OCS Study 

MMS 2009-013 

 

Coastal Marine Institute 

Foraminiferal Communities 

of Bathyal Hydrocarbon Seeps, 

Northern Gulf of Mexico: A Taxonomic, 

Ecologic, and Geologic Study 

Cooperative Agreement

Coastal Marine Institute 

Louisiana State University 

U.S. Department of the Interior

Minerals Management Service 

Gulf of Mexico OCS Region 
OCS Study 

 MMS 2009-013 

Coastal Marine Institute 

Foraminiferal Communities 

of Bathyal Hydrocarbon Seeps, 

Northern Gulf of Mexico: A Taxonomic, 

Ecologic, and Geologic Study 

Authors 

Barun K. Sen Gupta 

Melissa K. Lobegeier 

Lorene E. Smith 

April 2009 

Prepared under MMS Contract 

1435-01-99-CA-30951-18182 

by 

Louisiana State University 

Coastal Marine Institute 

Baton Rouge, Louisiana 70803 

Published by 

U.S. Department of the Interior Cooperative Agreement 

Minerals Management Service Coastal Marine Institute 

Gulf of Mexico OCS Region Louisiana State University 
DISCLAIMER 

This report was prepared under contract between the Minerals Management Service (MMS) and 

Louisiana State University. The report has been technically reviewed by MMS and approved for 

publication. Approval does not signify that the contents necessarily reflect the view and policies 

of the Service nor does mention of trade names or commercial products constitute endorsement 

or recommendation for use. It is, however, exempt from review and compliance with MMS 

editorial standards. 

REPORT AVAILABILITY 

Extra copies of the report may be obtained from the Public Information Office (MS 5034) at the 

following address: 

 U.S. Department of the Interior 

 Minerals Management Service 

 Public Information Office (MS 5034) 

 Gulf of Mexico OCS Region 

 1201 Elmwood Park Boulevard 

 New Orleans, Louisiana 70123-2394 

 Telephone Number: (504) 736-2519 

 1-800-200-GULF 

CITATION 

Suggested citation: 

Sen Gupta, B.K., M.K. Lobegeier, and L.E. Smith. 2009. Foraminiferal communities of bathyal 

hydrocarbon seeps, northern Gulf of Mexico: A taxonomic, ecologic, and geologic study. U.S. 

Dept. of the Interior, Minerals Management Service, Gulf of Mexico OCS Region, New 

Orleans, LA. OCS Study MMS 2009-013, 385 pp. 

ACKNOWLEDGMENTS 

Paul Aharon (University of Alabama), Joan Bernhard (Woods Hole Oceanographic Institution), 

and Harry Roberts helped greatly in sample collection; Erik Cordes (Pennsylvania State 

University) provided tubeworms from Green Canyon and Mississippi Canyon; Joan Bernhard 

provided control cores; Xiaogang Xie helped in scanning electron micrography; and Mary Lee 

Eggart helped with the art work. Submersible dives were funded by NOAA National Undersea 

Research Program (UNC-Wilmington NURC). Parts of a published article are included here with 

the permission of Elsevier (see Chapter 4 for details).

iii
 

ABSTRACT 

One hundred eighty-five species of benthic Foraminifera were identified in a study of sediment 

substrates and tubeworm surfaces in (a) Green Canyon, Garden Banks, and Mississippi Canyon 

(245-1081 m) and (b) Alaminos, Farnella, and De Soto Canyons (1848-2918 m); the samples 

were obtained from submersibles in both seep and non-seep (control) areas. One hundred twentytwo species were calcareous, 63 agglutinated. An atlas with illustrations of most species is 

included in this report. 

No species found in this study is endemic to seeps, but 21 species were previously unknown in 

the Gulf of Mexico. The imprint of water depth on foraminiferal assemblages is clearly 

detectable, because the species are recruited from the surrounding non-seep habitats. The two 

major surface-sample groups (clusters) recognized by numerical data analysis are separated by 

bathymetry. The shallower-water group contains all samples (seep and non-seep) from depths of 

245-1081 m; the deeper-water (deepest-bathyal) group contains all samples from 1848-2918 m. 

Foraminiferal species of wide-ranging morphologic and taxonomic affinities are able to maintain 

sizeable populations at sites of hydrocarbon seepage; the high bacterial productivity at the seeps 

is a major factor in the sustenance of these populations. The most conspicuous dominants at 

seep-influenced substrates (bacterial mats) in the shallower cluster are endobenthic species, 

especially Bolivina spp.; these are possibly facultative anaerobes. The pattern is not as clear in 

the deepest-bathyal group, because some epibenthic species (e.g., Nuttallides decorata) are 

present among the dominants. In the shallower areas, the diversity (species richness) of both 

calcareous and agglutinated Foraminifera is higher in non-seep than in seep substrates. This 

distinction too is not clear in the deepest-bathyal areas. The diversity at some sites may have 

been elevated by post-mortem mixing of species from different microhabitats. Fifteen sessile, 

epibenthic Foraminifera colonize surfaces of vestimentiferan tubeworms (and possibly other 

elevated microhabitats) at hydrocarbon seeps, centimeters to decimeters above the sedimentwater interface. These attachment points are sufficiently above locations of gas escape in the 

seafloor to provide the species with an oxic microhabitat with little or no H2S. Eight of these 

species have been found exclusively on tubeworms in this study. 

v
 

TABLE OF CONTENTS 

 

Page 

LIST OF FIGURES .......................................................................................................ix 

LIST OF TABLES.........................................................................................................xi 

1 INTRODUCTION .....................................................................................................1 

 1.1 Background and Scope of Study................................................................1 

 1.2 Material and Methods ................................................................................1 

2 SYSTEMATICS ........................................................................................................7 

2.1 Introduction................................................................................................7 

2.2 Foraminiferal Orders..................................................................................7 

 2.2.1 Order Astrorhizida .........................................................................7 

 2.2.2 Order Lituolida.............................................................................16 

 2.2.3 Order Trochamminida..................................................................34 

 2.2.4 Order Textulariida........................................................................36 

 2.2.5 Order Miliolida ............................................................................43 

 2.2.6 Order Spirillinida .........................................................................50 

 2.2.7 Order Lagenida ............................................................................52 

 2.2.8 Order Buliminida .........................................................................64 

 2.2.9 Order Rotaliida.............................................................................85 

 2.2.10 Order Robertinida ......................................................................110 

 

3 FORAMINIFERA IN SEDIMENT SAMPLES ....................................................113 

 3.1 Identified Species...................................................................................113 

 3.2 Rose-Bengal-Stained Foraminifera........................................................113 

 3.3 Species Richness....................................................................................120 

 3.4 Assemblage Density...............................................................................126 

 3.5 Cluster Analyses Results........................................................................126 

 3.6 Agglutinated Foraminifera.....................................................................134 

 3.7 Diagenesis..............................................................................................140 

 3.8 Stratigraphic Record ..............................................................................140 

vii
TABLE OF CONTENTS (continued)

 Page 

4 FORAMINIFERA IN TUBEWORM SAMPLES.................................................149 

 4.1 Introduction............................................................................................149 

 4.2 Tubeworm Foraminifera ........................................................................149 

5 DISCUSSION AND CONCLUSIONS .................................................................153 

 5.1 Dominant Calcareous Species................................................................153 

 5.2 Agglutinated Species .............................................................................154 

 5.3 Species Diversity ...................................................................................154 

 5.4 Absence of Endemics.............................................................................155 

 5.5 Substrate Effect......................................................................................155 

 5.6 Trophic Resources .................................................................................156 

 5.7 Effect of Oil in the Environment ...........................................................156 

 5.8 Bathymetric Imprint...............................................................................156 

 5.9 Tolerance of Anoxia and Sulfide ...........................................................157 

 5.1 Post-Mortem Assemblage Mixing .........................................................160 

 5.11 Conclusions............................................................................................160 

6 LITERATURE CITED ..........................................................................................163 

APPENDIX: ATLAS OF FORAMINIFERA............................................................ A-1 

viii
 

LIST OF FIGURES 

 

Figure Page 

1.1 Sampling areas and core locations.........................................................................2 

3.1 Species richness (number of species) in surface and near-surface samples 

 plotted against water depths...............................................................................125 

3.2 Assemblage density (individuals/cm3) in surface and near-surface samples 

 plotted against water depths...............................................................................127 

3.3 Q-mode clusters, surface samples......................................................................128 

3.4 Relative abundances of the more important species in minor clusters, 

 surface samples. .................................................................................................129 

3.5 Q-mode clusters, surface (0-1 cm) and near-surface (1-2 cm) samples. ...........130 

3.6 Relative abundances of the more important species in minor clusters, 

 surface and near-surface samples.......................................................................139 

3.7 Abundance variations of Bolivina lowmani (solid line) and 

 Nuttallides decorata (dashed line) in Alaminos Canyon core ACP05. .............145 

3.8 Q-mode cluster diagram (Ward's method) for stratigraphic data, 

 Alaminos Canyon core ACP05..........................................................................145 

3.9 Abundance variations of Bolivina lowmani (solid line), 

 Epistominella exigua (dotted line), and Nuttallides decorata (dashed line) 

in Alaminos Canyon core ACP15......................................................................146 

3.10 Q-mode cluster diagram (Ward's method) for stratigraphic data, 

 Alaminos Canyon core ACP15..........................................................................146 

3.11 Abundance variations of Cassidulina carinata (solid line) and Osangularia 

 culter (dashed line) in Mississippi Canyon core MCP73. .................................147 

3.12 Q-mode cluster diagram (Ward's method) for stratigraphic data, 

 Mississippi Canyon core MCP73.......................................................................147

ix
 

 LIST OF TABLES 

Table Page 

1-1 Core location, inventory of surface and near-surface samples, 

 submersible-dive reference, water depth, geographic coordinates, 

 and substrate character. .........................................................................................5 

2-1 Alphabetical list of species described or discussed in Chapter 2...........................8 

3-1 Geographic occurrence of species. ....................................................................114 

3-2 Counts of Rose-Bengal-stained individuals for the 32 species that contribute 

 more than one or two stained individuals at more than one site........................121 

3-3 Species richness (S) and assemblage density (AD) in sediment samples..........124 

3-4 Q-mode minor clusters for surface-sample data: Constituent samples, 

 water depths, substrates, and important foraminiferal species 

 (mean abundance >5% in minor cluster). ..........................................................131 

3-5 Q-mode minor clusters for combined surface- and near-surface-sample data: 

 Constituent samples, water depths, substrates, and important foraminiferal 

 species (mean abundance >5% in minor cluster)...............................................132 

3-6 List of the 15 most important calcareous species in cluster analysis.................135 

3-7 Counts (N) and cumulative percentages (Agg. %) of agglutinated 

 individuals in samples, with listing of dominant species (>5% of 

 agglutinated assemblage)...................................................................................136 

3-8 Core ACP05, Alaminos Canyon (Alvin dive 3624): Percentages of 

 calcareous species that constitute at least 1% of the assemblage in one 

 or more samples. ................................................................................................142 

3-9 Core ACP15, Alaminos Canyon (Alvin dive 3625): Percentages of 

 calcareous species that constitute at least 1% of the assemblage in 

 one or more samples. .........................................................................................143 

3-10 Core MCP73, Mississippi Canyon (Alvin dive 3631): Percentages of 

 calcareous species that constitute at least 1% of the assemblage in one 

 or more samples. ................................................................................................144 

4-1 Foraminifera attached to vestimentiferan tubeworms........................................151

xi 
1. INTRODUCTION

1.1. Background and Scope of Study 

Marine hydrocarbon seeps ("cold seeps"), especially those of the Gulf of Mexico, have been the 

subject of many studies because, like hydrothermal vents ("hot vents"), they support a complex 

community of organisms in a chemosynthesis-based ecosystem (e.g., Hecker, 1985; Kennicutt et 

al., 1985; MacDonald et al., 1989; Van Dover, 2000). The present study relates to a very large 

group of shelled protists, namely benthic Foraminifera, that live in seep-related substrates. This 

is frequently the most abundant meiofaunal group in diverse marine habitats (e.g., Snider et al., 

1984; Gooday, 1986; Gooday et al., 1992). Foraminiferal species respond rapidly to 

environmental disturbances, and their large extant populations and well-preserved shells make 

them excellent recorders of present and past marine biodiversity. Some species are capable of 

inhabiting environments where oxygen is depleted to extremely low levels, or even absent. 

Hydrocarbon seeps provide a special window to study the Foraminifera of such dysoxic and 

anoxic environments, and to assess long-term effects of petroleum pollution on foraminiferal 

communities. 

Our preliminary studies of the Foraminifera associated with Gulf of Mexico hydrocarbon seeps, 

especially those found in or under bacterial (Beggiatoa) mats, have indicated the existence of 

facultative anaerobes within this community (Sen Gupta and Aharon, 1994; Sen Gupta et al., 

1997). These studies, however, were too limited in scope for an understanding of spatial and 

temporal distribution trends, and for an assessment of diversity and abundance anomalies. In 

particular, the composition of seep foraminiferal communities in the Gulf of Mexico at depths 

greater than 700 m was wholly unknown at the commencement of the present study, although a 

later publication (Robinson et al., 2004) reported 32 living species (Rose-Bengal-stained 

individuals) from Alaminos Canyon and Atwater Canyon. In this context, our investigation 

involved a quantitative analysis of foraminiferal assemblages retrieved from various shallow and 

deep bathyal seepage areas in the northern Gulf of Mexico. The foraminiferal microhabitats 

included bacterial (Beggiatoa) mats, mussel beds, substrates of tubeworm colonies, and 

inorganic sediments around points of hydrocarbon seepage and at control sites away from seeps. 

In addition, a presence-absence census was also conducted for epibenthic species attached to 

tubeworm surfaces. Scanning electron micrographs were taken of all species for which suitable 

specimens were found; some digital photomicrographs were also taken, especially of species 

attached to tubeworms. All these illustrations are included in a taxonomic atlas that constitutes a 

major part of this report. 

1.2. Material and Methods 

Sediment Samples. The sampling locations are shown in Fig. 1.1. Most of the cores used in the 

present study were retrieved during dives of the submersible Alvin in October 2000. This 

sampling project included three separate research programs involving investigators from (1) LSU 

(B. K. Sen Gupta, P.I.) and University of South Carolina (USC, J. M. Bernhard, Co-P.I.), (2) 

Texas A & M University (TAMU, I. R. McDonald, P.I.), and (3) College of William and Mary 

(CWM, C. L. Van Dover, P.I.). 

1
Figure 1.1. Sampling areas and core locations. Top, sampling areas and bathymetry; bottom, 

approximate locations of coring sites on a latitude-longitude grid. Exact coordinates 

of sites are given in Table 1-1. 

2
The TAMU dives were funded by MMS (Gulf of Mexico Office) and DOE (National Energy 

Technology Laboratory); the LSU/USC and CWM dives were supported by NOAA (National 

Undersea Research Program). The cores retrieved on these Alvin dives included eight push cores 

and four box cores from Alaminos Canyon (water depth 2219-2238 m); five push cores from 

Mississippi Canyon (1067-1081 m); five push cores from Green Canyon (Block 272, 664-696 

m); and one push core from Farnella Canyon (2918m). The coordinates, water depth, and 

substrate information (seep-related habitat or non-seepage site) for each core are recorded in 

Table 1-1. The diameter of the push cores was 7 cm and the maximum length was about 25 cm. 

The box cores were taken (in Alaminos Canyon) with a four-compartment box (width 15 cm, 

depth 15 cm, height 18 cm). Besides the Alvin 2000 cores, three push cores were obtained earlier 

during dives of the Johnson Sea-Link in 1997: two from Green Canyon (Block 180, 245 m; 

Block 232, 569 m), and one from Garden Banks (640 m). Finally, two control cores from nonseep locations (De Soto Canyon, 1848 m; south of Mississippi Canyon, 1076 m), provided by 

Dr. Joan Bernhard were used for comparison. 

All cores were kept frozen until sectioned in the laboratory. They were split vertically into halves 

with a buzz saw, and one half was kept for archival purposes. The other half was sectioned in 

one-centimeter slices. Wherever possible, surface and near-surface samples (0-1 cm and 1-2 cm) 

from push cores and surface samples (0-1 cm) from box cores were studied. For three cores, 

samples below the 2-cm level were also studied. The surface samples from push core ACP12 (0-

1.5 cm and 1.5-2 cm) and box core ACB08 (0-2 cm) were larger, because the top surfaces were 

particularly uneven. The core slices were stained with a saturated solution of Rose Bengal (to 

distinguish between living and dead Foraminifera) and washed over a 63-μm sieve; the residues 

were dried at 60 ̊C. These residues were microsplit so that, if possible, 300 plus benthic 

foraminiferal specimens could be picked and counted. Pyrite-filled shells of calcareous and 

agglutinated Foraminifera and specimens with authigenic carbonate or barite overgrowth were 

not included in the total counts. In samples with excessive numbers of tube-like agglutinated 

Foraminifera, such as MCP73 from Mississippi Canyon, >100 calcareous benthic Foraminifera 

were picked from the split sample and numbers of tubular Foraminifera counted. 

Numerical data analyses. Our numerical database consists of species-level abundances of both 

living individuals and empty shells (unstained by Rose Bengal) of Foraminifera in core samples. 

The relative abundances of calcareous species were subjected to Q-mode cluster analysis, using 

Ward’s minimum variance method with squared Euclidean distance. Data on agglutinated 

Foraminifera were excluded from analysis because counts of agglutinated Foraminifera in these 

samples are seriously affected by taphonomic problems. In particular, tubular Foraminifera such 

as Hyperammina, Rhizammina, and Saccorhiza are frequently fragmented, and the correct 

number of specimens is difficult to estimate (see Licari et al., 2003; Mackensen et al., 1993; 

Schmiedl et al., 1997; Wollenburg and Mackensen, 1998; Kurbjeweit et al., 2000).

Two data sets were analyzed, one on just the surface samples (0-1 cm), the other on both surface 

and near-surface samples (0-1 cm and 1-2 cm). When performing Q-mode cluster analysis (CA) 

on the first data set (28 0-1 cm samples), six samples—all from Alaminos Canyon (ACP01, 

ACP10, ACP11, ACP12, ACB08, ACB09)—were excluded, because each of them contained 

less than 25 calcareous benthic individuals. Fifty-three variables, defined as species constituting 

3
at least 2% of the assemblage in at least one sample, were used in the clustering procedure. For 

CA of the second data set (49 0-1 and 1-2 cm samples), nine samples were excluded, because of 

the paucity of calcareous individuals; these were: ACP01a (= ACP01 0-1 cm), ACP10a, ACP10b 

(= ACP10 1-2 cm), ACP11a, ACP11b, ACP12a (= ACP12 0-1.5 cm), ACP12b (= ACP12 1.5-2 

cm), ACB08a (= ACB08 0-2 cm), and ACB09a. Fifty-four species (2% or more in one or more 

samples) were used. 

Tubeworm Foraminifera. Tubeworms from Alaminos Canyon were retrieved during our Alvin

dive of 2000; the others (from Green Canyon and Mississippi Canyon) were contributed by Erik 

Cordes of Pennsylvania State University. Air-dried tubeworms, with internal soft tissue removed, 

were first examined under a binocular microscope, photographed, and sketched, showing places 

of foraminiferal colonization. The tubes were then sectioned transversely by a Dremel tool. The 

attached Foraminifera were identified and documented by digital photomicrography and 

scanning electron micrography––in growth positions, and, if necessary, as detached specimens. 

4
Table 1-1 

Core Location, Inventory of Surface and Near-Surface Samples, Submersible-Dive Reference, 

Water Depth, Geographic Coordinates, and Substrate Character 

CORE 

NUMBER 

SAMPLES LOCATION DIVE 

NUMBER 

WATER 

DEPTH 

(m) 

LATITUDE & 

LONGITUDE 

SUBSTRATE 

ACP01 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3624 2226 26o21.355' N 

94o29.87156' W 

White mud mound 

ACP04 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3624 2227 26o21.35798' N 

94o29.87998' W 

White mud mound 

ACP05 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3624 2225 26o21.35744' N 

94o29.87757' W 

Mud next to 

tubeworm clump 

ACP10 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3625 2219 26o21.33252' N 

94o29.86254' W 

Orange bacterial 

mat with tubeworm 

roots 

ACP11 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3625 2219 26o21.33252' N 

94o29.86254' W 

Mud next to mussel 

bed 

ACP12 0-1.5 cm 

1.5-2 cm 

Alaminos 

Canyon 

Alvin 3625 2219 26o21.33252' N 

94o29.86254' W 

Mud next to mussel 

bed 

ACP13 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3625 2221 26o21.32115' N 

94o29.8938' W 

White, fluffy 

bacterial mat 

ACP15 0-1 cm 

1-2 cm 

Alaminos 

Canyon 

Alvin 3625 2221 26o21.32115' N 

94o29.8938' W 

Orange bacterial 

mat 

ACB07 0-1 cm Alaminos 

Canyon 

Alvin 3624 2227 26o21.35635' N 

94o29.87757' W 

Gray bacterial mat 

with tubeworm roots 

ACB08 0-2 cm Alaminos 

Canyon 

Alvin 3624 2227 26o21.35635' N 

94o29.87757' W 

Gray bacterial mat 

ACB09 0-1 cm Alaminos 

Canyon 

Alvin 3624 2227 26o21.3596' N 

94o29.87757' W 

Fluffy bacterial mat 

(H2S odor) 

ACB19 0-1 cm Alaminos 

Canyon 

Alvin 3625 2219 26o21.35910' N 

94o29.89801' W 

Non-seep gray mud 

(control) 

MCP02 0-1 cm 

1-2 cm 

Mississippi 

Canyon 

Alvin 3631 1067 28o7.58' N 

89o8.61' W 

Mud from ridge top, 

near mussel bed 

MCP04 0-1 cm 

1-2 cm 

Mississippi 

Canyon 

Alvin 3631 1072 28o7.24097' N 

89o8.4266' W 

Gray bacterial mat 

MCP08 0-1 cm 

1-2 cm 

Mississippi 

Canyon 

Alvin 3631 1071 28o7.235' N 

89o8.42' W 

Gray bacterial mat 

MCP69 0-1 cm 

1-2 cm 

Mississippi 

Canyon 

Alvin 3630 1081 28o7.59777' N 

89o8.62017' W 

Bacterial mat 

MCP73 0-1 cm 

1-2 cm 

Mississippi 

Canyon 

Alvin 3631 1070 28o7.218' N 

89o8.375' W 

Black sediment with 

bacterial mat 

GCP33 0-1 cm 

1-2 cm 

Green Canyon 

(Block 272) 

Alvin 3626 664 27o41.1582' N 

91o32.3409' W 

White bacterial mat 

GCP51 0-1 cm 

1-2 cm 

Green Canyon 

(Block 272) 

Alvin 3627 696 27o41.01363' N 

91o32.26124' W 

Bacterial mat on 

mound with clams 

GCP54 0-1 cm 

1-2 cm 

Green Canyon 

(Block 272) 

Alvin 3627 675 27o41.01' N 

91o32.25' W 

White bacterial mat 

GCP55 0-1 cm 

1-2 cm 

Green Canyon 

(Block 272) 

Alvin 3627 689 27o41.13654' N 

91o32.34759' W 

Non-seep, buffcolored mud 

(control) 

GCP57 0-1 cm 

1-2 cm 

Green Canyon 

(Block 272) 

Alvin 3627 675 27o41.19448' N 

91o32.28009' W 

Sand (control) 

5
Table 1-1. Core Location, Inventory of Surface and Near-Surface Samples, Submersible-Dive 

Reference, Water Depth, Geographic Coordinates, and Substrate Character (continued). 

CORE 

NUMBER 

SAMPLES LOCATION DIVE 

NUMBER 

WATER 

DEPTH 

(m) 

LATITUDE & 

LONGITUDE 

SUBSTRATE 

FCP61 0-1 cm Farnella 

Canyon 

Alvin 3628 2918 26o26.954' N 

90o46.421' W 

Tubeworm clump 

GCP12 0-1 cm 

1-2 cm 

Green Canyon 

(Block 232) 

JSL 2900 

(1997) 

569 27o44.44' N 

91o19.08' W 

White bacterial mat 

GCP15 0-1 cm 

1-2 cm 

Green Canyon 

(Block 180) 

JSL 2902 

(1997) 

245 27o47.8036' N 

91o43.6981' W 

Non-seep gray mud 

(control) 

GBP08 0-1 cm 

1-2 cm 

Garden Banks 

(Block 427) 

JSL 2896 

(1997) 

640 27o33.51' N 

92o25.26' W 

Non-seep gray mud 

(control) 

DCB-S36 0-1 cm De Soto 

Canyon 

Bernhard 1848 28o55.4' N 

87o39' W 

Non-seep (control) 

MCBC7 0-1 cm South of 

Mississippi 

Canyon 

Bernhard 1076 27o43.7' N 

89o58.7' W 

Non-seep (control) 

6
2. SYSTEMATICS 

2.1. Introduction 

The taxonomy of the large majority of benthic foraminiferal species encountered in this study is 

given in this chapter, with references to some previous uses of the chosen names and to 

significant synonyms. Where necessary, brief descriptions and reasons for accepting 

controversial names are given. Illustrations of 180 species, arranged alphabetically by their 

names, are given in the plates of the Atlas (Appendix); references to these plates accompany the 

species descriptions here. For suprageneric categories, we follow Loeblich and Tappan (1987), 

as modified by Sen Gupta (1999). Unless mentioned otherwise, the illustrations are scanning 

electron micrographs; the exceptions are digital photomicrographs. 

The species described or discussed in this section are listed in Table 2-1. Five rare species, 

although named in a table of occurrences later (Table 3-1), are not included here, because of a 

lack of specimens suitable for illustration. 

2.2. Foraminiferal Orders

2.2.1. Order Astrorhizida 

Superfamily Astrorhizacea 

Family Rhabdamminidae 

Genus Rhabdammina

Rhabdammina abyssorum Sars 

Rhabdammina abyssorum Sars, 1869, p. 248.

Rhabdammina abyssorum Sars – Barker, 1960, p. 42, pl. 21, figs. 1-13.

Rhabdammina abyssorum Sars – Jones, 1994, p. 32, pl. 21, figs. 1-8, 10-13. 

Morphology: Shell has three and sometimes four or five tubes radiating from a globular central 

cavity. 

Figures: 

Plate 152, fig. 1, Alaminos Canyon Boxcore B6 bulk sample, side view. 

Scale bars = 50 μm. 

Rhabdammina discreta Brady

Rhabdammina discreta Brady, 1881, p. 48, pl. 22, figs. 7-10. 

Rhabdammina discreta Brady – Brady, 1884, p. 268, pl. 22, figs. 11-13. 

Rhabdammina discreta Brady – Cushman, 1918, p. 21, pl. 11, fig. 1. 

Psammosiphonella discreta (Brady) – Barker, 1960, p. 44, pl. 22, figs. 7-10. 

7
Table 2-1 

Alphabetical List of Species Described or Discussed in Chapter 2 

SPECIES 

Abditodentrix pseudothalmanni (Boltovskoy and Giussani de Kahn) Cornuspira involvens (Reuss)

Adercotryma glomeratum (Brady) Cribromiliolinella subvalvularis (Parr)

Ammobaculites catenulatus Cushman and McCulloch Cribrostomoides subglobosus (Cushman)

Ammobaculites filiformis Earland Cystammina pauciloculata (Brady)

Ammodiscus tenuis (Brady) Dentalina sp. 

Ammolagena clavata (Jones and Parker) Deuterammina rotaliformis (Heron-Allen and Earland)

Ammoscalaria tenuimargo (Brady) Discammina compressa (Goës)

Amphicoryna hirsuta (d’Orbigny) Discanomalina semipunctata (Bailey)

Anomalinoides globulosus (Chapman and Parr) Dorothia scabra (Brady)

Anomalinoides mexicanus Parker Eggerella bradyi (Cushman)

Archimerismus subnodosus (Brady) Epistominella exigua (Brady)

Barbourinella atlantica (Bermúdez) Epistominella vitrea Parker

Bolivina alata (Seguenza) Eponides turgidus Phleger and Parker

Bolivina albatrossi Cushman Eubuliminella morgani (Andersen)

Bolivina barbata Phleger and Parker Fissurina fissa (Heron-Allen and Earland)

Bolivina daggarius Parker Fissurina sp. cf. F. incomposita (Patterson and Pettis)

Bolivina goesii Cushman Fursenkoina seminuda (Natland)

Bolivina sp. cf. B. hastata Phleger and Parker Gaudryina minuta Earland

Bolivina lowmani Phleger and Parker Gavelinopsis tranlucens (Phleger and Parker)

Bolivina minima Phleger and Parker Globocassidulina subglobosa (Brady) 

Bolivina ordinaria Phleger and Parker Glomospira gordialis (Jones and Parker)

Bolivina sp. cf. B. pusilla Schwager Glomospira irregularis (Grzybowski)

Bolivina subaenariensis Cushman Gyroidina bradyi (Trauth)

Bolivina translucens Phleger and Parker Gyroidina orbicularis (Parker, Jones, and Brady)

Bolivinita quadrilatera (Schwager) Gyroidinoides sp. cf. G. polius (Phleger and Parker)

Bulimina aculeata d’Orbigny Gyroidinoides sp. cf. G. regularis (Phleger and Parker)

Bulimina alazanensis Cushman Haplophragmoides sp. cf. H. kirki Wickenden

Bulimina marginata d’Orbigny Haynesina germanica (Ehrenberg)

Bulimina mexicana Cushman Hoeglundina elegans (d’Orbigny)

Buzasina ringens (Brady) Hormosina pilulifera (Brady)

Calcituba polymorpha von Roboz Hormosinella distans (Brady)

Cassidulina carinata Silvestri Hormosinella guttifera (Brady)

Cassidulina curvata Phleger and Parker Hyperammina friabilis Brady

Cassidulina obtusa Williamson Hyperammina laevigata Wright

Cassidulinoides tenuis Phleger and Parker Hyperammina sp. 

Chilostomella oolina Schwager Ioanella tumidula (Brady)

Cibicides lobatulus (Walker and Jacob) Karreriella bradyi (Cushman)

Cibicides wuellerstorfi (Schwager) Karrerulina apicularis (Cushman)

Cibicidoides pachyderma (Rzehak) Lagena chrysalis Heron-Allen and Earland

Cibicidoides robertsonianus (Brady) Lagena hispida Reuss

Cornuspira foliacea (Philippi) Lagena hispidula Cushman

8
Table 2-1. Alphabetical List of Species Described or Discussed in Chapter 2 (continued). 

SPECIES 

Lagenammina difflugiformis (Brady) Pseudoclavulina serventyi (Chapman and Parr)

Lagenammina laguncula Rhumbler Pseudogaudryina atlantica (Bailey)

Lagenammina tubulata (Rhumbler) Pseudoglandulina comatula (Cushman)

Laminononion tumidum (Cushman and Edwards) Pseudotrochammina sp. 

Laticarinina pauperata (Parker and Jones) Pullenia bulloides (d’Orbigny)

Lenticulina calcar (Linnaeus) Pullenia quinqueloba (Reuss)

Lenticulina gibba (d’Orbigny) Pyrgo lucernula (Schwager)

Lenticulina pliocaena (Silvestri) Pyrgo murrhina (Schwager)

Lenticulina thalmanni (Hessland) Pyrgo nasuta Cushman

Lituotuba lituiformis (Brady) ?Pyrgo sp. 

Martinottiella communis (d’Orbigny) Quinqueloculina bosciana d’Orbigny

Melonis affinis (Reuss) Reophax agglutinatus Cushman

Melonis pompilioides (Fichtel and Moll) Reophax scorpiurus Dénys de Montfort

Miliolinella warreni Andersen Rhabdammina abyssorum Sars

Miliolinella sp. Rhabdammina discreta Brady

Multifidella nodulosa (Cushman) Rhizammina algaeformis Brady

Neocrosbyia minuta (Parker) Rhizammina indivisa Brady

Nonionella iridea Heron-Allen and Earland Robertina subcylindrica (Brady)

Nuttallides decorata (Phleger and Parker) Robertinoides bradyi (Cushman and Parker)

Oolina apiopleura (Loeblich and Tappan) Rutherfordoides mexicanus (Cushman)

Oolina globosa (Montagu) Saccammina helenae (Rhumbler)

Oolina ovum (Ehrenberg) Saccorhiza ramosa (Brady)

Oolina squamosa (Montagu) Saracenaria altifrons (Parr)

Orectostomina camposi (Brönnimann and Beurlen) Seabrookia earlandi Wright

Oridorsalis umbonatus (Reuss) Sigmoilinita elliptica (Galloway and Wissler)

Osangularia culter (Parker and Jones) Sigmoilopsis schlumbergeri (Silvestri)

Osangularia rugosa (Phleger and Parker) Siphonina pulchra Cushman

Parafissurina botelliformis (Brady) Siphonodosaria calomorpha (Reuss)

Parafissurina kerguelenensis (Parr) Siphotextularia rolshauseni Phleger and Parker

Parafissurina lateralis (Cushman) Spirillina vivipara Ehrenberg

Paratrochammina challengeri Brönnimann and Whittaker Spiroplectammina sp. 

Parvigenerina arenacea (Heron-Allen and Earland) Stainforthia complanata (Egger)

Patellina corrugata Williamson Stainforthia compressa (Bailey)

Planispirinoides bucculentus var. placentiformis (Brady) Stainforthia pontoni (Cushman)

Planulina ariminensis d’Orbigny Subreophax monile (Brady)

Planulina foveolata (Brady) Tetrataxiella sp. 

Plectofrondicularia advena (Cushman) Textularia foliacea occidentalis Cushman

Portatrochammina antarctica (Parr) Textularia sp. cf. T. mayori Cushman

Praeglobobulimina ovata (d’Orbigny) Textularia mexicana Cushman

Praeglobobulimina ovula (d’Orbigny) Textulariella barrettii (Jones and Parker)

Procerolagena gracilis (Williamson) Trifarina bradyi Cushman

Prolixoplecta parvula (Cushman) Trifarina jamaicensis (Cushman and Todd)

Psammosphaera fusca Schulze Triloculina tricarinata d’Orbigny

Pseudoclavulina mexicana (Cushman) Usbekistania charoides (Jones and Parker)

9
Table 2-1. Alphabetical List of Species Described or Discussed in Chapter 2 (continued). 

SPECIES 

Uvigerina auberiana d’Orbigny Valvulineria glabra Cushman

Uvigerina bellula Bandy Valvulineria mexicana Parker

Uvigerina flintii Cushman Vasiglobulina reticulata Poag

Uvigerina peregrina Cushman Veleroninoides jeffreysii (Williamson)

Vaginulinopsis subaculeata (Cushman) Veleroninoides wiesneri (Parr)

10
Rhabdammina discreta Brady – Charnock and Jones, 1990, p. 152, pl. 1, fig. 25; pl. 2, fig. 23. 

Rhabdammina discreta Brady – Kaminski et al., 1990, p. 363-364, pl. 1 fig. 3. 

Rhabdammina discreta Brady – Jones, 1994, p. 32, pl. 22, figs. 7-10. 

Morphology: Tubular shell, straight with constrictions at irregular intervals. 

Remarks: As Berggren and Kaminski (1990) pointed out there is confusion about the 

morphology of the genus Rhabdammina. We are following Kaminski et al. (1990) in including 

single-branched, open agglutinated tubes under Rhabdammina. 

Figures: 

Plate 153, figs. 1-4, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Rhizammina 

Rhizammina algaeformis Brady 

Rhizammina algaeformis Brady, 1879, p. 39, pl. 4, figs. 16-17. 

Rhizammina algaeformis Brady – Brady, 1884, p. 274, pl. 28, figs. 1-11. 

Rhizammina algaeformis Brady – Barker, 1960, p. 58, pl. 28, figs. 1-11. 

Rhizammina algaeformis Brady – Jones, 1994, p. 36, pl. 28, figs. 1-11. 

Morphological comments: Shell tubular, branching dichotomously. 

Figures: 

Plate 154, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 2, Green Canyon 

(Block 272) Core P-57 Sample 0-1 cm, side view; fig. 3, Alaminos Canyon Core PC5 Sample 1-

2 cm, side view. 

Scale bars = 50 μm. 

Rhizammina indivisa Brady

Rhizammina indivisa Brady, 1884, p. 277, pl. 29, figs. 5-7. 

Rhizammina indivisa Brady – Cushman, 1918, p. 32, 33, pl. 12, figs. 7-10. 

Testulosiphon indivisa (Brady) – Avnimelech, 1952, p. 64, 66. 

Testulosiphon indivisa (Brady) – Barker, 1960, p. 60, pl. 29, figs. 5-7. 

Testulosiphon indivisa (Brady) – Jones, 1994, p. 36, pl. 29, figs. 5-7. 

Morphology: Shell an elongate tube tapering towards the ends; shell wall composed of two 

layers with varying numbers of planktonic Foraminifera overlying a finer clay layer. 

11
Remarks: The original generic description of Rhizammina by Brady (1879) is of a branching 

tube. Branching or simple tubes are described in Loeblich and Tappan (1964) and only 

branching tubes in Loeblich and Tappan (1987). Kaminski et al. (1990) decribed Rhizammina 

indivisa as “unbranched curved tubes with a thin, finely agglutinated wall.” However, in this 

study we have noted a wide variation in material picked up by the Foraminifera starting with a 

few planktonic shells on a fine clayey matrix to a complete layer of planktonic Foraminifera 

shells on finer material underneath. 

Figures: 

Plate 155, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, Mississippi 

Canyon Core P-69 Sample 0-1 cm, side view; fig. 3, De Soto Canyon control sample DGOMB2 

S36, side view; fig. 4, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

Family Psammosphaeridae 

Genus Psammosphaera 

Psammosphaera fusca Schulze 

Psammosphaera fusca Schulze, 1875, p. 113, pl. 2, figs. 8a-f. 

Psammosphaera fusca Schulze – Brady, 1884, p. 249, pl. 18, figs. 1-8. 

Psammosphaera fusca Schulze – Cushman, 1918, p. 34, 35, pl. 13, figs. 1-6; pl. 14, figs. 1-3. 

Psammosphaera fusca Schulze – Barker, 1960, p. 36, pl. 18, figs. 1, 5-8. 

Psammosphaera parva Flint – Barker, 1960, p. 36, pl. 18, figs. 2-4. 

Psammosphaera fusca Schulze – Jones, 1994, p. 31, pl. 18, figs. 1-8. 

Morphology: Shell a single, rough-textured, rounded chamber composed of loosely cemented 

large sand grains; no definite apertures. 

Remarks: This is the original type species of the genus Psammosphaera Schulze 1875. 

Figures: 

Plate 137, figs. 1-2, Green Canyon Core P-51 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Family Saccamminidae 

Genus Lagenammina 

Lagenammina difflugiformis (Brady) 

Reophax difflugiformis Brady, 1879, p. 51, pl. 4, figs. 3a, b. 

Reophax difflugiformis Brady – Brady, 1884, p. 289, pl. 30, figs. 1-3. 

Proteonina difflugiformis (Brady) – Cushman, 1918, p. 47, 48, pl. 21, figs. 1, 2. 

12
Proteonina difflugiformis (Brady) – Phleger and Parker, 1951, p. 2, pl. 1, figs. 4, 5. 

Reophax difflugiformis Brady – Barker, 1950, p. 62, pl. 30, figs. 1-3. 

Lagenammina difflugiformis Brady – Charnock and Jones, 1990, p. 146, pl. 1, fig. 2; pl. 13, fig. 

2. 

Lagenammina sp. – Kaminski et al., 1990, p. 365, pl. 2, fig. 2. 

Lagenammina difflugiformis Brady – Jones, 1994, p. 36, pl. 30, figs. 1-3. 

Morphology: Shell a single flask-shaped chamber; tubular neck tapering to terminal aperture. 

Remarks: Kaminski et al. (1990) referred this species to Lagenammina sp. stating that 

Lagenammina difflugiformis (Brady) is in need of taxonomic revision based on type material. 

However, Charnock and Jones (1990) noted the similarity of their fossil specimens with the 

Recent types of L. difflugiformis in the Brady collection in the British Museum (Natural History). 

Figures: 

Plate 91, figs. 1-3, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

Lagenammina laguncula Rhumbler 

Lagenammina laguncula Rhumbler, 1911, p. 92, 111, pl. 1, fig. 4. 

Proteonina longicollis Wiesner, 1931, p. 82, pl. 6, fig. 55. 

Morphology: Shell a single elongate chamber with a long thin neck and a round aperture. 

Remarks: Wiesner (1931) states that Proteonina longicollis resembles Lagenammina laguncula

Rhumbler but differs in its firm chamber (“fest Sandschale”). We accept L. laguncula due to 

priority. 

Figures: 

Plate 92, fig. 1, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, side view. 

Scale bar = 50 μm. 

Lagenammina tubulata (Rhumbler)

Saccammina tubulata Rhumbler, 1931, in Wiesner, 1931, p. 82, pl. 23, fig. 1. 

Proteonina bulbosa Chapman and Parr, 1937, p. 150, pl. 10, fig. 42. 

Lagenammina tubulata (Rhumbler) – Kuhnt et al., 2000, p. 279, pl. 1, fig. 14; pl. 10, fig. 2. 

Morphology: Shell a single globular chamber terminating in a long, slender neck. 

Remarks: The descriptions of Lagenammina tubulata (Rhumbler, in Wiesner, 1931) and 

Proteonina bulbosa by Chapman and Parr (1937) are essentially the same. We accept S. 

tubulata due to priority. 

13
 

Figures: 

Plate 93, figs. 1-3, De Soto Canyon control sample DGOMB2 S36, side view; fig. 4, Alaminos 

Canyon Boxcore B6 bulk sample, side view. 

Scale bars = 50 μm. 

Genus Saccammina 

Saccammina helenae (Rhumbler) 

Proteonina helenae Rhumbler, 1913, p. 380, pl. 2, figs. 16-17.

Proteonina helenae Rhumbler – Cushman, 1918, p. 50, pl. 20, figs. 6, 7. 

Morphology: Shell a single chamber, fusiform in shape, tapering towards the aperture; shell wall 

composed of tests of planktonic foraminifera; aperture circular in shape. 

Figures: 

Plate 159, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, Alaminos 

Canyon Boxcore B6 bulk sample, side view. 

Scale bars = 50 μm. 

Superfamily Hippocrepinacea 

Family Hippocrepinidae 

Genus Hyperammina 

Hyperammina friabilis Brady 

Hyperammina friabilis Brady, 1884, p. 258, pl. 23, figs. 1-3, 5, 6. 

Hyperammina friabilis Brady – Cushman, 1918, p. 75, 76, pl. 29, figs. 1-3. 

Hyperammina friabilis Brady – Barker, 1960, p. 46, pl. 23, figs. 1-3, 5, 6. 

Hyperammina friabilis Brady – Charnock and Jones, 1990, p. 150, pl. 1, fig. 14; pl. 13, fig. 15. 

Hyperammina friabilis Brady – Jones, 1994, p. 33, pl. 23, figs. 1-3, 5, 6. 

Morphology: Shell a straight, elongate tube which may taper slightly towards one end; 

proloculus large and rounded but frequently absent due to breakage. 

Remarks: Differs from H. elongata Brady in its larger size, stouter dimensions and more broadly 

rounded proloculus. 

14
Figures: 

Plate 81, fig. 1, Alaminos Canyon Core PC10 Sample 0-1 cm, side view; fig. 2, Alaminos 

Canyon Boxcore B19.1 Sample 0-1 cm, side view; fig. 3, Alaminos Canyon Core PC5 Sample 1-

2 cm, side view. 

Scale bars = 50 μm. 

Hyperammina laevigata Wright 

Hyperammina elongata var. laevigata Wright, 1891, p. 466, pl. 20, fig. 1. 

Hyperammina elongata Brady – Brady, 1884, p. 257, pl. 23, figs. 9, 10. 

Hyperammina laevigata Wright – Cushman, 1918, p. 77, 78, pl. 29, figs. 5, 6. 

?Hyperammina tubulosa Earland, 1934, p. 75, pl. 2, fig. 21. 

Hyperammina laevigata Wright – Barker, 1960, p. 46, pl. 23, figs. 9, 10. 

Hyperammina laevigata Wright – Jones, 1994, p. 33, pl. 23, figs. 9, 10. 

Morphology: Shell a simple elongate tube, straight to slightly curved; proloculous varies from 

elongate to slightly bulbous; shell wall fine grained, smooth and polished; aperture a round 

opening. 

Remarks: Barite-covered shells of H. laevigata are common in Mississippi Canyon Core P-73. 

Hyperammina tubulosa is probably synonymous with H. laevigata as the aboral end of H. 

laevigata is variable. 

Figures: 

Plate 82, figs. 1-2, De Soto Canyon control sample DGOMB2 S36, side view; fig. 3, Mississippi 

Canyon Core P-73 Sample 0-1 cm, side view; fig. 4, Mississippi Canyon Core P-73 Sample 1-2 

cm, side view; fig. 5, Mississippi Canyon Core P-73 Sample 0-1 cm, side view; figs. 6-8, bariteencrusted specimens, Mississippi Canyon Core P-73 Sample 0-1 cm, side view. 

Plate 83, barite-encrusted specimens, figs. 1, 3, 4, 8, Mississippi Canyon Core P-73 Sample 1-2 

cm, side view; figs. 2, 5, 6, 7, Mississippi Canyon Core P-73 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Hyperammina sp. 

Morphology: Shell a straight or slightly curved elongate tube; without a separate proloculus; 

shell wall composed of large sand grains, cemented closely together. 

Remarks: Somewhat similar to Hyperammina rugosa Verdenius and van Hinte, 1983, p. 187, pl. 

1, figs. 12-14. Kaminski et al. (1990) refer to this species as: “Straight tubular fragments with 

extremely coarse agglutination. A proloculus is sometimes present.” 

15
Figures: 

Plate 84, fig. 1, Alaminos Canyon Core PC10 Sample 0-1 cm (Rose-Bengal-stained), side view; 

fig. 2, Mississippi Canyon Core P-69 Sample 0-1 cm, side view; fig. 3, De Soto Canyon control 

sample DGOMB2 S36, side view; fig. 4, Green Canyon Core P-55 Sample 0-1 cm (RoseBengal-stained), side view. 

Scale bars = 50 μm. 

Genus Saccorhiza 

Saccorhiza ramosa (Brady) 

Hyperammina ramosa Brady, 1879, p. 33, pl. 3, figs. 14-15. 

Hyperammina ramosa Brady – Brady, 1884, p. 261, pl. 23, figs. 15-19. 

Saccorhiza ramosa (Brady) – Eimer and Fickert, 1899, p. 670. 

Saccorhiza ramosa (Brady) – Cushman, 1918, p. 81, 82, pl. 30, figs. 3, 4. 

Saccorhiza ramosa (Brady) – Barker, 1960, p. 46, pl. 23, figs. 15-19. 

Saccorhiza ramosa (Brady) – Poag, 1981, p. 82, pl. 15, fig. 4; pl. 16, figs. 4a, b. 

Saccoriza ramosa (Brady) – Denne, 1990, pl. 12, fig. 5. 

Saccorhiza ramosa (Brady) – Jones, 1994, p. 33, pl. 23, figs. 15-19. 

Morphology: Shell a tubular chamber, straight, rounded, or branched; rounded proloculus (rarely 

seen in our specimens); shell wall composed of cemented sand grains, and frequently, numerous 

sponge spicules. 

Figures: 

Plate 160, figs. 1-3, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 4, Alaminos 

Canyon Boxcore B19.1 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

2.2.2. Order Lituolida 

Superfamily Ammodiscacea 

Family Ammodiscidae 

Genus Ammodiscus 

Ammodiscus tenuis (Brady) 

Trochammina (Ammodiscus) tenuis Brady, 1884, p. 51, pl. 38, figs. 4-6. 

Ammodiscus incertus (d’Orbigny) – Cushman, 1918, p. 95, 96, pl. 39, figs. 1-8. 

Involutina tenuis (Brady) – Barker, 1960, p. 78, pl. 38, figs. 5, 6. 

Ammodiscus tenuis (Brady) – Charnock and Jones, 1990, p. 155, pl. 2, fig. 8; pl. 14, fig. 6. 

Ammodiscus tenuis (Brady) – Jones, 1994, p. 43, pl. 38, figs. 4-6. 

16
Morphology: Shell large; planispirally coiled with several overlapping convolutions. 

Remarks: Brady’s fig. 6 appears to show growth lines. None can be seen on any of our 

specimens. 

Figures: 

Plate 5, figs. 1-2, Alaminos Canyon Boxcore B7.1 Sample 0-1 cm, side view; figs. 3-5, Alaminos 

Canyon Boxcore B8.1 Sample 0-2 cm, side view; fig. 6, Alaminos Canyon Boxcore B6 bulk 

sample, side view. 

Scale bars = 50 μm. 

Genus Ammolagena 

Ammolagena clavata (Jones and Parker)

Trochammina irregularis var. clavata Jones and Parker, 1860, p. 304. 

Webbina clavata (Jones and Parker) – Brady, 1884, p. 349, pl. 41, figs. 12-16. 

Ammolagena clavata (Jones and Parker) – Eimer and Fickert, 1899, p. 673. 

Ammolagena clavata (Jones and Parker) – Cushman, 1918, p. 89, 90, pl. 34, figs. 2-5; pl. 35, 

figs. 1-3. 

Ammolagena clavata (Jones and Parker) – Barker, 1960, p. 84, pl. 41, figs. 12-16. 

Ammolagena clavata (Jones and Parker) – Charnock and Jones, 1990, p. 155, pl. 2, fig. 9; pl. 14, 

fig. 7. 

Ammolagena clavata (Jones and Parker) – Kaminski et al., 1990, p. 365, pl. 3, fig. 1. 

Ammolagena clavata (Jones and Parker) – Jones, 1994, p. 46, pl. 41, figs. 12-16. 

Ammolagena clavata (Jones and Parker) – Bender, 1995, p. 41, pl. 2, figs. 9a, 9b. 

Ammolagena clavata (Jones and Parker) – Kuhnt et al., 2000, p. 278, pl. 12, figs. 5-9. 

Morphology: Shell a single ovate chamber with a tubular extension; commonly attached to 

planktonic Foraminifera in our samples. 

Remarks: In some specimens the ovate chamber may be unattached (plate-figs. 2, 4) and may 

have been removed by mechanical breakage (plate-fig. 3) leaving only the tubular extension. In 

the figures of Kuhnt et al. (2000) the ovate chamber is also absent. Bender (1995) noted that 

some specimens show an ovoid central chamber with two tubular exits leading to terminal 

apertures. This is the type species by original monotypy of the genus Ammolagena Eimer and 

Fickert 1899. 

Figures: 

Plate 6, figs. 1-2, Green Canyon Core P-55 Sample 0-1 cm, side view; fig. 3, De Soto Canyon 

control sample DGOMB2 S36, side view; fig. 4, Alaminos Canyon Core PC5 Sample 1-2 cm, 

side view. 

Scale bars = 50 μm. 

17
Genus Glomospira 

Glomospira gordialis (Jones and Parker) 

Trochammina squamata var. gordialis Jones and Parker, 1860, p. 304. 

Ammodiscus gordialis (Jones and Parker) – Brady, 1884, p. 333, pl. 38, figs. 7-9. 

Glomospira gordialis (Jones and Parker) – Cushman, 1918, p. 99, 100, pl. 36, figs. 7-9. 

Glomospira gordialis (Jones and Parker) – Barker, 1960, p. 78, pl. 38, figs. 7-9. 

Glomospira gordialis (Jones and Parker) – Charnock and Jones, 1990, p. 156, pl. 2, fig. 12; pl. 

14, fig. 9. 

Glomospira gordialis (Jones and Parker) – Jones, 1994, p. 43, pl. 38, figs. 7-9. 

Morphology: Shell coiled irregularly rather than streptospirally; coiling in some specimens 

tending to be planispiral. 

Remarks: This is the type species by original monotypy of the genus Glomospira Rzehak1885. 

It was lectotypified by Berggren and Kaminski (1990). 

Figures: 

Plate 69, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Glomospira irregularis (Grzybowski) 

Ammodiscus irregularis Grzybowski, 1898, p. 285, pl. 11, figs. 2-3. 

Ammodiscus demarginatus Grzybowski, 1898, p. 284-285, pl. 10, fig. 34. 

Glomospira irregularis (Grzybowski) – Charnock and Jones, 1990, p. 157, pl. 2, fig. 14; pl. 14, 

fig. 11. 

Glomospira irregularis (Grzybowski) – Kaminski et al., 1990, p. 365-366, pl. 3, fig. 3. 

Morphology: Shell coiled haphazardly. 

Remarks: Charnock and Jones (1990) retain the name Glomospira irregularis over G. 

demarginatus despite the latter having priority by page, because according to current I.C.Z.N. 

rules, this is not a means of establishing seniority. 

Figures: 

Plate 70, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 2, De Soto Canyon 

control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

18
Genus Usbekistania 

Usbekistania charoides (Jones and Parker) 

Trochammina squamata var. charoides Jones and Parker, 1860, p. 304. 

Ammodiscus charoides (Jones and Parker) – Brady, 1884, p. 334, pl. 38, figs. 10-16. 

Glomospira charoides (Jones and Parker) – Cushman, 1918, p. 100, 101, pl. 36, figs. 10-15. 

Glomospira charoides (Jones and Parker) – Barker, 1960, p. 78, pl. 38, figs. 10-16. 

Usbekistania charoides (Jones and Parker, 1860) – Charnock and Jones, 1990, p. 158, pl. 2, figs. 

17-19; pl. 14, fig. 13. 

Glomospira charoides (Jones and Parker) – Denne, 1990, pl. 6, fig. 15. 

Glomospira charoides (Jones and Parker) – Kaminski et al., 1990, p. 365, pl. 3, fig. 2. 

Usbekistania charoides (Jones and Parker, 1860) – Jones, 1994, p. 43, pl. 38, figs. 10-16. 

Morphology: Shell streptospirally coiled to form a subglobular mass. 

Remarks: Jones states that this species was referred to Usbekistania by Charnock and Jones 

(1990) and comments: “It has recently been lectotypified by Berggren and Kaminski (1990). It 

is also, by synonymy with the originally designated Glomospirella (Usbekistania) mubarakensis

Suleymanov 1960, the type species of Usbekistania Suleymanov 1960. Usbekistania differs 

from the irregularly coiled Glomospira in being strictly streptospiral, that is in being 

characterized by regular changes in the axis of coiling.” 

Figures: 

Plate 183, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, Alaminos 

Canyon Core PC5 Sample 1-2 cm, side view; fig. 3, De Soto Canyon control sample DGOMB2 

S36, side view; figs. 4-6, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Superfamily Hormosinacea 

Family Hormosinidae 

Genus Hormosinella 

Hormosinella distans (Brady) 

Lituola (Reophax) distans Brady, 1881, p. 50. 

Reophax distans (Brady) – Brady, 1884, p. 296, pl. 31, figs. 18-22. 

Reophax distans (Brady) – Cushman, 1920, 12, pl. 3, figs. 5, 6. 

Reophax distans delicatulus Cushman – Parker, 1954, p. 482, pl. 1, fig. 4. 

Reophax distans (Brady) – Barker, 1960, p. 64, pl. 31, figs. 18-22. 

Hormosinella distans (Brady) – Charnock and Jones, 1990, p. 163-164, pl. 4, figs. 10-11; pl. 15, 

fig. 11. 

19
Hormosinella distans (Brady) – Kaminski et al., 1990, p. 366, pl. 3, fig. 10. 

Hormosinella distans (Brady) – Jones, 1994, p. 38, pl. 31, figs. 18-22. 

Morphology: Chambers vary from rounded and globular to fusiform and tapering; linked by 

stoloniferous tubes. 

Remarks: Specimens of this study were all unilocular. The specimen illustrated in plate 79, fig. 

1 matches Cushman’s Reophax distans var. delicatulus (1920), and that in plate 79, fig. 2 

matches his R. distans var. turbo (1920). This is the type species of the genera Hormosinella

Shchedrina 1969 and Cadminus Saidova 1970. 

Figures: 

Plate 79, figs. 1-3, De Soto Canyon control sample DGOMB2 S36, side view; fig. 4, Green 

Canyon Core P-57 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Hormosinella guttifera (Brady)

Lituola (Reophax) guttifera Brady, 1881, p. 49. 

Reophax guttifera (Brady) – Brady, 1884, p. 295, pl. 31, figs. 10-15. 

Reophax guttifera (Brady) – Barker, 1960, p. 64, pl. 31, figs. 10-15. 

Hormosinella guttifera (Brady) – Jones, 1994, p. 38, pl. 31, figs. 10-15. 

Morphology: A series of pyriform chambers tapering into stoloniferous tubes which connect the 

chambers together. 

Figures: 

Plate 80, figs. 1-2, south of Mississippi Canyon control sample DGOMB2 C7, side view; fig. 3, 

Green Canyon Core P-57 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Reophax 

Reophax agglutinatus Cushman

Reophax scorpiurus Dénys de Montfort – Brady, 1884, pl. 30, fig. 13. 

Reophax agglutinatus Cushman, 1913, p. 637, pl. 79, figs. 6a, 6b. 

Reophax agglutinatus Cushman – Cushman, 1920, p. 9, 10, pl. 2, figs. 4, 5. 

Reophax agglutinans Cushman – Barker, 1960, p. 62, pl. 30, fig. 13. 

Reophax agglutinans Cushman – Jones, 1994, p. 37, pl. 30, fig. 13. 

Morphology: 3 to 4 gradually enlarging pear-shaped chambers; shell wall composed largely of 

planktonic foraminiferal tests, dominantly Globigerina. 

20
Remarks: Brönnimann and Whittaker (1980) comment that the Challenger specimen shown by 

pl. 30 fig. 13 differs from R. scorpiurus Dénys de Montfort s.s. and R. bradyi. 

Figures: 

Plate 150, figs. 1-2, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

Reophax scorpiurus Dénys de Montfort 

Reophax scorpiurus Dénys de Montfort, 1808, p. 331. 

Reophax scorpiurus Dénys de Montfort – Brady, 1884, p. 291, pl. 30, figs. 12, 15-17. 

Reophax scorpiurus Dénys de Montfort – Cushman, 1920, p. 6, 7, pl. 1, figs. 5-7. 

Reophax scorpiurus Dénys de Montfort – Phleger and Parker, 1951, p. 3, pl. 1, figs. 7, 8. 

Reophax scorpiurus Dénys de Montfort – Parker, 1954, p. 484, pl. 1, fig. 11. 

Reophax scorpiurus Dénys de Montfort – Barker, 1960, p. 62, pl. 30, figs. 12, 15-17. 

Reophax scorpiurus Dénys de Montfort – Brönnimann and Whittaker, 1980, p. 261, figs. 1-7, 12, 

17 (neotype). 

Reophax scorpiurus Dénys de Montfort – Jones, 1994 

Morphology: 3 to 4 pear-shaped asymmetric chambers increasing rapidly in size, in an irregular 

series; coarsely agglutinated shell wall; aperture terminal and rounded on a neck. 

Remarks: This species is conspecific with the neotype assigned by Brönnimann and Whittaker 

(1980). 

Figures: 

Plate 151, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Subreophax

Subreophax monile (Brady) 

Trochammina (Hormosina) monile – Brady, 1881, p. 52, pl. 39, figs. 10-13. 

Hormosina monile (Brady) – Brady, 1884, p. 328, pl. 39, figs. 10-13. 

Hormosina monile (Brady) – Cushman, 1920, p. 30, pl. 6, fig. 4. 

Hormosina monile (Brady) – Barker, 1960, p. 80, pl. 39, figs. 10-13. 

Subreophax monile (Brady) – Jones, 1994, p. 44, pl. 39, figs. 10-13. 

Morphology: Uniserial series of chambers, straight to curved; chambers rounded to globular, 

may be the same size throughout the series or increasing slightly in size; shell wall rough to 

hispid; round aperture is located on the top of a collar. 

21
Figures: 

Plate 174, fig. 1, Green Canyon Core P-51 Sample 0-1 cm, side view; figs. 2-3, Green Canyon 

Core P-51 Sample 1-2 cm, side view; figs. 4a-b, Green Canyon Core P-51 Sample 1-2 cm, 4a: 

side view, 4b: tilted side view. 

Scale bars = 50 μm. 

Genus Archimerismus 

Archimerismus subnodosus (Brady) 

Hyperammina subnodosa Brady, 1884, p. 259, pl. 23, figs. 11-14. 

Hyperammina subnodosa Brady – Cushman, 1918, p. 76, 77, pl. 29, figs. 7, 8. 

Hyperammina subnodosa Brady – Barker, 1960, p. 46, pl. 23, figs. 11-14. 

Archimerismus subnodosus (Brady) – Jones, 1994, p. 33, pl. 23, figs. 11-14. 

Morphology: Shell elongate; elongate tubular chambers, irregularly increasing in size, separated 

by constrictions in the shell wall that do not form true septa. 

Remarks: Hyperammina subnodosa is the type species of the genus Archimerismus (Loeblich 

and Tappan, 1984). Archimerismus differs from Hyperammina in having incipient septa. 

Figures: 

Plate 12, fig. 1, Alaminos Canyon Core PC5 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Hormosina 

Hormosina pilulifera (Brady) 

Reophax pilulifera Brady, 1884, p. 292, pl. 30, figs. 18-20. 

Reophax pilulifer Brady – Cushman, 1920, p. 7, pl. 2, fig. 1. 

Reophax pilulifer Brady – Barker, 1960, p. 62, pl. 30, figs. 18-20. 

Hormosina pilulifera Brady – Charnock and Jones, 1990, p. 162, pl. 4, fig. 5; pl. 15, fig. 7. 

Hormosina pilulifera Brady – Jones, 1994, p. 37, pl. 30, figs. 18-20. 

Morphology: Shell uniserial or slightly curved; symmetrical globular chambers increasing in size 

rapidly; aperture terminal and rounded on a short neck. 

Figures: 

Plate 78, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; figs. 2-3, south of 

Mississippi Canyon control sample DGOMB2 C7, side view; fig. 4, Green Canyon Core P-57 

22
Sample 1-2 cm, side view; fig. 5, Green Canyon Core P-55 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Superfamily Lituolacea 

Family Haplophragmoididae 

Genus Buzasina 

Buzasina ringens (Brady) 

Trochammina ringens Brady, 1879, p. 57, pl. 5, figs. 12a, b. 

Haplophragmoides ringens (Brady) – Cushman, 1920, p. 49, 50, pl. 9, fig. 2. 

Alveolophragmium ringens (Brady) – Parker, 1954, p. 487, pl. 1, fig. 19. 

Alveolophragmium ringens (Brady) – Barker, 1960, p. 82, pl. 40, figs. 17, 18.

Buzasina ringens (Brady) – Jones, 1994, p. 45, pl. 40, figs. 17, 18. 

Morphology: Shell planispirally enrolled with rapidly enlarging and strongly overlapping 

chambers; 5 chambers; shell wall thin; areal and slit-like aperture. 

Remarks: Five chambers have been reported to be in each whorl of this species and are clearly 

seen in Challenger pl. 40 fig. 18. Our specimens show only two to four chambers in the final 

whorl. This is the type species of the genus Buzasina Loeblich and Tappan 1985. 

Figures: 

Plate 31, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, Alaminos 

Canyon Boxcore B6 bulk sample, side view; fig. 3, De Soto Canyon control sample DGOMB2 

S36, side view. 

Scale bars = 50 μm. 

Genus Cribrostomoides 

Cribrostomoides subglobosus (Cushman) 

Lituola subglobosa Sars, 1868 (1869), p. 250 (nomen nudum); Sars, 1872, p. 253. 

Haplophragmium latidorsatum (Bornemann) - Brady, 1884, p. 307, pl. 34, figs. 8-10. 

Haplophragmoides subglobosum (Sars) – Cushman, 1910, p. 105, text figs. 162-164. 

Alveophragmium subglobosum (Sars) – Parker, 1954, p. 487, pl. 2, figs. 1, 2. 

Alveophragmium subglobosum (Sars) – Barker, 1960 

Cribrostomoides subglobosus (Cushman) – Jones, 1994, p. 40, pl. 34, figs. 8-10. 

Morphology: Shell planispiral; periphery broadly rounded; chambers highly inflated; aperture a 

slit lying obliquely at the base of the final chamber. 

23
Figures: 

Plate 50, figs. 1a-b, Green Canyon (Block 272) Core P-57 Sample 1-2 cm, 1a: side view, 1b: 

edge view. 

Scale bars = 50 μm. 

Genus Haplophragmoides

Haplophragmoides sp. cf. H. kirki Wickenden 

Haplophragmoides kirki Wickenden, 1932, p. 85, pl. 1, figs. 1a-c. 

Haplophragmoides kirki Wickenden – Verdenius and van Hinte, 1983, p. 193, pl. 5, fig. 13. 

Haplophragmoides kirki Wickenden – Charnock and Jones, 1990, p. 170, pl. 5, figs. 17-18; pl. 

16, fig. 10. 

Haplophragmoides cf. kirki Wickenden – Kaminski et al., 1990, p. 368, pl. 5, figs. 3a, b. 

Morphology: Shell planispiral, involute; 4-5 chambers in the final whorl; quadrate in outline; 

shell wall finely agglutinated; aperture a simple arch at the base of the final chamber. 

Remarks: As with Kaminski et al. (1990), our specimens are more robust than the typical H. 

kirki from the upper Cretaceous of Alberta. 

Figures: 

Plate 75, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: side 

view, c: apertural view. 

Scale bars = 50 μm. 

Genus Veleroninoides

Veleroninoides jeffreysii (Williamson) 

Haplophragmium canariensis d’Orbigny – Brady, 1884, p. 310, pl. 35, figs. 1-3, 5. 

Nonionina jeffreysii Williamson 1858, p. 34, figs. 72, 73. 

Haplophragmoides canariensis (d'Orbigny) - Cushman, 1920, p. 38, pl. 8, fig. 1. 

Cribrostomoides jeffreysii (Williamson) – Haynes, 1973. 

Veleroninoides jeffreysii (Williamson) – Jones, 1994, p. 41, pl. 35, figs. 1-3, 5. 

Morphology: Planispiral, discoid shell; lobulate periphery; broad and deep umbilicus. 

Remarks: This species is flatter than V. wiesneri. V. jeffreysii has been reported from Gulf of 

Mexico seafloor sediments, but in our samples it is found only attached to tubeworms. 

24
Figures: 

Plate 192, fig. 1, Mississippi Canyon tubeworm EC1, digital photomicrograph; fig. 2, Mississippi 

Canyon tubeworm MC2. 

Veleroninoides wiesneri (Parr) 

Trochammina trullissata Brady, 1884, p. 342, pl. 40, figs. 14, 15 (not figs. 13, 16).

Haplophragmoides trullissata (Brady) – Cushman, 1920, p. 43, pl. 9, fig. 5. 

Labrospira wiesneri Parr, 1950, p. 272, pl. 4, figs. 25-26. 

Alveolophragmium wiesneri (Parr) – Parker, 1954, p. 488, pl. 1, fig. 23. 

Alveolophragmium wiesneri (Parr) – Barker, 1960, p. 82, pl. 40, figs. 14,15. 

Cribrostomoides wiesneri (Parr) – Denne, 1990, pl. 4, fig. 11. 

Veleroninoides wiesneri (Parr) – Jones, 1994, p. 45, pl. 40, figs. 14-15. 

Morphology: Shell evolute planispiral; periphery slightly lobulate; about 3 whorls; chambers of 

the earlier whorls seen in the depressed umbilical region, about 7 chambers in the final whorl; 

aperture slit at the base of the final chamber. 

Figures: 

Plate 193, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; figs. 2-3, Green 

Canyon Core P-57 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Family Discamminidae 

Genus Ammoscalaria 

Ammoscalaria tenuimargo (Brady) 

Haplophragmium tenuimargo Brady, 1882, p. 715. 

Haplophragmium tenuimargo Brady – Brady, 1884, p. 303, pl. 33, figs. 13-16. 

Ammobaculites tenuimargo (Brady) - Cushman, 1920, p. 65, pl. 13, figs. 3-5. 

Ammoscalaria tenuimargo (Brady) – Höglund, 1947, p. 151. 

Ammoscalaria tenuimargo (Brady) – Jones, 1994, p. 40, pl. 33, figs. 13-16. 

Morpholgy: Coarsely agglutinated shell, with a planispiral initial coil followed by a rectilinear 

series of chambers; aperture terminal. 

Remarks: This is the type species of the genus Ammoscalaria. In our samples, it is found only 

attached to tubeworms. 

25
Figures: 

Plate 7, fig. 1, Alaminos Canyon tubeworm LS1, digital photomicrograph; fig. 2, Alaminos 

Canyon tubeworm LS4, digital photomicrograph; fig. 3, Alaminos Canyon tubeworm LS3; fig. 

4, higher magnification view of initial coil in fig. 3. 

Genus Discammina

Discammina compressa Goës 

Lituolina irregularis (Römer) var. compressa Goës, 1882, p. 141, pl. 12, figs. 421-423. 

Haplophragmium emaciatum Brady, 1884, p. 305, pl. 33, figs. 26-28. 

Haplophragmoides emaciatum (Brady) – Cushman, 1920, p. 40, 41, pl. 8, fig. 4. 

Discammina compressa (Goës) – Barker, 1960, p. 68, pl. 33, figs. 26-28. 

Discammina compressa (Goës) – Poag, 1981, p. 58, pl. 9, fig. 4; pl. 10, figs. 4a, b. 

Discammina compressa (Goës) – Jones, 1994, p. 40, pl. 33, figs. 26-28. 

Morphology: Shell planispiral; somewhat umbilicate; up to 3 whorls; later chambers inflated; 

shell wall composed of sand grains, roughly finished; aperture a narrow, elongate slit at the base 

of the final chamber. 

Remarks: This is the type species by synonymy with the originally designated D. fallax Lacroix 

1932, of the genus Discammina Lacroix 1932. Goës’ original description of D. compressa and 

Cushman’s description of H. emaciatum both mention a shell wall containing varying amounts of 

sponge spicules. Our specimens, however, contain no spicules. 

Figures: 

Plate 54, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 1a: side view, 1b: side 

view, 1c: edge view; fig. 2, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, edge view; 

figs. 3-4, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: edge view. 

Scale bars = 50 μm. 

Family Lituotubidae 

Genus Lituotuba 

Lituotuba lituiformis (Brady) 

Trochammina lituiformis Brady, 1879, p. 59, pl. 5, fig. 16. 

Trochammina lituiformis Brady – Brady, 1884, p. 342, pl. 40, figs. 4-7. 

Lituotuba lituiformis (Brady) – Cushman, 1920, p. 59, pl. 12, figs. 1, 2. 

Lituotuba lituiformis (Brady) – Barker, 1960, p. 82, pl. 40, figs. 4-7. 

Lituotuba lituiformis (Brady) – Jones, 1994, p. 44, 45, pl. 40, figs. 4-7. 

26
Morphology: Initial end coiled planispirally, followed by uniserial, rounded to globular 

chambers. 

Remarks: This is the type species by subsequent designation of the genus Lituotuba Rhumbler, 

1895. 

Figures: 

Plate 101, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

Family Lituolidae 

Genus Ammobaculites 

Ammobaculites catenulatus Cushman and McCulloch 

Ammobaculites catenulatus Cushman and McCulloch, 1939, p. 90, pl. 7, figs. 11-14. 

Morphology: Shell compressed; initally planispirally coiled, later uncoiled and rectilinear; 

umbilical region somewhat depressed; sutures indistinct; aperture terminal. 

Figures: 

Plate 3, figs. 1a-d, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

side view, 1c: edge view, 1d: apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 

Sample 0-1 cm, 2a: side view, 2b: side view. 

Scale bars = 50 μm. 

Ammobaculites filiformis Earland 

Haplophragmium agglutinans d’Orbigny – Brady, 1884, p. 301, pl. 32, figs. 22-23. 

Ammobaculites agglutinans var. filiformis Earland, 1934, p. 92, pl. 3, figs. 11-13. 

Ammobaculties filiformis Earland – Barker, 1960, p. 66, pl. 32, figs. 22-23. 

Ammobaculites filiformis Earland – Jones, 1994, p. 39, pl. 32, figs. 21?, 22-23.

Morphology: Shell very small; initial end coiled, followed by a rectilinear series of round 

chambers; number of chambers in the series ranges between 4 and 10; aperture terminal and 

rounded. 

Figures: 

Plate 4, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 2, Alaminos Canyon 

Core PC5 Sample 1-2 cm, side view; fig. 3, Alaminos Canyon Core PC5 Sample 0-1 cm, side 

view. 

Scale bars = 50 μm. 

27
Superfamily Haplophragmiacea 

Family Ammosphaeroidinidae 

Genus Adercotryma 

Adercotryma glomeratum (Brady) 

Lituola glomerata Brady, 1878, p. 433, pl. 20, figs. 1a-c. 

Haplophragmium glomerata (Brady) – Brady, 1884, p. 309, pl. 34, figs. 15-18. 

Haplophragmoides glomeratum (Brady) – Cushman, 1920, p. 47, 48, pl. 9, fig. 6. 

Adercotryma glomeratum (Brady) – Parker, 1954, p. 486, pl. 1, fig. 18.

Adercotryma glomeratum (Brady) – Poag, 1981, p. 35, pl. 15, fig. 2; pl. 16, figs. 2a, 2b. 

Adercotryma glomeratum (Brady) – Denne, 1990, pl. 1, fig. 2. 

Adercotryma glomeratum (Brady) – Jones, 1994, p. 41, pl. 34, figs. 15-18. 

 

Morphology: Shell subglobular to elongated in the axis of coiling; streptospirally enrolled; few 

whorls; few chambers in each whorl; chambers elongated in the axis of coiling; aperture a low 

interiomarginal arch located towards the umbilicus. 

Remarks: This is the type species of the genus Adercotryma Loeblich and Tappan 1952. 

Brönnimann and Whittaker (1987), who designated a lectotype for it, point out that the correct 

trivial name is the neuter glomerat-um rather than the feminine glomerat-a. 

Figures: 

Plate 2, figs. 1-3, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

Genus Cystammina 

Cystammina pauciloculata (Brady) 

Trochammina pauciloculata Brady, 1879, p. 58, pl. 5, figs. 13-14. 

Trochammina pauciloculata Brady – Brady, 1884, p. 344, pl. 41, figs. 1, 2. 

Ammochilostoma pauciloculata (Brady) – Cushman, 1920, p. 86. 

Cystammina pauciloculata (Brady) – Jones, 1994, p. 45. pl. 41, fig. 1. 

Morphology: Shell ovoid, obscurely spiral; early chamber hidden, only 3-4 chambers visible; 

sutures distinct, depressed; shell wall finely arenaceous, smooth; aperture an elongate arched slit 

at the base of the final chamber. 

Figures: 

Plate 51, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 2, Alaminos Canyon 

Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

28
Superfamily Spiroplectamminacea 

Family Spiroplectamminidae 

Genus Orectostomina 

Orectostomina camposi (Brönnimann and Beurlen) 

Spiroplectamminoides camposi Brönnimann and Beurlen, 1977, p. 87, 88, pl. 3, figs. 1-3, 6-8, 

10. 

Orectostomina camposi (Brönnimann and Beurlen) – Loeblich and Tappan, 1987, p. 111, pl. 

119, figs. 13-17. 

Morphology: Shell elongate; rounded periphery; early chambers arranged in a planispiral coil 

followed by a short biserial stage; biserial chambers increasing rapidly in height; shell wall finely 

agglutinated; aperture an oblong crescent-like opening surrounded by thin, strongly protruding 

lips in the center of the septal face of the final chamber. 

Figures: 

Plate 115, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: dorsal view, 

1b: ventral view, 1c: edge view. 

Scale bars = 50 μm. 

Genus Spiroplectammina 

Spiroplectammina sp. 

Morphology: Shell elongate, some specimens slightly curved; initial end broadly rounded and 

slightly tapered; early chambers arranged in a planispiral coil, later chambers biserially arranged; 

shell wall finely agglutinated; aperture at the base of the final chamber. 

Remarks: This species resembles Cushman’s Spiroplecta wrightii (1922, p. 5) but not Silvestri’s 

illustration of the type specimens (Loeblich and Tappan, 1987, pl. 120, figs. 1-6). 

Figures: 

Plate 170, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, De Soto 

Canyon control sample DGOMB2 S36, side view; fig. 3, De Soto Canyon control sample 

DGOMB2 S36, side view; fig. 4, De Soto Canyon control sample DGOMB2 S36, side view. 

Scale bars = 50 μm. 

29
Family Pseudobolivinidae 

Genus Parvigenerina 

Parvigenerina arenacea (Heron-Allen and Earland) 

Bifarina porrecta (Brady) var. arenacea Heron-Allen and Earland, 1922, p. 132, pl. 4, figs. 23-

26. 

Parvigenerina arenacea (Heron-Allen and Earland) – Loeblich and Tappan, 1987, p. 116, pl. 

123, figs. 13-16. 

Morphology: Shell elongate; early chambers arranged biserially, later chambers becoming 

uniserial; shell wall finely agglutinated; aperture terminal on a distinct neck. 

Remarks: Charnock and Jones (1990) noted that this genus differs from Plectinella Marie in 

becoming semi-uniserial to uniserial in later stages. 

Figures: 

Plate 123, figs. 1a-b, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Superfamily Verneuilinacea 

Family Prolixoplectidae 

Genus Karrerulina

Karrerulina apicularis (Cushman) 

Gaudryina siphonella Reuss - Brady, 1884, p. 382, pl. 46, figs. 17-19. 

Gaudryina apicularis Cushman, 1911, p. 70, textfig. 110. 

Gaudryina apicularis Cushman – Cushman, 1922, p. 72, 73, pl. 8, fig. 4. 

Plectina apicularis (Cushman) – Cushman, 1937, p. 110, pl. 12, figs. 22-26 

Plectina apicularis (Cushman) – Phleger and Parker, 1951, p. 6, pl. 3, fig. 3. 

Plectina apicularis (Cushman) – Parker, 1954, p. 495, pl. 3, fig. 18. 

Karreriella (Karrerulina) apicularis (Cushman) – Barker, 1960, p. 94, pl. 46, figs. 17-19. 

Karreriella apicularis (Cushman) – Pflum and Frerichs, 1976, pl. 1, fig. 2. 

Karreriella apicularis (Cushman) – Poag, 1981, pl. 15, fig. 5; pl. 16, figs. 5a, b. 

Karrerulina conversa (Grzybowski) – Charnock and Jones, 1990, p. 195, 196, pl. 12, fig. 19; pl. 

25, fig. 10. 

Karrerulina apicularis (Cushman) – Denne, 1990, pl. 7, fig. 10. 

Karrerulina conversa (Grzybowski) – Jones, 1994, p. 51, pl. 46, figs. 17-19. 

Karrerulina apicularis (Cushman) – Bender, 1995, p. 46, pl. 6, fig. 1; pl. 11, fig. 2. 

Karrerulina apicularis (Cushman) – Kuhnt et al., 2000, p. 279, pl. 8, figs. 6-9. 

30
Morphology: Shell elongate; early chambers arranged triserially, later chambers becoming 

biserial, biserial portion making up more than half of the shell; sutures depressed; shell wall 

coarsely agglutinated; aperture rounded at the end of a neck-like extension on the final chamber. 

Remarks: It is the type species, by synonymy with the originally designated Gaudryina 

apicularis Cushman 1911, of the genus Karrerulina Finlay 1940. 

Figures: 

Plate 87, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, De Soto 

Canyon control sample DGOMB2 S36, side view with a broken final chamber; figs. 3-4, 

Alaminos Canyon Boxcore B6 bulk sample, side view. 

Scale bars = 50 μm. 

Prolixoplecta parvula Cushman

Textularia parvula Cushman, 1922, p. 11, pl. 6, figs. 1, 2. 

Textularia parvula Cushman – Phleger and Parker, 1951, p. 5, pl. 2, fig. 8. 

Prolixoplecta parvula (Cushman) – Denne, 1990, pl. 10, fig. 13. 

Morphology: Shell elongate, tapering from pointed initial end to rounded apertural end; 

numerous distinct chambers; sutures depressed; wall agglutinated; aperture at the base of the 

final chamber. 

Figures: 

Plate 136, fig. 1, Mississippi Canyon Core P-69 Sample 0-1 cm, side view; fig. 2, Mississippi 

Canyon Core P-69 Sample 1-2 cm, side view; figs. 3-4, Green Canyon (Block 272) Core P-33 

Sample 0-1 cm, side view; fig. 5, Green Canyon (Block 272) Core P-51 Sample 0-1 cm, side 

view. 

Scale bars = 50 μm. 

Family Verneuilinidae 

Genus Gaudryina 

Gaudryina minuta Earland

Gaudryina minuta Earland, 1934, p. 121, pl. 5, figs. 45, 46. 

Gaudryina cf. minuta Parker, 1954, p. 493, pl. 15, 16. 

Morphology: Shell elongate, tapering to the initial end; early chambers arranged triserially, later 

chambers arranged biserially; sutures depressed; aperture a textularian slit on the inner margin of 

the final chamber. 

31
Remarks: Parker (1954) states that her Gaudryina cf. minuta specimens are larger and more 

elongate than Gaudryina exilis Cushman and Bronnimann and in some cases reach a greater 

length than that given by Earland. Gaudryina exilis Cushman and Bronnimann differs from G. 

minuta Earland in its larger size, much shorter triserial stage and shorter chambers. Some of our 

specimens reach 0.55 mm in length, thus exceeding the maximum length of 0.4 mm given by 

Parker (1954). 

Figures: 

Plate 66, figs. 1, 4, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2-3, Green 

Canyon (Block 272) Core P-57 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Barbourinella 

Barbourinella atlantica (Bermúdez) 

Barbourina atlantica Bermúdez, 1939, p. 10, pl. 1, figs. 1-4. 

Barbourinella atlantica (Bermúdez) – Loeblich and Tappan, 1987, p. 137, pl. 144, figs. 15, 16. 

Morphology: Shell triangular in section; triserial throughout; shell wall coarsely agglutinated; 

aperture located in the center of the final chamber, circular to elongate on a short neck. 

Remarks: Bermúdez’s original description of the aperture states that it is in the center of the 

terminal face of the final chamber, circular, and protected by a short and protracted neck. Our 

specimens match this description except the apertures are elongated rather than circular. We 

consider this to be within the natural range of morphological variation. 

Figures: 

Plate 13, figs. 1, 3, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: 

apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 2a: side 

view, 2b: apertural view. 

Scale bars = 50 μm. 

Superfamily Ataxophragmiacea 

Family Globotextulariidae 

Genus Tetrataxiella 

Tetrataxiella sp.

Morphology: Shell spiral, trochoid; 3-5 whorls with 4 chambers in each whorl; periphery 

subacute; ventral surface with a sunken umbilicus; sutures on ventral surface straight, slightly 

depressed; shell wall finely agglutinated with occasional larger sand grains; aperture 

interiomarginal and umbilical loop. 

32
Remarks: The species is placed in Tetrataxiella Seiglie 1964 (conical, quadriserial throughout, 

aperture an interiomarginal arch opening into the umbilicus; Caribbean). Charnock and Jones 

(1990) noted that Brönnimann and Whittaker (1988, p. 88) synonymised Tetrataxiella Seiglie 

with Gravellina Brönnimann, 1953. 

Figures: 

Plate 175, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

ventral view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 2, dorsal view; 

figs. 3a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 3a: side view, 3b: ventral 

view; fig. 4, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Family Textulariellidae 

Genus Textulariella 

Textulariella barrettii (Jones and Parker) 

Textularia barrettii Jones and Parker, 1876, p. 99, text-figs. 

Textularia trochus d’Orbigny – Brady, 1884, p. 367, pl. 43, fig. 17; pl. 44, fig. 3. 

Textularia barrettii Jones and Parker – Brady, 1884, p. 367, pl. 44, figs. 6-8. 

Textularia barrettii Jones and Parker – Cushman, 1922, p. 20, pl. 3, figs. 3-6. 

Textulariella barrettii (Jones and Parker) – Cushman, 1937, p. 66, pl. 7, figs. 5-8. 

Textulariella barrettii (Jones and Parker) – Barker, 1960, p. 88, pl. 43, fig. 17; p. 90, pl. 44, figs. 

3, 6-8. 

Textulariella barrettii (Jones and Parker) – Denne, 1990, pl. 13, fig. 10. 

Textulariella barrettii (Jones and Parker) – Jones, 1994, p. 48, pl. 43, fig. 17; p. 49, pl. 44, figs. 

3, 6-8. 

Morphology: Shell conical; circular in transverse section; sutures distinct, frequently slightly 

raised; aperture a low, elongate opening at the inner margin of the final chamber.

Remarks: This is the type species of the genus Textulariella Cushman 1927. It has been 

lectotypified by Loeblich and Tappan (1964). 

Figures: 

Plate 179, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

33
2.2.3. Order Trochamminida 

Superfamily Trochamminacea 

Family Trochamminidae 

Genus Paratrochammina 

Paratrochammina challengeri Brönnimann and Whittaker 

Lituola nautiloidea Lamarck var. globigeriniformis Parker and Jones, 1865, p. 407, pl. 15, figs. 

46-47; pl. 17, figs. 96-98. 

Trochammina globigeriniforme (Parker and Jones) – Brady, 1884, p. 312, pl. 35, figs. 10a-c (not 

fig. 11). 

Ammoglobigerina globigeriniformis (Parker and Jones) – Barker, 1960, p. 72, pl. 35, fig. 10. 

Paratrochammina challengeri Brönnimann and Whittaker, 1988, p. 43, 44, figs. 16H-K. 

Paratrochammina challengeri Brönnimann and Whittaker – Bender, 1995, p. 47, pl. 5, fig. 9. 

Paratrochammina challengeri Brönnimann and Whittaker – Jones, 1994, p. 41, 42, pl. 35, fig. 

10. 

Morphology: Shell trochospiral; lobulate outline, rounded periphery; 3 whorls, 4 chambers in 

final whorl; chambers subglobular, increasing rapidly in size; deep umbilical depression; 

depressed sutures; coarsely agglutinated; asymmetric, crescentic, intermarginal aperture. 

Remarks: Agglutinated particles in shells of of this species may be very coarse, and may include 

planktonic foraminiferal shells (e.g., plate 122, fig. 1). 

Figures: 

Plate 122, fig. 1, Alaminos Canyon Boxcore B6 bulk sample, ventral view; fig. 2, De Soto 

Canyon control sample DGOMB2 S36, dorsal view. 

Scale bars = 50 μm. 

Genus Portatrochammina

Portatrochammina antarctica (Parr) 

Trochammina inflata (Montagu) – Wiesner, 1931, p. 111, pl. 17, figs. 201a-c (not Nautilus 

inflatus Montagu, 1808).

Trochammina wiesneri Parr, 1950, p. 279, pl. 5, fig. 14. 

Trochammina antarctica Parr, 1950, p. 280, pl. 5, figs. 1, 2. 

Portatrochammina antarctica antarctica (Parr) - Brönnimann and Whittaker, 1988, p. 64-66, 

figs. 25A-C; 26A-G. 

Portatrochammina antarctica wiesneri (Parr) – Brönnimann and Whittaker, 1988, p. 68, 69, figs. 

25D-F; 26H-K; 27D-I. 

34
Morphology: Shell trochospiral; convex on dorsal side, shallow concave on ventral side; slightly 

lobulate outline, rounded periphery; 3 whorls, 5-6 chambers in final whorl; sutures depressed; 

aperture interiomarginal arch, covered by a large umbilical flap. 

Remarks: Previous umbilical flaps can be seen in some specimens (e.g., plate 132, fig. 5). Parr 

(1950) differentiated between Trochammina antarctica and T. wiesneri saying that T. wiesneri

has a smoother, flatter shell and lower chambers causing the periphery to be subacute rather than 

broadly rounded as it is in T. antarctica. Parr (1950) also noted the characteristic greater length 

of the final chamber. The SEM photograph of the holotype (fig. 25E) included in Brönnimann 

and Whittaker (1988) shows that the periphery of T. wiesneri is rounded, not subacute. The 

other features may also be variable and hence Brönnimann and Whittaker (1988) concluded that 

the morphological differences between the two species were not great and gave them subspecific 

status. Since these slightly diffrenet morphologies are found in co-occurring individuals, and 

since the roundness of the margin is a gradational feature, we see no justification for recognizing 

sympatric subspecies. However, as Bronnimann and Whittaker were the first revisers of the 

original nomenclature, their designated species name (Portatrochammina antarctica) remains 

valid. 

Figures: 

Plate 132, fig. 1, Alaminos Canyon Core PC15 Sample 0-1 cm, dorsal view; fig. 2, Alaminos 

Canyon Core PC15 Sample 0-1 cm, ventral view; fig. 3, Alaminos Canyon Core PC15 Sample 

10-11 cm, ventral view; fig. 4, De Soto Canyon control sample DGOMB2 S36, dorsal view; fig. 

5, De Soto Canyon control sample DGOMB2 S36, ventral view; fig. 6, Green Canyon (Block 

272) Core P-51 Sample 0-1 cm, ventral view. 

Scale bars = 50 μm. 

Genus Pseudotrochammina

Pseudotrochammina sp.

Morphology: Shell trochospiral; periphery rounded, lobulate; 7-8 chambers in the final whorl; 

depressed umbilicus; sutures depressed; aperture narrow, rimmed, elongate with everted border, 

situated on apertural face parallel to basal suture. 

Remarks: This partly resembles P. echolsi Brönnimann and Whittaker (1988), but it has a greater 

number of chambers in the final whorl and the location of the aperture. 

Figures: 

Plate 142, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: dorsal view, 

1b: ventral view, 1c: edge view. 

Scale bars = 50 μm. 

35
Genus Deuterammina 

Deuterammina rotaliformis (Heron-Allen and Earland) 

Trochammina rotaliformis Heron-Allen and Earland, 1911, p. 309. 

Trochammina rotaliformis Wright - Cushman, 1920, p. 77, pl. 16, figs. 1, 2. 

Deuterammina (Deuterammina) roatliformis (Heron-Allen and Earland) - Bronnimann and 

Whittaker, 1983, p. 348, text-figs. 1-3, 25. 

Morphology: Low-trochospiral, finely agglutinated shell, with slightly lobulate periphery; broad 

and deep umbilicus. 

Remarks: The shell is fragile. In our samples, the species is found only attached to tubeworms. 

Figures: 

Plate 53, fig. 1, Green Canyon tubeworm EC3, digital photomicrograph; fig. 2, Green Canyon 

tubeworm EC3, digital photomicrograph; fig. 3, Green Canyon tubeworm EC3; fig. 4, same 

specimen as in fig. 2. 

2.2.4. Order Textulariida 

Superfamily Textulariacea 

Family Eggerellidae 

Genus Dorothia 

Dorothia scabra (Brady) 

Gaudryina pupoides Brady, 1870, p. 300, pl. 8, fig. 5.Gaudryina scabra Brady, 1884, p. 381, pl. 

46, fig. 7. 

Textularia aspera Brady, 1884, pl. 44, figs. 12-13. 

Gaudryina scabra Brady – Cushman, 1922, p. 68, pl. 11, figs. 6, 7. 

Dorothia scabra (Brady) – Cushman, 1937, p. 98, pl. 11, figs. 3, 4. 

Dorothia scabra (Brady) – Barker, 1960, p. 90, pl. 44, figs. 12, 13; p. 94, pl. 46, pl. 46, fig. 7. 

Dorothia scabra (Brady) – Charnock and Jones, 1990, p. 190, pl. 12, fig. 2; pl. 24, fig. 2. 

Dorothia scabra (Brady) – Jones, 1994, p. 49, pl. 44, figs. 12, 13; p. 50, 51, pl. 46, fig. 7. 

“Dorothia” scabra (Brady) – Bender, 1995, p. 44, pl. 6, figs. 6, 7; pl. 10, fig. 3. 

Morphology: Shell elongate, tapering to the initial end; early chambers arranged triserially, later 

chambers becoming biserial; 3-5 pairs of chambers in biserial portion, chambers inflated; sutures 

distinct, depressed; shell wall rather coarsely arenaceous; aperture simple arch at the inner 

margin of the final chamber. 

Remarks: Both coccoliths (plate 57, figs. 1 and 2) and sand grains (plate 57, fig. 3) have been 

observed as the primary agglutinated particles in this species. Our specimens do not have the 

breadth of the figured specimens of other authors. 

36
Figures: 

Plate 57, fig. 1, De Soto Canyon control sample DGOMB2 S36, side view; fig. 2, De Soto 

Canyon control sample DGOMB2 S36, side view with a broken final chamber; fig. 3, Alaminos 

Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Eggerella 

Eggerella bradyi (Cushman) 

Verneuilina pygmaea Egger – Brady, 1884, p. 385, pl. 47, figs. 4-7. 

Verneuilina bradyi Cushman, 1911, p. 54, figs. 87a, b. 

Verneuilina bradyi Cushman – Cushman, 1922, p. 59, pl. 11, fig. 1. 

Verneuilina bradyi Cushman – Thalmann, 1932. 

Eggerella brady (Cushman) – Thalmann, 1937. 

Eggerella bradyi (Cushman) – Barker, 1960, p. 96, pl. 47, figs. 4-7. 

Eggerella bradyi (Cushman) – Denne, 1990, pl. 5, fig. 5. 

Eggerella bradyi (Cushman) – Jones, 1994, p. 51, 52, pl. 47, figs. 4-7. 

Morphology: Shell triserial and pyramidal; chambers rapidly increasing in size, those in final 

whorl inflated; aperture an elongate slit at the base of the final chamber, frequently with a 

thickened lip. 

Remarks: This is the type species of the genus Eggerella Cushman 1933. 

Figures: 

Plate 58, fig. 1, Alaminos Canyon Core PC4 Sample 0-1 cm, side view; fig. 2, Alaminos Canyon 

Core PC5 Sample 0-1 cm, side view; fig. 3, Green Canyon (Block 272) Core P-51 Sample 1-2 

cm, side view. 

Scale bars = 50 μm. 

Genus Karreriella 

Karreriella bradyi (Cushman) 

Gaudryina pupoides d’Orbigny – Brady, 1884, p. 378, pl. 46, figs. 1-4. 

Gaudryina bradyi Cushman, 1911, p. 67, text-fig. 107. 

Gaudryina bradyi Cushman – Cushman, 1922, p. 74, pl. 12, fig. 8. 

Karreriella bradyi (Cushman) – Cushman, 1937, p. 135, pl. 16, figs. 6-11.

Karreriella bradyi (Cushman) – Phleger and Parker, 1951, p. 6, pl. 3, figs. 4. 

Karreriella bradyi (Cushman) – Parker, 1954, p. 495, pl. 3, fig. 11. 

Karreriella bradyi (Cushman) – Barker, 1960, p. 94, pl. 46, figs. 1-4. 

37
Karreriella bradyi (Cushman) – Andersen, 1961, p. 28, pl. 3, figs. 2a, b. 

Karreriella bradyi (Cushman) – Denne, 1990, pl. 7, fig. 9. 

Karreriella bradyi (Cushman) – Jones, 1994, p. 50, pl. 46, figs. 1-4. 

Karreriella bradyi (Cushman) – Bender, 1995, p. 46, pl. 6, fig. 11. 

Karreriella bradyi (Cushman) – Kuhnt et al., 2000, p. 279, pl. 8, figs. 16, 18. 

Morphology: Shell elongate, tapering to blunt initial end; early chambers arranged triserially, 

later chambers becoming biserial, biserial portion making up about three-quarters of the shell; 

sutures depressed; aperture oval, raised lip, close to the inner margin of the final chamber. 

Figures: 

Plate 86, fig. 1, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view; fig. 2, Green 

Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view; fig. 3, Green Canyon (Block 272) 

Core P-55 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Martinottiella

Martinottiella communis (d’Orbigny)

Clavulina communis d’Orbigny, 1846, p. 196, pl. 12, figs. 1, 2. 

Clavulina communis d’Orbigny – Brady, 1884, p. 394, pl. 48, figs. 1-8 (not 9-13). 

Clavulina communis (d’Orbigny) – Cushman, 1922, p. 84, 85, pl. 16, figs. 4, 5. 

Martinottiella bradyana (Cushman) – Barker, 1960, p. 98, pl. 48, figs. 1, 2, 5. 

Martinottiella communis (d’Orbigny) – Barker, 1960, p. 98, pl. 48, figs. 3-4, 6-8. 

Martinottiella communis (d’Orbigny) – Papp and Schmid, 1985, p. 74, 75, pl. 66, figs. 1-8. 

Martinottiella communis (d’Orbigny) – Hermelin, 1989, p. 34, pl. 2, figs. 5-6. 

Martinottiella communis (d’Orbigny) – Denne, 1990, pl. 8, fig. 14. 

Martinottiella communis (d’Orbigny)– Jones, 1994, p. 52, pl. 48, figs. 1-2, ?3, 4-8. 

Martinottiella communis (d’Orbigny) – Bender, 1995, p. 46, pl. 6, fig. 16. 

Morphology: Shell elongate; early portion trochospiral, later becoming uniserial; chambers 

numerous; sutures indistinct, only slightly depressed; shell wall finely agglutinated; aperture 

terminal, rounded to arcuate slit, on a short neck. 

Remarks: Jones comments that Clavulina communis d’Orbigny 1846 is the type species of the 

genus Martinotiella Cushman 1933 (which differs from Clavulina d’Orbigny 1826 in its initially 

trochospiral rather than triserial chamber arrangement). 

Figures: 

Plate 102, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: 

apertural view. 

Scale bars = 50 μm. 

38
Genus Multifidella

Multifidella nodulosa (Cushman) 

Clavulina communis d’Orbigny – Brady, 1884, pl. 48, figs. 9-12 (part). 

Clavulina communis d’Orbigny var. nodulosa Cushman, 1922, p. 85, pl. 18, figs. 1-3. 

Listerella nodulosa (Cushman) – Cushman, 1937, p. 150, pl. 17, figs. 13-19. 

Martinottiella nodulosa (Cushman) – Barker, 1960, p. 98, pl. 48, figs. 9-13. 

Multifidella nodulosa (Cushman) – Loeblich and Tappan, 1964, C277, fig. 185, no. 4. 

Multifidella nodulosa (Cushman) – Loeblich and Tappan, 1987, p. 171, 172, pl. 190, figs. 8, 9. 

Multifidella nodulosa (Cushman) – Jones, 1994, p. 52, 53, pl. 48, figs. 9-13. 

Morphology: Shell elongate; early portion trochospiral, later becoming uniserial; shell wall 

finely agglutinated; aperture terminal, cribrate with elongate slits and bordering lips. 

Remarks: This is the type species of the genus Multifidella Loeblich and Tappan 1961. 

Multifidella differs from Martinottiella in having a cribrate aperture of slits with bordering lips. 

The chambers of our specimens are more rounded and less cylindrical than those of other 

authors. However, they are within the natural range of variation. 

Figures: 

Plate 107, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: 

apertural view. 

Scale bars = 50 μm. 

Family Textulariidae 

Genus Textularia 

Textularia foliacea occidentalis Cushman 

Textularia foliacea Heron-Allen and Earland var. occidentalis Cushman, 1922, p. 16, pl. 2, fig. 

13. 

Textularia foliacea Heron-Allen and Earland var. occidentalis Cushman – Phleger and Parker, 

1951, p. 5, pl. 1, figs. 28, 29. 

Textularia foliacea occidentalis Cushman – Parker, 1954, p. 491, pl. 2, fig. 10. 

Textularia occidentalis Cushman – Andersen, 1961, p. 23, pl. 1, figs. 2a, b. 

Textularia foliacea occidentalis Cushman – Denne, 1990, pl. 13, fig. 7. 

Morphology: Shell triangular to leaf-shaped, tapering to initial end; apertural end truncated; 7-9 

pairs of chambers; sutures frequently obscure; shell wall coarsely agglutinated; aperture elongate 

slit at the base of the inner margin of the final chamber. 

Remarks: Textularia foliacea occidentalis differs from T. foliacea in being broader and less 

tapered with a less acute initial end. 

39
Figures: 

Plate 176, figs. 1, 3, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: 

apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 2a: side 

view, 2b: apertural view. 

Scale bars = 50 μm. 

Textularia sp. cf. T. mayori Cushman 

Textularia mayori Cushman, 1922, p. 23, pl. 2, fig. 3. 

Textularia mayori Cushman – Cushman, 1922, p. 7. 

Textularia mayori Cushman – Phleger and Parker, 1951, p. 5, pl. 2, figs. 1-5. 

Textularia mayori Cushman – Parker, 1954, p. 491, pl. 2, fig. 11. 

Textularia mayori Cushman – Andersen, 1961, p. 22, 23, pl. 1, figs. 6-8. 

Textularia mayori Cushman – Denne, 1990, pl. 13, fig. 8. 

Morphology: Shell increasing rapidly in breadth; initial end rounded, apertural end truncated; 

chambers indistinct; spinose projections, with broken tips, frequently present; aperture an 

elongate opening on the inner margin of the final chamber. 

Remarks: Phleger and Parker (1954) noted that the spinose projections were a varible feature. 

Andersen (1961) illustrated three specimens showing the range of variation in the development 

of the spinose projections including a spineless variant. However, the spines on Cushman’s 

holotype are so much more pronounced (and texturally distinct from the rest of the shell) than 

those on our specimens that we cannot be sure of the identity of our species, which does not 

match the figures of Parker (1954) or Andersen (1961). 

Figures: 

Plate 177, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

Textularia mexicana Cushman 

Textularia mexicana Cushman, 1922, p. 17, pl. 2, fig. 9. 

Textularia mexicana Cushman – Phleger and Parker, 1951, p. 5, pl. 2, figs. 6, 7. 

Textularia mexicana Cushman – Andersen, 1961, p. 23, pl. 1, figs. 4a, b. 

Textularia mexicana Cushman – Denne, 1990, pl. 13, fig. 9. 

Morphology: Shell triangular; periphery sharp; chambers numerous; sutures distinct, raised, 

coalescing at center of shell to form a high ridge, especially in latter half of the shell; aperture 

elongate, semicircular slit at the base of the inner margin of the final chamber. 

40
Figures: 

Plate 178, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

Genus Siphotextularia 

Siphotextularia rolshauseni Phleger and Parker 

Siphotextularia rolshauseni Phleger and Parker, 1951, p. 4, pl. 1, figs. 23, 24. 

Siphotextularia rolshauseni Phleger and Parker – Parker, 1954, p. 492, pl. 2, fig. 14. 

Morphology: Periphery rounded, lobulate; early portion planispirally coiled, later becoming 

biserial; 4-5 pairs of chambers in biserial portion; sutures distinct, depressed, almost at right 

angles to central axis; shell wall finely arenaceous, some larger grains; aperture round, on a 

projecting lip. 

Remarks: Siphotextularia rolshauseni differs from S. affinis in being smaller with a more inflated 

shell, sutures at right angles to the central axis, chambers that do not overlap, and a smaller 

aperture. 

Figures: 

Plate 167, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 2, Alaminos 

Canyon Core PC15 Sample 10-11 cm, side view. 

Scale bars = 50 μm. 

Family Pseudogaudryinidae 

Genus Pseudoclavulina

Pseudoclavulina mexicana (Cushman) 

Clavulina humilis Brady var. mexicana Cushman, 1922, p. 83, 84, pl. 16, figs. 1-3. 

Pseudoclavulina mexicana (Cushman) – Phleger and Parker, 1951, p. 6, pl. 2, figs. 14-16. 

Pseudoclavulina mexicana (Cushman) – Parker, 1954, p. 493, pl. 3, fig. 8. 

Pseudoclavulina mexicana (Cushman) – Andersen, 1961, p. 27, 28, pl. 2, fig. 12. 

Clavulina mexicana (Cushman) – Denne, 1990, pl. 5, fig. 1. 

Pseudoclavulina mexicana (Cushman) – Bender, 1995, p. 48, pl. 7, fig. 15; pl. 12, fig. 3. 

Morphology: Shell elongate; apertural end subacute; early portion sharply triserial, later 

becoming uniserial and subcylindrical; sutures indistinct in triserial portion, slightly depressed in 

uniserial portion; shell wall coarsely agglutinated; aperture at the end of a tubular neck. 

41
Figures: 

Plate 138, fig. 1, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view; figs. 2-3, 

Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Pseudoclavulina serventyi (Chapman and Parr) 

Clavulina parisiensis d’Orbigny – Brady, 1884, pl. 48, figs. 14-16. 

Clavulina serventyi Chapman and Parr, 1935, p. 5, pl. 1, fig. 7. 

Pseudoclavulina serventyi (Chapman and Parr) – Barker, 1960, p. 98, pl. 48, figs. 14-16. 

Pseudoclavulina serventyi (Chapman and Parr) – Jones, 1994, p. 53, pl. 48, figs. 14-16. 

Morphology: Shell elongate; apertural end rounded; early portion triserial, later becoming 

uniserial; sutures depressed; shell wall coarsely agglutinated; aperture terminal, consisting of two 

to three pores. 

Remarks: The type figure and the Challenger specimens of this species show numerous 

chambers in the uniserial portion, whereas our specimens have a maximum of 3-4 such 

chambers. 

Figures: 

Plate 139, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Pseudogaudryina 

Pseudogaudryina atlantica (Bailey) 

Textularia atlantica Bailey, 1851, p. 12, pl., figs. 38-43. 

Verneuilina triquetra (Munster) – Brady, 1884, pl. 47, fig. 18 (not figs. 19-20). 

Gaudryina atlantica (Bailey) – Cushman, 1922, p. 70, pl. 13, figs. 1-3. 

Gaudryina (Pseudogaudryina) atlantica (Bailey) – Phleger and Parker, 1951, p. 6, pl. 2, figs. 

13a, b. 

Gaudryina (Pseudogaudryina) atlantica (Bailey) – Parker, 1954, p. 493, pl. 3, fig. 7. 

Gaudryina atlantica (Bailey) – Barker, 1960, p. 96, pl. 47, fig. 18. 

Pseudogaudryina atlantica (Bailey) – Denne, 1990, pl. 11, fig. 1. 

Pseudogaudryina atlantica (Bailey) – Jones, 1994, p. 52, pl. 47, fig. 18. 

Morphology: Shell elongate, triangular in section; early portion triserial, later portion biserial; 

triserial portion short, biserial portion triangular in shape; shell wall coarsely agglutinated; 

aperture an interiomarginal arch. 

Remarks: This is the type species of the genus Pseudogaudryina Cushman 1936. 

42
Figures: 

Plate 140, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, a: side view, b: 

apertural view. 

Scale bars = 50 μm in fig. 1; 100 μm in fig. 2. 

2.2.5. Order Miliolida 

Superfamily Cornuspiracea 

Family Cornuspiridae 

Genus Cornuspira 

Cornuspira foliacea (Philippi) 

Orbis foliaceus Philippi, 1844, p. 147, pl. 24, fig. 26. 

Cornuspira foliacea (Philippi) – Brady, 1884, p. 199, pl. 11, figs. 5, 6. 

Cornuspira foliacea (Philippi) - Cushman, 1929, p. 79, pl. 20. figs. 3-5. 

Cornuspira foliacea (Philippi) – Jones, 1994, p. 26, pl. 11, figs. 5, 6. 

Morphology: Shell with a proloculus and a coiled, tubular second chamber whose diameter 

increases progressively (and rapidly in adult stage) toward the terminal aperture; somewhat 

flattened. 

Remarks: This is the type species of the genus Cornuspira. In our samples, it is found only 

attached to tubeworms. 

Figures: 

Plate 46, fig. 1, Alaminos Canyon tubeworm LS6 hairy rootlet, digital photomicrograph; fig. 2, 

Alaminos Canyon tubeworm LS6, digital photomicrograph; fig. 3, Alaminos Canyon tubeworm 

LS7; fig. 4, same specimen as in fig. 2; fig. 5, Alaminos Canyon mussel MAC1; fig. 6, Alaminos 

Canyon mussel MAC1, attachment side of foraminifer. 

Cornuspira involvens (Reuss) 

Operculina involvens Reuss, 1850, p. 370, pl. 46, figs. 20a, b. 

Cornuspira involvens (Reuss) – Brady, 1884, p. 200, pl. 11, figs. 1-3. 

Cornuspira involvens (Reuss) – Cushman, 1929, p. 80-81, pl. 20, figs. 6, 8. 

Cornuspira involvens (Reuss) – Barker, 1960, p. 22, fig. 11, figs. 1-3. 

Cornuspira involvens (Reuss) – Jones, 1994, p. 26-27, pl. 11, figs. 1-3. 

Morphology: Shell with a proloculus and a long, coiled, planispiral second chamber of nearly 

equal diameter throughout; aperture an opening at the end of the tube. 

43
Remarks: Many living specimens were observed to be encysted, particularly in sample PC15 

from Alaminos Canyon (e.g., plate 47, figs. 1, 2, 4). On the other hand, many specimens were 

found attached to tubeworms (Plate 48). 

Figures: 

Plate 47, fig. 1, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, side view; fig. 2, living 

and encysted specimen from Alaminos Canyon Core PC15 Sample 0-1 cm, side view; fig. 3, 

living and encysted specimen from Alaminos Canyon Core PC15 Sample 0-1 cm, side view; fig. 

4, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, side view; fig. 5, Green Canyon (Block 

272) Core P-51 Sample 0-1 cm, side view; fig. 6, living and encysted specimen from Alaminos 

Canyon Core PC15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Plate 48, fig. 1, Green Canyon tubeworm EC5, digital photomicrograph; fig. 2, same specimen as 

in fig. 1; fig. 3, Green Canyon tubeworm EC3; fig. 4, Green Canyon tubeworm EC4. 

Superfamily Nubeculariacea 

Family Nubeculariidae 

Genus Calcituba

Calcituba polymorpha von Roboz 

Calcituba polymorpha von Roboz, 1884, p. 421, figs. 1-16. 

Calcituba polymorpha von Roboz – Arnold, 1967, p. 280-304, text-fig. 2a-m, pl. 1, figs. a-u, pl. 

2, figs. a-r. 

Morphology: Attached, thin, translucent, porcelaneous shell; early chambers in a milioline coil, 

later chambers extending as irregular (occasionally branching) tubes; aperture terminal. 

Remarks: This is the type species of the genus Calcituba. It has been previously reported only 

from a depth of about 1 m in the Gulf of Mexico, but the individuals in our samples are attached 

to a tubeworm from a depth of 562 m. We ascribe this discrepancy to the poor record of the 

species in sediments due to the "extreme fragility" of its shell (Arnold, 1967), and to a dearth of 

observations on attached Foraminifera beyond shelf depths. 

Figures:

Plate 32, fig. 1, Green Canyon tubeworm EC5, digital photomicrograph; fig. 2, same specimens 

as in fig. 1; fig. 3, specimen in fig. 2, surface pits; fig. 4; Green Canyon tubeworm EC5; fig. 5, 

Green Canyon tubeworm EC5, milioline coils. 

44
Superfamily Miliolacea 

Family Spiroloculinidae 

Genus Planispirinoides

Planispirinoides bucculentus var. placentiformis (Brady) 

Miliolina bucculenta Brady var. placentiformis Brady, 1884, p. 171, pl. 4, figs. 1, 2. 

Planispirinoides bucculentus (Brady) var. placentiformis (Brady) – Barker, 1960, p. 8, pl. 4, figs. 

1, 2. 

Planispirinoides bucculentus var. placentiformis (Brady) – Jones, 1994, p. 20, pl. 4, figs. 1-2. 

Morphology: Shell asymmetrical, subglobular; periphery rounded; final whorl with three 

chambers, enclosing all previous whorls; aperture either a crescentic slit near the inner margin of 

the final chamber, or a long, narrow, curved, transverse slit. 

Remarks: Brady (1884) commented that this variety differed from the typical form in its general 

asymmetry, flattened contour, and irregularity of the segments. Our specimens are not as 

flattened as those figured by Brady or as globular as Brady’s Miliolina bucculenta (pl. 114, fig. 

3). 

Figures: 

Plate 126, figs. 1a-c, Green Canyon (Block 272) Core P-54 Sample 0-1 cm, 1a: side view, 1b: 

apertural view, 1c: side view; figs. 2a-c, Green Canyon (Block 232) Core P-EP12 Sample 0-1 

cm, 2a: side view, 2b: apertural view, 2c: side view. 

Scale bars = 50 μm. 

Family Hauerinidae 

Genus Quinqueloculina 

Quinqueloculina bosciana d’Orbigny

Quinqueloculina bosciana d’Orbigny, 1839b, p. 191, pl. 11, figs. 22-24. 

Quinqueloculina bosciana d’Orbigny – Denne, 1990, pl. 11, fig. 14. 

Morphology: Shell elongate; basal end broadly rounded; aperture an opening located between the 

outer margin of the final chamber and the penultimate chamber with a bifid tooth. 

Figures: 

Plate 149, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

side view, 1c: apertural view. 

Scale bars = 50 μm. 

45
Genus Cribromiliolinella 

Cribromiliolinella subvalvularis (Parr)

Miliolinella valvularis (Reuss) – Brady, 1884, p. 161, pl. 4, figs. 4, 5. 

Triloculina subvalvularis Parr, 1950, p. 296, pl. 7, fig. 4. 

Cribromiliolinella subvalvularis (Parr) – Loeblich and Tappan, 1987, p. 337, pl. 348, figs. 16-18. 

Morphology: Shell ovate in outline; cryptoquinqueloculine, chambers one-half coil in length, 

with 3 chambers visible; aperture terminal, with a projecting flap, the opening itself irregularly 

triradiate, occasionally with additional slits. 

Remarks: This is the type species of the genus Cribromiliolinella of Saidova (1981). 

Figures: 

Plate 49, figs. 1a-b, Green Canyon (Block 272) Core P-51 Sample 1-2 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 272) Core P-51 Sample 1-2 cm, side view; figs. 3ac, Green Canyon (Block 272) Core P-51 Sample 0-1 cm, 3a: side view, 3b: side view, 3c: 

apertural view. 

Scale bars = 50 μm. 

Genus Miliolinella 

Miliolinella warreni Andersen

?Miliola oblonga (Montagu) – Flint, 1899, pl. 43, fig. 3. 

Miliolinella warreni Andersen, 1961, p. 37-38, pl. 7, figs. 4a-c. 

Morphology: Shell elongate, sub-circular in cross-section; three chambers visible; basal end 

broadly rounded, apertural end truncated; aperture a crescent-shaped slit located between the 

outer margin of the final chamber and a large, excavated tooth. 

Figures: 

Plate 105, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

side view, 1c: apertural view. 

Scale bars = 50 μm. 

Miliolinella sp. 

Miliolinella sp.

Remarks: This is a rare Miliolinella that we found attached to a tubeworm. The observable 

features do not permit species-level identification. 

46
Figures: 

Plate 106, fig. 1, Mississippi Canyon tubeworm MC1, digital photomicrograph.

Genus Pyrgo 

Pyrgo lucernula (Schwager)

Biloculina lucernula Schwager, 1866, p. 202, pl. 4, figs. 17a, b (not figs. 14a-c). 

Biloculina lucernula Schwager – Brady, 1884, p. 142, pl. 2, figs. 5, 6. 

Pyrgo lucernula (Schwager) – Barker, 1960, p. 4, pl. 2, figs. 5, 6. 

Pyrgo lucernula (Schwager) – Srinivasan and Sharma, 1980, p. 21, pl. 3, figs. 10, 11 (neotype). 

Pyrgo lucernula (Schwager) – Jones, 1994, p. 18, pl. 2, figs. 5, 6. 

Morphology: Shell oval in outline, periphery rounded; only two chambers visible; aperture 

terminal on the end of a short neck, rounded with a bifid tooth. 

Remarks: Srinivasan and Sharma (1980) made the distinction between Triloculina lucernula and 

Pyrgo lucernula, which were both named as Biloculina lucernula by Schwager. Triloculina 

lucernula resembles Pyrgo lucernula in the front view but the apertural view reveals a third 

chamber. 

Figures: 

Plate 145, fig. 1, De Soto Canyon control sample DGOMB2 S36, front view; fig. 2, Farnella 

Canyon Core P-61 Sample 0-1 cm, side view; fig. 3, Green Canyon (Block 272) Core P-33 

Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Pyrgo murrhina (Schwager) 

Biloculina murrhina Schwager, 1866, p. 203, pl. 4, fig. 15a-c. 

Biloculina depressa d’Orbigny var. murrhyna Schwager - Brady, 1884, p. 146, pl. 2, figs. 10, 11. 

Biloculina depressa d’Orbigny – Brady, 1884, p. 146, pl. 2, figs. 15. 

Pyrgo murrhina (Schwager) – Cushman, 1929, p. 71-72, pl. 19, figs. 6, 7. 

Pyrgo murrhina (Schwager) – Phleger and Parker, 1951, p. 7, pl. 3, fig. 11. 

Pyrgo murrhina (Schwager) – Parker, 1954, p. 501, pl. 5, fig. 7. 

Pyrgo murrhyna (Schwager) – Barker, 1960, p. 4, pl. 2, figs. 10, 11, 15. 

Pyrgo murrhina (Schwager) – Srinivasan and Sharma, 1980, p. 22, pl. 3, figs. 6, 7 (neotype). 

Pyrgo murrhina (Schwager) – Denne, 1990, pl. 11, fig. 8. 

Pyrgo murrhina (Schwager) – Jones, 1994, p. 18-19, pl. 2, figs. 10, 11, 15. 

Morphology: Shell circular to oval in outline, periphery extended and carinate, the carina 

interrupted by a sinus, frequently with two small spines opposite the aperture; aperture with an 

exserted tubular neck and a bifid tooth. 

47
Remarks: This common deep-sea species exhibits a lot of variability. 

Figures: 

Plate 146, fig. 1, Alaminos Canyon Boxcore B6 bulk sample, front view; fig. 2, De Soto Canyon 

control sample DGOMB2 S36, front view. 

Scale bars = 50 μm. 

Pyrgo nasuta Cushman 

Pyrgo nasutus Cushman, 1935, p. 7, pl. 3, figs. 1-4. 

Pyrgo cf. nasutus Cushman – Phleger and Parker, 1951, p. 7, pl. 3, figs. 12-14. 

Pyrgo cf. nasutus Cushman – Parker, 1954, pl. 5, fig. 4. 

Pyrgo nasuta Cushman – Andersen, 1961, p. 38, pl. 9, figs. 2a-c. 

Pyrgo phlegeri Andersen, 1961, p. 38-39, pl. 8, figs. 1a-c, 2a-c. 

Morphology: Shell oval in outline, with a carinate periphery that may be serrated; aperture on a 

slight, compressed neck, convex on the outer dorsal side and concave on the inner side; aperture 

an elongate curved slit without a tooth. 

Remarks: Phleger and Parker (1951) noted that their Gulf of Mexico specimens lack the bend of 

the test at the base of the aperture that causes the aperture to tip forward as seen in Cushman’s 

fig. 1b. Parker (1954) observed Gulf of Mexico specimens with a less serrate periphery. 

Andersen (1961) distinguished a new spcies, P. phlegeri, from P. nasuta on the basis of its nonserrate and more circular periphery and greater breadth. These gradational characters are 

unreliable for this ditinction, and we follow Culver and Buzas (1981) in considering P. phlegeri

a synonym of P. nasuta.

Figures: 

Plate 147, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, a: front view, b: 

apertural view. 

Scale bars = 50 μm. 

?Pyrgo sp. 

Morphology: Shell circular in outline, periphery rounded; only two chambers visible; shell 

surface agglutinated; aperture terminal on the end of a very slight neck, rounded with a bifid 

tooth. 

Remarks: This extremely rare species has the typical morphology of Pyrgo, but with a definitely 

agglutinated surface. We have not been able to match it with any known Pyrgo. The possibilties 

are: (a) it is a true Pyrgo with an agglutinated outer wall, and thus a new species; (b) it is not a 

hauerinid but a rzehakinid (i.e., an agglutinated form), and thus a new genus and species. So far, 

we have been able to study a single specimen, and the dilemma remains unsolved. 

48
Figures: 

Plate 148, figs. 1a-b, Alaminos Canyon Core PC4 Sample 1-2 cm, 1a: front view, 1b: close up of 

aperture. 

Scale bars = 50 μm. 

Genus Triloculina

Triloculina tricarinata d’Orbigny 

Triloculina tricarinata d’Orbigny, 1826, p. 299. 

Triloculina tricarinata d’Orbigny – Parker et al., 1865, pl. 1, fig. 8. 

Triloculina tricarinata d’Orbigny – Cushman, 1929, p. 56-57, pl. 13, figs. 3a-c. 

Morphology: Shell elongate, triangular in apertural view; periphery sharply angled; three 

chambers visible, triloculine arrangement; sutures depressed; aperture usually on the end of a 

cylindrical neck, with a simple or bifid tooth. 

Remarks: The aperture in our specimens lacks a neck and has a bifid tooth. 

Figures: 

Plate 182, figs. 1a-b, Canyon (Block 272) Core P-57 Sample 0-1 cm, 1a: side view, 1b: apertural 

view; fig. 2, Alaminos Canyon Core PC5 Sample 4-5 cm, 2, side view. 

Scale bars = 50 μm. 

Genus Sigmoilinita 

Sigmoilinita elliptica (Galloway and Wissler)

Sigmoilina elliptica Galloway and Wissler, 1927, p. 39, pl. 7, figs. 2a, b. 

Sigmoilinita elliptica (Galloway and Wissler) – Denne, 1990, pl. 12, fig. 9. 

Morphology: Shell elliptical in side view; five chambers visible; chambers long, tubular, 

diameter unchanging from end to end; aperture terminal at end of a neck, no tooth. 

Remarks: Sigmoilina tenuis (Czejak) appears from the type figures to be flat in profile while S. 

elliptica (Galloway and Wissler) is not. Some confusion exists about the generic name. Jones 

(1994) regards Sigmoilinita Seiglie 1965 as a junior synonym of Spirosigmoilina Parr 1942 and 

refers Sigmoilinita tenuis (Czjzek), the type species, to Spirosigmoilina tenuis. Both genera have 

initial chambers added in a sigmoiline series, and the later whorls are planispiral. Sigmoilinita

Seiglie may or may not have a tooth while Spirosigmoilina Parr has a tooth according to 

Loeblich and Tappan (1987). Parr’s generic description does not mention the aperture or the 

presence of a tooth. 

49
Figures: 

Plate 163, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

side view, 1c: apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 

cm, 2a: side view, 2b: side view. 

Scale bars = 50 μm. 

Genus Sigmoilopsis 

Sigmoilopsis schlumbergeri (Silvestri) 

Planispirina celata (Costa) – Brady, 1884, p. 197, pl. 8, figs. 1-4. 

Sigmoilina schlumbergeri Silvestri, 1904, p. 267. 

Sigmoilina schlumbergeri Silvestri – Cushman, 1929, p. 49, pl. 11, figs. 1-3. 

Sigmoilina schlumbergeri Silvestri – Phleger and Parker, 1951, p. 8, pl. 4, fig. 6. 

Sigmoilina schlumbergeri Silvestri – Parker, 1954, p. 499, pl. 4, fig. 18. 

Sigmoilopsis schlumbergeri (Silvestri) – Barker, 1960, p. 16, pl. 8, figs. 1-4. 

Sigmoilopsis schlumbergeri (Silvestri) – Andersen, 1961, p. 34, pl. 7, figs. 7a, b. 

Sigmoilopsis schlumbergeri (Silvestri) – Denne, 1990, pl. 12, fig. 10. 

Sigmoilopsis schlumbergeri (Silvestri) – Jones, 1994, p. 23-24, pl. 8, figs. 1-4. 

Morphology: Shell large, longer than wide; chambers arranged in a sigmoid manner with several 

visible from either side; aperture at the end of a short neck with a tooth; shell wall a calcareous 

base encrusted with sand grains. 

Remarks: This is the type species of the genus Sigmoilopsis Finlay 1947. 

Figures: 

Plate 164, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

side view; fig. 2, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

2.2.6. Order Spirillinida 

Family Spirillinidae 

Genus Spirillina 

Spirillina vivipara Ehrenberg 

Spirillina vivipara Ehrenberg, 1843, p. 323, 422, pl. 3, fig. 41. 

Spirillina vivipara Ehrenberg - Brady, 1884, p. 630, pl. 85, figs. 1-4. 

Spirillina vivipara Ehrenberg - Cushman, 1931, p. 3, pl. 1, figs. 1-4. 

Spirillina vivipara Ehrenberg - Jones, 1994, p. 92, pl. 85, figs. 1-4. 

50
Morphology: Hyaline shell with a round proloculus and a planispirally coiled, tubular, second 

chamber, tube width showing gentle increase from juvenile to mature stages; scattered large 

pores; terminal aperture. 

Remarks: This is the type species of the genus Spirillina. It has been widely reported from Gulf 

of Mexico sediments (0-1481 m water depth). In our samples, however, it is found only attached 

to tubeworms. 

Figures: 

Plate 168, digital photomicrographs, fig. 1, Mississippi Canyon tubeworm MC3; figs. 2-4, 

Mississippi Canyon tubeworm EC1. 

Plate 169, fig. 1, same specimens as in fig. 3 of plate168; figs. 2-3, Mississippi Canyon 

tubeworm EC1; fig. 4, Mississippi Canyon tubeworm EC1, attachment side; fig. 5, Mississippi 

Canyon tubeworm MC2. 

Family Patellinidae 

Genus Patellina 

Patellina corrugata Williamson

Patellina corrugata Williamson, 1858, p. 46, pl. 3, 86-89. 

Patellina corrugata Williamson – Brady, 1884, p. 634, pl. 86, figs. 1-7. 

Patellina corrugata Williamson – Cushman, 1931, p. 11, 12, pl. 2, figs. 6, 7. 

Patellina corrugata Williamson – Phleger and Parker, 1951, p. 23, pl. 12, figs. 4a, b. 

Patellina corrugata Williamson – Barker, 1960, p. 178, pl. 86, figs. 1-7. 

Patellina corrugata Williamson – Andersen, 1961, p. 100, pl. 21, figs. 2a, b. 

Patellina corrugata Williamson – Jones, 1994, p. 93, pl. 86, figs. 1-7. 

Morphology: Shell planoconvex to concavo-convex; early chambers spirally arranged, later 

chambers overlapping with a thickened keel; aperture elongate at the inner border of the 

chamber. 

Remarks: This is the type species of the genus Patellina. 

Figures: 

Plate 124, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, dorsal view; figs. 2a-b, Alaminos 

Canyon Core PC5 Sample 4-5 cm, 2a: dorsal view, 2b: ventral view; fig. 3, Alaminos Canyon 

Core PC5 Sample 4-5 cm, ventral view. 

Scale bars = 50 μm. 

Plate 125, fig. 1, Mississippi Canyon tubeworm EC1, digital photomicrograph; fig. 2, Green 

Canyon tubeworm EC2; fig. 3, Green Canyon tubeworm EC4; fig. 4, Green Canyon tubeworm 

EC3, specimen attached to Laminononion tumidum. 

51
2.2.7. Order Lagenida 

Superfamily Nodosariacea 

Family Nodosariidae 

Genus Dentalina 

Dentalina sp. 

Morphology: Shell elongate, uniserial, circular in cross-section; chambers gradually increasing in 

size, sutures slightly depressed; several initial spines; aperture terminal, radiate. 

Figures: 

Plate 52, figs. 1-2, Garden Banks (Block 427) Core P-EP8 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Pseudoglandulina 

Pseudoglandulina comatula (Cushman)

Nodosaria comata (Batsch) – Brady, 1884, p. 509, pl. 64, figs. 1-5. 

Nodosaria comatula Cushman, 1923, p. 83, pl. 14, fig. 5. 

Pseudoglandulina comatula (Cushman) – Phleger and Parker, 1951, p. 10, pl. 5, figs. 7-9. 

Pseudoglandulina comatula (Cushman) – Parker, 1954, p. 505, pl. 5, fig. 22. 

Rectoglandulina comatula (Cushman) – Barker, 1960, p. 134, pl. 64, figs. 1-5. 

Pseudonodosaria comatula (Cushman) – Andersen, 1961, p. 71, pl. 17, figs. 9a, b. 

Pseudonodosaria comatula (Cushman) – Denne, 1990, pl. 11, fig. 2 

Pseudoglandulina comatula (Cushman) – Jones, 1994, p. 76, pl. 64, figs. 1-5. 

Morphology: Shell elongate and stout, initial end may be rounded or tapered, sometimes with a 

short spine; sutures distinct, slightly depressed; surface ornamented by closely-spaced, 

continuous longitudinal costae that obscure previous sutures; upper part of the final chamber 

smooth; aperture central, terminal, radiate. 

Figures: 

Plate 141, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Plectofrondicularia 

Plectofrondicularia advena (Cushman)

Frondicularia inaequalis (Costa) – Brady, 1884, p. 521, pl. 66, figs. 8-12. 

Frondicularia advena Cushman, 1923, p. 141, pl. 20, figs. 1-2. 

52
Parafrondicularia advena (Cushman) – Barker, 1960, p. 138, pl. 66, figs. 8-12. 

Plectofrondicularia advena (Cushman) – Jones, 1994, p. 78, pl. 66, figs. 8-12. 

Morphology: Shell compressed with a subspherical proloculus followed by v-shaped chambers, 

chambers broadest in the center and then tapering to the pointed outer ends; slight peripheral keel 

present; sutures distinct and depressed; aperture circular in the center of the final chamber. 

Remarks: This is the type species of the genus Proxifrons Vella 1963, regarded by Jones (1994) 

as a junior synonym of Plectofrondicularia Liebus 1902. Loeblich and Tappan (1987) list both 

genera. 

Figures: 

Plate 131, fig. 1, Mississippi Canyon Core P-69 Sample 0-1 cm, side view; fig. 2, Garden Banks 

(Block 427) Core P-EP8 Sample 1-2 cm, side view; fig. 3, Mississippi Canyon Core P-IM2 

Sample 1-2 cm, side view. 

Scale bars = 100 μm in fig. 1; 50 μm in figs. 2 & 3. 

Family Vaginulinidae 

Genus Lenticulina 

Lenticulina calcar (Linnaeus)

Nautilus calcar Linneaus, 1767, p. 1162, no. 272. 

Cristellaria calcar (Linnaeus) – Brady, 1884, p. 55, pl. 70, figs. 9-12. 

Lenticulina calcar (Linneaus) – Barker, 1960, p. 146, pl. 70, figs. 9-12. 

Robulus calcar (Linneaus) – Andersen, 1961, p. 48-49, pl. 11, figs. 1-2. 

Lenticulina calcar (Linneaus) – Denne, 1990, pl. 8, fig. 2. 

Lenticulina calcar (Linneaus) – Jones, 1994, p. 81-82, pl. 70, figs. 9-12. 

Morphology: Shell biconvex; peripheral keel present; 4 to 6 chambers in the last whorl; spine 

projecting from each chamber except the final chamber; sutures slightly curved; apertural face 

depressed and flat; aperture produced at the angle of the final chamber, radiate with a median slit 

flanked by two plates directed downward, their inner margins bearing numerous spiny 

projections. 

Figures: 

Plate 97, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

edge view showing apertural face; figs. 2-3, Green Canyon (Block 180) Core P-EP15 Sample 1-2 

cm, a: side view, b: edge view showing apertural face. 

Scale bars = 50 μm. 

53
Lenticulina gibba (d’Orbigny) 

Cristellaria gibba d’Orbigny, 1826, p. 292, no. 17. 

Cristellaria gibba d’Orbigny – Brady, 1884, p. 546, pl. 69, figs. 8, 9. 

Cristellaria gibba d’Orbigny – Cushman, 1923, p. 105-106, pl. 25, fig. 4. 

Lenticulina gibba d’Orbigny – Barker, 1960, p. 144, pl. 69, figs. 8-9. 

Lenticulina gibba d’Orbigny – Denne, 1990, pl. 8, fig. 6. 

Lenticulina gibba d’Orbigny – Jones, 1994, p. 81, pl. 69, figs. 8-9. 

Morphology: Shell biconvex, elongate in side view; periphery acute; sutures flush with the 

surface; apertural face triangular, aperture radiate at the periphery. 

Figures: 

Plate 98, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 1a: side view, 1b: 

edge view showing apertural face, 1c: apertural view; figs. 2a-b, Alaminos Canyon Core PC4 

Sample 0-1 cm, 2a: side view, 2b: edge view showing apertural face. 

Scale bars = 50 μm. 

Lenticulina pliocaena Silvestri 

Cristellaria sp. Brady, 1884, pl. 69, fig. 5. 

Robulus pliocaenicus (Silvestri) – Barker, 1960, p. 144, pl. 69, fig. 5. 

Lenticulina pliocaena (Silvestri) – Jones, 1994, p. 80, pl. 69, fig. 5. 

Morphology: Two chambers, first chamber globular, second chamber triangular in cross-section; 

two prominent longitudinal ribs running from basal spine to apertural end, sometimes a third rib 

present; sutures depressed; aperture terminal with fissures radiating from a central point. 

Remarks: Barker's (1960) identification of Robulus pliocaenicus is based on Thalmann (1932, p. 

252). 

Figures: 

Plate 99, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 1a: side view, 1b: 

edge view showing apertural face, 1c: apertural view. 

Scale bars = 50 μm. 

Lenticulina thalmanni (Hessland)

Cristellaria rotulata (Lamarck) – Brady, 1884, pl. 69, fig. 13. 

Cristellaria rotulata (Lamarck)? – Cushman, 1923, p. 108, pl. 22, fig. 2; pl. 28, figs. 1, 2. 

Robulus thalmanni Hessland, 1943, p. 265, pl. 2, fig. 16 (cited Brady, pl. 69, fig. 13 as holotype). 

Robulus thalmanni Hessland – Barker, 1960, p. 144, pl. 69, fig. 13. 

Lenticulina thalmanii (Hessland) – Jones, 1994, p. 81, pl. 69, fig. 13. 

54
Morphology: Shell biconvex; periphery keeled; less than 8 chambers in the final whorl; sutures 

distinct, not extending into the center of the shell; large umbonate center; apertural face small, 

aperture is radiate with a median slit sided by two plates. 

Remarks: Hessland cites Brady’s pl. 69, fig. 13 as the holotype illustration. Cushman (1923) 

comments that specimens similar to Brady’s fig. 13 are found in the Gulf of Mexico and differ 

from those he described in having many more chambers and curved sutures. Our specimens 

have up to 8 chambers in the last whorl, and the sutures may be straight or slightly curved. 

Figures: 

Plate 100, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, a: side view, b: 

edge view showing apertural face; fig. 3, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, 

side view; fig. 4, Alaminos Canyon Core PC4 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Vaginulinopsis 

Vaginulinopsis subaculeata (Cushman) 

Cristellaria aculeata Brady (not d’Orbigny), 1884, p. 555, pl. 71, figs. 4, 5. 

Cristellaria subaculeata Cushman, 1923, p. 124, pl. 32, fig. 4; pl. 33, fig. 3; pl. 34, figs. 2, 3. 

Cristellaria subaculeata Cushman var. glabrata Cushman, 1923, p. 124, pl. 32, fig. 4; pl. 33, fig. 

3; pl. 34, figs. 2, 3. 

Marginulina subacuealata (Cushman) var. glabrata (Cushman) – Phleger and Parker, 1951, p. 9, 

pl. 5, fig. 4. 

Percultazonaria subaculeta (Cushman) – Loeblich and Tappan, 1987, p. 406, pl. 448, figs. 1-3. 

Marginulopsis glabrata (Cushman) – Denne, 1990, pl. 8, fig. 12. 

Vaginulinopsis subaculeata (Cushman) – Jones, 1994, p. 82, pl. 71, figs. 4, 5. 

Morphology: Shell elongate; early portion coiled planispirally, later uncoiled and rectilinear; 

periphery subacute to carinate; chambers broad and low; sutures marked by lines of raised beadlike nodes; shell wall between the sutures is smooth; apertural face truncate and smooth; aperture 

projecting and radiate, located at the peripheral angle of the final chamber. 

Remarks: Cushman (1923) recognized a variety, named by him as Cristellaria subaculeata var. 

glabrata, on the basis a larger proportion of the uncoiled part, a greater development of spines on 

the periphery and the lack of secondary ornamentation between the sutural lines. As revealed by 

illustrations of earlier workers, these features vary widely within populations of Vaginulinopsis 

subaculeata in the Gulf of Mexico. 

There is some confusion over the generic name of this species. Loeblich and Tappan (1987) 

named Cristellaria subaculeata Cushman 1923 as the type species of a new genus 

Percultazonaria Loeblich and Tappan 1986. They state that it differs from Vaginulinopsis

Silvestri 1904 in the more prominent early coil and in the elevated, costate, and nodose sutural 

55
ornamentation. Ornamentation is not a dependable clue to generic separation in vaginulinids, 

and we follow Jones (1994) in considering Percultazonaria as a junior synonym of 

Vaginulinopsis. 

Figures: 

Plate 188, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: 

edge view showing apertural face, c: apertural view; figs. 3a-b, Green Canyon (Block 180) Core 

P-EP15 Sample 0-1 cm, 3a: side view, 3b: edge view showing apertural face; fig. 4, Green 

Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Saracenaria 

Saracenaria altifrons (Parr) 

Cristellaria acutauricularis Fichtel and Moll – Brady, 1884, p. 543, pl. 114, fig. 17. 

Lenticulina (Robulus) altifrons Parr, 1950, p. 323, pl. 11, figs. 12a, b. 

Lenticulina altifrons Parr – Barker, 1960, p. 236, pl. 114, fig. 17. 

Saracenaria (?) sp. Andersen 1961, p. 59, pl. 14, figs. 2a-c.

Saracenaria altifrons (Parr) – Jones, 1994, p. 113, pl. 114, fig. 17. 

Morphology: Shell is coiled, 7 chambers in the final whorl, chambers increasing in size; 

triangular in cross-section; sutures depressed, almost straight; apertural face is long, broad and 

inflated, aperture terminal, radiate with a median slit. 

Figures: 

Plate 161, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

edge view showing apertural face, 1c: apertural view. 

Scale bars = 50 μm. 

Genus Amphicoryna 

Amphicoryna hirsuta (d’Orbigny)

Nodosaria hirsuta d’Orbigny, 1826, p. 252, no. 7. 

Nodosaria papillosa Silvestri? – Brady, 1884, p. 508, pl. 63, figs. 10, 11. 

Nodosaria hispida (d’Orbigny) – Brady, 1884, p. 508, pl. 63, figs. 12-16. 

Nodosaria hispida (d’Orbigny) – Parker, 1954, p. 505, pl. 6, fig. 1. 

Amphicoryna hirsuta (d’Orbigny) – Barker, 1960, p. 132, pl. 63, figs. 10-16. 

Amphicoryna hispida (d’Orbigny) – Denne, 1990, pl. 1, fig. 7. 

Amphicoryna hirsuta (d’Orbigny) – Jones, 1994, p. 74, pl. 63, figs. 12-15. 

56
Morphology: Shell elongate; early chambers close-set, later chambers connected by the 

cylindrical neck of the preceding chambers; ornamented with small spines, some delicate costae 

present on earlier chambers; aperture terminal at the end of a costate and hispid neck. 

Figures: 

Plate 8, fig. 1, Green Canyon (Block 272) Core P-55 Sample 1-2 cm, side view; fig. 2, Green 

Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view; fig. 3, Garden Banks (Block 427) 

Core P-EP8 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Family Lagenidae 

Genus Lagena 

Lagena chrysalis Heron-Allen and Earland

Lagena chrysalis Heron-Allen and Earland, 1913, p. 74, pl. 6, fig. 4.

Lagena chrysalis Heron-Allen and Earland – Cushman, 1923, p. 10, pl. 1, fig. 14. 

Morphology: Shell chrysalidiform; tapering at both ends but more so at the initial end; final 

chamber rounded, initial end pointed; aperture simple. 

Figures: 

Plate 88, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bar = 50 μm. 

Lagena hispida Reuss

Lagena hispida Reuss, 1863, p. 335, pl. 6, figs. 77-79. 

Lagena hispida Reuss – Brady, 1884, p. 459, pl. 57, figs. 1, 2 (not pl. 57 figs. 3, 4; pl. 59, figs. 2, 

5). 

Lagena hispida Reuss – Cushman, 1923, p. 26-27, pl. 4, figs. 7, 8. 

Lagena hispida Reuss – Barker, 1960, p. 116, pl. 57, figs. 1, 2. 

Lagena hispida Reuss – Jones, 1994, p. 63, pl. 57, figs. 1, 2. 

Morphology: Shell globular or pyriform in shape; spines distributed uniformly over the shell 

wall; aperture terminal on a thin, elongate neck with a lip. 

Remarks: Jones (1994) has limited Lagena hispida to the more elongate forms illustrated as figs. 

1 and 2 on Brady’s plate 57. 

Figures: 

Plate 89, fig. 1, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bar = 50 μm. 

57
Lagena hispidula Cushman 

Lagena laevis Brady (part) (not Lagena laevis (Montagu)), 1884, pl. 56, figs. 10, 11. 

Lagena hispidula Cushman, 1913, p. 14, pl. 5, figs. 2, 3. 

Lagena hispidula Cushman – Barker, 1960, p. 114, pl. 56, figs. 10, 11, ?13. 

Pygmaeoseistron hispidula (Cushman) – Loeblich and Tappan, 1987, p. 416, pl.455, figs. 3-5. 

Lagena hispidula Cushman – Jones, 1994, p. 62, pl. 56, figs. 10, 11, ?13. 

Morphology: Shell cylindrical in shape; broadly rounded at both ends; shell wall is very finely 

hispid; aperture is terminal on a thin, elongate neck. 

Remarks: This is the type species of the genus Pygmaeoseistron Patterson and Richardson in 

Loeblich and Tappan (1987) but we follow Jones (1994) in considering Pygmaeoseistron as a 

junior synonym of Lagena Walker and Jacob (in Kanmacher, 1798). 

Figures: 

Plate 90, fig. 1, Alaminos Canyon Core PC4 Sample 0-1 cm, side view; fig. 2, Alaminos Canyon 

Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Genus Procerolagena 

Procerolagena gracilis (Williamson)

Lagena gracilis Williamson, 1848, p. 13, pl. 1, fig. 5. 

Lagena gracilis Williamson – Brady, 1884, p. 464, pl. 58, fig. 9 (part). 

Lagena distoma Parker and Jones MS., Brady – Brady, 1884, pl. 58, figs. 11-15. 

Lagena distoma Parker and Jones MS., Brady – Cushman, 1923, p. 14, 15, pl. 3, figs. 2, 3. 

Lagena gracilis Williamson – Cushman, 1923, p. 22, figs. 3, 4. 

Lagena gracillima (Seguenza) var. mollis Cushman, 1944, p. 21, pl. 3, fig. 3. 

Lagena gracilis Williamson – Barker, 1960, p. 119, pl. 58, fig. 9 (part). 

Lagena distoma Parker and Jones MS., Brady – Barker, 1960, pl. 58, figs. 11-15. 

Procerolagena gracilis Williamson – Jones, 1994, p. 65, pl. 58, figs. 9, ?11-15. 

Morphology: Shell elongate, the main body of the shell varying from long and thin to fusiform, 

tapering at the initial end into an apical spine and at the apertural end into a long thin neck; 

surface ornamented by slightly raised longitudinal costae varying in number. 

Remarks: Jones (1994) regards figs. 9 and 11-15 from pl. 58 of the Challenger report as falling 

within the range of variability of L. gracilis Williamson (1848), and also includes L. gracillima

var. mollis Cushman 1944 within that range. The morphology of our specimens fits Jones’ 

concept of P. gracilis. Lagena gracilis Williamson (1848) is the type species of the genus 

Procerolagena Puri 1954, and Jones (1984) designated a lectotype for this species. 

58
Figures: 

Plate 135, fig. 1, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, side view; fig. 2, Green 

Canyon (Block 272) Core P-54 Sample 1-2 cm, side view; fig. 3, Green Canyon (Block 272) 

Core P-51 Sample 0-1 cm, side view; fig. 4, Green Canyon (Block 272) Core P-54 Sample 0-1 

cm, side view; fig. 5, Alaminos Canyon Core PC5 Sample 1-2 cm, side view; fig. 6, Green 

Canyon (Block 272) Core P-54 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Superfamily Polymorphinacea 

Family Polymorphinidae 

Genus Vasiglobulina 

Vasiglobulina reticulata Poag 

Globulina caribaea d’Orbigny, 1839b, p. 135, pl. 2, figs. 7, 8. 

Globulina caribaea d’Orbigny – Parker, 1954, p. 506, pl. 5, fig. 23. 

Vasiglobulina reticulata Poag, 1982, p. 266, figs. 5 G-I, 6 A-I, 7 A-I, 8 A-I, 9 G-I. 

Morphology: Shell ovate, globular; circular in section; sutures indistinct; aperture terminal, 

radiate, may be obscured by a fistulose growth. 

Figures: 

Plate 191, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; fig. 2, Alaminos Canyon Core PC5 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Family Ellipsolagenidae 

Genus Oolina 

Oolina apiopleura (Loeblich and Tappan, 1953) 

Lagena acuticosta Reuss – Brady, 1884, p. 464, pl. 57, figs. 32; pl. 58, fig. 21 (part). 

Lagena acuticosta Reuss – Cushman, 1923, p. 5-6, pl. 1, figs. 1-3. 

Lagena apiopleura Loeblich and Tappan, 1953, p. 59, pl. 10, figs. 14, 15. 

Lagena elegantissima (Bornemann) – Barker, 1960, p. 118, pl. 57, fig. 32. 

Lagena sp. nov.? Barker, 1960, p. 119, pl. 58, fig. 21. 

Oolina apiopleura (Loeblich and Tappan) – Jones, 1994, p. 65, 66, pl. 57, fig. 32; pl. 58, fig. 21. 

Morphology: Single chambered, pear shaped, with several strong ribs; terminal, rounded 

aperture; entosolenian tube difficult to see. 

59
Figures: 

Plate 111, figs. 1a-b, Alaminos Canyon Core PC5 Sample 0-1 cm, 1a: side view, 1b: apertural 

view. 

Scale bars = 50 μm. 

Oolina globosa (Montagu) 

Vermiculum globosum Montagu, 1803, p. 523. 

Lagena globosa (Montagu) – Brady, 1884, p. 452, pl. 56, figs. 1-3. 

Lagena globosa (Montagu) – Cushman, 1923, p. 20, 21, pl. 4, figs. 1, 2. 

Lagena globosa (Montagu) – Barker, 1960, p. 114, pl. 56, figs. 1-3. 

Oolina globosa (Montagu) – Jones, 1994, p. 61, pl. 56, figs. 1-2, ?3. 

Morphology: Shell globular to subspherical; shell wall smooth and unornamented; aperture 

rounded with an entosolenian tube. 

Remarks: This is the type species of the genus Entolagena Silvestri 1900 but Jones (1994) notes 

that this was considered a junior synonym of Oolina first and later as of uncertain status by 

Loeblich and Tappan (1964, 1987). 

Figures: 

Plate 112, figs. 1-2, Green Canyon (Block 272) Core P-51 Sample 0-1 cm, a: side view, b: 

apertural view; figs. 3a-b, Green Canyon (Block 272) Core P-51 Sample 1-2 cm, 3a: side view, 

3b: apertural view. 

Scale bars = 50 μm. 

Oolina ovum (Ehrenberg) 

Miliola ovum Ehrenberg, 1843, p. 166. 

Lagena ovum (Ehrenberg) – Brady, 1884, p. 454, pl. 56, fig. 5. 

Lagena ovum (Ehrenberg) – Cushman, 1923, p. 44, 45, pl. 8, figs. 9, 10. 

Oolina ovum (Ehrenberg) – Barker, 1960, p. 114, pl. 56, fig. 5. 

Oolina ovum (Ehrenberg) – Jones, 1994, p. 61, pl. 56, fig. 5. 

Morphology: Shell elongate, elliptical, circular in cross-section; both ends rounded; aperture 

round with a short entosolenian neck. 

Plate 113, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

60
Oolina squamosa (Montagu) 

Vermiculum suamosum Montagu, 1803, p. 526, pl. 14, fig. 2. 

Entosolenia squamosa (Montagu) var. α, catenulata Jeffreys MS in Williamson, 1848, p. 19, pl. 

2, fig. 20. 

Lagena catenulata Williamson – Cushman, 1923, p. 9, 10, pl. 1, fig. 11. 

Morphology: Shell globular; basal end truncated, apertural end slightly drawn out; shell wall 

ornamented by raised polygonal reticulations arranged in vertical rows; aperture round. 

Figures: 

Plate 114, figs. 1a-b, Green Canyon (Block 272) Core P-51 Sample 1-2 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

Genus Fissurina

Fissurina fissa (Heron-Allen and Earland) 

Lagena marginata (Montagu) var. fissa Heron-Allen and Earland, 1922, p. 157, pl. 5, figs. 24-

25. 

Morphology: Shell ovate; carinate periphery, keel increases in breadth over the basal quadrant, 

deflected just before reaching the aboral extremity and rapidly decreases in breadth and may 

vanish; aperture terminal, slit-like, within a fissure at the test apex and an entosolenian tube. 

Remarks: The keel does not completely disappear in all specimens; in some it just decreases in 

breadth. 

Figures: 

Plate 63, figs. 1-3, Alaminos Canyon Core PC4 Sample 0-1 cm, a: side view, b: apertural view. 

Scale bars = 50 μm. 

Fissurina sp. cf. F. incomposita (Patterson and Pettis) 

Lagena acuta (Reuss) – Brady, 1884, p. 474, pl. 59, fig. 6. 

Lagena marginata (Montagu) var. spinifera Earland, 1934, p. 156, pl. 7, figs. 3-4. 

Fissurina crebra (Matthes) – Barker, 1960, p. 122, pl. 59, fig. 6.

Lagenosolenia incomposita Patterson and Pettis, 1986, p. 74. 

Fissurina incomposita (Patterson and Pettis) – Jones, 1994, p. 67, pl. 59, fig. 6. 

Morphology: Shell ovate; narrow peripheral carina, extending into a basal spine; aperture 

terminal, slit-like, within a fissure at the test apex, entosolenian tube internal. 

61
Remarks: Lagena marginata var. spinifera Earland 1934, the name assigned by Thalmann 

(1937), is a primary junior homonym of L. aspera var. spinifera Chapman 1895. It was renamed 

Lagenosolenia incomposita by Patterson and Pettis (1986), but Jones (1994) regards 

Lagenosolenia as a junior synonym of Fissurina Reuss 1850. Our specimens differ from those 

of Brady (1884) and Earland (1934) by not showing a clear neck and the wide, opaque carina at 

the apertural end. 

Figures: 

Plate 64, figs. 1a-b, Green Canyon (Block 272) Core P-51 Sample 0-1 cm, 1a: side view, 1b: 

apertural view. 

Scale bars = 50 μm. 

Genus Parafissurina 

Parafissurina botelliformis (Brady)

Lagena botelliformis Brady, 1881, p. 60. 

Lagena botelliformis Brady – Brady, 1884, p. 454, pl. 56, fig. 6. 

Lagena botelliformis Brady – Cushman, 1923, p. 8, 9; pl. 1, fig. 10. 

Oolina botelliformis (Brady) – Barker, 1960, p. 114, pl. 56, fig. 6. 

Parafissurina botelliformis (Brady) – Jones, 1994, p. 61, pl. 56, fig. 6. 

Morphology: Shell elongate, arcuate, cylindrical with rounded ends; shell wall smooth and 

unornamented; aperture round. 

Figures: 

Plate 119, figs. 1a-b, Alaminos Canyon Core PC5 Sample 1-2 cm, 1a: side view, 1b: apertural 

view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Parafissurina kerguelenensis (Parr) 

Lagena staphyllearia (Schwager) – Brady, 1884, p. 474, pl. 59, figs. 8-11. 

Fissurina kerguelenensis Parr, 1950, p. 305, pl. 8, fig. 7. 

Fissurina kerguelenensis Parr – Barker, 1960, p. 122, pl. 59, figs. 8-11. 

Fissurina kerguelenensis Parr – Hermelin, 1989, p. 48, pl. 6, figs. 17-18. 

Fissurina staphyllearia Schwager – Jones, 1994, p. 67, 68, pl. 59, figs. 8-11. 

Morphology: Shell ovate; rounded in cross-section; narrow peripheral carina; 3-10 spines at the 

basal part of the test; aperture crescentic, dorsal margin projecting above the opening in a hoodlike extension; entosolenian tube internal. 

62
Remarks: Parr (1950) states that his Fissurina kerguelenensis differs from F. staphyllearia

Schwager as Schwager’s species has a thicker, more inflated test, lacks a keeled periphery, and 

has a narrower and more produced apertural end. Parr (1950) observed one specimen in which 

the sides of the aperture are of different heights and noted that in the absence of other specimens 

this species would have been placed in Parafissurina. 

Figures: 

Plate 120, figs. 1a-b, Alaminos Canyon Core PC4 Sample 0-1 cm, 1a: side view, 1b: apertural 

view. 

Scale bars = 50 μm. 

Parafissurina lateralis (Cushman) 

Lagena apiculata (Reuss) – Brady, 1884, pl. 56, figs. 17-18. 

Lagena lateralis Cushman, 1913, p. 9, pl. 1, fig. 1. 

Parafissurina lateralis (Cushman) – Barker, 1960, p. 116, pl. 56, figs. 17, 18. 

Parafissurina lateralis (Cushman) – Jones, 1994, p. 62, pl. 56, figs. 17, 18. 

Morphology: Shell elongate, somewhat tapering towards apertural end; both ends rounded, basal 

end may be truncated; aperture crescentic, dorsal margin projecting above the opening in a hoodlike extension; entosolenian tube internal. 

Figures: 

Plate 121, figs. 1-2, Alaminos Canyon Core PC5 Sample 1-2 cm, a: side view, b: apertural view; 

figs. 3a-b, Alaminos Canyon Core PC4 Sample 0-1 cm, 3a: side view, 3b: apertural view. 

Scale bars = 50 μm. 

Family Glandulinidae 

Genus Seabrookia 

Seabrookia earlandi Wright 

Seabrookia earlandi Wright, 1891, p. 477, pl. 20, figs. 6-7. 

Seabrookia earlandi Wright – Denne, 1990, pl. 12, fig. 7. 

Morphology: Shell elongate, ovate in outline, somewhat irregular in shape; planoconvex to 

unevenly biconvex in end view; five chambers; peripheral keel; aperture a terminal slit, bordered 

by a thick lip; with entosolenian tube. 

Figures: 

Plate 162, figs. 1a-b, south of Mississippi Canyon control sample DGOMB2 C7, 1a: side view, 

1b: apertural view; figs. 2a-b, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, 2a: side 

view, 2b: apertural view. 

63
2.2.8. Order Buliminida 

Superfamily Bolivinacea 

Family Bolivinidae 

Genus Bolivina 

Bolivina alata (Seguenza)

Vulvulina alata Seguenza, 1862, p. 115, pl. 2, figs. 5, 5a. 

Bolivina beyrichi Reuss var. alata (Seguenza) – Brady, 1884, p. 422, pl. 53, figs. 2-4. 

Bolivina beyrichi Reuss var. alata (Seguenza) – Cushman, 1922, p. 30, 31, pl. 8, fig. 3. 

Bolivina alata (Seguenza) – Phleger and Parker, 1951, p. 12, pl. 6, fig. 11. 

Bolivina alata (Seguenza) – Barker, 1960, p. 108, pl. 53, figs. 2-4. 

Bolivina alata (Seguenza) – Andersen, 1961, p. 92, pl. 20, figs. 9a, b. 

Brizalina alata (Seguenza) – Poag, 1981, p. 43, 44, pl. 23, fig. 2, pl. 24, figs. 2a-c. 

Bolivina alata (Seguenza) – Denne, 1990, pl. 2, fig. 2. 

Brizalina alata (Seguenza) – Jones, 1994, p. 58, pl. 53, figs. 2-4. 

Morphology: Periphery sharp, with an imperforate carina extending into a spinose projection at 

the edge of each chamber; chambers distinct, increasing in size rather abruptly in last few stages; 

strongly curved sutures; terminal aperture with lips and a tooth. 

Remarks: Differs from B. barbata in having a wider carina which extends along the entire 

periphery of the final chamber and forms an apertural lip. 

Figures: 

Plate 14, figs. 1a-c, Mississippi Canyon Core P-IM2 Sample 0-1 cm, 1a: side view, 1b: edge 

view, 1c: apertural view; fig. 2, Mississippi Canyon Core P-IM4 Sample 1-2 cm, side view. 

Carbonate overgrowth visible on the shell; figs. 3a-c, Green Canyon Core P-51 Sample 0-1 cm, 

3a: side view, 3b: edge view, 3c: apertural view; fig. 4, Green Canyon Core P-55 Sample 1-2 cm, 

side view. 

Scale bars = 50 μm. 

Bolivina albatrossi Cushman

Bolivina albatrossi Cushman, 1922, pt. 3, p. 31, pl. 6, fig. 4. 

Bolivina albatrossi Cushman – Phleger and Parker, 1951, p. 12, pl. 6, figs. 15-16. 

Bolivina albatrossi Cushman – Parker, 1954, p. 513, pl. 7, fig. 13. 

Bolivina albatrossi Cushman – Pflum and Frerichs, 1976, pl. 1, figs. 5-6. 

Brizalina albatrossi (Cushman) – Poag, 1981, p. 44, pl. 23, fig. 5; pl. 24, figs. 5a-c. 

Bolivina albatrossi Cushman – Denne, 1990, pl. 2, fig. 3. 

64
Morphology: Initial end broadly rounded; periphery rounded; early portion of shell with 

reticulate surface; last few chambers smooth; terminal aperture with lips and a tooth. 

Figures: 

Plate 15, figs. 1-2, south of Mississippi Canyon control sample DGOMB2 C7, a: side view, b: 

edge view, c: apertural view. 

Scale bars = 50 μm. 

Bolivina barbata Phleger and Parker

Bolivina barbata Phleger and Parker, 1951, p. 13, pl. 6, figs. 12a, b, 13. 

Bolivina barbata Phleger and Parker – Parker, 1954, p. 514, pl. 7, fig. 14. 

Bolivina barbata Phleger and Parker – Andersen, 1961, p. 93, pl. 20, figs. 6a, b. 

Brizalina barbata (Phleger and Parker) – Poag, 1981, p. 44, 45, pl. 23, fig. 1; pl. 24, figs. 1a-c. 

Bolivina barbata Phleger and Parker – Denne, 1990, pl. 2, fig. 4. 

Morphology: Shell tapering gradually; with a narrow imperforate carina extending into a spinose 

projection at the edge of each chamber; chambers distinct, increasing gradually in size as added; 

strongly curved imperforate sutures separating densely perforate chamber walls (see plate 16, fig. 

4). 

Remarks: Differs from B. alata in having much narrower and more numerous chambers, a 

narrower carina, and imperforate sutural bands. Differs from B. minima in having more 

numerous chambers, more strongly curved, imperforate sutures, and a distinct carina. 

Figures: 

Plate 16, figs. 1a-c, Green Canyon (Block 272) Core P-51 Sample 1-2 cm, 1a: side view, 1b: 

edge view, 1c: apertural view. Carbonate overgrowth visible on the shell; fig. 2, Mississippi 

Canyon Core P-IM4 Sample 1-2 cm, side view. Carbonate overgrowth visible on the shell; figs. 

3a-c, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 3a: side view; 3b: edge view, 3c: 

apertural view; fig. 4, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Bolivina daggarius Parker 

Bolivina lanceolata Parker, 1954, p. 514, pl. 7, figs. 17-20. 

Not Bolivina lanceolata di Napoli, 1952. 

Bolivina daggarius Parker, 1955, p. 52. 

Brizalina lanceolata (Parker) – Poag, 1981, p. 45, 46, pl. 25, fig. 5; pl. 26, figs. 5a-5c. 

Morphology: Shell tapering to a pointed initial end, often with a short spine; periphery acute, 

sometimes with a narrow keel, especially on the later chambers; sutures slightly limbate, straight, 

at 45° with the horizontal; coarse perforations in shell wall excluding the inner and upper portion 

65
of earlier chambers, a few fine costae present on earlier chambers; aperture narrow, keeled with a 

serrated tooth.

Remarks: Parker (1954) noted that B. daggarius differed from B. acerosa Cushman in having a 

larger size, clear areas on the inner portions of chambers and costae (when present) extending 

further up the shell. Parker’s specimens have a maximum length of 0.6 mm compared with 0.35 

– 0.45 mm of B. acerosa. Our specimens fit Parker’s description and are of a similar size (0.6 

mm). 

Figures: 

Plate 17, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 1a: side view, 1b: 

edge view, 1c: apertural view. 

Scale bars = 50 μm. 

Bolivina goesii Cushman

Bolivina goësii Cushman, 1922, p. 34, pl. 6, fig. 5. 

Bolivina goësii Cushman – Parker and Phleger, 1951, p. 13, pl. 6, fig. 17. 

Bolivina goësii Cushman – Parker, 1954, p. 514, pl. 7, fig. 16. 

Brizalina goësii (Cushman) – Poag, 1981, p. 45, pl. 23, fig. 4; pl. 24, figs. 4a-c. 

Bolivina goessi Cushman – Denne, 1990, pl. 2, fig. 6. 

Morphology: Shell tapering to blunt initial end; narrow peripheral keel; chambers arranged in a 

chevron pattern; depressed chamber walls flanked by sutural ridges; coarse perforations in the 

shell wall; aperture elongate with a serrated tooth, surrounded by wide lips. 

Figures: 

Plate 18, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

edge view, 1c: apertural view. 

Scale bars = 50 μm. 

Bolivina sp. cf. B. hastata Phleger and Parker 

Bolivina fragilis Phleger and Parker, 1951, p. 13, pl. 6, figs. 14, 23, 24a, b. 

Bolivina hastata Phleger and Parker, 1951, p. 13, pl. 6, figs. 18a, b, 19. 

Bolivina fragilis Phleger and Parker – Parker, 1954, p. 514, pl. 7, fig. 15. 

Bolivina fragilis Phleger and Parker – Andersen, 1961, p. 93, pl. 20, figs. 8a, b. 

Brizalina fragilis (Phleger and Parker) – Poag, 1981, p. 45, pl. 25, fig. 4; pl. 26, figs. 4a-c. 

Bolivina fragilis Phleger and Parker – Denne, 1990, pl. 2, fig. 5.

Bolivina hastata Phleger and Parker – Denne, 1990, pl. 2, fig. 7. 

Morphology: Periphery acute with a thin keel; chambers distinct; sutures narrow, almost straight; 

several low costae extending from initial end, which often has a small spine, about one third up 

the length of the shell. 

66
Remarks: Differs from B. fragilis in its smaller size, narrower keel and less curved chambers. 

More similar to B. hastata although the keel in this species extends fully up the shell and there 

are usually only one costae and the apical end is less ovate. 

Figures: 

Plate 19, figs. 1a-c, Green Canyon (Block 272) Core P-33 Sample 1-2 cm, 1a: side view, 1b: 

edge view, 1c: apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 

cm, 2a: side view, 2b: edge view; fig. 3, Mississippi Canyon Core P-IM4 Sample 1-2 cm, side 

view. Carbonate overgrowth visible on the shell. 

Scale bars = 50 μm. 

Bolivina lowmani Phleger and Parker

Bolivina lowmani Phleger and Parker, 1951, p. 13-14, pl. 6, figs. 20a, b, 21. 

Bolivina lowmani Phleger and Parker – Parker, 1954, p. 515, pl. 7, fig. 21. 

Brizalina lowmani (Phleger and Parker) – Poag, 1981, p. 46, pl. 25, fig. 3; pl. 26, figs. 3a-c. 

Bolivina lowmani Phleger and Parker – Denne, 1990, pl. 2, fig. 8. 

Morphology: Initial end slightly tapering to rounded and rather compressed, sometimes slightly 

twisted, periphery slightly lobulate. 

Remarks: A few scattered medium-sized pores were observed on Phleger and Parker’s holotype 

under the light microscope but were not seen on specimens of this study. 

Figures: 

Plate 20, figs. 1a-c, Alaminos Canyon Core PC5 Sample 1-2 cm, 1a: side view, 1b: edge view, 

1c: apertural view; figs. 2a-b, Alaminos Canyon Core PC5 Sample 1-2 cm, 2a: side view, 2b: 

edge view; fig. 3, Alaminos Canyon Core PC5 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Bolivina minima Phleger and Parker

Bolivina minima Phleger and Parker, 1951, p. 14, pl.7, figs. 22a, 22b, 25. 

Bolivina minima Phleger and Parker – Parker, 1954, p. 515, pl. 7, figs. 22, 23. 

Brizalina minima (Phleger and Parker) – Poag, 1981, p. 46, 47, pl. 23 fig. 1; pl. 24, figs. 1a-1c. 

Bolivina minima Phleger and Parker – Denne, 1990, pl. 2, fig. 9. 

Morphology: Gradually tapering with a subacute to broadly rounded initial end; chambers 

distinct, numerous, frequently with a spinose extension; sutures distinct, slightly curved; between 

one and three rows of perforations along the lower part of chambers; very small (~ 10 μm) sharp 

projections across sutures between chambers. 

67
Remarks: Differs from B. barbata in the absence of a carina. The presence of spines on 

chambers is variable in this species. Phleger and Parker (1951) also noted this feature and 

observed that chamber spines were absent from many specimens. 

Figures: 

Plate 21, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: edge 

view; figs. 3a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 3a: side view, 3b: 

edge view, 3c: apertural view; fig. 4, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 

side view. 

Scale bars = 50 μm. 

Bolivina ordinaria Phleger and Parker

Bolivina simplex Phleger and Parker, 1951, p. 14, pl. 7, figs. 4, 5a, b, 6. 

Not Bolivina interjuncta Cushman var. simplex Cushman and Renz, 1941 = Bolivina simplex

Cushman and Renz of authors. 

Bolivina ordinaria Phleger and Parker, 1952, p. 14. 

Bolivina ordinaria Phleger and Parker – Parker, 1954, p. 515, pl. 7, fig. 24. 

Brizalina ordinaria (Phleger and Parker) – Poag, 1981, p. 47, pl. 23, fig. 6; pl. 24, figs. 6a-c. 

Bolivina ordinaria Phleger and Parker – Denne, 1990, pl. 2, fig. 10. 

Morphology: Shell compressed and tapering; periphery acute at initial end, more rounded 

towards apertural end; sutures slightly depressed; one to two rows of pores clustering along the 

top of sutures, but more extensive in later chambers of larger specimens. 

Remarks: The rows of pores make the sutures appear broad, opaque and white under the light 

microscope. 

Figures: 

Plate 22, figs. 1a-c, south of Mississippi Canyon control sample DGOMB2 C7, 1a: side view, 1b: 

edge view, 1c: apertural view; figs. 2-3, south of Mississippi Canyon control sample DGOMB2 

C7, side view; fig. 4, Green Canyon (Block 232) Core P-EP12 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Bolivina sp. cf. B. pusilla Schwager 

Bolivina pusilla Schwager, 1866, p. 254, pl. 7, fig. 101. 

Bolivina pusilla Schwager – Parker, 1954, p. 516, pl. 7, fig. 31. 

Bolivina pusilla Schwager – Pflum and Frerichs, 1976, pl. 1, fig. 7. 

Bolivina pusilla Schwager – Boltovskoy, 1978, pl. 1, figs. 16-18. 

Brizalina pusilla (Schwager) – Srinivasan and Sharma, 1980, p. 44, pl. 6, fig. 21 (neotype). 

Bolivina pusilla Schwager – Hermelin, 1989, p. 59, 60. 

Bolivina pusilla Schwager – Denne, 1990, pl. 2, fig. 11. 

68
Morphology: Elongate shell; wall ornamented with numerous longitudinal striations which 

sometimes terminate in a spine, last few chambers smooth. 

Remarks: The terminating spine was not observed by Srinivasan and Sharma (1980) on the 

neotype or other Car Nicobar specimens of B. pusilla, and is also not seen in Schwager’s type 

illustration. The specimens illustrated by Parker (1954) and Pflum et al. (1976) do have a spine, 

and the specimens from this study match those of Parker (1954) and Pflum et al. (1976). 

Bolivina sp. cf. B. pusilla differs from Bolivina hastata in that the sutures are obscured by the 

costae, and the chambers appear indistinct. 

Figures: 

Plate 23, figs. 1a-c, Alaminos Canyon Core PC5 Sample 3-4 cm, 1a: side view, 1b: edge view, 

1c: apertural view; fig. 2, Alaminos Canyon Core PC15 Sample A10, side view; fig. 3, Alaminos 

Canyon Core PC5 Sample 1-2 cm, side view; fig. 4, De Soto Canyon control sample DGOMB2 

S36, side view. 

Scale bars = 50 μm. 

Bolivina subaenariensis Cushman 

Bolivina aenariensis Brady, 1882, p. 711. 

Bolivina aenariensis Brady – Brady, 1884, p. 423, pl. 53, figs. 10, 11. 

Bolivina subaenariensis Cushman, 1922, p. 46, 47, pl. 7, fig. 6. 

Bolivina subaenariensis Cushman var. mexicana Cushman, 1922, p. 47, pl. 8, fig. 1. 

Bolivina subaenariensis Cushman var. mexicana Cushman – Phleger and Parker, 1951, p. 15, pl. 

7, figs. 8-10. 

Bolivina subaenariensis mexicana Cushman – Parker, 1954, p. 517, pl. 7, fig. 33. 

Bolivina subaenariensis Cushman var. – Barker, 1960, p. 110, pl. 53, figs. 10, 11. 

Bolivina mexicana Cushman – Andersen, 1961, p. 94, pl. 20, figs. 7a, b. 

Brizalina subaenariensis mexicana (Cushman) – Poag, 1981, p. 48, pl. 25, fig. 1; pl. 26, figs. 1ac. 

Bolivina subaenariensis mexicana Cushman – Denne, 1990, pl. 2, fig. 13. 

Bolivina subaenariensis var. mexicana Cushman – Jones, 1994, p. 58, pl. 53, figs. 10, 11. 

Morphology: Shell compressed; periphery acute, carinate; a few long raised costae run from the 

apex towards the apertural end; a single spine is present at the apical end; sutures distinct and 

curved. 

Remarks: Cushman (1922) stated that B. subaenariensis var. mexicana differs from B. 

subaenariensis in (a) having a greater number of costae that run nearly to the apertural end of the 

test and (b) the more abruptly tapering shell. This difference, however, seems to be gradational, 

and within the natural range of variability for the species (compare plate-figs. 1a and 3) and, 

therefore, the varietal name is rejected. This species differs from B. hastata in its larger size and 

more distinct keel and costae. 

69
Figures: 

Plate 24, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 1-2 cm, 1a: side view, 1b: 

edge view, 1c: apertural view; figs. 2-3, Green Canyon (Block 180) Core P-EP15 Sample 0-1 

cm, side view; figs. 4a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 4a: side view, 

4b: edge view, 4c: apertural view; figs. 5a-b, Mississippi Canyon Core P-IM4 Sample 1-2 cm, 

5a: side view, 5b: edge view. Carbonate overgrowth visible on the shell. 

Scale bars = 50 μm. 

Bolivina translucens Phleger and Parker

Bolivina translucens Phleger and Parker, 1951, p.15, pl. 7, figs. 13, 14a, b. 

Bolivina translucens Phleger and Parker – Parker, 1954, p. 517, fig. 34. 

Brizalina translucens (Phleger and Parker) – Poag, 1981, p. 48, pl. 25, fig. 2; pl. 26, figs. 2a-c. 

Bolivina translucens Phleger and Parker – Denne, 1990, pl. 2, fig. 14. 

Morphology: Shell slightly tapering to a blunt initial end in megalospheric forms and to a 

subacute end in microspheric forms; numerous chambers; sutures slightly oblique, depressed; 

lower half of chambers with numerous fine perforations; aperture loop-shaped with a lip on one 

side and a tooth. 

Figures: 

Plate 25, figs. 1a-c, Mississippi Canyon Core P-69 Sample 1-2 cm, 1a: side view, 1b: edge view, 

1c: apertural view; figs. 2a-b, Mississippi Canyon Core P-69 Sample 0-1 cm, 2a: side view, 2b: 

edge view; fig. 3, Mississippi Canyon Core P-69 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Superfamily Bolivinitacea 

Family Bolivinitiae 

Genus Abditodentrix 

Abditodentrix pseudothalmanni (Boltovskoy and Giussani de Kahn)

Bolivinita pseudothalmanni Boltovskoy and Giussani de Kahn, 1981, p.44. 

Abditodentrix asketocomptella Patterson, 1985, p. 140, pl. 1, figs. 1-9. 

Abditodentrix pseudothalmanni Boltovskoy and Giussani de Kahn – Loeblich and Tappan, 1987, 

p. 503, pl. 554, figs. 1-5. 

Abditodentrix asketocomptella Patterson – Hermelin, 1989, p. 59, pl. 10, fig. 16. 

Abditodentrix pseudothalmanni Boltovskoy and Giussani de Kahn – Denne, 1990, pl. 1 fig. 1. 

Morphology: Truncate margins, raised sutures and reticulate surface ornamentation. On some 

specimens the ridges do not form a complete reticulate pattern; this is normal variation within the 

population. There is also a variation in thickness, with the more heavily reticulate specimens 

70
thicker in size. The aperture is peripheral with an apertural lip on one side and a reduced and 

infolded toothplate. 

Remarks: Abditodentrix differs from Bolivinita in its surface ornamentation. This ornamentation 

along with the truncate margin distinguishes it from Bolivina. Latibolivina displays a similar 

network of reticulations, but it lacks the truncate margin of Abditodentrix. See Hermelin (1989) 

and Patterson (1985) for additional comments. 

Figures: 

Plate 1, figs. 1a-c, south of Mississippi Canyon control sample DGOMB2 C7, 1a: side view, 1b: 

edge view, 1c: top view; fig. 2, Alaminos Canyon Core PC15 Sample 7-8 cm, side view; figs. 3ac, Mississippi Canyon Core P-69 Sample 1-2 cm, 3a: side view, 3b: edge view, 3c: apertural 

view; fig. 4, Alaminos Canyon Core PC15 Sample 8-9 cm, side view. 

Scale bars = 50 μm; fig. 3c scale bar = 20 μm. 

Genus Bolivinita 

Bolivinita quadrilatera (Schwager)

Textilaria quadrilatera Schwager, 1866, p. 253, pl. 7, fig. 103. 

Textularia quadrilatera Schwager – Brady, 1884, p. 358, pl. 42, figs. 8-12.

Bolivina quadrilatera (Schwager) – Cushman, 1922, p. 44, pl. 8, fig. 2. 

Bolivinita quadrilatera (Schwager) – Barker, 1960, p. 86, pl. 42, figs. 8-12 

Bolivinita quadrilatera (Schwager) – Srinivasan and Sharma, 1980, p. 42, 43, pl. 6, figs. 22, 23 

(neotype). 

Bolivinita quadrilatera (Schwager) – Jones, 1994, p. 47, pl. 42, figs. 8-12. 

Morphology: Quadrate in edge view, edges carinate; seven to eight chambers; occassional 

costae; aperture suboval with a tooth. 

Remarks: Cushman (1922) observed a spine at the initial end of the species but this spine was 

not noted by Srinivasan and Sharma (1980) in their re-description of the species. They did note 

that some specimens were more narrowly pointed than the neotype. This is the type species of 

the genus Bolivinita Cushman 1927. 

Figures: 

Plate 26, figs. 1a-c, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: side view, 1b: edge view, 1c: 

apertural view. 

Scale bars = 50 μm. 

71
Superfamily Cassidulinacea 

Family Cassidulinidae 

Genus Cassidulina 

Cassidulina carinata Silvestri 

Cassidulina laevigata d’Orbigny var. carinata Silvestri, 1896, p. 104, pl. 2, figs. 10 a-c. 

Cassidulina laevigata d’Orbigny var. carinata Cushman, 1922, p. 124, pl. 25, figs. 6, 7. 

Cassidulina neocarinata Thalmann, 1950, p. 44 (new name). 

Cassidulina laevigata d’Orbigny – Phleger and Parker, 1951, p. 27, pl. 14, figs. 6a, 6b. 

Cassidulina laevigata d’Orbigny var. carinata Cushman – Phleger and Parker, 1951, p. 27, pl. 

14, figs. 7a, 7b. 

Cassidulina neocarinata Thalmann – Parker, 1954, p. 536, pl. 11, fig. 3. 

Cassidulina neocarinata Thalmann – Andersen, 1961, p. 113, pl. 25, figs. 4a-c. 

Cassidulina neocarinata Thalmann – Denne, 1990, pl. 3, fig. 10. 

Cassidulina laevigata d’Orbigny var. carinata Silvestri – Jones, 1994, p. 60, pl. 54, figs. 2, 3. 

Morphology: Shell compressed; periphery acute with a distinct keel; aperture is a long narrow 

slit parallel to the periphery. 

Remarks: Smaller specimens are very similar to Phleger and Parker’s (1951) figures of 

Cassidulina laevigata, whereas larger specimens match their figures of C. laevigata var. 

carinata, but theese are all conspecific. An ontogenetic series of such growth forms is shown in 

plate 34. C. teretis Tappan 1951 is a similar species, but our specimens differ from it in having 

less ovate chambers and in being much smaller. According to Jones (1994), Cassidulina 

laevigata d’Orbigny var. carinata Cushman 1922 is “both homonymous and synonymous with 

Cassidulina laevigata d’Orbigny var. carinata Silvestri 1896, such that its replacement name 

Cassidulina neocarinata Thalmann 1950 was unnecessary.” We agree with this assessment. 

Figures: 

Plate 33, fig. 1, Mississippi Canyon Core P-69 Sample 0-1 cm, side view showing the aperture; 

figs. 2a-b, Mississippi Canyon Core P-IM8 Sample 1-2 cm, 2a: side view showing the aperture, 

2b: side view; fig. 3, south of Mississippi Canyon control sample DGOMB2 C7, side view 

showing the aperture; figs. 4a-b, Mississippi Canyon Core P-55 Sample 0-1 cm, 4a: side view 

showing the aperture, 4b: side view. 

Plate 34, ontogenetic series of growth forms, figs. 1-9, Mississippi Canyon Core P-IM8 Sample 

1-2 cm, side view showing the aperture; figs. 10-11, Mississippi Canyon Core P-55 Sample 0-1 

cm, side view showing the aperture; fig. 12, Mississippi Canyon Core P-69 Sample 0-1 cm, side 

view showing the aperture. 

Scale bars = 50 μm. 

72
Cassidulina curvata Phleger and Parker 

Cassidulina curvata Phleger and Parker, 1951, p. 26, pl. 14, figs. 5a-b. 

Cassidulina curvata Phleger and Parker – Andersen, 1961, p. 113, pl. 25, figs. 3a-c. 

Islandiella curvata (Phleger and Parker) – Poag, 1981, p. 69, 70, pl. 17, fig. 1; pl. 18 figs. 1a, 1b. 

Cassidulina curvata Phleger and Parker – Parker, 1954, p. 535, pl. 11, fig. 1. 

Cassidulina curvata Phleger and Parker – Denne, 1990, pl. 3, fig. 10. 

Morphology: Shell biconvex; periphery rounded, acute with a keel; curved chambers reaching 

almost to the umbilicus; aperture is somewhat elongate and located close to the periphery. 

Remarks: As Phleger and Parker (1951) observed, this species differs from C. norcrossi in 

possessing curving chambers and coarsely perforate shell, and from C. laevigata in having a 

rounded nonlobulate periphery and a less densely perforate shell. 

Figures: 

Plate 35, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: side view, b: side 

view, c: edge view showing the aperture. 

Scale bars = 50 μm. 

Cassidulina obtusa Williamson 

Cassidulina obtusa Williamson, 1858, p. 69, pl. 6, figs. 143, 144. 

Cassidulina crassa d’Orbigny – Brady, 1884, p. 429, pl. 54, figs. 5. 

Cassidulina crassa d’Orbigny – Cushman, 1922, p. 124, p. 124, 125, pl. 26, fig. 7. 

Cassidulina crassa d’Orbigny – Phleger and Parker, 1951, p. 26, pl. 14, figs. 4a, 4b. 

Cassidulina aff. crassa d’Orbigny – Parker, 1954, p. 535, pl. 10, fig. 31. 

Cassidulina crassa d’Orbigny – Barker, 1960, p. 110, pl. 54, figs. 5. 

Cassidulina crassa d’Orbigny – Denne, 1990, pl. 3, fig. 8. 

Cassidulina obtusa d’Orbigny – Jones, 1994, p. 60, pl. 54, fig. 5. 

Morphology: Shell compressed; lobulate outline; periphery rounded; aperture long narrow slit 

parallel to the periphery. 

Taxonomic remarks: Cassidulina crassa d’Orbigny is similar to C. obtusa Williamson but as 

Williamson (1858) notes, in his description of C. obtusa, C. crassa has an aperture that is not a 

simple arcuate slit but bends back upon itself towards the outer portion of the shell. Haynes 

(1973) commented that C. crassa is a large species, up to 1 mm in diameter, and has an aperture 

almost at right angles to the basal suture. He stated that many North Atlantic references, 

including Brady’s (1884) and Cushman’s (1922), to C. crassa relate to C. obtusa. 

Figures: 

Plate 36, fig. 1, Alaminos Canyon Core PC4 Sample 0-1 cm, side view showing the aperture; fig. 

2, Mississippi Canyon Core P-69 Sample 1-2 cm, side view showing the aperture; fig. 3, south of 

Mississippi Canyon control sample DGOMB2 C7, side view showing the aperture; fig. 4, De 

73
Soto Canyon control sample DGOMB2 S36, side view; fig. 5, Alaminos Canyon Core PC4 

Sample 0-1 cm, side view; fig. 6, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side 

view. 

Scale bars = 50 μm. 

Genus Cassidulinoides 

Cassidulinoides tenuis Phleger and Parker

Cassidulinoides tenuis Phleger and Parker, 1951, p. 27, figs. 14a, 14b, 15-17. 

Cassidulinoides tenuis Phleger and Parker – Parker, 1954, p. 537, pl. 11, fig. 14. 

Cassidulinoides tenuis Phleger and Parker – Boltovskoy, 1978, pl. 3, fig. 2. 

Rutherfordoides tenuis (Phleger and Parker) – Kohl, 1985, p. 89, 90, pl. 18, figs. 5a-f. 

Rutherfordoides tenuis (Phleger and Parker) – Hermelin, 1989, p. 75, pl. 14, figs. 3, 4. 

Rutherfordoides tenuis (Phleger and Parker) – Denne, 1990, pl. 12, figs. 4a, b. 

Morphology: Initial end often bent; chambers biserially enrolled, later uncoiled, somewhat 

inflated; sutures depressed; loop-shaped aperture extending from the base of the final chamber 

almost to the apex of the shell with a toothplate. 

Remarks: The shell wall has elongate pores (1 μm long and only a fraction of a μm wide) noted 

by Hermelin (1989, fig. 4). 

Figures: 

Plate 37, fig. 1, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, side view; fig. 2, Green 

Canyon (Block 272) Core P-51 Sample 1-2 cm, side view; fig. 3, Green Canyon (Block 180) 

Core P-EP15 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Genus Globocassidulina 

Globocassidulina subglobosa (Brady)

Cassidulina subglobosa Brady, 1881, p. 30; 1884, p. 430, pl. 54, figs. 17a-c. 

Cassidulina subglobosa Brady – Cushman, 1922, p. 127, 128, pl. 24, fig. 6. 

Cassidulina subglobosa Brady – Phleger and Parker, 1951, p. 27, figs. 11,12,13. 

Cassidulina subglobosa Brady – Parker, 1954, p. 536, pl. 11, figs. 4-9. 

Cassidulina subglobosa Brady – Barker, 1960, p. 112, pl. 54, figs. 17a-c. 

Cassidulina subglobosa Brady – Andersen, 1961, p. 113, pl. 25, figs. 2a-c. 

Globocassidulina subglobosa (Brady) – Belford, 1966, p. 149, pl. 25, figs. 11-16, text-figs. 17:1-

7, 18:1-4. 

Cassidulina subglobosa subglobosa Brady – Boltovskoy, 1978, p. 155, pl. 2, fig. 34. 

Islandiella subglobosa (Brady) – Poag, 1981, p. 70-71, pl. 17, fig. 3; pl. 18, figs. 3a-c. 

74
Islandiella sp. aff. I. subglobosa (Brady) – Poag, 1981, p. 71, pl. 17, fig. 4; pl. 18, figs. 4a-c. 

Globocassidulina subglobosa (Brady) – Hermelin, 1989, p. 74, 75. 

Cassidulina subglobosa Brady – Denne, 1990, pl. 3, fig. 12. 

Globocassidulina subglobosa (Brady) – Jones, 1994, p. 60, pl. 54, fig. 17. 

Morphology: A globose biconvex species with a less convex ventral side and an oblique looplike aperture on the face of the terminal chamber. The aperture can vary from a loop-like 

aperture to a V-shaped aperture with a toothplate. 

Remarks: Those with V-shaped apertures were placed in Islandiella sp. aff. I. subglobosa by 

Poag (1981), but it has been synonymized with Globocassidulina subglobosa here. Differs from 

Cassidulina obtusa, which has a more compressed shell and a slit-like aperture. 

Figures: 

Plate 68, figs. 1a-c, Green Canyon (Block180) Core P-EP15 Sample 0-1 cm, 1a: side view 

showing the aperture, 1b: side view, 1c: close up of V-shaped aperture; figs. 2-3, south of 

Mississippi Canyon control sample DGOMB2 C7, side view showing the aperture; fig. 4, south 

of Mississippi Canyon control sample DGOMB2 C7, side view. 

Scale bars = 50 μm. 

Superfamily Turrilinacea 

Family Turrilinidae 

Genus Eubuliminella 

Eubuliminella morgani (Andersen)

Buliminella morgani Andersen, 1961, p. 87, pl. 19, fig. 10. 

Buliminella morgani Andersen – Poag, 1981, p. 51, pl. 33, fig. 1; pl. 34, figs. 1a, 1b. 

Buliminella morgani Andersen – Denne, 1990, pl. 3, fig. 4. 

Morphology: Multiserial shell with 3-4 chambers in each whorl; sutures depressed; aperture 

loop-shaped with a small toothplate. 

Remarks: This species, endemic to the Gulf of Mexico, has been reported by various authors 

under Buliminella. There is very little similarity, however, between it and the type species of 

Buliminella, B. elegantissima (d’Orbigny). We place Andersen’s species under Eubuliminella

Revets 1993, because its features fit well the morphology of that genus. Eubuliminella, unlike 

Buliminella, is tetraserial or pentaserial, at least in the early part, and possesses an aperture with 

a toothplate. 

Figures: 

Plate 62, figs. 1a-b, Mississippi Canyon Core P-69 Sample 1-2 cm, 1a: side view, 1b: apertural 

view; fig. 2, Mississippi Canyon Core P-IM2 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

75
Family Stainforthiidae 

Genus Stainforthia 

Stainforthia complanata (Egger)

Virgulina schreibersiana Czjzek – Brady, 1884, p. 414, pl. 52, figs. 1-3. 

Virgulina schreibersiana Czjzek var. complanata Egger, 1893, p. 292, pl. 8, figs. 91, 92. 

Virgulina complanata Egger – Phleger and Parker, 1951, p. 19, pl. 9, figs. 1a, 1b, 2a, 2b, 3a, 3b. 

Virgulina complanata Egger – Parker, 1954, p. 512, pl. 7, fig. 6. 

Virgulina davisi Chapman and Parr – Barker, 1960, p. 106, pl. 52, figs. 1-3. 

Virgulina complanata Egger – Andersen, 1961, p. 91, pl. 20, figs. 2a, b. 

Stainforthia complanata (Egger) – Denne, 1990, pl. 13, fig. 3. 

Fursenkoina complanata (Egger) – Jones, 1994, p. 56, pl. 52, figs. 1-3. 

Morphology: Initial end pointed, often with an apical spine; chambers triserially arranged in 

early stages, later becoming biserial; large loop-shaped aperture. 

Remarks: Members of the genus Fursenkoina are biserial throughout and are therefore distinct 

from this species which is triserial in its early stages, although this is not obvious in all 

specimens. Loeblich and Tappan (1987) describe the genus of Stainforthia as triserial in its early 

stages and twisted biserial later. 

Figures: 

Plate 171, fig. 1, Alaminos Canyon Core PC5 Sample 0-1 cm, side view; fig. 2, Alaminos 

Canyon Core PC5 Sample 1-2 cm, side view; fig. 3, Alaminos Canyon Core PC15 Sample 10-11 

cm, side view; fig. 4, Mississippi Canyon Core P-IM2 Sample 0-1 cm, side view; fig. 5, Green 

Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Stainforthia compressa (Bailey) 

Bulimina compressa Bailey, 1851, p. 12, pl. 12, figs. 35-37. 

Virgulina compressa (Bailey) – Cushman, 1922, p. 116, pl. 24, figs. 2-3. 

Virgulina compressa (Bailey) – Phleger and Parker, 1951, p. 19, pl. 9, figs. 4a, b, 5a, b.

Fursenkoina compressa (Bailey) – Denne, 1990, pl. 6, fig. 8. 

Morphology: Shell elongate, slightly tapered, apical end rounded; early chambers arranged 

triserially, later chambers arranged biserially; sutures depressed; aperture elongate with a serrate 

toothplate. 

Remarks: As with Stainforthia complanata this species is not a member of the genus 

Fursenkoina. This species is triserial in its early stages, later becoming biserial. 

76
Figures: 

Plate 172, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Stainforthia pontoni (Cushman)

Virgulina pontoni Cushman, 1932, p. 17, pl. 3, fig. 7. 

Virgulina pontoni Cushman – Phleger and Parker, 1951, p. 19, pl. 9, figs. 9a, b, 10a, b. 

Virgulina pontoni Cushman – Andersen, 1961, p. 92, pl. 20, fig. 3. 

Fursenkoina pontoni Cushman – Denne, 1990, pl. 6, fig. 10. 

Morphology: Shell elongate, tapering, apical end rounded; early chambers arranged triserially, 

later chambers arranged biserially; chambers overlapping; sutures greatly depressed; aperture is 

elongate with a toothplate. 

Remarks: See the comments about generic assignation for the two previous species belonging to 

genus Stainforthia. Stainforthia pontoni differs from S. compressa in its smaller size, more 

tapered profile and greatly overlapping chambers. 

Figures: 

Plate 173, fig. 1, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, side view; fig. 2, Green 

Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view; fig. 3, from Green Canyon (Block 

272) Core P-55 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Superfamily Buliminacea 

Family Buliminidae 

Genus Bulimina 

Bulimina aculeata d’Orbigny 

Bulimina aculeata d’Orbigny, 1826, p. 269, no. 7. 

Bulimina aculeata d’Orbigny – Brady, 1884, p. 406, pl. 51, figs. 7-9. 

Bulimina aculeata d’Orbigny – Cushman, 1922, p. 96-98, pl. 22, figs. 1, 2. 

Bulimina aculeata d’Orbigny – Phleger and Parker, 1951, p. 15, pl. 7, fig. 20. 

Bulimina aculeata d’Orbigny – Parker, 1954, p. 510, pl. 6, fig. 19. 

Bulimina aculeata d’Orbigny – Barker, 1960, p. 104, pl. 51, figs. 7-9. 

Bulimina aculeata d’Orbigny – Pflum and Frerichs, 1976, pl. 1, fig. 8. 

Bulimina aculeata d’Orbigny – Poag, 1981, p. 48-49, pl. 21, fig. 1; pl. 22, figs. 1a, 1b. 

Bulimina aculeata d’Orbigny – Denne, 1990, pl. 2, fig. 17. 

Bulimina aculeata d’Orbigny – Jones, 1994, p. 56, pl. 51, figs. 7-9. 

77
Morphology: Tapering shell broadest at apertural end; many long spines fringing the lower 

portion of the uppermost chambers and covering the bottom chambers; aperture with partial lip. 

Figures: 

Plate 27, figs. 1a-b, south of Mississippi Canyon control sample DGOMB2 C7, 1a: side view, 

1b: apertural view; figs. 2-3, south of Mississippi Canyon control sample DGOMB2 C7, side 

view. 

Scale bars = 50 μm; except in plate 27, fig. 1b, scale bar is 20 μm. 

Bulimina alazanensis Cushman

Bulimina alazanensis Cushman, 1927, p. 161, pl. 25, fig. 4. 

Bulimina alazanensis Cushman – Phleger and Parker, 1951, p. 16, pl. 7, figs. 24, 29. 

Bulimina alazanensis Cushman – Parker, 1954, p. 510, pl. 6, fig. 21. 

Bulimina rostrata alazanensis Cushman – Pflum and Frerichs, 1976, pl. 1, fig. 9. 

Bulimina alazanensis Cushman – Poag, 1981, p. 49, pl. 21, fig. 3; pl. 22, figs. 3a, 3b. 

Bulimina alazanensis Cushman – Denne, 1990, pl. 2, fig. 18. 

Morphology: Chambers and sutures obscured by prominent longitudinal costae that extend down 

the shell ending in spinose projections; aperture with partial lip. 

Figures: 

Plate 28, figs. 1a-b, south of Mississippi Canyon control sample DGOMB2 C7, 1a: side view, 

1b: apertural view; figs. 2-3, south of Mississippi Canyon control sample DGOMB2 C7, side 

view. 

Scale bars = 50 μm; except in fig. 1b scale bar is 10 μm. 

Bulimina marginata d’Orbigny 

Bulimina marginata d’Orbigny, 1826, p. 269, pl. 7, figs. 27, 28. 

Bulimina marginata d’Orbigny – Brady, 1884, p. 405, pl. 51, figs. 3-5. 

Bulimina marginata d’Orbigny – Cushman, 1922, p. 91, 92, pl. 21, figs. 4, 5. 

Bulimina marginata d’Orbigny – Phleger and Parker, 1951, p. 16, pl. 7, figs. 27, 28. 

Bulimina marginata d’Orbigny – Parker, 1954, p. 510, pl. 6, fig. 20. 

Bulimina marginata d’Orbigny – Barker, 1960, p. 104, pl. 51, figs. 3-5. 

Bulimina marginata d’Orbigny – Andersen, 1961, p. 88, pl. 19, figs. 14-16. 

Bulimina marginata d’Orbigny – Poag, 1981, p. 49, pl. 21, fig. 2; pl. 22, figs. 2a, 2b. 

Bulimina marginata d’Orbigny – Denne, 1990, pl. 2, fig. 19. 

Bulimina marginata d’Orbigny – Jones, 1994, p. 55, pl. 51, figs. 3-5. 

Morphology: Slightly tapered; numerous inflated smooth chambers; later chambers overhanging 

earlier chambers, with protruding short spines; aperture with a distinct rim. 

Remarks: This is the type species of the genus Bulimina d’Orbigny 1826. 

78
Figures: 

Plate 29, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

close up of aperture; figs. 2a-b, Green Canyon (Block 272) Core P-33 Sample 1-2 cm, 2a: side 

view, 2b: close up of aperture; fig. 3, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 

side view. 

Scale bars = 50 μm; except in figs. 1b and 2b scale bars are 20 μm. 

Bulimina mexicana Cushman

Bulimina inflata Seguenza – Brady, 1884, p. 406, pl. 51, figs. 10-13. 

Bulimina inflata Seguenza var. mexicana Cushman, 1922, p. 95, pl. 21, fig. 2. 

Bulimina striata d’Orbigny var. mexicana Cushman – Phleger and Parker, 1951, p. 16, pl. 7, figs. 

26, 32. 

Bulimina striata mexicana Cushman – Parker, 1954, p. 511, pl. 6, fig. 24. 

Bulimina costata d’Orbigny? – Barker, 1960, p. 104, pl. 51, figs. 11, 13. 

Bulimina striata d’Orbigny var. mexicana Cushman – Barker, 1960, p. 104, pl. 51, figs. 10, 12. 

Bulimina striata mexicana Cushman – Poag, 1981, p. 50, pl. 21, fig. 4; pl. 22, figs. 4a, 4b. 

Bulimina mexicana Cushman – Hermelin, 1989, p. 63, pl. 11, fig. 6. 

Bulimina mexicana Cushman – Denne, 1990, pl. 2, fig. 21. 

Bulimina mexicana Cushman – Jones, 1994, p. 56, pl. 51, figs. 10-13. 

Morphology: Shell expanding from the apex with the final three or four chambers forming about 

half of the shell; final chambers relatively smooth, otherwise shell wall ornamented with costae 

that terminate in prominent spines overhanging the previous chambers; loop-shaped aperture 

with lip. 

Remarks: The costae on the specimens from this study seem to be less pronounced than those on 

the figures of Cushman (1927) and Phleger and Parker (1951). Hermelin (1989) discusses the 

differences between Bulimina inflata Seguenza and Bulimina striata d’Orbigny and considers 

Bulimina mexicana Cushman to be a separate taxon. The differences between Bulimina spicata

Phleger and Parker 1951 and B. mexicana are unclear; they may be the same species. Bulimina 

mexicana differs from B. marginata in having costae ornamenting the shell wall. The spines of 

B. marginata are restricted to the edge of the chamber overhang and do not extend up the 

chamber walls. 

Figures: 

Plate 30, figs. 1a-b, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: side view, 1b: apertural 

view; figs. 2a-b, south of Mississippi Canyon control sample DGOMB2 C7, 2a: side view, 2b: 

apertural view; fig. 3, Farnella Canyon Core P-61 Sample 0-1 cm, side view. 

Scale bars = 50 μm; except in figs. 1b and 2b scale bars are 20 μm. 

79
Genus Praeglobobulimina 

Praeglobobulimina ovata (d’Orbigny) 

Bulimina ovata d’Orbigny, 1846, p. 185, pl. 11, figs. 13, 14. 

Bulimina ovata d’Orbigny – Brady, 1884, p. 400, pl. 50, figs. 13a, b. 

Bulimina affinis d’Orbigny – Cushman, 1922, p. 103, 104, pl. 20, fig. 6. 

Bulimina affinis d’Orbigny – Phleger and Parker, 1951, p. 15, pl. 7, figs. 21, 22. 

Globobulimina affinis (d’Orbigny) – Parker, 1954, p. 511, pl. 6, fig. 25; pl. 7, figs. 1, 2. 

Bulimina notovata Chapman – Barker, 1960, p. 102, pl. 50, figs. 13a, b. 

Bulimina affinis d’Orbigny – Andersen, 1961, p. 88, pl. 19, figs. 11a, b. 

Globobulimina affinis (d’Orbigny) – Denne, 1990, pl. 6, fig. 13. 

Praeglobobulimina ovata (d’Orbigny) – Jones, 1994, p. 54, pl. 50, figs. 13a, b. 

Morphology: Conical in shape, often with an acutely pointed apical end; sutures slightly 

depressed; aperture loop-shaped with a folded and crenulate toothplate (see plate 133, fig. 1b). 

Figures: 

Plate 133, figs. 1a-b, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: side view, 1b: apertural 

view (partial blockage of the aperture caused by secondary growth of calcite); fig. 2, Farnella 

Canyon Core P-61 Sample 0-1 cm, side view; figs. 3a-b, Mississippi Canyon Core P-69 Sample 

0-1 cm, 3a: side view, 3b: apertural view; fig. 4, Alaminos Canyon Core PC15 Sample 11-12 cm, 

side view. 

Scale bars = 50 μm in 1a, 2, 3a, and 4; 20 μm in 1b and 3b. 

Praeglobobulimina ovula (d’Orbigny) 

Bulimina ovula d’Orbigny, 1839a, p. 51, pl. 1, figs. 10-11. 

Globobulimina ovula (d’Orbigny) – Denne, 1990, pl. 6, fig. 14. 

Morphology: Ovate in shape; broadly rounded apical end with 1-3 short three spines; loopshaped aperture with a folded toothplate. 

Figures: 

Plate 134, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 1a: side view, 1b: 

apertural view; fig. 2, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

80
Family Uvigerinidae 

Genus Uvigerina 

Uvigerina auberiana d’Orbigny

Uvigerina auberiana d’Orbigny, 1839b, p. 106, pl. 2, figs. 23, 24. 

Uvigerina proboscidea Schwager, 1866, p. 250, pl. 7, fig. 96. 

Uvigerina auberiana d’Orbigny – Cushman, 1923, p. 163, pl. 42, figs. 3, 4. 

Uvigerina auberiana d’Orbigny – Parker, 1954, p. 519, pl. 7, fig. 38, pl. 8, fig. 1. 

Uvigerina proboscidea Schwager – Boltovskoy, 1978, pl. 8, figs. 22, 23. 

Uvigerina auberiana d’Orbigny – Hermelin, 1989, p. 64, 65, pl. 12, figs. 4, 5. 

Uvigerina auberiana d’Orbigny – Denne, 1990, pl. 14, fig. 4. 

Morphology: Shell initially triserial, later biserial; small spines randomly distributed over the 

shell surface but more common on the early chambers; aperture terminal on a neck with a collar. 

Remarks: Hermelin (1989) noted that many small, finely pitted uvigerinid species have been 

described in the literature and most seem to be conspecific with U. auberiana and should be 

regarded as junior synonyms. 

Figures: 

Plate 184, fig. 1, Alaminos Canyon Core PC15 Sample 9-10 cm, side view; fig. 2, Alaminos 

Canyon Core PC15 Sample 8-9 cm, side view; fig. 3, Green Canyon (Block 180) Core P-EP15 

Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Uvigerina bellula Bandy

Uvigerina auberiana d’Orbigny var. laevis Goës, 1896, p. 51. 

Uvigerina auberiana d’Orbigny var. laevis Goës – Cushman, 1923, p. 163. 

Uvigerina auberiana d’Orbigny var. laevis Goës – Phleger and Parker, 1951, p. 18, pl. 8, figs. 

12-14. 

Uvigerina laevis Goës – Parker, 1954, p. 520, pl. 8, fig. 4. 

Uvigerina bellula Bandy – Andersen, 1961, p. 90, pl. 20, fig. 15. 

Uvigerina laevis Goës – Poag, 1981, p. 86, pl. 27, fig. 4; pl. 28, figs. 4a, b. 

Uvigerina laevis Goës – Denne, 1990, pl. 14, fig. 8. 

Morphology: Elongate with triserially arranged chambers; shell wall mostly smooth with a few 

small spines or warts; terminal aperture located at the end of a neck with a distinct lip. 

81
Remarks: Bandy (1956) created the name Uvigerina bellula to replace Uvigerina auberiana

d’Orbigny var. laevis Goës 1896 (= U. laevis Goës of later authors) because U. laevis Ehrenberg 

1845 is a different species and has priority (i.e., the name laevis is preoccupied and not 

available). Uvigerina bellula differs from U. auberiana in having a smoother shell surface and 

less inflated and more elongate chambers. 

Figures: 

Plate 185, fig. 1, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view; fig. 2, 

Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Uvigerina flintii Cushman

Uvigerina flintii Cushman, 1923, p. 165, pl. 42, fig. 13. 

Uvigerina flintii Cushman – Phleger and Parker, 1951, p. 18, pl. 8, figs. 15-16. 

Uvigerina flintii Cushman – Parker, 1954, p. 520, pl. 8, fig. 2. 

Uvigerina flintii Cushman – Andersen, 1961, p. 89, pl. 20, fig. 17. 

Uvigerina flintii Cushman – Pflum and Frerichs, 1976, pl. 7, fig. 9. 

Uvigerina flintii Cushman – Poag, 1981, p. 86, pl. 27, fig. 5; pl. 28, figs. 5a, b. 

Uvigerina flintii Cushman – Denne, 1990, pl. 14, fig. 5. 

Morphology: Elongate to ovate in shape; coiled triserially; sutures depressed and partially hidden 

by ornamentation; shell wall ornamented by longitudinal costae which cross the sutures and 

often fuse with the costae of later chambers; terminal aperture at the end of a spirally costate 

neck. 

Figures: 

Plate 186, fig. 1, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, side view; figs. 2-3, 

Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, side view. 

Scale bars = 50 μm. 

Uvigerina peregrina Cushman

Uvigerina peregrina Cushman, 1923, p. 166, pl. 42, figs. 7-10. 

Uvigerina peregrina Cushman – Phleger and Parker, 1951, p. 18, pl. 8, figs. 22, 24-26. 

Uvigerina peregrina Cushman – Parker, 1954, p. 521, pl. 8, fig. 5. 

Uvigerina peregrina mediterranea Hofker – Pflum and Frerichs, 1976, pl. 8, fig. 1. 

Uvigerina peregrina peregrina Cushman – Pflum and Frerichs, 1976, pl. 8, fig. 2, 3. 

Uvigerina peregrina dirupta Todd – Pflum and Frerichs, 1976, pl. 8, fig. 4, 5. 

Uvigerina peregrina Cushman – Hermelin, 1989, p. 66, pl. 12, figs. 6-8. 

Uvigerina peregrina Cushman – Denne, 1990, pl. 14, fig. 10. 

Uvigerina peregrina dirupta Todd – Denne, 1990, pl. 14, fig. 11. 

82
Morphology: Triserial; shell wall ornamented with plate-like costae which do not continue across 

the chambers, hispid ornamentation also apparent in many specimens; terminal aperture present 

at the end of a neck with a distinct lip. 

Remarks: Hermelin (1989) notes that some workers separate various species of Uvigerina and 

subspecies of U. peregrina on the basis of the portion of the hispid or costate ornamentation 

covering the test. This is now recognized to be related to ecophenotypic variation, and, 

therefore, his concept of U. peregrina includes most uvigerinids with a costate or aligned hispid 

ornamentation (Hermelin, 1989). We have followed Hermlin in this regard. 

Figures: 

Plate 187, figs. 1-4, Mississippi Canyon Core P-69 Sample 1-2 cm, side view; fig. 5, south of 

Mississippi Canyon control sample DGOMB2 C7, side view. 

Scale bars = 50 μm. 

Genus Trifarina 

Trifarina bradyi Cushman 

Rhabdogonium tricarinatum Brady, 1884, p. 525, pl. 67, figs. 1-3. 

Trifarina bradyi Cushman, 1923, p. 99, pl. 22, figs. 3-9. 

Trifarina bradyi Cushman – Phleger and Parker, 1951, p. 18, pl. 8, figs. 10, 11. 

Trifarina bradyi Cushman – Parker, 1954, p. 522, pl. 8, fig. 9. 

Trifarina bradyi Cushman – Barker, 1960, p. 140, pl. 67, figs. 1-3. 

Trifarina bradyi Cushman – Denne, 1990, pl. 13, fig. 12. 

Trifarina bradyi Cushman – Jones, 1994, p. 78, 79, pl. 67, figs. 1-3. 

Morphology: Triserial in early stages, later uniserial, often slightly twisted, triangular in crosssection and carinate; terminal aperture at the end of a neck with a lip. 

Remarks: This is the type species of the genus Trifarina Cushman 1923. 

Figures: 

Plate 180, figs. 1-2, Green Canyon Core P-51 Sample 1-2 cm, side view; fig. 3, Green Canyon 

Core P-51 Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Trifarina jamaicensis (Cushman and Todd) 

Angulogerina jamaicensis Cushman and Todd, 1945, p. 53, pl. 8, figs. 3. 

Angulogerina jamaicensis Cushman and Todd – Parker, 1954, p. 521, 522, pl. 8, fig. 8.

Angulogerina jamaicensis Cushman and Todd – Denne, 1990, pl. 1, fig. 11. 

Morphology: Triserial throughout, with pronounced but discontinuous costae and short spines. 

83
Figures: 

Plate 181, fig. 1, Mississippi Canyon Core P-69 Sample 1-2 cm, side view; fig. 2, Green Canyon 

(Block 272) Core P-33 Sample 1-2 cm, side view; fig. 3, Green Canyon (Block 272) Core P-55 

Sample 0-1 cm, side view. 

Scale bars = 50 μm. 

Superfamily Fursenkoinacea 

Family Fursenkoinidae 

Genus Fursenkoina 

Fursenkoina seminuda (Natland)

Virgulina seminuda Natland, 1938, p. 145, pl. 5, fig. 12. 

Virgulina tessellata Phleger and Parker, 1951, p. 19, pl. 9, figs. 15a, 15b, 16a, 16b. 

Virgulina tessellata Phleger and Parker – Parker, 1954, p. 513, pl. 7, fig. 12. 

Fursenkoina seminuda (Natland) – Denne, 1990, pl. 6, fig. 10. 

Morphology: Fusiform shell; sinuous axial suture; lower part of the chambers is semi-opaque 

(see plate 65, figs. 1a and 2a). 

Remarks: Phleger and Parker (1951) noted that V. tessellata differed from V. seminuda in being 

less elongate, having a more rounded initial end and fewer chambers. These differences are 

within the limits of natural variation and thus V. tessellata is a junior synonym. 

Figures: 

Plate 65, figs. 1a-c, Mississippi Canyon Core P-69 Sample 0-1 cm, 1a: side view, 1b: edge view, 

1c: apertural view; figs. 2a-d, Mississippi Canyon Core P-69 Sample 1-2 cm, 2a: side view, 2b: 

edge view, 2c: apertural view, 2d: top view. 

Scale bars = 50 μm in figs. 1a-b and 2a-b, d; 20 μm in figs. 1c and 2c. 

Genus Rutherfordoides 

Rutherfordoides mexicanus (Cushman)

Virgulina mexicana Cushman, 1922, p. 120, pl. 23, fig. 8. 

Virgulina mexicana Cushman – Phleger and Parker, 1951, p. 19, pl. 9, figs. 6a, b, 7a, b, 8a, b. 

Virgulina mexicana Cushman – Parker, 1954, p. 512, pl. 7, figs. 7, 8. 

Virgulina mexicana Cushman – Andersen, 1961, p. 91, pl. 20, figs. 4a-c. 

Fursenkoina mexicana (Cushman) – Poag, 1981, p. 65, 66, pl. 33, fig. 7; pl. 34, figs. 7a, b. 

Rutherfordoides mexicana (Cushman) – Denne, 1990, 12, figs. 3a, b. 

84
Morphology: Biserial with bent initial end; oblique chambers, broad and low on dorsal side; 

strongly overlapping and inflated on ventral side; elongate aperture. 

Figures: 

Plate 158, figs. 1a-b, Mississippi Canyon Core P-69 Sample 1-2 cm, 1a: side view, 1b: apertural 

view; figs. 2a-b, Mississippi Canyon Core P-IM2 Sample 0-1 cm, 2a: side view, 2b: side view; 

figs. 3a-b, Mississippi Canyon Core P-69 Sample 0-1 cm, 3a: side view, 3b: apertural view; figs. 

4a-b, Mississippi Canyon Core P-IM2 Sample 0-1 cm, 4a: side view, 4b: side view. 

Scale bars = 50 μm. 

Superfamily Stilostomellacea 

Family Stilostomellidae 

Genus Siphonodosaria 

Siphonodosaria calomorpha (Reuss)

Nodosaria calomorpha Reuss, 1866, p. 129, pl. 3, figs. 15-19. 

Nodosaria calomorpha Reuss – Brady, 1884, p. 497, pl. 61, fig. 7. 

Nodosaria calomorpha Reuss – Cushman, 1923, p. 67, pl. 12, fig. 13. 

Nodosaria calomorpha Reuss – Barker, 1960, p. 128, pl. 61, figs. 23-27. 

Glandulonodosaria calomorpha (Reuss) – Jones, 1994, p. 72, pl. 61, figs. 23-26, ?27. 

Siphonodosaria calomorpha (Reuss) – Denne, 1990, p. 324

Morphology: Shell elongate; frequently arcuate; both ends broadly rounded; 3-5 chambers; 

aperture a simple opening. 

Figures: 

Plate 166, figs. 1a-b, Alaminos Canyon Core PC4 Sample 0-1 cm, 1a: side view, 1b: apertural 

view; figs. 2-3, Alaminos Canyon Core PC5 Sample 1-2 cm, side view.

Scale bars = 50 μm. 

2.2.9. Order Rotaliida 

Superfamily Discorbacea 

Family Bagginidae 

Genus Neocrosbyia 

Neocrosbyia minuta (Parker)

Valvulineria minuta Parker, 1954, p. 527, pl. 9, figs. 4, 5, 6. 

Neocrosbyia minuta (Parker) – Denne, 1990, pl. 9, fig. 2. 

85
Morphology: Shell concavo-convex; rounded periphery; deep umbilicus on ventral side; 7-8 

chambers in final whorl, chambers increasing gradually in size, inflated, more so on ventral side; 

small, narrow flap extending a short way into the umbilicus, previous flaps, appearing as small 

angular or rounded projections, can be seen in the umbilicus, especially in earlier whorls (e.g., 

plate 108, figs. 1b and 2b); sutures distinct, slightly curved, depressed, more so on ventral side; 

aperture a slit running from the periphery to the umbilicus, sometimes extending onto dorsal side 

(e.g., plate 108, figs. 3a and 4). 

Figures: 

Plate 108, figs. 1a-b, Alaminos Canyon Core PC-5 Sample 1-2 cm, 1a: dorsal view, 1b: ventral 

view showing umbilical projections; figs. 2a-b, Green Canyon (Block 272) Core P-57 Sample 0-

1 cm, 2a: dorsal view, 2b: ventral view showing umbilical projections; figs. 3a-b, Green Canyon 

(Block 180) Core P-EP15 Sample 0-1 cm, 3a: dorsal view, 3b: ventral view with umbilical 

projections obscured by detritus, 3c: edge view showing apertural slit; fig. 4, Alaminos Canyon 

Boxcore B19.1 Sample 0-1 cm, dorsal view. 

Scale bars = 50 μm. 

Genus Valvulineria 

Valvulineria glabra Cushman

Valvulineria vilardeboana (d’Orbigny) var. glabra Cushman, 1927, p. 161, pl. 4, figs. 5, 6. 

Valvulineria sp. Parker, 1948, p. 240, pl. 4, figs. 13a, b. 

Valvulineria laevigata Phleger and Parker, 1951, p. 25, pl. 13, figs. 11a, 11b, 12a, 12b.

Rotamorphina laevigata (Phleger and Parker) – Parker, 1954, p. 537, pl. 11, figs. 10, 11. 

Rotamorphina glabra (Cushman) – Andersen, 1961, p. 114, pl. 23, figs. 2a-c.

Quadrimorphina laevigata (Phleger and Parker) - Denne, 1990, pl. 11, fig. 13. 

Morphology: Shell biconvex, composed of 2 to 21⁄2 whorls; 5 chambers in final whorl; chambers 

increasing in size; sutures slightly depressed, more so on ventral side, slightly curved; periphery 

rounded; aperture interiomarginal to umbilical hidden beneath a lobe-like plate with serrations 

extending from the final chamber over the umbilical area. 

Remarks. Valvulineria glabra was considered to be larger than V. laevigata by Phleger and 

Parker (1951). However, this size difference was considered to be inconsequential by Andersen 

(1961) who observed specimens spanning the size range between the two species. Therefore V. 

laevigata is synonymized with V. glabra. 

Figures: 

Plate 189, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: dorsal view, 

1b: ventral view, 1c: edge view showing apertural opening; fig. 2, Mississippi Canyon Core P-69 

Sample 0-1 cm, dorsal view; figs. 3a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 

3a: dorsal view, 3b: ventral view; fig. 4, south of Mississippi Canyon control sample DGOMB2 

C7, ventral view; fig. 5, Green Canyon Core P-51 Sample 1-2 cm, ventral view. 

86
Carbonate overgrowth visible on the shell. 

Scale bars = 50 μm. 

Valvulineria mexicana Parker 

Valvulineria cf. araucana (d’Orbigny) – Phleger and Parker, 1951 (not Rosalina araucana

d’Orbigny, 1839a), p. 25, pl. 13, figs. 7a, b, 8a, b. 

Valvulineria mexicana Parker, 1954, p. 526-527, pl. 9, figs. 1-3. 

Valvulineria mexicana Parker – Andersen, 1961, p. 104, pl. 23, figs. 1a-c. 

Morphology: Shell biconvex; periphery rounded; 6-7 chambers in final whorl; chambers 

increasing in size, later chambers inflated on ventral side, especially towards the umbilicus (e.g., 

plate-fig. 3a); sutures depressed, particularly on ventral side near the umbilicus, curved on dorsal 

side; shell wall coarsely perforate with more perforations on later chambers, perforations 

frequently aligned along sutures on dorsal side; aperture interiomarginal to umbilical, covered by 

large lobe-like flap extending from the final chamber over the umbilical area, the flap from the 

penultimate chamber often visible below and to one side of it (e.g., plate 190, figs. 1b and 2b), 

flaps of earlier chambers occasionally visible (e.g., plate 190, fig. 3b). 

Remarks: This species differs from V. glabra in its larger size, more inflated chambers on ventral 

side and visibility of earlier chamber flaps in the umbilical area. 

Figures: 

Plate 190, figs. 1-2, Mississippi Canyon Core P-61 Sample 0-1 cm, a: dorsal view, b: ventral 

view; figs. 3a-b, Mississippi Canyon Core P-61 Sample 0-1 cm, 3a: edge view showing apertural 

opening, 3b: ventral view. 

Scale bars = 50 μm. 

Family Eponididae 

Genus Eponides 

Eponides turgidus Phleger and Parker 

Eponides turgidus Phleger and Parker, 1951, p. 22, pl. 11, figs. 9a, b. 

Eponides turgidus Phleger and Parker – Parker, 1954, p. 530, pl. 9, figs. 22, 23. 

Eponides turgidus Phleger and Parker – Poag, 1981, p. 65, pl. 1, fig.4; pl. 2, figs. 4a, 4b. 

Alabaminella turgida Phleger and Parker – Denne, 1990, pl. 1, figs. 3a, 3b. 

Morphology: Rounded periphery; sutures slightly depressed on dorsal side, more depressed on 

ventral side; closed umbilicus on ventral side; aperture a low arched opening, covered by a flap, 

midway between the umbilicus and the periphery with small pustules around it. 

87
Remarks: In some specimens the periphery is lobulate, the spire is higher, and the sutures are 

more depressed (compare plate figures 2 and 3). This species was placed in the genus 

Alabaminella by Denne (1990). However, it does not possess a minute slitlike aperture which is 

a feature of Alabaminella and therefore it is retained in Eponides. This is a small species and Sen 

Gupta et al. (1987) noted that it was almost absent from the >125 μm and >250 μm fractions of 

an Antarctic Bottom Water assemblage in the Equatorial North Atlantic, although smaller 

specimens were common. 

Figures: 

Plate 61, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 1a: dorsal view, 1b: 

ventral view, 1c: edge view; figs. 2-3, Alaminos Canyon Core PC-5 Sample 1-2 cm, dorsal view; 

fig. 4, Alaminos Canyon Core PC-5 Sample 1-2 cm, ventral view. 

Scale bars = 50 μm. 

Genus Ioanella 

Ioanella tumidula (Brady)

Truncatulina tumidula Brady, 1884, p. 666, pl. 95, figs. 8a-d. 

Eponides tumidulus (Brady) – Phleger and Parker, 1951, p. 21, pl. 11, figs. 7a, 7b, 8a, 8b. 

Eponides tumidulus (Brady) – Parker, 1954, p. 529, pl. 9, figs. 19, 24. 

“Eponides” cf. “E” tumidulus (Brady) – Andersen, 1961, p. 103, pl. 24, figs. 5a-c. 

Eponides tumidulus (Brady) – Pflum and Frerichs, 1976, pl. 2, figs. 4, 5. 

Eponides tumidulus (Brady) – Poag, 1981, p. 64, 65, pl. 1, fig. 2; pl. 2, figs. 2a, 2b. 

Ioanella tumidula (Brady) – Loeblich and Tappan, 1987, p. 549, 550, pl. 595, figs. 4-9. 

Ioanella tumidula (Brady) – Denne, 1990, pl. 7, fig. 8. 

Morphology: Lobulate outline; dorsal side inflated with depressed sutures; umbilicus open on 

ventral side, umbilicus surrounded by secondary apertures (plate 85, figs. 1b, 2); aperture low, 

arched, open from umbilicus to edge. 

Remarks: This species was placed in genus Ioanella Saidova by Loeblich and Tappan (1987) on 

the basis of the secondary apertures surrounding the umbilicus (plate 85, figs. 1b, 2) which 

differentiate it from Eponides turgidus. 

Figures: 

Plate 85, figs. 1a-b, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: dorsal view, 1b: ventral 

view; fig. 2, Farnella Canyon Core P-61 Sample 0-1 cm, ventral view. 

Scale bars = 50 μm. 

88
Family Rosalinidae 

Genus Gavelinopsis 

Gavelinopsis translucens (Phleger and Parker)

“Rotalia” translucens Phleger and Parker, 1951, p. 24, pl. 12, figs. 11a, 11b, 12a, 12b. 

“Rotalia” translucens Phleger and Parker – Parker, 1954, p. 531, pl. 10, figs. 3, 7. 

Gavelinopsis translucens (Phleger and Parker) – Denne, 1990, pl. 6, figs. 12a, 12b. 

Morphology: Shell small and biconvex; periphery acute with a keel; sutures on dorsal side flush 

with the surface and are indistinct on SEM images; last two or more sutures on ventral side 

depressed; umbilical plug present on ventral side; shell wall finely perforate on dorsal side and 

more coarsely perforate on ventral side; aperture present midway between periphery and 

umbilicus. 

Remarks: Plate 67, figs. 2a and 2b show encrusting barite rosettes on shell walls. 

Figures: 

Plate 67, figs. 1a-b, Mississippi Canyon Core P-73 Sample 1-2 cm, 1a: dorsal view, 1b: ventral 

view; figs. 2a-b, Mississippi Canyon Core P-73 Sample 3-4 cm, 2a: dorsal view, 2b: ventral 

view; figs. 3a-b, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 3a: dorsal view, 3b: 

ventral view. 

Scale bars = 50 μm. 

Superfamily Siphoninacea 

Family Siphoninidae 

Genus Siphonina 

Siphonina pulchra Cushman

Siphonina pulchra Cushman, 1919, p. 42, pl. 14, figs. 7a-c. 

Siphonina pulchra Cushman – Cushman, 1931, p. 69, pl. 14, figs. 2, 3. 

Siphonina pulchra Cushman – Phleger and Parker, 1951, p. 24, pl. 12, figs. 15a, b. 

Siphonina pulchra Cushman – Parker, 1954, p. 532, pl. 10, fig. 11, 12. 

Siphonina pulchra Cushman – Andersen, 1961, p. 104, pl. 24, figs. 2a-c. 

Siphonina pulchra Cushman – Denne, 1990, pl. 12, fig. 12a-b. 

Morphology: Shell almost circular in outline, biconvex; periphery subacute; carinate juveniles 

but keel frequently absent or only partially developed in adult specimens; about 5 chambers in 

last whorl; shell wall perforate; aperture on a short neck with a lip. 

89
Remarks: Contradicting the original description of S. pulchra by Cushman (1919) that “S. 

pulchra has no fimbriated periphery and practically no keel,” the Jamaican specimen illustrated 

by Cushman (1931, pl. 14, fig. 3) clearly shows a fimbriated edge zone. The carina in S. pulchra

is a variable feature with young stages carinate (Cushman, 1931) but not as well developed as in 

S. bradyana. The development of the carina in our specimens is variable. 

Figures: 

Plate 165, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: dorsal view, 

1b: ventral view, 1c: edge view showing aperture; figs. 2a-b, Green Canyon (Block 180) Core PEP15 Sample 0-1 cm, 2a: dorsal view, 2b: ventral view; fig. 3, juvenile from Green Canyon 

(Block 180) Core P-EP15 Sample 0-1 cm, dorsal view. 

Scale bars = 50 μm. 

Superfamily Discorbinellacea 

Family Pseudoparrellidae 

Genus Epistominella 

Epistominella exigua (Brady)

Pulvinulina exigua Brady, 1884, p. 696, pl. 103, figs. 13, 14. 

Pseudoparrella exigua (Brady) – Phleger and Parker, 1951, p. 28, pl. 15, figs. 6a, 6b, 7a, 7b. 

Epistominella exigua (Brady) – Parker, 1954, p. 533, pl. 10, figs. 22, 23. 

Epistominella exigua (Brady) – Poag, 1981, p. 63, pl. 5, fig. 4, pl. 6, figs. 4a-c. 

Epistominella exigua (Brady) – Boltovskoy, 1978, pl. 3, figs. 37, 38. 

Epistominella exigua (Brady) – Hermelin, 1989, p. 67, 68. 

Epistominella exigua (Brady) – Denne, 1990, pl. 5, fig. 11. 

Alabaminoides exiguus (Brady) – Jones, 1994, p. 103, pl. 103, figs. 13-14. 

Morphology: Shell biconvex; sutures straight and flush with the surface, indistinct in SEM 

images. 

Remarks: This is a small species and Hermelin (1989) noted that it is often excluded or underrepresented in studies that use the >125 μm or larger size fraction. Sen Gupta et al. (1987) found 

reduced abundances of E. exigua in the >250 μm fraction of an Antarctic Bottom Water 

assemblage from the equatorial North Atlantic. 

Figures: 

Plate 59, figs. 1-2, Mississippi Canyon Core P-69 Sample 0-1 cm, a: dorsal view, b: ventral 

view; figs. 3a-b, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 3a: dorsal view, 3b: edge 

view showing the aperture. 

Scale bars = 50 μm. 

90
Epistominella vitrea Parker

Epistominella vitrea Parker in Parker et al., 1953, p. 9, pl. 4, figs. 34-36, 40, 41. 

Epistominella vitrea Parker – Parker, 1954, p. 534, pl. 10, figs. 20, 26. 

Epistominella vitrea Parker – Andersen, 1961, p. 104, 105, pl. 24, figs. 1a-c. 

Epistominella vitrea Parker – Poag, 1981, p. 63, 64, pl. 5, fig. 3; pl. 6, figs. 3a, 3b. 

Epistominella vitrea Parker – Denne, 1990, pl. 7, figs. 12a, b. 

Morphology: Shell biconvex, lobate periphery; sutures curved and depressed on dorsal side, 

almost straight and depressed on ventral side, difficult to see in SEM figures. 

Remarks: Epistominella vitrea Parker differs from E. exigua in having curved and depressed 

sutures. 

Figures: 

Plate 60, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 1a: dorsal view, 1b: 

ventral view, 1c: edge view showing the aperture. 

Scale bars = 50 μm. 

Family Discorbinellidae 

Genus Laticarinina 

Laticarinina pauperata (Parker and Jones) 

Pulvinulina repanda Fichtel and Moll var. menardii d’Orbigny subvar. pauperata Parker and 

Jones, 1865, p. 395, pl. 16, figs. 50, 51. 

Laticarinina pauperata (Parker and Jones) – Galloway and Wissler, 1927. 

Laticarinina pauperata (Parker and Jones) – Phleger and Parker, 1951, p. 32, pl. 18, fig. 3. 

Laticarinina pauperata (Parker and Jones) – Parker, 1954, p. 540, pl. 12, fig. 3. 

Laticarinina halphora (Stache) – Barker, 1960, p. 214, pl. 104, figs. 3-11. 

Laticarinina pauperata (Parker and Jones) – Pflum and Frerichs, 1976, pl. 1, fig. 10. 

Laticarinina pauperata (Parker and Jones) – van Morkhoven et al., 1986, p. 89, 90, pl. 26, figs. 

1a-c. 

Laticarinina halophora (Stache) – Hermelin, 1989, p.68, 69. 

Laticarinina pauperata (Parker and Jones) – Denne, 1990, pl. 7, fig. 16. 

Laticarinina pauperata (Parker and Jones) – Jones, 1994, p. 104, pl. 104, figs. 3-11. 

Morphology: Shell biconvex to planoconvex in edge view, dorsal side flattened, broad thin keel 

around the periphery. 

Remarks: Hermelin (1989) followed Barker (1960) in using the name Laticarinina halophora

(Stache) for this species, remarking that L. halophora and Brady’s L. pauperata were identical. 

However, Hornibrook (1971) has shown that Robulina halophora Stache belongs to the 

nodosariid genus Planularia. 

91
Figures: 

Plate 96, fig. 1, Alaminos Canyon Boxcore B6 bulk sample, dorsal view; figs. 2-3, De Soto 

Canyon control sample DGOMB2 S36, dorsal view. 

Scale bars = 50 μm. 

Superfamily Planorbulinacea 

Family Planulinidae 

Genus Planulina 

Planulina ariminensis d’Orbigny

Planulina ariminensis d’Orbigny, 1826, p. 280, pl. 14, figs. 1-3, 3bis; Modèles, no. 49. 

Anomalina ariminensis (d’Orbigny) – Brady, 1884, p. 674, pl. 93, figs. 10, 11. 

Planulina ariminensis d’Orbigny – Cushman, 1931, p. 110. 

Planulina ariminensis d’Orbigny – Phleger and Parker, 1951, p. 32, pl. 18, figs. 4a, b. 

Planulina ariminensis d’Orbigny – Parker, 1954, p. 540, pl. 11, figs. 27, 30. 

Planulina ariminensis d’Orbigny – Barker, 1960, p. 192, pl. 93, figs. 10, 11. 

Planulina ariminensis d’Orbigny – Poag, 1981, p. 75, pl. 43, fig. 3; pl. 44, figs. 3a, b. 

Planulina ariminensis d’Orbigny – van Morkhoven et al., 1986, p. 38, 40, pl. 10, figs. 1-4. 

Planulina ariminensis d’Orbigny – Denne, 1990, pl. 10, figs. 7a, b. 

Planulina ariminensis d’Orbigny – Jones, 1994, p. 98, pl. 93, figs. 10-11. 

Morphology: Shell planispiral, somewhat evolute on both sides; periphery truncate, sides of the 

test flattened and almost parallel; chambers distinct; sutures strongly curved, papillate, more 

frequently on ventral side; aperture peripheral extending onto dorsal side. 

Remarks: The sutures in some of our specimens are not as strongly curved as most mentioned 

and figured in the literature. This is the type species by subsequent designation of the genus 

Planulina d’Orbigny 1826. 

Figures: 

Plate 127, figs. 1a-c, Green Canyon (Block 272) Core P-55 Sample 1-2 cm, 1a: ventral view, 1b: 

dorsal view, 1c: peripheral view; figs. 2a-c, Garden Banks (Block 427) Core P-EP8 Sample 1-2 

cm, 2a: ventral view, 2b: dorsal view, 2c: peripheral view; fig. 3, Green Canyon (Block 272) 

Core P-51 Sample 0-1 cm, dorsal view. 

Scale bars = 50 μm. 

Plate 128, digital photomicrographs, fig. 1, Mississippi Canyon tubeworm MC2; fig. 2, Green 

Canyon tubeworm EC4; fig. 3, Green Canyon tubeworm EC5; fig. 4, Mississippi Canyon 

tubeworm MC5. 

Plate 129, fig. 1, Mississippi Canyon tubeworm MC3, specimen attached to hydroid (arrow), 

digital photomicrograph; fig. 2, same specimen as arrow in fig. 1; fig. 3, same specimen as fig. 1 

of plate 128; fig. 4, Mississippi Canyon tubeworm MC4, attachment side. 

92
Planulina foveolata (Brady)

Anomalina foveolata Brady, 1884, p. 674, pl. 94, figs. 1a-c. 

Planulina foveolata (Brady) – Cushman, 1931, p. 111, pl. 20, figs. 2, 3. 

Planulina foveolata (Brady) – Phleger and Parker, 1951, p. 33, pl. 18, figs. 9a, b, 10a, b. 

Planulina foveolata (Brady) – Parker, 1954, p. 540, pl. 11, figs. 25, 26. 

Planulina foveolata (Brady) – Barker, 1960, p. 194, pl. 94, fig. 1. 

Planulina faveolata (Brady) – Andersen, 1961, p. 125, pl. 29, figs. 4a-c. 

Planulina foveolata (Brady) – van Morkhoven et al., 1986, p. 26, 28, pl. 5, figs. 1-3. 

Planulina foveolata (Brady) – Poag, 1981, p. 76, pl. 43, fig. 1; pl. 44, figs. 1a, 1b. 

Planulina foveolata (Brady) – Denne, 1990, pl. 10, figs. 9a, b. 

Planulina foveolata (Brady) – Jones, 1994, p. 98, pl. 94, fig. 1. 

Morphology: Shell planispiral, somewhat evolute on both sides, plano-convex; periphery 

rounded; chambers distinct, 9 to 11 in the last whorl; sutures curved, strongly raised on ventral 

side; aperture peripheral, extending onto dorsal side. 

Remarks: The shells of our specimens are frequently not pristine and in some cases not wellpreserved. This accounts for the holes and broken chambers in the SEM images. 

Figures: 

Plate 130, figs. 1-2, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, a: ventral view, b: 

dorsal view, c: peripheral view; figs. 3a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 

cm, 3a: ventral view, 3b: dorsal view. 

Scale bars = 50 μm. 

Family Cibicididae 

Genus Cibicides 

Cibicides lobatulus (Walker and Jacob)

Nautilus lobatulus Walker and Jacob, in Kanmacher, 1798, p. 642, pl. 14, fig. 36. 

Truncatulina lobatula Walker and Jacob – Brady, 1884, p. 660, pl. 92, fig. 10; pl. 93, figs. 1, 4, 

5; pl. 115, figs. 4, 5. 

Cibicides lobatula (Walker and Jacob) – Cushman, 1931, p. 118, pl. 21, fig. 3. 

Cibicides lobatulus (Walker and Jacob) – Barker, 1960, p. 190, pl. 92, fig. 10; p. 192, pl. 93, figs. 

1, 4, 5; p. 236, pl. 115, figs. 4, 5. 

Cibicides lobatulus (Walker and Jacob) – Jones, 1994, p. 97, pl. 92, fig. 10; pl. 93, figs. 1, 4, 5; 

p. 114, pl. 115, figs. 4, 5. 

Morphology: Shell planoconvex; periphery acute with a thickened keel; dorsal side typically 

adherent, dorsal side falttened or concave, ventral side convex; sutures sigmoidal, depressed on 

ventral side and flush on dorsal side; aperture peripheral extending onto dorsal side. 

93
Remarks: Lobatula Fleming 1828, based on this species, is regarded as a valid genus by Loeblich 

and Tappan (1987). There is, however, no distinguishing character on which Lobatula can be 

unequivocally separated from Cibicides; we agree with Jones (1994) that Lobatula is a junior 

synonym of Cibicides. 

Figures: 

Plate 39, figs. 1a-b, Alaminos Canyon Core PC15 Sample 8-9 cm, 1a: ventral view, 1b: dorsal 

view. 

Scale bars = 50 μm. 

Cibicides wuellerstorfi (Schwager) 

Anomalina wuellerstorfi Schwager, 1866, p. 258, pl. 7, figs. 105, 107. 

Truncatulina wuellerstorfi (Schwager) – Brady, 1884, p. 662, pl. 93, figs. 8, 9. 

Planulina wuellerstorfi (Schwager) – Cushman, 1931, p. 110, 111; pl. 19, figs. 5, 6. 

Planulina wuellerstorfi (Schwager) – Phleger and Parker, 1951, p. 33, pl. 18, figs. 11a, 11b; pl. 

19, figs. 1a, 1b, 2a, 2b, 3a, 3b. 

Cibicides wuellerstorfi (Schwager) – Parker, 1954, p. 544, pl. 13, figs. 3, 6. 

Planulina bradii Tolmachoff – Barker, 1960, p. 192, pl. 93, fig. 8. 

Planulina wuellerstorfi (Schwager) – Barker, 1960, p. 192, pl. 93, fig. 9. 

Cibicides wuellerstorfi (Schwager) – Pflum and Frerichs, 1976, pl. 4, figs. 2-4. 

Cibicides wuellerstorfi (Schwager) – Srinivasan and Sharma, 1980, p. 56, 57, pl. 8, figs. 11-13 

(neotype). 

Cibicides wuellerstorfi (Schwager) – Poag, 1981, p. 52, pl. 3, fig. 1; pl. 4, figs. 1a, 1b. 

Planulina wuellerstorfi (Schwager) – van Morkhoven et al., 1986, p. 48-50, pl. 14, figs. 1-2. 

Cibicidoides wuellerstorfi (Schwager) – Hermelin, 1989, p. 87. 

Cibicides wuellerstorfi (Schwager) – Sen Gupta, 1989, p. 706, pl. 1, figs. 1-7; pl. 2, figs. 1-6; pl. 

3, figs. 1-6; pl. 4, figs. 1-7. 

Cibicides wuellerstorfi (Schwager) – Denne, 1990, pl. 4, figs. 1a, 1b. 

Cibicidoides wuellerstorfi (Schwager) – Jones, 1994, p. 98, pl. 93, figs. 8, 9. 

Morphology: Shell planoconvex to slightly biconvex (in a small minority); periphery with a 

thickened keel; sutures strongly sigmoidal on both sides, raised to flush with the surface on 

dorsal side, raised with some surface roughening on ventral side; sutures meet near central plug 

on ventral side; central plug more pronounced in variants (plate 40, fig. 3); aperture peripheral 

extending onto dorsal side. 

Remarks: There has been considerable confusion over the generic placement of Anomalina 

wuellerstorfi Schwager 1866. It has generally been assigned to Planulina d’Orbginy 1826, 

Cibicides Dénys de Montfort 1808, or Cibicidoides Thalmann 1939. In addition, it is the type 

species of the genus Fontbotia González-Donoso and Linares 1970 (based on a 

misunderstanding of the wall structure). Fontbotia is regarded by Sen Gupta (1989) as a junior 

synonym of Cibicides Dénys de Montfort 1808 and by Jones (1994) as a junior synonym of 

Cibicidoides Thalmann 1939. Anomalina wuellerstorfi is not a member of Planulina as only the 

94
final whorl is visible on ventral side, and it is not a member of Cibicidoides as it is mainly planoconvex and not biconvex (Sen Gupta, 1989). All significant features of this species are 

characteristic of Cibicides Dénys de Montfort 1808 (Sen Gupta, 1989). 

Figures: 

Plate 40, figs. 1a-c, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: ventral view, 1b: dorsal 

view, 1c: edge view; figs. 2a-b, Farnella Canyon Core P-61 Sample 0-1 cm, 2a: ventral view, 2b: 

dorsal view; fig. 3, juvenile from De Soto Canyon control sample DGOMB2 S36, ventral view. 

Scale bars = 50 μm. 

Plate 41, fig. 1, Alaminos Canyon tubeworm LS4, digital photomicrograph. 

Genus Cibicidoides 

Cibicidoides pachyderma (Rzehak) 

Truncatulina pachyderma Rzehak, 1886, p. 87, pl. 1, figs. 5a-c. 

Truncatulina ungeriana (d’Orbigny) – Brady, 1884, p. 664, pl. 94, figs. 9a-c. 

Not Rotalina ungeriana d’Orbigny, 1826 and 1846. 

Cibicides pseudoungeriana (Cushman) – Cushman, 1931, p. 123-124, pl. 22, figs. 3-7. 

Cibicides aff. floridanus (Cushman) – Phleger and Parker, 1951, p. 30, pl. 16, figs. 1a, 1b, 2a, 2b, 

3a, 3b, 4a, 4b. 

Cibicides aff. floridanus (Cushman) – Parker, 1954, p. 541, pl. 12, figs. 5, 9. 

Cibicides cf. pseudoungerianus (Cushman) – Pflum and Frerichs, 1976, pl. 2, fig. 9. 

Cibicidoides “floridanus” forma bathyalis Poag, 1981, p. 53, pl. 29, fig. 1; pl. 30, figs. 1a, 1b. 

Cibicidoides “floridanus” forma sublittoralis Poag, 1981, p. 53, 54, pl. 29, fig. 2; pl. 30, figs. 2a, 

2b. 

Cibicidoides pachyderma (Rzehak) – van Morkhoven et al., 1986, p. 68-70, pl. 22, figs. 1a-c. 

Cibicidoides pachydermus bathyalis (Poag) – Denne, 1990, pl. 4, figs. 5a, 5b. 

Cibicidoides pachydermus sublittoralis (Poag) – Denne, 1990, pl. 4, figs. 6a, 6b. 

Morphology: Shell biconvex; periphery moderately to sharply carinate; 10-12 chambers in final 

whorl; sutures flush to slightly depressed on ventral side, flush on dorsal side, gently curved in 

smaller specimens, more sharply curved in larger specimens; coarsely perforate shell wall on 

dorsal side; flat to slightly convex umbonal boss; aperture an interiomarginal arch with a lip, 

continuing onto dorsal side. 

Remarks: Specimens of C. pachyderma differ from similar-size specimens of C. robertsonianus

in its more sharply curved sutures. The only difference between Cibicides and Cibicidoides is in 

the shape of the shell—planoconvex in the former and biconvex in the latter, and this is a 

gradational feature in some species. Thus, the placement of Cibicidoides in a separate family 

(Paralleloididae) by Loeblich and Tappan (1987) is not justified; we keep it in the Cibicididae. 

95
Figures: 

Plate 42, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 1-2 cm, 1a: ventral view, 1b: 

dorsal view, 1c: edge view; figs. 2a-b, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, 2a: 

ventral view, 2b: dorsal view; fig. 3, Alaminos Canyon Core PC4 Sample 0-1 cm, ventral view; 

figs. 4a-c, Green Canyon (Block 272) Core P-55 Sample 1-2 cm, 4a: ventral view, 4b: dorsal 

view, 4c: edge view. 

Scale bars = 50 μm. 

Plate 43, digital photomicrographs, fig. 1, Mississippi Canyon tubeworm MC1, d – dislodged 

specimen (dorsal surface), s – attachment scar; figs. 2a-c, specimen in fig. 1, 2a: ventral surface 

before cleaning, 2b: dorsal surface, 2c: ventral surface after cleaning. 

Cibicidoides robertsonianus (Brady)

Truncatulina robertsoniana Brady, 1881, p. 65; 1884, p. 664-665, pl. 95, figs. 4a-c. 

Cibicides robertsonianus (Brady) – Phleger and Parker, 1951, p. 31, pl. 16, figs. 10a, 10b, 11a, 

11b, 12a, 12b, 13a, 13b. 

Cibicides robertsonianus (Brady) – Parker, 1954, p. 543, pl. 13, figs. 2, 5. 

Cibicides robertsonianus (Brady) – Barker, 1960, p. 196, pl. 95, fig. 4. 

Cibicides robertsonianus (Brady) – Pflum and Frerichs, 1976, pl. 3, figs. 3-5. 

Cibicidoides robertsonianus (Brady) – Poag, 1981, p. 54, 55, pl. 5, fig. 1; pl. 6, figs. 1a, 1b. 

Cibicidoides robertsonianus (Brady) – van Morkhoven et al., 1986, p. 41-43, pl. 11, figs. 1a-c. 

Cibicidoides robertsonianus (Brady) – Hermelin, 1989, p. 86. 

Cibicidoides robertsonianus (Brady) – Denne, 1990, pl. 4, figs. 7a, 7b. 

Cibicidoides robertsonianus (Brady) – Jones, 1994, p. 99, pl. 95, fig. 4. 

Morphology: Shell planoconvex to biconvex; periphery angular and slightly lobulate; not 

entirely involute ventral side with visible earlier whorls; sutures gently curved, flush with 

surface, final two may be slightly depressed; coarsely perforate shell wall on dorsal side; 

aperture an interiomarginal arch with a lip, continuing onto dorsal side. 

Remarks: The imperforate keel mentioned in Hermelin (1989) and van Morkhoven et al. (1986) 

is not present in all our specimens. 

Figures: 

Plate 44, figs. 1a-d, Alaminos Canyon Core PC4 Sample 1-2 cm, 1a: ventral view, 1b: dorsal 

view, 1c: ventral view, digital photomicrograph, 1d: dorsal view, digital photomicrograph. 

Plate 45, figs. 1a-c, Garden Banks (Block 427) Core P-EP8 Sample 1-2 cm, 1a: ventral view, 1b: 

dorsal view, 1c: edge view showing apertural opening; figs. 2a-b, Green Canyon (Block 272) 

Core P-55 Sample 1-2 cm, 2a: ventral view, 2b: dorsal view. 

Scale bars = 50 μm. 

96
Superfamily Asterigerinacea 

Family Epistomariidae 

Genus Nuttallides 

Nuttallides decorata (Phleger and Parker) 

Pseudoparrella (?) decorata Phleger and Parker, 1951, p. 28, pl. 15, figs. 4a, b, 5a, b. 

Epistominella decorata (Phleger and Parker) – Parker, 1954, p. 533, pl. 10, figs18, 19. 

Alabamina decorata (Phleger and Parker) – Pflum and Frerichs, 1976, pl. 6, figs. 8, 9. 

Nuttallides decorata (Phleger and Parker) – Poag, 1981, p. 74, pl. 5, fig. 2; pl. 6, figs. 2a, 2b. 

Nuttallides decorata (Phleger and Parker) – Denne, 1990, pl. 9, figs. 9a, 9b. 

Morphology: Shell biconvex, trochospiral; periphery subacute to rounded; 21⁄2 whorls visible on 

dorsal side; 5 to 6 chambers in last whorl; sutures depressed and slightly curved; pores visible 

around the edge of some specimens on ventral side (see plate 110, fig. 5) and on the last whorl 

on dorsal side; shell wall ornamented with raised processes, occassionally clustered and 

coalescing on dorsal side of the species. 

Remarks: Raised processes ornamenting the shell walls give this species a roughened appearance 

under the light microscope. 

Figures: 

Plate 110, figs. 1, 3, 5, Alaminos Canyon Core PC15 Sample A10, 1: dorsal view, 3: dorsal view, 

5: ventral view; figs. 2, 4, Alaminos Canyon Core PC15 Sample A8, 2: dorsal view, 4: ventral 

view; fig. 6, Alaminos Canyon Core PC4 Sample 0-1 cm, cross-sectional view showing pyrite 

framboids filling the calcareous shell indicative of a reducing environment. 

Scale bars = 50 μm. 

Superfamily Nonionacea 

Family Nonionidae 

Genus Nonionella 

Nonionella iridea Heron-Allen and Earland 

Nonionella iridea Heron-Allen and Earland, 1932, p. 438, pl. 16, figs. 14-16. 

Morphology: Shell coiled; periphery rounded, slightly lobulate; 8-9 chambers present, only 6-7 

exposed on one side, chambers with a flaplike projection slightly overhanging the umbilicus; 

proloculus appears as a central boss; sutures depressed, curved; aperture an interiomarginal arch 

extending onto ventral side. 

Remarks: Heron-Allen and Earland in their original (1932) description of this species note that it 

consists of 12-14 chambers. However, their type figure shows 9 chambers on dorsal side and 7 

97
chambers visible on ventral side. Our specimens have 8 to 9 chambers, and in all other respects 

match the description of Heron-Allen and Earland (1932); hence we use the name N. iridea. 

Figures: 

Plate 109, figs. 1, 3, south of Mississippi Canyon control sample DGOMB2 C7, a: dorsal view, 

b: ventral view, c: edge view; figs. 2a-b, south of Mississippi Canyon control sample DGOMB2 

C7, 2a: dorsal view, 2b: edge view. 

Scale bars = 50 μm. 

Genus Laminononion 

Laminononion tumidum (Cushman and Edwards) 

Nonionina stelligera (d’Orbigny) – Brady, 1884, p. 728, pl. 109, fig. 5. 

Astrononion tumidum Cushman and Edwards, 1937, p. 33, pl. 3, fig. 17. 

Astrononion tumidum Cushman and Edwards – Barker, 1960, pl. 109, fig. 5. 

Astrononion (Laminononion) tumidum Cushman and Edwards – Hornibrook, 1964, p. 335. 

Astrononion tumidum Cushman and Edwards – Denne, 1990, pl. 1, fig. 15. 

Astrononion tumidum Cushman and Edwards – Jones, 1994, p. 107, pl. 109, fig. 5. 

Morphology: Shell planispiral; periphery broadly rounded; 6-7 chambers in final whorl; sutures 

depressed; large rhomboidal to subtriangular plate extending from lateral ends of the chamber 

across the umbilical margin to attach to the posterior border, opening along the posterior 

peripheral margin; aperture an interiomarginal, equatorial slit. 

Remarks: This is the type species of the subgenus Astrononion (Laminononion) Hornibrook 

1964; this subgenus was raised to the rank of genus (Laminononion) by Loeblich and Tappan, 

1987. 

Figures: 

Plate 94, fig. 1, Green Canyon (Block 272) Core P-55 Sample 0-1 cm, side view. 

Scale bar = 50 μm. 

Plate 95, fig. 1, Mississippi Canyon tubeworm EC1, digital photomicrograph; fig. 2, Green 

Canyon tubeworm EC5; fig. 3, Mississippi Canyon tubeworm EC1. 

Genus Melonis 

Melonis affinis (Reuss)

Nonionina affine Reuss, 1851, p. 72, pl. 5, figs. 32a-b. 

Nonionina barleeana Williamson, 1858, p. 32, pl. 3, figs. 68-69. 

Nonionina umbilicatula (Montagu) – Brady, 1884, p. 726, pl. 109, figs. 8-9. 

98
Nonion barleeanum (Williamson) – Cushman, 1930, p. 11, pl. 4, fig. 5. 

Nonion formosum (Seguenza) – Phleger et al., 1953, p. 30, pl. 6, fig. 5. 

Nonion formosum (Seguenza) – Parker, 1954, p. 506, pl. 6, fig. 3.

Gavelinonion barleeanum (Williamson) – Barker, 1960, p. 224, pl. 109, figs. 8-9. 

“Nonion” barleeanus (Williamson) – Andersen, 1961, p. 82, 83, pl. 18, figs. 6a, b. 

Melonis barleeanus (Williamson) – Pflum and Frerichs, 1976, pl. 7, figs. 5, 6. 

Nonion affine (Reuss) – Boltovskoy, 1978, p. 162, pl. 5, figs. 1-2. 

Melonis barleeanum (Williamson) – Hermelin, 1989, p. 88, pl. 17, fig. 12. 

Melonis barleeanum (Williamson) – Denne, 1990, pl. 8, fig. 16. 

Melonis affinis (Reuss) – Van Marle, 1991, p. 186-187, pl. 20, figs. 1-3. 

Melonis affinis (Reuss) – Jones, 1994, p. 107-108, pl. 109, figs. 8-9. 

Morphology: Shell planispiral, involute with deeply excavated umbilici; periphery broadly 

rounded; usually more than 10 chambers in the final whorl; sutures flush with the surface, 

curved; walls coarsely perforate; aperture a low arch extending from umbilicus to umbilicus. 

Remarks: Van Marle (1991) noted that Melonis barleeanum (Williamson) is a junior synonym of 

Melonis affinis (Reuss) with which we agree. Melonis affinis differs from M. pompilioides in 

being more compressed in edge view. 

Figure: 

Plate 103, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; figs. 2a-b, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 2a: side 

view: 2b: apertural view. 

Scale bars = 50 μm. 

Melonis pompilioides (Fichtel and Moll)

Nautilus pompilioides Fichtel and Moll, 1798, p. 31, pl. 2, figs. a-c. 

Nonionina pompilioides (Fichtel and Moll) – Brady, 1884, p. 727, pl. 109, figs. 10-11. 

Nonion pompilioides (Fichtel and Moll) – Cushman, 1930 (part), p. 4, pl. 1, figs. 7-8, 10-11.

Nonion pompilioides (Fichtel and Moll) – Phleger and Parker, 1951, p. 11, pl. 5, figs. 19, 20. 

Nonion pompilioides (Fichtel and Moll) – Parker, 1954, p. 506, 507, pl. 6, fig. 4. 

Melonis sphaeroides Voloshinova, 1958, p. 153, pl. 3, figs. 1a, 1b. 

Melonis pompilioides (Fichtel and Moll) – Voloshinova, 1958, p. 158, pl. 3, figs. 8-9. 

Nonion (?) pompilioides (Fichtel and Moll) – Barker, 1960, p. 224, pl. 109, figs. 10-11. 

Melonis pompilioides (Fichtel and Moll) – Pflum and Frerichs, 1976, pl. 7, figs. 7, 8. 

Nonion pompilioides (Fichtel and Moll) – Boltovskoy, 1978, pl. 5, figs. 3-4. 

Melonis pompilioides (Fichtel and Moll) – Hermelin, 1989, p. 88, 89, pl. 17, figs. 13-14. 

Melonis pompilioides (Fichtel and Moll) – van Morkhoven, Berggren and Edwards, 1986, p. 72-

80, pl. 23A, figs. 1-2; pl. 23C, figs. 1a-d; pl. 23D, figs. 1a-d, pl. 23E, figs. 1a-c. 

Melonis pompilioides (Fichtel and Moll) – Jones, 1994, p. 108, pl. 109, figs. 10-11. 

99
Morphology: Shell planispiral, involute with deeply excavated umbilici; periphery broadly 

rounded; 8 to 10 chambers in the final whorl; sutures flush with the surface; walls coarsely 

perforate; aperture a low arch extending from umbilicus to umbilicus. 

Remarks: Melonis pompilioides differs from M. affinis in a being a broader species in edge view. 

These two species also occur as two separate populations with no intergradation. Melonis 

pompilioides is found in the deeper water locations of Alaminos and Farnella Canyons and M. 

affinis occurs in the shallower Green Canyon and Garden Banks. Parker and Phleger (1951) also 

observed this depth separation. They stated that there were two forms of Nonion pompilioides, a 

typical form and a more compressed form that resembles N. barleeanum and is confined to 

shallower depths. Hermelin (1989) noted several different species names that appear to be 

ecophenotypic variants and therefore junior synonyms of M. pompilioides. He stated that more 

investigation is required before it can be decided if M. barleeanum and M. pompilioides are 

ecophenotypic variants. We have kept M. barleeanum (= M. affinis in this study) and M. 

pompilioides separated as two species. There remains some uncertainty surrounding Cushman’s 

(1930) figured specimens of N. pompilioides from the Atlantic. Those resembling our figured 

specimens are recognized by us as M. pompilioides; the thinner specimens (Cushman, 1930; pl. 1 

fig. 9, pl. 2, figs. 1-2) belong to a different species. 

Figures: 

Plate 104, figs. 1, 3, Alaminos Canyon Core PC5 Sample 2-3 cm, 1: side view, 3: apertural view; 

fig. 2, Alaminos Canyon Core PC5 Sample 4-5 cm, side view. 

Scale bars = 50 μm. 

Genus Pullenia 

Pullenia bulloides (d’Orbigny) 

Nonionina bulloides d’Orbigny, 1826, p. 293, model no. 2 (nomen nudum). 

Nonionina bulloides d’Orbigny, 1846, p. 107, pl. 5, figs. 9-10. 

Pullenia sphaeroides (d’Orbigny) – Brady, 1884, p. 615-616, pl. 84, figs. 12-13. 

Pullenia bulloides (d’Orbigny) – Phleger and Parker, 1951, p. 29, pl. 15, fig. 11. 

Pullenia bulloides (d’Orbigny) – Parker, 1954, p. 538, pl. 11, fig. 17. 

Pullenia bulloides (d’Orbigny) – Barker, 1960, p. 174, pl. 84, figs. 12-13. 

Pullenia bulloides (d’Orbigny) – Andersen, 1961, p. 115, pl. 25, figs. 8a, b. 

Pullenia bulloides (d’Orbigny) – Papp and Schmid, 1985, p. 45, pl. 34, figs. 6-9 (lectotypes). 

Pullenia bulloides (d’Orbigny) – Hermelin, 1989, p. 78, 79, pl. 15, figs. 4-5. 

Pullenia bulloides (d’Orbigny) – Denne, 1990, pl. 11, figs. 5a, b. 

Pullenia bulloides (d’Orbigny) – Jones, 1994, p. 92, pl. 84, figs. 12-13. 

Morphology: Shell planispiral, involute, sphaeroidal; 4 chambers in the last whorl; sutures flush 

with surface to slightly depressed; aperture an interiomarginal slit extending from one umbilicus 

to the other, with a lip. 

100
Remarks: Nonionina bulloides d’Orbigny 1846 is the type species of the genus Pullenia Parker 

and Jones (in Carpenter et al., 1862. It has been lectotypified by Papp and Schmid (1985). 

Figures: 

Plate 143, figs. 1a-b, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: side view, 1b: 

apertural view; figs. 2a-b, Alaminos Canyon Boxcore B19.1 Sample 0-1 cm, 2a: side view, 2b: 

apertural view. 

Scale bars = 50 μm. 

Pullenia quinqueloba (Reuss) 

Nonionina quinqueloba Reuss, 1851, p. 71, pl. 5, fig. 31. 

Pullenia quinqueloba (Reuss) – Brady, 1884, p. 616, pl. 84, figs14, 15. 

Pullenia subcarinata (d’Orbigny) – Heron-Allen and Earland, 1932, p. 403-404, pl. 13, figs. 14-

18. 

Pullenia quinqueloba (Reuss) – Phleger and Parker, 1951, p. 29, pl. 15, figs. 12, 13. 

Pullenia quinqueloba (Reuss) – Parker, 1954, p. 538, pl. 11, fig. 16. 

Pullenia subcarinata (d’Orbigny) – Barker, 1960, p. 174, pl. 84, figs. 14, 15. 

Pullenia subcarinata (d’Orbigny) – Andersen, 1961, p. 115, pl. 25, figs. 9a, b. 

Pullenia subcarinata subcarinata (d’Orbigny) – Boltovskoy, 1978, p. 167, pl. 6, figs. 21-22. 

Pullenia subcarinata quinqueloba (Reuss) – Boltovskoy, 1978, p. 166, pl. 6, figs. 23, 24. 

Pullenia subcarinata (d’Orbigny) – Hermelin, 1989, p. 79, 80, pl. 15, figs. 6, 7. 

Pullenia subcarinata (d’Orbigny) – Denne, 1990, pl. 11, figs. 6a, 6b. 

Pullenia quinqueloba (Reuss) – Jones, 1994, p. 92, pl. 84, figs. 14-15. 

Morphology: Shell planispiral, involute, compressed; 5 to 6 chambers in the last whorl; sutures 

slightly depressed and slightly curved; aperture an interiomarginal slit extending from one 

umbilicus to the other, with a lip. 

Remarks: Pullenia quinqueloba differs from P. bulloides in having more compressed shape and a 

greater number of chambers in the final whorl. 

Figures: 

Plate 144, figs. 1a-b, Mississippi Canyon Core P-69 Sample 0-1 cm, 1a: side view, 1b: apertural 

view; fig. 2, Alaminos Canyon Core PC4 Sample 1-2 cm, side view; fig. 3, Alaminos Canyon 

Boxcore B19.1 Sample 0-1 cm, side view; fig. 4, Alaminos Canyon Core PC15 Sample 12-13 

cm, side view; fig. 5, Alaminos Canyon Core PC15 Sample 7-8 cm, side view. 

Scale bars = 50 μm. 

101
Genus Haynesina 

Haynesina germanica (Ehrenberg) 

Nonionina germanica Ehrenberg, 1840, p. 23. 

Nonionina germanica Ehrenberg – Ehrenberg, 1841, pl. 2, fig. 1a-g. 

Nonionina depressula (Walker and Jacob) – Brady, 1884, p. 725, 726, pl. 109, fig. 6a, b. 

Nonion germanicum (Ehrenberg) – Cushman, 1930, p. 8, pl. 3, figs. 5a, b, ?4a, b. 

Nonion depressulum (Walker and Jacob) – Barker, 1960, p. 224, pl. 109, figs. 6a, b. 

Haynesina germanica (Ehrenberg) – Banner and Culver, 1978, p. 191-200, pl. 4, figs. 1-6; pl. 5, 

figs. 1-8; pl. 6, figs. 1-7; pl. 7, figs. 1-6; pl. 8, figs. 1-10; pl. 9, figs. 1-11, 15, 17, 18. 

Morphology: Shell planispiral, involute with depressed umbilici; periphery rounded; 8-10 

chambers in the final whorl, chamber walls bending inward posteriorly and fusing with 

preceding septal face; sutures curved, depressed, deeply incised near umbilicus; prominent 

tubercles along sutures and over umbilicus; primary aperture a low interiomarginal arch, 

occasionally obscured by abundant pustules; intercameral apertures present. 

Figures: 

Plate 76, fig. 1, Green Canyon (Block 272) Core P-57 Sample 1-2 cm, side view; fig. 2, Green 

Canyon (Block 272) Core P-54 Sample 0-1 cm, side view; fig. 3, Mississippi Canyon Core PIM4 Sample 1-2 cm, side view. 

Superfamily Chilostomellacea 

Family Chilostomellidae 

Genus Chilostomella 

Chilostomella oolina Schwager

Chilostomella oolina Schwager, 1878, p. 527, pl. 1, fig 16. 

Chilostomella ovoidea Reuss – Brady, 1884 (part), p. 436, pl. 55, figs. 12-14, 17-18 (not 15-16, 

19-23). 

Chilostomella oolina Schwager – Phleger and Parker, 1951, p. 29, pl. 15, fig. 10. 

Chilostomella oolina Schwager – Parker, 1954, p. 537, pl. 11, fig. 15. 

Chilostomella oolina Schwager – Barker, 1960, p. 112, pl. 55, figs. 12-14, 17-18. 

Chilostomella oolina Schwager – Andersen, 1961, p. 114, pl. 25, fig. 7. 

Chilostomella oolina Schwager – Hermelin, 1989, p. 76, pl. 14, fig. 5. 

Chilostomella oolina Schwager – Denne, 1990, pl. 3, fig. 13. 

Chilostomella oolina Schwager – Jones, 1994, p. 61, pl. 55, figs. 12-14, 17-18. 

102
Morphology: Shell elongate; ends broadly rounded; chambers coiled, each 180° apart from the 

preceding chamber; last formed chamber comprising more than half the test; aperture a narrow 

slit at the suture between the ultimate and penultimate chambers. 

Remarks: Our specimens are fragile and often have holes in the final chamber. 

Figures: 

Plate 38, fig. 1, Farnella Canyon Core P-61 Sample 0-1 cm, side view; fig. 2, Mississippi Canyon 

Core P-IM4 Sample 1-2 cm, side view. Carbonate overgrowth visible on shell. 

Scale bars = 50 μm. 

Family Osangulariidae 

Genus Osangularia 

Osangularia culter (Parker and Jones)

Planorbulina farcta (Fichtel and Moll) var. ungeriana (d’Orbigny) subvar. culter Parker and 

Jones, 1865, p. 421, pl. 19, figs. 1a-c. 

Truncatulina culter (Parker and Jones) – Brady, 1884, p. 668, pl. 96, figs. 3a-c. 

Parrella cultur (Parker and Jones) – Phleger and Parker, 1951, p. 23, pl. 12, figs. 3a, 3b. 

Osangularia cultur (Parker and Jones) – Parker, 1954, p. 530, 531, pl. 9, figs. 29, 30. 

Osangularia bengalensis (Schwager) – Barker, 1960, p. 198, pl. 96, figs. 3a-c. 

Osangularia culter (Parker and Jones) – Hermelin, 1989, p. 84, pl. 16, figs. 11-13. 

Osangularia culter (Parker and Jones) – Denne, 1990, pl. 10, figs. 1a, 1b. 

Osangularia bengalensis (Schwager) – Jones, 1994, p. 100, pl. 96, fig. 3. 

Morphology: Shell biconvex; periphery with an irregular, jagged keel; sutures depressed and 

curved on ventral side and flush and straight to curved on dorsal side; aperture an interiomarginal 

slit along the base of the ultimate chamber on ventral side and may be obscured by detritus; 

additional aperture bending at an oblique angle up the apertural face and surrounded by a lip. 

Remarks: Hermelin (1989) regards Osangularia bengalensis (Schwager, 1866) as a junior 

synonym of O. culter. Some confusion exists regarding the separation of O. bengalensis and O. 

culter. However, as Revets (1996) demonstrates, O. bengalensis of Schwager has multiple 

secondary openings on the apertural face and has been placed under the genus Cribroparrella. 

These secondary openings are not present in our specimens (e.g., plate 117, figs. 1b and 2b) or in 

Brady’s specimen. 

Figures: 

Plate 117, figs. 1a-b, Mississippi Canyon Core P-69 Sample 1-2 cm, 1a: ventral view, 1b: edge 

view; figs. 2a-b, south of Mississippi Canyon control sample DGOMB2 C7, 2a: ventral view, 2b: 

edge view; figs. 3-4, De Soto Canyon control sample DGOMB2 S36, dorsal view. 

Scale bars = 50 μm. 

103
Osangularia rugosa (Phleger and Parker)

Pseudoparrella (?) rugosa Phleger and Parker, 1951, p. 28, 29, pl. 15, figs. 8a, 8b, 9a, 9b. 

Epistominella rugosa (Phleger and Parker) – Parker, 1954, p. 533, pl. 10, figs. 24, 25. 

Osangularia rugosa (Phleger and Parker) – Pflum and Frerichs, 1976, pl. 7, figs. 2-4. 

Osangularia rugosa (Phleger and Parker) – Denne, 1990, pl. 10, figs. 2a, 2b. 

Morphology: Shell biconvex; 7 to 8 chambers in final whorl, later ones can be inflated; sutures 

straight to curved, sometimes raised on dorsal side, later ones depressed on ventral side; walls 

with small raised processes giving a roughened effect; medium-sized pores mostly restricted to 

the periphery; periphery angular with a thickened keel; aperture low, arched, open from the 

umbilicus to periphery along the base of the ultimate chamber on ventral side. 

Figures: 

Plate 118, figs. 1a-c, Mississippi Canyon Core P-73 Sample 0-1 cm, 1a: dorsal view, 1b: ventral 

view, 1c: edge view; figs. 2a-b, Alaminos Canyon Core PC4 Sample 0-1 cm, 2a: ventral view, 

2b: edge view; fig. 3, Mississippi Canyon Core P-73 Sample 0-1 cm, ventral view. 

Scale bars = 50 μm. 

Family Oridorsalidae 

Genus Oridorsalis 

Oridorsalis umbonatus (Reuss)

Rotalina umbonata Reuss, 1851, p. 75, pl. 5, figs. 35a-c. 

Pulvinulina umbonata (Reuss) – Brady, 1884, p. 695, 696, pl. 105, figs. 2a-c. 

Truncatulina tenera Brady, 1884, p. 665, pl. 95, figs. 11a-c. 

Pulvinulina umbonata (Reuss) – Heron-Allen and Earland, 1932, p. 430, pl. 15, figs. 16-18. 

Eponides umbonatus (Reuss) – Phleger and Parker, 1951, p. 22, pl. 11, figs. 10a, 10b, 13a, 13b, 

14a, 14b. 

Pseudoeponides umbonatus (Reuss) – Parker, 1954, p. 530, pl. 9, figs. 20, 21. 

Eponides (?) tenera (Brady) – Barker, 1960, p. 196, pl. 95, fig. 11. 

Eponides umbonatus (Reuss) – Barker, 1960, p. 216, pl. 105, fig. 2. 

Oridorsalis umbonatus (Reuss) – Todd, 1965, p. 23, pl. 6, fig. 2. 

Oridorsalis umbonatus (Reuss) – Boltovskoy, 1978, pl. 5, figs. 5, 6. 

Oridorsalis tener tener (Brady) – Pflum and Frerichs, 1976, pl. 6, figs. 2-4. 

Oridorsalis tener umbonatus (Reuss) – Pflum and Frerichs, 1976, pl. 6, figs. 5-7. 

Oridorsalis umbonatus (Reuss) – Hermelin, 1989, p. 81, 82, pl. 16, figs. 1-5. 

Oridorsalis umbonatus (Reuss) – Denne, 1990, pl. 9, figs. 14a, 14b. 

Oridorsalis umbonata (Reuss) – Jones, 1994, p. 99, pl. 95, fig. 11. 

Oridorsalis umbonata (Reuss) – Jones, 1994, p. 104, pl. 105, fig. 2. 

104
Morphology: Shell biconvex, periphery angular, last chambers slightly lobulate; sutures flush 

with surface or slightly depressed, radial and slightly curved on dorsal side and sinusoidal on 

ventral side; aperture an interiomarginal slit extending from the umbilical area to periphery, with 

lip and pustules surrounding it. Supplementary openings at the base of the suture on dorsal side 

(plate 116, figs. 3a, 4) and occasionally following the base of the last formed chamber on dorsal 

side (plate 116, fig. 1a) and along the suture between the ultimate and penultimate chambers on 

ventral side (plate 116, figs. 1b, 2). 

Remarks: Two forms of this species are observed in the study samples. One form is typical (e.g., 

plate 116, figs. 1a-c, 2 and 4), the other is represented by much smaller specimens (e.g., plate 

116, figs. 3a-c). Phleger and Parker (1951) and Parker (1954) also observed these smaller 

specimens, whose features, except for size, are almost identical to the typical form. As noted by 

Phleger and Parker (1951), such smaller specimens may be the “Truncatulina tenera” of Brady 

(1884, pl. 95 fig. 11). Parker (1954) did not observe supplementary apertures in her small 

specimens, and Barker (1960) could not find supplementary apertures on any of his Pacific 

material. However, Jones (1994) placed Brady’s Truncatulina tener specimen (pl. 95, fig. 11) in 

Oridorsalis umbonata stating that Brady’s figured specimens lack the supplementary spiral 

sutural apertures characteristic of Oridorsalis Andersen, although these features are developed in 

some unfigured Challenger specimens. Supplementary apertures are difficult to see under the 

light microscope in such small specimens but are obvious in SEM micrographs (e.g., plate 116, 

fig. 3a). Hence, in agreement with Hermelin (1989), we consider Oridorsalis tener to be a junior 

synonym of O. umbonatus. 

Figures: 

Plate 116, figs. 1a-c, Farnella Canyon Core P-61 Sample 0-1 cm, 1a: dorsal view, 1b: ventral 

view, 1c: edge view; fig. 2, Farnella Canyon Core P-61 Sample 0-1 cm, ventral view; figs. 3a-c, 

Alaminos Canyon Core PC5 Sample 1-2 cm, 3a: dorsal view, 3b: ventral view, 3c: edge view; 

fig. 4, Farnella Canyon Core P-61 Sample 0-1 cm, dorsal view. 

Scale bars = 50 μm. 

Family Heterolepidae 

Genus Anomalinoides 

Anomalinoides globulosus (Chapman and Parr)

Anomalina grosserugosa (Guembel) – Brady, 1884, p. 673, pl. 94, figs. 4-5. 

Anomalina globulosa Chapman and Parr, 1937, p. 117, pl. 9, fig. 27. 

Anomalina globulosa Chapman and Parr – Barker, 1960, p. 194, pl. 94, figs. 4-5. 

Anomalinoides globulosus (Chapman and Parr) - van Morkhoven et al., 1986, p. 36-38, pl. 9, 

figs. 1-3. 

Anomalinoides globulosus (Chapman and Parr) – Hermelin, 1989, p. 84-85, pl. 17, figs. 1, 5. 

Cibcidoides globulosus (Chapman and Parr) – Jones, 1994, p. 98, pl. 94, figs. 4-5. 

Morphology: Shell trochospiral, umbilcus depressed; 6-7 chambers in final whorl; chambers 

globular, increasing in size; sutures depressed, curved; wall coarsely perforate, especially on 

105
ventral side; aperture a low interiomarginal slit extending from the periphery to the umbilicus, 

with a distinct lip. 

Figures: 

Plate 9, figs. 1a-c, Garden Banks (Block 427) Core P-EP8 Sample 1-2 cm, 1a: dorsal view, 1b: 

ventral view, 1c: edge view showing apertural opening. 

Scale bars = 50 μm. 

Plate 10, fig. 1, Green Canyon tubeworm EC5, digital photomicrograph. 

Anomalinoides mexicanus Parker 

Anomalinoides mexicana Parker, 1954, p. 539, pl. 11, figs. 21-23. 

Morphology: Shell involute, trochospiral; broadly rounded periphery, early portion non-lobulate, 

later portion slightly lobulate; 7-8 chambers in final whorl; sutures flush with surface, later ones 

slightly depressed, slightly curved; wall finely perforate; aperture a low interiomarginal slit 

extending from the periphery to the umbilicus, with a distinct lip. 

Figures: 

Plate 11, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 1-2 cm, 1a: ventral view, 

1b: dorsal view, 1c: edge view showing apertural opening. 

Scale bars = 50 μm. 

Family Gavelinellidae 

Genus Gyroidinoides 

Gyroidinoides sp. cf. G. polius (Phleger and Parker) 

Eponides polius Phleger and Parker, 1951, p. 21, pl. 11, figs. 1a, b, 2a, b. 

Eponides polius Phleger and Parker - Parker, 1954, p. 528-529, pl. 9, figs. 11, 12. 

Eponides polius Phleger and Parker – Poag, 1981, p. 64, pl. 1, fig. 1; pl. 2, figs. 1a, 1b. 

Gyroidinoides polius (Phleger and Parker) – Denne, 1990, pl. 7, figs. 2a, b. 

Morphology: Shell biconvex; periphery slightly lobulate; about 3 whorls in adult, 9-11 chambers 

in final whorl; sutures on dorsal side slightly curved to sharply angled, sutures on ventral side 

slightly depressed and almost straight; aperture a narrow slit-like opening between periphery and 

umbilicus on ventral side and covered by an apertural flap. 

Remarks: Parker (1954) notes that this species has “sutural” foramina, and Poag (1981) notes an 

arch in the aperture at the peripheral end. Neither of these features is apparent in our specimens. 

106
Figures: 

Plate 73, figs. 1a-c, Alaminos Canyon Core PC-5 Sample 3-4 cm, 1a: dorsal view, 1b: ventral 

view, 1c: edge view; figs. 2a-b, south of Mississippi Canyon control sample DGOMB2 C7, 2a: 

dorsal view, 2b: ventral view; fig. 3, south of Mississippi Canyon control sample DGOMB2 C7, 

ventral view. 

Scale bars = 50 μm. 

Gyroidinoides sp. cf. G. regularis (Phleger and Parker) 

Eponides regularis Phleger and Parker, 1951, p. 21, pl. 11, figs. 3a, 3b, 4a-c. 

Eponides regularis Phleger and Parker – Parker, 1954, p. 529, pl. 9, figs. 16, 17. 

Neoeponides regularis (Phleger and Parker) – Andersen, 1961, p. 103, pl. 24, figs. 5a-c. 

Eponides regularis Phleger and Parker – Pflum and Frerichs, 1976, pl. 2, figs. 1-3. 

Eponides regularis Phleger and Parker – Poag, 1981, p. 64, pl. 1, fig. 3; pl. 2, figs. 3a, 3b. 

Hansenisca regularia (Phleger and Parker) – Denne, 1990, pl. 7, figs. 5a, b. 

Morphology: Shell biconvex; nonlobulate periphery; about 4 whorls in adult, 8 or 9 chambers in 

final whorl; sutures on dorsal side slightly curved; aperture a low, arched opening between 

periphery and umbilicus on ventral side; pustules present below the aperture. 

Remarks: Our specimens are smaller and more biconvex than the holotype and have less sharply 

angled sutures on dorsal side. The pustules below the aperture are not seen on the holotype or in 

Poag’s figures. 

Figures: 

Plate 74, figs. 1a-c, Alaminos Canyon Core PC-4 Sample 1-2 cm, 1a: dorsal view, 1b: ventral 

view, 1c: edge view; fig. 2, Alaminos Canyon Core PC-4 Sample 0-1 cm, dorsal view; figs. 3a-c, 

Alaminos Canyon Core PC-13 Sample 1-2 cm, 3a: dorsal view, 3b: ventral view, 3c: edge view; 

fig. 4, Green Canyon (Block 272) Core P-57 Sample 0-1 cm, ventral view. Carbonate 

overgrowth visible on the shell. 

Scale bars = 50 μm. 

Genus Discanomalina 

Discanomalina semipunctata (Bailey) 

Rotalina semipunctata Bailey, 1851, p. 11, figs. 17-19. 

Anomalina polymorpha Costa, 1856, p. 252, pl. 21, figs. 17-19. 

Anomalina polymorpha Costa - Brady, 1884, p. 676, pl. 97, figs. 3-6. 

Anomalina semipunctata (Bailey) - Cushman, 1931, p. 106, pl. 18, figs. 1-2. 

Discanomalina japonica Asano, 1951, p. 13, figs. 3-5. 

Discanomalina japonica Asano - LeRoy and Levinson, 1974, p. 16, pl. 9, figs. 3-5. 

107
Discanomalina semipunctata (Bailey) - Medioli and Scott, 1978, p. 298-300, pl. 1, figs. 1-20, pl. 

2, figs. pl. 2, figs. 1-20. 

Discanomalina semipunctata (Bailey) - Jones, 1994, p. 100, pl. 97, figs. 3-6. 

Morphology: Shell (in our specimens) attached by ventral side and irregular in form; frequently 

with one or more spine-like projections from the rounded margin; surface coarsely perforate; 

broad ventral aperture. 

Remarks: This is the type species of the genus Discanomalina. As demonstrated by Medioli and 

Scott (1978), it has been reported under multiple names from various seas. Its habit is known to 

be free or attached (Medioli and Scott, 1978; Loeblich and Tappan, 1987). All individuals in our 

samples, however, are attached to hydroid stalks growing on tubeworms. The curved and 

concave ventral side of D. semipunctata, illustrated and described by various authors (e.g., 

Cushman, 1931), is probably related to attachment to comparable tubular surfaces. The species 

has previously been found in Pleistocene sediments from the Gulf of Mexico (LeRoy and 

Levinson, 1974), but ours is the first report of a living population. 

Figures: 

Plate 55, digital photomicrographs, figs. 1, 2, Mississippi Canyon tubeworm MC5, specimens 

attached to hydroids; figs. 3, 4, Mississippi Canyon tubeworm MC3, specimens attached to 

hydroids. 

Plate 56, fig. 1, Mississippi Canyon tubeworm MC3, specimen attached to hydroid; figs. 2, 3, 

Mississippi Canyon tubeworm MC5, specimens attached to hydroids; figs. 4, 5, Mississippi 

Canyon tubeworm MC3, specimens attached to hydroids, attachment side. 

Genus Gyroidina 

Gyroidina bradyi (Trauth)

Truncatulina dutemplei (d’Orbigny) – Brady, 1884, p. 665, pl. 95, figs. 5a-c. 

Not Rotalina dutemplei d’Orbigny, 1846. 

Truncatulina bradyi Trauth, 1918, p. 235. 

Cibicides hyalina Hofker, 1951, p. 359, figs. 244-245. 

Cibicides bradyi (Trauth) – Barker, 1960, p. 196, pl. 95, fig. 5. 

Cibicides bradyi (Trauth) – Pflum and Frerichs, 1976, pl. 3, figs. 6, 7. 

Cibicides bradyi (Trauth) – Boltovskoy, 1978, pl. 3, figs. 6-8. 

Cibicidoides bradyi (Trauth) – Corliss, 1979, p. 9-10, pl. 3, figs. 1-3. 

Cibicidoides bradyi (Trauth) – van Morkhoven et al., 1986, p. 100-102, pl. 30, figs. 1-2. 

Cibicidoides bradyi (Trauth) – Hermelin, 1989, p. 85, 86, pl. 17, figs. 2-4. 

Gyroidina bradyi (Trauth) – Jones, 1994, p. 99, pl. 95, fig. 5. 

Morphology: Shell planoconvex to slightly biconvex; rounded periphery and no keel; sutures 

depressed on ventral, flush on dorsal side; dorsal side coarsely perforate; aperture extends onto 

dorsal side. 

108
Remarks: Phleger and Parker (1951) in their description of Cibicidoides robertsonianus mention 

a complete gradation between the typical form of that species and a much smaller form with a 

rounded periphery which appears to be close to the species described by Brady as Truncatulina 

dutemplei, and called Cibicides/Cibicidoides bradyi by most later workers. Both Hermelin 

(1989) and van Morkhoven et al. (1986) distinguish C. robertsonianus from C. bradyi by noting 

the larger test, more angular periphery and imperforate keel of C. robertsonianus. Jones (1994) 

lists C. bradyi as Gyroidina bradyi, as Gyroidina differs from Cibicidoides in possessing an 

aperture more or less restricted to the umbilical side rather than in an equatorial position. The 

extent of the apertural slit is somewhat more variable in our specimens, but we follow Jones in 

placing the species under Gyroidina. 

Figures: 

Plate 71, figs. 1a-c, Green Canyon (Block 272) Core P-57 Sample 1-2 cm, 1a: ventral view, 1b: 

dorsal view, 1c: edge view; fig. 2, Alaminos Canyon Core PC4 Sample 1-2 cm, ventral view; 

figs. 3a-b, Alaminos Canyon Core PC4 Sample 0-1 cm, 3a: ventral view, 3b: edge view; figs. 4ab, Green Canyon (Block 272) Core P-33 Sample 0-1 cm, 4a: ventral view, 4b: dorsal view; fig. 5, 

Green Canyon (Block 272) Core P-57 Sample 1-2 cm, ventral view. 

Scale bars = 50 μm. 

Gyroidina orbicularis (sensu Parker, Jones, and Brady) 

Rosalina orbicularis Parker, Jones, and Brady, 1865. 

Rotalia orbicularis d’Orbigny – Brady, 1884, p. 706, pl. 107, figs. 5a-c, pl. 115, figs. 6a-c. 

Rotalia orbicularis d’Orbigny – Cushman, 1915, p. 68, 69, pl. 29, figs. 3a-c, text-figs. 62a-c. 

Gyroidina orbicularis d’Orbigny – Phleger and Parker, 1951, p. 22, pl. 11, figs. 11a, 11b, 12a, 

12b. 

Gyroidina orbicularis d’Orbigny – Parker, 1954, p. 528, pl. 9, figs. 13, 18. 

Gyroidina orbicularis d’Orbigny – Pflum and Frerichs, 1976, pl. 5, figs. 5-7. 

Gyroidinoides orbicularis d’Orbigny – Hermelin, 1989, p. 83, 84, pl. 16, figs. 7-9. 

Gyroidinoides orbicularis d’Orbigny – Denne, 1990, pl. 7, figs. 4a, 4b. 

Gyroidina orbicularis (sensu Parker, Jones, and Brady) – Jones, 1994. 

Morphology: Shell unequally biconvex, ventral side is strongly convex, dorsal side is slightly 

convex, sutures radial and flush with the surface on dorsal side, spiral suture is slightly 

channeled, last few sutures on ventral side depressed and slightly recurved, aperture 

interiomarginal slit extending almost from the periphery to the umbilicus with a lip. 

Remarks: This is a widely distributed species with a long history of nomenclatural confusion (see 

Jones, 1994), primarily because the features of d’Orbigny’s original Gyroidina orbicularis are 

unknown. The name, with d’Orbigny as the author, has been used by numerous workers, but they 

have actually followed the concept of Parker et al. (1865) and Brady (1884) regarding the 

diagnostic characters of the species. Pending a thorough study of its comparative morphology, 

and possibly the designation of a new name, we follow the safe procedure of Jones (1994) in 

calling the species "Gyroidina orbicularis (sensu Parker, Jones, and Brady)". 

109
Figures: 

Plate 72, figs. 1-2, Alaminos Canyon Core PC5 Sample 2-3 cm, a: dorsal view, b: ventral view, 

c: edge view showing aperture. 

Scale bars = 50 μm. 

2.2.10. Order Robertinida 

Superfamily Ceratobuliminacea 

Family Epistominidae 

Genus Hoeglundina 

Hoeglundina elegans (d’Orbigny) 

Rotalina (Turbinulina) elegans d’Orbigny, 1826, p. 276, (not Rotalia elegans, Ibid., p. 272, no. 

6). 

Pulvinulina partschiana (d’Orbigny) – Brady, 1884, p. 699, pl. 105, figs. 3a-c. 

Pulvinulina elegans (d’Orbigny) – Brady, 1884, p. 699, pl. 105, figs. 4-6. 

Epistomina elegans (d’Orbigny) – Cushman, 1931, p. 65, 66, pl. 13, figs. 6a-c. 

Höglundina elegans (d’Orbigny) – Phleger and Parker, 1951, p. 22, pl. 12, figs. 1a, b.

Höglundina elegans (d’Orbigny) – Parker, 1954, p. 531, pl. 10, figs. 4, 8. 

Hoglündina elegans (d’Orbigny) – Barker, 1960, p. 216, pl. 105, figs. 3-6. 

Epistomina elegans (d’Orbigny) – Andersen, 1961, p. 112, pl. 28, figs. 4a-c. 

Hoeglundina elegans (d’Orbigny) – Poag, 1981, p. 69, pl. 19, fig. 3; pl. 20, figs. 3a, b, c. 

Hoeglundina elegans (d’Orbigny) – van Morkhoven, Berggren and Edwards, 1986, p. 97, 98, pl. 

29, figs. 1a, b, 2a, b. 

Hoeglundina elegans (d’Orbigny) – Denne, 1990, pl. 7, figs. 7a, b. 

Hoeglundina elegans (d’Orbigny) – Jones, 1994, p. 104, 105, pl. 105, figs. 3-6. 

Morphology: Shell biconvex; typically 7-9 chambers in final whorl; sutures limbate, flush with 

surface; aperture an arched slit that extends the breadth of the chamber within and parallel to the 

periphery on ventral side; apertures of earlier chambers may remain open or be secondarily 

closed and visible as scars along the periphery. 

Remarks: This is the type species of the genus Hoeglundina Brotzen 1948. 

Figures: 

Plate 77, figs. 1a-c, Green Canyon (Block 180) Core P-EP15 Sample 0-1 cm, 1a: ventral view, 

1b: dorsal view, 1c: side view showing aperture; figs. 2a-b, Green Canyon (Block 180) Core PEP15 Sample 0-1 cm, 2a: ventral view, 2b: dorsal view; fig. 3, Alaminos Canyon Core PC15 

Sample 6-7 cm, ventral view. 

Scale bars = 50 μm. 

110
Superfamily Robertinacea 

Family Robertinidae 

Genus Robertina 

Robertina subcylindrica (Brady)

Bulimina subcylindrica Brady, 1881, p. 56. 

Bulimina subcylindrica Brady – Brady, 1884, p. 404, pl. 50, figs. 16a, b. 

Robertina subcylindrica (Brady) – Cushman and Parker, 1936, p. 95, 96, pl. 16, figs. 10a, b. 

Robertina subcylindrica (Brady) – Barker, 1960, p. 102, pl. 50, figs. 16a, b. 

Robertina subcylindrica (Brady) – Jones, 1994, p. 55, pl. 50, figs. 16a, b. 

Morphology: Shell is subcylindrical and elongate in shape; 4 pairs of chambers in final whorl; 

aperture is slitlike from the base of the final chamber. 

Figures: 

Plate 156, fig. 1, Alaminos Canyon Core PC4 Sample 0-1 cm, front view. 

Scale bar = 50 μm. 

Genus Robertinoides 

Robertinoides bradyi (Cushman and Parker) 

Bulimina subteres Brady, 1881, p. 55. 

Bulimina subteres Brady – Brady, 1884, p. 404, pl. 50, figs. 18a, b. 

Robertina bradyi Cushman and Parker, 1936, p. 99, 100, pl. 16, figs. 9a, b. 

Robertinoides bradyi (Cushman and Parker) – Barker, 1960, p. 102, pl. 50, figs. 18a, b. 

Robertinoides bradyi (Cushman and Parker) – Andersen, 1961, p. 86, pl. 19, fig. 4. 

Robertinoides bradyi (Cushman and Parker) – Denne, 1990, pl. 11, fig. 21.

Robertinoides bradyi (Cushman and Parker) – Jones, 1994, p. 55, pl. 50, figs. 18a, b. 

Morphology: Test tapering towards initial end; 4-5 pairs of chambers in final whorl; sutures 

slightly depressed in most specimens; aperture elongate from the base of the final chamber; 

supplementary aperture present, elongate and short. 

Figures: 

Plate 157, figs. 1-2, Alaminos Canyon Boxcore B6 bulk sample, front view; fig. 3, Alaminos 

Canyon Core Sample 0-1 cm, front view. 

Scale bars = 50 μm.

111

3. FORAMINIFERA IN SEDIMENT SAMPLES 

3.1. Identified Species 

A total of 177 species––60 agglutinated, 13 porcelaneous, and 104 hyaline––have been identified 

from the sediment samples. In addition, 8 species were found only on tubeworm surfaces. Table 

3-1 lists the species alphabetically within each of the three primary morphological groups. The 

species indicated with superscripts 1 and 2 were utilized in the cluster analyses. All species in 

our samples, except Heronallenia sp., Miliolinella subrotunda (Seguenza), Neoconorbina 

terquemi (Rzehak), Procerolagena gracillima (Seguenza), and Trochamminoides coronatus

(Brady) are illustrated in the plates of the Atlas (Appendix) and discussed in the systematics 

section (Chapter 2). The excluded species are rare; we could not find specimens that were 

suitable for illustrations. 

Twenty-one modern species found in the course of this study were previously unreported from 

the Gulf of Mexico. Two of these, Deuterammina rotaliformis and Discanomalina semipunctata, 

were found only on tubeworms; D. semipunctata has been previously reported from Pleistocene 

sediments of the Gulf. The rest were found only in the sediment samples; these are: 

Ammobaculites catenulatus, Barbourinella atlantica, Cribromiliolinella subvalvularis, Fissurina 

fissa, Glomospira irregularis, Lagena chrysalis, Lagenammina laguncula, L. tubulata, 

Lenticulina pliocaena, Nonionella iridea, Orectostomina camposi, Parafissurina kerguelenensis, 

Parvigenerina arenacea, Planispirinoides bucculentus var. placentiformis, Portatrochammina 

antarctica, Pseudoclavulina serventyi, Robertina subcylindrica, Saccammina helenae, and 

Saracenaria altifrons. 

3.2. Rose-Bengal-Stained Foraminifera 

Approximate estimates of living populations were obtained by counting Rose-Bengal-stained 

specimens. Although widely used, the reliability of Rose Bengal stain for distinguishing between 

living and dead foraminiferal specimens is in some doubt (Bernhard and Sen Gupta, 1999; 

Bernhard et al., 2006). It has been shown that less than half of the Rose-Bengal-stained 

Foraminifera in certain samples from bathyal marine sediments may be living when collected 

(Bernhard et al., 2006). The main criticism of Rose Bengal is that it is a non-vital stain and will 

adhere to dead as well as living cytoplasm (Bernhard, 1988, 1989; Corliss and Emerson, 1990; 

Jorissen et al., 1995), and therefore will mark recently dead specimens in addition to living 

specimens. Thus, dependence on Rose Bengal leads to overestimation of the abundance of living 

hyaline Foraminifera. In contrast, because the stain can be difficult to see in opaque specimens 

(Bernhard, 2000), the number of living agglutinated and miliolid Foraminifera may be 

underestimated. Despite such problems, however, Rose Bengal has been used in numerous 

studies to recognize Foraminifera collected alive (e.g., Corliss, 1985; Lutze and Altenbach, 1991; 

Gooday, 1993, 1996; Gooday and Rathburn, 1999; Rathburn et al., 2000; Rathburn et al., 2003). 

The consensus is that a conservative use, with only deeply stained foraminiferal shells, should 

yield a reasonable estimate of living assemblages (Lutze and Altenbach, 1991; Gooday and 

Rathburn, 1999; Rathburn et al., 2003), especially for comparative studies of seep and non-seep 

assemblages. 

113
Table 3-1 

Geographic Occurrence of Species 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

AGGLUTINATED SPECIES 

Adercotryma glomeratum X X X X X 

Ammobaculites catenulatus X 

Ammobaculites filiformis X X X 

Ammodiscus tenuis X X X X X 

Ammolagena clavata X X X X X X X 

Ammoscalaria tenuimargo T 

Archimerismus subnodosus X X X X X X 

Barbourinella atlantica X 

Buzasina ringens X X 

Cribrostomoides subglobosus X 

Cystammina pauciloculata X X X 

Deuterammina rotaliformis T T 

Discammina compressa X X X X 

Dorothia scabra X X 

Eggerella bradyi X X X X X X X 

Gaudryina minuta X X X X X X X 

Glomospira gordialis X X 

Glomospira irregularis X X X X X 

Haplophragmoides sp. cf. H. kirki X X X X 

Hormosina pilulifera X X X X 

Hormosinella distans X X X X 

Hormosinella guttifera X X X X 

Hyperammina friabilis X X 

Hyperammina laevigata X X X X X 

Hyperammina sp. X X X X X X 

Karreriella bradyi X X X X 

Karrerulina apicularis X X X X 

Lagenammina difflugiformis X X X X X X 

Lagenammina laguncula X 

Lagenammina tubulata X X X X X 

Lituotuba lituiformis X X X 

Martinottiella communis X X X X X 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

114
Table 3-1. Geographic Occurrence of Species (continued). 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

AGGLUTINATED SPECIES 

Multifidella nodulosa X 

Orectostomina camposi X 

Paratrochammina challengeri X X 

Parvigenerina arenacea X X 

Portatrochammina antarctica X X X X X X X 

Prolixoplecta parvula X X X X X 

Psammosphaera fusca X X 

Pseudoclavulina mexicana X 

Pseudoclavulina serventyi X 

Pseudogaudryina atlantica X 

Pseudotrochammina sp. X X 

Reophax agglutinatus X X 

Reophax scorpiurus X X X X 

Rhabdammina abyssorum X 

Rhabdammina discreta X X 

Rhizammina algaeformis X X X 

Rhizammina indivisa X X X X X 

Saccammina helenae X X 

Saccorhiza ramosa X X X X X X X X 

Siphotextularia rolshauseni X X X 

Spiroplectammina sp. X X X X 

Subreophax monile X X 

Tetrataxiella sp. X 

Textularia foliacea occidentalis X 

Textularia mexicana X 

Textularia sp. cf. T. mayori X 

Textulariella barrettii X 

Trochamminoides coronatus X X X X X 

Usbekistania charoides X X X X X X X X 

Veleroninoides jeffreysii T 

Veleroninoides wiesneri X X X X X X 

PORCELANEOUS SPECIES 

Calcituba polymorpha T 

Cornuspira foliacea T 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

115
Table 3-1. Geographic Occurrence of Species (continued). 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

PORCELANEOUS SPECIES 

Cornuspira involvens1, 2 X X X 

Cribromiliolinella subvalvularis 1, 2 X X X 

Miliolinella subrotunda1, 2 X 

Miliolinella warreni X 

Miliolinella sp. T T 

Planispirinoides bucculentus var. placentiformis1, 2 X 

Pyrgo lucernula X X X X 

Pyrgo murrhina X X X X 

Pyrgo nasuta X 

?Pyrgo sp. X 

Quinqueloculina bosciana X 

Sigmoilinita elliptica1, 2 X X X X 

Sigmoilopsis schlumbergeri1, 2 X X X X 

Triloculina tricarinata X X 

HYALINE SPECIES 

Abditodentrix pseudothalmanni1, 2 X X X X X 

Amphicoryna hirsuta X X X 

Anomalinoides globulosus T X 

Anomalinoides mexicanus X X 

Bolivina alata X X X 

Bolivina albatrossi1, 2 X X X X X X X X 

Bolivina barbata1, 2 X X 

Bolivina daggarius X 

Bolivina goesii X 

Bolivina sp. cf. B. hastata X X 

Bolivina lowmani1, 2 X X X X X X X X 

Bolivina minima2 X X X 

Bolivina ordinaria1, 2 X X X X X X 

Bolivina sp. cf. B. pusilla X X 

Bolivina subaenariensis1, 2 X X X 

Bolivina translucens1, 2 X X X X X X 

Bolivinita quadrilatera X 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

116
Table 3-1. Geographic Occurrence of Species (continued). 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

HYALINE SPECIES 

Bulimina aculeata1, 2 X X X X X X X 

Bulimina alazanensis1, 2 X X X X X 

Bulimina marginata1, 2 X X X 

Bulimina mexicana1, 2 X X X X X X X X 

Cassidulina carinata1, 2 X X X X X X X 

Cassidulina curvata1, 2 X X X X 

Cassidulina obtusa1, 2 X X X X X X X X 

Cassidulinoides tenuis X X X 

Chilostomella oolina X X X X 

Cibicides lobatulus X 

Cibicides wuellerstorfi1, 2 X X X 

Cibicidoides pachyderma1, 2 X X X X X X X 

Discanomalina semipunctata T 

Cibicidoides robertsonianus X X X X X 

Dentalina sp. X 

Epistominella exigua1, 2 X X X X X X X X 

Epistominella vitrea1, 2 X X X X X 

Eponides turgidus1, 2 X X X X X X X X 

Eubuliminella morgani X X X 

Fissurina fissa X 

Fissurina sp. cf. F. incomposita X X 

Fursenkoina seminuda1, 2 X X X X X 

Gavelinopsis tranlucens1, 2 X X X X X 

Globocassidulina subglobosa1, 2 X X X X X X X X 

Gyroidina bradyi1, 2 X X X X X X X X 

Gyroidina orbicularis (sensu lato)1, 2 X X X X X X X X 

Gyroidinoides sp. cf. G. polius1, 2 X X X X X X 

Gyroidinoides sp. cf. G regularis1, 2 X X X X X X X 

Haynesina germanica X X 

Heronallenia sp. X 

Hoeglundina elegans1, 2 X X X X X X 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

117
Table 3-1. Geographic Occurrence of Species (continued). 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

HYALINE SPECIES 

Ioanella tumidula1, 2 X X X X 

Lagena chrysalis X 

Lagena hispida X 

Lagena hispidula X X X 

Laminononion tumidum X T 

Laticarinina pauperata X X X X X X 

Lenticulina calcar1, 2 X X X 

Lenticulina gibba X X 

Lenticulina pliocaena X X X X 

Lenticulina thalmanni X X X 

Melonis affinis X X 

Melonis pompilioides X X 

Neoconorbina terquemi X X X X 

Neocrosbyia minuta X X X X 

Nonionella iridea1, 2 X X X X X 

Nuttallides decorata1, 2 X X X 

Oolina apiopleura X 

Oolina globosa1, 2 X X X 

Oolina ovum X X X X 

Oolina squamosa X 

Oridorsalis umbonatus1, 2 X X X X X X X X 

Osangularia culter1, 2 X X X X X X 

Osangularia rugosa1, 2 X X X X X X X X 

Parafissurina botelliformis X X X X 

Parafissurina kerguelenensis X X 

Parafissurina lateralis X X 

Patellina corrugata1, 2 X X X X 

Planulina ariminensis X X 

Planulina foveolata X 

Plectofrondicularia advena X X 

Praeglobobulimina ovata1, 2 X X X X X 

Praeglobobulimina ovula X X 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

118
Table 3-1. Geographic Occurrence of Species (continued). 

SPECIES AREA 

 1 2 3 4 5 6 7 8 

HYALINE SPECIES 

Procerolagena gracilis1, 2 X X X 

Procerolagena gracillima X X X 

Pseudoglandulina comatula X 

Pullenia bulloides X X X X X 

Pullenia quinqueloba1, 2 X X X X X X X 

Robertina subcylindrica X 

Robertinoides bradyi X X X 

Rutherfordoides mexicanus1, 2 X X X X X X 

Saracenaria altifrons X 

Seabrookia earlandi1,2 X X X X X X X 

Siphonina pulchra1, 2 X X X 

Siphonodosaria calomorpha X X X X 

Spirillina vivipara T 

Stainforthia complanata X X X X X X X 

Stainforthia compressa X X 

Stainforthia pontoni X X 

Trifarina bradyi1, 2 X X X X 

Trifarina jamaicensis X X 

Uvigerina auberiana X X X X 

Uvigerina bellula X 

Uvigerina flintii1, 2 X 

Uvigerina peregrina1, 2 X X X X X X X X 

Vaginulinopsis subaculeata X X X 

Valvulineria glabra X X X X X X 

Valvulineria mexicana1, 2 X X X X 

Vasiglobulina reticulata X X 

Areas: 1, Shallow Green Canyon (245 m); 2, Deep Green Canyon (562-696 m); 3, Garden Banks (640 m); 4, South of Mississippi 

Canyon (~1000 m); 5, Mississippi Canyon (620 m, 1067-1081 m); 6, De Soto Canyon (1850 m); 7, Alaminos Canyon (2218-2227 

m); 8, Farnella Canyon (2918 m).

T = found exclusively on tubeworms (and not in sediments).

Superscripts 1 and 2 indicate species used in cluster analyses 1 and 2, respectively.

119
About 80 species in our samples caught the Rose Bengal stain. Of these, only 32 species are 

represented by more than one or two Rose-Bengal-stained specimens in more than two samples, 

and 8 such species are present in more than 2% of the samples. Also, only three stained species 

constitute >2% of the calcareous assemblage (Table 3-2). Several of these species are important 

at multiple sites. These include Cornuspira involvens, Eponides turgidus, Nuttallides decorata, 

Saccorhiza ramosa, Hyperammina sp., and Portatrochammina antarctica. The Rose-Bengalstained Eponides turgidus and Nuttallides decorata individuals are mostly confined to non-seep 

samples. Stained Cornuspira involvens specimens are restricted to bacterial mat seep samples 

and are most abundant in samples ACP15a and ACP15b (Table 3-2). Stained specimens of 

Portatrochammina antarctica are also abundant in ACP15a (23 individuals), comprising 9% of 

the total assemblage. Small numbers of Rose-Bengal-stained Portatrochammina antarctica are 

found in several other samples, mostly from bacterial mat substrates. Rose-Bengal-stained 

specimens of Saccorhiza ramosa are limited to non-seep samples, while stained Hyperammina

sp. specimens are found in both seep and non-seep samples. Of the 27 benthic Foraminifera 

individuals present in ACP10a, 19 are Rose-Bengal-stained Hyperammina sp. specimens. 

The percentage of Rose-Bengal-stained Foraminifera in most bacterial mat samples is less than 

5%, with some notable exceptions. In sample ACP15a (orange bacterial mat), 83% of the total 

specimens were stained, with encysted Cornuspira involvens (the sole calcareous species in the 

sample, 183 specimens) constituting 71%. The percentage of stained individuals (all C. 

involvens) decreased in ACP15b to 24% of the total and 32% of the calcareous assemblage. The 

other two samples with anomalously high numbers of Rose-Bengal-stained individuals include 

ACP10a and GCP54a. Sample ACP10a has 19 stained Hyperammina sp. specimens that 

comprise 70% of the total Foraminifera. Twenty-two percent of the calcareous Foraminifera in 

GCP54a caught the stain; they comprise 83% of all stained individuals in the sample. In all, 13% 

of all Foraminifera in GCP54a were stained by Rose Bengal. These stained individuals belong 

mainly to Cribromiliolinella subvalvularis, Bulimina marginata, Portatrochammina antarctica, 

Cornuspira involvens, and Planispirinoides bucculentus var. placentiformis.

3.3. Species Richness 

The diversity of benthic Foraminifera, as represented by species richness (number of species in a 

sample), is high in the majority of samples analyzed in this study. Much variablity in species 

richness is observed in Green Canyon, Mississippi Canyon, and Alaminos Canyon samples 

(Table 3-3, Fig. 3.1). GCP15a and GCP15b, the two samples from shallow Green Canyon, have 

the highest species richness of 133 and 152, respectively. Samples from deep Green Canyon and 

Garden Banks show species richness values between 46 and 102. In the deeper locations of 

Mississippi Canyon and Alaminos Canyon, however, several samples have lower values, with 

ranges of 20-74 for Mississippi Canyon and 0-114 for Alaminos Canyon; species richness seems 

to be more erratic at these deeper sites. The samples with the lowest diversities at any site are the 

those from bacterial mats, but there are also four non-mat samples in Alaminos Canyon with 

very low diversiy (Fig. 3.1). Those four samples are probably from sites of active seepage (see 

later). 

120
Table 3-2 

Counts of Rose-Bengal-Stained Individuals for the 32 Species That Contribute More Than One 

or Two Stained Individuals at More Than One Site 

Species in which Rose-Bengal-stained individuals represent greater than 2% of the assemblage at a site are in bold; 

species for which Rose-Bengal-stained individuals represent greater than 2% of the calcareous assemblage are in 

underlined bold. Total counts for all specimens (stained and unstained) are given in the last two rows. 

SPECIES AC AC AC AC AC AC AC AC AC AC 

P15a P15b P04a P05a P05b P10a P11a P12b B07a B08a

Bolivina albatrossi 

Bolivina ordinaria 

Bulimina aculeata 

Bulimina marginata 

Cassidulina carinata 

Globocassidulina subglobosa 

Hoeglundina elegans 1

Cibicides wuellerstorfi 1 2 

Epistominella exigua 

Eponides turgidus 15 4 

Gavelinopsis translucens 

Nuttallides decorata 14 8 1

Oridorsalis umbonatus 1 2 

Osangularia culter 

Osangularia rugosa 

Cornuspira involvens 183 14 2 

Cribromiliolinella subvalvularis 

Planispirinoides bucculentus var. 

placentiformis 

 

Hyperammina sp. 2 19 

Lagenammina difflugiformis 

Lagenammina tubulata 

Rhizammina algaeformis 4 

Saccorhiza ramosa 8 

Rhizammina indivisa 

Adercotryma glomerata 

Ammodiscus tenuis 4 1 

Gaudryina minuta 

Reophax scorpiurus 

Usbekistania charoides 

Portatrochammina antarctica 23 1 1 3 2 

Trochamminoides coronatus 1 

Unidentified trochamminids 3 4 

Total calcareous individuals 183 44 1290 261 1056 2 5 0 26 2

Total individuals 257 59 1324 515 2450 27 8 11 456 291

121
Table 3-2. Counts of Rose-Bengal-Stained Individuals for the 32 Species That Contribute More 

Than One or Two Stained Individuals at More Than One Site (continued). 

Species in which Rose-Bengal-stained individuals represent greater than 2% of the assemblage at a site are in bold; 

species for which Rose-Bengal-stained individuals represent greater than 2% of the calcareous assemblage are in 

underlined bold. Total counts for all specimens (stained and unstained) are given in the last two rows. 

SPECIES AC AC DC MC MC MC MC MC MC FC 

B09a B19a B-S36 P02a P02b P04b P08a P08b B-C7 P61a 

Bolivina albatrossi 3 

Bolivina ordinaria 4 

Bulimina aculeata 1 1 

Bulimina marginata 

Cassidulina carinata 1 2 1 

Globocassidulina subglobosa 2 

Hoeglundina elegans 3 1 

Cibicides wuellerstorfi 

Epistominella exigua 2 3 1 3 2 2 

Eponides turgidus 42 5 2 3 

Gavelinopsis translucens 1 

Nuttallides decorata 33 1 

Oridorsalis umbonatus 1 1 

Osangularia culter 3 

Osangularia rugosa 1 1 

Cornuspira involvens 1 

Cribromiliolinella 

subvalvularis

 2 

Planispirinoides bucculentus 

var. placentiformis 

 

Hyperammina sp. 1 

Lagenammina difflugiformis 1 4 

Lagenammina tubulata 4 

Rhizammina algaeformis 2 

Saccorhiza ramosa 47 13 4 6 

Rhizammina indivisa 5 

Adercotryma glomerata 6 

Ammodiscus tenuis 

Gaudryina minuta 3 1 

Reophax scorpiurus 1 3 

Usbekistania charoides 4 

Portatrochammina antarctica 4 1 1 2 

Trochamminoides coronatus 1 4 

Unidentified trochamminids 2 6 1 

Total calcareous individuals 4 932 2246 340 410 106 78 243 444 438 

Total individuals 631 1642 1070 465 460 314 186 4721 962 558 

122
123

Table 3-2. Counts of Rose-Bengal-Stained Individuals for the 32 Species That Contribute More 

Than One or Two Stained Individuals at More Than One Site (continued). 

Species in which Rose-Bengal-stained individuals represent greater than 2% of the assemblage at a site are in bold; 

species for which Rose-Bengal-stained individuals represent greater than 2% of the calcareous assemblage are in 

underlined bold. Total counts for all specimens (stained and unstained) are given in the last two rows. 

SPECIES GC GC GC GC GC GC GC GC GC 

P33a P33b P51a P54a P54b P55a P57a P57b P12a 

Bolivina albatrossi 2 2 

Bolivina ordinaria 2 

Bulimina aculeata 3 

Bulimina marginata 7 

Cassidulina carinata 1 

Globocassidulina subglobosa 1 1 

Hoeglundina elegans 1 1 

Cibicides wuellerstorfi 

Epistominella exigua 1 1 3 

Eponides turgidus 1 5 2 

Gavelinopsis translucens 1 1 1 

Nuttallides decorata 

Oridorsalis umbonatus 

Osangularia culter 

Osangularia rugosa 1 

Cornuspira involvens 5 

Cribromiliolinella subvalvularis 1 10 10 

Planispirinoides bucculentus var. 

placentiformis 

 3 

Hyperammina sp. 3 3 1 

Lagenammina difflugiformis 

Lagenammina tubulata 1 

Rhizammina algaeformis 

Saccorhiza ramosa 

Rhizammina indivisa 

Adercotryma glomerata 

Ammodiscus tenuis 

Gaudryina minuta 

Reophax scorpiurus 1 

Usbekistania charoides 

Portatrochammina antarctica 2 6 1 

Trochamminoides coronatus 

Unidentified trochamminids 

Total calcareous individuals 180 199 237 139 110 284 501 410 293 

Total individuals 458 434 288 275 288 400 623 520 298 
Table 3-3 

Species richness (S) and Assemblage Density (AD) in Sediment Samples 

LOCATION SAMPLES DEPTH (m) S (calc) S (agg) S (total) AD 

Green Canyon (shallow) GCP15a 

GCP15b 

245 84 

112 

49 

40 

133 

152 

86 

382 

Garden Banks GBP08a 

GBP08b 

640 46 

67 

21 

18 

67 

85 

66 

83 

Green Canyon (deep) GCP12a 

GCP12b 

GCP33a 

GCP33b 

GCP51a 

GCP51b 

GCP54a 

GCP54b 

GCP55a 

GCP55b 

GCP57a 

GCP57b 

569 

664 

696 

675 

689 

675 

44 

45 

46 

53 

52 

50 

37 

27 

63 

60 

68 

60 

4 

5 

17 

14 

11 

9 

15 

19 

30 

30 

34 

38 

48 

50 

63 

67 

63 

59 

52 

46 

93 

90 

102 

98 

8 

13 

48 

23 

8 

7 

7 

8 

166 

185 

65 

27 

Mississippi Canyon MCP04a 

MCP04b 

MCP08a 

MCP08b 

MCP73a 

MCP73b 

MCP02a 

MCP02b 

MCP69a 

MCP69b 

1072 

1071 

1070 

1067 

1081 

22 

37 

18 

20 

18 

16 

48 

53 

38 

44 

5 

9 

2 

1 

6 

4 

18 

16 

21 

30 

27 

46 

20 

21 

24 

20 

66 

69 

59 

74 

2 

8 

5 

6 

30 

40 

48 

48 

46 

84 

South of Mississippi Canyon MCB-C7 1076 57 33 90 174 

De Soto Canyon DCB-S36 1848 64 50 114 102 

Alaminos Canyon ACP01a 

ACP01b 

ACP04a 

ACP04b 

ACP13a 

ACP13b 

ACB07a 

ACB08a 

ACB09a 

ACP10a 

ACP10b 

ACP11a 

ACP11b 

ACP12a 

ACP12b 

ACP15b 

ACP15a 

ACP05a 

ACP05b 

ACB19a 

2226 

2227 

2221 

2227 

2219 

2221 

2225 

2219 

8 

26 

72 

47 

16 

32 

9 

2 

4 

2 

1 

3 

0 

0 

0 

6 

1 

30 

73 

55 

0 

0 

6 

5 

1 

0 

8 

13 

7 

4 

0 

2 

0 

1 

3 

4 

9 

30 

41 

32 

8 

26 

78 

52 

17 

32 

17 

15 

11 

6 

1 

5 

0 

1 

3 

10 

10 

60 

114 

87 

0.2 

3 

34 

43 

1 

5 

9 

12 

52 

0.7 

0.03 

0.2 

0 

0.02 

0.6 

2 

7 

13 

64 

134 

Farnella Canyon FCP61a 2918 53 21 74 29 

S (calc) = number of calcareous species; S (agg) = number of agglutinated species; S (total) = total number of species; 

AD = number of individuals/cm3

. 

124
Figure 3.1. Species richness (number of species) in surface and near-surface samples plotted 

against water depths. Open squares, samples from bacterial mat substrates; closed 

circles, others. 

125
High species richness values have been observed previously from non-seep sites in the 

northwestern Gulf of Mexico (Gibson and Buzas, 1973; Denne, 1990; Jones and Sen Gupta, 

1995). Gibson and Buzas (1973), using data from Phleger (1951), noted species richness values 

of 10-50 at depths comparable to those of our samples, and Denne (1990) recorded species 

richness values of 38-51. Jones and Sen Gupta (1995) observed species richness values of 20-60 

at depths of 1500-2500 m. None of these studies recorded the high species richness values seen 

in our study (Table 3-3). Judging by our observation on species present on tubeworms and in 

sediments at the base of tubeworm bushes, it is likely that the species richness in some of our 

sediment samples is elevated by post-mortem mixing of foraminiferal shells. This problem is 

discussed more fully in Chapter 4. 

3.4. Assemblage Density 

Assemblage densities (Table 3-3, Fig. 3.2) were measured by calculating the number of 

individuals per cm3 of the surface or near-surface sediment. There is a slight trend towards 

decreasing densities with increasing depths. Six samples, none from a bacterial mat substrate, 

show assemblage densities greater than 100. The highest density (382) is observed in sample 

GCP15b from shallow Green Canyon. 

3.5. Cluster Analyses Results 

 

Surface samples. The Q-mode CA dendrogram for surface sample data for calcareous species 

(Fig. 3.3) shows two major clusters, A and B, and one outlier sample (ACP15). The two clusters 

are bathymetrically distinct. Cluster A includes deepest-bathyal samples from Alaminos Canyon 

(2219-2227 m water depth), Farnella Canyon (2918 m), and De Soto Canyon (1848 m). Cluster 

B includes shallower bathyal samples from Green Canyon (245 m, 569 m, 664-696 m), Garden 

Banks (640 m), Mississippi Canyon (1067-1081 m), and a location south of Mississippi Canyon 

(1076 m). Several minor clusters (A1, B1, etc.) can be recognized within these two large clusters. 

Table 3-4 lists the samples and the species with 5% or greater mean abundance in these clusters; 

Figure 3.4 also shows the relative abundances of the more important species in the minor 

clusters. 

Surface and near-surface samples. In the combined data set for calcareous species in surface 

and near-surface samples, the 0-1 cm samples are indicated by the suffix "a" (e.g., ACP05a) and 

the 1-2 cm samples by the suffix "b" (e.g., ACP05b). Here too, two major clusters (X and Y) are 

recognized (Fig. 3.5), although the number of samples and that of minor clusters (X1, Y1, etc.) 

are larger. Table 3-5 lists the samples and the species with 5% or greater mean abundance in 

these clusters; Fig. 3.6 shows the relative abundances of the more important species in the minor 

clusters.

Cluster X, as with cluster A, consists of samples from Alaminos Canyon (2219-2227 m) and two 

other samples, DCB-S36 from De Soto Canyon (1848 m) and FCP61 from Farnella Canyon 

(2918 m). Samples in cluster Y are from Mississippi Canyon (1067-1081 m), south of 

Mississippi Canyon (1076 m), Garden Banks (640 m), and Green Canyon (245 m, 568-696 m). 

All near-surface samples belong to the same minor clusters as their matching surface samples. 

126
Figure 3.2. Assemblage density (individuals/cm3

) in surface and near-surface samples plotted 

against water depths. Open squares, samples from bacterial mat substrates; closed 

circles, others. 

127
Figure 3.3. Q-mode clusters, surface samples. 

128
Figure 3.4. Relative abundances of the more important species in minor clusters, surface 

samples. 

129
Figure 3.5. Q-mode clusters, surface (0-1 cm) and near-surface (1-2 cm) samples. Surface 

samples are indicated by suffix a, near-surface samples by b. 

130
Table 3-4 

Q-Mode Minor Clusters for Surface-Sample Data: 

Constituent Samples, Water Depths, Substrates, 

and Important Foraminiferal Species (Mean Abundance >5% in Minor Cluster) 

MINOR 

CLUSTER 

SAMPLES DEPTH 

(m) 

SUBSTRATE IMPORTANT SPECIES % 

A1 ACP04a 

ACP05a 

ACB19a 

DCB-S36 

2227 

2225 

2219 

1848 

White mud mound 

Mud 

Non-seep gray mud 

Non-seep 

Eponides turgidus 

Nuttallides decorata 

Bolivina lowmani

33 

26 

16 

A2 ACB07a 

ACP13a 

FCP61a 

2227 

2221 

2918 

Gray bacterial mat 

White bacterial mat 

Tubeworm clump 

Nuttallides decorata 

Cibicides wuellerstorfi 

Praeglobobulimina ovata 

Hoeglundina elegans 

Eponides turgidus 

Cornuspira involvens 

22 

14 

9 

8 

7 

5 

B1 MCP08a 

MCP73a 

GCP12a 

GCP33a 

GCP51a 

1071 

1070 

569 

664 

696 

Gray bacterial mat 

Bacterial mat 

White bacterial mat 

White bacterial mat 

Bacterial mat on mound 

with clams 

Bolivina ordinaria 

Cassidulina carinata 

Gavelinopsis translucens 

Epistominella exigua 

Bolivina albatrossi 

19 

12 

10 

6 

5 

B2.1 MCP02a 

MCP69a 

MCB-C7 

1067 

1081 

1076 

Mud from ridge top 

Bacterial mat 

Non-seep 

Eponides turgidus 

Nonionella iridea 

Bolivina lowmani 

Bulimina aculeata 

Uvigerina peregrina 

Epistominella exigua 

Bolivina albatrossi 

14 

10 

9 

7 

7 

6 

5 

B2.2 MCP04a 

GBP08a 

GCP15a 

GCP55a 

GCP57a 

1072 

640 

245 

689 

675 

Gray bacterial mat 

Non-seep gray mud 

Non-seep gray mud 

Non-seep mud 

Sand 

Bolivina ordinaria 

Bolivina lowmani 

Globocassidulina subglobosa 

Epistominella exigua 

Bolivina albatrossi 

Cassidulina carinata 

Uvigerina peregrina 

9 

7 

6 

6 

5 

5 

4 

B3 GCP54a 675 White bacterial mat Cribromiliolinella subvalvularis

Miliolinella subrotunda 

Bulimina marginata 

Osangularia rugosa 

Cassidulina carinata 

Bolivina ordinaria 

Cornuspira involvens 

24 

8 

8 

7 

6 

6 

4 

C 

(outlier) 

ACP15a 2221 Orange bacterial mat Cornuspira involvens 100 

131
Table 3-5 

Q-Mode Minor Clusters for Combined Surface- and Near-Surface-Sample Data: 

Constituent Samples, Water Depths, Substrates, 

and Important Foraminiferal Species (Mean Abundance >5% in Minor Cluster) 

MINOR 

CLUSTER 

SAMPLES DEPTH 

(m) 

SUBSTRATE IMPORTANT SPECIES % 

X1.1 ACP05a 

ACP05b 

ACB19a 

DCB-S36 

2225 

2225 

2219 

1848 

Mud 

Mud 

Non-seep gray mud 

Non-seep 

Eponides turgidus 

Nuttallides decorata 

Bolivina lowmani

31 

23 

21 

X1.2 ACP01b 

ACP04a 

ACP04b 

ACP13a 

ACP13b 

2226 

2227 

2227 

2221 

2221 

White mud mound 

White mud mound 

White mud mound 

White bacterial mat 

White bacterial mat 

Nuttallides decorata 

Eponides turgidus 

Hoeglundina elegans 

Globocassidulina subglobosa 

33 

24 

5 

5 

X2 ACB07a 

ACP15b 

FCP61a 

2227 

2221 

2918 

Gray bacterial mat 

Orange bacterial mat 

Tubeworm clump 

Nuttallides decorata 

Cornuspira involvens 

Cibicides wuellerstorfi 

Bolivina lowmani 

Praeglobobulimina ovata 

16 

16 

14 

11 

8 

X3 ACP15a 2221 Orange bacterial mat Cornuspira involvens 100 

Y1.1 GCP12a 

GCP12b 

569 

569 

White bacterial mat 

White bacterial mat 

Bolivina ordinaria 

Gavelinopsis translucens 

Cassidulina carinata 

23 

19 

10 

Y1.2 MCP08a 

MCP08b 

MCP73a 

MCP73b 

1071 

1071 

1070 

1070 

Gray bacterial mat 

Gray bacterial mat 

Bacterial mat 

Bacterial mat 

Bolivina ordinaria 

Cassidulina carinata 

Epistominella exigua 

Gavelinopsis translucens 

Bolivina albatrossi 

Osangularia culter 

Bulimina alazanensis 

Osangularia rugosa

21 

20 

9 

8 

7 

7 

6 

6 

Y2.1 MCP02a 

MCP02b 

MCP69a 

MCP69b 

MCB-C7 

1067 

1067 

1081 

1081 

1076 

Mud from ridge top 

Mud from ridge top 

Bacterial mat 

Bacterial mat 

Non-seep 

Eponides turgidus 

Uvigerina peregrina 

Nonionella iridea 

Bolivina lowmani 

Bulimina aculeata 

Epistominella exigua 

Bolivina albatrossi

13 

9 

8 

8 

8 

6 

5 

Y2.2 GCP54a 

GCP54b 

675 

675 

White bacterial mat 

White bacterial mat 

Cribromiliolinella subvalvularis 

Bolivina ordinaria 

Cornuspira involvens 

Procerolagena gracilis 

Bolivina lowmani 

22 

9 

6 

5 

5 

Y2.3 MCP04a 

MCP04b 

GCP33a 

GCP33b 

GCP51a 

GCP51b 

1072 

1072 

664 

664 

696 

696 

Gray bacterial mat 

Gray bacterial mat 

White bacterial mat 

White bacterial mat 

Bacterial mat on mound 

with clams 

Bolivina ordinaria 

Bulimina aculeata 

Bolivina lowmani 

Epistominella exigua 

Cassidulina carinata 

Gavelinopsis translucens 

Cribromiliolinella subvalvularis 

15 

8 

6 

6 

5 

5 

5 

132
Table 3-5. Q-Mode Minor Clusters for Combined Surface- and Near-Surface-Sample Data: 

Constituent Samples, Water Depths, Substrates, 

and Important Foraminiferal Species (Mean Abundance >5% in Minor Cluster) (continued). 

MINOR 

CLUSTER 

SAMPLES DEPTH 

(m) 

SUBSTRATE IMPORTANT SPECIES % 

Y2.4 GCP15a 

GCP15b 

245 

245 

Non-seep gray mud 

Non-seep gray mud 

Globocassidulina subglobosa 

Bolivina subaenariensis 

Bulimina marginata 

Bolivina ordinaria 

8 

6 

6 

6 

Y2.5 GBP08a 

GBP08b 

GCP55a 

GCP55b 

GCP57a 

GCP57b 

640 

640 

689 

689 

675 

675 

Non-seep gray mud 

Non-seep gray mud 

Non-seep mud 

Non-seep mud 

Sand 

Sand 

Globocassidulina subglobosa 

Bolivina ordinaria 

Epistominella exigua 

Eponides turgidus 

Bolivina lowmani 

Gavelinopsis translucens 

Bolivina albatrossi 

Cassidulina carinata 

Cibicidoides bradyi 

Uvigerina peregrina 

8 

8 

7 

6 

6 

6 

6 

5 

5 

5 

133
However, several surface samples that clustered together in the CA of the first dataset, do not 

cluster together in this second CA (Tables 3.4 and 3.5). 

Table 3-6 lists the 15 most important calcareous species affecting cluster formation, the minor 

clusters in both dendrograms (Figs. 3.3 and 3.5) to which they contribute greater than 5%, and 

their previously recorded depth ranges in the Gulf of Mexico. Nuttallides decorata is a defining 

species in clusters A and X, but is absent from shallower-bathyal samples of clusters B and Y 

(Figs. 3.4 and 3.6). Eponides turgidus and Bolivina lowmani also are important species in 

clusters A and X, although they are not restricted to deepest bathyal samples. Bolivina ordinaria, 

Cassidulina carinata, Gavelinopsis translucens, Osangularia rugosa, and Bolivina albatrossi are 

important in the minor clusters within B and Y, and are absent or negligible in the deepest 

bathyal samples (Figs. 3.4 and 3.6). These species have all been previously recorded from 

comparable depths within the Gulf of Mexico (Sen Gupta et al., in press). Cribromiliolinella 

subvalvularis, the dominant species in minor clusters B3 (Table 3-4, Figs. 3.4) and Y2.2 (Table 

3-5, Fig. 3.6), has not been previously identified from the Gulf of Mexico. 

3.6. Agglutinated Foraminifera 

We identified 60 species of agglutinated Foraminifera, belonging to four orders, at the sampling 

sites. The dominant species belong to the genus Hyperammina. Other important agglutinated 

taxa include Saccorhiza ramosa, Portatrochammina antarctica, Prolixoplecta parvula, 

Trochamminoides coronatus, Ammodiscus tenuis, and Lagenammina tubulata. 

Agglutinated Foraminifera are an important component of the assemblage in many samples, but 

estimations of their number, particularly those for tubular species such as Hyperammina and 

Saccorhiza (which are frequently broken into segments), can be hindered by taphonomic 

problems. Estimates of agglutinated Foraminifera as a percentage of the total assemblage are 

provided in Table 3-7, but these are possibly overestimates, because a significant proportion of 

the assemblage may be composed of tubular species. In 28 samples, agglutinated species 

constitute more than 15% of the foraminiferal assemblage; each of these samples contain >50 

agglutinated individuals. The species that make up >5% of the agglutinated assemblage are listed 

in Table 3-7. 

One sample, ACP11b, is barren of Foraminifera. Five samples contain only calcareous species, 

whereas two samples contain only agglutinated species; all but two of these seven samples 

(ACP01b, 2.7 individuals/cm3

; ACP13b, 4.7 individuals/cm3) contain less than 1 individual/cm3

(Table 3-3). The assemblage in 12 samples is 85-100% calcareous, with <50 agglutinated 

individuals in each sample; two samples from Alaminos Canyon contain <25 agglutinated 

individuals. No dominant agglutinated species is listed for these agglutinant-poor samples in 

Table 3-7. 

Twelve samples have >49% agglutinated Foraminifera; three have >90%. (Nine of the 12 

samples are from bacterial mat substrates.) Hyperammina is the dominant genus in 7 samples––

three from Alaminos Canyon (ACB07a, ACB08a, ACB09a), two from Mississippi Canyon 

(MCP04b, MCP08a), and two from Green Canyon (GCP33a, GCP33b); Hyperammina sp. is the 

dominant species in all these samples but MCP04b, in which Hyperammina laevigata is 

dominant. Other species that contribute >5% to these agglutinated assemblages include 

134
Table 3-6 

List of the 15 Most Important Calcareous Species in Cluster Analysis 

These species have >5% mean abundance in the minor clusters noted. 

The previous records are taken from Sen Gupta et al. (in press). 

SPECIES OCCURRENCE 

(CURRENT STUDY) 

PREVIOUS 

RECORD IN 

GULF (m) 

MINOR CLUSTERS 

Nuttallides decorata DC, AC, FC 155-3850 A1, A2 

X1.1, X1.2, X2 

Eponides turgidus GCs, GCd, GB, SMC, 

MC, DC, AC, FC 

1-3850 A1, A2, B2.1 

X1.1, X1.2, Y2.1 

Bolivina lowmani GCs, GCd, GB, SMC, 

MC, DC, AC, FC 

0-3632 A1, B2.1, B2.2 

Y2.1, Y2.2, Y2.3, Y2.5 

Cibicides wuellerstorfi DC, AC, FC 457-3850 A2 

X2 

Praeglobobulimina 

ovata 

GCs, GCd, MC, AC, FC 10-2432 A2 

X2 

Cornuspira involvens GCd, SMC, AC 1-1426 A2, C 

X2, X3, Y2.2 

Hoeglundina elegans GCs, GCd, SMC, DC, 

AC, FC 

36-3632 A2 

X1.2 

Bolivina ordinaria GCs, GCd, GB, SMC, 

MC, DC 

18-3164 B1, B2.2, B3 

Y1.1, Y1.2, Y2.2, Y2.3, Y2.4, Y2.5 

Cassidulina carinata GCs, GCd, GB, SMC, 

MC, AC, FC 

21-3237 B1, B2.2, B3 

Y1.1, Y1.2, Y2.3, Y2.5 

Gavelinopsis translucens GCs, GCd, GB, SMC, 

MC 

0-3246 B1 

Y1.1, Y1.2, Y2.3, Y2.5 

Osangularia rugosa GCs, GCd, GB, SMC, 

MC, DC, AC, FC 

256-3850 B3 

Y1.2 

Bolivina albatrossi GCs, GCd, GB, SMC, 

MC, DC, AC, FC 

8-3237 B1, B2.1, B2.2 

Y1.2, Y2.1, Y2.5 

Cribromiliolinella 

subvalvularis 

GCd, MC, AC None B3 

Y2.2, Y2.3 

Nonionella iridea GCd, GB, SMC, MC, 

DC 

None B2.1 

Y2.1 

Globocassidulina 

subglobosa 

GCs, GCd, GB, SMC, 

MC, DC, AC, FC 

2-3850 B2.2 

X1.2, Y2.4, Y2.5 

GCs = Green Canyon shallow (245 m); GCd = Green Canyon deep (569-696 m); GB = Garden Banks (640 m); SMC = 

South of Mississippi Canyon (1076 m); MC = Mississippi Canyon (1067-1081 m); DC = De Soto Canyon (1848 m); AC = 

Alaminos Canyon (2219-2227 m); FC = Farnella Canyon (2918 m). 

135
Table 3-7 

Counts (N) and Cumulative Percentages (Agg. %) of Agglutinated Individuals in Samples, With 

Listing of Dominant Species (>5% of Agglutinated Assemblage) 

SAMPLE 

NUMBER 

WATER 

DEPTH 

(m) 

SUBSTRATE AGG. 

% 

DOMINANT AGGLUTINANT N 

ACP01a 2226 White mud mound 0 0 

ACP01b 2226 White mud mound 0 0 

ACP04a 2227 White mud mound 3 6 

ACP04b 2227 White mud mound 2 5 

ACP05a 2225 Mud next to tubeworm clump 49 Saccorhiza ramosa 30 

ACP05b 2225 Mud next to tubeworm clump 57 Saccorhiza ramosa, Hyperammina 

sp., Portatrochammina antarctica 

41 

ACP10a 2219 Orange bacterial mat with 

tubeworm roots 

93 4 

ACP10b 2219 Orange bacterial mat with 

tubeworm roots 

0 0 

ACP11a 2219 Mud next to mussel bed 38 2 

ACP11b 2219 Mud next to mussel bed 0 0 

ACP12a 2219 Mud next to mussel bed 100 1 

ACP12b 2219 Mud next to mussel bed 100 3 

ACP13a 2221 White, fluffy bacterial mat 3 1 

ACP13b 2221 White, fluffy bacterial mat 0 0 

ACP15a 2221 Orange bacterial mat 29 Portatrochammina antarctica, 

Ammodiscus tenuis 

9 

ACP15b 2221 Orange bacterial mat 25 Portatrochammina antarctica 4 

ACB07a 2227 Gray bacterial mat with 

tubeworm roots 

94 Hyperammina sp., Ammodiscus 

tenuis, Portatrochammina 

antarctica 

8 

ACB08a 2227 Gray bacterial mat 99 Hyperammina sp., Ammodiscus 

tenuis, Portatrochammina 

antarctica 

13 

ACB09a 2227 Fluffy bacterial mat 99 Hyperammina sp., 

Portatrochammina antarctica 

7 

ACB19a 2219 Non-seep gray mud 43 Saccorhiza ramosa, Hyperammina 

sp. 

32 

MCP02a 1067 Mud from ridge top, near mussel 

bed 

27 Prolixoplecta parvula, 

Trochamminoides coronatus, 

Portatrochammina antarctica, 

Hyperammina laevigata 

18 

MCP02b 1067 Mud from ridge top, near mussel 

bed 

11 Prolixoplecta parvula, 

Trochamminoides coronatus 

16 

MCP04a 1072 Gray bacterial mat 32 Prolixoplecta parvula, 

Portatrochammina antarctica 

5 

MCP04b 1072 Gray bacterial mat 66 Hyperammina laevigata, Textularia 

sp. Portatrochammina antarctica 

9 

MCP08a 1071 Gray bacterial mat 58 Hyperammina sp. 2 

MCP08b 1071 Gray bacterial mat 1 1 

136
Table 3-7. Counts (N) and Cumulative Percentages (Agg. %) of Agglutinated Individuals in 

Samples, With Listing of Dominant Species (>5% of Agglutinated Assemblage) (continued). 

SAMPLE 

NUMBER 

WATER 

DEPTH 

(m) 

SUBSTRATE AGG. 

% 

DOMINANT AGGLUTINANT N 

MCP69a 1081 Bacterial mat 37 Prolixoplecta parvula, 

Trochamminoides coronatus, 

Lagenammina difflugiformis, 

Portatrochammina antarctica, 

Hyperammina sp. 

21 

MCP69b 1081 Bacterial mat 33 Prolixoplecta parvula, 

Trochamminoides coronatus, 

Portatrochammina antarctica, 

Hyperammina sp.

30 

MCP73a 1070 Bacterial mat 7 6 

MCP73b 1070 Bacterial mat 6 4 

GCP33a 664 White bacterial mat 60 Hyperammina sp., Prolixoplecta 

parvula, Portatrochammina 

antarctica, Hyperammina laevigata, 

unidentified trochamminids 

17 

GCP33b 664 White bacterial mat 54 Hyperammina sp., Hyperammina 

laevigata, Prolixoplecta parvula, 

Portatrochammina antarctica, 

unidentified trochamminids

14 

GCP51a 696 Bacterial mat on mound with 

clams 

18 Portatrochammina antarctica, 

Prolixoplecta parvula, 

Psammosphaera fusca, Discammina 

compressa, unidentified 

trochamminids 

11 

GCP51b 696 Bacterial mat on mound with 

clams 

8 9 

GCP54a 675 White bacterial mat 49 Prolixoplecta parvula, 

Hyperammina sp., 

Portatrochammina antarctica, 

unidentified trochamminids

15 

GCP54b 675 White bacterial mat 62 Prolixoplecta parvula, 

Hyperammina sp., 

Portatrochammina antarctica, 

unidentified trochamminids 

19 

GCP55a 689 Non-seep mud 29 Hyperammina sp., Lagenammina 

tubulata, Haplophragmoides sp. cf. 

H. kirki, Saccorhiza ramosa, 

unidentified trochamminids 

30 

GCP55b 689 Non-seep mud 21 Lagenammina tubulata, Gaudryina 

minuta, Hyperammina sp., 

Hyperammina friabilis, Reophax 

guttifera, unidentified 

trochamminids

30 

GCP57a 675 Non-seep sand 20 Lagenammina tubulata, 

Psammosphaera fusca, Rhizammina 

algaeformis, Hyperammina 

friabilis, Saccorhiza ramosa, 

unidentified trochamminids

34 

137
Table 3-7. Counts (N) and Cumulative Percentages (Agg. %) of Agglutinated Individuals in 

Samples, With Listing of Dominant Species (>5% of Agglutinated Assemblage) (continued). 

SAMPLE 

NUMBER 

WATER 

DEPTH 

(m) 

SUBSTRATE AGG. 

% 

DOMINANT AGGLUTINANT N 

GCP57b 675 Non-seep sand 21 Hyperammina friabilis, Gaudryina 

minuta, Lagenammina tubulata, 

Psammosphaera fusca, 

Hyperammina sp., 

Portatrochammina antarctica, 

unidentified trochamminids

38 

GCP12a 569 White bacterial mat 1 4 

GCP12b 569 White bacterial mat 1 5 

GBP08a 640 Non-seep gray mud 12 Prolixoplecta parvula 21 

GBP08b 640 Non-seep gray mud 11 Haplophragmoides sp. cf. H. kirki 18 

GCP15a 245 Non-seep gray mud 41 unidentified trochamminids

Lagenammina difflugiformis, 

Pseudogaudryina atlantica, 

Pseudoclavulina serventyi, 

Tetrataxiella sp. 

49 

GCP15b 245 Non-seep gray mud 30 Lagenammina difflugiformis, 

Tetrataxiella sp., unidentified 

trochamminids

40 

MCB-C7 1076 Non-seep 54 Rhizammina indivisa, Lagenammina 

difflugiformis, Hyperammina 

friabilis, Lagenammina tubulata, 

Portatrochammina antarctica, 

Trochamminoides coronatus

33 

DCB-S36 1848 Non-seep 36 Saccorhiza ramosa, Lagenammina 

tubulata, Trochamminoides 

coronatus, Usbekistania charoides, 

Reophax scorpiurus, Hyperammina 

friabilis 

50 

FCP61 2918 Tubeworm clump 21 Saccorhiza ramosa, Hyperammina

sp., Martinottiella communis 

21 

138
Figure 3.6. Relative abundances of the more important species in minor clusters, surface and 

near-surface samples. 

139
Portatrochammina antarctica (all areas), Ammodiscus tenuis (Alaminos Canyon), and 

Prolixoplecta parvula (Green Canyon). As regards the other five samples of this group (>49% 

agglutinated), GCP54a and GCP54b are bacterial mat samples from Green Canyon in which 

Prolixoplecta parvula is dominant. Saccorhiza ramosa is dominant in non-seep samples 

(ACP05a, ACP05b) from Alaminos Canyon, and Rhizammina indivisa is dominant in a non-seep 

sample (MCB-C7) from south of Mississippi Canyon. 

Saccorhiza ramosa is the dominant agglutinated species in non-seep, deepest-bathyal samples 

and Portatrochammina antarctica is dominant in deepest-bathyal bacterial mats. Prolixoplecta 

parvula is dominant in both bacterial mat and non-seep sites from bathyal Mississippi Canyon. 

Portatrochammina antarctica is important in most of these samples. Diverse agglutinated 

species, including several unidentified trochamminids, constitute a significant part of the 

assemblage in both shallow and deep Green Canyon samples. Other important species include 

Portatrochammina antarctica, Lagenammina tubulata, and Hyperammina friabilis. 

Samples from bacterial mat substrates have a smaller number of agglutinated species than nonseep samples at comparable depths (Table 3-7). In Alaminos and De Soto Canyons, non-seep 

samples contain 30-50 agglutinated species, compared to 4-13 species in bacterial mats. Nonseep samples from Mississippi Canyon and south of Mississippi Canyon have 18-33 species, 

compared to 2-9 species in mats. Bacterial mat samples MCP69a (21 species) and MCP69b (30 

species) will be discussed later. Non-seep Green Canyon samples have 30-49 agglutinated 

species, compared to 4-19 species in bacterial mat samples. 

3.7. Diagenesis 

Samples MCP73a, MCP73b, and MCP08b contain numerous shells of Hyperammina sp. that 

show overgrowths of barite or are infilled with barite rosettes (Appendix, Plate 84). Also, barite 

chimneys are known to be associated with hydrocarbon venting in Mississippi Canyon at 500 m 

water depth (Fu et al., 1994). Besides barite, the constituents of these chimneys include minor 

amounts of pyrite, iron oxide, Mg calcite, and detrital silicates. Our finding of barite rosettes 

cemented by iron oxide in shells of Hyperammina at an 1100 m deep site extends the water depth 

of barite formation in Mississippi Canyon by about 600 m. 

Pyrite-filled shells of calcareous and agglutinated Foraminifera have been recovered from 

Alaminos Canyon cores. Figure 6, Plate 110 (Appendix) shows pyrite framboids infilling a 

calcareous Nuttallides decorata shell. The presence of pyrite is an indicator of anoxia in 

sediments, and pyritized foraminiferal shells have been found in diverse oxygen-deficient 

environments. Of particular relevance to our study, however, is the reported occurrence of pyrite 

framboids within shells of Uvigerina peregrina in cold-seep sediments of Monterey Bay, 

California (Stakes et al., 1999), attesting to anoxia and bacterial reduction of sulfate there, as is 

the case at Alaminos Canyon seep sites. 

3.8. Stratigraphic Record 

In order to investigate recent temporal variation in seepage intensity––as indicated by the 

foraminiferal assemblage––we examined species frequencies in death assemblages in multiple 

140
downcore samples from two Alaminos Canyon cores and one Mississipi Canyon core. For 

numerical analyses, we considered just the calcareous species that constitute at least 1% of the 

assemblage in one or more samples (Tables 3.8, 3.9, 3.10). The Alaminos Canyon cores, ACP05 

and ACP15, were from adjacent locations and similar water depths (about 2220 m); the 

Mississippi Canyon core, MCP73, was from a depth of 1070 m (see Table 1.1 for coordinates). 

Twenty-two species or congeneric species groups (identified at generic level) provided the data 

for ACP05 and ACP15, whereas the MCP73 data came from 13 such taxa. 

Alaminos Canyon core ACP05. This core was taken through the sediment next to a tubeworm 

bush. The top five samples (each 1-cm thick) were examined. Taken separately, the stratigraphic 

trends in the relative abundances (percentages) of most species seem erratic, but two major 

species, Bolivina lowmani and Nuttallides decorata, show a definite unidirectional change. B. 

lowmani, an endobenthic species, putatively more tolerant to sulfide emission and seepagerelated dysoxia, shows a marked upcore increase between 5 and 2 cm. In contrast, N. decorata, 

an epibenthic species that dominates samples of non-seep clusters, and is possibly less tolerant of 

seepage effects, shows a sharp upcore decrease between 5 and 3 cm (Fig. 3.7). A Q-mode cluster 

analysis of the assemblage data (Table 3-8) corroborates this change. Two major clusters are 

present; one with the three uppermost samples, the other with the two samples farther down (Fig. 

3.8). We surmise that a higher level of seep gases is reflected in the assemblage composition in 

the top 3 cm of the core. 

Alaminos Canyon core ACP15. This core was taken through a bacterial mat at the sedimentwater interface. All 1-cm samples were examined from the core top down to 13 cm. The 

variations in abundances of three species, in all but the surface (0-1 cm) sample, are shown in 

Fig. 3.9. The surface sample is anomalous, because it contains just one calcareous species, 

Cornuspira involvens, which disappears below the 3-cm level in the core. The abundance of the 

endobenbenthic species Bolivina lowmani stays between 4 and 11% from 13 to 7 cm, and then 

rises steeply and linearly to 39% at 3 cm, undergoing a small decrease in the next 2 cm. The 

abundance of Epistominella exigua (also endobenthic) attains a high of 14% in the 4-5 cm 

sample, but is <6% in all other samples. The trend of the epibenthic Nuttallides decorata is 

clearly opposed to that of B. lowmani; it shows a decline from 13 cm (30%) to 7 cm (7%). The 

cluster diagram (Fig. 3.10) shows two main clusters, each with very similar samples (at small 

Euclidean distances). One cluster consists of the 1-2 and 2-3 cm samples; the other includes the 

rest, except an outlier (the 0-1 cm sample). We infer an increase of seepage in the time interval 

represented by the 3-7 cm segment of the core. 

Mississippi Canyon core MCP73. The surface sediment at the coring site was black, with an 

orange bacterial mat on top. Seven selected samples––a continuous series from 0 to 5 cm, and 

the 6-7 and 14-15 cm samples––were examined for Foraminifera. Percentage plots for 

Cassidulina carinata (endobenthic) and Osangularia culter (epibenthic) are shown in Fig. 3.11. 

The distributions do not show any conspicuous trends, possibly because of sampling gaps. 

However, the lowest abundance of C. carinata (5%) and the greatest abundance of O. culter

(20%) are in the oldest sample (14-15 cm); younger sediments may have been deposited under 

conditions of increased seepage. The cluster diagram (Fig. 3.12) does not show clusters of 

contiguous samples; the 14-15 cm sample is an outlier. 

141
Table 3-8 

Core ACP05, Alaminos Canyon (Alvin Dive 3624): 

Percentages of Calcareous Species That Constitute at Least 1% of the Assemblage in One or 

More Samples. 

SAMPLE 

0-1 cm 1-2 cm 2-3 cm 3-4 cm 4-5 cm 

SPLIT EXAMINED 1 1 1/4 1/4 1/4 

TOTAL CALCAREOUS SPECIMENS 261 1056 226 218 328 

SPECIES PERCENTAGES 

Bolivina lowmani 22.6 23.5 21.7 18.8 12.5

Eponides turgidus 31.0 27.6 25.7 20.2 25.6

Globocassidulina subglobosa 3.5 3.8 4.9 2.8 1.8

Gyroidina orbicularis 1.5 0.8 2.2 4.1 1.2

Nuttallides decorata 26.8 24.9 25.2 30.3 36.6

Quinqueloculina spp. 1.5 1.6 2.7 0.9 0.3

142
Table 3-9 

Core ACP15, Alaminos Canyon (Alvin Dive 3625): Percentages of Calcareous Species That 

Constitute at Least 1% of the Assemblage in One or More Samples. 

SAMPLE 

 

0-1 

cm 

1-2 

cm 

2-3 

cm 

3-4 

cm 

4-5 

cm 

5-6 

cm 

6-7 

cm 

7-8 

cm 

8-9 

cm 

9-10 

cm 

10-11 

cm 

11-12 

cm 

12-13 

cm 

SPLIT 

EXAMINED 

1 1 1 1 1 1 1 1 1 1 1 1 1 

TOTAL 

CALCAREOUS 

SPECIMENS 

183 44 32 18 7 28 74 133 140 231 227 268 220 

SPECIES PERCENTAGES 

Cornuspira 

involvens 100 31.8 18.8 0 0 0 0 0 0 0 0 0 0 

Epistominella 

exigua 0 0 0 5.6 14.3 0 5.4 3.8 2.9 3 1.3 2.6 0 

Epistominella 

sp. 0 0 12.5 0 0 0 0 0 0 0 0 0 0 

Eponides sp. cf.

E. regularis 0 0 0 0 0 0 0 0.8 0 0.4 0.9 0.8 2.3 

Eponides 

turgidus 0 2.3 3.1 5.6 0 28.6 32.4 11.3 29.3 32 30.4 28 16.8 

Fissurina spp. 0 0 0 0 0 3.6 1.4 2.3 1.4 2.2 2.2 1.1 0.9 

Gyroidina 

orbicularis 0 0 0 0 0 0 1.4 3.8 0.7 2.6 1.3 0.4 1.4 

Hoeglundina 

elegans 0 0 0 0 14.3 17.9 9.5 12 8.6 4.8 9.3 7.1 9.1 

Lagena 

hispidula 0 0 0 0 0 0 4.1 0 0 0 0 0.4 0 

?Lagena spp. 0 0 0 0 0 3.6 1.4 1.5 0.7 0.9 0.9 0.4 1.8 

Neoconorbina 

terquemi 0 0 3.1 0 0 0 0 0 0 0 0.4 0 0 

Nonion sp. 0 22.7 21.9 16.7 28.6 0 0 0 0 0.4 0 0 0 

Nonionellina sp. 0 0 3.1 0 0 0 1.4 0 0 0 0 0.4 0.5 

Nuttallides 

decorata 0 9.1 3.1 0 0 10.7 6.8 21.8 17.9 21.2 18.9 25.4 30 

Oridosalis 

umbonatus 0 0 0 5.6 14.3 3.6 0 3 0 1.3 0.9 0.8 1.4 

Osangularia 

culter 0 0 0 0 0 3.6 2.7 1.5 4.3 0.9 5.3 2.2 2.7 

Parafissurina 

kerguelenensis 0 0 0 0 0 3.6 0 1.5 0.7 0.4 0 0 0 

Pullenia 

quinqueloba 0 0 0 5.6 0 3.6 10.8 4.5 2.9 6.5 4.4 2.2 4.1 

Quinqueloculina 

spp. 0 0 0 0 0 0 1.4 3 1.4 1.7 0.4 0.8 0.5 

Siphonodosaria 

calomorpha 0 0 0 0 0 3.6 0 0 0.7 0.9 0 0.4 0 

143
Table 3-10 

Core MCP73, Mississippi Canyon (Alvin Dive 3631): Percentages of Calcareous Species That 

Constitute at Least 1% of the Assemblage in One or More Samples. 

SAMPLE 

 

0-1 cm 1-2 cm 2-3 cm 3-4 cm 4-5 cm 6-7 cm 14-15 

cm 

SPLIT EXAMINED 

1/8 1/16 1/16 1/16 1/8 1/4 3/16 

TOTAL CALCAREOUS 

SPECIMENS 133 90 80 86 103 75 121 

SPECIES PERCENTAGES 

Bolivina albatrossi 3.76 4.44 2.5 6.98 0.97 2.67 7.44 

Bolivina lowmani 8.27 4.44 8.75 5.81 8.74 10.67 7.44 

Bolivina ordinaria 13.53 25.56 30 20.93 31.07 17.33 22.31 

Bulimina alazanensis 5.26 7.78 2.5 10.47 0.97 1.33 5.79 

Bulimina mexicana 0.75 0 0 0 0 2.67 0 

Cassidulina carinata 24.06 13.33 21.25 20.93 19.42 22.67 4.96 

Globocassidulina subglobosa 4.511 2.22 2.5 1.16 3.88 1.33 2.48 

Epistominella exigua 7.52 8.89 10 4.65 7.77 12 13.22 

Eponides turgidus 0.75 4.44 0 1.163 0.97 0 0.83 

Gavelinopsis translucens 9.023 8.89 7.5 4.65 4.85 1.33 4.96 

Osangularia culter 9.77 8.89 3.75 5.814 5.83 14.67 19.83 

Osangularia rugosa 7.52 4.44 2.5 5.81 4.85 4 2.48 

Quinqueloculina spp. 0 2.22 2.5 0 0 0 0 

144
Figure 3.7. Abundance variations of Bolivina lowmani (solid line) and Nuttallides decorata

(dashed line) in Alaminos Canyon core ACP05. 

Figure 3.8. Q-mode cluster diagram (Ward’s method) for stratigraphic data, Alaminos Canyon 

core ACP05. 

145
Figure 3.9. Abundance variations of Bolivina lowmani (solid line), Epistominella exigua (dotted 

line) and Nuttallides decorata (dashed line) in Alaminos Canyon core ACP15. 

Figure 3.10. Q-mode cluster diagram (Ward’s method) for stratigraphic data, Alaminos Canyon 

core ACP15. 

146
Figure 3.11. Abundance variations of Cassidulina carinata (solid line) and Osangularia culter

(dashed line) in Mississippi Canyon core MCP73. 

Figure 3.12. Q-mode cluster diagram (Ward’s method) for stratigraphic data, Mississippi 

Canyon core MCP73.

147

4. FORAMINIFERA IN TUBEWORM SAMPLES 

[Note: Much of this chapter has been published as an article (Sen Gupta et al., 2007) in Marine Micropaleontology, 

an Elsevier journal (copyright 2006). Excerpts are included here and in Chapter 5, and species illustrations in the 

Atlas (Appendix), with permission from Elsevier.] 

4.1. Introduction 

Overall, many endobenthic species, in the Gulf of Mexico and elsewhere, are known to be 

adapted to the typical environmental constraints of cold-seep seafloor (Akimoto et al., 1994; Sen 

Gupta and Aharon, 1994; Sen Gupta et al., 1997; Rathburn et al., 2000, 2003; Bernhard et al., 

2001; Hill et al., 2003; Robinson et al., 2004; Heinz et al., 2005; Panieri, 2005), but the presence 

of certain epibenthic species (e.g., Cibicides spp.) has remained perplexing, because these 

species are generally known to be restricted to oxic waters (e.g., Kaiho, 1994). To test the 

hypothesis that such species may actually live in oxic microhabitats very close to points of 

seepage, we examined surfaces of vestimentiferan tubeworms (Escarpia laminata and 

Lamellibrachia luymesi). These tubeworms were retrieved from three seepage areas during dives 

of the submersible Alvin in Alaminos Canyon (depth 2225 m) in 2000 (labeled LS in Table 4-1, 

collected by Lorene Smith) and those of the submersible Johnson Sea-Link in Mississippi 

Canyon (620 m) and Green Canyon (562 m) in 2002 (labeled EC and MC in Table 4-1, collected 

by Erik Cordes, Pennsylvania State University). 

4.2. Tubeworm Foraminifera

The identified foraminiferal species are named in Table 4-1; all three major groups of 

Foraminifera (hyaline, porcelaneous, and agglutinated) are represented by these 15 species. They 

are illustrated, mostly in life positions, in the plates of the Atlas (Appendix); the plate references 

are given in Table 4-1. Judging by the edges of the shells as seen in electron micrographs, the 

species are attached by protoplasm or by organic adhesive, and not by mineral cement or special 

shell extensions (see Poag, 1982). Although the tubeworms are endemic to cold seeps, none of 

the identified Foraminifera are endemic to either the seeps or the Gulf of Mexico. 

These foraminiferal species attached to seep tubeworms generally conform to their known waterdepth limits in non-seep areas of the Gulf of Mexico (Sen Gupta et al., in press). The thin-walled 

Calcituba polymorpha (Appendix, Plate 32) is a striking exception. It has been previously 

reported only from a depth of about 1 m in the gulf, but we find it on a tubeworm at 562 m. We 

ascribe this discrepancy to the poor record of the species in sediments due to the "extreme 

fragility" of its shell (Arnold, 1967), and to a dearth of observations on attached Foraminifera 

beyond shelf depths. 

Microhabitat considerations. The escape of methane and sulfide in Gulf of Mexico cold seeps 

is localized and episodic (Roberts and Carney, 1997). Although vestimentiferan tubeworms such 

as Lamellibrachia and Escarpia need sulfide for sustenance, their presence or growth does not 

imply escape of H2S from the seafloor. At the first stage of colony formation, the settlement and 

aggregation of vestimentiferans depend on sulfide emission at the sediment-water interface, but 

at later stages, as H2S in seafloor water is reduced, the colony growth is maintained by the 

tapping of H2S from sediment pore water by downward-extending vestimentiferan roots (Julian 

149
et al., 1999; Freytag et al., 2001; Bergquist et al., 2003). For much of the time during which 

tubeworms grow, the water above the base is oxic and almost sulfide-free, thus providing an 

environment in which many epibenthic foraminifers can settle on tubeworm surfaces. One of 

these taxa, Cornuspira involvens (Appendix, Plate 48), is known to exist as an endobenthic 

species in a non-seep muddy substrate from 0 to 2 cm, but 2-3 cm above the redox boundary 

(Murray, 2003). Apparently, it is opportunistically sessile on vestimentiferan tubes. 

Our surface-sediment samples from the base of Gulf of Mexico tubeworm bushes contain shells 

of some species reported here. One of these samples (FCP61), from a locality in Farnella 

Canyon, contains an unusual assemblage of calcareous Foraminifera. Among the 53 species of 

this group, Praeglobobulimina ovata, an endobenthic species known to live in H2S-containing 

seep sediments (Robinson et al., 2004), is dominant (23%), but Cibicides wuellerstorfi (14%) 

ranks second. This admixture shows that a confusing environmental signal can be introduced in 

the foraminiferal thanatocoenose of cold-seep sediments simply by post-mortem dislodgment of 

attached species from elevated microhabitats (which were not affected by seep gases), even 

without sediment movement by bottom currents. 

150
Table 4-1 

Foraminifera Attached to Vestimentiferan Tubeworms 

SITE 

Alaminos 

Canyon 

Mississippi 

Canyon 

Green 

Canyon 

VESTIMENTIFERAN SPECIES 

(with specimen numbers and heights of tubes) 

Escarpia 

laminata 

Lamellibrachia 

luymesi 

L. luymesi 

FORAMINIFERAL SPECIES 

(with atlas plate numbers) 

(8 tubes, 

LS1-8, 19-

37 cm) 

(6 tubes; EC1, 

26 cm; MC1-5, 

18-32 cm) 

(4 tubes, 

EC2-5, 14-

24 cm) 

(1) CALCAREOUS PERFORATE (HYALINE) GROUP 

Anomalinoides globulosus (Chapman & Parr), Plate 10 T 

Cibicides wuellerstorfi (Schwager), Plate 41 x 

Cibicidoides pachyderma (Rzehak), Plate 43 x x x 

Discanomalina semipunctata (Bailey), Plates 55, 56 T 

Laminononion tumidum (Cushman & Edwards), Plate 95 x x 

Patellina corrugata Williamson, Plate 125 x x 

Planulina ariminensis d'Orbigny, Plates 128, 129 x x 

Spirillina vivipara Ehrenberg, Plates 168, 169 T 

(2) CALCAREOUS IMPERFORATE (PORCELANEOUS) GROUP 

Calcituba polymorpha von Roboz, Plate 32 T 

Cornuspira foliacea (Philippi), Plate 46 TM 

Cornuspira involvens (Reuss), Plate 48 x x 

Miliolinella sp., Plate 106 x x 

(3) AGGLUTINATED GROUP 

Ammoscalaria tenuimargo (Brady), Plate 7 T 

Deuterammina rotaliformis (Heron-Allen & Earland), 

Plate 53

 T T 

Veleroninoides jeffreysii (Williamson), Plate 192 T 

T = only attached to tubeworms, not present in examined seep sediments; M = attached to mussels. 

151

5. DISCUSSION AND CONCLUSIONS 

5.1. Dominant Calcareous Species 

One hundred seventeen species of calcareous Foraminifera were present in surface or nearsurface sediment samples from seep and non-seep (control) sites. Of these, 54 species constituted 

2% or more of the assemblage in one or more samples. The distributions of several dominant or 

common species provide clues to their tolerance to environmental stresses of seep microhabitats. 

Bolivina ordinaria, Cassidulina carinata, Gavelinopsis translucens, Osangularia rugosa, and 

Bolivina albatrossi are important in the shallower-bathyal clusters of B and Y. All of these 

species have been reported as living under Beggiatoa mats in Green Canyon and have been 

considered to be tolerant of anoxia and sulfide toxicity (Sen Gupta et al., 1997). In the present 

study, Rose-Bengal-stained individuals of these five species were not common and were not 

restricted to bacterial mat samples. Stained specimens of Bolivina ordinaria were found in 

bacterial mat samples from Mississippi Canyon (MCP04b) and Green Canyon (GCP54a), and 

stained specimens of Osangularia rugosa were found in bacterial mat samples from Mississippi 

Canyon (MCP08a and MCP08b). Stained individuals of Bolivina albatrossi were found in nonseep samples from Mississippi Canyon (MCP02b) and Green Canyon (GCP55a and GCP57a). (It 

is possible that some sites recorded as "non-seep" during seafloor observations were close to 

points of seepage and the substrate actually contained seep fluids.) Stained individuals of 

Gavelinopsis translucens and Cassidulina carinata were found in both bacterial mat and nonseep samples from Mississippi and Green Canyons. Bolivina ordinaria is the most abundant of 

the five species. It is present in all minor clusters and is the dominant species in many. The other 

four species are important in many of the minor clusters but are never dominant. Cassidulina 

carinata is present in every minor cluster of B and Y and is important in many. Gavelinopsis 

translucens and Bolivina albatrossi are important in most minor clusters. 

Bolivina ordinaria is the dominant species in minor clusters B1, B2.2, Y1.1, Y1.2, and Y2.3. 

Besides the four associates of B. ordinaria mentioned above, Globocassidulina subglobosa, 

Epistominella exigua, Bolivina lowmani, and Uvigerina peregrina are important in these clusters. 

All of these species, except Bolivina lowmani, have been previously identified as living under 

Beggiatoa mats (Sen Gupta et al., 1997). Globocassidulina subglobosa has also been recorded as 

abundant in seep samples from the Gulf of Mexico (Sen Gupta and Aharon, 1994; Sen Gupta et 

al., 1997; Robinson et al., 2004) and the Rockall Trough (Panieri, 2005). It is generally known to 

be abundant in organic-rich, oxygen-depleted sediments (Sen Gupta and Machain-Castillo, 

1993). In our study, however, Rose-Bengal-stained specimens of Globocassidulina subglobosa

were found only in processed non-seep samples from Alaminos and Green Canyons (ACB19a, 

GCP55a, and GCP57a). Uvigerina peregrina, a worldwide deep-sea species, has also been 

recorded as living in severely oxygen-depleted environments (Bernhard and Sen Gupta, 1999), 

such as the Pacific margin of El Salvador (Smith, 1964), Sagami Bay, Japan (Ohga and Kitazato, 

1997), northern Arabian Sea (Jannink et al., 1998), and California margin (Bernhard, 1992; 

Rathburn et al., 2000; Rathburn et al., 2003). 

Bolivina ordinaria is important, but not dominant, in minor clusters B3, Y2.2, Y2.4, and Y2.5; 

Y2.4 also contains Bolivina subaenariensis and Bulimina marginata. Bolivina subaenariensis

153
has been previously recorded as abundant under bacterial mats of Green Canyon (Sen Gupta and 

Aharon, 1994) and also in an oxygen-deprived upwelling region on the Florida Atlantic slope 

(Sen Gupta et al., 1981). Bulimina marginata was observed in shallow-water oxygen-depleted 

samples from Drammensfjord, Norway (Alve, 1990; 1995a; Bernhard and Alve, 1996). In our 

study seven Rose-Bengal-stained specimens of Bulimina marginata were identified from the 

bacterial mat sample GCP54a. 

5.2. Agglutinated Species 

Of the 14 samples that contained >49% agglutinated Foraminifera, 10 were from seep sites. 

Hyperammina is dominant in all but two of these. Ammodiscus tenuis is an important species, 

behind Hyperammina sp., in bacterial mat samples from Alaminos Canyon. In non-seep samples, 

Hyperammina and Saccorhiza dominate. These large tubular Foraminifera are generally 

considered to be fast growing and opportunistic, and are probably able to tolerate some degree of 

oxygen depletion (Gooday et al., 1997). However, Rose-Bengal-stained specimens of 

Hyperammina sp. are found in both seep and non-seep samples, while stained specimens of 

Saccorhiza ramosa are limited to non-seep samples. Apparently, Hyperammina sp. is more 

tolerant of anoxia than Saccorhiza ramosa. In Misissippi Canyon, shells of Hyperammina sp. 

may be encrusted or infilled with barite; formation of authigenic barite here indicates 

hydrocarbon venting (Fu et al., 1994). 

Portatrochammina antarctica is the dominant agglutinated species in samples ACP15a and 

ACP15b; 65% of the specimens in ACP15a caught the Rose Bengal stain. Portatrochammina 

antarctica is likely to be epifaunal, because this mode of life is characteristic of many 

trochospiral agglutinated Foraminifera (Gooday, 1990; Mackensen and Douglas, 1989; Jones and 

Charnock, 1985). 

Two seep samples, GCP54a and GCP54b, contain >49% agglutinated Foraminifera, but the 

dominant species is Prolixoplecta parvula, not Hyperammina sp. This species is also dominant 

in several Mississippi Canyon samples (including MCP04a, a seep sample) that contain 15-49% 

agglutinated individuals. Non-seep samples from Green Canyon have diverse assemblages, and 

include species of the genera Hyperammina, Lagenammina, Saccorhiza, and Reophax.

5.3. Species Diversity 

Mississippi Canyon samples in non-seep minor cluster Y2.1 have more calcareous species (38-

57) than seep samples from this area (16-37). Green Canyon non-seep samples also have higher 

numbers of calcareous species (46, 60-112), compared to bacterial mat samples (27-53). This 

distinction is not as clear with the deepest-bathyal samples. Calcareous species richness is highly 

variable in samples from Alaminos, Farnella, and De Soto Canyons. Sample ACP05a in nonseep minor cluster X1.1 has 30 calcareous species, whereas the other samples in this cluster have 

55-73 species. Samples ACP04a and ACP04b, in minor cluster X1.2 (probably non-seep), have 

72 and 47 calcareous species, respectively, but other samples in X1.2 have 16-32 species. In seep 

minor cluster X2, sample FCP61a contains 53 calcareous species, whereas the other two samples 

have 6-9 species. ACP15a, the only sample in X3, has just one calcareous species. Except for 

154
ACPO4a and ACP05a, the number of calcareous species (55-73) in minor cluster X1.1 is higher 

than in X1.2, X2, and X3 (1-53). 

The agglutinated species richness may be a more reliable indicator of seepage. Alaminos Canyon 

samples in non-seep minor cluster X1.1 have 30-41 species of agglutinated Foraminifera, 

whereas those in seep cluster X2 have 4-8; the outlier seep sample ACP15a has 9. Non-seep 

samples from Mississippi Canyon minor cluster Y2.1 have 16-33 agglutinated species, compared 

to 1-9 species in seep clusters Y1.2 and Y2.3. Green Canyon samples in seep clusters Y1.1, 

Y2.2, and Y2.3 have 4-19 agglutinated species, whereas those in non-seep Y2.4 and Y2.5 have 

30-49 species. 

There is evidence that species richness in some samples may be elevated by post-mortem mixing 

of species from different microhabitats. This problem will be discussed later in this chapter. 

5.4. Absence of Endemics

Data from the present study and all previous reports (Sen Gupta and Aharon, 1994; Sen Gupta et 

al., 1997; Robinson et al., 2004) demonstrate that foraminiferal species from various 

microhabitats of Gulf of Mexico hydrocarbon seeps are not exotic taxa, but recruits from the 

seep-free surrounding area. A similar conclusion is reached when species lists from other cold 

seeps are examined (Akimoto et al., 1994; Rathburn et al., 2000, 2003; Bernhard et al., 2001; 

Torres et al., 2003; Hill et al., 2003; Panieri, 2003, 2005, 2006; Levin, 2005). Thus, unlike the 

macrofauna (annelids, molluscs), no identified foraminiferal taxa are endemic to cold seeps. In 

our present data set, no species identified from the non-seep De Soto Canyon sample (DCB-S36) 

is restricted to that control sample. Twenty-one species found in seep-associated microhabitats in 

this study were previously unknown from the Gulf of Mexico, but they have been reported from 

non-seep areas in other marine basins. 

5.5. Substrate Effect

A major aspect of macrofaunal distribution at Gulf of Mexico seeps (mussel beds versus 

tubeworm clusters) is related to geochemical differences in the environment, because of the 

physiological dependence of seep mussels (Bathymodiolus sp.) on methane (e.g., Childress et al., 

1986) and that of tubeworms (Lamellibrachia sp., Escarpia sp.) on hydrogen sulfide (e.g., 

Brooks et al., 1987). These relationships are reflected in the areal separations of these two types 

of macrofauna in the Bush Hill area (Green Canyon, MMS Block 184/185). MacDonald et al. 

(1989) found a significant correlation (p<0.05) here between (a) the ranked abundance of 

mussels and the concentration of methane in water samples and (b) the ranked abundance of 

tubeworms and the concentration of extractable organic material (related to sulfide level) in 

sediment samples. A comparable pattern has been reported for three foraminiferal species, 

Rutherfordoides cornuta, Nonionella auris, and Bulimina striata, living at Pacific seeps (but not 

at Gulf of Mexico seeps). Apparently, the distributions of the first two species are related to high 

CH4 in the sediment, and that of the third species to high H2S content (Akimoto et al., 1994; 

Wefer et al., 1994). Our seep samples are mainly from under bacterial mats, but the set also 

includes a few samples from muddy sediment adjacent to mussel beds or tubeworm bushes. 

155
However, we found no evidence of any significant difference between the foraminiferal 

communities of these substrates. 

 

5.6. Trophic Resources 

For foraminiferal species that are able to survive the geochemical constraints of seep 

environments, the available food is abundant. Besides particulate food matter in the water and 

sediment, the bacterial productivity is generally orders of magnitude greater around hydrocarbon 

seeps than in non-seep marine environments (Ducklow and Carlson, 1992). Known values of 

bacterial biomass from Green Canyon seeps fit this pattern. Daily production can be as high as 

18 g C m-3 (Hyun, 1994), and microbial doubling time in the seep water column can be as fast as 

about an hour, thus creating a major effect on the carbon resource of the entire benthic 

community (LaRock et al., 1994). In addition, the formation of large bacterial aggregates from 

individual cells possibly makes it easier for the seep community of the Louisiana slope to utilize 

the bacterial food (Hyun et al., 1997). 

5.7. Effect of Oil in the Environment 

One potential source of stress for the Foraminifera in hydrocarbon seeps is the liquid petroleum 

present in bottom water and sediments. In our previous studies (essentially on bacterial mats), we 

could not detect any effect of these natural pollutants on the structure of the foraminiferal 

community (Sen Gupta et al., 1997). Samples with an extraordinary amount of oil were not 

barren of Foraminifera, and did not contain deformed individuals, as reported for a gigantic oil 

spill from a tanker (Vénec Péyre, 1981), or for environments polluted with heavy metals (e.g., 

Alve, 1995b; Yanko et al., 1999). The present study led to the same conclusion. Also, the 

influence of petroleum pollution on foraminiferal communities is reported to be minor or 

undetectable in habitats near Louisiana offshore drilling platforms (Lockin and Maddocks, 1982) 

and in the North Sea (Murray, 1985). As regards non-foraminiferal meiofauna, a community 

survives in a shallow-marine petroleum seep off California, because the "stressful effects....are 

more than offset by dynamic trophic processes associated with the fringes of spots of active 

seepage" (Montagna and Spies, 1985). Furthermore, laboratory experiments on metazoan 

meiofauna indicate an adaptation to increased levels of petroleum hydrocarbons, if the natural 

environment of the community is one of chronic contamination by such chemicals (Carman et 

al., 1995). 

5.8. Bathymetric Imprint 

Like other marine benthos, seafloor foraminiferal communities show conspicuous patterns of 

depth segregation in virtually all marine basins. As demonstrated by numerous studies, the Gulf 

of Mexico is no exception (see, e.g., Phleger, 1960). This bathymetric imprint, caused by a notfully-understood combination of many factors, is also observed in the assemblages described in 

this report. Regardless of whether the constituent samples are from seep or non-seep substrates, 

the two major clusters produced by either cluster analysis are partitioned by bathymetry. Both 

clusters A (just surface samples) and X (surface and near-surface) include all samples from 1848 

to 2918 m, whereas clusters B (surface) and Y (surface and near-surface) include all samples 

156
from 245 to 1081 m. The separation between seep and non-seep samples is seen only in minor 

clusters. 

Overall, the depth distributions of sediment-dwelling species in our samples do not show any 

marked departure from the known depths of their occurrence in the Gulf of Mexico (Sen Gupta 

et al., in press), although our findings have extended these known depth ranges in some cases. 

Epibenthic Foraminifera of seep tubeworms also conform to their known water-depth limits in 

non-seep areas, but the thin-walled porcelaneous species Calcituba polymorpha (Appendix, Plate 

32) provides a striking exception. It has been previously reported only from a depth of about 1 m 

in the gulf, but we find it on a tubeworm at 562 m. We ascribe this discrepancy to the poor 

record of the species in sediments due to the "extreme fragility" of its shell (Arnold, 1967), and 

to a dearth of observations on attached Foraminifera beyond shelf depths (Sen Gupta, et al., 

2007). 

5.9. Tolerance of Anoxia and Sulfide 

Environment of bacterial (Beggiatoa) mats. Beggiatoa is a large, chemolithotrophic, sulfideoxidizing bacterium. Many samples of the sediment under mats of Beggiatoa were examined in 

this and earlier investigations of Gulf of Mexico seep Foraminifera. A critical oxic-anoxic 

boundary zone is present within the mats, with H2S present in the underlying sediment (because 

of sulfate-reducing bacteria), but O2 in overlying water (e.g., Jørgensen, 1977; Spies and Davis, 

1979; Larkin and Strohl, 1983). In laboratory experiments, the disappearance of O2 and 

appearance of H2S have been detected within Beggiatoa mats that were a fraction of a millimeter 

in thickness (e.g., Jørgensen, 1977, 1982; Jørgensen and Revsbech, 1983; Nelson et al., 1986). In 

the Gulf of Mexico, the larger Beggiatoa mats may be several meters wide (Sassen et al., 1993), 

but even the easily recognizable (i.e., apparently well established) mats are no more than a few 

mm thick, and frequently less than one mm thick. The exceptions are the network-like Beggiatoa

mats that follow fissures in hard ground, and are 1 mm to 3 cm thick (Larkin et al., 1994). None 

of our samples, however, were from this type of mat; they were all obtained from round patches 

of mats (1-2 m across) overlying a mud substrate. We surmise that in our studied cores, an oxicanoxic boundary zone was present within about 1 mm from the core top, and Foraminifera living 

below this sediment depth are microaerophiles or facultative anaerobes. In a previous study (Sen 

Gupta et al., 1997), a few living individuals were found at a substrate depth of about 3 cm, 

where the sediment was black and the pore fluid sulfidic. Preliminary carbon isotope data, 

however, have not resolved the question whether species of Foraminifera invading the anoxic 

layers under Beggiatoa mats can construct tests in these microhabitats (Sen Gupta and Aharon, 

1994; Sen Gupta et al., 1997). 

From the vertical distribution of living (Rose-Bengal-stained) individuals of foraminiferal 

species in sediment under Green Canyon Beggiatoa mats, Sen Gupta et al. (1997) concluded that 

several species, including Bolivina albatrossi, B. ordinaria, Cassidulina carinata, and

Gavelinopsis translucens, are facultative anaerobes. Several minor sample clusters recognized in 

the present study (B1, B2.2, Y1.1, Y1.2, and Y2.3) are dominated by these species, especially by 

Bolivina ordinaria; these are regarded as seep clusters. Overall, species of Bolivina are common 

in areas with dysoxic/anoxic conditions and high food supply (Bernhard et al., 1997; Bernhard 

157
and Reimers, 1991; Panieri, 2006), and this adaptation is reflected in their distribution at and 

around seeps. 

In contrast, Nuttalides decorata and Eponides turgidus, common in waters deeper than 1200 m in 

the Gulf of Mexico (Phleger and Parker, 1951), are generally rare or absent at seep locations. All 

Rose-Bengal-stained specimens of these species are from non-seep samples, except for one 

specimen of E. turgidus in the bacterial mat sample GCP51a and one of N. decorata in the 

bacterial mat sample ACB08a. Together, these two species constitute >50% of the assemblage in 

minor clusters A1 and X1 which we interpret as non-seep clusters (although two samples in X1.2 

were collected from white bacterial mats). The control sample from De Soto Canyon, DCB-S36, 

is a component of both clusters. Because of post-mortem mixing of foraminiferal shells, 

however, a relatively high abundance of these species in a sediment sample does not necessarily 

indicate absence of seepage. Minor clusters B2.1 and Y2.1, containing both seep and non-seep 

samples, have dominant Eponides turgidus (14% in B2.1 and 13% in Y2.1). The species 

Nonionella iridea is also important in these minor clusters (10% in B2.1 and 8% in Y2.1). Other 

important species in these minor clusters include the anoxia/dysoxia tolerant Uvigerina 

peregrina, Bulimina aculeata, Epistominella exigua, and Bolivina albatrossi. Nonionella iridea

has not been previously recorded from the Gulf of Mexico, but other species of Nonionella have 

been listed as common in oxygen-depleted environments (Bernhard and Sen Gupta, 1999), 

including N. stella, which was abundant in samples from the Santa Barbara Basin, California 

(Bernhard et al., 1997) and the northern California margin (Rathburn et al., 2000). 

Protistan ecologists generally accept that no "real anaerobic species" are known among the 

Foraminifera (Fenchel and Finlay, 1995). However, in benthic marine habitats, foraminifers 

generally withstand hypoxia better than other meiofauna (Josefson and Widbom, 1988), and 

species that are tolerant to extremely low levels of oxygen have been reported from diverse water 

depths (e.g., Sen Gupta and Machain-Castillo, 1993; Bernhard and Sen Gupta, 1999). In the 

central Santa Barbara Basin, living Foraminifera (determined by ATP assay) with intact 

organelles (determined by electron microscopy) have been found in anoxic pore waters 3 cm 

below Beggiatoa mats, but even these species do not tolerate anoxia indefinitely (Bernhard and 

Reimers, 1991). Such evidence would suggest that the survival of most Foraminifera under 

Beggiatoa mats at Gulf of Mexico hydrocarbon seeps is a short-term phenomenon, but this 

hypothesis has not been tested. Compared to the isobathyal but much larger Beggiatoa mats of 

the silled Santa Barbara Basin, those of the Green Canyon hydrocarbon seeps (data from Sen 

Gupta et al., 1997) support much smaller foraminiferal populations. This discrepancy can be 

explained by (a) the size difference of the mats in the two areas, and (b) the inherently ephemeral 

nature of the Green Canyon mats, whose occurrence is controlled by the shifting point sources of 

sea-floor hydrocarbon emission. Such shifts are connected with the complex interplay of various 

geological and geochemical factors, such as locations of hydrocarbon (methane and petroleum) 

reservoirs, faulting, erosion, and sedimentation (Roberts and Aharon, 1994). 

Porcelaneous species. Very few porcelaneous (miliolid) species have been reported from 

dysoxic to anoxic environments (Bernhard and Sen Gupta, 1999). However, using the Rose 

Bengal stain, living populations of Triloculina trigonula have been recorded in Green Canyon 

from under Beggiatoa mats, and those of Quinqueloculina spp. from under Thioploca mats (Sen 

Gupta et al., 1997; Robinson et al., 2004). Also, living Pyrgo sp. has been reported from seep 

158
habitats in Blake Ridge (Robinson et al., 2004). In this context, we draw attention to 

Cribromiliolinella subvalvularis, previously identified from the Gulf of Mexico. This miliolid 

dominates the surface and near-surface samples (24% and 21%, respectively) in Green Canyon 

core GCP54, taken through a white bacterial mat; 30% of its individuals in the 0-1 cm sample 

(GCP54a) and 43% in the 1-2 cm sample (GCP54b) caught the Rose Bengal stain. Thus, we 

recognize C. subvalvularis to be a facultative anaerobe. Another porcelaneous species in our 

samples that may belong to this category is Cornuspira involvens; the unusually variable 

substrate preference of this species will be discussed in the next paragraph. 

Elevated epibenthos. As indicated above, benthic Foraminifera that actually live in sediments of 

Gulf of Mexico cold seeps need to be tolerant of severe oxygen deprivation and the presence of 

methane and hydrogen sulfide. Judging by the growth environments of sulfide-oxidizing bacteria 

(e.g., Jørgensen, 1982) and foraminiferal abundances in non-seep but sulfide-enriched, bacterial 

mat environments elsewhere (Bernhard and Buck, 2004), cold-seep species living under bacterial 

mats (Sen Gupta et al., 1997) must be particularly tolerant of H2S. This constraint, however, does 

not apply to species that do not live in or on the sediment, but grow attached to vestimentiferan 

tubeworms––centimeters to decimeters above the seafloor––thus avoiding the oxygen depletion 

and H2S toxicity at the sediment-water interface. In this study, we have found 8 species that live 

exclusively on tubeworms. Two of them, Cibicides wuellerstorfi and Planulina ariminensis, have 

been found affixed to elevated microhabitats (rock fragments, hydroids, etc.) in several non-seep 

areas outside the Gulf of Mexico, and it has been argued that this adaptation gives such 

suspension-feeding species a special advantage in a slow bottom current (Lutze and Thiel, 1989; 

Linke and Lutze, 1993). At seep sites, where bacterial and particulate food is plentiful (Levin, 

2005), the attachment to tubeworm surfaces above the seafloor also offers freedom from hypoxia 

and sulfide toxicity (Freytag et al., 2001; Cordes et al., 2003). There is ample documentation that 

the cosmopolitan species Cibicides wuellerstorfi, widely reported (under 5 generic names, see 

Sen Gupta, 1989) from deep-bathyal and abyssal waters (especially from the North Atlantic 

Deep Water), is restricted to oxic environments (e.g., Douglas and Woodruff, 1981; Kaiho, 

1994). Besides our findings in the Gulf of Mexico, C. wuellerstorfi is also known to be attached 

to tubeworms “above the sediment-water interface” at an abyssal seep site on the Aleutian 

margin (Adamic et al., 2005). From these observations, we infer that (1) many sessile 

Foraminifera, of diverse taxonomic affinities and bathymetric preferences, colonize surfaces of 

vestimentiferan tubeworms (and possibly other elevated microhabitats) at hydrocarbon seeps, 

and (2) in many seepage areas, these attachment points are sufficiently above locations of gas 

escape in the seafloor to provide the species with an oxic microhabitat with little or no H2S (Sen 

Gupta et al., 2007). 

One species living attached to Alaminos Canyon tubeworms surfaces is a facultative epibenthos. 

This species, Cornuspira involvens, together with Cibicides wuellerstorfi, is abundant in sample 

ACB07a taken from a gray bacterial mat with tubeworm roots. Arguably, for both species, this 

abundance reflects post-mortem dislodgment from tubeworm stalks. C. involvens is also the sole 

calcareous species in sample ACP15a (also from Alaminos Canyon) collected from an orange 

bacterial mat with tubeworm roots. The individuals, however, were all living (as judged by Rose 

Bengal) and encysted when collected; they did not seem dislodged from tubeworms. 

Foraminifera may become dormant in adverse conditions (Bernhard and Sen Gupta, 1999) and 

encystment may reflect dormancy; cysts of the well-known shelf species Elphidium incertum

159
have been reported from anoxic environments (Linke and Lutze, 1993). Abundant Rose-Bengalstained individuals of C. involvens are also present in ACP15b, 1-2 cm below the sediment-water 

interface. Furthermore, it is known to exist as a shallow endobenthic species in a non-seep 

muddy substrate from 0 to 2 cm, although 2-3 cm above the redox boundary (Murray, 2003). Our 

results indicate that C. involvens is opportunistically sessile on vestimentiferan tubes but may 

also survive as a facultative anaerobe in seep sediments. 

5.10. Post-Mortem Assemblage Mixing 

Some unusual features of our sample assemblages can be explained by post-mortem mixing of 

foraminiferal shells. A notable example comes from the Farnella Canyon surface sample 

(FCP61a), the deepest-water sample (2918 m) in our set. The calcareous species richness (53) is 

one of the highest for samples in the deepest-bathyal major cluster. Praeglobobulimina ovata is 

dominant (23%), but Cibicides wuellerstorfi (14%) ranks second. This combination is enigmatic, 

because the endobenthic P. ovata is known to live at the sedimentary oxic-anoxic boundary 

(Corliss, 1985; Corliss and Emerson, 1990) and even in H2S-containing seep sediments 

(Robinson et al., 2004), whereas C. wuellerstorfi, an elevated-epibenthic species, is restricted to 

oxic waters (Kaiho, 1994; Douglas and Woodruff, 1981; Murray, 1995). Because (a) C. 

wuellerstorfi, a deepest-bathyal to abyssal species, is sessile on vestimentiferan tubeworms, and 

(b) FCP61a was taken from sediment at the base of a tubeworm bush, we conclude that the 

presence of the species there is due to post-mortem dislodgment from tubeworms. Such postmortem mixing of foraminiferal species from entirely different microhabitats could significantly 

raise the number of species (live and dead) present in a sediment. Thus, an abundance of the 

shells of aerophilic, epibenthic Foraminifera in a sediment sample does not necessarily mean 

absence of gas seepage at the seafloor. The data on epibenthic (especially, sessile) species need 

to be treated with caution when particular species or assemblages of fossil Foraminifera are used 

as clues to ancient cold-seep environments (e.g., Barbieri and Panieri, 2004). 

5.11. Conclusions

(1) There are no seep endemics among the 185 identified species of benthic Foraminifera, even 

among the 21 that were previously unknown in the Gulf of Mexico. 

(2) The imprint of water depth on foraminiferal assemblages is clearly detectable because the 

species are recruited from the surrounding non-seep habitats. The two major surface-sample 

groups (clusters) recognized by numerical data analysis are separated by bathymetry. The 

shallower-water group contains all samples (seep and non-seep) from Green Canyon, Garden 

Banks, and Mississippi Canyon (245-1081 m); the deeper-water (deepest-bathyal) group contains 

all samples from Alaminos, Farnella, and De Soto Canyons (1848-2918 m). 

(3) Foraminiferal species of wide-ranging morphologic and taxonomic affinities are able to 

maintain sizeable populations at sites of hydrocarbon seepage. The most conspicuous dominants 

at seep-influenced substrates (bacterial mats) in the shallower cluster are endobenthic species, 

especially Bolivina spp.; these are possibly facultative anaerobes. The pattern is not as clear in 

the deepest-bathyal group, because some epibenthic species (e.g., Nuttallides decorata) are 

included among the dominants. 

160
(4) The high bacterial productivity at northern Gulf of Mexico hydrocarbon seeps is a major 

factor in sustaining the foraminiferal populations. 

(5) The presence of oil in surface and subsurface sediments has no conspicuous effect on the 

dominant foraminiferal species of hydrocarbon seeps. 

(6) The abundance of Cibicides wuellerstorfi in seep sediments is an evidence of post-mortem 

assemblage mixing; C. wuellerstorfi is an elevated-epibenthic species of deep bathyal to abyssal 

depths, and is intolerant of anoxia or dysoxia. 

(7) In the shallower sampling areas, the diversity (species richness) of both calcareous and 

agglutinated Foraminifera is higher in non-seep than in seep substrates. This distinction is not 

clear in the deepest-bathyal areas. The diversity at some sites may have been elevated by postmortem mixing of species from different microhabitats. 

(8) Fifteen epibenthic species were found attached to vestimentiferan tubeworms, centimeters to 

decimeters above the seafloor. Such species are not affected by the oxygen depletion and H2S 

toxicity at the sediment-water interface. Eight of these species were found exclusively on 

tubeworms and not in sediments.

161
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A-1

APPENDIX: ATLAS OF FORAMINIFERA

Explanatory Notes

One hundred eighty species are illustrated in this atlas in 193 plates. The plates are

arranged alphabetically by species names. Each plate contains one or more illustrations

of a single species; most are scanning electron micrographs, but some digital

photomicrographs (mainly of species attached to tubeworms) are also included.

All relevant information on these species, including taxonomy and brief descriptions, is

given in Chapter 2 (Systematics). Sample references for illustrated specimens are also

given. The digital photomicrographs are identified under species names in Chapter 2; all

others are scanning electron micrographs.

Five rare species are not shown on the atlas plates, because of a lack of specimens

suitable for illustration. These species are: Heronallenia sp., Miliolinella subrotunda

(Seguenza), Neoconorbina terquemi (Rzehak), Procerolagena gracillima (Seguenza),

and Trochamminoides coronatus (Brady).

































































































































































































The Department of the Interior Mission

As the Nation's principal conservation agency, the Department of the Interior has responsibility 

for most of our nationally owned public lands and natural resources. This includes fostering 

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