GNS Science Monograph 28

LATE MIOCENE PLIOCENE PLEISTOCENE

WAI- lower upper lower up OPOITIAN WAIP- MANGA- NUKU- CASTLE- HAW-
AUAN TONGAPORUTUAN KAPITEAN IPIAN PANIAN MARUAN CLIFFIAN ERAN

Available from -
G. H. Scott
B. compressa

NZ$20 (pdf) NZ$30 (CD)

B. finlayi
Quaternary)

B. obconica
B. pohana

B. urenuia B. pseudocompressa

B. pliozea
B. medialis

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B. pliobliqua
B. striata
(Foraminifera; Neogene -

B. elegantissima

B. sp. striate

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B. quadrilatera granttaylori
?

B. quadrilatera
Guide to New Zealand Bolivinita
GUIDE TO NEW ZEALAND BOLIVINITA
( FORAMINIFERA; NEOGENE - QUATERNARY)

DRAFT
G.H. SCOTT

GNS Science Monograph 28

(New Zealand Geological Survey Paleontological Bulletin 78)

GNS Science Lower Hutt, New Zealand

2017
 DRAFT
BIBLIOGRAPHIC REFERENCE

Scott, G.H. 2017. Guide to New Zealand Bolivinita (Foraminifera; Neogene - Quaternary). Institute
of Geological & Nuclear Sciences monograph 28. 127 p. Lower Hutt, New Zealand: Institute of
Geological & Nuclear Sciences Limited.

George H. Scott Institute of Geological & Nuclear Sciences Limited, P.O. Box 30368, Lower Hutt,
New Zealand

Typeset with MikTeX by the author

ISBN 978-1-98-850018-8 (CD)

ISBN 978-1-98-85019-5 (Online)
ISSN 2537-6853 (Online)

©Copyright Institute of Geological & Nuclear Sciences Limited 2017

FRONT COVER

Biostratigraphic summary of Bolivinita spp. in New Zealand. Specimens are shown in

spiral orientation.
 Abstract

DRAFT
The biserial benthic foraminiferal genus Bolivinita is widely represented in New Zealand Late Miocene - Recent
shelf and bathyal environments and several of its species are useful in regional biostratigraphy. To advance the
stratigraphic application of the genus, the external morphology and population variation of several of the more widely
distributed species are reviewed, and their biogeography and stratigraphy are summarized. Detailed accounts are
given of Bolivinita compressa Finlay, B. finlayi Kennett, B. medialis Scott n. sp., B. pliobliqua Vella, B. pliozea Finlay,
B. pohana Finlay, B. pseudocompressa Crundwell n. sp., B. quadrilatera (Schwager), B. quadrilatera granttaylori
Vella and B. urenuia Scott n. sp. Bolivinita pseudocompressa Crundwell n. sp., B. obconica Crundwell n. sp. and
B. striata Crundwell n. sp. are proposed in the appendices. Bolivinita elegantissima Boomgaart, B. subangularis
(Brady) and an undescribed species are illustrated and noted; they are not examined in the main study.

Taxa are described and illustrated in three standard orientations (spiral, axial, distal) and terminology is introduced
for the major architectural elements, especially those seen in the shell outline. Important features are identified on
plates that illustrate intra-sample variation. The spiral outline is regarded as particularly useful because of its clear
presentation of chamber components and shell geometry. It is the best orientation for differentiating between taxa
with bilaterally symmetrical shells (Bolivinita quadrilatera, B. medialis, B. pliozea) and those with rhomboidal shells
(B. compressa, B. finlayi, B. pohana, B. pliobliqua, B. pseudocompressa, B. urenuia).

Ordinations and cluster analyses of coefficients of elliptical Fourier curves fitted to outline coordinates of shells in
spiral orientation are used to portray intra- and inter-sample variation, and to assist in the discrimination of taxa.
Stratigraphically, attention is focused on the lowest and highest occurrences of taxa and their regional distributions.

A principal biogeographic feature of the group is the appearance of Bolivinita compressa, B. medialis, B. pohana and
B. quadrilatera in close proximity (probably as migrants) in the New Zealand record at the base of the Tongaporutuan
Stage (base of Late Miocene, c. 11 Ma). These taxa represent most of the shell designs found in the group. In upward
order Bolivinita urenuia, B. pliobliqua, B. pliozea, B. finlayi and B. pseudocompressa appeared in the Late Miocene
- earliest Pliocene. Architecturally, most resemble earlier taxa and several may be their descendants. Intra- and
inter-sample variation is often large and complicates the recognition and discrimination of taxa. Several species are
interpreted as polytypic although some populations presently interpreted as intra-specific variants may warrant formal
recognition. The record of speciation events is poor and the phyletic history of Bolivinita in New Zealand is obscure.

Bolivinita was widely distributed in onshore Late Miocene bathyal environments in New Zealand. Generally, species
of Bolivinita were minor components of Late Miocene benthic foraminiferal assemblages but were often persistent.
Localized blooms of rhomboidal taxa (e.g., B. compressa, B. pohana) suggest that they could be opportunists. In
the Kapitean Stage (latest Miocene) the distribution of the principal rhomboidal species (B. compressa) contracted,
although the record may be biased by the prevalence of shelf sedimentation in New Zealand basins at this time. Con-
currently, B. pliozea, a bilaterally symmetrical design, evolved from a weakly rhomboidal ancestor (B. pliobliqua). It
became widespread in Pliocene outer shelf and upper bathyal assemblages and persisted into the Pleistocene. There
are few younger records and it is unlikely that it is a member of the Recent fauna. Bolivinita pseudocompressa,
the principal Pliocene rhomboidal species, was largely restricted to one basin. Records of the globally-distributed
Bolivinita quadrilatera (one of the original migrants), are sporadic through its long presence in the New Zealand
region.

i
Preface

DRAFT
Research for this study concluded in 1997; more recent records of Bolivinita in the New Zealand Fossil
Record File (http://www.fred.org.nz) have not been considered. Expectedly, these newer data
may affect the taxonomy and observed stratigraphic ranges of taxa reported here. The elapsed time be-
tween research and publication has also impinged on statistical analysis of the data. This was completed
prior to the wide availability of tools for geometric morphometrics, particularly from the R Project for Sta-
tistical Computing (https://www.r-project.org/). These enable better portrayal of intra-sample
variation, and fuller analysis of shape transformations than is reported here. In retrospect, the study con-
tributes more to demonstrating the variability of Bolivinita populations than to resolving their classification
as morphospecies.

The format and medium of this publication represent an effort to improve the transfer of information to other
taxonomists. While images potentially are more effective for information transfer, words are needed to guide
their interpretation. Yet in many publications images and their related text are often widely separated. Here
an attempt is made to more closely associate and cross-reference them. However, the portable document
format used here is a paper-orientated medium, and is not ideal for juxtaposing images and text. This
is better achieved with a split-screen layout whereby text and images are juxtaposed and are separately
moveable.

Acknowledgments

GNS Science provided facilities for the completion of this work. Several staff gave valued assistance: Martin
Crundwell (collections from Wairarapa), Des Darby and Roger Williams (computation), Philip Carthew
(cover design), and Wendy St George (imagery in the Appendices). I am grateful to Bruce Hayward and
Hugh Grenfell (Geomarine Research), and James Crampton for reviewing the manuscript.

Keywords

Foraminifera; benthic; Bolivinitidae; Bolivinita; taxonomy; New Zealand; Neogene; Quaternary; biostratig-
raphy; morphometrics; evolutionary history

ii
Contents

DRAFT
I Preamble 1

1 Introduction 2
1.1 Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.3 Samples . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.4 Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.5 Spiral Outline Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2 Morphology 6
2.1 Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2.2 Significance of Spiral Orientation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Bolivinita quadrilatera - A Bilaterally Symmetrical Design . . . . . . . . . . . . . . . . . 6
Bolivinita compressa - A Rhomboidal Design . . . . . . . . . . . . . . . . . . . . . . . . . 8

3 Interpretation 9
3.1 Shell Design . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3.2 Comparative Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3.3 Modelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3.4 Functional Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.5 Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.6 Evolutionary History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

II Taxonomy 19

4 Bolivinita compressa Finlay 20

4.1 Bolivinita compressa Reference Population X18/f38 . . . . . . . . . . . . . . . . . . . . . 20
4.2 Bolivinita compressa Comparative Population T27/f399 . . . . . . . . . . . . . . . . . . . 22
4.3 Bolivinita compressa Comparative Population P29/f11 . . . . . . . . . . . . . . . . . . . 23
4.4 Bolivinita compressa Comparative Population T26/f139 . . . . . . . . . . . . . . . . . . . 24
4.5 Overview of Variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
4.6 Bolivinita compressa Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
4.7 Bolivinita compressa Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . 27

5 Bolivinita pseudocompressa Crundwell 30

5.1 Bolivinita pseudocompressa Reference Population T27/f78 . . . . . . . . . . . . . . . . . 30
5.2 Bolivinita pseudocompressa Comparative Population X19/f9493 . . . . . . . . . . . . . . 32
5.3 Bolivinita pseudocompressa Comparative Population W21/f8521 . . . . . . . . . . . . . . 33
5.4 Discrimination Bolivinita pseudocompressa : B. compressa . . . . . . . . . . . . . . . . . 34
5.5 Discrimination Bolivinita pseudocompressa : B. finlayi . . . . . . . . . . . . . . . . . . . 36
5.6 Bolivinita pseudocompressa Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . 37
5.7 Bolivinita pseudocompressa Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . 37

6 Bolivinita finlayi Kennett 39

6.1 Bolivinita finlayi Reference Populations U23/f142, U24/f697-8 . . . . . . . . . . . . . . . 39
6.2 Discrimination Bolivinita compressa : B. finlayi . . . . . . . . . . . . . . . . . . . . . . . 41
6.3 Bolivinita finlayi Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
6.4 Bolivinita finlayi Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . . . 43

iii
7 Bolivinita urenuia Scott n. sp. 44
7.1 Bolivinita urenuia Type Population Q19/f102 . . . . . . . . . . . . . . . . . . . . . . . . . 44

DRAFT
7.2 Bolivinita urenuia Comparative Population J32/f54 . . . . . . . . . . . . . . . . . . . . . 46
7.3 Bolivinita urenuia Comparative Population P29/f14 . . . . . . . . . . . . . . . . . . . . . 47
7.4 Discrimation Bolivinita urenuia : B. compressa . . . . . . . . . . . . . . . . . . . . . . . 48
7.5 Discrimination Bolivinita urenuia : B. pohana . . . . . . . . . . . . . . . . . . . . . . . . 50
7.6 Bolivinita urenuia Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
7.7 Bolivinita urenuia Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . . . 52

8 Bolivinita pohana Finlay 54

8.1 Bolivinita pohana Type Population Y18/f7479 . . . . . . . . . . . . . . . . . . . . . . . . 55
8.2 Bolivinita pohana Comparative Population Q18/f13 . . . . . . . . . . . . . . . . . . . . . 56
8.3 Discrimination Bolivinita pohana : B. compressa . . . . . . . . . . . . . . . . . . . . . . . 57
8.4 Bolivinita pohana Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
8.5 Bolivinita pohana Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . . . 59

9 Bolivinita pliobliqua Vella 61

9.1 Bolivinita pliobliqua Type Population T26/f6910 (Recollection) . . . . . . . . . . . . . . . 61
9.2 Bolivinita pliobliqua Comparative Population P29/f16 . . . . . . . . . . . . . . . . . . . . 64
9.3 Bolivinita pliobliqua Comparative Population P29/f18 . . . . . . . . . . . . . . . . . . . . 65
9.4 Bolivinita pliobliqua Comparative Population T26/f139 . . . . . . . . . . . . . . . . . . . 65
9.5 Discrimination Bolivinita pliobliqua : B. pohana . . . . . . . . . . . . . . . . . . . . . . . 65
9.6 Discrimination Bolivinita pliobliqua : B. urenuia . . . . . . . . . . . . . . . . . . . . . . . 66
9.7 Discrimination Bolivinita pliobliqua : B. pliozea . . . . . . . . . . . . . . . . . . . . . . . 67
9.8 Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
9.9 Bolivinita pliobliqua Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
9.10 Bolivinita pliobliqua Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . 70

10 Bolivinita pliozea Finlay 71

10.1 Bolivinita pliozea Reference Population R22/f7516 . . . . . . . . . . . . . . . . . . . . . . 71
10.2 Bolivinita pliozea Comparative Population P29/f29 . . . . . . . . . . . . . . . . . . . . . 73
10.3 Bolivinita pliozea Comparative Population J32/f44 . . . . . . . . . . . . . . . . . . . . . 74
10.4 Discrimination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
10.5 Bolivinita pliozea Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
10.6 Bolivinita pliozea Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . . . 77

11 Bolivinita medialis Scott n. sp. 79

11.1 Bolivinita medialis Type Population T27/f551 . . . . . . . . . . . . . . . . . . . . . . . . . 79
11.2 Bolivinita medialis Comparative Population T27/f495 . . . . . . . . . . . . . . . . . . . . 82
11.3 Discrimination Bolivinita medialis : B. pliozea . . . . . . . . . . . . . . . . . . . . . . . 83
11.4 Bolivinita medialis Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
11.5 Bolivinita medialis Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
11.6 Bolivinita medialis Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . . . 87

12 Bolivinita quadrilatera (Schwager) 88

12.1 Description of neotype . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
12.2 Bolivinita quadrilatera Reference Population T27/f460 . . . . . . . . . . . . . . . . . . . 89
12.3 Discrimination Bolivinita quadrilatera : B. urenuia . . . . . . . . . . . . . . . . . . . . . 91
12.4 Bolivinita quadrilatera Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
12.5 Bolivinita quadrilatera Stratigraphic Distribution . . . . . . . . . . . . . . . . . . . . . . 93

iv
13 Bolivinita quadrilatera granttaylori Vella 95
13.1 Bolivinita quadrilatera granttaylori Type Population . . . . . . . . . . . . . . . . . . . . . 95

DRAFT
13.2 Bolivinita quadrilatera granttaylori Biogeography . . . . . . . . . . . . . . . . . . . . . . 96
13.3 Bolivinita quadrilatera granttaylori Stratigraphic Distribution . . . . . . . . . . . . . . . 97

III Additional Studies 99

14 Bolivinita elegantissima Boomgaart 100

15 Bolivinita subangularis (Brady) 102

16 Bolivinita sp. aff. pohana striate 104

Appendices by M.P. Crundwell: New species of Bolivinita 106

A Bolivinita pseudocompressa Crundwell n. sp. 107

B Bolivinita striata Crundwell n. sp. 109

C Bolivinita obconica Crundwell n. sp. 111

Taxon Index 113

v
 DRAFT
Part I

Preamble

The scope, objectives, and methods of the study are outlined. Shell archi-
tectures are introduced and designs are compared. Taxon biogeography and
evolutionary history are summarized.

1
Chapter 1

DRAFT
Introduction

1.1 Overview

The appearance of several species of Bolivinita, including B. quadrilatera, near the base of the Late Miocene, and their
wide distribution subsequently, are major features of New Zealand Late Neogene benthic foraminiferal biogeography.
The biostratigraphic utility of Bolivinita was early recognized by ?, who described several species, and by ?, who cited
the entry of B. pohana as a primary foraminiferal event for recognition of the Tongaporutuan Stage. ? and ? named
additional species. ??? discussed variation in some species, and research on the genus was summarized by ? .

Primarily, the study examines the systematics of some Bolivinita that are useful in Late Neogene biostratigraphy.
Included are local populations of the globally-distributed B. quadrilatera (?), and species proposed by ?, B. compressa,
B. pliozea, B. pohana, ?, B. quadrilatera granttaylori, ?, B. pliobliqua, and ?, B. finlayi. Also studied are B.
pseudocompressa Crundwell n. sp., B. medialis Scott n. sp., and B. urenuia Scott n. sp.

To provide a fuller reference to New Zealand Bolivinita, original descriptions, with some additional data, of B. ele-
gantissima Boomgaart and B. subangularis (Brady) are included. Bolivinita pseudocompressa Crundwell n. sp., B.
obconica Crundwell n. sp. and B. striata Crundwell n. sp. are described in appendices.

Currently, the Late Miocene sequence in New Zealand is poorly resolved by foraminifera. That only two units (Ton-
gaporutuan, Kapitean Stages) are recognized for an interval of >5 m.y. (Fig. 1.1) justifies a search for more closely
spaced biostratigraphic events suitable as time proxies. In this view, Bolivinita merits attention because of its wide
dispersal in shelf through slope environments, and its significant diversity.

From another perspective, biostratigraphic events are only as robust as the systematics on which they are based. If
good dispersal and moderate diversity are potential merits of Bolivinita for the biostratigrapher then demerits are our
poor understanding of the variation of some taxa, their stratigraphy, and the criteria for species discrimination.

This study focuses on these latter topics. Some of the species considered here were only briefly introduced in the
original proposals. Little attention was given to intra- and inter-sample variation, and few specimens were illus-
trated. Here, more detailed accounts are given of shell morphology in standard orientations. Intra-sample variation
is described qualitatively and, for the most informative orientation (spiral), the accounts are supplemented by cluster
analyses and ordinations using Fourier coefficients for specimen outlines. Where possible, topotypic material is con-
sidered. However, as species are viewed as sets of variable, possibly polytypic populations, attention is given also to
samples that differ from the topotypic and add to the understanding of intra-specific variation. Because the primary
role of holotypes is that of name-bearer, not exemplar, they are not given special attention (?).

The primary aim is simply to aid the qualitative identification of taxa and to assess their utility in biostratigraphy.
Taxa are examined only at the basic level as groups of populations whose morphology is sufficiently unified to enable
their discrimation from others. The higher classification of these morphotaxa is not pursued. It will be apparent that
highly diverse morphotaxa have been referred to Bolivinita. A classification that reflects the evolutionary histories of
bolivinitid taxa requires a much wider survey than is reported here.

2
1.2 Perspective

DRAFT
Following ?, the focus is primarily on the outline of the shell and the shape of its major elements. These data are re-
garded as the most informative in qualitative visual perception. The shape of significant outline elements is described
in the text and the relevant features are identified on the accompanying images. The description is deliberately quali-
tative (e.g., "convex", "angular", "linear"), in keeping with the way taxonomists rapidly scan and compare specimens.
Metric data might improve recognition but are unlikely to be applied by taxonomists in routine work. The objective
is to guide the way in which the identifier views the material, and the descriptive technique reflects current levels of
instrumentation. This methodology will change as image databases become available and pattern recognition systems
are better integrated with microscopy.

Viewed as an exercise in pattern recognition, a principal problem of paleobiological taxonomy based on morphology
is the architectural variability of taxa in space and time. There may be typical individuals but there is no archetype
to serve as an arbiter of group membership. Thus morphologically, the boundaries of taxa are commonly difficult to
locate. To help resolve this problem, ordinations and cluster analyses were performed on data representing the spiral
outline of specimens. These data provide metrics for display of intra-sample variation and help identify outliers. As
pooled data they are applied to inter-sample comparisons and used as aids to assessment of disjunction.

1.3 Samples

Much of the material had been already mounted on assemblage slides by several paleontologists at GNS Science
and its predecessors, or for previous studies by ???. Some residues were re-examined for additional specimens.
Generally, the material is from the >200 µm fraction and, while not strictly random, is likely to be representative.
Where possible, intact specimens were used in the study of spiral outlines. Collection identifiers prefixed by A-Z are
registered in the New Zealand Fossil Record File (http://www.fred.org.nz). Well names and depths (in feet
or meters, according to the method of curation) are used for subsurface collections. All material is curated at GNS
Science, Lower Hutt.

1.4 Figures

Images were recorded on a Philips PSEM500 scanning electron microscope. Standard magnifications were used to
facilitate comparisons among taxa. Spiral views were recorded at a nominal magnification of x200; axial and distal
views were recorded at x100. Images were edited using PhotoStyler (Aldus Corporation). Figures were composed
with CorelDraw (Corel Corporation) and saved as high resolution .jpg files.

1.5 Spiral Outline Data

Specimen images were recorded digitally (800x600 pixel format, 128 grey levels) at a standard magnification (nom-
inally x200 on the monitor). Specimens were positioned in spiral orientation (Fig. 1.2). The aim was to record an
orthogonal view of the last-formed pair of chambers (i.e., normal to the direction of the coiling axis). This is possible if
the coiling axis is linear, and late-formed chambers expand in a geometric series. In some specimens departures from
these conditions lead to the outline being a composite of elements from several pairs of late-formed chambers. Manual
editing of images to eliminate outline elements derived from chambers preceding the last pair was not attempted.

Photostyler was used to adjust contrast in the recorded images, and to create a uniformly black background. The latter
step can be largely automated using a flood paint tool, and results in sharp definition of the specimen boundary when
the image is thresholded (converted to binary format). As the direction of coiling affects the shape of rhomboidal

3
 DRAFT
Figure 1.1: Stratigraphic ranges of Bolivinita spp. Correlation of New Zealand stages with the geomagnetic polarity
time scale (?) is based on ?, ? and unpublished data. ? provide an updated chronostratigraphy. It is available at
http://www.gns.cri.nz/content/download/10887/58333/file/SR2012-39/

4
taxon.
DRAFT
Figure 1.2: Standard orientations and terminology for Bolivinita quadrilatera granttaylori, a bilaterally-symmetrical

outlines, specimens coiling anticlockwise (spiral view, last chamber uppermost) were mirrored to the clockwise posi-
tion. Thresholding of images and extraction of outline x,y coordinates were carried out using NIH-Image (National
Institute of Health, USA). Coordinates were plotted in CorelDraw and outlines inspected and edited.

Thinned sets of c.100 - 200 coordinates per specimen were transferred to a stand-alone version of the elliptical Fourier
routine EFOUR in NTSYS v. 1.8 (Exeter Software). Specimen data were standardized for size and starting point. Five
harmonics were fitted to the data and the resultant 20 coefficients/specimen formed the data set for the presentation
of variation. Principal components analysis of the variance-covariance matrices was used for ordinations, which are
presented as 2 or 3 dimensional plots. In the cluster analyses average taxonomic distances were used to measure spec-
imen similarities. The sorting strategy employed was the unweighted pair-group method with arithmetic averaging.
Both were performed with NTSYS.

Fourier analyses are sensitive to the starting positions in the x,y data. A study of several of the data sets by ? indicates
that starting positions for the coordinate data were poorly standardized. This may have degraded the ordinations and
cluster analyses based on the Fourier coefficients and reduced their discriminatory resolution.

5
Chapter 2

DRAFT
Morphology

2.1 Overview

Chambers in Bolivinita are arranged in a biserial spiral, typically with each chamber larger than its predecessor (a
slowly expanding geometric series). A chamber partially overlaps its predecessor so that the series interfingers. The
overlap provides space for the toothplate, a principal internal structure (??) in the vicinity of the coiling axis, to run
between apertures.

Based on the symmetry of the shell in spiral orientation, species studied here fall into two primary architectural groups.
Bolivinita quadrilatera, B. medialis and B. pliozea have bilaterally symmetrical shells. Bolivinita quadrilatera is used
as an exemplar for this group (Fig. 1.2).

In a larger group, exemplified by Bolivinita compressa (Fig. 2.1) simple mirror-image bilateral symmetry is lost be-
cause of differentiation in the length, shape, and orientation of the axial walls of chambers. This group is characterized
in spiral orientation by its rhomboidal outlines. These are strongly developed in B. compressa, B. finlayi and B. pseu-
docompressa and, to lesser extents, in B. pliobliqua, B. pohana, and B. urenuia.

2.2 Significance of Spiral Orientation

In residue strews, most shells will be viewed at or near their natural positions of repose (axial in bilaterally symmetrical
taxa, oblique axial in rhomboidal taxa). Some taxa can be reliably identified in this position but it provides inadequate
information on others, especially those with rhomboidal architecture.

Several considerations justify the extra effort needed to mount specimens in spiral orientation. It is the best position
for observation of symmetry. It provides a plan view that includes the two axial walls as well as the distal wall of
chambers. The locations and amplitudes of keels and ribs are well defined in this view. Functionally, shell shape in
this orientation is critical for an infaunal organism tunnelling through sediment.

Although axial and distal orientations provides essential data on the ontogeny of individuals, some of these data are
a source of potential noise if, as in this study, the focus is on assessment of intra-sample variation and discrimination
of taxa. In these contexts, spiral orientation is preferable as a source of data as it represents morphology only of a
restricted (late ontogeny) age segment of individuals. Also, differences between megalospheric and microspheric shell
morphologies (Fig. 4.2 #6-#8) are much less apparent in spiral than in axial and distal orientations. This buffering of
ontogenetic and generational (megalospheric - microspheric) variation of specimens, when viewed in spiral orienta-
tion, makes it particularly valuable for both qualitative and quantitative descriptions of shape. Its principal demerit for
routine use is the effort required to mount specimens perpendicularly to their positions of natural repose.

Bolivinita quadrilatera - A Bilaterally Symmetrical Design

Significant elements

• Spiral orientation - bilateral symmetry, axial walls are mirror images

• Keel at each axial - distal wall junction

6
Spiral orientation: The view is normal to the coiling axis. In plan, the last-formed chamber resembles a quadrilateral,
with the axial edges inclined towards the coiling axis. The shell outline reflects the shape of the last two (opposed)

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chambers and varies from rectangular to octagonal, depending on the inclination of axial walls. Major chamber
components visible are the distal and axial walls, aperture of the last-formed chamber, and keels at junctions between
axial and distal walls. Axial walls of the opposed chambers are aligned to form the longer sides of the shell. Keels
protrude at wall junctions as ridges. They are constructed by close-folding of the chamber walls and are analogous
to crimps used to strengthen mechanical thin shells The aperture is an orifice for cytoplasm and is the termination of
the internal toothplate (?). This tube-like, folded plate is twisted spirally in sympathy with the 180o rotation between
successive chambers. It runs from aperture to aperture, through the chamber sequence. The tooth visible in the
aperture is the free edge of the plate (in cross-sectional view).

Axial orientation: The full ontogenetic record of individuals is visible and the interfingering nature of the chamber
sequence is shown. In axial orientation, so named because axial walls are viewed orthogonally, the coiling axis is in
a longitudinal position. Several components of the outline are discriminated. The prolocular section is adjacent to
the initial chamber. In Bolivinita quadrilatera the lateral sections are defined by the axial - distal wall junctions. In
some taxa they represent the crests of the distal walls of chambers (particularly if they are significantly convex), or a
composite of distal wall crests and axial - distal wall junctions. Lobulation refers to small lobes along the lateral section
that represent crests of the distal wall of individual chambers. The gross shape of the shell outline is largely determined
by the angular divergence of the lateral sections during growth. The size of the proloculus is an important influence.
In most species the lateral sections of shells with a small (microspheric) proloculus diverge through ontogeny at near
constant angles, so that maximum width is reached adjacent to the last pair of chambers. In specimens with a large
(megalospheric) proloculus, the initial divergence of the lateral sections often decreases in ontogeny and the sections
become almost parallel. The apertural section above the aperture joins the lateral sections. Ribs refer to narrow,
usually linear, calcified ridges that run along chambers.

Distal orientation: This view (Fig. 1.2) is obtained by rotating the shell 90o about the coiling axis from the axial
orientation. The coiling axis remains in a longitudal position. The orientation is termed 'distal' as the wall, which is
radially distant from the coiling axis, is in full view. This distal wall completes the chamber envelope by linking the
axial walls. Because chambers lie 180o apart, only distal walls of alternate chambers are visible. The aperture of the
last-formed chamber is shown in one of the two distal positions (Fig. 1.2). Sections of the shell outline equivalent to
those described for the axial orientation are recognized. Generally, shell outlines are narrower in this view as the view
is of single chambers rather than pairs. Contrasts due to variation in the size of the proloculus are muted.

Remarks: Critical features of the architecture of Bolivinita quadrilatera are similarities in the length and orientation
of the axial walls of chambers. The plan view of the shell (spiral orientation) is symmetrical about an axis through
the aperture. This bilateral symmetry is maintained in B. pliozea and B. medialis, although the shape of the chamber
envelope differs. Chambers in these taxa have axial walls of equal length and, although they are not parallel, they
usually lie at equal angles relative to the reference axis.

7
Bolivinita compressa - A Rhomboidal Design

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Figure 2.1: Standard orientations and terminology for Bolivinita compressa, a rhomboidal species.

Significant elements

• Spiral orientation - length, orientation of axial walls leads to rhomboidal spiral outline
• Keel at longer axial - distal wall junction
• Generally rounded junction between shorter axial and distal wall

Spiral orientation: Chamber design differs significantly from that in Bolivinita quadrilatera and leads to changes in
shell architecture. In this orientation (plan view) the basic innovation is that the axial walls differ in length, shape,
and orientation (Fig. 2.1). The longer axial wall meets the distal wall at an acute angle; there is a prominent keel at
this junction. The opposite axial wall is shorter and makes a curved junction with the distal wall. In some chambers
the junction is marked by a narrow arc in which curvature visibly increases; in others, the axial and distal walls form
a continuous curve and their junction is marked by the turning point. Generally in late-formed chambers there is no
keel, or remnant thereof, at this junction.

These modifications to the axial walls distort a rectangular shell outline to a grossly rhomboidal, sometimes alate,
outline (Fig. 2.1). A measure of the distortion is the offset between mid-points of the distal walls of the opposed
chambers with the shell in a standard orientation. Simple (mirror) bilateral symmetry about an axis through the
aperture is lost, but consecutive chambers are symmetrical when rotated through 180o . The major axis of the shell lies
through the opposed keels. ? used the ratio of the length of this axis to one intersecting the junctions of the shorter
axial - distal walls as a measure of distortion ('obliquity').

Axial, distal orientations: One axial face and one distal face are seen when Bolivinita compressa is viewed at its
natural angle of free repose on a planar surface; the position is identified as oblique axial. The view compares with
other rhomboidal objects resting on one of their wider faces. A strictly axial orientation, comparable to that for
bilaterally symmetrical shells, is obtained by rotating the shell to bring the profiles of both distal walls into view. Distal
orientation is obtained by further rotation to bring the profiles of both axial walls into view (Fig. 2.1). Rhomboidal
taxa are described in these strictly axial and distal orientations as they allow the discrimination of the same elements
visible in bilaterally symmetrical taxa.

8
Chapter 3

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Interpretation

3.1 Shell Design

As an introduction to terminology, two basic designs, bilaterally symmetrical and rhomboidal, were identified. Al-
though any individual can be classified in this way, the systematic data indicate that the designs are not always strictly
exclusive at the population level. Particularly, taxa regarded as possessing bilateral symmetry (e.g., Bolivinita pliozea,
B. quadrilatera) often have individuals that show small departures from this plan. Similarly, a weakly rhomboidal
species like B. pliobliqua has some individuals that are nearly bilaterally symmetrical. By contrast, such specimens
are not found in more strongly rhomboidal taxa such as B. compressa. These data suggest that the designs are not
exclusive and raise the possibility of transformations between them. Perhaps there is a spectrum in spiral design with
taxa showing adaptive preferences for a particular architecture.

Figure 3.1: Comparison of quadrate, bilaterally-symmetrical shells. Other characters in common are re-entrants
formed by axial walls and keels at axial-distal wall junctions. Note comparative width of chambers in Bolivinita
quadrilatera. Bolivinita quadrilatera- T27/f460, Wairarapa, Bells Creek Mudstone. Loxostomum truncatum-
D45/8670, Clifden, Southland. Plectofrondicularia parri- J41/f9499, Oamaru, Rifle Butts Formation.

3.2 Comparative Morphology

In principle, the stratigraphic record of Bolivinita is the primary guide to the evolutionary development of the principal
designs. However, in view of the imperfect record presently available, it is useful to consider the designs, compara-
tively. For this purpose three taxa are taken as exemplars. Bolivinita quadrilatera represents a rectangular, bilaterally
symmetrical design, B. pliozea represents an oval to sub-circular, bilaterally symmetrical design and B. pseudocom-
pressa represents an advanced, rhomboidal design. The focus is on the spiral orientation of the shell. Comparisons
are made principally, but not exclusively, within the Bolivinitidae, particularly Bolivina.

9
 DRAFT
Figure 3.2: Oval to sub-circular spiral outlines in Bolivinita. Circular form is closely approximately by B. elegan-
tissima. Comparable outlines are common in Bolivina and many uniserial and biserial taxa. Re-entrants formed by
inwardly inclined axial walls interrupt the oval to sub-circular outlines in Bolivinita pliozea and B. subangularis.
Bolivinita elegantissima- T27/f122, Wairarapa, Bells Creek Mudstone. Bolivinita subangularis- D45/f8646, Clifden,
Southland. Bolivinita pliozea- R22/f7516, Wanganui, Pinnacle Sand.

Bolivinita quadrilatera

The quadrilateral to rectangular spiral form of Bolivinita quadrilatera is unusual in Bolivina and Loxostomum. There
are no close analogues in ?. Quadrate form, with stout keels at junctions of axial and distal walls, is developed in the
early ontogeny of Loxostomum truncatum, but the width of chambers is relatively much narrower than in B. quadrilat-
era. Interestingly, chambers junctions become rounded and keels atrophy in late ontogeny. Beyond the Bolivinitidae,
narrowly quadrate spiral profiles, again with a keel at each wall junction, occur in some Plectofrondicularia (e.g., P.
parri). In this lagenid genus chambers are arranged uniserially for much of ontogeny.

In overview, quadrate designs are rare in the Bolivinitidae, and relatively rare among other biserial and uniserial benthic
taxa. However, Bolivinita quadrilatera shares features with its analogues: rectangular shape in spiral orientation,
shallow re-entrants formed by axial walls, and angular junctions between axial and distal walls, often with a keel.
Primarily, Bolivinita quadrilatera is distinctive in its use of this type of design in large, wide chambers.

Bolivinita pliozea

Disregarding the re-entrants defined by the axial walls, the gross outline of the shell is oval to sub-circular (Fig. 3.2,
Fig. 15.1). This form is created primarily by the convex curvature of the distal wall. Similar oval to sub-circular
profiles are quite common in Bolivina (e.g., B. acerosa, B. advena, B. silvestrina). Bolivinita elegantissima (Fig. 14.1)
is another example. In all of these taxa the chamber envelope is a continuous curve; axial and distal segments are not
discriminated.

The bounding ellipse over the spiral outline of Bolivinita pliozea approaches a circle on some specimens. This shape
is generated solely by the strongly convex distal walls; it is considerably interrupted (often >120o of arc) by re-entrants
formed by the inward inclination of the axial walls. Chamber walls in many Bolivina are convex and a shallow median
depression is formed along chamber junctions. The re-entrant in Bolivinita pliozea is set apart from these by its
much greater width and depth, and by the almost planar surfaces of axial walls, which usually abut to create a regular
channel. Much closer to Bolivinita pliozea than any Bolivina is Bolivinita subangularis (Fig. 3.2). However, in this

10
species the distal wall, although elevated as in B. pliozea, is not uniformly curved but is formed from two angular
facets.

DRAFT
Bolivinita pliozea may be interpreted as a partial implementation of the circular spiral design, which is common in
Bolivina. The quasi-circular profile is achieved by inflating the distal wall and inward tucking of the axial walls.

Figure 3.3: Alate spiral form in Bolivinita (bilaterally asymmetrical) compared with Bolivina alata (bilaterally sym-
metrical). Axial, distal walls weakly distinguished in last chamber of Bolivinita pseudocompressa, not identifiable
in Bolivinita sp. or Bolivina alata. Compare disposition of aperture and toothplate in Bolivinita sp. with Bolivina
alata. Bolivina alata- #9000, Kumai Dome, Koramasarik, west of Madang, PNG, upper Ouba Formation. Bolivinita
sp.- F100155 H.S. Ladd station 371, Suva, Fiji, Suva Soapstone. Bolivinita pseudocompressa- T27/f78, Wairarapa,
Mangaopari Mudstone.

Bolivinita pseudocompressa

Bolivinita pseudocompressa and B. compressa construct rhomboidal shells by differentiating the length and orientation
of the axial walls of chambers (Fig. 3.3). By this means the major axis of the spiral profile through opposite keels is
significantly increased relative to the minor axis through the alternate chambers. Relative to the sharply rectangular,
box-like spiral profile of B. quadrilatera, these taxa have compressed profiles, acutely angular at the extremities of the
long major axis but curved near the extremities of the minor axis.

Narrow, elongate spiral profiles that are compressed progressively towards the periphery are common in Bolivina.
They are termed 'alate' after B. alata, an advanced example. In alate shells the wall segment equivalent to the distal
wall in Bolivinita is completely eliminated and the axial walls meet acutely, sometimes with a flange at the junction.
The greatest spiral diameter of alate Bolivina normally runs between the extremities of the axial walls and passes
through the aperture. The profile is symmetrical about this axis. The minor axis of the profile, considered as an
ellipse, is in the vicinity of the aperture.

Although the spiral profile of some Bolivinita pseudocompressa is compressed towards the periphery and alate, several

11
components differ significantly from alate Bolivina. A distal wall element is distinguishable in many specimens. Axial
walls differ in length and sometimes orientation. Although the greatest spiral diameter is similarly located, it does not

DRAFT
define an axis of bilateral symmetry as in Bolivina.

While rhomboidal taxa like Bolivinita pseudocompressa have gross spiral profiles resembling alate Bolivina, and are
close analogues, some of their architectural elements are substantially different. The incomplete fusion of axial and
distal wall elements may be a clue to the ancestry of rhomboidal Bolivinita. Speculatively, rhomboidal form might be
a strategy by which a rectangular profile, as in B. quadrilatera, is transformed into an alate profile.

3.3 Modelling

Although the wall elements involved in a transformation from one design to another are readily identified, it is useful
to model the transformation using digital image morphing techniques that resolve intermediate stages. SPMORF (S.H.
Pultz, 114-168th Ave NE, Bellevue WA, USA) uses landmarks regarded as homologous on source and target images
as inputs to spline functions, which map the transformation. Colour interpolation progressively fades out the source
image and fades in the target image. Mid-points on keels, axial and distal walls, and points at the the base and apex
of the aperture were identified on typical specimens of Bolivinita quadrilatera and B. pliozea (Fig. 3.4 #a,#e) and
used as landmarks for the progressive transformation of a rectangular to a sub-circular profile. The principal feature
(Fig. 3.4 #h) is inward movement of the axial walls to create progressively deeper re-entrants. The curvature of these
walls decreases slightly. Convexity of the distal walls mainly increases in the final stages of the transformation.

Figure 3.4: Transformation of Bolivinita quadrilatera (spiral orientation, #a) to Bolivinita pliozea (#e) using landmarks
with interpolation by spline functions. #b-#d are computed intermediate morphs. For comparison are specimens of
Bolivinita medialis (#f, #g) that resemble #c-#d. #h summarizes the deformation pattern.

Of interest is the form of the intermediate morphs and their correspondence with real specimens. In the source
population (T27/f460, Fig. 12.2) there are specimens with slightly convex distal walls and weakly concave axial walls
like Fig. 3.4 #b, which is an early morph. More advanced morphs like Fig. 3.4 #c-#d occur in neither source nor
target populations but have some similarity with B. medialis from T27/f551 (Fig. 3.4 #f-#g). In these specimens the
re-entrant formed by axial walls is greater than in B. quadrilatera, as is the convexity of distal walls.

Although there are more informative tools now available for analysis and portrayal of shape deformations (e.g.
https://cran.r-project.org/web/packages/shapes/shapes.pdf), the model suggests that a re-
alistic transformation between exemplars of the principal designs can be obtained by progressive movement of walls,

12
particularly the axial walls, and requires few landmarks. Although the transformation appears to be simple, it does not
imply that an evolutionary path is mapped. Stratigraphic data suggest that B. pliozea descended from B. pliobliqua, a

DRAFT
species in which convexity of distal walls was already well developed. The principal architectural change in the evo-
lution of B. pliozea was development of bilateral symmetry from a weakly rhomboidal predecessor by modification
to the geometry of axial walls of chambers. It did not involve deformation of existing bilaterally symmetrical archi-
tecture. In an evolutionary context, it is inappropriate to model the architectural development of B. pliozea directly
from B. quadrilatera. However, it is of interest that some morphs compare well with the bilaterally symmetrical B.
medialis. Convexity of distal walls is quite variable in this species and there are individuals (Fig. 11.4 #3) in which
convex walls rise only slightly above well developed keels. Such morphotypes resemble B. quadrilatera. Whether
they represent inherited features and provide clues to ancestry is unknown. No stratigraphic data on the evolution of
B. medialis were found in this study.

3.4 Functional Morphology

In the absence of useful data on Bolivinita ecology, the functional significance of the exemplar designs is highly spec-
ulative. Some analogies are again provided by Bolivina. The great majority of its taxa are infaunal and burrow in
the upper few centimeters of sediment. Infaunal foraminifers are mobile organisms (?) and show statistical, rather
than absolute, preferences for particular microhabitats (?). The decline in oxygen levels below the sediment interface
is an important control on the vertical distribution of the foraminiferal infauna. However, some Bolivina are oppor-
tunists (??) and flourish in dysoxic environments eschewed by equilibrium taxa. These data may be relevant to the
understanding of several Bolivinita designs.

Many uniserial and biserial infaunal taxa in addition to Bolivina have oval to sub-circular spiral profiles (?). Movement
of infaunal foraminifers through sediment is still poorly understood (??) but in some taxa it is achieved by tunnelling
ahead with pseudopodia to clear a cavity into which the shell is then dragged. No data were found on the relationship
between tunnel shape and the spiral shape of the shell but they are likely to be correlated. A quasi-circular cross-
section is an efficient design for constructing a stable tunnel that is resistant to deformation by the overload. This
shape also minimizes its cross-sectional area and the required energy to build it. From this interpretation, the partially
realized sub-circular spiral profile of Bolivinita pliozea represents an adaptation which facilites shell movement within
sediment. The spiral profile of B. elegantissima (Fig. 14.1) approximates the circular paradigm more closely than B.
pliozea but it may not be a close relative. ? suggested that its placement in Bolivinita is uncertain.

The area:perimeter relation of an alate shell (e.g., Bolivinita pseudocompressa (Fig. 3.3) is the opposite to that of
a circular shell. While a narrow, elongated cross-section might assist tunnelling in, say, laminated sediment, more
important may be the extension of shell area that is obtained. Pores allow gas exchange through the mineralized shell
wall (?). Extension of wall area, and hence number of pores relative to chamber volume, will improve the exchange.
This might be important as foraminifers tunnel into progressively more dysoxic environments below the sediment
interface. Bolivina alata is identified as a deep infaunal species by ?, who found that it was abundant in low diversity
assemblages from oxygen-poor environments. Although not all species exploiting dysoxic environments have alate
shells, the inequitable structure of some assemblages containing rhomboidal Bolivinita suggests that, like Bolivina
alata, they were successful in such environments.

Review

Relative to Bolivina designs, sub-circular and alate profiles are only partially realized by the Bolivinita exemplars
considered here. These retain wall elements e.g., re-entrants formed by axial walls in B. pliozea and remnant distal
walls in B. pseudocompressa, which depart from the simple circular and alate designs found in Bolivina. From this
perspective, the implementation of sub-circular and alate designs in some Bolivinita may have been constrained by
architectural elements in ancestral taxa. Nevertheless, modelling suggests that quite simple changes in the shape and
orientation of chamber walls are required to transform substantially the spiral profile of a quadrate shell. This plasticity
may have contributed to the variability apparent in preserved populations, facilitated their adaptation to specialized
niches, and the evolutionary diversification of the group.

13
Functional interpretation of the shell features of Bolivinita quadrilatera requires research on the ecology of its modern
populations; there are no close analogues in other groups for comparison. However, sub-circular and alate designs in

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Bolivinita are more amenable to interpretation via comparative morphology. They appear to be partial implementations
of designs more fully developed in Bolivina. Their spiral profiles are consistent with those commonly developed in
infaunal biserial foraminifera.

3.5 Biogeography

The geographic distribution of each species in the main study is noted in the descriptions and summarized graphically
(Fig. 3.5). The species distributions are sketched as cartoons. As such, they are inexact and likely to significantly
underestimate actual distributions. Their extension beyond New Zealand is not considered.

• The outstanding feature is the entry of Bolivinita compressa, B. medialis, B. pohana and B. quadrilatera in New
Zealand within a short interval near the base of the Tongaporutuan Stage (Late Miocene, c.11 Ma; Fig. 1.1). As no
potential ancestors are known in the region the taxa are regarded as migrants, probably from the tropical province
where ? reported the evolutionary appearance of Bolivinita quadrilatera in the Globorotalia fohsi robusta Zone (Mid-
dle Miocene, c.12.3-11.9 Ma; ?) Factors promoting southward dispersal of the taxa are speculative. They seem not
to have been accompanied by other taxa in the Foraminifera, or indeed in the wider biota (?). Paleoclimatically,
the migration occurred within a long interval of progressive cooling following the Middle Miocene Climatic Opti-
mum. However, about the time of the postulated migrations ? mapped an intense warm event of global significance
(c.10.7-10.8 Ma) in the deep ocean at ODP Site 1146 (west Pacific Ocean). Regionally, changes in upper water mass
hydrography at this time may be signalled by the change in coiling direction in Globorotalia miotumida (Kaiti coiling
zone). Nevertheless, if they were significant for the dispersal of Bolivinita, these events apparently had little effect on
the dispersal of other biota. That is the enigma.

• Species that appeared near the base of Tongaporutuan Stage (Bolivinita compressa, B. medialis, B. pohana, B.
quadrilatera) are most persistent in the Eastern Basin. It is inferred that their major populations were in that basin.

• Of the taxa identified above, Bolivinita compressa, B. pohana and B. quadrilatera occur in Tongaporutuan strata
in Taranaki Basin but all are significantly less persistent there than in the Eastern Basin, and populations sizes are
commonly smaller. As yet, Bolivinita medialis has not been identified from Taranaki Basin. All of the taxa were
primarily bathyal to abyssal dwellers. Bathyal biotopes are extensively developed in both basins in Tongaporutuan
time and there is no macro-environmental explanation for the inter-basin contrast. Detailed comparison of assemblage
compositions in the basins might help resolve this problem.

• From latest Miocene (Kapitean Stage) to the Pleistocene Bolivinita pliozea was persistent and widely distributed in
all basins. In part, its high record reflects the wider distribution of upper bathyal and shelf sediments in the Pliocene
and Pleistocene than in the Late Miocene. Bolivinita pliozea flourished in these environments. However, the species
is unique in its rather uniform occupancy of all marine basins. The contrast in population density between Eastern
and Taranaki Basins shown by most taxa is not seen in the distribution of B. pliozea. No other species achieved a
comparable distribution, yet it seems unlikely that it survived the Pleistocene.

• Among factors influencing biogeography are adaptive range and population strategy. Bolivinita pliozea occurs in
moderately diverse benthic foraminiferal assemblages in which taxa are typically quite evenly distributed, numerically.
It is seldom dominant or even sub-dominant. In these respects it is identifiable as an equilibrium species (?); Scott
in ?, fig. 37. In contrast, B. compressa and to a lesser extent B. pohana, occasionally dominate assemblages. Such
blooms are a feature of opportunist species that can expand population sizes to take advantage of unusual resource
scenarios. An example is the ability of some Bolivina to thrive when oxygen availability falls below levels survivable
by most equilibrium species. The data suggest that rhomboidal Bolivinita might act more opportunistically than
bilaterally symmetrical Bolivinita. However, neither group seems to occur in very low diversity, strongly inequitable
assemblages in the manner of some Bolivina (?).

14
3.6 Evolutionary History

DRAFT
The preserved record of Bolivinita in New Zealand is extensive and is regarded as the most informative source of data
on its evolutionary history. Considered here is the Late Miocene - Recent history of the taxa reported in the main
study. Bolivinita subangularis (Fig. 15.1) and B. elegantissima (Fig. 14.1) are recorded from Middle Miocene strata
but their phyletic connection with B. quadrilatera and allied taxa is unresolved.

Fig. 3.6 shows several reconstructions developed from stratigraphic data; they reflect improved understanding of
species variation and distributions. Significant features of the current interpretation are noted.

• Bolivinita compressa, B. medialis, B. pohana and B.quadrilatera appeared in New Zealand near the base of the
Tongaporutuan Stage (Late Miocene, c.11 Ma; Fig. 1.1). As no potential ancestors are known in the region the taxa
are regarded as migrants. They represent the major architectures found subsequently. The nature of their appearance
suggests that that the group had diversified significantly prior to its entry into the New Zealand region and that we have
only a partial record of its history. ? reported the evolutionary appearance of B. quadrilatera from B. elegantissima in
the Globorotalia fohsi robusta Zone (Andaman Islands). ? dated this zone between 12.3 -11.9 Ma. But in the same
province ?, Table 13 recorded it from Zone N4 (c.21-23 Ma) at ODP Site 715A. Research is required to reconcile
these disparate records. If confirmed, the extended Miocene record of B. quadrilatera mapped by ? in the tropical
Indian Ocean would be strong evidence that it was a founder taxon.

• Architectural innovation within Bolivinita is minor. Rather than diversification of morphology, the pattern is
one in which a few basic designs either persist or are repeated. This scenario poses problems for the assessment of
ancestry. Character convergences severely constrain cladistic methods while questions about the completeness of the
preserved record constrain reconstructions based on stratigraphy. Simple cladistic analysis might resolve B. pliozea
as a descendant, or a sister taxon, of B. medialis because of the large number of characters in common, but did these
characters evolve uniquely? Stratigraphy shows that the taxa are well separated in time, but what is the fidelity of
the record? In this instance, parallel evolution (acquisition of very similar structures by related species) is inferred
because of characters present in B. pliobliqua, which immediately preceded B. pliozea and was probably its ancestor.

• As Fig. 3.6 shows, little is known about speciation processes. Cladistic (lineage splitting) events, if present, are
poorly defined in the record. Bolivinita compressa extended into the Pliocene, after the appearance of the similarly
compressed rhomboidal species B. finlayi and B. pseudocompressa. Possibly they originated from B. compressa via
lineage bifurcations. In contrast, B. pliozea is inferred to have evolved via phyletic anagenesis from B. pliobliqua, an
immediate predecessor. Bolivinita pliobliqua includes a spectrum of weakly rhomboidal forms that are architectural
prototypes of the bilaterally symmetrical B. pliozea. The absence of B. pliobliqua following the appearance of B.
pliozea supports the latter's origin by phyletic transformation rather than divergence.

• It is overly simple to assume that all originations following the basal Tongaporutuan migrations are speciation
events. Later migrations may have occurred and knowledge of the history of Bolivinita in adjoining regions is neces-
sary for accurate reconstruction of its evolutionary history in New Zealand.

• Of the taxa studied, generally 3-4 were present in New Zealand at any one time. Possibly, gross diversity was almost
static (but recall that the present study is not comprehensive). Rather than a regional radiation (?), the pattern was
one of maintenance of a limited set of architectures, either by long-lived species or by replacements. The longevity of
some species (Fig. 1.1) suggests evolutionary stasis in well-adapted taxa. Although there is significant and possibly
complex variation at the species level, this did not lead to diversification.

• Minor innovation in architecture, some long-lived species, static diversity, and broad spectrum rather than special-
ist niche occupancy are macroevolutionary features that might identify Bolivinita as an equilibrium or "status quo"
strategist.

• However, this overlooks its two-phase history in New Zealand. Rhomboidal taxa, particularly B. compressa, were
often dominant in the Tongaporutuan Stage and, to a lesser extent, in the Kapitean Stage. They became a minor
component of Pliocene - Pleistocene assemblages. Their place was taken by the bilaterally symmetrical B. pliozea

15
whose unique biogeography has been noted. The changeover from "Miocene" to "post-Miocene" assemblages seems
to have occurred in the Kapitean Stage, a correlative of the latest Miocene Messinian Stage. The complexity of

DRAFT
paleoclimatic events in that interval, which included a change in carbon cycling (?), precludes useful speculation
about their effects on the evolutionary history of Bolivinita.

Due to its restricted focus, this study provides only a partial conspectus of Bolivinita evolution and little about its
mechanisms. Interior morphology, particularly of the toothplate, is not considered. Many other features of the local
record of Bolivinita require further study and documentation. They include the sources and timing of the original mi-
grations, environmental factors facilitating these migrations, explanation of convergences in shell design, and reasons
for significant intra-specific variation yet only limited diversification.

16
 DRAFT
Figure 3.5: Distributions of Bolivinita spp. on modern coordinates. The mappings (shaded areas) are generalizations
of known distributions and are conservative. They do not identify possible changes in distribution of species through
the Late Neogene. Most species occupied all or parts of several basins. Basins are shown as cartoons on maps for
B. finlayi and B. medialis. Connections between basins are implicit in the distribution of most species but inter-basin
migration routes are not shown because of the magnitude of the time slice (c.11 m.y.) and the need for multiple
palinspastic maps.

17
 DRAFT
Figure 3.6: A comparison of evolutionary histories for Bolivinita inferred from stratigraphic distributions and mor-
phology. Although the order of Bolivinita events in the Tongaporutuan Stage shown in ? and ? is preserved, their
relation to those of the present study is approximate. Particularly, the relation of Bolivinita spp. to the highest occur-
rence of Globoquadrina dehiscens was not considered in the earlier studies. Here, this event is used to partition the
stage into informal lower and upper zones.

18
 DRAFT
Part II

Taxonomy

Principal taxa are described, illustrated and compared. Their biogeographies and
stratigraphic distributions are summarized.

19
Chapter 4

DRAFT
Bolivinita compressa Finlay

1939 Bolivinita compressa Finlay, Transactions of the Royal Society of New Zealand 69: p. 319, pl. 27, fig. 101-2.

Original Description

"..wide extremely compressed shell (one side angle running almost up middle of front), absence of sculpture on sides,
and slightly flanged keels; size 1.4 mm." (Finlay 1939, p. 319)

4.1 Bolivinita compressa Reference Population X18/f38

Figure 4.1: Bolivinita compressa Finlay. Variation in the spiral outline of specimens from X18/f38. Two specimens
identified as Bolivinita pliobliqua Vella from this locality are included.

Material

The study of spiral morphology is based on imagery of 20 specimens from X18/f38. Observations on axial and distal
morphology are based on c. 120 specimens. X18/f38 is in the vicinity of G146 (?), c. 650 m below the base of
Opoiti Limestone in Hangaroa River section. X18/f38 is classified as upper Tongaporutuan as it lies above the LAD of
Globoquadrina dehiscens. The type locality of Bolivinita compressa (X18/f7466 F5018) is shown on a map (compiled
by Vacuum Oil Co Ltd 25 April 1934, held in Micropaleontology Laboratory, GNS Science) about 1600 m east of
X18/f38. The entry for X18/f7466 in the Micropaleontology Laboratory sample register states that the horizon is
c. 120 m below Opoiti Limestone. The accompanying specimens of Bolivinita pliobliqua more closely resemble B.
pliozea than do those in X18/f38; it probably represents a higher level in upper Tongaporutuan, and may be near the
boundary with the Kapitean Stage.

20
Spiral Orientation X18/f38

DRAFT
Chamber arrangement: Chambers are always offset (Fig. 4.1 #18) and the shell outline is rhomboidal. The major
axis of the shell lies through the keels at the junction of the longer axial and distal walls and, for a given orientation,
there is rotational symmetry at 180o intervals.

Distal wall: The boundary adjacent to the longer axial wall is sharply defined by the keel. At the opposite end, the
distal wall usually merges with the shorter axial wall in a continuous curve (Fig. 4.1 #17) but the junction is sometimes
angular (Fig. 4.1 #4). The outline of the wall is weakly convex and often is little elevated above the keel (Fig. 4.1
#18).

Axial walls: The longer wall is adjacent to the keel and has a linear outline. The profile of the shorter wall is variable.
Commonly, it is slightly convex (Fig. 4.1 #18) and merges outward with the distal wall. Some chambers appear
bulbous when both distal and shorter axial walls are convex. The re-entrant formed in the outline by opposed axial
walls is typically shallow, although wide. Commonly, it is asymmetrical, due to the differing lengths of the axial walls.
Occasionally, there is no re-entrant; this enhances the rhomboidal appearance of some shells (Fig. 4.1 #6, left side).

Keels: There is a prominent, often strong, keel at the junction of the longer axial wall and the distal wall. The keel runs
via the aperture to the equivalent structure on the opposed chamber. Generally, this keel lies along the major axis of the
shell. In spiral orientation, a keel is not visible at the junction of the distal wall with the shorter axial wall. However,
an angular junction between these walls (Fig. 4.1 #4) may be associated with a weak, sometimes discontinuous, keel
at this position on preceding chambers (visible in axial orientation).

Aperture: In most specimens apertural morphology is poorly preserved. In some, the opening is an asymmetrical
A-shape, with the side adjacent to the keel curving over to parallel the latter.

Axial Orientation X18/f38

Shell outline: The prolocular section is broadly rounded in some megalospheric specimens (Fig. 4.2 #1). Its shape
is affected by keel morphology. The lateral sections are defined by the distal wall outlines and are lobulate; the keel
is only slightly inset from one of the lateral sections. In most specimens the lateral sections diverge at a low angle
throughout growth but they may become almost parallel (Fig. 4.2 #1). The lateral sections taper to a narrow apex
above the aperture.

Chambers: There are up to 8 pairs of moderately inclined, wedge-shaped chambers. The number is often greatest
in microspheric specimens. Chambers inter-digitate laterally at low angles. They are often close-packed throughout
ontogeny but occasionally there is a lumen (seen with light microscopy) between late-formed chambers. The surface
of the longer axial wall is almost planar; the shorter axial wall is often convex, particularly where it merges with the
distal wall. Sutures are depressed. There is a strong keel at the distal end of the longer axial wall. It tapers outwards
to a thin, wide flange, best observed in oblique axial orientation (Fig. 4.2 #3). Some specimens have a much weaker
keel (without a flange) at the junction of the shorter axial and distal walls on early chambers. It seldom persists onto
the last pair. Often there are very weak ribs on the proloculus which may extend over the earliest chambers. There are
no ribs on later chambers. Pores are poorly preserved in this sample.

Distal Orientation X18/f38

Shell outline: Usually, the prolocular section is narrower than in axial orientation but its shape is similarly affected
by prolocular diameter and keel development (Fig. 4.2 #4). The keel sharply defines one lateral section; it is non-
lobulate. The opposite lateral section is defined by the turning point of the curve joining the shorter axial wall with
the distal. It may be slightly lobulate. If a keel is present at this site, definition of the outline is sharper and lobulation
is dampened. Angular divergence of the lateral sections is slight and they appear almost parallel in the late ontogeny
of some specimens (Fig. 4.2 #5). The apertural section is almost linear in some specimens.

21
tions.
DRAFT
Figure 4.2: Guide to morphology of specimens from X38/f38 and T27/f399 in axial, oblique axial and distal orienta-

Chambers: Height increases much more rapidly than width during ontogeny. The lower margin of a chamber is almost
linear but appears to curve upward at its junction with the shorter axial wall. At this location the lateral margins of
chambers are moderately convex (generating lobulation in the shell outline). The opposite margin (adjacent to the
keel) is less convex. Chamber surfaces are convex, particularly in late ontogeny, and sutures are depressed. Chamber
surfaces are featureless, apart from weak ribs on the proloculus and earliest chambers.

4.2 Bolivinita compressa Comparative Population T27/f399

Material: The locality is c. 90 m below the top of Bells Creek Mudstone in Ngawaka Creek, southern Wairarapa. It
is classified as lower Tongaporutuan as the assemblage includes Globoquadrina dehiscens. Twenty-four specimens
were studied in spiral orientation; c. 30 in axial and distal orientations.

Spiral orientation: The sample is dominated by specimens in which the shorter axial and distal walls merge in a curve.
There is a wide and prominent keel at the junction of the distal wall with the longer axial wall. Occasionally, the distal
wall is broadly convex and axial walls are divergent, producing a bulbous chamber (Fig. 4.3 #4). These features are
more strongly developed than in X18/f38 and axial re-entrants tend to be better defined, and deeper. The sides of the
aperture are curved and meet in a broad arch. In some specimens they are symmetrical (Fig. 4.3 #18); in others they
are asymmetrical, and flexed to follow the curvature of the adjacent keel (Fig. 4.3 #17).

Axial orientation: On a few specimens there is a weak keel at the junction of the shorter axial and distal walls of early
chambers. It does not extend up to the last pair of chambers. Its absence there may be related to the more bulbous
outlines (spiral orientation) of late-formed chambers. Oval pores are irregularly distributed over wall surfaces. On

22
 DRAFT
Figure 4.3: Bolivinita compressa Finlay. Variation in the spiral outline of specimens from T27/f399.

Figure 4.4: Bolivinita compressa T27/f399. Wall topog-

raphy.

early chambers they are raised as low domes and partially occluded (Fig. 4.4 #13).

4.3 Bolivinita compressa Comparative Population P29/f11

Material: The horizon is in Upton Formation, c. 200 m above basement in Blind River, Marlborough (?) and is
classified as upper Tongaporutuan. The magnetostratigraphy (after ?) is shown in Fig. 4.5. ? identified Bolivinita cf.
pohana from this sample.

Spiral Orientation: Intra-sample variation is greater than in T27/f399. Several specimens ( Fig. 4.6 #1, #12) probably
represent Bolivinita urenuia (p. 44). Specimens like Fig. 4.6 #5, #16 compare with many in T27/f399 (e.g., Fig. 4.3
#6) as the curved junction of the shorter axial wall with the distal wall creates a bulbous chamber, with one prominent
keel. However, in the majority of specimens, there is a narrowly rounded junction between these walls (Fig. 4.6 #8,
#15). Rarely, there is a very low keel (Fig. 4.6 #2) at this position.

Specimens from P29/f11 intermingle with those from T27/f399 in a cluster analysis and ordination of the combined
data on spiral outlines (Fig. 4.10 (B-C)). This suggests architectural similarity, although the data possibly do not
resolve adequatedly the detail of the junction between the shorter axial and distal walls. Relative to material from
X18/f38 (e.g., Fig. 4.1 #6), some specimens from P29/f11 tend to be less elongated (e.g., Fig. 4.10 (A) #C). However,

23
variation in the shape of shorter axial and distal walls, and in the shape of the re-entrant formed by axial walls, is
similar.

DRAFT
Figure 4.5: Bolivinita spp., magnetostratigraphy, geochronology, and biostratigraphy of the lower part of the Blind
River section, Marlborough. Column, magnetostratigraphy after ?; time scale from ?.

Axial orientation: The keel at the junction of the longer axial and distal walls is prominent and commonly tapers to
a thin, translucent flange. Development of a keel at the shorter axial and distal walls is variable. It is absent from
chambers in which these walls merge in a curve, and this architecture sometimes occurs throughout ontogeny. In
others there is a weak keel (without a flange) that extends through to the base of the last chamber. It is not visible in
spiral orientation, but may be indicated there by an angular junction between the distal and shorter axial walls.

4.4 Bolivinita compressa Comparative Population T26/f139

Material: About 70 m below the top of Bells Creek Mudstone, Te Kowhai Stream, southern Wairarapa (? appendix
1 section 2), upper Tongaporutuan. Nine specimens were studied in spiral orientation and 5 in axial orientation. The
assemblage also includes Bolivinita pliobliqua and B. quadrilatera. Most specimens are readily allocated among the
taxa, although the identity of individuals like Fig. 4.7 #8 is equivocal.

24
 DRAFT
Figure 4.6: Bolivinita compressa Finlay. Variation in the spiral outline of specimens from P29/f11. Also shown are
specimens tentatively identified as Bolivinita urenuia from this assemblage.

Spiral orientation: Variation in the small sample is comparable with that in P29/f11, with the junction between the
shorter axial and distal walls varying between rounded and angular (Fig. 4.7 (A) #20, #22). The keel at the junction
between the longer axial and distal walls is robust and tapers to a wide flange on some specimens (Fig. 4.7 (A) #20).
Outlines are less consistently elongated than in X18/f38 and, as in P29/f11, there are compact forms (Fig. 4.7 (A)
#24).

Axial orientation: There is a weak keel at the junction between shorter axial and distal walls on the early chambers
of most specimens but it dies out in later ontogeny.

4.5 Overview of Variation

Commonly, the junction between the shorter axial and distal walls is angular in early chambers and a weak keel is
present. The extent to which the junction is modified to a curve in late ontogeny and the keel is lost, is a major aspect of
variation among specimens viewed in spiral orientation. In the reference population, Fig. 4.1 #6 shows an individual
in which little or no modification has occurred by late ontogeny. In contrast, the shorter axial and distal walls of the
last-formed chamber in Fig. 4.1 #17 are significantly modified and merge in a broad curve. Similar variation occurs
in other upper Tongaporutuan populations (Fig. 4.6, Fig. 4.7). In another comparative population (T27/f399 Fig. 4.3)
the junction is modified to a curve in most specimens and in a few the last-formed chambers are bulbous (Fig. 4.3 #4).

'Advanced' specimens with inflated, late-formed chambers are present in earliest populations (e.g., Y18/f560 Fig. 4.8).
Their representation thereafter is quite variable. When the morphotype is dominant, as in T27/f399 (Fig. 4.3), the
populations are distinctive. More commonly, it is an intra-population variant. Although this morphotype is most
similar, architecturally, to compressed Bolivina, no bioseries from angular to bulbous profiles has been noted. A
conjecture is that the design is related to infaunal existence. Present knowledge of the distribution of the morphotype
does not justify its recognition in taxonomy.

25
 DRAFT
Figure 4.7: (A) Bolivinita compressa Finlay. Bolivinita compressa Finlay, B. pliobliqua Vella and B. quadrilatera
(Schwager). Variation in the spiral outline of specimens from T26/f139. (B) Specimens (standardized for size) are
plotted on 3-dimensional coordinates based on principal components analysis of 20 coefficients for 5 elliptical Fourier
harmonics fitted to outlines; 96% of variation is shown.

Figure 4.8: Bolivinita compressa Finlay and B. pohana Finlay. Specimens from Y18/f560, basal Tongaporutuan Stage.

4.6 Bolivinita compressa Biogeography

Although Bolivinita compressa is widely distributed through the Eastern Basin, it is less persistent in the southern
extension of the basin in northeastern South Island. In North Canterbury it occurs at Cobolds Creek (N33/f9694-

26
9696) but has not been found in apparently suitable facies in the coastal sequences at Gore Bay and Motunau. Its local
extinction at Blind River, Marlborough (Fig. 4.5) might be related to a decline in basin bathymetry.

DRAFT
Typically, Bolivinita compressa is a prominent member of middle bathyal or deeper assemblages in the Eastern Basin.
However, in the west of the basin (Napier - Taupo Road section) there are large populations (e.g., V20/f390) in
upper bathyal sediments. Its relative abundance (c. 30% of benthic specimens in the >200 µm fraction) in the latter
collection, and in assemblages in the vicinity of the type locality at Hangaroa River, suggest that it might have been
an opportunist. Blooms of infaunal taxa commonly occur in oxygen-deficient environments (?).

Beyond the Eastern Basin the record is sparse (Fig. 4.9). In Taranaki Basin it is impersistent, and rare, in middle -
lower bathyal Tongaporutuan sequences adjacent to the Taranaki Fault but has not been recorded in Tongaporutuan
strata elsewhere in the basin, even in deep water facies. However, it is present in bathyal Opoitian strata in Tangaroa-1.
There are no records in the New Zealand Fossil Record File from western South Island, although this is not conclusive
evidence that the species is absent. In Southland it is present in deep bathyal Kapitean assemblages (C46/f53-54)
from Wairaurahiri River. The records are notable because B. compressa is seldom found in Kapitean assemblages (?).
However, it was not found in middle - lower bathyal lower Tongaporutuan assemblages (e.g., C46/f46) from Southland
that include B. medialis.

In overview, the record of Bolivinita compressa indicates that it dispersed widely in New Zealand region but only in
the Eastern Basin was it persistent, with large populations. The distribution does not seem to be related to bathymetric
differences between basins and its explanation is a principal enigma of Bolivinita biogeography.

Figure 4.9: Distribution of Bolivinita compressa on

modern coordinates. Shaded areas are generaliza-
tions of known distributions and are conservative.
They do not identify possible changes in its distri-
bution through the Late Neogene.

4.7 Bolivinita compressa Stratigraphic Distribution

Base of Tongaporutuan Stage to upper Opoitian Stage; ?Mangapanian Stage. (Fig. 1.1)

Lowest occurrence: ? and ? wrote that the first appearance of Bolivinita compressa marked the base of the upper
Tongaporutuan Stage. Upper Tongaporutuan is an informal usage and ? and ? may not have referred to the same
events for defining upper Tongaporutuan. In the Tongaporutuan stratotype (north Taranaki coast, ?), the lowest record
(Q18/f8512) of B. compressa by ? was from near the base of Urenui Formation. The horizon is near the top of the
informal lower Tongaporutuan unit of Scott (in ? p. 92), which is marked by the highest occurrence of Globoquadrina
dehiscens.

27
This review indicates that Bolivinita compressa first occurs lower, near the base of Tongaporutuan Stage. At Kaiti
Beach (Gisborne City) the lowest record is a single specimen in Y18/f554, c. 6 m above the lowest record of B.

DRAFT
pohana (Y18/f552) and just below the base of the Globorotalia miotumida Kaiti coiling zone (?). More specimens
occur in Y18/f560 (Fig. 4.8), above the base of this zone. In the Tongaporutuan stratotype B. compressa is present in
Q18/f36 from the lower part of an interval that Scott (in ? fig. 23) identified as the Kaiti coiling zone. It is probably
present near the base of the stratotype in Q18/f72 (one individual recorded as B. pohana by Scott in ?.

Highest occurrence: ? noted that Bolivinita compressa was present in the Opoitian Stage stratotype at Mangapoike
River. In records at Micropaleontology Laboratory, GNS Science, Hornibrook identified it up to X19/f7758 (c. 560
m below the top of the Opoitian stratotype). The presence of Globorotalia crassaconica, G. inflata (morphotypes),
G. puncticulata and G. pliozea indicate that the horizon is in the upper part of the stage. This record of B. compressa
is accepted, as are specimens from a lower collection (X19/f7749). Bolivinita pseudocompressa is present higher
in the sequence (X19/f7763) and appears to replace B. compressa. At Willow Flat (Mohaka River) both taxa are
identified in W19/f64 while only B. pseudocompressa is present in W19/f65, probably higher in the section. Specimens
with a very large megalospheric proloculus occur in Taranga-1 at 1600 m (Mangapanian; middle bathyal) and are
tentatively assigned to B. compressa. If valid, this record might indicate that B. compressa was reduced to small,
isolated populations in deep environments in the Late Pliocene and that it does not have a well-defined extinction
event.

28
 DRAFT
Figure 4.10: Bolivinita compressa Finlay. (A) Variation in the spiral outline of specimens from P29/f11 and T27/f399.
(B) Specimens (standardized for size) are plotted on 3-dimensional coordinates based on principal components anal-
ysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 89 % of variation is shown and the
minimum spanning tree is drawn. Refer to (A) for micrographs of specimens #A-#E, #10, #18, #21. (C) Cluster
analysis of the data.

29
Chapter 5

DRAFT
Bolivinita pseudocompressa Crundwell

2016 Bolivinita pseudocompressa Crundwell, This volume: p. 107, fig. A.1

Original Description

"Test medium sized, broad, moderately compressed, about 2 to 2.5 times longer than broad, rhomboidal in cross-
section, expanding rapidly initially, then parallel sided. Chambers arranged biserially, although opposed at 180o
intervals. Axial walls strongly differentiated in length to form broad, shallow, convex axial surfaces. Distal walls
generally smooth, slightly convex. Sutures inclined backward at 45o to the axial plane, generally flush except for
slightly depressed short-axial wall sutures. Wide flange-like keels developed at junctions of long-axial and distal
walls, often broken to produce a serrated edge. Short-axial/distal wall keels are generally incipient or may be absent
in some specimens. Aperture interiomarginal, loop-shaped, with inner facing tooth plate projecting from junction of
thickened keels bordering the apertural distal wall." (p. 107)

5.1 Bolivinita pseudocompressa Reference Population T27/f78

Material

All specimens are from c. 200 m above the base of Mangaopari Mudstone in the Mangaoriki Stream - Ruakiwi Stream
section, southern Wairarapa. ?, p. 96 placed the horizon in the upper Opoitian - Waipipian Stages. Twenty specimens
were examined in spiral orientation; c. 50 were studied in axial and distal orientations.

Spiral Orientation T27/f78

Figure 5.1: Bolivinita pseudocompressa Crundwell. Variation in the spiral outline of specimens from T27/f78.

Chamber arrangement: Chambers are offset and the outline of most shells resembles a rhomboid. Commonly, the
major axis of the shell is through the keels. Symmetry of the outline is retained by 180o rotation about this axis.

30
Distal wall: At one side, adjacent to the longer axial wall, the wall makes a sharp boundary with the keel. At the
opposite side the junction with the shorter axial wall varies from angular (Fig. 5.1 #3) to narrowly rounded (Fig. 5.1

DRAFT
#19). At some angular junctions there is a low keel (Fig. 5.1 #8). At the opposite extreme, the distal wall of the
penultimate chamber in Fig. 5.1 #17 merges with the shorter axial in a wide curve. The outline of the wall varies from
slightly convex (Fig. 5.1 #3, #15) to almost linear (Fig. 5.1 #8). Commonly, the crest of the wall is lower than the
major keel.

Axial walls: Shorter and longer walls are usually well differentiated in length and diverge at different angles. Opposed
axial walls often create a wide, shallow, asymmetrical, V-shaped re-entrant in the outline (e.g., Fig. 5.1 #11). Rarely,
the walls are almost parallel (Fig. 5.1 #17 last chamber) and the re-entrant is barely defined. Rhomboidal outlines are
well-defined in such shells.

Keels: There is always a strong, elevated keel at the junction between the distal and longer axial walls. On some
specimens it tapers distally to a thin flange (Fig. 5.1 #17). This keel links, via the rim of the aperture to its equivalent
on the opposed chamber. The combined keels form a weakly sigmoidal structure closely aligned to the major axis
of the spiral outline. Commonly, there is a much less prominent, weaker keel at the junction between the distal
and shorter axial walls (Fig. 5.1 #8). Exceptionally, this keel may be prominent (Fig. 5.1 #18) but, like the weaker
exemplars, it does not extend up to the aperture.

Aperture: Commonly, the opening varies from an asymmetrical A-shape (Fig. 5.1 #3) to oval (Fig. 5.1 #15). The
opening is exceptionally high in Fig. 5.1 #16. Asymmetry in the shape of the opening develops from the greater
curvature of the side adjacent to the major keel. The rim of the aperture is wide and welded to the major keels. The
tooth has a wide base at the top of the opening (Fig. 5.1 #19) and rises to about one half of the height of the opening.

Axial Orientation T27/f78

Figure 5.2: Bolivinita pseudocompressa Crundwell. Guide to morphology of specimens from T27/f78 in axial, oblique
axial and distal orientations.

Shell outline: The prolocular section is wider in megalospheric than in microspheric forms (Fig. 5.2 #1, #3), but a
wide flange at the extremity of keels in some microspheric specimens may greatly reduce the contrast. Lateral sections
are mainly defined by the crests of distal walls; their convexity is sufficient to create a slightly lobulate outline. As the
axial walls come very close to the periphery (Fig. 5.2 #1), they may show in the troughs between lobes in the outline.
In oblique axial orientation (Fig. 5.2 #4, #5), lobulation usually is completely masked by the wide keels. The lateral
sections diverge very little in some megalospheric specimens (Fig. 5.2 #1). More commonly, width gradually expands
through ontogeny. The lateral sections taper to a narrow apex above the aperture. The apex is considerably wider in
oblique axial orientation (Fig. 5.2 #4, #5).

Chambers: There are up to c. 8 pairs of biserially-arranged chambers. Chambers are closely packed and interdigitate
laterally at low angles. This produces the offset that is apparent in spiral and oblique axial orientations. Axial wall

31
 DRAFT
Figure 5.3: Bolivinita pseudocompressa Crundwell
T27/f78. Wall topography.

surfaces are nearly planar but the outer section of some shorter axial walls become convex as they meld with distal
walls (Fig. 5.2 #4). Sutures are moderately depressed under SEM imagery (Fig. 5.2 #3) but this is less obvious with
light microscopy. The keel at the junction between longer axial and distal walls is prominent and commonly tapers out
to a wide flange (Fig. 5.2 #4, #5). There is considerable variation in keel development at the junction between shorter
axial and distal walls. In a few specimens none of the chambers have a keel at this site. At the other limit of variation
are specimens that have a keel running continuously along the chamber sequence. It may be quite prominent, but is
weaker than the major keel at the longer axial - distal junction; it never develops a flange. Between these limits are
specimens with weak, usually discontinuous keels. In Fig. 5.2 #4 the keel is present on early chambers, attenuates
about mid-ontogeny and is absent on the nth and (n-2)th chambers. In Fig. 5.2 #5 the keel on the (n-1)th chamber
is not connected with its equivalent on the early chambers. On some specimens each of the later chambers has a
weak keel that is discrete and not linked to its neighbours. These data might suggest that the keel at the shorter axial
- distal junction is a relict structure. There are no ribs on axial walls. Pores are small, oval-shaped, and irregularly
distributed on the last chamber (Fig. 5.3 #8). Their density reduces (?progressively) on earlier chambers but their size
and elongation increases (Fig. 5.3 #9, #10).

Distal Orientation T27/f78

Shell outline: Some prolocular sections are narrow (Fig. 5.2 #7) but keels and flanges may considerably augment their
width (Fig. 5.2 #6). Lateral sections diverge very gradually and shell profiles are narrower than in axial orientation.
One of the lateral sections is sharply defined by the major keel and is non-lobulate. The other, formed by the junction
of shorter axial and distal walls, is often less well-defined, particularly if a keel is absent or discontinuous. This section
is lobulate in some specimens. The apertural section is narrow.

Chambers: Outlines are rectangular but change significantly in shape through ontogeny due to the much more rapid
increase in height than width. The lower margin is linear, or slightly concave. Chamber surfaces are gently convex
and sutures are weakly depressed. Generally, ribs are inconspicuous, or absent. Specimens in Fig. 5.2 #6, #7 have a
rib over the proloculus and there is a longer rib on the distal wall of Fig. 5.2 #5. Very weak ribs occur on the distal
wall of Fig. 5.2 #4 and are most obvious over the suture between the nth and (n-2)th chambers.

5.2 Bolivinita pseudocompressa Comparative Population X19/f9493

Material

State Highway 2 at junction with Tahaenui Road (west of Nuhaka); c. 70 m above Tahaenui Limestone (unpublished
column, Micropaleontology Laboratory, GNS Science, Lower Hutt). Waipipian Stage. Most of the 25 specimens are
damaged.

Spiral orientation: Some specimens have compact outlines. Chambers are offset, but the amount is sometimes only
moderate. The junction between the shorter axial and distal walls is narrowly rounded to angular. Re-entrants in axial
walls are well-defined (Fig. 5.4 #2).

Axial, distal orientations: The majority of specimens have a weak keel at the junction betweeen the shorter axial and
distal walls that runs through the chamber sequence (Fig. 5.4 #8). Axial walls are featureless. Some specimens have

32
 DRAFT
Figure 5.4: Bolivinita pseudocompressa Crundwell. Selected specimens in spiral and axial or oblique axial orienta-
tions.

several weak ribs on the distal walls of early chambers (Fig. 5.4 #8). Flanges are weaker than in T27/f78.

Remarks: Relatively compact spiral outlines, only moderate offset of chambers, and common development of a
(continuous) keel at the junction between shorter axial and distal walls are features that distinguish many specimens
from those in T27/f78.

5.3 Bolivinita pseudocompressa Comparative Population W21/f8521

Material

Maraetotara River section, 0-6 m below the unconformity with Castlecliffian strata; c. 40 specimens. Upper Opoitian.
? described the biostratigraphy of the lower part of this sequence and ? recorded the presence of Bolivinita finlayi.

Spiral orientation: Chambers are offset, but possibly less so than in T27/f78. Axial walls are differentiated and often
define asymmetrical V-shaped re-entrants (Fig. 5.4 #4) although, as in T27/f78, the re-entrants are very weak on some
specimens (Fig. 5.4 #5). There is a strong keel at the junction of the longer axial and distal walls. A flange is often
present but is often less conspicuous than in T27/f78. The junction between the shorter axial and distal walls varies
from narrowly rounded to angular. The convexity of the distal wall in some specimens (Fig. 5.4 #4) is greater than in
T27/f78. Commonly, its crest is as high, or higher, than the level of the keel.

Axial, distal orientations: There is a prominent keel at the junction of the longer axial and distal walls. It tapers to a
flange that is often narrower and less continuous than in specimens from T27/f78. Keel development at the junction
of the shorter axial and distal walls is variable; Fig. 5.4 #10 shows a specimen in which the structure is present on
chambers prior to the last. It is less extensive in Fig. 5.4 #11. Generally, keel development at this locus is weaker than
in T27/f78. Most axial walls appear featureless under optical microscopy but very weak, close-spaced ribs are visible
on some distal walls. Scanning electron microscopy shows that ribs are present on some axial walls (Fig. 5.4 #10).

33
5.4 Discrimination Bolivinita pseudocompressa : B. compressa

DRAFT
Material

Bolivinita compressa fromX18/f38 and T27/f399. Bolivinita pseudocompressa fromT27/f78, X19/f9493 and W21/f8521.

Spiral orientation: Although Bolivinita pseudocompressa is commonly smaller (e.g., Fig. 5.5 (A) #32, #37), its design
resembles B. compressa. Chambers are significantly offset; the lengths of axial walls are unequal; the keel between
longer axial and distal walls is robust and prominent; shell outlines are rhomboid-like. Principal differences are in
second-order features. Offsets between chambers are sometimes greater in B. pseudocompressa from T27/f78. The
shorter axial - distal wall junction in B. pseudocompressa commonly is angular (Fig. 5.1), and sometimes has a keel.
In B. compressa from T27/f399 the junction is broadly rounded. A related difference is the weaker convexity of distal
wall profiles in B. pseudocompressa (Fig. 5.5 (A) cf. #32, #A. This is quite marked in the T27/f78 sample (Fig. 5.1),
although it is less conspicuous in specimens from W21/f8521 (Fig. 5.4). Re-entrants in axial walls often are deeper
and better defined than in B. compressa.

The cluster analysis of spiral outline data from T27/f78 and T27/f399 (Fig. 5.5 (C)) is structured, with one low level
group formed solely of B. pseudocompressa and the remaining 5 specimens distributed in the clusters for B. compressa.
There is minor overlap between the taxa in the ordination. However, there is greater overlap in the comparison with B.
compressa from X18/f38. All of the low level clusters are admixtures, although one or other of the taxa is dominant
in each.

Axial, distal orientations: These show that the often angular junction between shorter axial and distal walls, seen
in spiral orientation, sometimes persists from early ontogeny in Bolivinita pseudocompressa (Fig. 5.2). Although
specimens with a continuous keel at this site are a minority in B. pseudocompressa, there are no equivalents among
specimens of B. compressa from T27/f399. Similarly, 'relict' structures (weak, impersistent keels) are rarely encoun-
tered in T27/f399. In both samples the major keel tapers to a flange, but it is typically wider, and more fragile, in
B. pseudocompressa (Fig. 5.2). Lateral sections of the outline are less lobulate in B. pseudocompressa (Fig. 5.2)
than in B. compressa (Fig. 4.2). Wall topography is featureless in B. pseudocompressa (Fig. 5.3) whereas there are
small domes around pores in B. compressa (Fig. 4.4). Short, megalospheric specimens of B. compressa, often with
<5 chambers and a with very broad prolocular section (Fig. 4.2 #8), do not occur among B. pseudocompressa from
T27/f78.

Remarks: Specimens in T27/f95 (Fig. 5.4) #6, #12; Mangaopari Mudstone in Mangawhangki Stream, southern
Wairarapa; lower Opoitian with Globorotalia mons and G. puncticulata) are identified by M.P. Crundwell (pers.comm.
1996) as early Bolivinita pseudocompressa, or a possible ancestor. In Fig. 5.4 #6 the axial and distal walls form a con-
tinuous curve, as in B. compressa from T27/f399.

In W19/f64 (Willow Flat, Mohaka River, upper Opoitian) Bolivinita pseudocompressa has weak ribs on axial and
distal walls. These are absent from accompanying specimens tentatively identified as B. compressa. Further work
on the discrimination of these taxa in upper Opoitian assemblages is needed and will help resolve the ancestry of B.
pseudocompressa.

34
 DRAFT
Figure 5.5: Bolivinita pseudocompressa Crundwell and B. compressa Finlay. (A) Comparison of specimens in spi-
ral orientation from T27/f78 and T27/f399. (B) Specimens of B. pseudocompressa (T27/f78) and B. compressa
(T27/f399), standardized for size, plotted on 3-dimensional coordinates from principal components analysis of 20
coefficients for 5 elliptical Fourier harmonics fitted to the outlines (94% of variation represented); a minimum span-
ning tree is fitted. (C) The same data are used in the cluster analysis. (D) Cluster analysis of spiral outline data for B.
pseudocompressa (T27/f78) and B. compressa (X18/f38); the latter sample includes 3 specimens of B. pliobliqua.

35
5.5 Discrimination Bolivinita pseudocompressa : B. finlayi

DRAFT
Material: Bolivinita finlayi from U23/f142, U24/f697 and U24/f698. Bolivinita pseudocompressa from T27/f78,
X19/f9493 and W21/f8521.

Figure 5.6: (A) Bolivinita pseudocompressa Crundwell and B. finlayi Kennett. Comparison of specimens in spiral
and axial orientations. (B) Cluster analysis of specimens from T27/f78 and U23/f142 using 20 coefficients for 5
elliptical Fourier harmonics (specimens standardized for size) fitted to outlines in spiral orientation. Refer to (A) for
micrographs of specimens #A-#C, #2, #8, #13.

Spiral orientation: Major architectural features compare with those of Bolivinita finlayi: rhomboid outline, offset
chambers, axial walls differentiated in length and keel between longer axial and distal walls. However, there are
differences in detail. The keel between longer axial and distal walls is generally much more prominent in B. pseudo-
compressa and may extend as a flange (Fig. 5.6 #8; compare Fig. 5.6 #A-#C). A flange is not developed in B. finlayi.
The junction between the shorter axial and distal walls is usually better defined in B. pseudocompressa, as are the
re-entrants in axial walls. These features contribute to the more angular outline of B. pseudocompressa. In some
populations of B. pseudocompressa (e.g., T27/f78), the offset between chambers is greater than in most B. finlayi. The
taxa are largely separated in the cluster analysis of spiral outline data (Fig. 5.6 (B)), but the distribution of outliers
suggests that the variation fields show minor overlap. Note that the analysis neglects the generally larger size of B.
pseudocompressa relative to B. finlayi.

Axial, distal orientations: In Bolivinita pseudocompressa the prominence of the major keel and flange, and lobulate
lateral outline of one of the distal walls contrast with B. finlayi. Particularly, B. pseudocompressa lacks the narrow,
low ribs that are occur on axial and distal walls of B. finlayi.

Remarks: Hornibrook (in ?, p. 100) listed as Bolivinita aff. compressa material from Maraetotara River (W21/f8521 =
N135/f521; Fig. 5.4 #4, #5, #10, #11) that ? had included in Bolivinita finlayi. The species was figured from Tangoio
River (Nukumaruan Fig. 5.4 #1, #7) as Bolivinita cf. compressa by ? who showed that it ranged from Opoitian Stage

36
to Nukumaruan Stage. They noted that it has faint ribs ("costae") but considered that B. finlayi was "distinguished
particularly by the finely but consistently costate test". Here the Maraetotara River and Tangoio River material are

DRAFT
included in B. pseudocompressa.

Although most specimens in the samples of B. pseudocompressa and B. finlayi compared above are readily separated,
the architectural similarity of the taxa is such that some material, particularly if poorly preserved, may be difficult to
allocate. Bolivinita finlayi and B. pseudocompressa appear to be successional taxa but their phyletic relationship is
unknown.

5.6 Bolivinita pseudocompressa Biogeography

Figure 5.7: Distribution of Bolivinita

pseudocompressa on modern coordi-
nates. Shaded areas are generalizations
of known distributions and are conser-
vative. They do not identify possible
changes in its distribution through the
Late Neogene.

Bolivinita psedocompressa is more localized, less persistent, and usually much less abundant than B. pliozea, with
which it sometimes occurs. In the Eastern Basin it is present from northern Hawke's Bay to Wairarapa. It is relatively
persistent, and sometimes common, in bathyal upper Opoitian - Waipipian assemblages in Wairoa Syncline and in
southern Wairarapa). In the west of the basin it is common in U20/f8547 (near Kuripapango) in upper bathyal Opoitian
sediments. The Nukumaruan records from Petane Group are from significantly shallower environments with shelf
dwellers (e.g., V20/f149, with Notorotalia zelandica, Florilus parri).

In Turakina River sequence, Wanganui Basin, it is a minor species in Taihape Mudstone, with largest populations
from upper bathyal assemblages near the top of the unit (e.g., S21/f9, S21/f12; Waipipian). It is impersistent in the
overlying Utiku Sandstone (Waipipian - Mangapanian).

In Taranaki Basin Bolivinita pseudocompressa is very rare in upper bathyal Opoitian assemblages from Kapuni-2 1820
ft, 335.09 m, Te Kiri-1 350 m but is common at 1100 m in Witiora-1 (Waipipian). It is tentatively identified from a
Nukumaruan shelf assemblage in Taimana-1 at 670 m. Overall, the rarity of the species in Taranaki Basin parallels
the record of the architecturally similar B. compressa.

5.7 Bolivinita pseudocompressa Stratigraphic Distribution

Opoitian Stage to Nukumaruan Stage (Fig. 1.1).

37
Lowest occurrence: There are several lower Opoitian occurrences (T27/f95, V24/f10) that might represent Bolivinita
pseudocompressa. However, typical specimens first occur in upper Opoitian assemblages, associated with Globoro-

DRAFT
talia inflata. Specimens from X18/f7763 in the Opoitian lectostratotype at Mangapoike River are shown in Fig. 5.4
#3, #9.

Highest occurrence: There are Nukumaruan records from the Petane Group of Hawke's Bay, including V20/f149
(Fig. 5.4 #1, #7) and V21/f6005 (Napier City). A caveat is that specimens are rare and insufficient for assessment of
variation.

38
Chapter 6

DRAFT
Bolivinita finlayi Kennett

1967 Bolivinita finlayi Kennett, Transactions of the Royal Society of New Zealand, Geology 4: p. 997, 1000; fig. 23,
24.

Original Description

"Test of medium size, strongly compressed, fairly broad, tapering gradually to a rounded end. Cross section com-
pressed rhomboid, the acute angles carinate, the obtuse angles gently rounded and non-carinate; the broader faces
slightly concave, the narrower faces slightly convex. Sculpture of fine, numerous, irregular longitudinal costae along
most of the length of the narrow faces and in most specimens less conspicuous along the broader faces. About 7
chambers on each side, often slightly inflated. Sutures fairly heavily limbate, slightly depressed or flush, very gently
curved to straight, and moderately oblique. Aperture large, widest at base." (?, p. 997, 1000; fig. 23, 24)

6.1 Bolivinita finlayi Reference Populations U23/f142, U24/f697-8

Material

Data on spiral morphology are based on 20 specimens from U23/f142 (interbed in fine sandstone, Waikopiro Stream;
upper Kapitean - Opoitian). Descriptions of axial and distal morphology refer to c. 20 specimens from U24/f697
and U24/f698, both in the vicinity of the type locality (U24/f6049, upper Kapitean) in Mangapuka Stream, southern
Hawke’s Bay.

Spiral Orientation U23/f142, U24/f697-8

Chamber arrangement: Chambers are always offset (Fig. 6.1 (A) #20); the arrangement leads to the rhomboidal
shape of the shell outline. The major axis of the shell is usually through the keels and symmetry of the outline is
maintained by 180o rotation about this axis.

Distal wall: There is a sharp boundary at the junction with the longer axial wall. The boundary with the shorter axial
wall is very weakly defined in the great majority of specimens because these sections join in a curve (Fig. 6.1 (A) #4).
In a minority of specimens the junction is angular (Fig. 6.1 (A) #15). The distal wall profile is convex but its extent
and form are variable (Fig. 6.1 (A) #4, #6). The wall often rises above the adjacent keel (Fig. 6.1 (A) #6, #10).

Axial walls: Typically, they are of unequal length, with the longer wall adjacent to the keel. They diverge out from the
base of the chamber, usually at a small angle. The outline of the shell most closely resembles a rhomboid when axial
walls in both chambers are only slightly divergent, and distal walls are only slightly convex (Fig. 6.1 (A) #11). Axial
wall profiles vary from linear to slightly convex. The re-entrant in the shell outline formed by pairs of axial walls of
opposed chambers is usually shallow and may be imperceptible (Fig. 6.1 (A) #10, #15).

Keels: A topographic ridge is always present at one distal - axial wall junction on the last chamber and links via the
aperture with an equivalent ridge on the opposing chamber (e.g., Fig. 6.1 (A) #1). Usually, these keels lie close to the
major axis of the shell profile and are well-defined. Occasionally, the keel is very minor topographic feature (Fig. 6.1
(A) #15). A keel at the junction of shorter axial and distal walls is not visible in spiral orientation.

Aperture: The opening varies from A-shaped (Fig. 6.1 (A) #7) to sub-oval (Fig. 6.1 (A) #4). One side tends to align

39
with the keels (Fig. 6.1 (A) #6). The toothplate wraps over the opening and contributes to the wide rim (Fig. 6.1 (A)
#7). The curved tooth in the opening represents a free margin of the toothplate.

DRAFT
Figure 6.1: Bolivinita finlayi Kennett. (A) Variation in the spiral outline of specimens from U23/f142. (B) Speci-
mens (standardized for size) are plotted on 3-dimensional coordinates based on principal components analysis of 20
coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 93% of variation is shown.

Axial Orientation U23/f142, U24/f697-8

Shell outline: In most specimens the prolocular section is broadly rounded. There are rare specimens in which the
prolocular section is narrower, but microspheric individuals have not been definitely identified. The distal walls define
the lateral outlines but the margins of the axial wall are in close proximity (Fig. 6.2 #1). The lateral sections diverge out
from the proloculus but angles commonly decline in mid-ontogeny and the sections become almost parallel (Fig. 6.2
#2).

Chambers: In mature specimens there are at least 6 pairs of biserial chambers. They incline downward and become
wedge-shaped near the periphery. The maximum number is uncertain because many specimens are broken. For most
of ontogeny, chambers are closely packed but there may be clear inter-chamber areas between some later chambers.
Axial walls are flat, or almost so. They abut with flush sutures (Fig. 6.2 #3), although some sutures between late-
formed chambers are depressed. There is a keel at the margin of the longer axial wall (Fig. 6.2 #4). The shorter
axial and distal walls usually join in a continuous curve, but some specimens have a weak keel (Fig. 6.2 #3). Ribs are
always present on axial walls but are variable in number and continuity. One rib in Fig. 6.2 #3 runs over most pairs of
chambers. Another is present only on later chambers. In Fig. 6.2 #1 about four sinuous ribs arise near the proloculus;
only one reaches the last chamber. There are numerous shorter ribs, some of which are very weak. Small, oval-shaped
pores are irregularly distributed over the smooth wall (Fig. 6.3 #7) but are not clearly preserved on some specimens.
Some pores may be enlarged by solution (Fig. 6.3 #8).

40
 DRAFT
Figure 6.2: Bolivinita finlayi Kennett. Guide to morphology of specimens from U23/f142 and U24/f697 in axial,
oblique axial and distal orientations.

Distal Orientation U32/f142, U24/f697-8

Shell outline: The keel sharply defines the outline at the junction between the distal and longer axial walls. The
opposite margin is usually less well defined as it lies on the curved surface joining distal and shorter axial walls.
Flexure of the coiling axis is rare and weak; very little of the axial wall is visible in this orientation (Fig. 6.2 #5,
#6).The prolocular section is broadly rounded. The lateral margins diverge at low angles from this section. Slight
expansion in width often continues through ontogeny. Lobulation is always very weak but is more apparent on the
side without the keel. The apertural section is almost flat (Fig. 6.2 #5, #6).

Chambers: During ontogeny, height increases much more rapidly than width. The lower margin of a chamber is
mostly linear and meets the side adjacent to the keel almost at right angles. On the side without a keel the lower
margin appears to curve into the lateral section. Lateral sections are weakly convex. Junctions between chambers
formed early in ontogeny are almost flush, signifying only slight curvature of the distal wall. Later chambers are
weakly convex and chamber junctions are slightly depressed (Fig. 6.2 #5). Walls are covered by up to 8 very weak
ribs that often are slightly oblique to the longitudinal axis of the shell.

Figure 6.3: Bolivinita finlayi Kennett U23/f142. Wall to-

pography.

6.2 Discrimination Bolivinita compressa : B. finlayi

Material

Bolivinita compressa from X18/f38 and T27/f399. Bolivinita finlayi from U23/f142, U24/f697-8.

Spiral orientation: Bolivinita finlayi is significantly smaller and is relatively less elongated (Fig. 6.4 (A) #C, #14).
Distal walls tend to be more convex in B. finlayi than in B. compressa from X18/f38. However, this does not apply
to populations like T27/f399 (Fig. 4.3). The keel is less prominent in B. finlayi than in B. compressa. Some major

41
architectural features are closely similar. These include the amount of offset between chambers, the common absence
of a keel between shorter axial and distal walls, and the variably weak re-entrants in axial walls.

DRAFT
Figure 6.4: (A) Discrimination of Bolivinita finlayi (U23/f142) from B. compressa (X18/f38) using characters seen in
spiral and axial orientations. (B) Discrimination of Bolivinita finlayi (U23/f142) from B. compressa (X18/f38) using
spiral outline data. Several specimens of Bolivinita pliobliqua are included. The cluster analysis uses 20 coefficients
for 5 elliptical Fourier harmonics (specimens standardized for size) fitted to outlines in spiral orientation.

The cluster analysis (Fig. 6.4 (B)) indicates that there is only minor overlap in spiral shape. Remember that the analysis
neglects the marked difference in gross size. The relative elongation of Bolivinita compressa outlines contributes to
the distinction.

Axial, distal orientations: Bolivinita finlayi is commonly significantly shorter than B. compressa (Fig. 6.4 (A) #i);
this probably reflects differences both in number and height of chambers. Outlines (axial orientation) are usually
non-lobulate in B. finlayi and often distinctly lobulate in B. compressa (Fig. 6.4 (A) #ii). A flange does not develop
on the keels of B. finlayi. Particularly, the presence of ribs on axial and distal walls of B. finlayi distinguish it from B.
compressa.

6.3 Bolivinita finlayi Biogeography

In the Eastern Basin Bolivinita finlayi has been recorded from southern Hawke's Bay and adjacent northern Wairarapa,
and from Marlborough (Fig. 6.5). In Taranaki Basin it occurs about the Kapitean - Opoitian boundary in the peninsula
region. Good understanding of the biogeography of B. finlayi is made difficult by its sparse, possibly impersistent,
populations. It probably extended beyond the distribution shown in Fig. 6.5. At Blind River it is preceded by B.
compressa, which occurs rarely in middle bathyal assemblages with Sigmoilopsis schlumbergeri; see ? for bathymetric
calibration. The lowest record (P29/f32 Fig. 4.5) is near the initiation of progressive shallowing of the site through the

42
upper bathyal zone (?). Other records of B. finlayi are from upper bathyal or shelf environments. The data suggest that
B. finlayi occupied a shallower niche than B. compressa. It was never as populous, or persistent, as the latter species.

DRAFT
6.4 Bolivinita finlayi Stratigraphic Distribution
Figure 6.5: Distribution of Bolivinita finlayi on
modern coordinates. Shaded areas are general-
izations of known distributions and are conser-
vative. They do not identify possible changes
in its distribution through the Late Neogene.

Figure 6.6: Bolivinita finlayi Kennett Specimen from

P29/f32, Blind River.

Kapitean Stage to Opoitian Stage (Fig. 1.1).

Lowest occurrence: P29/f32 (Fig. 6.6) Blind River, lower Kapitean (Fig. 4.5). Rare specimens occur in cuttings from
an undifferentiated upper Tongaporutuan - lower Kapitean interval in Titihaoa-1 (SE40176/f1) but might be caved.
The type locality (U24/f6049) is the highest Kapitean collection of ? in Mangapuka Stream, and is adjacent to basal
Opoitian.

Highest occurrence: Bolivinita finlayi occurs in small numbers in Opoitian strata in southern Hawke's Bay (e.g.,
U23/f143). In upper Opoitian assemblages it is replaced by B. pseudocompressa. ? recorded B. finlayi from Opoitian
strata in the Maraetotara River section (?). Specimens in W21/f8521 and W21/f8545 are here referred to B. pseudo-
compressa (Fig. 5.4 #4, #5, #10, #11).

No Waipipian records of B. finlayi have been found.

43
Chapter 7

DRAFT
Bolivinita urenuia Scott n. sp.

Bolivinita cf. pohana Finlay. ?, fig. 15-22, 67-71, 89-91.

Holotype: TF1675/1 (GNS Science type foraminiferal collection).

7.1 Bolivinita urenuia Type Population Q19/f102

Material

Q19/f102 (type locality), west end of Onaero Beach, 3 m west of concrete protection wall, adjacent to semi-tabular
concretions. Urenui Formation, upper weakly bedded unit. The location is adjacent to Q19/f56 (? appendix 1 map G)
and near the top of measured section 39 (? appendix 2). Twenty-seven specimens were examined in spiral orientation;
c. 45 in axial and distal orientations.

Spiral Orientation Q19/f102

Figure 7.1: Bolivinita urenuia Scott n. sp. Variation in the spiral outline of specimens from Q19/f102.

Chamber arrangement: The offset between the biserially-arranged chambers is well-defined in most specimens (e.g.,
Fig. 7.1 #6) but is reduced in some (Fig. 7.1 #2, #18). Rhomboid-like outlines are typical. Symmetry is preserved
when the outline is rotated 180o . The major diagonal of the outline is through the stronger pair of keels. A few
specimens show minor flexure of the coiling axis during ontogeny.

Distal wall: Usually, both junctions with axial walls are well-defined by a keel. Rarely, the junction with the shorter
axial wall is narrowly rounded, and without a keel (Fig. 7.1 #27). Most outlines are weakly convex and rise only
slightly above the keels. However, convexity is greater in some specimens (Fig. 7.1 #14).

44
Axial walls: In most specimens the lengths of the shorter and longer walls are clearly differentiated (Fig. 7.1 #6,
#23). The difference is most marked in specimens in which chambers are well offset. The wall profiles are linear and

DRAFT
are always divergent. The V-shaped re-entrant in the outline formed by opposed axial walls is commonly wide and
asymmetrical. Although often shallow, it is present in all specimens.

Keels: The keel at the junction of the longer axial wall with the distal wall is wide and robust. It links, via the aperture,
with the equivalent keel on the opposed chamber (Fig. 7.1 #17). On the last chamber the keel at the junction of the
shorter axial wall with the distal wall is narrower (Fig. 7.1 #7) and typically dies away on the upper surface of the
chamber (Fig. 7.1 #17). However, on the penultimate chamber this keel commonly runs up to the rim of the aperture.

Aperture: The base is defined by the distal wall of the chamber and is almost flat. The sides are curved to form a
broad, arched opening. Often, curvature is greater on the side adjacent to the primary keel and follows the direction of
the latter (Fig. 7.1 #6). This asymmetry tends to be less obvious in specimens in which the offset between chambers
is minor (Fig. 7.1 #18). The tooth is curved, robust, and projects into the upper part of the opening.

Axial Orientation Q19/f102

Figure 7.2: Bolivinita urenuia Scott n. sp. Guide to morphology of specimens from Q19/f102 in axial, oblique axial
and distal orientations.

Shell outline: The radius of curvature of the prolocular section varies with the size of the proloculus (Fig. 7.2 #2, #3)
but may be increased by flanges at the margins of the keels. As the keel and crests of distal walls are almost aligned,
lateral sections are often defined by a combination of their profiles, with the keel bridging between the weakly convex
crests of distal walls. Lateral sections are slightly lobulate (Fig. 7.2 #2) but this is not apparent in oblique axial
orientation. The sections diverge at greater angles in microspheric than in megalospheric specimens. Lateral sections
in some of the latter are almost parallel in late ontogeny (Fig. 7.2 #1). The apex above the aperture is narrow.

Chambers: Some microspheric specimens have up to 9 pairs of wedge-shaped chambers. Often they are close-packed
but some specimens show small lumina between chambers. Laterally, chambers interdigitate at low angles; this forms
part of the offset seen in spiral orientation. Surfaces of both shorter and longer axial walls are almost planar and many
sutures are only slightly depressed (Fig. 7.2 #2). However, because of the angle at which chambers interdigitate, a
shallow, poorly-defined depression sometimes forms between the two sets of chambers (Fig. 7.2 #1). The keel at the
junction between the longer axial and distal walls is robust and continuous from the prolocular area to the aperture.
Commonly, it tapers outward to a thin, translucent flange (best seen in oblique axial orientation Fig. 7.2 #4). Generally,
the keel at the junction between the shorter axial and distal walls is narrower, and without a flange. It runs continuously
from the prolocular region to the last chamber but often becomes very weak on the upper surface of that chamber and
may not extend to the aperture. Relict keels are conspicuous on late-formed chambers of some specimens (Fig. 7.2
#3). Characteristically, ribs are absent from axial walls but one or more weak ribs extend from the proloculus to the

45
 DRAFT
Figure 7.3: Bolivinita urenuia Scott n. sp. Q19/f102.
Wall topography.

earliest chambers of a few specimens.

Distal Orientation Q19/f102

Shell outline: The prolocular section is narrowly rounded in megalospheric specimens and is pointed in some micro-
spheric specimens. The keels significantly modify the outline around the proloculus in many specimens. The lateral
outlines are sharply defined by the keels and are usually non-lobulate. They diverge immediately above the proloculus
but the angle declines in the early chambers (Fig. 7.2 #6) and may reduce to zero. Near the base of the last chamber
the lateral outlines taper inward towards the linear section above the aperture.

Chambers: The height of chambers increases relative to their width during ontogeny. The lower margin of a chamber
is linear. The lateral margins are slightly convex, but any lobulation of the outline is suppressed by the wide keels.
In some specimens the slightly convex wall surfaces generate shallow sutural depressions (Fig. 7.2 #5). They are
not apparent on specimens with very weakly convex wall surfaces. Ribs are more common on distal walls than on
axial walls and extend further (up to the base of the penultimate chamber in Fig. 7.2 #5). They are always weak and
sometimes run slightly oblique to the coiling axis. Otherwise, wall surfaces (Fig. 7.3 #7) are featureless. This applies
also to the axial walls.

7.2 Bolivinita urenuia Comparative Population J32/f54

Material

J32/f54 samples Stillwater Mudstone c. 125 m below Callaghan's Greensand in Callaghan's Creek, Westland. It is
located in the Tongaporutuan Stage. The absence of Globoquadrina dehiscens from the assemblage suggests that the
horizon is in the informal upper unit. ?, fig. 7 showed the regional stratigraphy. Twenty-two specimens were examined
in spiral orientation and c. 40 in axial and distal orientations.

Spiral orientation: Most characters compare with those in Q18/f102. On the last chamber, the keel at the junction
between the longer axial and distal walls is robust and links, via the rim of the aperture, with its equivalent on the
opposing chamber (Fig. 7.4 #5, #18). The keel at the junction between the shorter axial and distal walls is usually
narrower and dies out on the upper surface of the last chamber (Fig. 7.4 #7). However, often on the penultimate
chamber it continues to the aperture. Distal walls are slightly convex and are often little higher than the keels (Fig. 7.4
#7). Axial walls differ slightly in length and angles of divergence.

The offset between chambers is commonly less than in Q19/f102 (Fig. 7.5 (A) #A, #13). This feature contributes to
the separation of the samples in Fig. 7.5 (B). In some specimens (Fig. 7.4 #4, #13) the offset is neglible and their gross
architecture resembles Bolivinita quadrilatera. ? so identified similar material from Corehole D. Yet even in these
specimens the minor keel on the last chamber does not continue to the aperture and distal walls retain their convexity.

Axial, distal orientations: Resemblance with Q19/f102 is close, although the smaller offset of chambers is seen in
some specimens, particularly in oblique axial view. Both keels are prominent. The flange is more conspicuous on the
major keel (at the junction of longer axial and distal walls) than on the minor keel. Generally, the keel defines the

46
 DRAFT
Figure 7.4: Bolivinita urenuia Scott n. sp. Variation in the spiral outline of specimens from J32/f54.

Figure 7.5: (A) Bolivinita urenuia Scott n. sp. Variation in the spiral outline of specimens from J32/f54 and Q19/f102.
(B) Specimens (standardized for size) are plotted on 3-dimensional coordinates based on principal components anal-
ysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 94% of variation is shown and the
minimum spanning tree is drawn.

lateral outline, or is slightly inset. Wall surfaces are usually featureless and without ribs.

7.3 Bolivinita urenuia Comparative Population P29/f14

Material

Blind River section (Fig. 4.5) is sampled by P29/f14, 270 m above base of Upton Formation. It is located in the
informal upper unit of the Tongaporutuan Stage. Twenty-four specimens were examined in spiral orientation and c.
90 in axial and distal orientations. Bolivinita compressa is present, as well as some specimens tentatively identified as

47
B. pliobliqua.

DRAFT
Spiral Orientation: Some specimens (Fig. 7.6 #3) compare closely with those in Q19/f102 (Fig. 7.4 #14) in the extent
of chamber offset, curvature of distal walls, differentiation of axial walls and keels, and shape of the re-entrants formed
by the axial walls. Specimens in which the offset between chambers is minor, as in some from J32/f54, are absent.

Figure 7.6: Bolivinita urenuia Scott n. sp. Variation in the spiral outline of specimens from P29/f14. Also shown are
Bolivinita compressa and specimens tentatively identified as Bolivinita pliobliqua.

Intra-sample variation is greater in this sample than in J32/f54 and Q19/f102. Axial wall re-entrants are considerably
reduced in some specimens and absent in others (Fig. 7.6 #19). These specimens are identified as Bolivinita compressa.
More common are specimens like Fig. 7.6 #16 in which chamber length is reduced, relative to width. This creates a
compact outline, as in B. pliobliqua and B. pliozea, although the offset between chambers is greater.

Axial, Distal Orientations: The keel at the junction between the longer axial and distal walls is robust and commonly
tapers to a flange. Usually, the keel at the junction of the shorter axial and distal walls is weaker. Very rarely, it is not
continuous between chambers. A few specimens have very weak ribs on the distal walls of early chambers.

7.4 Discrimation Bolivinita urenuia : B. compressa

Material

Bolivinita compressa from X18/f38, Hangaroa River, Gisborne, and T27/f399, Ngawaka Creek, southern Wairarapa,
are compared with B. urenuia from the type locality (Q19/f102).

Spiral orientation: Chambers are significantly offset in both taxa but there are detailed differences in the shape of wall
profiles (Fig. 7.7 (A) #A). Axial walls diverge at greater angles in Bolivinita urenuia than in B. compressa and wide,
V-shaped re-entrants are created by the linear profiles of the walls. The V-form is suppressed in B. compressa (Fig. 7.7
(A) #B). This is shown clearly in specimens from T27/f399 (Fig. 4.3 #18) but is seen also in the X18/f38 material
(Fig. 4.1 #17). Re-entrants are sometimes very weak or absent in B. compressa (Fig. 4.1 #6); this does not occur in B.
urenuia. Associated with the linear profile of the shorter axial wall in B. urenuia is a keel at the junction with the distal
wall (Fig. 7.7 (A) #13). This structure is not seen in spiral orientation in B. compressa where the shorter axial and
distal walls commonly merge in a curve. The outline data for B. compressa from T27/f399 show only minor overlap
with those for Q19/f102 (Fig. 7.7 (C-D)) and support the visual perception of significant architectural differences.

48
 DRAFT
Figure 7.7: Bolivinita urenuia Scott n. sp. and B. compressa Finlay. (A-B) Discrimination of Bolivinita urenuia
(Q19/f102) from B. compressa (T27/f399) using characters seen in spiral, axial, oblique axial and distal orientations.
(C) Bolivinita urenuia Scott n. sp. and B. compressa Finlay. Discrimination of Bolivinita urenuia (Q19/f102) from
B. compressa T27/f399) using spiral outline data. Specimens (standardized for size) are plotted on 3-dimensional
coordinates based on principal components analysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the
outlines; 92% of variation is shown and the minimum spanning tree is drawn. (D) The same data are used in the cluster
analysis.

49
Axial, distal orientations: Because of the amount of the offset between chambers, the normal repose of specimens
of both taxa is intermediate between axial and distal orientations. In this oblique axial position, maximum width of

DRAFT
some shells of Bolivinita compressa is greater than in B. urenuia (Fig. 7.7 (B) #iv). Generally, a keel is conspicuous
at the junction of shorter axial and distal walls in B. urenuia. In B. compressa a comparable structure is missing in
populations like that from T27/f399. When present in material from X18/f38, it is weak and does not extend onto the
last chamber.

7.5 Discrimination Bolivinita urenuia : B. pohana

Material

Bolivinita pohana from Q18/f13, Taranaki coastal section, and Y18/f7479, Kaiti Beach, Gisborne, are compared with
B. urenuia from the type locality (Q19/f102).

Spiral orientation: In topotypes of Bolivinita urenuia the amount of offset between chambers is often similar to B.
pohana from Q18/f13 (Fig. 7.8 (A) #B, #27). However, specimens with only a minor offset are much more common
among topotypes of B. pohana (e.g., Fig. 7.8 (A) #xi) than in topotypes of B. urenuia. Typically, distal walls are
bounded by keels in B. urenuia, and well-defined. In some specimens of B. pohana a keel is absent at the junction of
the distal with the shorter axial wall, and the outline is angular to rounded (Fig. 8.4 #22; Fig. 7.8 (A) #A). Distal walls
are less convex in B. urenuia than in topotypes of B. pohana; the difference is less marked in the comparison with
specimens from Q18/f13 but still perceptible in some (Fig. 7.8 (A) #B, #27). Axial walls commonly form V-shaped
re-entrants in both taxa, and their depth is often similar. Keels have greater amplitude, and are more conspicuous in B.
urenuia than in B. pohana. This applies particularly to populations like that at the type locality (Fig. 8.1).

Bolivinita urenuia (topotypes) are only partially separated from B. pohana (Q18/f13) in the cluster analysis of spiral
outline data (Fig. 7.8 (B)), with only 1 of 3 clusters at the 0.032 level being restricted to 1 taxon. The comparison
with Q18/f13 is presented because the (minor) convexity of distal walls in that population resembles the design in
B. urenuia. It provides a more severe test of outline correspondence between the taxa than would an analysis using
topotypes of B. pohana. The classification suggests that spiral outline data alone will not satisfactorily separate B.
urenuia from some populations of B. pohana. Other characters, particularly keels, should be considered.

Axial, distal orientations: Axial outlines are less lobulate in Bolivinita urenuia than in topotypes of B. pohana (Fig. 7.8
(C) #v, #ix). However, the contrast with B. pohana from Q18/f13 (Fig. 7.8 (C) #i) is minor. Another effect of the
weaker convexity of distal walls in B. urenuia is that sutures are less depressed (distal orientation) than in B. pohana.
The contrast also applies to axial walls, particularly in a comparison with topotypes of B. pohana (Fig. 7.8 (C) #ix).

The major keel (between longer axial and distal walls) is stronger in B. urenuia than in B. pohana. A flange, present
in some topotypes of B. urenuia (Fig. 7.2 #4), is absent from B. pohana. The minor keel extends through the chamber
sequence in B. urenuia; it is weaker in B. pohana and may be discontinuous (Fig. 7.8 (C) #ii), or absent. Ribs on distal
walls are more extensive, particularly in topotypes (Fig. 7.8 (C) #x) than in B. urenuia.

50
 DRAFT
Figure 7.8: Bolivinita urenuia Scott n. sp. and B. pohana Finlay from Q19/f102, Q18/f13 and Y18/f7479. (A)
Comparison of specimens in spiral orientation. (B) Discrimination of Bolivinita urenuia (Q19/f102) from B. pohana
(Q18/f13) using spiral outline data. The cluster analysis uses 20 coefficients for 5 elliptical Fourier harmonics (spec-
imens standardized for size) fitted to outlines in spiral orientation. (C) Comparison of specimens (axial and distal
orientation) of Bolivinita urenuia (Q19/f102) with B. pohana (Q18/f13, Y18/f7479) using characters seen in axial and
distal orientations.

51
 DRAFT
Figure 7.9: Distribution of Bolivinita ure-
nuia on modern coordinates. Shaded areas
are generalizations of known distributions
and are conservative. They do not identify
possible changes in its distribution through
the Late Neogene.

7.6 Bolivinita urenuia Biogeography

In western New Zealand the species occurs south of Taranaki Basin in Westland (Stillwater Mudstone, figured as B. cf.
pohana by ?, fig. 67-71 and in Waiau Basin (C45/f163, C45/f9511). As with some Stillwater Mudstone specimens,
chamber offsets in the latter samples are minor.

In the southern sector of the Eastern Basin Bolivinita urenuia is present in Blind River section, Marlborough (P29/f11
- P29/f14, (Fig. 4.5) and in southern Wairarapa at Palliser Bay (S28/f46 - S28/f48). Its distribution northward in the
basin is still poorly known. It is tentatively recorded from X18/f14 (Hangaroa River in Gisborne district, above the
highest occurrence of Globoquadrina dehiscens and from Z14/f87 (Orutua River near East Cape, base of Tokomaru
Sandstone; G. dehiscens is present in underlying Mapiri Formation). Although B. urenuia seems to have dispersed
throughout the Eastern Basin, its record there is that of a minor species in comparison with its principal contemporary,
B. compressa.

Bathymetrically, Bolivinita urenuia is present in upper middle bathyal assemblages (Blind River) and in upper bathyal
assemblages in the north Taranaki coastal sequence. It has not been recorded in deeper bathyal environments com-
monly occupied by B. compressa in the Eastern Basin.

7.7 Bolivinita urenuia Stratigraphic Distribution

Tongaporutuan Stage (informal upper unit, Fig. 1.1).

Lowest occurrence: In the Tongaporutuan stratotype the first record is in Q19/f21 (Urenui Formation, middle weakly
bedded unit (?, appendix 2 sheet 32). The locality is c. 60 m above the highest occurrence of Globoquadrina dehiscens,
which marks the base of the informal upper Tongaporutuan unit. ?, fig. 89-91 illustrated B. urenuia from this section
as B. cf. pohana. Elsewhere, I have not identified it in assemblages below the highest occurrence of G. dehiscens.

Highest occurrence: Bolivinita urenuia does not persist in the low diversity assemblages that occur in the vicinity of
the Urenui Formation middle channelled unit, but occurs again in the upper weakly bedded unit at Q19/f9. It continues
into the upper channelled unit up to Q19/f4 (?, appendix 2 section 24). Scott in ?, appendix 2 section 24 identified
most of the material from the upper weakly bedded unit as B. pohana. Specimens tentatively identified as B. pliobliqua

52
occur in the highest assemblages (Q19/f3, Q19/f8540) from Urenui Formation in the coastal sequence.

DRAFT
Specimens resembling B. urenuia are present in P29/f73 (Opoitian Stage, Blind River section, ?). Their taxonomic
status is uncertain.

53
Chapter 8

DRAFT
Bolivinita pohana Finlay

1939 Bolivinita pohana Finlay, Transactions of the Royal Society of New Zealand 69: p.319, pl. 27, fig. 99-100.

Original Description

"Shell differing from quadrilatera Schwager in considerably greater compression and obliquity (compressed rhomboid
in section instead of squarish), blunter and unflanged keels, simple proloculum, presence of a few ridges at sutures on
sides and much smaller aperture, occupying half of terminal face instead of nearly all. Size, 1.5 mm." (?)

Figure 8.1: (A) Bolivinita pohana Finlay and B. compressa Finlay. Variation in the spiral outline of specimens from
Y18/f7479. (B) Bolivinita pohana Finlay and B. compressa Finlay. Variation in the spiral outline of specimens
from Y18/f7479. Specimens (standardized for size) are plotted on 3-dimensional coordinates based on principal
components analysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 94% of variation is
shown and the minimum spanning tree is drawn.

54
8.1 Bolivinita pohana Type Population Y18/f7479

DRAFT
Material

Y18/f7479 (type collection), Kaiti Beach, Gisborne (near yacht club) c. 33 m above the Cyclammina tuff (Taumat-
apoupou Formation); lower Tongaporutuan Stage. Thirty topotypes were studied in spiral orientation and c. 55,
including some individuals identified as Bolivinita compressa, in axial and distal orientations.

Spiral Orientation Y18/f7479

Chamber arrangement: Although all specimens show some offset between the biserially-arranged chambers, the
amount is minor in some (Fig. 8.1 #4). Usually, the major axis of the shell lies through the keels at the junction of
the longer axial and distal walls. A few outlines are compact (Fig. 8.1 #14); others are elongated (Fig. 8.1 #18), with
higher chambers.

Distal wall: The wall is sharply defined when a keel is present at either end (Fig. 8.1 #15). However, in some
specimens (Fig. 8.1 #14) one of the keels is either very weak or absent, and the distal wall merges with an axial wall
in a zone of rapid curvature. Typically, the distal wall is regularly convex and rises well above the bordering keel(s)
(Fig. 8.1 #15).

Axial walls: Usually, these are of unequal length (Fig. 8.1 #7), with the longer wall adjacent to the major axis of the
shell. Walls incline outward from the base of the chamber and their outlines are nearly linear throughout, or contain
significant linear segments. Convex outlines are very rare. The outlines of adjoining axial walls form a wide, shallow,
weakly asymmetrical, V-shaped re-entrant in the outline of the shell (Fig. 8.1 #14). These re-entrants considerably
modify the grossly rhomboidal shape of the shell outline.

Keels: A low, well-defined ridge is formed at the junction between the longer axial and distal walls of the last chamber
and is linked to a similar ridge at the opposing junction on the penultimate chamber. The link segment, adjacent to the
aperture, is much broader than the chamber segment. A second keel is present at the shorter axial - distal wall junction
in some specimens. It is typically weaker than the primary keel and attenuates towards the aperture (Fig. 8.1 #18).

Aperture: The base, defined by the distal wall of the penultimate chamber, is weakly convex. Near the base the sides
are subparallel but in the upper half of the opening they curve inward to form an arch (Fig. 8.1 #4). The opening is
wide, and often weakly asymmetrical. A tooth projects into it from the crest of the arch and the plate folds over the
sides to form a thick rim.

Axial Orientation Y18/f7479

Shell outline: Most of the lateral outline is defined by the crests of distal walls (Fig. 8.2 #1). From the proloculus,
width expands gradually to a maximum near the base of the last chamber, above which it reduces to a rounded apex
above the aperture. Expansion in width is greater in microspheric individuals (Fig. 8.2 #2) than in megalospheric and
is minor in some of the latter. The outline is clearly lobulate, reflecting the convexity of distal walls.

Chambers: Biserial arrangement, with up to c.7 pairs. Chambers are wedge-shaped and steeply inclined (Fig. 8.2 #3).
Axial walls are slightly convex and sutures are depressed (Fig. 8.2 #2). Rarely, a relict section of the major keel is
preserved at the base of some late-formed chambers. A strong keel runs the length of the shell at one margin of the
axial walls. The opposite margin is usually marked by a much weaker keel, which may not be continuous. It is absent
on a few specimens. Both margins are well inset from the periphery. On some individuals the proloculus has several
weak ribs that may extend on to the earliest chambers. There are no ribs on the axial walls of later chambers. Surfaces
of the last 1-2 chambers are smooth but there is progressive secondary calcification around pores in earlier chambers
(Fig. 8.3 #6-#8).

55
 DRAFT
Figure 8.2: Bolivinita pohana Finlay. Guide to morphology of specimens from Y18/f7479 in standard orientations.

Figure 8.3: Bolivinita pohana Finlay. Y18/f7479 Wall

topography.

Distal Orientation Y18/f7479

Shell outline: The boundaries of distal walls define lateral sections of the outline. Expansion in width during ontogeny
is often slight, especially in megalospheric individuals (Fig. 8.2 #4). Often the outline resembles a narrow rectangle,
completed by a blunt apex above the aperture and a weakly convex prolocular section. The lateral section defined by
the major keel is usually less lobulate than the opposite margin and may include linear sections (Fig. 8.2 #5).

Chambers: Lower and upper margins are linear while the lateral margins are slightly convex. Often chambers are
almost quadrate in shape. Chamber height expands considerably more than width during ontogeny. Surfaces are
convex and some incline away from the major keel (reflecting the offset of the biserial chamber sequence and unequal
length of axial walls). Sutures are depressed (Fig. 8.2 #4). Secondary calcification around pores follows the pattern
found on axial walls. Ribs are weak, although often significantly higher where they bridge chamber boundaries
(Fig. 8.3 #6). Commonly, 2-3 ribs run longitudinally over most of the chamber sequence.

8.2 Bolivinita pohana Comparative Population Q18/f13

Material: Q18/f13 lies c. 400 m above the base of Mount Messenger Formation (?) in the north Taranaki coastal
section and closely above the top of a unit that ? identified as the Kaiti coiling zone. Probably, it represents a
higher horizon in the lower Tongaporutuan Stage than Y18/f7479 (type locality of Bolivinita pohana). Twenty-four
specimens were examined in spiral orientation; c. 100 in axial and distal orientations.

Spiral orientation: The range in the amount of offset between chambers is similar to that in topotypes but specimens
with minor offset are less common. The convexity of distal walls is often less (Fig. 8.4 #7) than in topotypes and
there are no strongly convex examples comparable with Fig. 8.1 #15. Keels are differentiated similarly to topotypes,

56
 DRAFT
Figure 8.4: Bolivinita pohana Finlay. Variation in the spiral outline of specimens from Q18/f13.

with the stronger structure between the longer axial and distal walls (Fig. 8.4 #18). Specimens like the latter show the
minor keel in spiral orientation but it is less obvious than in topotypes. In some specimens there is only an angular
junction (Fig. 8.4 #7, #16) between the shorter axial and distal walls. Re-entrants in axial walls are often shallower
than in topotypes and good examples of V-shapes like Fig. 8.1 #14 are rare. Apertural shape is often comparable with
topotypes but in some specimens (Fig. 8.4 #7) the upper arch is asymmetrical. Some of the distinctions between the
populations are summarized in Fig. 8.5.

Clusters formed from the pooled data (Fig. 8.5 (C)) contain specimens from both samples, although most are domi-
nated by one sample. This result is supported by the ordination (Fig. 8.5 (B)), which suggests that the variation fields
overlap. Specimens like the topotypes shown in Fig. 8.1 #4 (minor offset between chambers, elongated outlines) are
not present in Q18/f13.

Axial orientation: The axial surfaces of chambers are less inflated in Q18/f13 and sutures are nearly flush rather than
depressed. Throughout the chamber sequence the distal:shorter axial wall junction is often more angular and the keel
is more commonly continuous.

Distal orientation: There are no specimens in Q18/f13 with 2-3 ribs running the length of the distal walls. If present,
ribs occur only on the earliest chambers and are very weak.

Remarks: The distribution of some characters in populations beyond Kaiti Beach section often do not closely match
topotypes. For example, in spiral orientation specimens in which chamber offsets are minor, or distal walls are
moderately convex, are more common among topotypes than elsewhere. Angular junctions between shorter axial
and distal walls are rarely seen in topotypes but are common elsewhere. Some other differences are noted above.
Qualitatively, the most similar specimens occur in lower Tongaporutuan populations, particularly from Gisborne to
East Cape. Inclusion of populations (e.g., Q18/f13) from other basins, or even from the southern region of the Eastern
Basin, expands the concept of the species beyond that suggested by topotypes. Regional variation is suggested but
further study is needed to define the pattern. Possibly, populations like that in Q18/f13 should be discriminated from
B. pohana.

8.3 Discrimination Bolivinita pohana : B. compressa

Material: Although not remarked upon by ? in his original proposal, Bolivinita pohana is accompanied at its type
locality (Y18/f7479) by B. compressa. The separation of the species in this assemblage is discussed with reference to
the configuration in Fig. 8.1.

57
 DRAFT
Figure 8.5: (A) Bolivinita pohana Finlay. Variation in the spiral outline of specimens from Q18/f13 (specimens
#A-#C) and Y18/f7479 (specimens #1, #14). (B) Specimens (standardized for size) are plotted on 3-dimensional
coordinates based on principal components analysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the
outlines; 92% of variation is shown and the minimum spanning tree is drawn. (C) The same data are used in the cluster
analysis.

Spiral orientation: Several groups occur in the ordination of specimens using spiral outline data (Fig. 8.1 (B)). At the
right (including #4, #18) are specimens in which the offset between chambers is small and the outline is elongated,
due to the height of chambers. This group links with a larger group (including #14, #15, #17, #20) in which chamber
offset is again often minor but outlines are more compact (lower chambers). Distal walls in both groups are convex,
sometimes strongly so. Specimens in both groups are identified as Bolivinita pohana.

The offset between chambers sometimes is greater in the adjacent group (linked by Fig. 8.1 (B) #9) and some axial
walls show greater variation in relative length. Chambers are moderately offset in Fig. 8.1 (A) #7 and there is consid-
erable extension of the outline on the axis of the major keels (a feature of B. compressa). Yet its axial wall re-entrants
are still prominent and it has two keels. This specimen is assigned to Bolivinita pohana. Specimen #2 has weaker
axial re-entrants and its distal walls are less inflated. As the ordination indicates (Fig. 8.1 (B)), it resembles specimens
in the left group (#16, #22, #24, #25). Tentatively, it is assigned to Bolivinita compressa.

58
Specimens shown in Fig. 8.1 (B) as Bolivinita compressa were identified by qualitative inspection. They feature
significant offset of chambers, weakly convex distal walls; their axial re-entrants are very shallow, or absent. Inter-

DRAFT
estingly, only #16, #22, #24, #25 and #2 are nearest neighbours in the graph defined by the minimum spanning tree.
Three other specimens (#8, #12, #27) have nearest neighbours that I identify as B. pohana. However, all specimens
qualitatively identified as B. compressa map high on the y-axis. The data might suggest gradation in spiral shape
between B. pohana and B. compressa (Fig. 8.1 (A) #2 is a possible example).

Remarks: Chamber offsets are generally less in Bolivinita pohana than in B. compressa and there is lesser extention
of the outline along the major axis of the spiral outline; distal walls show greater convexity; re-entrants in axial walls
are deeper and V-shaped; differences in keel strength are smaller. In oblique axial orientation, the alate megalospheric
morphotype of B. compressa (Fig. 4.2 #8) has no counterpart in B. pohana populations.

8.4 Bolivinita pohana Biogeography

Figure 8.6: Distribution of Bolivinita pohana on

modern coordinates. Shaded areas are generaliza-
tions of known distributions and are conservative.
They do not identify possible changes in its distri-
bution through the Late Neogene.

Bolivinita pohana occurs throughout the Eastern Basin in lower Tongaporutuan assemblages. Although this basin
seems to have been its primary domain, it is not highly persistent and populations vary considerably in size. In the
section at Kaiti Beach (type locality) it is common up to Y18/f506 (?) but is very rare, or absent, in higher samples.
At some horizons (e.g., Y18/f539), it is replaced by B. compressa. South of Hawke's Bay it occurs with B. medialis
(e.g., V22/f8637, Tuki Tuki River; N32/f9207 and N32/f9209, Waiau River, north Canterbury) but the association is
irregular. Bolivinita pohana is absent from some assemblages that include B. medialis, although B. compressa may be
present (e.g., T27/f503 Karamu South Stream, (?). The apparently complex biogeographies of B. pohana, B. medialis
and B. quadrilatera complicate their use in the lower Tongaporutuan biostratigraphy of the basin.

In western New Zealand Bolivinita pohana is present in Taranaki Basin, Westland and western Southland, but popula-
tions are often smaller, less persistent, and less typical, architecturally, than those in Eastern Basin. However, Scott (in
?, p. 91; fig. 29 reported several assemblages from Mount Messenger Formation (Taranaki Basin) in which Bolivinita
pohana is strongly dominant and perhaps an opportunist.

Typical populations of Bolivinita pohana occur in middle and lower bathyal assemblages. In the Tongaporutuan
stratotype its replacement by B. urenuia is possibly related to upward shallowing of the basin.

59
8.5 Bolivinita pohana Stratigraphic Distribution

DRAFT
Tongaporutuan Stage, extending from the base into the lower part of the informal upper zone whose base is defined
by the highest occurrence of Globoquadrina dehiscens (Fig. 1.1).

Lowest occurrence: Scott (in ?) reviewed literature on criteria for the base of the Tongaporutuan Stage and, following
?, considered that the lowest occurrence of Bolivinita pohana was an important defining event. ? showed that the
event precedes the base of a coiling excursion in Globorotalia miotumida (Kaiti coiling zone).

Highest occurrence: In the Tongaporutuan stratotype (north Taranaki coast) the highest definite record is Q18/f54
(shown in ?, appendix 2 sheet 31). This horizon is c. 5 m above the highest occurrence of Globoquadrina dehiscens
(Q18/f1). ? identified Bolivinita pohana from upper Tongaporutuan strata at Blind River, Marlborough but this
material is here identified as B. pliobliqua. In the northern Hawke's Bay sequence at Hangaroa River B. pohana occurs
with the highest record of G. dehiscens (X18/f15). It is replaced in the next collection (X18/f14, c. 230 m higher) by a
population tentatively referred to B. urenuia. In the same region at Mangapoike River B. pohana is present in X19/f55
(c. 250 m above the highest occurrence of G. dehiscens).

Whereas ? and ? considered that Bolivinita pohana extended to the Kapitean boundary, this review restricts it primar-
ily to lower Tongaporutuan assemblages (below the highest occurrence of Globoquadrina dehiscens). Difficulty in
discriminating some late populations from B. urenuia compromises the utility of the highest occurrence of B. pohana
in biostratigraphy. In this survey it does not show a consistent relationship with the highest occurrence of Globoquad-
rina dehiscens.

60
Chapter 9

DRAFT
Bolivinita pliobliqua Vella

1963 Bolivinita pliobliqua Vella, Transactions of the Royal Society of New Zealand, Geology 2: p. 7, pl. 1, fig.
12-13.

Original Description

"Shell similar to pliozea Finlay but rhomboid in transverse section and generally narrower and longer; slightly twisted,
tapering gently from the adult end to the bluntly rounded initial end. Sides smooth, concave, each bounded by its own
pair of prominent longitudinal keels. Periphery separated from sides by the keels, gently convex, ornamented by two
to four longitudinal ribs which are continuous across the sutures." (?)

9.1 Bolivinita pliobliqua Type Population T26/f6910 (Recollection)

Material

Specimens are from a recollection of T25/f6910 (type locality), Mangaoranga Formation, Makakahi River near Eke-
tahuna. ? placed the formation in the Tongaporutuan Stage. Twenty-five specimens (including 6 that are damaged)
were studied in spiral orientation; c. 100 specimens were studied in axial and distal orientations.

Spiral Orientation

Figure 9.1: Bolivinita pliobliqua Vella. Variation in the spiral outline of specimens from T26/f6910.

Chamber arrangement: Chambers in the biserial arrangement are always offset although the amount is sometimes
small (Fig. 9.1 #22). Normally, the coiling axis is linear but significant flexure is shown in Fig. 9.1 #10. The major
diameter of the shell lies in the direction of the offset. The spiral outline of specimens is rhomboidal, but is usually
deformed by re-entrants formed by the axial walls.

61
Distal wall: The boundaries are well-defined when a keel is present at each junction with an axial wall (Fig. 9.1 #1). In
some specimens the boundary with the shorter axial wall may not have an obvious keel but is still well-defined by an

DRAFT
angular junction (Fig. 9.1 #13 upper). More rarely, the angular junction is replaced by a curve (Fig. 9.1 #19). The distal
wall is always convex, although curvature is variable. Usually, the wall rises above the bordering keels (Fig. 9.1 #25).
The wall profile is commonly featureless in this orientation; occasionally, ribs are visible (Fig. 9.1 #25).

Axial walls: Walls are unequal in length, although the difference may be minor (Fig. 9.1 #17). They diverge out from
the base of the chamber and are often linear in profile. The longer wall is adjacent to the keel at the major axis of the
shell. The re-entrant in the shell outline formed by adjacent axial walls varies from an an open, weakly asymmetrical
V (Fig. 9.1 #17) to a very shallow depression (Fig. 9.1 #16). Rarely, the re-entrant is not visible (Fig. 9.1 #24) but this
may be affected by the precise orientation of the shell.

Keels: The major keel at the junction of the longer axial and distal walls is well-defined on larger specimens and is
broad on some (Fig. 9.1 #19). It extends to the aperture where it links with its equivalent on the opposite chamber
(Fig. 9.1 #17). On some smaller specimens (Fig. 9.1 #7, #16) this keel is very low and does not form a topographic
prominence. Generally, the minor keel at the junction of the shorter axial and distal walls is weaker. It may not
extend to the aperture (Fig. 9.1 #25). In this orientation the structure does not appear in small specimens in which the
major keel is very low (Fig. 9.1 #7). However, the development of major and minor keels is not always correlated: in
Fig. 9.1 #19, a large individual, the major keel is robust but there is no minor keel on the last chamber.

Aperture: Although there are some examples of symmetrical, inverted-U openings (Fig. 9.1 #16), most are asymmet-
rical, with the side adjacent to the major keel showing greater curvature (Fig. 9.1 #19). The base is flat. The tooth in
the upper part of the opening is robust (Fig. 9.1 #7).

Axial Orientation

Figure 9.2: Bolivinita pliobliqua Vella. Guide to morphology of specimens from T25/f6910 in standard orientations.

Shell outline: The prolocular section of megalospheric specimens is usually significantly wider than that of micro-
spheric specimens. Lateral sections are defined by the crest of the distal walls, although keels at the distal - axial
wall junctions lie close to the periphery in some specimens (Fig. 9.2 #3). Lateral sections tend to diverge at a greater
angle in microspheric specimens and maintain the angle through most of ontogeny (Fig. 9.2 #2). In megalospheric
specimens the angle often declines after about the third chamber pair and the section is slightly convex (Fig. 9.2 #3)
with maximum width near the base of the last pair of chambers. Lobulation is best defined on late chambers but is
commonly weaker than is shown in Fig. 9.2 #1. The shape of the apex above the aperture is variable (Fig. 9.2 #2, #3).

Chambers: There are up to c. 8 pairs of wedge-shaped, steeply inclined, closely packed chambers. Axial walls incline
towards the coiling axis; their surfaces include planar areas but most are slightly convex near the margins, creating
weakly depressed sutures. In some specimens the chambers are flush (Fig. 9.2 #3). Keels at the margins of axial
walls extend through the chamber sequence. Commonly, they are differentiated, with the major keel (at the junction
of longer axial and distal walls) tapering outward to a thin flange while the minor keel may be discontinuous, and not
extend fully over the last, or penultimate chambers (Fig. 9.2 #1 right). Pores are subcircular, small, and distributed

62
 DRAFT
Figure 9.3: Bolivinita pliobliqua Vella T25/f6910. Wall
topography.

fairly evenly over walls (Fig. 9.3 #6-#8). There is no secondary calcification around pores. A weak, short rib occurs
in the prolocular region of some specimens (Fig. 9.2 #2); it does not extend onto later chambers.

Figure 9.4: Bolivinita pliobliqua Vella. Variation in the spiral outline of specimens from P29/f16.

Distal Orientation

Shell outline: The lateral sections are defined by keels at the margins of the distal walls. Weak flexure of the coiling
axis may bring a little of the axial wall into view (Fig. 9.2 #4). Lateral sections diverge outward from the proloculus
to the base of the last chamber. Over this interval their form is almost linear, with only very weak re-entrants at some
sutures. The expansion in width of the last chamber relative to its predecessor in Fig. 9.2 #4 is greater than usual.
The lateral outlines of the last chamber contract towards the apertural section. This is often flat and well-defined
(Fig. 9.2 #4, #5); occasionally it is reduced to a narrow apex. The prolocular section is rounded, and narrower in
microspheric than in megalospheric specimens.

Chambers: The lower margin is linear and usually normal to the direction of the coiling axis; some are oblique
(Fig. 9.2 #4). Sides of chambers are slightly convex and their height increases rapidly through ontogeny. Wall surfaces
are gently convex and sutures are slightly depressed. There are 1-3 low, narrow ribs on the proloculus and earliest
chambers. Commonly, at least one runs up to the base of the last pair of chambers (Fig. 9.2 #4) but on some specimens
all are restricted to the early chambers. Occasionally, ribs run tangentially to the coiling axis.

63
9.2 Bolivinita pliobliqua Comparative Population P29/f16

DRAFT
Material

P29/f16 (Upton Formation, Blind River Fig. 4.5) is the lowest sample in the Blind River sequence in which Bolivinita
pliobliqua is identified. ? identified the material as B. pohana and noted that this was the lowest horizon with speci-
mens having some characters of B. pliozea.

Spiral orientation: As in the topotype population (T26/f6910), many shells are compact. Chambers are offset in all
specimens, creating rhomboidal outlines. Occasionally, the amount of offset is slight (Fig. 9.4 #17). There are only
small differences in the length and orientation of axial walls. The V-shaped re-entrants formed by axial walls are
wide and well-defined on most specimens. They resemble topotypes like Fig. 9.1 #17 but are more common than in
that population. The deepest re-entrants in the sample are seen in Fig. 9.4 #10 which, like a comparable topotype
(Fig. 9.1 #10), occur on a specimen in which the coiling axis is flexed. Weakly defined re-entrants, seen in some
topotypes (Fig. 9.1 #7, #16), do not occur in the sample.

The keel between the shorter axial and distal walls is often better developed (Fig. 9.4 #6), and more commonly present,
than in topotypes. Distal walls are gently convex and rise little higher than the keels. Commonly, walls are less convex
than in topotypes (e.g., Fig. 9.1 #25).

Axial, distal orientations: Keels at the margins of axial walls are well-defined. Generally, they are more prominent
than in topotypes. The keel at the longer axial - distal wall junction often tapers outward to a flange. Usually,
distal walls have a median rib on early chambers; it weakens on later chambers and may not extend to the last pair
(cf. Fig. 9.2 #5). Less commonly, there are 2-3 such ribs.

Figure 9.5: Bolivinita pliobliqua Vella. Variation in the spiral outline of specimens from P29/f18. A specimen of
Bolivinita compressa from this locality is included.

64
9.3 Bolivinita pliobliqua Comparative Population P29/f18

DRAFT
Material

Blind River, c. 30 m above P29/f16 (Fig. 4.5).

Spiral orientation: Some major features (rhomboidal outline, chamber offset, axial wall re-entrants, keel at both axial
- distal wall junctions) compare with those in P29/f16. However, distal walls on some specimens are more convex and
rise further above the keels (Fig. 9.5 #4). Compact outlines remain common and, as in P29/f16, small specimens with
weak or inconspicuous keels are absent. Some individuals (e.g., Fig. 9.5 #16) anticipate forms seen in early B. pliozea
(e.g., Fig. 10.4 #13). Specimens like Fig. 9.5 #9 are referred to B. compressa.

9.4 Bolivinita pliobliqua Comparative Population T26/f139

Material

Te Kowhai Stream section (?, appendix 1, section 2). Bells Creek Mudstone, c. 70 m below top. Twelve specimens
were studied.

Spiral orientation: The convexity of distal walls in some specimens (Fig. 4.7 (A) #3) compares with topotypes
(Fig. 9.1 #25) and the amount of offset between chambers is similarly small (Fig. 4.7 (A) #14). Axial wall re-entrants
are generally V-shaped but often shallower (e.g., Fig. 4.7 (A) #14) than in P29/f16 and P29/f18 (e.g., Fig. 9.5 #4).
Note that the narrow, deep V-shaped re-entrant on Fig. 4.7 (A) #1 is associated with a strongly convex distal wall, an-
ticipating characters found in B. pliozea. Small, compact specimens with weak keels (Fig. 4.7 #12) have counterparts
among topotypes (Fig. 9.1 #7). Specimens like Fig. 4.7 (A) #8 have some features of the accompanying B. compressa
(relatively elongate outline, strong major keel) but have deeper axial re-entrants and chambers that are only moderately
offset.

9.5 Discrimination Bolivinita pliobliqua : B. pohana

Material

Bolivinita pliobliqua from T25/f6910, Eketahuna (type population). Bolivinita pohana from Y18/f7479 (Kaiti Beach,
Gisborne, (type population) and Q18/f13 (Taranaki coastal section).

Spiral orientation: Outlines in both taxa are rhomboidal (Fig. 9.6 (A) #B). The amount of offset between chambers
in topotypes is often similar, although very minor offsets are more common among topotypes of Bolivinita pliobliqua
than among topotypes of B. pohana. Convex distal walls are developed in both topotypic populations. In B. pohana
re-entrants formed by axial walls are often significantly asymmetrical and may be shallow. V-shaped re-entrants in
axial walls are better developed in some B. pliobliqua because of the small difference in lengths of opposed shorter
and longer axial wall segments, and their greater linearity. Specimens of B. pliobliqua as in Fig. 9.1 #25 do not have
counterparts in B. pohana populations. If present, the keel between the shorter axial and distal walls is often weaker
in B. pohana.

Analysis of spiral outline data for topotypes of the taxa (Fig. 9.6 (B)) produced one low-level cluster of Bolivinita
pohana but the main cluster is a mixture of both species. The result confirms the visual impression that many outlines
are closely similar. In spiral orientation, discrimination of the taxa relies on the presence of distinctive specimens of
B. pliobliqua (e.g., Fig. 9.1 #25; Fig. 9.6 (A) #22). These may be rare.

65
 DRAFT
Figure 9.6: (A) Discrimination of Bolivinita pliobliqua (T25/f6910) from Bolivinita pohana (Y18/f7479) using char-
acters seen in spiral and axial orientations. (B) Cluster analysis using 20 coefficients for 5 elliptical Fourier harmonics
(specimens standardized for size) fitted to outlines in spiral orientation.

Axial and distal orientations: In a comparison of topotypes, lobulation of the outline is greater in Bolivinita pohana
because of the greater convexity of distal walls. To a lesser extent, this applies to axial walls. Thus sutures are
depressed in B. pohana (Fig. 9.6 (A) #i) but almost flush in B. pliobliqua (Fig. 9.6 (A) #ii). Secondary calcification
around pores is not present in topotypes of B. pliobliqua (Fig. 9.3 #6-#8, cf. Fig. 8.3 #6-#8) but the value of this
character as a discriminant is uncertain.

9.6 Discrimination Bolivinita pliobliqua : B. urenuia

Material

Bolivinita pliobliqua from P29/f16, Blind River (lowest record, Fig. 4.5) and B. urenuia from an underlying sample
(P29/f14) are compared in spiral orientation. Topotypes of both taxa are considered in the discussion of axial and
distal orientations.

Spiral orientation: Outlines are rhomboidal in both samples but shapes tend to be more elongated in Bolivinita urenuia
(e.g., Fig. 9.7 (A) #B, #15) and the lengths of axial walls are better differentiated. Offset of chambers is greater in
many B. urenuia. Distal walls in some specimens of B. pliobliqua (Fig. 9.7 (A) #B) are more convex than in B.
urenuia, although the variation fields overlap. Re-entrants formed by axial walls in B. urenuia are shallower than in
B. pliobliqua and V-shapes are often less regular. Particularly, there are no specimens like Fig. 9.7 (A) #B that have

66
 DRAFT
Figure 9.7: A) Discrimination of B. pliobliqua (P29/f16) from B. urenuia (P29/f14) using characters seen in spiral
orientation. (B) Specimens (standardized for size) are plotted on 3-dimensional coordinates based on principal com-
ponents analysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 90% of variation is shown
and the minimum spanning tree is drawn.

an indistinct U-shaped re-entrant (subsequently better developed in B. pliozea). The keel at the junction between the
longer axial and distal walls is often more prominent in B. urenuia. The ordination (Fig. 9.7 (B)) indicates that the
variation fields are adjacent, with only minor overlap.

Axial, distal orientation: Lobulation of lateral sections (axial orientation) is more conspicuous in some Bolivinita
pliobliqua (Fig. 9.2 #1) than in B. urenuia Fig. 7.2 #1. The natural repose position of many B. urenuia is significantly
oblique to the axial position and resembles that in Fig. 7.2 #4. Lesser offsets in B. pliobliqua result in positions of natu-
ral repose of shells closer to the axial orientation (Fig. 9.2 #3). Thus in a strew, the width of specimens of B. pliobliqua
is often narrower than in B. urenuia. The flange on the major keel of B. urenuia is more commonly present, wider,
and more conspicuous than on B. pliobliqua. The minor keel (between shorter axial and distal walls) is consistently
present in B. urenuia and often robust. In B. pliobliqua the structure is weaker and sometimes discontinuous. Ribs,
sometimes multiple, are more extensive on distal walls of B. pliobliqua (Fig. 9.2 #4) than on B. urenuia (Fig. 7.2 #6).

9.7 Discrimination Bolivinita pliobliqua : B. pliozea

Material

Bolivinita pliobliqua from T25/f6910, Eketahuna (type population) is compared with B. pliozea in an early population
(P29/f29, Blind River). Comparative features of some later populations of B. pliozea are noted.

Spiral Orientation: Symmetry of the spiral outline is an important discriminator. Typically, chambers in Bolivinita
pliobliqua are offset and the outline of shells resembles a rhomboid (Fig. 9.8 (A) #20). In Bolivinita pliozea offsets
between chambers are usually small and outlines are often bilaterally symmetrical (Fig. 10.4 #25).The difference in

67
chamber offset affects the shape of the aperture, which is often more asymmetrical in B. pliobliqua than in B. pliozea.
The elongate outlines of some B. pliobliqua (Fig. 9.8 (A) #15) do not develop in B. pliozea populations. Re-entrants

DRAFT
formed by axial walls are usually deeper and narrower in B. pliozea (Fig. 9.8 (A) #A), but note the shallow re-
entrants of the Kapitea Creek populations (Fig. 10.6 #2). Distinctive, weakly U-shaped re-entrants feature in Pliocene
- Quaternary populations of B. pliozea (Fig. 10.4 #13) but do not occur in B. pliobliqua. Keels in B. pliobliqua
are commonly unequal and project obliquely. In B. pliozea they are not differentiated and often project laterally in
Pliocene - Quaternary populations. However, this feature is poorly developed in early populations like P29/f29. Early
populations of Bolivinita pliozea show closest resemblance with B. pliobliqua, particularly in convexity of distal walls
and form of axial wall re-entrants. However, ordination of topotypes of B. pliobliqua and a sample of early B. pliozea
(Fig. 9.8 (B)) indicates very minor intersection of their variation fields. Characters related to shell symmetry in spiral
orientation are useful discriminators.

Axial and distal orientations: The axial walls are narrower in many Pliocene - Quaternary specimens of Bolivinita
pliozea, sometimes conspicuously so (Fig. 10.2 #1). However, this character does not discriminate some earlier
specimens from B. pliobliqua (Fig. 9.8 (A) #A). Similarly, while the number of ribs on the distal walls (3-4) helps
distinguish many Pliocene - Quaternary populations, the early populations often have up to two, as in B. pliobliqua.

Figure 9.8: (A) Discrimination of early specimens of B. pliozea (P29/f29) from topotypes of B. pliobliqua (T25/f6910)
using characters seen in spiral orientation. Specimens in axial orientation also shown. (B) Specimens (standardized
for size) are plotted on 3-dimensional coordinates based on principal components analysis of 20 coefficients for 5
elliptical Fourier harmonics fitted to the outlines; 95% of variation is shown and the minimum spanning tree is drawn.

9.8 Overview

? synonymized Bolivinita pliobliqua with B. pohana. This interpretation, followed by ?, is supported by the archi-
tectural resemblance (spiral orientation) of some topotypes (e.g, Fig. 9.6 (A) #B, #15) but is less tenable when more

68
aspects of morphology are considered. These data, together with better understanding of the distribution of B. pohana
and its variation, suggest that B. pliobliqua may be a homoplastic design that arose subsequent to the extinction of B.

DRAFT
pohana, or after its migration from the New Zealand region.

In the sequence at Blind River (Fig. 4.5) ? identified Bolivinita cf. pohana in P29/f11 and P29/f13, B. pohana from
P29/f16 to P29/f26, and placed the lowest occurrence of B. pliozea at P29/f29. This study identifies B. compressa in
P29/f11 (Fig. 4.6 #16), B. urenuia in P29/f14 (Fig. 7.6 #3), B. pliobliqua in P29/f16 and P29/f18 (Fig. 9.5 #16), and
B. pliozea in P29/f29 (Fig. 10.4 #13).

The principal revision is the assignment of populations between P29/f16 - P29/f28 to Bolivinita pliobliqua. A few
topotypes of the latter (e.g., Fig. 9.1 #25) quite closely resemble some in P29/f16 (Fig. 9.4 #6). Significant features
in common are compact spiral outline, minor offset of chambers, convex distal walls, minor differentiation of axial
walls, and well-defined V-shaped re-entrants formed by axial walls. Such individuals might be envisaged as prototypes
of B. pliozea but are distinguished architecturally by the offset of chambers and consequent departure from bilateral
symmetry. There are also rare specimens (Fig. 9.1 #10, Fig. 9.4 #10) in which a pliozea-like spiral profile occurs in
a shell with a flexed coiling axis. Specimens shown in Fig. 9.1 #1 and Fig. 9.4 #18 represent another morphotype
(distal walls only slightly convex) that occurs in topotypes and at Blind River. Their spiral outlines resemble some
weakly rhomboidal B. urenuia (Fig. 7.4 #4). However, the principal contrast with topotypes is the absence from
Blind River populations of specimens with very weak keels (Fig. 9.1 #7). Bolivinita pliobliqua is interpreted as a
polymorphic species with several morphotypes, not all of which may be present in each population. Further research
on this interpretation is warranted.

? summarized changes in morphology in Blind River material now referred to Bolivinita pliobliqua and wrote of
progressively greater resemblance with B. pliozea. High resolution mappings of morphology are required to confirm
this view and it may be an oversimplification. The evolution of B. pliozea from B. pliobliqua may not closely match
the classical phyletic model of directional trends in simple multivariate normal populations. Prototypes of B. pliozea
appeared early in the history of B. pliobliqua at Blind River; some have abnormal shell geometry (flexed coiling
axis). Bolivinita pliobliqua shows a spectrum of morphotypes and none is strongly dominant. The major architectural
changes (development of bilateral symmetry, more compact chambers, inflation of distal walls, V- to U-shaped re-
entrants in axial walls) form only weakly defined trends at Blind River. Changes in the relative dominance of partially
discrete morphotypes may have played a role in its history.

Bolivinita pliobliqua had a short history (c. 0.5 Ma) relative to other taxa. Its populations record architectural trans-
formations that led to a new, stable, and long-lived design (B. pliozea). There is no evidence of lineage splitting.
From an evolutionary perspective, Bolivinita pliobliqua possibly represents an episode of relatively rapid phyletic
transformation (punctuated anagenesis (?). Principal uncertainties are the identity of the ancestor and the population
mechanisms by which the transformation was achieved. Although Bolivinita urenuia precedes B. pliobliqua in the
Blind River sequence, this may not identify it as the ancestor. A specimen (Fig. 7.6 #16) in P29/f14 resembling B.
pliobliqua maps as an outlier in the ordination.

At Blind River the range of Bolivinita pliobliqua coincides closely with a shift towards lighter values of 13 C recorded
in foraminiferal carbonate (?). ? considered that the event (their Messinian carbon shift) primarily reflected exchange
between the ocean and an external organic carbon reservoir, with fractionation between the Indo-Pacific and Atlantic
basins, and increased productivity as secondary causes. However, its effects on bottom environments are poorly
understood and any connection with the evolution of B. pliobliqua is speculative.

9.9 Bolivinita pliobliqua Biogeography

In the north of the Eastern Basin (Fig. 9.9) Bolivinita pliobliqua occurs in the base of Tokomaru Sandstone (Z14/f8) in
a middle bathyal assemblage. It is also present in Hangaroa River section (X18/f12), at Mangapoike River (X19/f7732,
Mapiri Formation; ?, fig. 100-102) and in the west of the basin (V20/f386) in an upper bathyal assemblage. It has
not been recorded from southern Hawke's Bay but further south there are records from northern Wairarapa through to
Marlborough. This sector of the basin has the largest and most persistent populations of B. pliobliqua. In addition to

69
 DRAFT
Figure 9.9: Distribution of Bolivinita pliobliqua on
modern coordinates. Shaded areas are generalizations
of known distributions and are conservative. They
do not identify possible changes in its distribution
through the Late Neogene.

records near the top of the Tongaporutuan stratotype (e.g., Q19/f3), there are subsurface records in Taranaki Basin in
Kiwa-1 (1640 m), Kahawai-1 (1945 m) and possibly in Maui-2 (3090 ft).

Bolivinita pliobliqua occupied shelf to mid-bathyal environments. Usually, it is rare in assemblages in which Bolivinita
compressa is a dominant member (e.g., Hangaroa River); conversely, B. compressa is rare in assemblages (e.g., as at
Blind River) in which B. pliobliqua is common. Possibly, this reflects their differing bathymetric preferences.

9.10 Bolivinita pliobliqua Stratigraphic Distribution

Tongaporutuan Stage (informal upper zone Fig. 1.1).

Lowest occurrence: The best resolved record is at Blind River where the species appears in the vicinity of P29/f16
(Fig. 4.5) in strata classified as upper Tongaporutuan. Extensive sampling has not revealed the occurrence of Globo-
quadrina dehiscens at Blind River and it is inferred that the sequence lies above its last occurrences. Paleomagnetic
data (?; ?) support this interpretation. ? and Scott (in ?) considered that Bolivinita pohana occurred at the top of
Urenui Formation in the Tongaporutuan stratotype. Although the paucity of specimens constrains assessment, the ma-
terial figured by ?, fig. 92-94 from N99/f540 and three similar specimens in Q19/f3 are here assigned to B. pliobliqua.
These upper Tongaporutuan horizons overlie those with B. urenuia.

Highest occurrence: At Blind River this event is marked by the appearance of Bolivinita pliozea and is close to the
appearance of Kapitean Mollusca. Note that some variants (usually small) in B. pliozea populations (e.g., Fig. 10.1
#27) have counterparts among B. pliobliqua topotypes (Fig. 9.1 #7). On a strictly morphotype basis (not followed
here) the species would extend into the Quaternary.

70
Chapter 10

DRAFT
Bolivinita pliozea Finlay

1939 Bolivinita pliozea Finlay, Transactions of the Royal Society of New Zealand 69: p. 319, pl. 27, fig. 101-2.

Original Description

"..differing from pohana in having hardly any obliquity and very convex sides, with 3-5 sharp linear equidistant ridges
between the main keels, which are much closer together and enclose a deeply hollowed median smooth area." (?)

10.1 Bolivinita pliozea Reference Population R22/f7516

Material

R22/f7516 is from Pinnacle Sand (?, fig. 38) in the section at Castlecliff, Wanganui; Castlecliffian Stage. It is c. 10 m
below the type locality of Bolivinita pliozea (F5214). Spiral morphology was studied on 27 specimens; >40 specimens
were examined in axial and distal orientations.

Spiral Orientation R22/f7516

Figure 10.1: Bolivinita pliozea Finlay. Variation in the spiral outline of specimens from R22/f7516.

Chamber arrangement: Offsets between chambers in the biserial shells are often very small or negligible. Because of
this, many shell outlines are almost bilaterally symmetrical (Fig. 10.1 #6). Some are slightly asymmetrical (Fig. 10.1

71
#3) due to unequal axial walls. The convexity and extent of the distal walls contribute to the oval to sub-circular
appearance of some outlines (Fig. 10.1 #4), although they are disrupted by the axial wall re-entrants.

DRAFT
Distal wall: The outline is a regular convex curve, with the crest located centrally above the aperture (Fig. 10.1 #26).
There are angular junctions with the axial walls. On most specimens the outline is interrupted by end sections of 2-4
ribs (approximately equidistant, but often located nearer to the centre than to the margins).

Axial walls: Typically, their length and orientation is similar in both chambers and the profiles vary from linear to
weakly concave. The inclination of the walls creates a distinctive deep, relatively narrow channel-like V- to U-shaped
re-entrant (e.g., Fig. 10.1 #6). In a minority of specimens (e.g., Fig. 10.1 #3) the re-entrant is shallower and broader.

Keels: In most specimens each axial - distal wall junction is marked by a narrow but protruding keel. Typically, it
projects almost laterally as a rounded ridge above the adjacent wall surfaces and links with the aperture. Keels are
commonly of equal size. However, in some small individuals (Fig. 10.1 #27) only one low structure is visible. There
are comparable and usually small specimens in populations of Bolivinita pliobliqua (Fig. 9.1 #7).

Aperture: The opening is relatively high and narrow on some individuals (Fig. 10.1 #4). The base, which is defined
by the distal wall of the (n-1)th chamber, is weakly convex. The sides are usually linear for over 60% of their length
and incline gently inward. The top section varies from a well-defined low arch (occasionally almost flat), to a poorly-
defined high arch which merges with the sides. On some, a symmetrical "A" shape is suggested by the inwardly
inclined high sides and flattish top (Fig. 10.1 #17). The tooth, which represents the folded edge of the toothplate, rises
from the top section and is long (often >50% of the height of the opening), narrow, and weakly curved to linear.

Axial Orientation R22/f7516

Figure 10.2: Bolivinita pliozea Finlay. Guide to morphology of specimens from R22/f7516 in standard orientations.

Shell outline: The lateral sections are defined by the rib(s) nearest to the apex of each distal wall. Although the
distal walls have lobulate profiles due to their convexity, the ribs bridge the major re-entrants at chamber junctions
and considerably dampen lobulation in the shell outline. Lateral outlines diverge outward from the rounded prolocular
section. Divergence is greatest in microspheric specimens (Fig. 10.2 #2) and maximum width is reached about the base
of the last pair of chambers. This also applies to some megalospheric specimens. In others (Fig. 10.2 #1) maximum
width is reached in mid-ontogeny and changes little over the last 2-3 chamber pairs. The apex above the aperture is
broadly rounded.

Chambers: Up to c. 9 pairs of wedge-shaped chambers, biserially arranged and steeply inclined. The axial walls
are weakly concave, particularly in specimens in which these walls form a deep, channel-like re-entrant in the spiral
outline. The base of late-formed chambers often preserves a relict keel. Sutures are weakly depressed or almost flush.
Pores are elliptical to circular (Fig. 10.3 #6) and often large. Axial walls are generally featureless, without ribs or

72
 DRAFT
Figure 10.3: Bolivinita pliozea Finlay R22/f7516. Wall
topography.

secondary calcification around pores. However, several ribs on distal walls are seen in this orientation.

Distal Orientation R22/f7516

Shell outline: The lateral sections are defined by keels at the junction of axial and distal walls. The sections diverge
out from the prolocular section at lower angles than in axial orientation, but the location of maximum width of the shell
is similarly variable and related to the size of the proloculus. Flexure of the coiling axis occurs in a few specimens
and is more obvious than in axial orientation. The apertural section is sometimes narrow (Fig. 10.2 #5) but there are
specimens in which it is broad, with little curvature.

Chambers: The gross shape is quadrate. In detail, lateral margins are slightly convex and lower and upper margins are
either linear or slightly concave. Upward concavity reflects overlap by a later chamber. Surfaces are convex in both
lateral and longitudinal directions. Sutures are weakly depressed (Fig. 10.2 #4). Principal features on walls are 2-4
(commonly 3-4) longitudinal ribs spaced approximately equidistantly across the surface (Fig. 10.2 #5). Ribs near the
centre of chambers are continuous throughout the chamber sequence. Some ribs near the periphery may be restricted
to several late-formed chambers. There is a strong keel at each lateral margin. Pores resemble those on axial walls.
There is considerable variation in size (Fig. 10.3 #7) and some are notably large.

10.2 Bolivinita pliozea Comparative Population P29/f29

Figure 10.4: Bolivinita pliozea Finlay. Variation in the spiral outline of specimens from P29/f29.

73
Material

DRAFT
Blind River section, Marlborough, (?) c. 550 m above base of Upton Formation (Fig. 4.5; ?). ? described a gradation
in the upper Tongaporutuan sequence at Blind River between Bolivinita pohana (populations here regarded as B.
pliobliqua) and B. pliozea. He identified P29/f29 as the lowest horizon in which the majority of specimens were
identifiable as B. pliozea.

Figure 10.5: Bolivinita pliozea Finlay. (A) Variation in the spiral outline of specimens from P29/f29 and R22/f7516.
(B) Variation in the spiral outline of specimens from P29/f29 and R22/f7516. Specimens (standardized for size) are
plotted on 2-dimensional coordinates based on principal components analysis of 20 coefficients for 5 elliptical Fourier
harmonics fitted to the outlines; 93% of variation is shown and the minimum spanning tree is drawn.

Spiral orientation: As in R22/f7516, there is little or no offset between chambers and little differentiation in the
length and orientation of the axial walls. Many outlines are bilaterally symmetrical or nearly so (Fig. 10.4 #25). The
ordination of the combined data on spiral outlines (Fig. 10.5 (B)) indicates that the variation fields of the samples
partially overlap.

Axial, distal orientations: The margins of the axial walls are often less inset from the lateral outline than in R22/f7516.
This is due to weaker inflation of the distal walls. Two ribs on the distal walls are typical in P29/f29, whereas 3-4
occur in R22/f7516. Some specimens in P29/f29 are distinguished by a robust rib that is centrally located on the distal
wall.

10.3 Bolivinita pliozea Comparative Population J32/f44

Material

J32/f44 samples Eight Mile Formation c.10 m above its base in the Kapitean stratotype (?, fig. 2). It is the lowest

74
 DRAFT
Figure 10.6: Bolivinita pliozea Finlay. Variation in the spiral outline of specimens from J32/f44.

collection available with sufficient material for quantitative study. ? identified Bolivinita pliozea from an adjacent
horizon. Comparisons are made with P29/f29, Blind River, as both are regarded as early populations of B. pliozea.

Spiral orientation: Offsets between chambers are small (Fig. 10.6 #24) and most shells approach bilateral symmetry.
Many shells are small and have compact outlines. Re-entrants formed by axial walls are V-shaped but are often shallow
(Fig. 10.6 #2). Distal walls are distinctly convex (Fig. 10.6 #10). Keels are orientated obliquely, as in P29/f29, but
they are commonly low and barely projecting (Fig. 10.6 #17). This feature occurs occasionally in other populations of
B. pliozea (e.g., Fig. 10.1 #27), but it is significantly more frequent in this population. The ordination analysis of spiral
outline data from J32/f44 and P29/f29 (Fig. 10.7 (B)) indicates that their variation fields are adjacent and marginally
intersect.

Axial, distal orientations: Commonly, keels are located near the lateral periphery (Fig. 10.7 (A) #e). They are further
inset in Fig. 10.7 (A) #f, as is common in R22/f7516 (Fig. 10.2 #1).

Remarks: Shells like those in J32/f44 continue into Opoitian strata at Kapitea Creek (e.g., J32/f9775, ?, fig. 78).
Possibly, they are only local variant populations in the Westland region as typical specimens of B. pliozea (Fig. 10.7
(A) #a-#d) occur nearby in a lower Kapitean assemblage from Kumara Anticline (Corehole D, 35 ft). At Blind River,
specimens like the near-topotypes of R22/f7516 (e.g., axial re-entrants form deep, channel-like re-entrants, keels
project laterally) become common in upper Kapitean horizons (e.g., P29/f60, P29/f62). These reconnaissance data
suggest that several variant populations might have existed in the lower Kapitean. Their status needs further study.

10.4 Discrimination

Refer to page 67 for discrimination from Bolivinita pliobliqua.

Refer to page 83 for discrimination from Bolivinita medialis.

75
 DRAFT
Figure 10.7: Bolivinita pliozea Finlay. (A) Variation in the spiral, axial and distal outlines of specimens from J32/f44
and P29/f29. (B) Variation in the spiral outline of specimens from J32/f44 and P29/f29. Specimens (standardized
for size) are plotted on 3-dimensional coordinates based on principal components analysis of 20 coefficients for 5
elliptical Fourier harmonics fitted to the outlines; 95% of variation is shown and the minimum spanning tree is drawn.

10.5 Bolivinita pliozea Biogeography

Bolivinita pliozea was distributed from Southland to East Cape in Kapitean time and in the Pliocene - Pleistocene
occupied all marine basins in the New Zealand region (Fig. 10.8). It extended from shelf down to middle bathyal sites,
with central populations on the upper slope. ? inferred that the Pinnacle Sand (source of the reference population)
might have been deposited at about 180 m, although shallower-dwelling Mollusca are also present. It is commonly
absent from shallow shelf assemblages. However, the Recent record in Pauatahanui Inlet near Wellington (?), if
authentic, would indicate the existence of some inner shelf peripheral populations. In the Pliocene and Pleistocene the
species was widely distributed and almost ubiquitous in outer shelf and upper bathyal assemblages. There are a few
records from middle bathyal or deeper loci (Taranga-1 1600m; DSDP Site 592, (?); ODP Site 1119, (B.W. Hayward
June 2000 pers. comm.).

76
 DRAFT Figure 10.8: Distribution of Bolivinita pliozea on modern
coordinates. Shaded areas are generalizations of known
distributions and are conservative. They do not identify
possible changes in its distribution through the Late Neo-
gene.

In overview, the biogeographic features that distinguish Bolivinita pliozea from others in this study are its good
distribution in all basins and persistence within its main bathymetric range. It may also have followed a different
population strategy from taxa like B. pohana and B. compressa. Blooms of B. pliozea have not been observed and
population sizes are usually modest. This suggests that it was an equilibrium species rather than an opportunist. ?
showed that its disappearance was part of a global extinction of various uniserial and biserial benthic taxa during the
mid-Pleistocene Climate Transition.

10.6 Bolivinita pliozea Stratigraphic Distribution

Kapitean Stage to Castlecliffian Stage (Fig. 1.1).

Lowest occurrence: Important molluscan events for defining the base of the Kapitean Stage are the lowest occurrences
Austrofusus coerulescens and Sectipecten wollastoni (?). At Upton Brook 8 km west of Blind River, ? showed that
these events almost coincide with the base of Chron 5 (=C3An, Fig. 4.5). The molluscan events do not occur in the
deeper water sequence at Blind River, but ? inferred that the base of C3An is about 600 m above the base of Upton
Formation, in a poorly exposed interval. This datum is c. 30 m above the lowest record of Bolivinita pliozea (P29/f29).
The data suggest that the appearance of B. pliozea is in the vicinity of the base of the Kapitean Stage. However, note
that the lowest occurrence of B. pliozea is poorly resolved in this rapidly deposited sequence, and that the position of
C3An.2n at Blind River is only inferred. Similarly, in the Mangapoike River sequence ?, fig. 5.2 was unable to resolve
the magnetostratigraphy in the vicinity of the lowest occurrence of B. pliozea (N106/f735 in ?, fig. 95. While it is
suggested that the entry of B. pliozea is a criterion for the base of the Kapitean Stage, more data about the association
between its appearance, the molluscan events, and magnetostratigraphic signatures are needed to confirm the result at

77
Blind River. Bolivinita pliozea is present in lower Kapitean sequences near East Cape (e.g., Z14/f19) but its relation
there with the molluscan events is unknown.

DRAFT
Highest occurrence: ? did not specify the highest occurrence of Bolivinita pliozea but ? placed it in the Castlecliffian
Stage. ? recorded rare specimens, which he regarded as reworked, in 6 dredge samples from Cook Strait. ? found
rare specimens in a shallow (< 10 m) inlet near Wellington. ? did not list it from a transect off southern Hawkes Bay.
? interpreted the Cook Strait specimens as part of the Recent and ? maintained this view.

In collections at GNS Science mounted from the Quaternary sequence at Wanganui (?), the highest assemblage in
which Bolivinita pliozea is common is R22/f7516 (type locality) from lower Shakespeare Cliff Silt. There are no
specimens in another collection (R22/f6368) from this unit. The only specimen noted in assemblages mounted
from higher formations is in R22/f7391 (Landguard Formation). However, this specimen is equivocal as it was found
loose in a slide that originally included two assemblages, one of which (S23/f6485) has been removed subsequently.
? correlated Shakespeare Cliff Silt, the highest reliable record of the species, with oxygen isotope stage 13 and
Landguard Formation with oxygen isotope stage 9. These data support the conclusion of ? and ? that the species is
not present in New Zealand Recent assemblages. At ODP Site 1119 off southeastern South Island, its highest record
(?) is at 133 meters composite depth and prior to the base of oxygen isotope stage 11 at 108 meters composite depth.
? dated its highest occurrence in ODP Site 1119 at 0.46 Ma.

Although some data are equivocal, it is likely that Bolivinita pliozea did not live in the New Zealand region beyond
the Pleistocene.

78
Chapter 11

DRAFT
Bolivinita medialis Scott n. sp.

Holotype: TF1676/1 (GNS Science type foraminiferal collection).

11.1 Bolivinita medialis Type Population T27/f551

Figure 11.1: (A) Bolivinita medialis Scott n. sp. Variation in the spiral outline of specimens from T27/f551. Included
are some specimens of B. compressa Finlay. (B) Specimens (standardized for size) are plotted on 3-dimensional
coordinates based on principal components analysis of 20 coefficients for 5 elliptical Fourier harmonics fitted to the
outlines; 98% of variation is shown.

Material

T27/f551 Bells Creek Mudstone in Karamu South Stream, Wairarapa, between T27/f431 and T27/f432 (?, fig. 5).
The horizon is in the upper part of an interval characterized by dextrally-coiled shells of Globorotalia miotumida

79
which ? identified as the Kaiti coiling zone, lower Tongaporutuan Stage. Eighteen topotypes were studied in spiral
orientation and c. 100 in axial and distal orientations.

DRAFT
Spiral Orientation

Chamber arrangement: In most shells there is little or no offset between chambers and outlines are often nearly
bilaterally symmetrical (Fig. 11.1 (A) #15). An outlier (Fig. 11.1 (A) #5) is weakly rhomboidal due to the amount
of offset between chambers. In a subcluster (e.g., Fig. 11.1 (A) #7, Fig. 11.1 (A) #16) the upper left of the outline
is raised. More significant is the contrast with the strongly rhomboidal outlines of the accompanying specimens of
Bolivinita compressa (Fig. 11.1 (A) #19, #21).

Distal wall: Boundaries with the axial walls are usually well-defined by keels. Rarely, one of the junctions with an
axial wall is rounded (Fig. 11.1 (A) #7) either because a keel is not present, or does not extend sufficiently high on the
wall of the chamber to be visible. The distal wall is moderately convex (Fig. 11.1 (A) #13) and usually rises above the
bordering keels. The profile of a strong rib is sometimes seen near the crest of the wall (Fig. 11.1 (A) #3).

Axial walls: Differences in the length and orientation of axial walls are usually minor. Note that the weakly rhom-
boidal outline of Fig. 11.1 (A) #15 occurs in an individual in which axial walls are differentiated in length and orien-
tation. Some profiles are linear throughout (Fig. 11.1 (A) #13 upper right). Others are weakly concave, particularly
near the junction with the opposed axial wall (Fig. 11.1 (A) #7 right). This variation is reflected in the shape of the
re-entrant formed by axial walls. In Fig. 11.1 (A) #18 left the re-entrant is almost regular and relatively deep. Shallow
concave re-entrants occur in Fig. 11.1 (A) #3 right, #17 right; in Fig. 11.1 (A) #7 right the concave re-entrant is deeper.

Keels: Typically, there is a keel at each axial - distal wall junction. Most have an oblique orientation, intersecting the
angle made by the walls. However, in Fig. 11.1 (A) #16 the upper right keel projects laterally, as in many Bolivinita
pliozea. Keels are usually wide but the extent to which they project above the walls is quite variable (cf. Fig. 11.1 (A)
#7, #17). Some attenuate towards the aperture (Fig. 11.1 (A) #18).

Aperture: The opening is often U-shaped, high, with only minor asymmetry (Fig. 11.1 (A) #13). The rim is broad but
may project little above the adjacent walls. There is a tooth in the upper part of the opening.

Axial Orientation T27/f551

Shell outline: The prolocular section is narrowly rounded in microspheric specimens and more broadly so in mega-
lospheric specimens (Fig. 11.2 #1, #2). The difference is affected by the width of keels on the earliest chambers.
Lateral sections are defined by the outlines of the distal walls. Keels at the distal - axial wall junctions are inset from
the periphery, but often only slightly (Fig. 11.2 #2). In many specimens the lateral sections diverge almost uniformly
throughout ontogeny but in some megalospheric forms the amount is very minor in late ontogeny and the sections
become almost parallel. Lobulation along the lateral sections is often weak, due to the dampening effect of ribs on
distal walls (Fig. 11.2 #3). The apex above the aperture is often rounded (Fig. 11.2 #2).

Chambers: Biserial arrangement, with up to c. 9 pairs. Chambers are wedge-shaped and inclined (Fig. 11.2 #3). They
are closely packed in nearly all specimens. Axial walls incline towards the coiling axis and are flat to very weakly
concave. Generally, sutures are slightly depressed (Fig. 11.2 #2), or flush. On some specimens a relict section of the
keel is preserved as a wide ridge at the base of late-formed chambers (Fig. 11.2 #1). There is a strong keel at each
axial - distal junction. Ribs are absent from the axial walls. Late-formed chambers have smooth walls and small, oval
pores that are fairly evenly distributed (Fig. 11.3 #6). Domes are formed around pores on earlier chambers and are
wider and higher on the earliest chambers (Fig. 11.3 #8).

80
tions.

DRAFT
Figure 11.2: Bolivinita medialis Scott n. sp. Guide to morphology of specimens from T27/f551 in standard orienta-

Figure 11.3: Bolivinita medialis Scott n. sp. T27/f551.

Wall topography.

Distal Orientation

Shell outline: The outline is defined by keels at the margins of the distal walls. The prolocular section is broadly
rounded in megalospheric specimens; lateral sections in such forms expand only slightly in ontogeny (Fig. 11.2 #5).
There is greater divergence near the proloculus in microspheric specimens (Fig. 11.2 #4). Lateral sections are weakly
curved to linear. Lobulation is almost entirely suppressed by the strong keels. The section across the top of the aperture
is often flat (Fig. 11.2 #5).

Chambers: The height of chambers expands through ontogeny much faster than their width. Much of the base of a
chamber is linear but it may curve downward at the margins (Fig. 11.2 #5). The lateral sections are convex, although
they are partly obscured by the keels. Chamber walls are moderately convex and the suture between adjacent chambers
is depressed. Ribs are often wide and robust. A median rib, sometimes very strong (Fig. 11.2 #4) is distinctive, but
specimens with 2 ribs (Fig. 11.2 #5) are quite common. Some specimens have up to 3 weak ribs. Rarely, ribs die out
by mid-ontogeny.

81
 DRAFT
Figure 11.4: Bolivinita medialis Scott n. sp. Variation in the spiral outline of specimens from T27/f495.

Figure 11.5: Bolivinita medialis Scott n. sp. Guide to morphology of specimens from T27/f495 in axial and distal
orientations.

11.2 Bolivinita medialis Comparative Population T27/f495

Material

T27/f495 samples Bells Creek Mudstone in Karamu South Stream c. 55 m below T27/f551 (?, fig. 5) This Tonga-
porutuan horizon is below the base of the Kaiti coiling zone. Twenty-four specimens were studied in spiral orientation
and c. 60 specimens in axial and distal orientations.

Spiral orientation: As with some topotypes, chambers are slightly offset (Fig. 11.4 #16). This, together with small dif-
ferences in lengths and orientations of axial walls, result in outlines that are not exactly bilaterally symmetrical. Some
outlines are very weakly rhomboidal (Fig. 11.4 #1). Many of the features seen in topotypes are present. Distal walls
are moderately convex, as in topotypes, and axial wall re-entrants are sometimes similar (Fig. 11.4 #6 right, Fig. 11.1

82
 DRAFT
Figure 11.6: Bolivinita medialis Scott n. sp. (A) Variation in the spiral outline of specimens from T27/f495 and
T27/f551. (B) The cluster analysis includes several specimens of B. compressa and uses 20 coefficients for 5 elliptical
Fourier harmonics (specimens standardized for size) fitted to outlines in spiral orientation.

#3 right). However, in a few specimens (Fig. 11.4 #5) there are combinations of characters e.g., compact cham-
bers, relatively strongly convex distal walls and laterally-projecting keels that are better developed than in topotypes.
Some of these specimens closely resemble Bolivinita pliozea. Cluster analysis of spiral outline data for T27/f495
and T27/f551 (Fig. 11.6) formed groups consisting predominately of specimens from one or other of the localities.
Contributing to the partial separation is the greater prevalence of compact outlines in T27/f495 (Fig. 11.6 (A) #A).
Some outlines in T27/f551 are relatively elongated (Fig. 11.6 (A) #1). The data indicate appreciable inter-population
variation in spiral shape.

Axial, distal orientations: In a few specimens the axial walls are narrow, well inset from the periphery (Fig. 11.5 #3),
and form channels resembling those of Bolivinita pliozea. Their keels project almost laterally. The resemblance is
heightened by 3-4 ribs on distal walls. More common are specimens with 1-2 strong ribs on distal walls (Fig. 11.5 #4,
#5).

11.3 Discrimination Bolivinita medialis : B. pliozea

Material

R22/f7516 is adjacent to the type locality of Bolivinita pliozea; P29/f29 is the lowest horizon in Blind River section in
which B. pliozea is identified. Both are compared with T27/f495 as this population includes specimens more closely
resembling B. pliozea than do topotypes of B. medialis.

83
 DRAFT
Figure 11.7: (A) Discrimination of typical Bolivinita pliozea (R22/f7516) from B. medialis (T27/f495) using charac-
ters seen in spiral orientation. (B) The cluster analysis uses 20 coefficients for 5 elliptical Fourier harmonics (speci-
mens standardized for size) fitted to outlines in spiral orientation.

R22/f7516

Spiral orientation: Variation in the offset between chambers is similar in both samples. Distal walls are commonly
more convex in Bolivinita pliozea than in the B. medialis sample, although convexity is closely similar in some
specimens (Fig. 11.7 (A) #18, #C). Axial walls are often longer in Bolivinita medialis from T27/f495 and specimens
with linear elements in axial wall outlines are common (Fig. 11.7 (A) #3). While there are counterparts in B. pliozea
(Fig. 11.7 (A) #A), weakly concave profiles are better developed (Fig. 11.7(A) #B, #C). This difference affects the
form of axial wall re-entrants. Specimens with narrow, deep, V- to U-shaped re-entrants are common in R22/f7516
(e.g., Fig. 10.1 #6) but U-shapes are barely developed in B. medialis from T27/f495. Both samples have specimens
with keels that project almost laterally. However, keels are more robust in some B. medialis (Fig. 11.7 (A) #3). The
latter morphotype, with robust, obliquely projecting keels, weakly convex distal walls, and wide re-entrants in the
axial walls, is not found in B. pliozea from R22/f7516. In contrast, both samples have a few specimens, usually small,
in which keels are low, or not visible in this orientation (Fig. 11.7 (A) #4, #A).

Most of the low-level (c. 0.05) groups determined by cluster analysis of spiral outline data (Fig. 11.7 (B)) include
specimens from both taxa; intersection of variation fields is suggested. Note the assignment of the specimen shown in
Fig. 11.7 (A) #18 to a group that includes quite typical morphotypes of B. pliozea (Fig. 11.7 (A) #B-#C). A rare, and
rather atypical specimen of B. pliozea (Fig. 11.7 (A) #A) is grouped in a cluster largely consisting of B. medialis.

Axial, distal orientations: Specimens of Bolivinita pliozea with deep, narrow re-entrants formed by the axial walls
(Fig. 10.2 #1) are quite common in R22/f7516. There are rare counterparts in B. medialis (Fig. 11.5 #3). Specimens
with one strong, median rib on distal walls occur in the B. medialis sample (Fig. 11.5 #5) but are absent from B.
pliozea in R22/f7516.

84
P29/f29

DRAFT
Spiral orientation: Compact outlines are more common than in Bolivinita medialis from T27/f495. Many other
characters in the samples are closely similar (Fig. 11.8 (A) #B). These include moderately convex distal walls, near
linear axial wall profiles, wide, V-shaped axial wall re-entrants, and a keel at each axial - distal wall junction. Although
keels project obliquely in both samples, the structure is sometimes wider and more robust in B. medialis than in B.
pliozea. Overall, in spiral view the differences the samples are few, subtle, and apparent only on some specimens.

Figure 11.8: (A) Discrimination of early specimens of Bolivinita pliozea (P29/f29) from B. medialis (T27/f495)
using spiral outline data. (B) The cluster analysis uses 20 coefficients for 5 elliptical Fourier harmonics (specimens
standardized for size) fitted to outlines in spiral orientation.

Cluster analysis of outline data (Fig. 11.8 (B)) identifies one, large group at the 0.06 level which is an admixture of
specimens from the two samples. The analysis supports the visual impression that the gross features of spiral outlines
in the two samples are closely similar.

Axial, distal orientations: Lateral sections of axial profiles are moderately convex in both samples. However, in
Bolivinita pliozea from P29/f29 the sutural depressions are fully bridged by distal wall ribs. This masking results
in lateral sections that are non-lobulate (Fig. 11.9 #f). Some B. medialis in T27/f495 are lobulate (Fig. 11.9 #a, #b),
although the amplitude is usually dampened by ribs. Specimens in which axial walls are inset from the periphery
and form a relatively narrow, channel-like depression (Fig. 11.9 #c) occasionally occur in B. medialis from T27/f495.
They match a morphotype widespread in B. pliozea populations (e.g., Fig. 10.7 (A) #f). Distally, both samples have
specimens with a strong median rib, but it is usually wider and more robust in T27/f495 (Fig. 11.9 #e). Rib numbers
vary between 1-4; 2 ribs are typical in P29/f29 and 1-2 in T27/f495.

11.4 Bolivinita medialis Overview

Bolivinita pliozea and B. medialis are closely similar morphospecies that share a common design. Although most
specimens from the type populations are readily separable, the number of individuals misidentified is likely to be much
higher when populations like P29/f29 and T27/f495 are considered. Why then is B. medialis recognized when much

85
 DRAFT
Figure 11.9: Discrimination of early specimens of Bolivinita pliozea (P29/f30) from B. medialis (T27/f495) using
characters seen in axial and distal orientations.

greater intra-specific variation has been allowed in taxa such as B. pohana? Bolivinita medialis is proposed because
of evidence suggesting that B. pliozea evolved from B. pliobliqua. Bolivinita pliobliqua immediately precedes B.
pliozea and much of its design suggests that it is a prototype of B. pliozea. Lineage splitting is not indicated by the
stratigraphic distributions of the taxa. Rather, B. pliozea possibly arose by phyletic transformation from B. pliobliqua.

In contrast with the stratigraphic juxtaposition of Bolivinita pliobliqua and B. pliozea near the Tongaporutuan -
Kapitean boundary, B. medialis occurs significantly earlier and is not known to have extended above the informal
lower Tongaporutuan zone. The taxa are separated by c. 3 Ma (Fig. 1.1). Data on stratigraphic distributions are equiv-
ocal for estimation of ancestry as it might be argued that B. medialis temporarily withdrew from the New Zealand
region. However, this does not account for the evidence of morphological transformations between B. pliobliqua and
B. pliozea.

No evidence about the origin of B. medialis has been found in New Zealand strata and it is viewed as a ?migrant species
that appeared, along with several other Bolivinita, in the New Zealand region near the base of the Tongaporutuan Stage.

11.5 Bolivinita medialis Biogeography

The species is principally distributed in the Eastern Basin between Hawke's Bay and north Canterbury (Fig. 11.10).
Presently, there no records known from northern sequences in this basin. A significant boundary in its biogeography
might be indicated by its absence from apparently suitable biofacies in the sequence at Kaiti Beach, Gisborne. There
are no records of Bolivinita medialis from Taranaki Basin as yet, but it occurs in Westland (Kawhaka-1 890 ft; identi-
fied as B. pliozea in ?) and Southland. Most occurrences are from middle bathyal or deeper assemblages, sometimes
in association with B. quadrilatera or B. pohana. Population sizes are usually modest.

In overview, Bolivinita medialis differs from B. pliozea in its more restricted geographic distribution, preference for
deeper bathyal environments, and local impersistence. Resemblances in the shell design of the taxa are not paralleled
in their gross biogeographies and population strategies.

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Figure 11.10: Distribution of Bolivinita me-
dialis on modern coordinates. Shaded areas
are generalizations of known distributions
and are conservative. They do not identify
possible changes in its distribution through
the Late Neogene.

11.6 Bolivinita medialis Stratigraphic Distribution

Lower Tongaporutuan Stage (Fig. 1.1).

Lowest occurrence: The Globorotalia miotumida Kaiti coiling zone (?) provides a useful reference datum near the
base of Tongaporutuan Stage that is unrelated to Bolivinita events. In southern Wairarapa (Karamu Nouth Stream)
Bolivinita medialis appears between the lowest occurrence of B. quadrilatera (T27/f460) and the base of Kaiti coiling
zone (T27/f464). In southern Hawke's Bay B. medialis occurs with B. quadrilatera and B. pohana in an assemblage
(V22/f8772) that may be below the Kaiti coiling zone. At Gore Bay (North Canterbury) B. medialis is present in an
assemblage (O33/f13) from the base of Kaiti coiling zone. The horizon is c. 7 m above the first record of Bolivinita
quadrilatera (O33/f15). The available data, some of which are not adequately constrained, indicate that B. medialis
appeared subsequent to the lowest record of B. quadrilatera but prior to the base of Kaiti coiling zone.

Highest occurrence: No records are known of Bolivinita medialis above the highest occurrence of Globoquadrina
dehiscens (datum defining the top of the informal lower unit of the Tongaporutuan Stage).

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Chapter 12

DRAFT
Bolivinita quadrilatera (Schwager)

1866 Textilaria quadrilatera Schwager. Fossile Foraminiferen von Kar Nikobar. Reise der Osterreichischen Fregatte
Novara um die Erde in den Jahren 1857, 1858, 1859. Palaontologische Mitteilungen, 2: p. 253, taf. vii, fig. 103.

12.1 Description of neotype

"Test elongate, about two and half times as long as broad, slightly twisted at the initial-end, compressed, quadrate in
apertural view, tapering to rounded initial-end; the broader faces moderately concave, angles carinate, chambers seven
to eight on each side on broader faces; sutures broad, prominent, slightly raised; wall calcareous, finely and densely
perforate; aperture suboval, at the inner margin of the last-formed chamber." (?, p. 43)

Variation: "Some individuals are more narrowly pointed initially than specimen illustrated here. Variation is also
observed in the curvature of the broader face of the test. It varies from wholly concave to initial portion concave and
later part nearly flat. A few specimen show a slight inflation in the adult chambers." (?, p. 43) They refer to specimens
from neotype locality Mf/42, Passa Bridge section, Car Nicobar, Andaman Islands.

Figure 12.1: Bolivinita quadrilatera holotype (?, taf. vii, fig. 103) and neotype (?, pl. 6, fig. 22-23).

Note on systematics: Bolivinita is the type genus of the Bolivinitidae and B. quadrilatera is the type species of the
genus. In a revision of the systematics of the Bolivinitidae ? examined topotypes of B. quadrilatera deposited in the
collections of the Natural History Museum, London. He noted that their toothplate morphology differed from Recent
specimens and suggested that the latter should be referred to B. pliozea Finlay. He did not figure any Recent specimens
he would so identify.

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 DRAFT
Figure 12.2: Bolivinita quadrilatera (Schwager). Variation in the spiral outline of specimens from T27/f460. Speci-
mens (standardized for size) are plotted on 3-dimensional coordinates based on principal components analysis of 20
coefficients for 5 elliptical Fourier harmonics fitted to the outlines; 90% of variation is shown.

12.2 Bolivinita quadrilatera Reference Population T27/f460

Material

T27/f460 is from North Karamu Stream, Wairarapa, about 10 m above the base of Bells Creek Mudstone. Twenty-four
specimens were studied in spiral orientation and 64 in axial and distal orientations. It is located in the informal lower
unit of the Tongaporutuan Stage.

Spiral Orientation

Chamber arrangement: Commonly, the biserial shell is nearly symmetrical (Fig. 12.2 (A) #20). Weakly asymmetrical
outlines (Fig. 12.2 (A) #18) occur when chambers are slightly offset and axial walls differ in length. Flexure of the
axis of coiling during ontogeny may also cause departure from simple bilateral symmetry (e.g., Fig. 12.2 (A) #4).

89
Distal wall: Margins are well-defined by keels at the junction with axial walls. The profile is often almost linear and
lies below (Fig. 12.2 (A) #12), or at (Fig. 12.2 (A) #23), the level of the bordering keels. However, if sufficiently

DRAFT
convex (Fig. 12.2 (A) #18), it rises above the latter. Occasionally, the profile varies markedly between successive
chambers; note the concave profile of the penultimate (lower) chamber in Fig. 12.2 (A) #15.

Axial walls: Their length and orientation are often similar, leading to bilaterally symmetrical outlines (Fig. 12.2
(A) #20). However, departures from exact matches are common and produce outlines that are weakly asymmetrical
(Fig. 12.2 (A) #18). Wall surfaces are often weakly concave, but there are almost flat sections. The walls of opposed
chambers usually abut to create a wide, shallow, concave re-entrant (Fig. 12.2 (A) #1). In some it is nearly symmetrical
(e.g., Fig. 12.2 (A) #15).

Keels: These are robust and occur at each distal - axial wall junction. Often they project obliquely above the adjacent
walls but there is significant variation: compare Fig. 12.2 (A) #9, #12. Commonly, keels are equal in size and are
linked across the rim of the aperture. But in Fig. 12.2 (A) #9 one of the keels attenuates towards the aperture.

Aperture: Often the base is almost flat, reflecting planar surfaces in the distal wall of the preceding chamber on which
it is built. The opening is arched, and commonly almost symmetrical about its median longitudinal axis (Fig. 12.2
(A) #23). The sides are gently curved and form a weakly-defined, broad apex. However, some openings are weakly
asymmetrical (Fig. 12.2 (A) #15), while the sides are almost parallel in Fig. 12.2 (A) #16. A robust, curved tooth
projects from the apical region of the opening.

Axial Orientation

Figure 12.3: Bolivinita quadrilatera (Schwager) Guide to morphology of specimens from T27/f460 in axial and distal
orientations.

Shell outline: Usually this is defined by the keels at junctions of distal and axial walls. Distal walls are out of
view (Fig. 12.3 #1), except in specimens in which the coiling axis is flexed during ontogeny. The lateral margins
diverge out from the prolocular section. In some megalospheric specimens maximum width is approached by mid-
ontogeny (Fig. 12.3 #1) and there is only slight, or no subsequent expansion. In other megalospheric specimens, and
in microspheric specimens (Fig. 12.3 #2), shell width expands through ontogeny, although the rate may decline. The
apex above the aperture varies from rounded (Fig. 12.3 #3) to a narrow peak. Due to the presence of robust keels at
the lateral margins, lobulation is almost entirely suppressed.

Chambers: Up to c. 10 pairs of biserially-arranged chambers. They are wedge-shaped and incline at a steep angle
downward from the coiling axis (Fig. 12.3 #3). Overall, chamber surfaces are weakly concave, but may include planar
regions. In early ontogeny, junctions between chambers are flush but a remnant section of the keel is preserved as a
strong ridge at the base of late-formed chambers (Fig. 12.3 #1). The remnant section represents a part of the keel that
is overriden when a new chamber is formed. In late-formed chambers the axial wall appears inset within the bounding
ridges of the keel and its remnants. The keel at the distal margin of chambers is wide and very prominent. Sometimes

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 DRAFT
Figure 12.4: Bolivinita quadrilatera T27/f460. Wall to-
pography.

it tapers to a thin flange. There are no ribs on the axial walls. Pore distributions on axial and distal walls are similar.

Shell outline: The lateral outline is usually defined by the keels at the margins of chambers. Axial walls may be
partially visible in specimens in which the coiling axis is flexed (Fig. 12.3 #5). Keels are broad, and often taper
to a flange. There is good differentiation between the narrow, almost pointed, prolocular region of microspheric
specimens and the flattish curve of megalospheric specimens. Although the width of specimens is much less than
in axial orientation, the same differences between generations are seen. Lobulation of the lateral margins is either
indistinct or absent. The outline above the aperture is often flat (Fig. 12.3 #4).

Chambers: Lower and upper margins are usually linear, although a few are weakly concave. Lateral margins are
weakly convex (Fig. 12.3 #4). However, the shape of chambers is approximately rectangular. During ontogeny,
chamber height expands much more rapidly than does width. Sections across chambers are very weakly convex to
linear; longitudinally, convexity is greater and is reflected in depressed sutures, especially in late-formed chambers.
Pores are small, often oval, and irregularly distributed (sometimes over 5 diameters apart, (Fig. 12.4 #7). There is
minor secondary calcification around pores in early chambers (Fig. 12.4 #8).

12.3 Discrimination Bolivinita quadrilatera : B. urenuia

Material

Bolivinita urenuia from J32/f54, Callaghan's Creek, Westland; similar populations were identified as B. quadrilatera
by ?.

Spiral Orientation: A principal feature separating specimens of Bolivinita quadrilatera (T27/f460) from B. urenuia
(J32/f54) is the smaller, sometimes negligible, offset between chambers. The spiral outline is often almost bilaterally
symmetrical (Fig. 12.5 (A) #2, #5), whereas outlines in B. urenuia are usually rhomboidal (Fig. 12.5 (A) #B). A rare
specimen of the latter in which the offset is slight is shown in Fig. 12.5 (A) #C. In the dendrogram this specimen
lies adjacent to the B. quadrilatera cluster (Fig. 12.5 (C)). While variable, distal walls are usually less convex in B.
quadrilatera. The axial wall re-entrant is often shallower, and forms a continuous curve (Fig. 12.5 (A) #5), rather than
an open V, as in B. urenuia (Fig. 12.5 (A) #B). In spiral orientation both keels link with the aperture in B. quadrilatera
(Fig. 12.5 (A) #2), whereas in B. urenuia one keel often dies away on the upper surface of the last chamber (Fig. 12.5
(A) #B).

Axial, distal orientations: In axial orientation the shell outline of Bolivinita urenuia (Fig. 12.5 (B) #a) resembles
Bolivinita quadrilatera. Many characters are similar: shell outlines; nearly planar axial walls with barely depressed
sutures; relict keels preserved on later chambers; weakly convex distal walls with depressed sutures; strong keels and
absence of ribs (Fig. 12.5 (B) #c, #d). Depending on the extent to which chambers are offset, part of the distal wall is
visible in Bolivinita urenuia in oblique axial orientation e.g., Fig. 7.2 #4. Typically, there is no comparable view in B.
quadrilatera.

Remarks: Specimens in J32/f54 often show lesser offset between chambers than in topotypes of Bolivinita urenuia.
While groupings in the cluster analysis (Fig. 12.5 (C)) indicate that B. urenuia from J32/f54 occupies a morphospace
adjacent to B. quadrilatera, there are no stratigraphic data that suggest that B. urenuia evolved from B. quadrilatera.

91
 DRAFT
Figure 12.5: Bolivinita quadrilatera (Schwager) and B. urenuia Scott n. sp. from T27/f460 and J32/f54. (A)
Discrimination of specimens in spiral orientation. (B) Discrimination of specimens in axial orientation. (C) Variation
in the spiral outline of specimens from T27/f460 and J32/f54. The cluster analysis uses 20 coefficients for 5 elliptical
Fourier harmonics (standardized for size) fitted to outlines in spiral orientation. Refer to (A) for micrographs of
specimens #2, #5, #17, #A-#C.

12.4 Bolivinita quadrilatera Biogeography

This was reviewed by ? who noted its preference for bathyal environments and its patchy distribution. This pattern
is confirmed by more recent work (Fig. 12.6). ? found it only in 1 (DSDP Site 594) of 3 bathyal oceanic sequences
adjacent to New Zealand and there only in post-Miocene strata). Like Bolivinita compressa and B. pohana, its Tonga-
porutuan record (persistence, population size) is best in the Eastern Basin. ? found that B. quadrilatera was common
to abundant on the steep wall of a canyon system south of Kaikoura. Perhaps significantly, there were large populations
in the same region at the base of the late Miocene, over 10 m.y. previously.

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 DRAFT
Figure 12.6: Distribution of Bolivinita quadrilat-
era on modern coordinates. Shaded areas are gen-
eralizations of known distributions and are conser-
vative. They do not identify possible changes in its
distribution through the Late Neogene.

12.5 Bolivinita quadrilatera Stratigraphic Distribution

Base of Tongaporutuan Stage - Recent (Fig. 1.1).

Lowest occurrence: ? considered that the lowest occurrence of Bolivinita quadrilatera was the primary foraminiferal
event for recognizing the base of the Tongaporutuan Stage. Hornibrook (in ?) noted that Loxostomum truncatum,
considered by ? as the index species for Waiauan (i.e., restricted to that stage), occurred with B. quadrilatera in
some southern Hawke’s Bay assemblages. He rationalized these observations by selecting the lowest record of B.
quadrilatera as the boundary event, in preference to the highest record of L. truncatum. Scott (in ?) did not find B.
quadrilatera in the Tongaporutuan stratotype (north Taranaki coastal section) and considered that ? probably intended
the lowest occurrence of B. pohana to mark the base of the stage.

In Taranaki Basin a single specimen of Bolivinita quadrilatera was found in Kahawai-1 (core at 2760 m), c. 200 m
below strata with dextrally-coiled Globorotalia miotumida which ? identified as the basal Tongaporutuan Kaiti coiling
zone (?). The species is rarely present in Taranaki Basin.

In the Eastern Basin Bolivinita compressa, B. medialis, B. pohana and B. quadrilatera all occur below the base of the
Kaiti coiling zone, a datum near the base of the Tongaporutuan Stage. In the Kaiti Beach sequence (?) B. compressa
and B. pohana occur below this datum but B. quadrilatera has not yet been identified. Some of the southern Hawke’s
Bay assemblages with B. quadrilatera and Loxostomum truncatum discussed by Hornibrook (in ?) have sinistrally-
coiled specimens of Globorotalia miotumida and most likely lie below the base of the Kaiti coiling zone. No consistent
pattern in their stratigraphic distribution below this datum is yet apparent. V22/f8772 includes B. medialis and B.
pohana; V22/f8850 has B. pohana; V22/f8842 has neither, as does V22/f8660.

In North Karamu Stream (Wairarapa), the lowest record of Bolivinita quadrilatera (T27/f460) includes rare specimens
tentatively identified as B. compressa. The horizon is c. 90 m below the base of the Kaiti coiling zone (M.P. Crundwell
pers. comm. 22 August 1996). Bolivinita medialis appears closer to this datum. The pattern at Gore Bay (northern
Canterbury Basin) is similar. Bolivinita quadrilatera is common at its lowest record (O33/f15) and is accompanied
by rare B. medialis. This horizon is below the base of the Kaiti coiling zone (first identified in O33/f13, 7 m higher).
In that assemblage B. quadrilatera is ?entirely replaced by B. medialis. Further south at Motunau, B. quadrilatera is
again common at its lowest record (N34/f7563) but it is accompanied by B. pohana.

93
While many of these data are poorly constrained because of inadequate sample resolution, they indicate that in eastern
sequences south of Hawke Bay, Bolivinita quadrilatera is often common at its lowest occurrence, below the base

DRAFT
of Kaiti coiling zone. The data suggest, but do not clearly establish, that it was the first of the group to enter the
New Zealand record. Several other Bolivinita entered the region within a short interval; their earliest distributions are
poorly known, and were possibly localized.

Highest occurrence: Records from Recent sediments include ?, ?, ? and ?.

Remarks: ? noted the episodic occurrence of Bolivinita quadrilatera in the Neogene record. The presence of suitable
environments may have been significant. There is a fair record of the species in the Eastern Basin in the Tongaporutuan
Stage, particularly in the informal lower zone. It is uncertain whether it was present in the latest Miocene (Kapitean
Stage). One specimen is present in Z14/f9503 from a locality with Kapitean Mollusca, but might be reworked (Globo-
quadrina dehiscens tentatively identified, also). Following this possible break in occupancy come the Early Pliocene
(Opoitian Stage) records of B. quadrilatera granttaylori.

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Chapter 13

DRAFT
Bolivinita quadrilatera granttaylori Vella

1957 Bolivinita grant-taylori Vella. New Zealand Geological Survey Paleontological Bulletin 28: p. 33, pl. 8 fig.
157-159.

Original Description

"Shell large, thick, translucent, elongate, tapering gradually to a blunt rounded initial end; about half the specimens
slightly twisted, the remainder not twisted. Cross section quadrate, the broader faces deeply concave, the narrower
faces lightly convex and sculptured at the adult end with weak, irregular longitudinal ribs. Angles prominent, acutely
carinate except at the initial end where they are limbate but rounded. Proloculum large, followed by eight chambers
on each side, slightly inflated on the broader faces, more so on the narrower faces. Sutures heavily limbate, slightly
raised, gently curved, and moderately oblique. Aperture large with a distinct, projecting tooth plate. Length 1.12 mm,
width 0.33 mm, thickness 0.22 mm." (?)

Figure 13.1: Bolivinita quadrilatera granttaylori Vella. Variation in the spiral outline of specimens from X19/f9509.

13.1 Bolivinita quadrilatera granttaylori Type Population

Material

X19/f9509 is from the Nukumaruan sequence along SH2 at Te Uhi Hill, just east of Wairoa (?, fig. 3) . It is the
type locality of Bolivinita granttaylori Vella. Twenty-three specimens were studied in spiral orientation and c. 20

95
specimens in axial and distal orientations. They are compared with the reference sample for Bolivinta quadrilatera
(T27/f460).

DRAFT
Spiral orientation: Most profiles are nearly bilaterally symmetrical (Fig. 13.1 #13). None has a profile as asym-
metrical as that in Fig. 12.2 #18 from T27/f460. Concave re-entrants formed by axial walls are less common than
in T27/f460 and often shallower (Fig. 13.1 #2). Specimens in which the re-entrants are absent (Fig. 13.1 #6) are
more common than in T27/f460. Distal walls have linear (Fig. 13.1 #19) to convex (Fig. 13.1 #4) profiles. While
the amount of convex curvature is little greater than in T27/f460, the effect is exaggerated by the lesser projection
of keels beyond the walls (Fig. 13.1 #3). Generally, keels are less conspicuous than in T27/f460 (Fig. 13.2 (A) #1).
? shows exceptions. In the original description, ? noted that about half of the specimens were "twisted" (pairs of
chambers progressively rotated about the coiling axis); few specimens show this feature in spiral orientation, and then
only slightly (Fig. 13.1 #11).

The dendrogram (Fig. 13.2 (C)) for spiral outline data standardized for size shows clusters that are largely sample-
specific. Specimens from T27/f460 that are weakly rhomboidal (Fig. 13.2 (A) #12) are relatively distant from the
larger clusters. Most specimens of B. quadrilatera granttaylori lie in one cluster which includes a specimen from
T27/f460 (Fig. 13.2 (A) #1) with weak axial wall re-entrants.

Axial, distal orientations: Megalospheric shells are dominant in X19/f9509 whereas T27/f460 has many microspheric
individuals. Megalospheric proloculi in X19/f9509 are commonly larger than in T27/f460. Commonly, gross shell
length for a given number of chambers is greater in X19/f9509 (Fig. 13.2 (B) #c, #d) than in T27/f460; the increase
in size results from greater inclination of chambers (Fig. 13.2 (B) #d), and possibly higher rates of expansion in
chamber dimensions during ontogeny. The lesser divergence of lateral margins (axial view) during growth probably
is correlated with the larger size of the proloculus. Sutures are conspicuous in both populations. They are wider in
X19/f9509 but relict keels are more prominent in T25/f460 (Fig. 13.2 (B) #b).

Whereas there are no ribs on distal walls of specimens from T27/f460, there are 1-4 very weak ribs on some from
X19/f9509. They are best developed on specimens with convex walls (Fig. 13.1 #3). Some in Fig. 13.2 (B) #f are
discontinuous or restricted to a single chamber.

Remarks: ?, fig. 43 studied the symmetry of the spiral outline of specimens from X19/f9509 (=N116/f509) and
concluded that it fell within the variation of other collections of Bolivinita quadrilatera. Characters such as size and
rib development were discounted as significant in taxonomy and B. granttaylori was regarded as a synonym of B.
quadrilatera.

This study confirms the basic architectural resemblance of Bolivinita quadrilatera granttaylori to other populations of
the species in New Zealand. However, it shows that there are differences in gross size, wall shapes, keels, sutures and
size of the proloculus that ? either did not identify or discounted (e.g., the connection between gross length of shells
and geometry of the chamber sequence). That B. quadrilatera granttaylori is restricted to the Pliocene - Pleistocene is
also significant. In combination, the morphological and distribution data suggest that B. granttaylori Vella should be
recognized as a subspecies of B. quadrilatera. The distinction is useful in biostratigraphy. ? discussed some features
of a Pliocene population from the vicinity of the type locality of B. quadrilatera.

The origin of Bolivinita quadrilatera granttaylori is obscure. Bolivinita quadrilatera seems to have been largely
absent from New Zealand during the latest Miocene Kapitean Stage. There was at least one other group of variant
populations (Bolivinita angelina Church) in the Pacific during the Pliocene. The possibility that B. quadrilatera
granttaylori represents another invasion of the New Zealand region needs to be investigated.

13.2 Bolivinita quadrilatera granttaylori Biogeography

Bolivinita quadrilatera granttaylori occupied foreset units of the Giant Foresets Formation in Taranaki Basin (Fig. 12.6),
particularly in the northern sector (e.g., Kahawai-1, ?). Yet it has not been recorded from equivalent strata in several
sequences west of Taranaki Peninsula (e.g., Taimana-1, ?). Localized distributions along the same foreset front are

96
suggested.

DRAFT
13.3 Bolivinita quadrilatera granttaylori Stratigraphic Distribution

Bolivinita quadrilatera granttaylori: Opoitian Stage to Nukumaruan Stage (Fig. 1.1).

Lowest occurrence: The earliest records of Bolivinita quadrilatera granttaylori are from Opoitian strata in southern
Marlborough (e.g., P30/f7594).

Highest occurrence: Typical populations of Bolivinita quadrilatera granttaylori have not been found above Nuku-
maruan Stage but further study is needed.

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 DRAFT
Figure 13.2: (A) Bolivinita quadrilatera (Schwager) and B. quadrilatera granttaylori Vella. Variation in specimens
in spiral orientation from T27/f460 and X19/f9509. (B) Variation in specimens in axial and distal orientations. (C)
Variation in specimens in spiral orientation. The cluster analysis uses 20 coefficients for 5 elliptical Fourier harmonics
(specimens standardized for size) fitted to outlines in spiral orientation. Refer to (A) for micrographs of specimens
#1-#2, #12, #A-#B.

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Part III

Additional Studies

Original descriptions of Bolivinita elegantissima Boomgaart, B. subangularis

(Brady) and B. sp. aff. pohana striate (?) are given and specimens illustrated.

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Chapter 14

DRAFT
Bolivinita elegantissima Boomgaart

Description (?, p. 95, pl. XI, fig. 3a-c, D)

"Test small, slender, tapering, in some specimens slightly twisted; initial end bluntly pointed; subquadrate in cross-
section; chambers biserial throughout, five to six pairs, slightly inflated; sutures straight, more or less limbate, some-
what depressed; wall smooth, but with costae along the peripheral edges and also on the lateral faces; aperture subter-
minal, not in median line, large, nearly round."

"Our form resembles Bolivina subangularis Brady (1884, pl. 53, fig. 32-33), but is more slender than Brady's spec-
imens, whereas also the sides are not concave but rather convex due to the inflatedness of the chambers. This last
characteristic places this species between Bolivina and Bolivinita".

Addendum (G.H. Scott):

Stratigraphy: Waiauan Stage (Late Middle Miocene) to Opoitian Stage (Early Pliocene, ?). Bathyal environments,
impersistent.

Remarks: Chamber architecture differs significantly from species considered in the systematic study. The chamber
envelope (spiral orientation) is a uniform curve; distal and axial wall elements are not defined and keels are absent. Of
interest is the report in ? that Bolivinita quadrilatera evolved from this species in the Middle Miocene Globorotalia
fohsi robusta Zone. This implies cladogensis that involved major changes in chamber design.

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 DRAFT
Figure 14.1: Bolivinita elegantissima Boomgaart. T27/f122 Ruakiwi Stream, true right bank 111 m upstream from
junction with Mangariki Stream, Wainuioru Valley, east Wairarapa. Bells Creek Mudstone. Imagery by W. St George.

101
Chapter 15

DRAFT
Bolivinita subangularis (Brady)

Description (?, p. 59)

"Test oblong, tapering, stoutly built, more or less angular, somewhat concave or excavated on both sides; inferior
extremity obtusely pointed. The angular contour is determined by the prominence of superficial costae, the principal
of which, six in number, are placed, one down each lateral margin and two down each face of the test. Aperture
comma-shaped. Length 1/55 inch (0.45 mm)."

Addendum (G.H. Scott)

Systematics: ? did not illustrate the species or designate its type locality. ?, pl. LIII fig. 32-33 illustrated specimens
from Challenger Station 209, 95-100 fathoms, Philippine Islands.

Previous references of this species to Bolivinita are summarized by ? who considered that its generic identity is
questionable.

Several features of a Middle Miocene specimen from New Zealand (Fig. 15.1) compare closely with ?, pl. LIII fig.
32-33.
• Both are bilaterally symmetrical (Fig. 15.1 #b, #B) with inflated distal walls which extend well beyond their
junctions with axial walls.
• In the spiral outline, the distal chamber wall has two linear segments that meet centrally at a keel.
• There are strong keels at the junctions of axial and distal walls, and centrally on the distal walls.
• Axial walls incline inwards to form well-defined re-entrants in the spiral outline.
• Particularly, the lower axial and distal walls of chambers are inclined relative to the coiling axis so that the base of
the overlying chamber overhangs the top of its predecessor. This offset creates prominent projecting sutures (Fig. 15.1
#A, #C, #32).
• Chambers expand rapidly (Fig. 15.1 #32, #A, #33) from a small proloculus.

There are differences.

• The rate of chamber expansion appears to be greater in the Miocene specimen (Fig. 15.1 #A) which generates
greater flare in the axial outline.
• ? refers to 6 costae (keels in this study). While there are corresponding structures on the Miocene specimen, (Brady's
(1884) illustrations also show a rib located centrally on each facet of the distal wall on later chambers. These are not
present on the Miocene specimen.

Referral of the New Zealand Middle Miocene material to Bolivinita subangularis is tentative. Comparisons are based
on Brady's illustrations. A Recent Indo-Pacific population has not been examined and specimens are very rare in
New Zealand Middle Miocene assemblages. There is little knowledge of population variation and no information on
toothplate morphology. Light micrographs of Bolivinita subangularis from the Late Miocene in Papua New Guinea
(?, fig. 8-11) suggest that his material resembles B. pliozea. Late Pliocene-Pleistocene specimens from Timor (?, pl. 6
fig. 7-8) are closer to those illustrated here, although the absence of spiral and distal views constrains the assessment.

Stratigraphy: The lowest record found in collections at GNS Science is in D45/f8625, upper part of the Altonian
stratotype (latest Early Miocene), Clifden, Southland. The illustrated specimen (Fig. 15.1 #A-D) is from the Clifdenian
stratotype (Middle Miocene) in the same section. This assemblage includes Praeorbulina circularis. The highest
record known (D45/f8466) is from the Waiauan stratotype (Middle Miocene), also in the Clifden section. The Clifden
records are from shelfal assemblages, as are several other occurrences. ? cited B. subangularis as a shelf facies
indicator.

102
 DRAFT
Figure 15.1: Bolivinita subangularis (Brady). 32-33: Challenger Station 209 Philippine Islands, 95-100 fathoms.
Images scanned from Jones’ (1994) reproduction of Brady (1884, pl. LIII). #A-#D: D45/f8646 Clifdenian Stage
(Middle Miocene), Clifden, Southland.

103
Chapter 16

DRAFT
Bolivinita sp. aff. pohana striate

Description (?, p.41)

"In the Taranaki Coastal Section the first Bolivinita to appear is a small slightly oblique form with longitudinal ribs
on both the peripheral walls and the sides of the test. These small forms appear to be intermediate between B.
elegantissima and B. pohana. They range from the base of Unit 1 of the Mohakatino Formation to Unit 6 of the
Urenui Formation, but occur in few samples above the top of Unit 7 of the Tongaporutu Formation. Rare specimens
of this form occur with B. pohana in two samples from near the top of the Tongaporutuan massive mudstone in the
Palliser Section, and with the lowest B. pohana in the Tamumu Section."

"This form is probably a new species but more specimens are needed to determine its relationship to other species,
especially B. pohana."

104
 DRAFT
Figure 16.1: Bolivinita sp. aff. pohana striate. Q18/f18 Tongaporutu River estuary, north bank. Tawariki Formation
(?, fig. 23 section 6D). Specimens selected by G.H. Scott. Imagery by W. St George.

105
 DRAFT
Appendices (M.P. Crundwell)

Bolivinita pseudocompressa, Bolivinita striata and Bolivinita obconica are

proposed from ?.

106
Appendix A

DRAFT
Bolivinita pseudocompressa Crundwell n. sp.

Description: Test medium sized, broad, moderately compressed, about 2 to 2.5 times longer than broad, rhomboidal
in cross-section, expanding rapidly initially, then parallel sided. Chambers arranged biserially, although opposed at
180o intervals. Axial walls strongly differentiated in length to form broad, shallow, convex axial surfaces. Distal
walls generally smooth, slightly convex. Sutures inclined backward at 45o to the axial plane, generally flush except
for slightly depressed short-axial wall sutures. Wide flange-like keels developed at junctions of long-axial and distal
walls, often broken to produce a serrated edge. Short-axial/distal wall keels are generally incipient or may be absent
in some specimens. Aperture interiomarginal, loop-shaped, with inner facing tooth plate projecting from junction of
thickened keels bordering the apertural distal wall.

Variability: Microspheric forms are more strongly tapered and often have a slight axial twist. Fine, incipient, longitu-
dinal distal wall costae are present on the first formed chambers of some specimens, although they are generally few
in number and cover less than a quarter of the test.

Types: Holotype (VF 1160) and 64 paratypes (VF 1161) lodged at Victoria University of Wellington, 16 paratypes
(TF 1610/2-17) lodged at GNS Science, Lower Hutt.

Dimensions of holotype: Length 0.69 mm, breadth 0.37 mm (axial orientation).

Type locality: T27/f139, Westmere Road, 1 km north of intersection with Tupurupuru/Te Wharau Road, Wainuioru
Valley, east Wairarapa (grid reference T27/4112 0858).

Type level: Mangaopari Mudstone 465 m above Makara Greensand, Globorotalia inflata Zone, late Opoitian to Waip-
ipian (Middle Pliocene).

Remarks: This species is related to B. compressa, which is distinguished by its more squat form and much weaker
development or total absence of short-axial/distal wall keels. The new name B. pseudocompressa is used for popula-
tions of compressed Bolivinita where more than 50% of the specimens have short-axial/distal wall keels extending for
at least half of the length of the test.

Distribution: The species occurs frequently in massive fine grained sediments of Mangaopari Mudstone, cropping out
along the western side of the Wainuioru Valley.

Observed stratigraphic range: Opoitian to Waipipian (Early to Late Pliocene).

Observed ecological range: middle to outer shelf.

107
 DRAFT
Figure A.1: Bolivinita pseudocompressa Crundwell n. sp. #A-#K T27/f139 (type locality). Imagery by W. St George.

108
Appendix B

DRAFT
Bolivinita striata Crundwell n. sp.

Description: Test small, subrounded, rhomboidal in spiral profile, slender, about three times longer than broad, ta-
pering gradually to a sharply rounded initial end. Chambers arranged biserially, although opposed at 180o intervals.
Axial surfaces gently concave, the walls differentiated slightly in length, with depressed acutely angled limbate su-
tures, separating chambers that become increasingly inflated towards the adult end. Distal walls convex, divided by
depressed transverse sutures. Walls finely perforate, with numerous, slightly sinuous longitudinal costae, which be-
come obsolescent on later formed chambers, covering the lower half of axial surfaces and all except the ultimate and
penultimate chambers of the distal surfaces. Keels are weakly developed in microspheric forms being strongest at
the junctions of distal and axial walls. Although they are absent from megalospheric forms, costae along junctions of
axial and distal walls are slightly less prominent. Aperture interiomarginal, loop-shaped, with inner facing tooth plate
projecting from junction of keels bordering the apertural distal wall.

Variability: Microspheric forms have less well developed costae and tend to be more rhomboidal in spiral profile, and
often have a slight axial twist. Costae tend to become weaker and cover less of the test of specimens from higher
stratigraphic horizons.

Types: Holotype (VF1162) and 32 paratypes (VF1163) lodged at Victoria University of Wellington, 14 paratypes
(TF1611/2-15) lodged at GNS Science, Lower Hutt.

Dimensions of holotype: Length 0.47 mm, breadth 0.19 mm (axial orientation).

Type locality: T26/f133, Airstrip Creek, 170 m downstream from culvert on farm track, Wainuioru Valley, east
Wairarapa, (grid reference T26/43871124).

Remarks: This species has in the past been mistakenly identified as B. elegantissima, which is distinguished by its
relatively cylindrical cross-section, more inflated chambers, and fewer and stronger longitudinal costae. ? recognised
a form of Bolivinita intermediate between B. elegantissima and B. pohana, which he referred to as Bolivinita sp. aff.
pohana striate, but as it was never figured and reference material is unavailable it is uncertain how it relates to the
new species. The new species contrasts with its long-ranging, morphologically stable ancestor B. elegantissima, by
changing rapidly in form, becoming more elongate and rhomboidal with well developed flanges at the junctions of
long-axial and distal walls. Such forms are referred to as B. obconica.

Distribution: B. striata is a common species occurring mostly in the northern part of the Wainuioru Valley, in massive
fine grained lower Mangaopari Mudstone (late Kapitean to middle Opoitian).

Observed stratigraphic range: Late Kapitean to middle Opoitian (early Pliocene).

Observed ecological range: Outer shelf to mid bathyal.

109
 DRAFT
Figure B.1: Bolivinita striata Crundwell n. sp. #A-#I T27/f139 Westmere Road 1 km north of intersection with Te
Wharau Road, Wainuioru Valley, east Wairarapa. Mangaopari Mudstone. Imagery by W. St George.

110
Appendix C

DRAFT
Bolivinita obconica Crundwell n. sp.

Description: Test small, rhomboidal in spiral profile, elongate, gradually tapered to a sharply rounded initial end.
About 2.5 to 3 times longer than broad, often with a slight axial twist. Chambers arranged biserially, although opposed
at 180o intervals. Axial walls differentiated slightly in length, meeting at an obtuse angle to form shallow re-entrant
axial surfaces. Distal walls convex, subdivided by depressed, slightly oblique transverse sutures, inclined towards
short axial walls. Chambers slightly inflated posteriorly and separated by limbate sutures. Keels are developed at the
four junctions of axial and distal walls, being strongest adjacent to long-axial walls. Walls finely perforate, with weak
longitudinal costae becoming obsolescent on the distal walls of later formed chambers, and often restricted to the early
formed chambers of axial walls. Aperture interiomarginal, loop-shaped, bordered by a thickened rim of calcite formed
by the extension of the apertural distal wall keels that meet at an inner projecting toothplate.

Variability: Megalospheric forms are slightly less rhomboidal in spiral profile and some specimens have an incipient
keel adjacent to the apertural-distal wall, which often only extends to the base of the penultimate chamber. Costae tend
to become weaker and may be absent in higher stratigraphic horizons. Such specimens lacking costae may warrant
separation as a new taxon.

Types: Holotype (VF1158 and 64 paratypes (VF1159) lodged at Victoria University of Wellington, 16 paratypes
(TF1609/2-17) lodged at Institute of Geological & Nuclear Sciences, Lower Hutt.

Dimensions of holotype: Length 0.58 mm, breadth 0.23 mm (axial orientation).

Type locality: T26/f131, Maungatepopo Stream, 370 m downstream from Westmere Road, Wainuioru Valley, east
Wairarapa, (grid reference T26/43371113).

Type level: Lower Mangaopari Mudstone, Globorotalia crassaformis Zone, Opoitian (Early Pliocene).

Remarks: This species evolved from B. striata, which is distinguished by its relatively squat form, inflated rhomboidal
cross-section and more prominent costae.

Distribution: The species occurs throughout the Wainuioru Valley in massive fine grained hemipelagic sediments of
lower and middle Mangaopari Mudstone.

Observed stratigraphic range: Early Opoitian to Waipipian (Early Pliocene).

Observed ecological range: Upper to middle bathyal.

111
 DRAFT
Figure C.1: Bolivinita obconica Crundwell n. sp. #A-#L T27f131 (type locality). Imagery by W. St George.

112
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Locality Index

DRAFT
xxx/fxxx refer to collections in the New Zealand Fossil Record File http://www.fred.org.nz

C45/f163, 52 Q18/f102, 46
C45/f9511, 52 Q18/f13, 56, 57, 65
C46/f46, 27 Q18/f18, 104
C46/f53, 27 Q18/f36, 28
C46/f54, 27 Q18/f38, 50
Challenger Station 209, Philippine Islands, 102 Q18/f72, 28
Corehole D, 46 Q18/f8512, 27
Corehole D, 35 ft, 75 Q19/f102, 44, 48
Q19/f21, 52
D45/f8466, 102 Q19/f3, 53, 69, 70
D45/f8625, 102 Q19/f4, 52
D45/f8646, 9, 102 Q19/f8540, 53
D45/f8670, 9 Q19/f9, 52
DSDP Site 592, 76
R22/f7516, 71, 74, 75, 83, 84
J32/f44, 74
J32/f54, 46, 48, 91–92 S21/f12, 37
J41/f9499, 9 S21/f9, 37
S28/f46, 52
Kahawai-1, 69, 93
S28/f48, 52
Kapuni-2, 37
SE40176/f1, 43
Kawhaka-1 890 ft, 86
Kiwa-1, 69 T25/f6910, 65, 67, 68
T26/f131, 111
Manutahi-1, 37
T26/f133, 109
Maui-26, 69
T26/f139, 24, 26, 65
Mf/42, 88
T26/f6910, 61, 62, 64
N32/f9207, 59 T27/f122, 9, 100
N32/f9209, 59 T27/f139, 25, 107
N33/f9694, 26 T27/f399, 22, 23, 25, 34, 35, 41, 48
N33/f9696, 27 T27/f431, 79
N34/f7563, 93 T27/f432, 79
N99/f540, 70 T27/f460, 9, 12, 87, 89–91, 96
T27/f464, 87
O33/f13, 87, 93 T27/f495, 81–85, 87
O33/f15, 87, 93 T27/f503, 59
ODP Site 1119, 76 T27/f551, 12, 79, 80, 82, 83
T27/f78, 30–32, 34–36
P29/f11, 23, 24, 52, 69 T27/f95, 34, 38
P29/f13, 69 Taimana-1, 37
P29/f14, 47, 52, 66, 69 Taranga-1 1600 m, 76
P29/f16, 63–66, 69, 70 Te Kiri-1, 37
P29/f18, 64, 65, 69
P29/f26, 69 U20/f8547, 37
P29/f28, 69 U23/f142, 36, 39–41
P29/f29, 67–69, 73, 74, 83, 85 U23/f143, 43
P29/f30, 85 U24/f6049, 39, 43
P29/f32, 42, 43 U24/f697, 36, 39, 41
P29/f60, 75 U24/f698, 36, 39
P29/f62, 75
P29/f73, 53 V20/f149, 37, 38
P30/f7594, 97 V20/f386, 69

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V20/f390, 27
V21/f6005, 38

DRAFT
V22/f8637, 59
V22/f8660, 93
V22/f8772, 87, 93
V22/f8842, 93
V22/f8850, 93
V24/f10, 38

W19/f64, 28, 34
W19/f65, 28
W21/f8521, 33, 34, 36, 43
W21/f8545, 43
Witiora-1, 37

X18/f12, 69
X18/f38, 20–22, 25, 34, 35, 41, 48
X18/f7466, 20
X18/f7763, 38
X19/f7732, 69
X19/f7749, 28
X19/f7758, 28
X19/f7763, 28
X19/f9493, 32, 34
X19/f9509, 95, 96

Y18/539, 59
Y18/f506, 59
Y18/f552, 28
Y18/f554, 28
Y18/f560, 25, 26, 28
Y18/f7479, 50, 54–57, 65

Z14/f8, 69
Z14/f87, 52
Z14/f9503, 94

118
Taxon Index

DRAFT
Not exhaustive, please supplement with reader search tool

Austrofusus coerulescens, 77
Austrofusus wollastoni, 77
Bolivina alata, 11
Bolivina, 9, 10, 12, 13, 25, 100
Bolivinita compressa, i, 2, 6, 8, 9, 11, 14–16, 20, 24, 26, 27, 34, 37, 41, 42, 48, 52, 54, 55, 57–59, 65, 69, 70, 77, 79,
80, 82, 92–93, 107
Bolivinita elegantissima, i, 2, 9, 15, 100, 104, 109
Bolivinita finlayi, i, 2, 14, 15, 33, 36, 39–43
Bolivinita medialis, i, 2, 6, 7, 12–15, 27, 59, 75, 80–87, 93
Bolivinita obconica, i, 2, 109, 111
Bolivinita pliobliqua, i, 2, 6, 9, 13, 15, 20, 24, 42, 48, 53, 61–70, 72, 74, 75, 86
Bolivinita pliozea, i, 2, 6, 7, 9–16, 20, 37, 48, 61, 64, 65, 67–76, 78, 80, 83–86
Bolivinita pohana, i, 2, 6, 14, 15, 26, 50, 53–60, 64–66, 68–70, 77, 86, 87, 92–93, 104, 109
Bolivinita pseudocompressa, i, 2, 6, 9, 11–13, 15, 28, 30–32, 34, 36, 37, 43, 107
Bolivinita quadrilatera granttaylori, i, 2, 94, 96–97
Bolivinita quadrilatera, i, 2, 6–15, 24, 46, 54, 59, 86, 87, 89–94, 96–97
Bolivinita striata, i, 2, 109, 111
Bolivinita subangularis, i, 2, 9, 15, 100, 102
Bolivinita urenuia, i, 2, 6, 23, 24, 44–53, 59, 66, 67, 69, 70
Bolivinita cf. compressa, 36
Bolivinita cf. pohana, 23
Bolivinita sp. aff. pohana striate, 104, 109
Florilus parri, 37
Globoquadrina dehiscens, 20, 22, 28, 46, 52, 60, 70, 87
Globorotalia crassaconica, 28
Globorotalia fohsi robusta, 15, 100
Globorotalia inflata, 28, 38
Globorotalia miotumida, 80, 87, 93
Globorotalia mons, 34
Globorotalia pliozea, 28
Globorotalia puncticulata, 28, 34
Loxostomum truncatum, 93
Notorotalia zelandica, 37
Praeorbulina circularis, 102
Sigmoilopsis schlumbergeri, 42

119