Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends

Figures & data

Figure 1. A, schematic tectonic map of western and central Cuba (after Iturrlade-Vinent 1994), with locations of the stratigraphical sections and samples. B, stratigraphical relations of Eocene units in western and central Cuba, slightly modified from García-Delgado & Torres-Silva (1997); stratigraphical ranges of the studied sections: A, 98LC-2; B, 98LC-1; C, LM-52; D, NOR-UN; E, 98MT-1; F, E-126; G, CA-215.

Figure 2. Measurements of characters in equatorial section. A, embryonic apparatus; B, marginal test spiral; C, chamber measurements; D, example individual, Operculinoides floridensis, specimen 98LC-1H-648.

Table 1. Growth-independent and growth-invariant characters.

Character		Attribute	Code	Equation	Growth
Proloculus	1	Proloculus mean diameter	PD	$PD =\frac{PW+PH}{2}$	Independent
Deuteroloculus	2	Deuteroloculus ratio	DR	$DR =\frac{DW}{PW}$	Independent
First chamber	3	First chamber length	FCL	Directly measured	Independent
Marginal test spiral	4	Initial radius	IMR	$MR = IMR\cdot e^{MRInc\theta }$	Independent
RL	5	Radius increase	MRinc		Invariant
Spiral chamber height	6	Initial spiral chamber height	ICH	$CH = CHinc\cdot j+ICHex$	Independent
CH	7	Spiral chamber height increase	CHinc		Invariant
Backbend angle BBA	8	Backbend angle	BBA	$BBA =(1/n )\sum _{j = 1}^{j = n}{BBA}_j$	Invariant
Chamber lengths	9	Initial final chamber length	ICL	$CL = ICL\cdot e^{CLinc}$	Independent
	10	Final chamber length increase	CLinc		Invariant
Perimeter ratio PerR	11	Perimeter ratio of chambers	PerR	$PerR =(1/n )\sum _{j = 1}^{j = n}{PerR}_j$	Invariant

Figure 3. A, two-dimensional ordination of studied specimens; colours accord with the results of the K-means cluster analysis. Numbers indicate the measured type material: 1, Nummulites stritoreticulatus, holotype; 2, N. macgillavry (from Butterlin 1981); 3, Operculinoides trinitatensis, holotype; 4, O. spiralis, holotype; 5. O. kugleri, holotype; 6, N. trinitatensis (from Butterlin 1961); 7, O. willcoxi (from Barker 1939); 8, O. willcoxi (from Cole 1941); 9, O. floridensis (from Frost & Langenheim 1974); 10, O. floridensis (from Cole 1941); 11, O. floridensis (from Cole 1941); 12, O. soldadensis (from Vaughan & Cole 1941); 13, O. suteri (from Caudri 1996); 14, N. floridensis (from Butterlin 1961). B, three-dimensional ordination of the studied specimens emphasizes the variation in the third component, highlighting the differentiation between Nummulites from 98LC-2 and Palaeonummulites from 98LC-1. C, discriminant analysis between the interpreted species: Nummulites striatoreticulatus, Palaeonummulites trinitatensis, Operculinoides floridensis and Operculinoides soldadensis; parameters are sorted in order of their importance as discriminators.

Figure 4. Discriminant analysis of nummulitid species, where the important discriminators are ranked along the discriminant functions. A, discriminant analysis between Operculinoides and Palaeonummulites species; B, discriminant analysis within Nummulites striatoreticulatus from different localities; C, discriminant analysis within O. floridensis from different localities; D, discriminant analysis within O. soldadensis from different localities.

Figure 5. Ordinations and discriminant analysis. A, two-dimensional ordination of studied specimens; genera are separated by different shapes (squares = Operculinoides; polygons = Palaeonummulites; triangles = Heterostegina). B, three-dimensional ordination of the studied specimens emphasizes the variation in the third component, highlighting the differentiation between Heterostegina sp. indet. and Operculinoides. C, discriminant analysis of Heterostegina species and Operculinoides or Palaeonummulites species; parameters are sorted in order of their importance as discriminators.

Figure 6. Stratigraphical ranges of the nummulitid species from the Cuban sections and their correlation with the standard planktonic zones. A, Pearson et al. (2006); B, Berggren et al. (1995); C, Martini (1971); D, Agnini et al. (2014).

Figure 7. A, schematic diagram showing the Eocene depositional environments in sections from western and central Cuba (modified from Cotton 2012). B, schematic diagram showing depth zonation of nummulitid species and larger benthic foraminifera (LBF) present in the Eocene section across the depositional gradient (modified from Beavington-Penney & Racey 2004).

Figure 8. Palaeogeographical distribution of the Eocene nummulitid species found at the Cuban localities. Map adapted from Pindell (2009).

Figure 9. Distribution of larger benthic foraminifera (LBF) in the Angelita Quarry section, western Cuba.

Figure 10. Distribution of larger benthic foraminifera (LBF) in the Entronque de Herradura section, western Cuba.

Figure 11. Distribution of larger benthic foraminifera (LBF) in the Loma Candelaria section, western Cuba (modified from Torres-Silva et al. 2017).

Figure 12. Distribution of larger benthic foraminifera (LBF) in the Loma El Santo section, central Cuba.

Figure 13. Distribution of larger benthic foraminifera (LBF) in the Noroña section, western Cuba (modified from Torres-Silva et al. 2017).

Figure 14. Distribution of larger benthic foraminifera (LBF) in the Loma Vigía section, central Cuba.

Table 2. Characters and attributes (means and standard deviations, SD, in mm) for Nummulites and comparisons with Palaeonummulites and Operculinoides. Symbol key: ++, strong positive differences with < 1% error probability; +, differences with < 5% error probability; 0, no significant differences; −−, strong negative differences with < 1% error probability.

Nummulites	Mean	SD	Operculinoides	Palaeonummulites
First chamber length	448.7	220.50	0	++
Proloculus nominal diameter	334.2	152.84	++	++
Deuteroloculus ratio	0.999	0.1297	−−	0
Initial marginal radius	355.9	143.51	++	++
Marginal radius increase	0.062	0.0081	−−	0
Spiral chamber height increase	3.8	1.47	++	++
Initial spiral chamber height	111.0	58.49	++	++
Backbend angle	0.164	0.0513	−−	+
Initial chamber length	246.6	125.21	0	++
Chamber length increase	0.009	0.0065	−−	0
Perimeter ratio	1.101	0.0595	−−	0

Figure 15. Nummulites striatoreticulatus Rutten. A–C, Entronque de Herradura; A, 98LC-2-686; B, 98LC-2-687; C, 98LC-2-1a. D–F, Loma Candelaria; D, 98LC-1-660; E, 98LC-1-630; F, 98LC-1-806. G–K, La Esperanza; G, E-126-474; H, E-126-466; I, E-126-458; J, E-126-470, gaps in the septa between adjacent alar prolongations of the chambers; K, E-126-459; L, M, Loma El Santo; L, CA-215- 865; M, CA-215- 65. A, B, E, F, H, I, L and M are A forms in equatorial section; C, D, G and J are A forms in axial section.

Table 3. Characters and attributes (means and standard deviations, SD, in mm) for Nummulites striatoreticulatus and comparisons with Palaeonummulites trinitatensis, Operculinoides floridensis (tightly coiled), O. floridensis (loosely coiled) and O. soldadensis. Symbol key: ++, strong positive differences with < 1% error probability; +, differences with < 5% error probability; 0, no significant differences; -, negative differences with < 5% error probability; −−, strong negative differences with < 1% error probability.

N. striatoreticulatus	Mean	SD	O. floridensis (tightly coiled)	O. floridensis (loosely coiled)	O. soldadensis	P. trinitatensis
First chamber length	454.7	221.33	0	−−	++	++
Proloculus nominal diameter	335.0	151.10	++	++	++	++
Deuteroloculus ratio	1.003	0.1318	0	-	−−	0
Initial marginal radius	360.2	144.63	++	0	++	++
Marginal radius increase	0.062	0.0080	−−	−−	−−	−−
Spiral chamber height increase	3.8	1.45	0	0	++	++
Initial spiral chamber height	112.9	59.18	+	0	++	++
Backbend angle	0.167	0.0527	−−	−−	−−	−−
Initial chamber length	253.2	131.05	0	−−	++	++
Chamber length increase	0.009	0.0064	−−	−−	−−	0
Perimeter ratio	1.102	0.0591	−−	−−	−−	−−

Table 4. Characters and attributes (means and standard deviations, SD, in mm) for Palaeoummulites and comparisons with Nummulites and Operculinoides. Symbol key: +, differences with < 5% error probability; 0, no significant differences; –, strong negative differences with < 1% error probability.

Palaeonummulites	Mean	SD	Operculinoides	Nummulites
First chamber length	243.3	38.59	−−	−−
Proloculus nominal diameter	113.0	25.84	0	−−
Deuteroloculus ratio	0.888	0.1258	−−	0
Initial marginal radius	140.1	28.76	0	−−
Marginal radius increase	0.094	0.0195	0	0
Spiral chamber height increase	1.7	0.07	−−	−−
Initial spiral chamber height	46.5	10.02	0	−−
Backbend angle	0.495	0.1320	0	+
Initial chamber length	139.0	17.17	−−	−−
Chamber length increase	0.028	0.0175	0	0
Perimeter ratio	1.277	0.1076	0	0

Table 5. Characters and attributes (means and standard deviations, SD, in mm) for Palaeonummulites trinitatensis and comparisons with Nummulites striatoreticulatus, Operculinoides floridensis (tightly coiled), O. floridensis (loosely coiled) and O. soldadensis. Symbol key: ++, strong positive differences with < 1% error probability; +, differences with < 5% error probability; 0, no significant differences; -, negative differences with < 5% error probability; −−, strong negative differences with < 1% error probability.

P. trinitatensis	Mean	SD	O. floridensis (tightly coiled)	O. floridensis (loosely coiled)	O. soldadensis	N. striatoreticulatus
First chamber length	218.2	53.81	−−	−−	0	−−
Proloculus nominal diameter	99.0	28.91	−−	−−	0	−−
Deuteroloculus ratio	0.950	0.1376	0	−−	-	0
Initial marginal radius	126.8	32.48	−−	−−	0	−−
Marginal radius increase	0.099	0.0172	0	−−	−−	++
Spiral chamber height increase	1.7	0.30	−−	0	0	−−
Initial spiral chamber height	35.5	17.71	−−	−−	0	−−
Backbend angle	0.445	0.1165	0	-	−−	++
Initial chamber length	151.9	26.76	−−	−−	0	−−
Chamber length increase	0.025	0.0149	0	0	−−	0
Perimeter ratio	1.263	0.0859	0	-	−−	+

Table 6. Characters and attributes (means and standard deviations, SD, in mm) for Operculinoides and comparisons with Nummulites and Palaeoummulites. Symbol key: ++, strong positive differences with < 1% error probability; 0, no significant differences; −−, strong negative differences with < 1% error probability.

Operculinoides	Mean	SD	Palaeonummulites	Nummulites
First chamber length	369.0	165.69	++	0
Proloculus nominal diameter	146.3	67.08	0	−−
Deuteroloculus ratio	1.166	0.1684	++	++
Initial marginal radius	192.0	88.59	0	−−
Marginal radius increase	0.126	0.0191	0	++
Spiral chamber height increase	2.8	1.15	++	−−
Initial spiral chamber height	62.5	32.84	0	−−
Backbend angle	0.636	0.1120	0	++
Initial chamber length	224.0	126.60	++	0
Chamber length increase	0.048	0.0220	0	++
Perimeter ratio	1.392	0.1130	0	++

Figure 16. Operculinoides floridensis (Heilprin). A–C, Loma Candelaria; A, 98LC-1-651; B, 98LC-1-667; C, 98LC-1-815. D, E, Loma Vigía; D, CA-216-F3-16; E, CA-216-D1a. F, Loma El Santo, CA-215-852. G, Loma Jabaco, LM-52-759. H, Angelita Quarry, 98MT-1. A–D, F and G are A forms in equatorial section; E and F are A forms in axial section.

Table 7. Characters and attributes (means and standard deviations, SD, in mm) for Operculinoides floridensis (tightly coiled) and comparisons with Nummulites striatoreticulatus, Palaeonummulites trinitatensis, O. floridensis (loosely coiled) and Operculinoides soldadensis. Symbol key: ++, strong positive differences with < 1% error probability; 0, no significant differences; -, negative differences with < 5% error probability; −−, strong negative differences with < 1% error probability.

O. floridensis (tightly coiled)	Mean	SD	O. floridensis (loosely coiled)	O. soldadensis	P. trinitatensis	N. striatoreticulatus
First chamber length	431.2	92.32	−−	++	++	0
Proloculus nominal diameter	192.8	44.16	0	++	++	−−
Deuteroloculus ratio	1.080	0.0797	-	0	0	0
Initial marginal radius	255.3	51.51	0	++	++	−−
Marginal radius increase	0.113	0.0132	-	−−	0	++
Spiral chamber height increase	3.7	0.68	0	++	++	0
Initial spiral chamber height	77.1	23.95	0	++	++	-
Backbend angle	0.563	0.1055	0	−−	0	++
Initial chamber length	287.2	78.67	-	++	++	0
Chamber length increase	0.033	0.0117	0	−−	0	++
Perimeter ratio	1.302	0.0758	0	−−	0	++

Table 8. Characters and attributes (means and standard deviations, SD, in mm) for Operculinoides floridensis (loosely coiled) and comparisons with Nummulites striatoreticulatus, Palaeonummulites trinitatensis, O. floridensis(tightly coiled) and Operculinoides soldadensis. Symbol key: ++, strong positive differences with < 1% error probability; +, differences with < 5% error probability; 0, no significant differences; −−, strong negative differences with < 1% error probability.

O.floridensis (loose coiled)	Mean	SD	O.floridensis (tight coiled)	O.soldadensis	P.trinitatensis	N.striatoreticulatus
First chamber length	655.5	116.47	++	++	++	++
Proloculus nominal diameter	213.6	43.52	0	++	++	−−
Deuteroloculus ratio	1.317	0.1965	+	0	++	+
Initial marginal radius	292.5	50.03	0	++	++	0
Marginal radius increase	0.136	0.0151	+	0	++	++
Spiral chamber height increase	3.2	1.35	0	0	0	0
Initial spiral chamber height	93.0	33.38	0	++	++	0
Backbend angle	0.632	0.0866	0	0	+	++
Initial chamber length	434.4	104.66	+	++	++	++
Chamber length increase	0.042	0.0187	0	0	0	++
Perimeter ratio	1.435	0.1299	0	0	+	++

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	++	++

Figure 17. A–G, Operculinoides soldadensis Vaughan & Cole; A, Loma El Santo, CA-215-871; B, Loma Candelaria, 98LC-1-669; C–F, Noroña; C, NOR-UN 24; D–F, NOR-UN 15/14; G, holotype, Trinidad. H, I, Palaeonummulites trinitatensis (Nutall); H, Loma Candelaria, 98LC-1ICT3; I, holotype of Operculinoides kugleri Vaughan & Cole, Trinidad. J, Operculinoides ocalanus (Cushman), Loma Jabaco, CA-4-724. A–D, G–J, A forms in equatorial section; E, A form in axial section; F, external view.

Figure 18. A, B, D–H, Heterostegina ocalana Cushman; A, Loma Vigía, CA-216-D1a; B, Noroña, NOR-UN 15/14; D, Loma Vigía, CA-216-79; E, F, Loma Jabaco; E, LM-52-756; F, LM-52-752; G, H, Noroña, NOR-UN 24. C, Heterostegina cubana Cizancourt, Loma candelaria, 98LC-1H-809. I, Heterostegina sp. indet., Loma Candelaria, 98LC-1H-808. A, B, A forms in axial section; C–G, I, A forms in equatorial section; H, external view.

Figure 19. Discriminant analysis of nummulitid species, where the important discriminators are ranked along the discriminant functions. Orange arrows indicate possible source of morphological changes. A, discriminant analysis within Nummulites striatoreticulatus at localities 98LC-2, 98LC-2 and E-126. B, discriminant analysis within Operculinoides floridensis at localities 98 LC-1, CA-215 and CA-216. C, discriminant analysis within O. soldadensis at localities 98LC-1, CA-125 and NOR-UN.

Iturralde‐Vinent, M. A. 1994. Cuban geology: a new plate‐tectonic synthesis. Journal of Petroleum Geology, 17, 39–69.

 Web of Science ®Google Scholar

García-Delgado, D. & Torres-Silva, A. I. 1997. Sistema Paleógeno. Pp. 115–140 in K. E. Nuñez & G. F. Furrazola-Bermúdez (eds) Estudios sobre Geología de Cuba. Centro Nacional de Información Geológica, La Habana.

 Google Scholar

Butterlin, J. 1981. Claves para la determinación de macroforaminíferos de México y del Caribe, del Cretácico Superior al Mioceno Medio. Instituto Mexicano del Petróleo, Mexico, 219 pp.

 Google Scholar

Butterlin, J. 1961. Grandes foraminíferos del pozo Palizada núm. 2, Municipio de Palizada, Estado de Campeche. Paleontología Mexicana, 10, 1–59.

 Google Scholar

Barker, R. W. 1939. Species of the foraminiferal family Camerinidae in the Tertiary and Cretaceous of Mexico. Proceedings of the United States National Museum, 86, 305–330.

 Google Scholar

Cole, W. S. 1941. Stratigraphic and paleontologic studies of wells in Florida. Florida Geological Survey Bulletin, 19, 1–53.

 Google Scholar

Frost, S. H. & Langenheim, R. L. 1974. Cenozoic reef biofacies. Tertiary larger foraminifera and scleractinian corals from Chiapas, Mexico. Northern Illinois University Press, DeKalb, 387 pp.

 Google Scholar

Vaughan, T. W. & Cole, W. S. 1941. Preliminary report on the Cretaceous and Tertiary larger foraminifera of Trinidad British West Indies. Geological Society of America Special Papers, 30, 1–137.

 Google Scholar

Caudri, C. M. B. 1996. The larger Foraminifera of Trinidad (West Indies). Eclogae Geologicae Helvetiae, 89, 1137–1310.

 Google Scholar

Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben. C. & Breggren, W. A. (eds) 2006. Atlas of Eocene planktonic Foraminifera. Cushman Foundation for Foraminiferal Research, 41, 1–513.

 Google Scholar

Berggren, W. A., Kent, D. V., Swisher III, C. C. & Aubry, M. P. 1995. A revised Cenozoic geochronology and chronostratigraphy. SEPM Special Publication, 54, 129–212.

 Google Scholar

Martini, E. 1971. Standard Tertiary and Quaternary calcareous nannoplankton zonation. Pp. 739–785 in A. Farinacci (ed) Proceedings of the Second Planktonic Conference, Roma 1970, Tecnoscienza.

 Google Scholar

Agnini, C., Fornaciari, E., Raffi, I., Catanzariti, R., Pälike, H., Backman, J. & Rio, D. 2014. Biozonation and biochronology of Paleogene calcareous nannofossils from low and middle latitudes. Newsletters on Stratigraphy, 47, 131–181.

 Web of Science ®Google Scholar

Cotton, L. 2012. Paleogene larger benthic foraminifera of Tanzania and the Eocene–Oligocene transition. Unpublished PhD thesis, Cardiff University, Cardiff, 162 pp.

 Google Scholar

Beavington-Penney, S. J. & Racey, A. 2004. Ecology of extant nummulitids and other larger benthic foraminifera: applications in palaeoenvironmental analysis. Earth-Science Reviews, 67, 219–265.

 Web of Science ®Google Scholar

Torres-Silva, A. I., Hohenegger, J., Ćorić, S., Briguglio, A. & Eder, W. 2017. Biostratigraphy and evolutionary tendencies of Eocene heterostegines in Western and Central Cuba based on morphomteric analyses. Palaios, 32, 44–60.

 Web of Science ®Google Scholar