Five new species and one new genus of recent miliolid foraminifera from Raja Ampat (West Papua, Indonesia)

Abstract

Raja Ampat is an archipelago of about 1,500 small islands located northwest off the Bird’s Head Peninsula of Indonesia’s West Papua province. It is part of the Coral Triangle, a region recognized as the “epicenter” of tropical marine biodiversity. In the course of a large-scale survey on shallow benthic foraminifera we have discovered one new genus and five new species of recent miliolid benthic foraminifera from the highly diverse reefal and nearshore environments. The new fischerinid genus
Dentoplanispirinella
is characterized by its planispiral coiling and by the presence of a simple tooth, that differentiate it from
Planispirinella
Wiesner. It is represented in our sample material by the new species
Dentoplanispirinella occulta
. The other four species described herein are
Miliolinella moia, Miliolinella undina, Triloculina kawea
and
Siphonaperta hallocki.
All new species are comparatively rare and occur sporadically in the sample material. Detailed morphological descriptions, scanning electron microscopy pictures of complete and dissected specimens as well as micro-computed tomography images are provided.

Full text

Submitted 27 February 2016
Accepted 31 May 2016
Published 23 June 2016
Corresponding author
Martin R. Langer,
martin.langer@uni-bonn.de
Academic editor
James Reimer
Additional Information and
Declarations can be found on
page 15
DOI 10.7717/peerj.2157
Copyright
2016 Förderer and Langer
Distributed under
Creative Commons CC-BY 4.0
OPEN ACCESS
Five new species and one new genus of
recent miliolid foraminifera from Raja
Ampat (West Papua, Indonesia)
Meena Förderer and Martin R. Langer
Steinmann Institut, Paleontology, University of Bonn, Bonn, NRW, Germany
ABSTRACT
Raja Ampat is an archipelago of about 1,500 small islands located northwest off the
Bird’s Head Peninsula of Indonesia’s West Papua province. It is part of the Coral
Triangle, a region recognized as the ‘‘epicenter’’ of tropical marine biodiversity. In
the course of a large-scale survey on shallow benthic foraminifera we have discovered
one new genus and five new species of recent miliolid benthic foraminifera from
the highly diverse reefal and nearshore environments. The new fischerinid genus
Dentoplanispirinella is characterized by its planispiral coiling and by the presence of
a simple tooth, that differentiate it from Planispirinella Wiesner. It is represented
in our sample material by the new species Dentoplanispirinella occulta. The other
four species described herein are Miliolinella moia, Miliolinella undina, Triloculina
kawea and Siphonaperta hallocki. All new species are comparatively rare and occur
sporadically in the sample material. Detailed morphological descriptions, scanning
electron microscopy pictures of complete and dissected specimens as well as micro-
computed tomography images are provided.
Subjects Biodiversity, Paleontology, Taxonomy
Keywords Foraminifera, Coral Triangle, Raja Ampat, Indonesia, Tropical reefs, Benthic, Protists
INTRODUCTION
The Raja Ampat Archipelago (West Papua, Indonesia) off the northwestern coast of New
Guineas Bird’s Head Peninsula (Fig. 1B) is one of the most species rich marine environments
(Erdmann & Pet,2002;McKenna, Allen & Suryadi,2002), situated in the Indo-Pacific’s
‘‘epicenter’’ of biodiversity, commonly referred to as the Coral Triangle (Hoeksema,2007).
The Coral Triangle encompasses a large part of the tropical marine waters of Indonesia,
the Philippines, Malaysia, the Solomon Islands, Papua New Guinea and Timor-Leste
(Fig. 1A). It includes ecoregions that are each home to at least 500 species of hermatypic
corals and also show extraordinary diversity among coral associated species (Veron,1995;
Veron et al.,2009;Roberts et al.,2002;Bellwood & Hughes,2001;Tittensor et al.,2010).
The region is recognized as a ‘‘species factory’’ and functions as the most significant net
exporter of biodiversity for adjacent reef regions (Briggs & Bowen,2013;Ekman,1953).
Comprehensive studies on benthic foraminifera from the central Indo-Pacific region
began with marine scientific expeditions in the late 1800s with the report on the Challenger
Foraminifera by Brady (1884), and the work of Millett (1898–1904) from the Malay
Archipelago. In the 20th century, systematic surveys were conducted around the Philippines
How to cite this article Förderer and Langer (2016), Five new species and one new genus of recent miliolid foraminifera from Raja Am-
pat (West Papua, Indonesia). PeerJ 4:e2157; DOI 10.7717/peerj.2157

Figure 1 Maps of the sampling area. (A) Area of the Coral Triangle (shaded) in the Central Indo-Pacific;
(B) location of Raja Ampat northwest of the Bird’s Head Peninsula (West Papua, Indonesia); (C) location
of sample stations where the species described herein occur (for details see Table 1).
(Cushman,1921;Graham & Militante,1959), in the Papuan Lagoon near Port Moresby,
Papua New Guinea (Haig,1988a;Haig,1988b;Haig,1993), in the Timor Sea and Sahul
Shelf (Loeblich & Tappan,1994), in Madang, eastern Papua New Guinea (Langer,1992;
Langer & Lipps,2003) and more recently in the Ningaloo Reef area at Australia’s northwest
coast (Parker,2009), at Chuuk Island of the Caroline reefs (Makled & Langer,2011) and
around New Caledonia (Debenay,2012). Recent environmental and biogeographic studies
on larger benthic foraminifera in the tropical waters of the central Indo-Pacific were
conducted by Langer & Hottinger (2000), Renema (2003), Renema & Hohenegger (2005),
Renema (2006), Renema (2010), Renema & Troelstra (2001), Hohenegger (2004), Hohenegger
(2011), Weinmann et al. (2013) and Prazeres, Uthicke & Pandolfi (2016).
To date, however, large-scale systematic studies on benthic foraminifera from Raja
Ampat are lacking. The archipelago consists of the four main islands Waigeo, Batana,
Salawati, and Misool, and hundreds of small satellite islets, which are largely uninhabited.
Due to its remote location and difficult access conditions the coral reefs of the region
remained relatively unexplored and pristine. However, increasing exposure to exploitation
have required the establishment of several marine protected areas (Agostini et al.,2012).
The first and to date only report on benthic foraminifera from Raja Ampat is that of Hofker
(1927) and Hofker (1930), who examined the material taken by the Siboga Expedition
(1899–1900) that included five samples from Raja Ampat. He documented nine species of
benthic foraminifera including eight rotalid taxa and the miliolid symbiont bearing species
Peneroplis pertusus (Forskål).
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 2/20

Table 1 Collection sites. Sample site information for collection locations from Raja Ampat (Indonesia) including environmental information on
reefal habitat type.
Station Depth (m) Latitude Longitude Reef- or eco-type Reef-zone
B14 41 0◦5022.4000N 130◦13035.2900 E Fringing reef Fore-reef slope
B15 43 0◦5022.4000N 130◦13035.2900 E Fringing reef Fore-reef slope
E23 24 0◦16025.8700S 130◦18059.1900 E Fringing reef Fore-reef slope
FW 49 0◦35019.8600S 130◦17045.5400 E Fringing reef Fore-reef slope
M21 27 0◦29050.4000S 130◦43037.6200 E Patch reef Fore-reef slope
MG 18 0◦35023.4000S 130◦18054.5400 E Patch reef Platform
MI05 32 0◦16025.8700S 130◦18059.1900 E Fringing reef Fore-reef slope
MI06 32 0◦16025.8700S 130◦18059.1900 E Fringing reef Fore-reef slope
MR17 12 0◦5047.5800S 130◦1409.6600 E Fringing reef Fore-reef slope
MR18 18 0◦5047.5800S 130◦1409.6600 E Fringing reef Fore-reef slope
MS03 16 0◦34047.8800S 130◦32032.0400 E Sand channel, sparse coral cover Channel
MS04 14 0◦34047.8800S 130◦32032.0400 E Sand channel, sparse coral cover Channel
N18 30 0◦10022.7400N 130◦0022.3800 E Fringing reef Fore-reef slope
U16 45 0◦5049.1300N 130◦13059.0800 E Fringing reef Fore-reef slope
W07 24 0◦15021.7200S 130◦17032.1600 E Fringing reef Fore-reef slope
Y24 26 0◦4708.6400S 130◦45025.6200 E Fringing reef Fore-reef slope
Y25 26 0◦4708.6400S 130◦45025.6200 E Fringing reef Fore-reef slope
MATERIAL AND METHODS
This study was conducted with 30 sediment samples from the Raja Ampat Archipelago (New
Guinea, Indonesia) from around the islands of Waigeo, Batana, Kawe, Fam and adjacent
small islets in an area that covers about 2,500 km2(Fig. 1C). The archipelago is located in
the central Indo-Pacific warm pool with an average annual sea surface temperature of 29 ◦C
(Mangubhai et al.,2012). Raja Ampat is further situated in the passage way of the Indonesian
Throughflow, a major ocean current that leads water masses from the western Pacific to
the eastern Indian Ocean. Previous studies have shown that the reef fauna of Raja Ampat
is strongly current dependent (Devantier, Turak & Allen,2009;Turak & Souhoka,2003).
The samples were collected by snorkeling and SCUBA diving in September 2011 by M
Langer. Sediment surface samples from the top 2 cm were collected from the fore-reef
slope of fringing reefs, with two samples from a patch reef, and two samples from a sandy
channel with sparse coral cover (Table 1). The sediment was predominantly carbonaceous
(∼90%) and included fine-grained sediments as well as coarse reef rubble. All samples
were washed through a 63 µm sieve and dried at 50 ◦C in an oven overnight. Foraminifera
were picked from each sample and the best preserved specimens were imaged using a
Tescan VEGA MV2300 Scanning Electron Microscope (SEM) at the Steinmann Institute
of the University of Bonn. Digital plates were assembled using Adobe Photoshop CS6.
Micro-computer tomography (CT) scan imaging was conducted using a phoenix v|tome|x
s computed tomography system at the Steinmann Institute and visualization was carried
out with Avizo 7.1.0. The new species and the new genus are described in detail using the
supra-generic classification of Loeblich & Tappan (1987).
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 3/20

Repository of the Material: the holotypes and topotypic paratypes of the new species are
deposited in the micropaleontology collection of the Steinmann Institute of Paleontology
at the University of Bonn, Germany (MaLaPNG 2011–10, MaLaPNG 2011–11, MaLaPNG
2011–12, MaLaPNG 2011–13, MaLaPNG 2011–14).
The electronic version of this article in Portable Document Format (PDF) will represent
a published work according to the International Commission on Zoological Nomenclature
(ICZN), and hence the new names contained in the electronic version are effectively
published under that Code from the electronic edition alone. This published work and the
nomenclatural acts it contains have been registered in ZooBank, the online registration
system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the
associated information viewed through any standard web browser by appending the LSID
to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:
pub:FB001C3C-AEA9-45D5-9224-EDD084378897. The online version of this work is
archived and available from the following digital repositories: PeerJ, PubMed Central and
CLOCKSS.
RESULTS
Smaller miliolid benthic foraminifera are typical dwellers in surface sediments of shallow
water reefal and lagoonal habitats. By studying the highly diverse assemblages of benthic
foraminifera from Raja Ampat, taken from different locations around the islands (Fig. 1C),
we recorded a total of 455 species among them 249 miliolid species, of which five are
described here as new. Four species belong to the widely distributed miliolid genera of
Miliolinella Wiesner, Triloculina d’Orbigny and Siphonaperta Vella. As the morphological
properties of the fifth species differentiate it from any previously known genera, we
designate and describe it as the new genus Dentoplanispirinella.
SYSTEMATIC DESCRIPTIONS
Subclass Miliolana Saidova,1981
Order Miliolida Delage & Hérouard,1896
Suborder Miliolina Delage & Hérouard,1896
Superfamily Cornuspiracea Schultze,1854
Family Fischerinidae Millett, 1898
Subfamily Fischerininae Millett, 1898
Genus Dentoplanispirinella Förderer and Langer gen. nov.
urn:lsid:zoobank.org:act:98A1DD41-C0AE-4401-830B-0D189E70661A
Description. Test small, broadly circular in outline, discoidal to slightly biconvex. Periphery
with a weakly developed subrounded keel that encircles the entire test margin. Wall thick,
calcareous, porcelaneous, imperforate. Coiling involute, throughout planispirally enrolled
with 2.5 to 3.5 tubular chambers per whorl, each whorl slightly offset to the proceeding coil
with a tendency to become sigmoiline (axial section as seen in CT scan, Fig. 2H). Lateral
wall extensions of the adult chambers entirely cover the earliest chambers and tend to
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 4/20

Figure 2 Holotype, paratype, CT scans and details of Dentoplanispirinella gen. nov. occulta sp. nov.
(A) Side view and (B) apertural view of a more juvenile specimen with a nearly triangular aperture and
weakly developed tooth (paratype); (C) side view and (D) apertural view (holotype); (E) apertural view
of a specimen with a well-developed tooth and elongated aperture; (F) detail of a well-developed periph-
eral keel; (G) CT scan reconstruction of the chamber cavities revealing the presence of 2.5 – 3.5 chambers
per whorl in an adult specimen (note that penultimate chamber is broken); (H) CT scan showing planispi-
rally arranged chambers; (I) detail of an aperture with a very well-developed tooth; (J) detail of the striate
surface ornamentation; (K) detail of the construction of the outer wall layer showing randomly arranged
calcite needles in the lower part (test surface removed) and longitudinally arranged calcite needles on the
outer test surface. Scale bar is 100 µm (unless indicated).
overlap the umbilical region in each whorl. Sutures oblique, thin and irregular. Aperture
arch-shaped, triangular in juvenile specimens, high and subtriangular in adult specimens,
tapering apically, on the base connected with the peripheral margin of the proceeding
chamber and provided with a very small and thin tooth. In juvenile specimens the tooth
appears just like a little knob or slightly raised spine.
Type species. Dentoplanispirinella occulta sp. nov.
Remarks. Dentoplanispirinella gen. nov. resembles Planispirinella Wiesner,1931 in having
a discoidal shape, a high aperture and a planispiral chamber arrangement, but differs
from Planispirinella by the presence of a tooth and the more biconvex test shape in apical
view (Fig. 2B). The apertural features and the coiling mode of Dentoplanispirinella further
distinguish it from Nummoloculina Steinmann 1881, which has an apertural flap and an
early quinqueloculine coiling.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 5/20

Dentoplanispirinella occulta Förderer and Langer sp. nov.
Figures 2A–2K
urn:lsid:zoobank.org:act:7E132939-5284-484D-9B50-BC79A0B52D0A
Etymology. From the Latin ‘‘occultare’’ meaning for ‘‘hiding.’’
Material. 28 specimens from nine samples (MR18, MI05, MI06, MS03, MS04, MG, M21,
U16, Y24; Fig. 1C;Table 1), recent.
Holotype. The specimen illustrated here as Figs. 2C and 2D (sample MS03; MaLaPNG
2011–10).
Paratype. The specimen illustrated here as Figs. 2A and 2B (sample MS03; MaLaPNG
2011–10).
Type locality. The holotype and the paratype are from sample station MS03 (16m), a
sand channel between Arborek Island and Pulau Mansuar; Raja Ampat, New Guinea
(Indonesia).
Diagnosis. A species of Dentoplanispirinella gen. nov. with a discoidal to biconvex test
shape, a slightly keeled periphery, a radial oriented, finely striate surface ornamentation
and an arch-shaped, triangular aperture, provided with a small tooth.
Dimensions. Observed range of test dimensions: diameter 285–704 µm (lateral view), test
width 100–193 µm (apertural view).
Occurence.Dentoplanispirinella occulta is widely distributed in the Raja Ampat area in fine
to coarse coral rubble samples from depths of 14 to 45 m.
Description. Test porcelaneous and imperforate. Almost circular in lateral view, lenticular
and biconvex in apertural view with a slightly developed, subrounded keel and weakly
inflated chambers. Coiling planispiral and involute. Two and a half to three and a half
chambers visible from the exterior. Lateral wall extensions of the adult chambers entirely
cover the earliest chambers and tend to overlap the umbilical region; the final chamber
covers approximately half of the test surface. Sutures oblique, thin, irregular and recurved
near the periphery. Test surface ornamented with radial oriented, fine, sub-parallel to
anastomosing striae that are straight to slightly curved backwards, towards the outer
margins of the chambers. Umbilical region and test periphery more weakly ornamented.
Outer wall layer constructed of longitudinally aligned needle-shape crystals, oriented
perpendicular to direction of ornamentation. The test appears matte white under the
light microscope with a slightly translucent periphery. Apertural face not ornamented.
Aperture arch-shaped and triangular in juvenile specimens, high and subtriangular in
adult specimens, tapering apically, on the base connected with the peripheral margin to
the preceding chamber and provided with a peristomal rim. Aperture provided with a very
small and thin tooth, with the flat side oriented in lateral direction.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 6/20

Remarks. Dentoplanispirinella occulta sp. nov. differs from Planispirinella involuta Collins
(1958, p. 374, pl. 4, Figs. 2A and 2B) by its more lenticular biconvex shape in horizontal
section, the subtriangular shape of the aperture, the presence of a small tooth, and the
striate surface ornamentation.
Superfamily Miliolacea Ehrenberg,1839
Family Hauerinidae Schwager,1876
Subfamily Hauerininae Schwager,1876
Genus Miliolinella (Wiesner,1931)
Miliolinella moia Förderer and Langer sp. nov.
Figures 3A–3L
urn:lsid:zoobank.org:act:D8184E0C-2805-40D7-BCCB-492D74216168
Etymology. The new species is named after the indigeneous Moi people from Malaumkarta,
a Papuan tribe from the north coast near Sorong.
Material. 11 specimens from six samples (B14, B15, E23, MR17, N18, U16; Fig. 1C;
Table 1), recent.
Holotype. The specimen illustrated here as Figs. 3A–3C (sample B14; MaLaPNG 2011–11).
Paratypes. The specimens illustrated here as Figs. 3D–3F (sample B14), Figs. 3G–3I and
Figs. 3J–3J (sample ER23; MaLaPNG 2011–11).
Type locality. The holotype and the paratype are from sample station B14 (41 m), Bag
Island, east of Pulau Uranie; Raja Ampat, New Guinea (Indonesia).
Diagnosis. A slightly enlongated, medium-sized species of Miliolinella Wiesner,1931 with
a compressed, angular and slightly slanted outline, a smooth and shiny wall, and a high
subcircular opening.
Dimensions. Observed range of test dimensions: test height 409–554 µm, test width
278–396 µm (lateral view), 166–250 µm (apertural view).
Occurence. This species is widely distributed in the Raja Ampat area in fine to coarse coral
rubble samples and occurs at depths between 12 and 45 m.
Description. Test porcelaneous and imperforate, ovate in outline and slightly higher than
broad. Test weakly compressed and flattened, subtriangular in apertural view. Chamber
arrangement quinqueloculine with five chambers visible from the exterior. In some
specimens only three to four chambers are visible. Periphery rounded to subrounded,
chambers slightly inflated. Sutures curved, distinct and weakly depressed. Chambers
tend to be off-centered, giving them a slanted appearance. Test wall smooth, translucent
to opaque and glossy under the light microscope. Aboral end of the chambers slightly
constricted. Aperture in basal position, a Miliolinella-type large subcircular opening with
an everted peristomal rim and a semicircular, slightly excavated flap, that covers more than
half of the opening.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 7/20

Figure 3 Holotype and paratypes of Miliolinella moia sp. nov. (A–C) Holotype, five chambers visible
from the exterior: (A) lateral view of more evolute side; (B) top view; (C) lateral view of more involute
side; (D–F) paratype, a specimen with a broken ultimate chamber showing three chambers visible from
the exterior: (D) lateral view of more evolute side; (E) top view; (F) lateral view of more involute side; (G–
I) a specimen with four chambers visible from the exterior: (G) lateral view of more evolute side; (H) top
view; (I) lateral view of more involute side; (J–L) a specimen with four chambers visible from the exte-
rior: (J) lateral view of more involute side; (K) top view; (L) lateral view of more evolute side. Scale bar is
100 µm.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 8/20

Remarks. Miliolinella moia sp. nov. differs from Miliolinella pilasensis McCulloch,1977 (p.
566, pl. 238, Fig. 16 and Loeblich & Tappan,1994, p. 57, pl. 99, Figs. 1–9) in its angular
and more compressed outline, and the large subcircular opening. Millet (1898) depicted
a species of Miliolina valvularis (Reuss) from the Malay Archipelago (p. 11, Figs. 5A–5C)
that shows a high degree of similarity to Miliolinella moia, but his specimen has a more
rounded periphery. The original description of Triloculina valvularis by Reuss (1851, p. 85,
pl. 7, Fig. 56) shows a specimen with a broadly rounded periphery and inflated chambers
without angles. Miliolinella sp. 2 figured in Parker,2009 from Ningaloo Reef, Australia (p.
128, Figs. 92A–92I, 93A–93J, 94A–94K) differs from Miliolinella moia by the low apertural
opening and the broadly rounded and more inflated chambers.
Miliolinella undina Förderer and Langer sp. nov.
Figure 4A–4I
urn:lsid:zoobank.org:act:D11E1426-9DCC-41B8-A992-27D974A92520
1988a Miliolinella sp. B—Haig, Papuan Lagoon, Port Moresby, p. 224, pl. 2,
Figs. 23 and 24.
1992 Miliolinella sp.—Hatta & Ujiié, Ryukyu Islands, p. 72, pl. 10, Fig. 6.
?2012 Miliolinella cf. M. semicostata (Wiesner)—Debenay, New Caledonia,
p. 110, 275.
Etymology. After the undulate ornamentation of the test. From the Latin ‘‘unda’’ meaning
wave and mythological ‘‘Undine,’’ a term established by the Renaissance alchemist
Paracelcus for water spirits.
Material. Three specimens from three samples (MR18, N18, U16; Fig. 1C;Table 1), recent.
Holotype. The specimen illustrated here as Figs. 4A–4C (sample MR18; MaLaPNG
2011–12).
Paratypes. The specimens illustrated here as Figs. 4D–4F (sample N18) and Figs. 4G–4I
(sample U16; MaLaPNG 2011–12).
Type locality. The holotype is from sample station MR18 (18 m), east of Kawe Island. The
paratypes are from sample stations N18 (30 m), south-west coast of Pulau Wayag, and U16
(45 m), between Pulau Uranie and Bag Island; Raja Ampat, New Guinea (Indonesia).
Diagnosis. A small quinqueloculine species of Miliolinella Wiesner with inflated chambers,
a rounded outline and an undulate to reticulate surface ornamentation.
Description. Test porcelaneous and imperforate, small, ratio of height and width variable
but usually slightly higher than broad. Periphery rounded and chambers slightly inflated.
Chamber arrangement quinqueloculine, with five chambers visible from the exterior.
Aboral end rounded, flush with the surface in the holotype to slightly raised in paratypes.
Wall smoothly finished, matte, translucent under the light microscope. Sutures curved and
depressed. Test surface ornamented with numerous irregular, predominantly longitudinal,
somehow honeycombed reticulate to undulate low anastomosing costae that are covering
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 9/20

Figure 4 Holotype and paratypes of Miliolinella undina sp. nov. (A–C) Holotype: (A) oblique apertu-
ral view; (B) apertural view; (C) lateral view of more involute side; (D–F) a specimen with the final cham-
ber missing: (D) lateral view of more evolute side; (E) top view; (F) lateral view of more involute side; (G–
I) a specimen with an erratic growth stage in the final chambers: (G) side view; (H) top view; (I) lateral
view of more involute side. Scale bar is 50 µm.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 10/20

large parts of the test. Outer-wall layer constructed of needle-shaped crystals that are
primarily aligned in longitudinal direction. Aperture basal, a large semicircular Miliolinella-
type opening, provided with a thickened and everted peristomal rim and a broad, slightly
excavated basal flap.
Dimensions. Observed range of test dimensions: test height 146–162 µm, test width
114–224 µm (lateral view), 81–119 µm (apertural view).
Occurence. Miliolinella undina is present with one specimen in each of three highly diverse,
miliolid-rich, fine coral rubble samples from depths of 18 to 45 m.
Remarks. Specimens of Miliolinella undina sp. nov. have been previously documented
by Haig,1988a as Miliolinella sp. B from the Papuan Lagoon, Port Moresby and by
Hatta & Ujiié,1992 as Miliolinella sp. from the Ryukyus. Hatta & Ujiié mentioned the
species to occur rarely in their assemblages. The new species has also been recorded in
samples from northern Palawan (M Förderer, 2016, unpublished data). Miliolinella cf.
M. semicostata (Wiesner) depicted by Debenay from New Caledonia (2012, p. 110, 275)
may also belong to Miliolinella undina, but shows a less undulated test ornamentation.
Test shape, apertural and ornamental features are more similar to our holotype (Figs.
4A–4C) than to Miliolinella semicostata (Wiesner,1923) from the Mediterranean Sea (see
Cimerman & Langer,1991, p. 42, pl. 38, Figs. 10–15). Miliolinella semicostata has less inflated
chambers and the ornamentation is not reticulate but longitudinally striate and restricted
to the angles. Miliolinella undina also resembles Miliolinella sp. 4 depicted by Parker from
the Ningaloo Reef in Western Australia (2009, p. 136, Figs. 97A–97H), but his specimen
has a less undulated and more striate alignment of costae. The new species also resembles
Miliolinella flintiana (Cushman,1932) (p. 55, pl. 12: 4A–4C) in size, test shape, chamber
arrangement and apertural features. However it differs in its surface ornamentation, that
is distinctly longitudinal costate in Miliolinella flintiana and undulate and more irregular
in Miliolinella undina.Miliolinella flintiana also occurs in our assemblages.
Genus Triloculina D’Orbigny,1826
Triloculina kawea Förderer and Langer sp. nov.
Figs. 5A–5H
urn:lsid:zoobank.org:act:6F5B38CE-88B3-4FBE-9329-8483756158E1
2009 Triloculina? sp. 2—Parker, Ningaloo Reef, p. 372, Figs. 271F–271J.
Etymology. This species is named in honor of the indigeneous people of West Papua after
the Kawe tribe, that owns and protects a highly diverse marine protected area of Raja
Ampat.
Material. 12 specimens from seven samples (B15, FW, M05, MS04, N18, U16, Y25; Fig.
1C;Table 1), recent.
Holotype. The specimen illustrated here as Figs. 5A–5C (sample FW; MaLaPNG 2011–13).
Paratype. The specimen illustrated here as Figs. 5E–5G (sample FW; MaLaPNG 2011–13).
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 11/20

Figure 5 Holotype, paratype, cross section and detail of Triloculina kawea sp. nov. (A–C) Holotype:
(A) lateral view of more involute side; (B) apertural view; (C) lateral view of more evolute side; (D) cross
section of a specimen; (E–G) Paratype: (E) lateral view of more evolute side; (F) apertural view; (G) lateral
view of more involute side; (H) detail of the irregular test surface. Scale bar is 100 µm (unless indicated).
Type locality. The holotype and the paratype are from sample station FW (49 m), south-east
Penemu, Fam Islands; Raja Ampat, New Guinea (Indonesia).
Diagnosis. A medium-sized species of Triloculina d’Orbigny with a slightly elevated
‘‘Lachlanella’’-type aperture, rounded periphery, blunt angles and a roughly textured wall.
Dimensions. Observed range of test dimensions: test height 377–439 µm, test width
200–245 µm (lateral view), 162–195 µm (apertural view).
Occurence. This species is widely distributed in our sampling area in fine to coarse coral
rubble samples from depths of 14 to 49 m.
Description. Test porcelaneous and imperforate, about one and a half times longer
than broad. Broadly triangular in apertural view, ovate in outline. Chamber arrangement
triloculine, periphery rounded to subrounded, chambers inflated with blunt angles. Sutures
distinct and depressed. Surface ornamented with elongated, irregular longitudinal arranged
short striae covering the entire test surface, giving the appearance of a matte and roughly
textured wall under the light microscope. Outer wall layer consisting of longitudinally
aligned plate shaped crystals. Aboral end rounded and slightly produced, oral end
produced and connected with the peripheral margin of the preceeding chamber. Aperture
basal, ‘‘Lachlanella’’-type with a long slender tooth that becomes thickened at the tip.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 12/20

Remarks. The species Triloculina? sp. 2 reported by Parker,2009 from Western Australia
differs from Triloculina kawea sp. nov. in its less triangular shape and less elongated outline.
We consider Parker’s specimen a juvenile individual of Triloculina kawea. The aperture of
Parker’s specimen is not intact but resembles very well the apertural features of Triloculina
kawea. The outer wall layer appears identical (Fig. 5H). Parker mentioned the species to
be possibly cryptoquinqueloculine. Figures 5B and 5F and the horizontal section (5D)
show the triloculine chamber arrangement. Triloculina sp. 1, reported by Debenay 2012
from New Caledonia (p. 139, 278) is very similar in shape and surface ornamentation to
Triloculina kawea, but has significantly more acute angles and a Y-shaped tooth. Triloculina
kawea further differs from Triloculina linneiana d’Orbigny depicted by Baccaert,1987 from
the Great Barrier Reef (p. 128, pl. 57, Figs. 3 and 4) in the less striate ornamentation and
more acute angles.
Subfamily Siphonapertinae Saidova,1975
Genus Siphonaperta Vella,1957
Siphonaperta hallocki Förderer and Langer sp. nov.
Figs. 6A–6F
urn:lsid:zoobank.org:act:DD4F0DB3-1355-4BB1-841A-FFE32E0F6455
?1988a Quinqueloculina sp. C—Haig, Papuan Lagoon, Port Moresby, p. 234, pl. 9,
Figs. 7–10.
?2009 Quinqueloculina sp. 13—Parker, Ningaloo Reef, p. 311, Figs. 224A–224J,
225A–225G.
Etymology. In honor of Pamela Hallock Muller for her extensive work on tropical
foraminifera.
Material. Four specimens from three samples (MS03, N18, W07; Fig. 1C;Table 1), recent.
Holotype. The specimen illustrated here as Figs. 6A–6C (sample MS03; MaLaPNG
2011–14).
Paratype. The specimen illustrated here as Figs. 6D–6F (sample N18; MaLaPNG 2011–14).
Type locality. The holotype is from sample station MS03 (18 m), a sand channel between
Arborek Island and Pulau Mansuar. The paratype is from sample station N18 (30 m),
south-west coast of Pulau Wayag; Raja Ampat, New Guinea (Indonesia).
Diagnosis. A medium-sized species of Siphonaperta Vella with a finely agglutinated wall,
carinate shoulders, a short neck and a circular aperture with a small bifid tooth.
Description. Test porcelaneous and imperforate, medium-sized, about two times longer
than broad, and ovate in outline. Outer layer of the calcareous test wall covered with
finely agglutinated mostly biogenic grains. Agglutinated grains are particularly frequent
along the sutures. Periphery carinate to subacute. Chamber arrangement quinqueloculine
with five chambers visible from the exterior. Sutures slightly curved, incised and depressed.
Chambers weakly inflated and angular in section, with weakly developed longitudinal striae
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 13/20

Figure 6 Holotype and paratype of Siphonaperta hallocki sp. nov. (A–C) Holotype: (A) lateral view of
more evolute side; (B) apertural view; (C) lateral view of more involute side; (D–F) paratype: (D) lateral
view of more evolute side; (E) apertural view; (F) lateral view of more involute side. Scale bar is 100 µm
(unless indicated).
(in well preserved specimens). Aboral end rounded and produced; oral end becoming more
slender and leading into a short produced neck. Aperture terminal, a wide circular opening
with a short T-shaped, bifid tooth, that reaches more than one third of the apertural
diameter. Apertural opening surrounded by a slightly thickened and everted peristomal rim.
Dimensions. Observed range of test dimensions: test height 240–442 µm, test width
132–233 µm (lateral view), 87–119 µm (apertural view).
Occurence. Siphonaperta hallocki occurs sporadically in fine to coarse coral rubble samples
from depths of 16 to 30 m.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 14/20

Remarks. Very similar specimens were previously documented as Quinqueloculina sp. C
from the Papuan Lagoon (Haig,1988a) and Quinqueloculina sp. 13 from Ningaloo Reef
(Parker,2009). Test shape, wall texture and apertural features appear to be identical to
our specimens from Raja Ampat. Quinqueloculina sp. 4 documented by Parker,2009 from
the Ningaloo Reef appears very similar to Siphonaperta hallocki, but differs in its more
elongated shape, more rounded and inflated chambers and the cryptoquinqueloculine
coiling. In addition, Parker describes the wall as roughly textured with some agglutinated
grains. Quinqueloculina tropicalis Cushman from Samoa (1924, p. 63, pl. 23, Figs. 9 and 10)
differs from our new species by its more compressed shape and more elongated broadly
rounded chambers without any angles or costae. Quinqueloculina polygona D’Orbigny
(1839, p. 198,pl. 12, Figs. 21–23) differs from Siphonaperta hallocki in its smooth and shiny
surface, the pronounced carinae and the less inflated chambers. Langer et al.,2013 depicted
a specimen of Quinqueloculina polygona d’Orbigny from Bazaruto (Langer et al.,2013,
p. 163, Fig. 5: 14) that resembles our new species in size, shape and apertural features.
However, it is unlikely that this species from Bazaruto belongs to Siphonaperta hallocki, as
its outer wall layer is not agglutinated.
ACKNOWLEDGEMENTS
The authors gratefully acknowledge Dr. Stephanie Pietsch for assistance with the collection
of the samples, Georg Oleschinski for help with the SEM and Peter Göddertz and Kai Jäger
for support with CT scan imaging. We thank Justin H. Parker, Tomas Cedhagen and an
anonymous reviewer for constructive and helpful comments on the manuscript.
ADDITIONAL INFORMATION AND DECLARATIONS
Funding
Collection and study of the material was supported by grants from the German Science
Foundation (La-884/10-1/13- 1) and the University of Bonn. The funders had no role
in study design, data collection and analysis, decision to publish, or preparation of the
manuscript.
Grant Disclosures
The following grant information was disclosed by the authors:
German Science Foundation: La-884/10-1/13-1.
University of Bonn.
Competing Interests
The authors declare there are no competing interests.
Author Contributions
•Meena Förderer performed the experiments, analyzed the data, contributed
reagents/materials/analysis tools, wrote the paper, prepared figures and/or tables,
reviewed drafts of the paper.
Förderer and Langer (2016), PeerJ, DOI 10.7717/peerj.2157 15/20

•Martin R. Langer conceived and designed the experiments, performed the experiments,
analyzed the data, contributed reagents/materials/analysis tools, wrote the paper,
reviewed drafts of the paper.
Data Availability
The following information was supplied regarding data availability:
1) Micropaleontological Collection Steinmann Institute of Paleontology at the University
of Bonn, Germany
2) MaLaPNG 2011-10, MaLaPNG 2011-11, MaLaPNG 2011-12, MaLaPNG 2011-13,
MaLaPNG 2011-14.
New Species Registration
The following information was supplied regarding the registration of a newly described
species:
Dentoplanispirinella gen. nov. urn:lsid:zoobank.org:act:98A1DD41-C0AE-4401-830B-
0D189E70661A
Dentoplanispirinella occulta sp. nov.
urn:lsid:zoobank.org:act:7E132939-5284-484D-9B50-BC79A0B52D0A
Miliolinella moia sp. nov.
urn:lsid:zoobank.org:act:D8184E0C-2805-40D7-BCCB-492D74216168
Miliolinella undina sp. nov.
urn:lsid:zoobank.org:act:D11E1426-9DCC-41B8-A992-27D974A92520
Triloculina kawea sp. nov.
urn:lsid:zoobank.org:act:6F5B38CE-88B3-4FBE-9329-8483756158E1
Siphonaperta hallocki sp. nov.
urn:lsid:zoobank.org:act:DD4F0DB3-1355-4BB1-841A-FFE32E0F6455
The LSID for this publication is: urn:lsid:zoobank.org:pub:FB001C3C-AEA9-45D5-
9224-EDD084378897.
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