of the Loma Candela Formation; late middle Eocene, CNE 13, Arroyo Blanco Formation;?early Oligocene O1/P18 and NP 21 /CP 16, Jabaco Formation. Remarks. Cole (1958) considered O. soldadensis to be synonymous with O. floridensis ; however, our morphometric analysis based on growth-independent and growth-invariant characters clearly distinguished the two species ( Table 3 ): Operculinoides soldadensis shows fewer morphological variations (ecophenotypes) at distinct depositional gradients than O. floridensis . Table 9. Characters and attributes (means and standard deviations, SD, in mm) for Operculinoides soldadensis and comparisons with Nummulites striatoreticulatus , Palaeonummulites trinitatensis , Operculinoides floridensis (tightly coiled) and O. floridensis (loosely coiled). Symbol key: ++, strong positive differences with <1% error probability; 0, no significant differences; ――, strong negative differences with <1% error probability.

O. soldadensis	Mean	SD	O. floridensis (tightly coiled)	O. floridensis (loosely coiled)	P. trinitatensis	N. striatoreticulatus
First chamber length	252.0	92.62	――	――	0	――
Proloculus nominal diameter	92.2	29.93	――	――	0	――
Deuteroloculus ratio	1.191	0.1834	0	0	+	++
Initial marginal radius	116.9	44.90	――	――	0	――
Marginal radius increase	0.133	0.0193	++	0	++	++
Spiral chamber height increase	2.0	0.96	――	0	0	――
Initial spiral chamber height	38.5	15.47	――	――	0	――
Backbend angle	0.687	0.1008	++	0	++	++
Initial chamber length	129.1	43.24	――	――	0	――
Chamber length increase	0.060	0.0202	++	0	++	++
Perimeter ratio	1.448	0.0809	++	0	++	++

Stratigraphical and geographic dialstribution. Middle to late Eocene (Lutetian to Priabonian); Cuba Trinidad , Mexico .