Micronesica 35-36:38-53. 2003
An annotated checklist of Foraminifera of Guam
SUSAN L. RICHARDSON
Smithsonian Marine Station at Fort Pierce
701 Seaway Dr.
Fort Pierce, FL 34949
email: richardson@sms.si.edu
RUSSELL N. CLAYSHULTE
Denver Regional Council of Governments
2480 Wt 26th Ave. Ste. 200B
Denver,CO 80211
Abstract—This paper presents a checklist of 303 species of benthic and
planktic Foraminifera from the marine waters off Guam, based on
literature records and new collections. Notations on taxonomic
synonymies and depositories of type and identified specimens are
included in the list.
Introduction
The checklist given below is comprised of 303 species representing all of the
major subclades of the clade Foraminifera (Pawlowski 2000). This species
checklist was compiled from various resources in the published literature
(Cushman 1917, 1932, 1933a, 1933b, 1934, 1942, Todd 1960, 1965, 1966,
Matsumaru & Matsuo 1976, Lessard 1980) using the preliminary checklist of 150
species published by Clayshulte (1981a) as a foundation. In addition, several
species identified from field collections by the authors were integrated into the
list (Clayshulte 1981a, b, S. Richardson, unpublished observations). As much as
possible, species identified in the published literature (Cushman 1917, 1932,
1933a, 1933b, 1934, 1942, Todd 1960, 1965, 1966) were checked against
specimens housed in the Cushman Collection of Foraminifera, Department of
Paleobiology, National Museum of Natural History, Smithsonian Institution,
Washington, DC. USNM numbers are listed for those specimens housed in the
Smithsonian primary and secondary type collections; however, many of
specimens listed by Todd (1966) are stored on assemblage (faunal) microslides
that are kept in separate slide cabinets adjacent to the type collections. These
microslides are labeled with the locality numbers given in Table 4 of Todd
(1966), rather than the USNM numbers assigned to individual slides in the type
collections. Attempts to locate the specimens used by Lessard (1980) were
unsuccessful. Accession numbers are also listed for those species for which DNA
sequences have been deposited in the GenBank Database (Holzmann et al. 2001).
Richardson and Clayshulte: Foraminifera of Guam
39
The 303 species of Foraminifera listed below were collected from various
shallow-water habitats around Guam, including: beach sands (Todd 1966,
Matsumaru & Matsuo 1976, Lessard 1980), sediments (Cushman 1917, 1932,
1933a, 1933b, 1934, 1942, Todd 1960, 1965, 1966), phytal substrata (Lessard
1980, S. Richardson, unpublished observations), and other solid substrata
(Clayshulte 1981b). The foraminiferal species identified in the beach sands and
sediment samples represent total (dead + live) foraminiferal populations, as no
attempt was made in any of the published studies to distinguish between the tests
of living foraminiferans and the empty tests of dead organisms. Field studies
conducted by Clayschulte (1981b) and S. Richardson (unpublished observations);
however, included observations of live individuals attached to settlement plates
and other natural substrata, such as reef rock and seagrasses.
Many of the foraminiferal species identified from Guam, such as
Baculogypsina sphaerulata (Parker & Jones, 1860) (Fig. 1), are distributed
widely throughout the Indo-Pacific (Todd 1960, Langer & Hottinger 2000,
Belasky 1996). Of the 303 species of Foraminifera identified from Guam, 129 are
also known from Saipan (Todd 1957). In addition, several of the Recent species
of large, endosymbiont-bearing foraminiferans living today in the shallow waters
off Guam are also known from the fossil record (Cole 1939, 1963, Hanzawa
1957, Matsumaru 1976, Todd 1957, 1966).
Currently, the most widely used classification of Foraminifera is that
published by Loeblich & Tappan (1987, 1992). In this scheme, Foraminifera is
assigned the Linnaean rank of a Class within the Phylum Granuloreticulosa.
More recently, molecular phylogenetic studies have challenged traditional ideas
on the position of Foraminifera in the tree of life, as well as hypotheses of the
evolutionary relationships within Foraminifera itself. While Foraminifera has
been consistently shown to comprise a monophyletic clade of single-celled
eukaryotes, the position of this clade within Eukaryota is still unresolved. In
molecular studies based on 18S ribosomal DNA sequences, Foraminifera appears
to be a deep-branching clade of basal mitochondrial eukaryotes, perhaps the
earliest extant group with mitochondria (Pawlowski et al. 1996; Pawlowski et al.
1997; Pawlowski et al. 1999 a, b; Pawlowski 2000); however, in a recent
molecular analysis based on actin gene sequences, Foraminifera emerges near the
eukaryotic crown as the sister group to cercomonads and chlorachniophytes
(photosynthetic amoebaflagellates) (Keeling 2001).
Molecular studies have also shed light on the taxonomic composition and
relationships of the major subclades within Foraminifera, both confirming and
invalidating earlier groupings. It is expected that future studies, both
morphological and molecular, will result in further rearrangements. For these
reasons, the checklist outlined below is organized in an untraditional manner.
40
Micronesica 35-36, 2003
Figure 1. Baculogypsina sphaerulata (Parker & Jones, 1860), a benthic foraminiferal species that
hosts intracellular diatom endosymbionts. All SEM images are of specimens found in beach
sand collected near the entrance to Lost Pond, Guam. a close-up view of test surface, radialpatterned projections represent surface expression of imperforate conical pillars; scale bar = 10
µm. b close-up view of ornamentation on spine surface; scale bar = 10 µm. c view of whole
test showing spines radiating from central body of chambers; scale bar = 100 µm. d view of
early coiled chambers in sectioned test; scale bar = 100 µm. e close-up view of peripheral
chambers in sectioned test showing perforated chamber walls and imperforate conical pillar;
scale bar = 10 µm.
Foraminifera and its inclusive subclades (as listed in Appendix 1 below), are
not assigned to traditional Linnaean ranks, but are given names based on the most
recent molecular phylogeny of Foraminifera (Pawlowski 2000), as is consistent
with the principles of phylogenetic taxonomy (de Queiroz & Gauthier 1990,
Richardson and Clayshulte: Foraminifera of Guam
41
1992, 1994). The Linnaean system of binomial nomenclature is retained,
however, for ease of information retrieval, and species are ordered alphabetically
by genus and species within each subclade.
Acknowledgments
We thank John J. Lee and an anonymous reviewer for their helpful
suggestions on revisions to the checklist. Susan Richardson would also like to
acknowledge Marty Buzas, Mark Florence, and Jennifer Jett for their help in
accessing the Cushman Collection of Foraminifera (Department of Paleobiology,
National Museum of Natural History, Smithsonian Institution, Washington, DC).
This is Contribution Number 532 of the Smithsonian Marine Station at Fort
Pierce, Florida.
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Cushman, J. A. 1942. The Foraminifera of the tropical Pacific collections of the
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de Queiroz, K. & J. Gauthier. 1990. Phylogeny as a central principle in
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307-322.
de Queiroz, K. & J. Gauthier. 1992. Phylogenetic taxonomy. Annual Review of
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de Queiroz, K. & J. Gauthier. 1994. Toward a phylogenetic system of
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Hanzawa, S. 1957. Cenozoic Foraminifera of Micronesia. Geological Survey of
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Holzmann, M., J. Hohenegger, P. Hallock, W. E. Piller & J. Pawlowski. 2001.
Molecular phylogeny of large miliolid foraminifera (Soritacea Ehrenberg
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Keeling, P. J. 2001. Foramnifera and Cercozoa are related in actin phylogeny:
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Received 10 Aug. 2001
44
Micronesica 35-36, 2003
Appendix 1. Checklist of foraminiferal species
Catalogue numbers with an USNM-prefix are given for type specimens (holo=holotype,
para(s)=paratypes(s)) of those species deposited in the Cushman Collection of Foraminifera,
Department of Paleobiology, National Museum of Natural History of the Smithsonian
Institution in Washington, DC. Accession numbers are listed for those species for which
sequences have been deposited in the GenBank Database. USNM a.s.: USNM assemblage
slide(s).
Refs: references: numbered references follow at the end of the Appendix
Notes: numbered notes follow at the end of the Appendix
Taxon
CLADE ASTRORHIZIDA+ALLOGROMIIDA
Bdelloidina aggregata Carter, 1877
Haddonia torresiensis Chapman, 1898
Halyphysema bayeri Loeblich, 1958
Halyphysema cf. H. harryi Loeblich, 1958
Hemidiscella cf. H. palabunda Bock, 1971
Iridia diaphana Heron-Allen & Earland, 1915
Saccammina sp. Todd, 1966
Sagenina frondescens (Brady, 1879)
Spiculidendron corallicolum Richardson & Rützler, 1996
CLADE MILIOLIDA
Ammomassilina alveoliniformis (Millett, 1898)
Amphisorus cf. A. kudakajimensis (Gudmundsson, 1994)
Articulina pacifica Cushman, 1944
Articulina sp. Todd, 1966
Biloculinella globula (Bornemann, 1855)
Borelis pulchrus (d'Orbigny, 1839)
Cornuspira planorbis Schultze, 1854
Cornuspiramia antillarum Cushman, 1922
Coscinospira cf. C. pertusus (Forsskål, 1775)
Dendritina ellipticus (d'Orbigny, 1904)
Flintina bradyana Cushman, 1921
Fischerina pellucida Millett, 1898
Hauerina diversa Cushman, 1946
Hauerina involuta Cushman, 1946
Hauerina milletti Cushman, 1954
Hauerina ornatissima (Karrer, 1868)
Hauerina pacifica Cushman, 1917
Massilina durrandi (Millett, 1898)
Massilina reticulata Heron-Allen & Earland, 1915
Massilina secans (d'Orbigny, 1826)
Massilina planata Cushman, 1932
Catalogue #
USNM a.s.
USNM a.s.
USNM a.s.
USNM 642328a
USNM a.s.
GenBank AJ404316
AJ278050
USNM a.s.
USNM 642321,
USNM a.s.
USNM 642315,
USNM a.s.
USNM a.s.
USNM CC14825
USNM 642328b
USNM a.s.
USNM 642308
USNM slide missing
USNM a.s.
USNM a.s.
USNM a.s.
USNM slide missing
USNM CC14971,
USNM CC46252
USNM a.s.
USNM a.s.
USNM a.s.
USNM CC14629
(holo); USNM
CC15031 (paras)
Refs.
Notes
9,12
9,12
14
14
14
14
9
9,12
14
9
15
16
9,12
9
9
9,12
2,9,11
12
9,14
11
9,12
9,12
2
9,12
9,12
9,12
2
2,9,12
9,12
9
9
2
17
18
19
20
Richardson and Clayshulte: Foraminifera of Guam
45
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Refs.
Miliolinella australis (Parr, 1932)
USNM a.s.
9,11
12
11
9,12
9,12
9,12
9
9
9,10
11,12
15
2,9,12
3,9,12
15
9,11
12
9
Miliolinella circularis Bornemann, 1855
Miliolinella labiosa (d'Orbigny, 1839)
Miliolinella oceanica (Cushman, 1932)
Miliolinella flintiana (Cushman, 1932)
Nubeculina advena Cushman, 1924
Nubeculina divaricata (Brady, 1879)
Orbitolites vertebralis (Quoy & Gaimard in de
Blainville, 1834)
Parrina bradyi (Millett, 1898)
Parasorites cf. P. carpenteri (Smout, 1963)
Planispirinella exigua (Brady, 1879)
Pseudomassilina reticulata Heron-Allen & Earland,
1915
Puteolina malayensis Hofker, 1951
USNM a.s.
USNM a.s.
USNM a.s.
USNM 642303
USNM a.s.
USNM a.s.,
GenBank AJ27805962
USNM CC15077
USNM CC19973,
GenBank AJ404305
USNM a.s.
USNM a.s.
Pyrgo denticulata (Brady, 1884)
USNM a.s.,GenBank
AJ404300
USNM CC15095
Pyrgo striolata (Brady, 1884)
Quinqueloculina agglutinans d'Orbigny, 1839
Quinqueloculina agglutinans (Wiesner in Heron-Allen &
Earland, 1915)
Quinqueloculina apicula Cushman, 1921
Quinqueloculina arenata Said, 1949
Quinqueloculina berthelotiana d'Orbigny, 1839
Quinqueloculina bidentata d'Orbigny, 1839
Quinqueloculina bosciana d'Orbigny, 1839
USNM a.s.
USNM a.s.
USNM CC14722,
USNM 642310
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM 642311
Quinqueloculina ferussacii d'Orbigny, 1826
Quinqueloculina lamarckiana d'Orbigny, 1839
Quinqueloculina parkeri (Brady, 1881)
USNM a.s.
USNM a.s.
USNM 13070,
USNM 13071,
USNM CC14735
Quinqueloculina plana d'Orbigny, 1839
Quinqueloculina poeyana d'Orbigny, 1839
Quinqueloculina polygona d'Orbigny, 1839
Quinqueloculina samoaensis Cushman, 1924
Quinqueloculina cf. Q. seminulum (Linné, 1758)
Quinqueloculina subarenaria Cushman, 1917
Quinqueloculina subcuneata Cushman, 1921
Quinqueloculina tubus Todd, 1957
Schlumbergerina alveoliniformis (Brady, 1879)
Siphonaperta distorqueata Cushman, 1954
Sorites cf. S. orbiculus (Forsskål, 1775)
USNM 642307
USNM a.s.
USNM CC14994
USNM a.s.
USNM a.s.
USNM 642301
USNM a.s.
USNM a.s.
USNM a.s.
GenBank AJ27805355
Notes
21
22
23
24
25
9,14*
26
2,9,11
12
9
9,12
2
27
9,12
9,12
9,12
9,12
9,11
12
9,12
9,12
1,2,9
11,12
11
9,12
9,12
2
9,12
9,12
9,12
9,12
9
9,12
10,14
15
28
29
30
46
Micronesica 35-36, 2003
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Refs.
Spirolina acicularis (Batsch, 1791)
USNM a.s.
Spirolina arietina (Batsch, 1791)
Spirolina okinawaensis Ujiéé & Hatta, 1994
Spiroloculina angulata Cushman, 1917
USNM a.s.
9,11
12
9,12
14
2,9,11
12
9,12
2,9
9,12
1
9,12
9
9,12
1
2
9,12
9
Spiroloculina caduca Cushman, 1922
Spiroloculina clara Cushman, 1932
Spiroloculina communis Cushman & Todd, 1944
Spiroloculina depressa d'Orbigny, 1826
Spiroloculina eximia Cushman, 1922
Spiroloculina folia Todd, 1957
Spiroloculina foveolata Egger, 1893
Spiroloculina grata Terquem, 1878
Spiroloculina grateloupi d'Orbigny, 1826
Spiroloculina hadai Thalmann, 1933
Spiroloculina marshallana Todd in Cushman, Todd &
Post, 1954
Spiroloculina rugosa Cushman & Todd, 1944
Spirophthalmidium acutimargo (Brady, 1884)
Triloculina affinis d'Orbigny, 1826
Triloculina cf. T. bassensis Parr, 1945
Triloculina cf. T. bicarinata d'Orbigny, 1839
Triloculina bikiniensis Todd in Cushman, Todd & Post,
1954
Triloculina earlandi Cushman, 1954
Triloculina fichteliana d'Orbigny, 1839
Triloculina incisura Todd, 1957
Triloculina involuta Todd, 1954
Triloculina irregularis (d'Orbigny, 1839)
Triloculina kerimbatica (Heron-Allen & Earland, 1915)
Triloculina marshallana Todd in Cushman, Todd &
Post, 1954
Triloculina oblonga (Montagu, 1803)
Triloculina oceanica Cushman, 1932
Triloculina spinata Cushman, 1932
Triloculina striolata Cushman, 1918
Triloculina subgranulata Cushman, 1918
Triloculina subplanciana Cushman, 1954
Triloculina terquemiana (Brady, 1884)
Triloculina transversestriata (Brady, 1881)
Triloculina tricarinata d'Orbigny, 1826
Triloculina trigonula (Lamarck, 1804)
Varidentella neostriatula (Thalmann, 1950)
Vertebralina striata d'Orbigny, 1826
CLADE SPIRILLINIDA
Conicospirillina semiinvoluta Cushman, 1933
USNM CC14937,
USNM a.s.
USNM a.s.
USNM slide missing
USNM a.s.
USNM I388044
USNM a.s.
USNM a.s.
USNM a.s.
USNM I388945
USNM slide missing
USNM a.s.
USNM a.s.
USNM a.s.
USNM I388032
USNM CC14876
USNM a.s.
USNM a.s.
9,12
1
2
9
9,12
11
USNM a.s.
USNM CC14618
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
9,12
2
9,12
9
9,12
9,11
12
9,12
USNM a.s.
USNM CC14924
USNM a.s.
USNM 642304
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM CC14964
USNM a.s.
2,9,11
12
11
9
11
9,12
9,12
9,12
9,12
9,12
9,12
9,12
2,9,12
9
Notes
31
32
33
Richardson and Clayshulte: Foraminifera of Guam
47
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Conicospirillina trochoidea Cushman, 1933
USNM CC19284
(holo)
USNM a.s.
USNM CC17552
USNM CC17548,
USNM a.s.
USNM 517045
USNM a.s.
USNM a.s.
USNM 517050
USNM 517049
USNM a.s.
Conicospirillina sp.
Patellina advena Cushman, 1922
Patellina altiformis Cushman, 1933
Patellinella inconspicua (Brady, 1884)
Spirillina decorata Brady, 1884
Spirillina denticulogranulata Chapman, 1907
Spirillina revertens Rhumbler, 1906
Spirillina spinigera Chapman, 1899
Spirillina vivipara Ehrenberg, 1843
CLADE NODOSARIIDA
Amphicoryna scalaris (Batsch, 1791)
Astacolus bradyi (Cushman, 1917)
Astacolus aff. A. californicus Galloway & Wissler, 1927
Astacolus planulatus Galloway & Wissler, 1927
Darbyella sp.
Dentalina mucronata Neugeboren, 1856
Fissurina marginata (Montagu, 1803)
Frondicularia repanda Cushman, 1913
USNM a.s.
USNM 642320,
USNM a.s.
USNM a.s.
USNM 642317,
USNM a.s.
USNM a.s.
USNM a.s.
USNM 642329,
USNM a.s.
Lagena desmophora Rymer-Jones, 1872
USNM 15725
Lagena marginato-perforata Seguenza, 1880
USNM 19837
Lagena spiralis Brady, 1884
USNM 15727
Lenticulina orbicularis (d'Orbigny, 1902)
USNM 642316,
USNM a.s.
Lenticulina vortex (Fichtel & Moll, 1798)
USNM a.s.
Nodosaria calomorpha Reuss, 1866
USNM 642323,
USNM a.s.
CLADE GLOBIGERINIDA+ROTALIIDA+TEXTULARIIDA,
SUBCLADE GLOBIGERINIDA
Candeina nitida d'Orbigny, 1839
USNM 642286,
USNM a.s.
Globigerinella aequilateralis (Brady, 1884)
USNM a.s.
Globigerinella adamsi (Banner & Blow, 1959)
USNM slide missing
Globigerinita glutinata (Egger, 1893)
USNM 517040,
USNM a.s.
Globigerinoides conglobatus (Brady, 1879)
Globigerinoides sacculifer (Brady, 1884)
USNM 517041,
USNM a.s.
Globorotalia hirsuta (d'Orbigny, 1839)
USNM a.s.
Globorotalia menardii (Parker, Jones & Brady, 1865)
USNM a.s.
Globorotalia tumida (Brady, 1877)
USNM a.s.
Globorotalia ungulata Bermudez, 1960
USNM slide missing
Neogloboquadrina dutertrei (d'Orbigny, 1839)
USNM a.s.
Orbulina universa d'Orbigny, 1839
USNM a.s.
Refs.
Notes
4
8
8
8,9
8
9
9,12
8,9,12
8
8,9,12
9
9
34
35
36
9
9
9
9
11
9
37
3
3
3
9
38
9
9
9
8
8
8,9
11
8,9,11
9
9
9
8
9
9
39
40
41
48
Micronesica 35-36, 2003
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Pulleniatina obliquiloculata (Parker & Jones, 1865)
USNM 642285,
USNM a.s.
Sphaeroidinella dehiscens (Parker & Jones, 1865)
USNM a.s.
CLADE GLOBIGERINIDA+ROTALIIDA+TEXTULARIIDA,
SUBCLADE ROTALIIDA
Acervulina inhaerans Schultze, 1856
USNM 517032
Acervulina sp.
USNM a.s.
Alliatina translucens (Cushman, 1933)
USNM a.s.
Ammonia beccarii (Linné, 1758)
USNM a.s.
Ammonia beccarii (Linné, 1758) var. Todd, 1957
Ammonia tepida (Cushman, 1926)
USNM 642306
USNM a.s.
Amphistegina lessonii d'Orbigny, 1826
USNM a.s.
Amphistegina lobifera Larsen, 1976
Amphistegina madagascarensis d'Orbigny, 1826
Amphistegina papillosa Said, 1949
Angulogerina albatrossi Cushman, 1932
Anomalina coronata Parker & Jones, 1857
Anomalina glabrata Cushman, 1924
Anomalina maculosa Todd, 1957
Baculogypsina sphaerulata (Parker & Jones, 1860)
Baggina indica (Cushman, 1921)
Baggina totomiensis Makiyami, 1931
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM 517033
USNM a.s.
Biarritzina proteiformis (Goes, 1882)
Bolivina compacta Sidebottom, 1905
Bolivina hantkeniana Brady, 1881
USNM 642332,
USNM a.s.
USNM a.s.
Bolivina pseudopygmaea Cushman, 1933
Bolivina rhomboidalis (Millett, 1899)
USNM CC17412
USNM 642330,
USNM a.s.
USNM a.s.
USNM a.s.
Bolivina spinescens Cushman, 1911
Bolivina robusta Brady, 1881
Bolivina tortuosa Brady, 1881
USNM a.s.
USNM a.s.
USNM a.s.
Bolivina vadescens Cushman, 1933
Bolivinella folia (Parker & Jones, 1865)
Brizalina striatula Cushman, 1922
USNM 17437
USNM a.s.
USNM a.s.
Bronnimannia haliotis (Heron-Allen & Earland, 1924)
Bulimina costata d'Orbigny, 1852
Buliminella milletti Cushman, 1933
USNM 642314,
USNM a.s.
USNM a.s.
USNM CC17324
Buliminoides williamsoniana (Brady, 1884)
Calcarina hispida Brady, 1884
USNM a.s.
USNM 64944
Refs.
Notes
9
9
8,9-13
9
9
9,11
12
9,11
8,9,11
12
9,11
12
9,12
9
9
9
9
9,12
8,9,12
7,9-12
11
9
9,12
13
6,9,12
9,12
9,12
9,11
12
9
9,12
9,11
12
4
9
9,11
12
9
9
4,6,9
11,12
9,12
8
42
43
44
45
46
47
48
Richardson and Clayshulte: Foraminifera of Guam
49
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Calcarina spengleri (Gmelin, 1791)
USNM a.s.
Cancris auriculus (Fichtel & Moll, 1798)
Cancris sagra (d'Orbigny, 1839)
USNM 517034
USNM 642309,
USNM a.s.
Carpenteria monticularis (Carter, 1877)
Carpenteria utricularis (Carter, 1876)
Cassidulina delicata Cushman, 1927
Cassidulina minuta Cushman, 1933
Chrysalidinella earlandi Cushman, 1945
Cibicides cicatricosus (Schwager, 1866)
Cibicides floridanus (Cushman, 1918)
Cibicides lobatulus (Walker & Jacob, 1798)
Cibicides mayori (Cushman, 1924)
Cibicides pseudoungerianus (Cushman, 1922)
Cibicidella variabilis (d'Orbigny, 1826)
Cycloclypeus carpenteri Brady, 1881
Cymbaloporella tabellaeformis (Brady, 1884)
Cymbaloporetta bradyi (Cushman, 1915)
Cymbaloporetta squammosa (d'Orbigny, 1839)
Discorbis orbicularis (d'Orbigny in Fornasini, 1906)
Discorbis nipponica Husezima & Maruhasi, 1944
Elphidium advenum (Cushman, 1922)
Elphidium craticulatum (Fichtel & Moll, 1798)
Elphidium crispum (Linné, 1758)
Elphidium hyalocostatum Todd, 1957
Elphidium jenseni (Cushman, 1924)
Elphidium sagrum (d'Orbigny, 1839)
Elphidium simplex Cushman, 1933
Elphidium striatopunctatum (Fichtel & Moll, 1798)
Epistominella pulchra (Cushman, 1933)
Epistominella tubulifera (Heron-Allen & Earland, 1915)
Epistomaroides polystomelloides (Parker & Jones, 1865)
Epistomaroides rimosa (Parker & Jones in Carpenter,
1862)
Epistominella cf. E. obtusa (Burrows & Holland in
Jones, 1896)
Epistominella rugosa (Phleger & Parker, 1951)
Eponides orientalis Asano, 1937
Eponides repandus (Fichtel & Moll, 1798)
Fontbotia wuellerstorfi (Schwager, 1866)
USNM 642326,
USNM a.s.
USNM 642327,
USNM a.s.
USNM a.s.
USNM a.s.
USNM 517037
USNM 517038
USNM 64948
USNM a.s.
USNM 517035
USNM 642328,
USNM a.s.
USNM 64946
USNM 47270,
USNM CC47272
USNM a.s.
USNM a.s.
USNM 19892
USNM a.s.
USNM CC19896
USNM a.s.
USNM a.s.
USNM 517039
USNM 64945,
USNM a.s.
USNM a.s.
Refs.
12,13
12,13
9
9,11
9
9,12
8
8,9,11
12
8,9,12
9
8,9,12
9
8,9,12
8,9,11
12
9,11
12
11
11
9,12
11
3,9,12
9,12
3,9,12
11
9,12
9,12
8
8,9,12
9,12
11
USNM a.s.
9
USNM a.s.
9
11
8,9,12
USNM CC47320,
USNM 642312,
USNM a.s.
USNM 517047
Notes
9,1012
8
9,12
8
49
50
Micronesica 35-36, 2003
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Gavelinopsis praegeri (Heron-Allen & Earland, 1913)
Glabratella orientalis (Cushman, 1925)
Glabratella patelliformis (Brady, 1884)
USNM a.s.
USNM a.s.
Glabratella tabernacularis (Brady, 1881)
Glandulina laevigata d'Orbigny, 1826
Globocassidulina subglobosa (Brady, 1881)
Gypsina vesicularis (Parker & Jones, 1860)
USNM a.s.
USNM CC15731
USNM a.s.
USNM a.s.
Hanzawaia concentrica (Cushman, 1918)
Heronallenia lingulata (Burrows & Holland in Jones,
1896)
Heterostegina depressa d'Orbigny, 1826
Heterostegina suborbicularis d'Orbigny, 1904
Hoeglundina elegans (d'Orbigny, 1826)
Homotrema rubra (Lamarck, 1816)
USNM 517043
USNM 641043
Loxostomum costulatum (Cushman, 1922)
Loxostomum limbatum (Brady, 1884)
Loxostomum mayori (Cushman, 1922)
Loxostomum porrectum (Brady, 1881)
Miniacina miniacea (Pallas, 1766)
Neoconorbina crustrata (Cushman, 1933)
Neoconorbina fulva (Todd, 1957)
Neoconorbina terquemi (Rzehak, 1888)
Neoconorbina tuberocapitata (Chapman, 1899)
Nodogenerina milletti Cushman, 1933
Nonion boueanum (d'Orbigny, 1846)
Nonion japonicum Asano, 1938
Nonion pacificum (Cushman, 1927)
Nonionella translucens Cushman, 1933
Operculina granulosa Leymerie, 1846
Oridorsalis umbonatus (Reuss, 1851)
Paumotua terebra (Cushman, 1933)
Pararotalia calcar (d'Orbigny, 1826)
Pegidia dubia (d'Orbigny in Fornasini, 1908)
USNM CC15684
USNM a.s.
USNM a.s.
USNM a.s.
USNM CC38414
USNM a.s.
USNM 17380
USNM a.s.
USNM 642324,
USNM a.s.
USNM a.s.
USNM a.s.
USNM CC47234
USNM CC19017
(holo), USNM
CC19018 (paras)
USNM a.s.
USNM a.s.
USNM a.s.
USNM CC19870
USNM CC15685
USNM specimen
missing
USNM a.s.
Planispirinella exigua (Brady, 1879)
Planodiscorbis rarescens (Brady, 1885)
Planorbulina acervalis Brady, 1885
USNM 642319,
USNM a.s.
USNM a.s.
USNM a.s.
USNM 641049
Planorbulinella larvata (Parker & Jones, 1865)
Planorbulinoides retinaculatus (Parker & Jones, 1862)
Planogypsina squamiformis (Chapman, 1901)
USNM 517046
USNM a.s.
USNM a.s.
Rectobolivina striatula (Cushman, 1937)
USNM a.s.
Refs.
9
11
9,11
12
9,12
3
9
9,1113
8
8
3,11
9,12
9
9,12
13
6,9
9
6,9
9
12,13
9
Notes
50
51
52
54
53
55
56
9,12
8,9,12
8,9
4,6
9,12
9,12
9
3
3
8
9
11
9,12
9
9
8,9,11
12
8,9,12
9,12
9,12
13
9
57
58
Richardson and Clayshulte: Foraminifera of Guam
51
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Reusella pulchra Cushman, 1945
USNM 642331,
USNM a.s.
USNM a.s.
Reusella simplex (Cushman, 1929)
Reussella spinulosa (Reuss, 1850)
Rosalina cf. R floridana (Cushman, 1922)
Rosalina micens (Cushman, 1933)
Rosalina opima (Cushman, 1933)
Rosalina rugosa d'Orbigny, 1839
Rosalina subbertholoti (Cushman, 1924)
Siphogenerina raphana (Parker & Jones, 1865)
USNM a.s.
USNM 517031
USNM CC17482
Siphonina tubulosa Cushman, 1924
Siphoninoides echinatus (Brady, 1897)
Sphaeridia papillata Heron-Allen & Earland, 1923
Sphaerogypsina globulus (Reuss, 1848)
USNM a.s.
USNM 517048
USNM a.s.
USNM 517042
Sphaeroidina bulloides d'Orbigny, 1826
Sporadotrema aff. S. cylindrica (Carter, 1880)
Sporadotrema rubra (d'Orbigny in Fornasini, 1908)
Stomatorbina concentrica (Parker & Jones in Brady,
1884)
Svratkina australiensis (Chapman, Parr & Collins, 1956)
Tretomphalus clarus Cushman, 1934
USNM a.s.
Tretomphalus concinnus (Brady, 1884)
Tretomphalus grandis Cushman, 1934
Tretomphalus milletti (Heron-Allen & Earland, 1915)
Tretomphalus planus Cushman, 1924
Trifarina bradyi Cushman, 1923
Trochulina mira (Cushman, 1922)
USNM CC17467
USNM CC47250
USNM CC47271
USNM 642325,
USNM a.s.
USNM a.s.
USNM CC21562
(holo)
USNM 21566,
USNM a.s.
USNM 51730
USNM 641047
USNM a.s.
USNM 642313,
USNM a.s.
Uvigerina ampullacea Brady, 1881
USNM a.s.
Uvigerina porrecta Brady, 1879
USNM a.s.
Uvigerina vadescens Cushman, 1933
USNM CC17515
(holo)
CLADE GLOBIGERINIDA+ROTALIIDA+TEXTULARIIDA
SUBCLADE TEXTULARIIDA
Carterina spiculotesta (Carter, 1879)
USNM a.s.
Clavulina difformis Brady, 1884
USNM 14708,
USNM 14709,
USNM a.s.
Clavulina multicamerata Chapman, 1907
Gaudryina angulata Cushman, 1924
USNM a.s.
Gaudryina rugulosa Cushman, 1932
USNM 14701,
USNM a.s.
Gaudryina siphonifera (Brady, 1884)
USNM a.s.
Gaudryina transversaria (Brady, 1884)
USNM a.s.
Refs.
Notes
9
9,11
12
6
8,9,12
8,9
11
9,12
8
6,9,11
12
9
8,9,12
9,12
8,9,12
13
9
12,13
12,13
9
59
60
7,9
5
8,9,11
12
11
8
8,9,12
9
9,11
12
9
9
4,9
61
62
63
9
2,9,12
11
9,12
2,9,11
12
9,12
9
64
52
Micronesica 35-36, 2003
Appendix 1. Checklist of foraminiferal species / (continued)
Taxon
Catalogue #
Gaudryina trullissata Todd in Cushman, Todd & Post,
1954
Liebusella bradyi (Cushman, 1911)
Rotaliammina mayori Cushman, 1924
Textularia agglutinans d'Orbigny, 1839
Textularia alveata Todd, 1957
Textularia candeiana d'Orbigny, 1839
Textularia conica d'Orbigny, 1839
USNM a.s.
Refs.
9,12
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
Textularia corrugata Heron-Allen and Earland, 1915
Textularia dupla Todd in Cushman, Todd & Post, 1954
Textularia foliacea Heron-Allen and Earland, 1915
Textularia kerimbaensis Said, 1949
Textularia oceanica Cushman, 1932
Textularia semialata Cushman, 1913
Valvulina davidiana Chapman, 1900
USNM CC14666
USNM a.s.
USNM 14658
USNM a.s.
USNM a.s.
USNM a.s.
USNM a.s.
9
9
9,12
9,12
11
9,11
12
2
9,12
2,9,12
9,12
9
9,12
9,12
Notes
65
66
References: 1. Cushman 1917; 2. Cushman 1932; 3. Cushman 1933a; 4. Cushman 1933b; 5.
Cushman 1934; 6. Cushman 1942; 7. Todd 1960; 8. Todd 1965; 9. Todd 1966; 10. Matsumaru
& Matsuo 1976; 11. Lessard 1980; 12. Clayschulte 1981a; 13. Clayschulte 1981b; 14. S.
Richardson, unpublished observations; 15. Holzmann et al. 2001
Notes: 16. Synonym: Marginopora cf. M. kudakajimensis (Gudmundsson, 1994) (Ref. 15) 17.
Synonym: Cornuspira involvens Reuss, 1850 (Ref. 2) 18. Synonym: Placopsilina? sp. (Ref. 9)
19. Synonym: Peneroplis pertusus (Forsskål, 1775) (Ref. 11) 20. Synonym: Peneroplis
ellipticus (d'Orbigny, 1904) (Refs. 9, 12) 21. Synonym: Miliolinella oceanica var. flintiana
(Cushman, 1932) (Refs. 9, 12) 22. Synonym: Nubeculina divaricata advena Cushman, 1924
(Ref. 9) 23. Synonym: Marginopora vertebralis (Quoy and Gaimard in de Blainville, 1834)
(Refs. 9-12, 15) 24. Synonyms: Sorites marginalis (Lamarck, 1801) (Refs. 3, 9, 12);
Parasorites sp. (Ref. 15) 25. Synonym: Massilina secans reticulata Heron-Allen & Earland,
1915 (Ref. 9) 26. Synonyms: Peneroplis proteus d'Orbigny, 1839 (Ref. 9); Laevipeneroplis sp.
(Ref. 15) 27. Synonym: Quinqueloculina anguina Terquem var. agglutinans (Wiesner in
Heron-Allen & Earland, 1915) (Ref. 2) 28. Synonym: Quinqueloculina anguina var. arenata
Said, 1949 (Refs. 9, 12) 29. Synonym: Quinqueloculina distorqueata Cushman, 1954 (Refs. 9,
12) 30. Synonym: Sorites marginalis (Lamarck, 1801) (Ref. 10). J. J. Lee (personal
communication, December 2001) believes this is a new species. It may also be conspecific
with a new unnamed species from Pohnpei, Micronesia identified by the senior author as New
clade I n. sp. in Richardson (2001). 31. Synonym: Spiroloculina grata var. angulata Cushman,
1917 (Ref. 2) 32. Synonym: Spiroloculina acutimarginata Brady, 1884 (Ref. 1)33. Synonym:
Quinqueloculina neostriatula Thalmann, 1950 (Refs. 9, 12) 34. Synonym: Patellina advena
altiformis Cushman, 1933 (Refs. 8, 9) 35. Synonym: Spirillina vivipara revertens Rhumbler,
1906 (Refs. 8, 9, 12) 36. Synonym: Lagenonodosaria scalaris (Batsch, 1791) (Ref. 9) 37.
Synonym: Frondicularia robusta var. repanda Cushman, 1913 (Ref. 9) 38. Synonym: Robulus
orbicularis (d'Orbigny, 1902) (Ref. 9) 39. Synonym: Globigerinoides triloba (Reuss, 1850)
(Ref. 11) 40. Synonym: Globorotalia menardii ungulata Bermudez, 1960 (Ref. 8) 41.
Synonym: Globigerina eggeri Rhumbler, 1911 (Ref. 9) 42. Synonym: Streblus beccarii
(Linné, 1758) (Refs. 9, 12) 43. Synonym: Streblus beccarii (Linné, 1758) var. Todd, 1957
(Refs. 9, 12) 44. Synonym: Streblus beccarii var. tepida (Cushman, 1926) (Refs. 9, 12) 45.
Synonym: Amphistegina radiata var. papillosa Said, 1949 (Ref. 9) 46. Synonym:
Amphistegina radiata (Fichtel & Moll, 1798) 47. Synonym: Carpenteria proteiformis Goes,
Richardson and Clayshulte: Foraminifera of Guam
53
1882 (Refs. 9, 12, 13) 48. Synonym: Bolivina striatula Cushman, 1922 (Refs. 9, 12, 13) 49.
Synonym: Planulina wuellerstorfi (Schwager, 1866) (Ref. 8) 50. Synonym: Discorbis
orientalis Cushman, 1925 (Ref. 11) 51. Synonym: Neoconorbina patelliformis (Brady, 1881)
(Refs. 9, 12); Pileolina (?) patelliformis (Brady, 1884) (Ref. 11) 52. Synonym: Neoconorbina
tabernacularis (Brady, 1881) (Refs. 9, 12) 53. Synonym: Bolivina (Loxostomum) limbata
Brady, 1884 (Ref. 9) 54. Synonym: Bolivina (Loxostomum) limbata var. costulata (Cushman,
1922) (Refs. 6, 9) 55. Synonym: Bolivina (Loxostomum) mayori Cushman, 1922 (Refs. 6, 9)
56. Synonym: Bolivina (Loxostomum) porrecta Brady, 1881 (Ref. 9) 57. Synonym: Tinoporus
calcar (d'Orbigny, 1826) (Ref. 11) 58. Synonym: Rectobolivina bifrons var. striatula
(Cushman, 1937) (Ref. 9) 59. Synonym: Discorbis opima Cushman, 1933 (Ref. 11) 60.
Synonym: Gypsina globula (Reuss, 1848) (Ref. 8, 9, 13) 61. Synonym: Rosalina concinna
Brady, 1884 (Refs. 9, 12); Tretomphalus bulloides (d'Orbigny, 1839) (Ref. 11) 62. Synonyms:
Rotorbinella? mira (Cushman, 1922) (Refs. 9, 12); Discorbis mirus (Cushman, 1922) (Ref.
11) 63. Synonym: Uvigerina proboscidea Schwager var. vadescens Cushman, 1933 (Refs. 4,
9) 64. Synonym: Gaudryina triangularis var. angulata Cushman, 1924 (Refs. 9, 12) 65.
Synonym: Clavulina bradyi Cushman, 1911 (Ref. 9) 66. Synonym: Textularia foliacea var.
oceanica Cushman, 1932 (Ref. 9)