[Palaeontology, Vol. 52, Part 2, 2009, pp. 401–427]
RE-ASSESSMENT OF THE TYPE COLLECTIONS OF
FOURTEEN CORALLINALEAN SPECIES
(CORALLINALES, RHODOPHYTA) DESCRIBED
BY W. ISHIJIMA (1942–1960)
by YASUFUMI IRYU*§, DAVIDE BASSI and WILLIAM J. WOELKERLINGà
*Institute of Geology and Paleontology, Graduate School of Science, Tohoku University, Aobayama, Sendai 980-8578, Japan;
e-mail: [email protected]
Dipartimento di Scienze della Terra, Università degli Studi di Ferrara, via Saragat 1, I-44100 Ferrara, Italy; e-mail: [email protected]
àSchool of Botany, La Trobe University, Bundoora, Victoria, Australia 3083; e-mail: [email protected]
§Present address: Department of Earth and Planetary Sciences, Graduate School of Environmental Studies, Nagoya University, Nagoya 464-8601, Japan
Typescript received 5 August 2007; accepted in revised form 23 May 2008
Abstract: Many species of fossil coralline algae have been
separated on the basis of only a few characters with slight or
doubtful taxonomic significance. Analysis of the type material of the fossil coralline algal species is, therefore, needed in
order to assess the taxonomic status and circumscription of
these taxa. Wataru Ishijima has been one of the most prolific
Japanese palaeontologists who worked on fossil calcareous
algae from the western Pacific region. Ishijima described a
large number of new taxa of fossil calcareous algae mostly
belonging to the Corallinales from Eocene to Pleistocene sedimentary successions. An analysis of some of Ishijima’s types
of Corallinales (Rhodophyta) described from 1942 to 1960,
currently housed in the Institute of Geology and Paleontology, Graduate School of Science, Tohoku University, Sendai
(Japan), is presented. For fourteen taxa, changes were made
to previously published statements of typification. Re-examination of the types from a modern perspective of coralline
algal taxonomy reveals that Hydrolithon taishakuensis (Ishijima) comb. nov (= Lithothamnium taishakuensis), Sporolithon
kobamazimensis (Ishijima) comb. nov. (= Archaeolithothamnium kobamazimensis), Sporolithon kuboiensis (Ishijima)
comb. nov. (= Archaeolithothamnium kuboiensis) and Sporoli-
T he taxonomy of fossil coralline red algae (Corallinales)
has undergone marked changes in recent years as a consequence of a number of major studies involving presentday counterparts. The group is now treated as a separate
order of Rhodophyta (Silva and Johansen 1986), and is
considered to represent a major evolutionary line within
the red algae (Bailey and Chapman 1996). Concepts of
families, subfamilies, genera and species have changed as
a result of extensive new information regarding vegetative
morphology, anatomy and reproduction (e.g. Woelkerling
ª The Palaeontological Association
thon taiwanensis (Ishijima) comb. nov. (= Archaeolithothamnium taiwanensis) showed significant features justifying their
use as species names. No diagnostic characters occur in two
types (L. nishiwadai, L. otsukiensis) and therefore the taxonomic disposition of these taxa remains unknown. Lithophyllum quadratum Ishijima is considered as a species of
uncertain status within Lithophyllum pending further study.
Lithoporella crassa Ishijima and Lithoporella hayasakai Ishijima are considered heterotypic synonyms of Lithophyllum
prototypum (Foslie) Foslie, while Porolithon hanzawai Ishijima is a heterotypic synonym of Hydrolithon onkodes (Heydrich) Penrose and Woelkerling. Mesophyllum arakuraensis
Ishijima is co-specific to, and has nomenclatural priority
over, M. contii Ishijima. However, M. arakuraensis and
M. contii do not show evidence whether either might belong
to Synarthrophyton instead of Mesophyllum. Lithothamnion
ishizuchiensis Ishijima and L. kasedaensis Ishijima have no
diagnostic features that can be used to ally them to any
particular genus or species of Melobesioideae.
Key words: Ishijima’s type collections, Corallinales, Rhodo-
phyta, Japan, Taiwan.
1988, 1996a, b, c; Woelkerling and Lamy 1998; Harvey
et al. 2003a). New genera have been described and some
genera commonly used in the older literature have been
relegated to synonymy.
These developments have led to a significant, ongoing
taxonomic re-evaluation of fossil coralline algae. A number
of studies have aimed to re-assess various 19th and 20th
century fossil type specimens in the context of our current
understanding of coralline systematics (e.g. Braga et al.
1993; Rasser and Piller 1999; Braga 2003). As a result, many
doi: 10.1111/j.1475-4983.2009.00850.x
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PALAEONTOLOGY, VOLUME 52
taxa have been transferred to different genera and ⁄ or subfamilies and ⁄ or families, while other taxa have found to be
of uncertain taxonomic affinities (e.g. Piller 1994; Rasser
and Piller 1994; Braga and Aguirre 1995; Aguirre et al.
1996; Aguirre and Braga 1998; Basso et al. 1997, 1998; Bassi
et al. 2000, 2005; Bassi 2003; Braga et al. 2005).
Moreover, formal recognition of hundreds of fossil species is questionable because they are based only on cell
sizes in small fragments of thallus or on other characters
of uncertain taxonomic significance (e.g. Braga and Aguirre 1995; Bassi et al. 2000, 2005). Vague and inconsistent
species circumscriptions have made it very difficult to
interpret many fossil species, and most such species have
been cited only by the describing author (Aguirre and
Braga 2005).
Wataru Ishijima (1906–1980; http://en.wikipedia.org/
wiki/Ishijima_Wataru) was one of the most prolific
researchers of fossil calcareous algae. After graduating from
the Imperial Fisheries Institute (Tokyo; current Tokyo
University of Marine Science and Technology) in 1927,
Ishijima joined the Institute of Geology and Paleontology,
Faculty of Science, Tohoku Imperial University (Sendai)
from 1927 to 1931. He then worked at the Institute of
Geology, Taihoku Imperial University (Taipei) during
1942–1945 and then at the St. Paul’s University (Rikkyo
University, Tokyo) from1945 to 1980. His doctoral dissertation was submitted to Tohoku University and was privately
published by Y
uhod
o (Ishijima 1954). He described a total
of 139 taxa of fossil calcareous algae (Iryu 2004) including
at least 114 species of Corallinales, and he produced more
than 45 publications on coralline algal taxonomy.
Ishijima described taxa from Eocene to Pleistocene
sedimentary successions occurring in the western Pacific.
Most of Ishijima’s papers were published during the
1940s and 1960s, the main period of productivity in fossil
coralline taxonomy, which corresponds to the peak introduction of new names (Aguirre and Braga 2005).
Ishijima deposited portions of his material in four different institutions: the Institute of Geology and Paleontology,
Faculty of Science (current Graduate School of Science),
Tohoku University; the Faculty of Science, St. Paul’s
University (Rikkyo University); the Institute of Geology,
Taihoku Imperial University; and the Geological Institute
of the Tokyo University of Education (current Institute of
Geoscience, Tsukuba University). The collection at the Faculty of Science, St. Paul’s University is now deposited in
the Institute of Geology and Paleontology, Graduate
School of Science, Tohoku University. The assessment of
these collections is in progress by the authors.
The aim of this study is to re-assess those 14 Ishijima
species of Corallinales for which original material is
retained in the Institute of Geology and Paleontology,
Graduate School of Science, Tohoku University. Our reassessment was focused on determining the taxonomic
placement of this material in a modern context, and the
relationships of Ishijima’s taxa to other described
species.
MATERIAL AND METHODS
Data were obtained from the examination of thin sections
of Ishijima’s original material using light microscopy.
Anatomical and morphological terms follow Woelkerling
(1988), Braga et al. (1993), Woelkerling et al. (1993),
Harvey et al. (2002) and Braga (2003).
For some species, Ishijima mentioned an ‘IGPS coll.
cat. no.’ under the heading ‘Repository’ in his publications. These Repository numbers are listed in relevant
species accounts below. According to Ishijima (1954, p.
20, footnote) the acronym IGPS is an abbreviation for the
Institute of Geology and Paleontology, Tohoku University. The Institute has a catalogue and register cards in
which collections were recorded and given catalogue
numbers. Ishijima, however, did not record these numbers on his slides, and it is uncertain what the numbers
present on some slides mean.
Herbarium abbreviations are those used in Index Herbarium, formerly in print (Holmgren et al. 1990) and
more recently online electronically [Holmgren and Holmgren 1998 (continuously updated)]. The International
Code of Botanical Nomenclature (ICBN) cited is the
Vienna edition (McNeill et al. 2006). In accordance with
ICBN Art. 61 (also see Rec. 50F.1 under Art. 50), published orthographic variants of taxa have been corrected
to the validly published form of the name. Original incorrect variants are given in parentheses, as for example
Archaeolithothamnion kobamazimensis Ishijima, 1942b (as
Archaeolithothamnium), which in this case indicates that
Ishijima used the incorrect variant Archaeolithothamnium
in his 1942b paper.
SYSTEMATIC PALAEONTOLOGY
The taxonomic disposition of the 14 Ishijima taxa dealt with
below is summarized in Table 1 using the classification scheme
proposed by Harvey et al. (2003a). Current concepts have been
used in assigning or re-assigning Ishijima taxa to genera; brief
remarks are included for each genus. Ten species are known
only from the fossil record, while four described as distinct
fossil species by Ishijima have been found to be heterotypic
synonyms of currently living species of Corallinales. As a result,
the fossil species Lithophyllum quadratum is considered as a
species of uncertain status within Lithophyllum pending further
study, the fossil species Lithoporella crassa Ishijima and Lithoporella hayasakai Ishijima are treated here as synonyms of the living species Lithophyllum prototypum, and the fossil species
Porolithon hanzawai is treated here as a synonym of the living
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
TABLE 1.
403
Proposed taxonomic disposition of Ishijima’s species.
Ishijima’s placement
Family Corallinaceae
Subfamily Melobesiae
Lithophyllum quadratum Ishijima, 1954
Lithoporella crassa Ishijima, 1942a
Lithoporella hayasakai Ishijima, 1942a
Porolithon hanzawai Ishijima, 1954
Lithothamnium taishakuensis Ishijima, 1960a
Mesophyllum arakuraensis Ishijima, 1954
Mesophyllum Contii Ishijima, 1954
Lithothamnium ishizuchiensis Ishijima, 1960b
Lithothamnium kasedaensis Ishijima, 1954
Archaeolithothamnium kobamazimensis Ishijima, 1942b
Archaeolithothamnium kuboiensis Ishijima, 1942b
Archaeolithothamnium taiwanensis Ishijima, 1942c
Lithophyllum Nishiwadai Ishijima, 1944
Lithophyllum otsukiensis Ishijima, 1954
This paper
Order CORALLINALES Silva and Johansen, 1986
Family CORALLINACEAE Lamouroux, 1812
Subfamily LITHOPHYLLOIDEAE Setchell, 1943
Lithophyllum quadratum Ishijima
Lithophyllum prototypum (Foslie) Foslie, 1905
Subfamily MASTOPHOROIDEAE Setchell, 1943
Hydrolithon onkodes (Heydrich) Penrose and Woelkerling, 1992
Hydrolithon taishakuensis (Ishijima) comb. nov.
Family HAPALIDIACEAE Gray, 1864
Subfamily MELOBESIOIDEAE Bizzozero, 1885
Mesophyllum arakuraensis Ishijima
Uncertain generic placement
Uncertain generic placement
Family SPOROLITHACEAE Verheij, 1993
Sporolithon kobamazimensis (Ishijima) comb. nov.
Sporolithon kuboiensis (Ishijima) comb. nov.
Sporolithon taiwanensis (Ishijima) comb. nov.
Family uncertain
Uncertain generic placement
Uncertain generic placement
Classification system in left-hand column is that used by Ishijima and in common use during the period of Ishijima’s studies. In that
scheme all corallines were placed in a single family (the Corallinaceae) and all species lacking genicula were referred to a single subfamily (the Melobesioideae). Classification system in right-hand column follows Harvey et al. (2003a) and is used in the present study.
species Hydrolithon onkodes. For species of uncertain genus or
uncertain family placement, Ishijima’s original names (basionyms) are used.
Order CORALLINALES Silva and Johansen, 1986
Remarks. Prior to 1993, all coralline red algae were
referred to a single family, the Corallinaceae. Recent morphological and molecular studies, however, have provided
evidence (summarized in Harvey et al. 2003a) that three
distinct lineages involving extant taxa occur, each representing a separate family (i.e. the Corallinaceae, Sporolithaceae and Hapalidiaceae). Additional remarks appear
under each family name.
Molecular data are lacking for the exclusively fossil Graticulaceae, established by Brooke and Riding (1998, 2000)
for the middle Silurian genus Graticula. On morphological
grounds, the family is allied to the Sporolithaceae.
Family CORALLINACEAE Lamouroux, 1812
Remarks. The Corallinaceae contains those members of
the Corallinales that produce tetrasporangia in uniporate
conceptacles. The tetrasporangia lack apical plugs, and
each sporangium contains zonately arranged spores. Male
and female structures also are produced in uniporate conceptacles. The family contains four subfamilies (see Harvey et al. 2003a, p. 994 for details), all of which have
been reported from the fossil record. Two subfamilies are
represented amongst the types examined during this
study.
Subfamily LITHOPHYLLOIDEAE Setchell, 1943
Remarks. The Lithophylloideae contains those members
of the Corallinaceae in which cells of adjacent vegetative filaments are linked exclusively or principally by
secondary pit-connections (Harvey et al. 2003a, p. 994).
Secondary pit-connections commonly are not preserved
in fossil coralline specimens, but the absence of cell
fusions provides indirect evidence that such specimens
possess secondary pit-connections and thus belong to
the Lithophylloideae. The distinguishing features of genera with nonfossil species are summarized by Woelkerling et al. (2002, p. 370, table 2). The status and
disposition of most exclusively fossil genera referable to
the Lithophylloideae requires reassessment in a modern
context.
404
PALAEONTOLOGY, VOLUME 52
Genus LITHOPHYLLUM Philippi, 1837
Remarks. Species of Lithophylloideae that lack genicula,
lack haustoria, do not have an isobilaterial internal organization and do not have alternating tiers of long and
short cells are placed in the Lithophyllum–Titanoderma
complex (Woelkerling et al. 2002, p. 370). As noted by
Woelkerling et al. (2002) and Braga (2003), molecular
evidence suggests that Lithophyllum and Titanoderma represent distinct genera, but generic boundaries remain
blurred on morphological grounds, and thus they have
been treated as a complex using the oldest generic name
Lithophyllum. Lithophyllum includes species that are
exclusively fossil, or exclusively nonfossil, or both fossil
and nonfossil. Both species dealt with here are known
from fossil and from nonfossil specimens.
Lithophyllum quadratum Ishijima, 1954
Plate 1, figures 1–3; Text-figure 1A
Basionym. Lithophyllum quadratum Ishijima, 1954, Privately Published, Yuhodo, Tokyo, pp. 37–38, pl. 22, fig. 5a–b; pl. 23, figs 1–4.
1954 Lithophyllum quadratum Ishijima, pp. 37–38, pl. 22,
fig. 5a–b; pl. 23, figs 1–4.
? 1961 Lithophyllum quadratum Ishijima; Johnson, p. 930,
pl. 273, fig. 2.
1965 Lithophyllum cfr. quadratum Ishijima; Ishijima, p. 15,
pl. 3, fig. 1.
? 1975 Lp. quadratum Ishijima; Bellini and Mastrorilli, p. 55.
Repository data. Lithophyllum quadratum Ishijima. ‘IGPS coll.
cat. no. 79439’ (Ishijima 1954, p. 38).
Derivation of Ishijima name. No derivation of the specific epithet
quadratum was provided in the protologue, but it probably
refers to the more or less quadrate cells of the thallus.
Lectotype of Lithophyllum quadratum. Ishijima (1954, p. 36) did
not designate a holotype, and the only original material found
and illustrated in the protologue is one unnumbered thin section
(Text-fig. 1A). In accordance with ICBN Art. 9.13 (McNeill
et al. 2006), we designate here as the lectotype the material in
the thin section labelled ‘R.L.S. Daihanroku’ (R.L.S., Riukiu
Limestone) (Text-fig. 1A). The lectotype was illustrated in the
protologue by Ishijima (1954) by two photos (pl. 22, fig. 5a–b;
pl. 23, fig. 1) and by five drawings (pl. 23, figs 1–5).
Age and locality. Pleistocene; Daihanroku, southern Taiwan.
Daihanroku is now called South Bay, located in southernmost
Taiwan. The limestone cropping out in this area was assigned to
the Riukiu Limestone defined by Hanzawa (1935). This formation is currently named the Hengchun Limestone.
Examination of the lactotype. The lectotype thin section (Textfig. 1A) contains a large fragment of an abraded, subrounded
nodule about 10 mm long and 8 mm wide. The nodule is made
up of a single plant with thin encrusting thalli associated with
bryozoans and encrusting foraminifera. The thallus shows only
dimerous construction. Well-defined cell walls separate adjacent
filaments, and the absence of cell fusions indicate that only secondary pit connections are present. Cells are square in section;
their size ranges from 10 to 15 lm in diameter and from 10 to
15 lm in length, and cells of adjacent filaments generally are
more or less aligned horizontally. Uniporate conceptacles are
200–230 lm in diameter and 70–80 lm in height. The conceptacle floor is characterized by a weak columella mostly 1 to 1.5
cells above conceptacle floor. The pore canal is conical in shape
(with a base 30–40 lm in diameter and 60–80 lm in length)
and is lined by filaments parallel to it. The occurrence of a columella suggests that the conceptacles are probably tetrasporangial.
Remarks. The lectotype of Lithophyllum quadratum
belongs the family Corallinaceae, subfamily Lithophylloideae, genus Lithophyllum as currently circumscribed by
Woelkerling and Campbell (1992), Woelkerling (1996b)
and Braga (2003). Its status as a distinct species, however,
EXPLANATION OF PLATE 1
Figs 1–3. Lithophyllum quadratum Ishijima, 1954, lectotype, thin section ‘R.L.S. Daihanroku’, Ishijima’s Collection; Pleistocene,
Daihanroku, southern Taiwan. 1, part of a lumpy protuberance with abundant uniporate conceptacles. 2–3, longitudinal sections
of uniporate conceptacles, with a poorly developed columella (arrow) and a conical pore canal.
Figs 4–7. Lithophyllum prototypum (Foslie) Foslie, 1905. 4–5, [= Lithoporella crassa Ishijima], lectotype, thin section ‘no. 26 R. Aoki’,
Ishijima’s Collection; Late Miocene, Minami-daito-jima, Okinawa Prefecture, south-western Japan. 4, single plant consisting of
numerous layers of dimerous branches. 5, detail of figure 4 illustrating the uniporate conceptacles which arise from the surface of
individual dimerous branches and become overgrown by other dimerous branches. 6–7, [= Lithoporella hayasakai Ishijima],
lectotype, thin section ‘Acervulina flava’, Ishijima’s Collection; Pleistocene, Kohama-jima, Okinawa Prefecture, south-western
Japan. 6, encrusting dimerous plant on a coral fragment. 7, detail of figure 6 showing the oblique tangential sections of the
conceptacles and dimerous branches composed of ventral palisade cells (p) with each bearing a small epithallial cell (e).
Fig. 8. Hydrolithon onkodes (Heydrich) Penrose and Woelkerling, 1992 [= Porolithon hanzawai Ishijima], lectotype thin section
‘Archaeolithoth. Porolithon Halimeda’, Ishijima’s Collection; Pleistocene, Jun-ko-shi, Pingtung County, southern Taiwan. The small
abraded and rounded fragment characterized by occurrence of horizontally aligned rows of trichocytes (arrows).
Scale bars represent 1 mm in 4, 0.50 mm in 1, 6, and 0.20 mm in 2–3, 5, 7–8.
PLATE 1
1
2
3
4
5
6
7
8
e
p
e
IRYU et al., Lithophyllum, Hydrolithon
406
PALAEONTOLOGY, VOLUME 52
requires further study. Plants referred by Keats (1997) to
the nonfossil species Lithophyllum insipidum Adey, Townsend and Boykins also show horizontal alignment of cells
and conceptacles also have conical shaped pore canals.
Keats (1997), however, did not examine the type of L. insipidum, and the original account (Adey et al. 1982, p. 44,
figs 23, 29) contains no information on pore canal shape
and does not mention horizontal alignment or include
anatomical sections of the holotype (although three sections of other plants mentioned in the protologue occur).
Minor differences also occur in the ranges of cell sizes
and conceptacle dimensions between the specimens
described by Adey et al. (1982), by Keats (1997) and in
the type of L. quadratum (reported above). Lythophyllum
insipidum has been also recognized off Lanai (Hawaii;
Webster et al. 2007). Should further study of the types
show that L. quadratum and L. insipidum are conspecific,
the specific epithet insipidum, which is based on a nonfossil species, has priority over the epithet quadratum,
which is based on a fossil species (ICBN Art. 11.8; see
McNeill et al. 2006, p. 25).
Plants identified as Lithophyllum quadratum also have
been recorded from Kita-Daito-Jima (Johnson 1961) and
from Corsica (Bellini and Mastrorilli 1975), but these
identifications are problematic. The specimen described
and illustrated by Johnson (1961) lacks conceptacles while
Bellini and Mastrorilli (1975) did not illustrate their
specimen.
Lithophyllum prototypum (Foslie) Foslie, 1905
Plate 1, figures 4–7; Text-figure 1B–C
Basionym. Lithothamnion prototypum Foslie, 1897, Det Kongelige
Norske Videnskabers Selskabs Skrifter, 1897 (1), p. 18.
1897
1898a
1900
1905
Lithothamnion prototypum Foslie, p. 18.
Melobesia prototypa (Foslie); Foslie, p. 11.
Dermatolithon prototypum (Foslie); Foslie, p. 22.
Lithophyllum prototypum (Foslie); Foslie, p. 129.
1943
1970
1985
1990
1992
1992
1998
2006
Goniolithon prototypum (Foslie); Setchell and
Mason, p. 89.
Tenarea prototypa (Foslie); Adey, p. 7.
Titanoderma prototypum (Foslie); Woelkerling,
Chamberlain and Silva, p. 333.
Lithophyllum prototypum (Foslie) Foslie; Campbell
and Woelkerling, text-fig. 3.
Lithophyllum prototypum (Foslie) Foslie; Woelkerling
and Campbell, pp. 67–78, figs 42–49; includes a
summary of many earlier records.
Lithophyllum tesellatum Lemoine; Iryu, p. 1171.
Titanoderma prototypum (Foslie); Woelkerling,
Chamberlain and Silva; Baba, pp. 619–620, pl. 3–36,
figs A–B.
Lithophyllum prototypum (= Lithophyllum tessellatum); Camoin et al., p. 284.
Heterotypic synonyms. Lithoporella hayasakai Ishijima, 1942a;
Taiwan Tigaku Kizi (Bulletin of Formosan Geography), 13(4), pp.
129–130, pl. 9, fig. 1; 1 text-fig. (unnumbered). Lithoporella
crassa Ishijima, 1942a, Taiwan Tigaku Kizi (Bulletin of Formosan
Geography), 13(4), pp. 131–132, pl. 10, fig. 8; pl. 11, figs 9–10;
text-figs 1–2.
1942a Lithoporella hayasakai Ishijima, pp. 129–130, pl. 9,
fig. 1; 1 text-fig. (unnumbered).
1942a Lithoporella crassa Ishijima, pp. 131–132, pl. 10,
fig. 8; pl. 11, figs 9–10; text-figs 1–2.
1951 Lithoporella crassa Ishijima; Ishijima, text-fig. 2.
1954 Lithoporella crassa Ishijima; Ishijima, p. 50, pl. 48,
figs 1–3; pl. 49, figs 1–3.
1961 Lithoporella cf. L. crassa Ishijima; Johnson, pp. 936–
937, pl. 276, fig. 6.
1961 Lithoporella longicella Johnson n. sp., p. 937, pl. 276,
figs 3–4.
1967 Lithoporella crassa Ishijima; Ishijima, pl. 2, fig. 3.
Further heterotypic synonyms are listed in Woelkerling and
Campbell (1992, p. 67).
Repository data. Lithoporella hayasakai Ishijima, 1942a; not indicated in protologue. Lithoporella crassa Ishijima, 1942a; ‘IGPS
coll. cat. no. 79451’ (in Ishijima 1954, p. 50).
T E X T - F I G . 1 . Thin sections of types; Ishijima’s collection; Institute of Geology and Paleontology, Graduate School of Science,
Tohoku University, Sendai, Japan. Scale bar represents 2 cm. A, Lithophyllum quadratum Ishijima, 1954; ‘R.L.S. Daihanroku’,
lectotype. B, Lithoporella hayasakai Ishijima, 1942a; ‘Acervulina flava’, lectotype. C, Lithoporella crassa Ishijima, 1942a; no. 26 R. Aoki,
lectotype. D, Porolithon hanzawai Ishijima, 1954; ‘Archaeolithoth. Porolithon Halimeda’; lectotype. E, Lithothamnion taishakuensis
Ishijima, 1960a; ‘Archaeolithothamnium megamiensis Ishi. (8)’, lectotype. F, Lithothamnion ishizuchiensis Ishijima, 1960b; ‘4’,
isolectotype. G, Mesophyllum arakuraensis Ishijima, 1954; ‘498 ⁄ 3’, lectotype. H, Mesophyllum arakuraensis Ishijima, 1954; ‘499’. I,
Mesophyllum contii Ishijima, 1954; ‘510’, lectotype. J, Lithothamnion ishizuchiensis Ishijima, 1960b; ‘14’, lectotype. K, Lithothamnion
kasedaensis Ishijima, 1954; ‘no. 154’, lectotype. L, Archaeolithothamnium kobamazimensis Ishijima, 1942b; ‘Pellatispira A’, lectotype. M,
Archaeolithothamnium kobamazimensis Ishijima, 1942b; ‘Pellatispira madraszi douvillei’, syntype. N, Archaeolithothamnium kuboiensis
Ishijima, 1942b; ‘Archaeolithothamnium kuboiensis n. sp. Ishijima’, lectotype. O, Archaeolithothamnium taiwanensis Ishijima, 1942c;
21222, lectotype. P, Lithophyllum nishiwadai Ishijima, 1944; ‘Lithophyllum nishiwadai n. sp.’, lectotype. Q, Lithophyllum otsukiensis
Ishijima, 1954; ‘Lithoporella sp. nov.’, lectotype. R, Lithophyllum otsukiensis Ishijima, 1954; ‘no. 143’, syntype.
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
Derivation of Ishijima names. No derivations of the specific
epithets hayasakai and crassa were provided in the protologues. The epithet hayasakai, however, almost certainly was
chosen by Ishijima to honour Prof. Ichiro Hayasaka (Taihoku
Imperial University, Sendai and Shimane University, Matsue).
The epithet crassa, derived from the Latin word ‘crassus’
meaning thick, probably refers to the overall thickness of
lectotype plant material.
B
A
G
M
Holotype of Lithophyllum prototypum. TRH A18-1226 (Woelkerling et al. 2005, pp. 164–165). An account of the type (from
the U.S. Virgin Islands) and of nonfossil specimens from southern Australia is provided by Woelkerling and Campbell (1992,
pp. 67–78, figs 42–49).
Lectotype of Lithoporella hayasakai. Ishijima (1942a) did not
typify Lithoporella hayasakai. In accordance with ICBN Art. 9.13
C
H
N
D
I
O
407
J
P
E
K
F
L
Q
R
408
PALAEONTOLOGY, VOLUME 52
(McNeill et al. 2006), we designate here as lectotype the material
in the thin section labelled ‘Acervulina flava’ (Text-fig. 1B). The
lectotype is illustrated in the protologue (Ishijima 1942a, pl. 10,
fig. 8).
Lectotype of Lithoporella crassa. Ishijima (1942a) did not designate a type. In accordance with ICBN Art. 9.13 (McNeill et al.
2006), we designate here as lectotype the material in the thin
section labelled ‘no. 26 R. Aoki’ (Text-fig. 1C). The lectotype is
illustrated in the protologue (Ishijima 1942b, pl. 9, fig. 1).
Age and locality. Lithoporella hayasakai Ishijima, Pleistocene;
north of Ufu-daki, Kohama-jima, Ryukyu Islands, Okinawa Prefecture, south-western Japan. Lithoporella crassa Ishijima, Late
Miocene (as ‘Plio-Pleistocene’ in Ishijima 1942a); Ôta-ana,
Minami-daito-jima, Daito Islands, Okinawa Prefecture, southwestern Japan. Iryu et al. (in press) showed that most of reef
deposits occurring over the island surface at Daito Islands
formed during the Late Miocene.
Examination of the lectotype of Lithoporella hayasakai Ishijima. The lectotype (Pl. 1, figs 6–7) consists of a thin encrusting,
dimerous plant, 12 mm long and 1 mm thick. The plant occurs
on a coral fragment within an encrusting sequence composed of
corallines, bryozoans and acervulinids. Ventral filaments consist
of palisade cells (10 lm in diameter and 30 lm in length). Epithallial cells (12–15 lm in diameter and 10 lm long) are locally
preserved. Palisade cells possess pit connections but no cell
fusions. Four random oblique tangential sections of buried conceptacles (270–330 lm in diameter and 100–110 lm high) are
present. The sections do not include conceptacle pore canals or
pores.
Examination of the lectotype of Lithoporella crassa Ishijima. The
lectotype (Pl. 1, figs 4–5) illustrated by Ishijima (1942a) is a part
of a large fragment about 70 mm long and 45 mm high. This
specimen consists of superimposed encrusting dimerous thalli
which locally form warty protuberances. Each superimposed
layer is composed of a single layer of palisade cells (15–20 lm
in diameter and 40–60 lm in length) each bearing an epithallial
cell (subspheroidal in shape, 10–15 lm in diameter). The palisade cells show pit connections. No cell fusions are evident. The
uniporate conceptacles present have a flat floor and an arcuate
roof (320 lm in diameter and 100–120 lm high); they are ellipsoidal in section. The pore canal is more or less cylindrical (30–
40 lm in diameter and 50–70 lm high) and is flanked by the
conceptacle roof.
Remarks. The lectotype specimens of Lithoporella hayasakai Ishijima and Lithoporella crassa Ishijima clearly belong
to Lithophyllum Philippi, 1837 as currently circumscribed
by Woelkerling (1996b), Braga et al. (1993), Braga and
Aguirre (1995), Woelkerling et al. (2002) and Braga
(2003). Lithoporella, the genus to which Ishijima referred
his species, is a currently recognized genus in the Corallinaceae, subfamily Mastophoroideae, and is characterized
by the occurrence of cell fusions. Both Ishijima lectotypes,
however, possess pit connections and lack cell fusions.
Within Lithophyllum, all observable features of both lectotypes are concordant with those of the earlier described
L. prototypum (see Woelkerling and Campbell 1992, pp.
67–78; and Braga and Aguirre 2004), and consequently
both are treated here as heterotypic synonyms of that
species (Table 1). Lithoporella longicella Johnson (1961),
described from Pleistocene-Recent deposits of Eniwetok,
is another possible heterotypic synonym of Lithophyllum
prototypum, but this requires confirmation via a study of
the type. Additional information on Lithoporella and its
type species is provided by Woelkerling (1988, pp. 124–
128; 1996c, pp. 251–255).
Subfamily MASTOPHOROIDEAE Setchell, 1943
Remarks. The Mastophoroideae contains those members
of the Corallinaceae in which cells of adjacent vegetative
filaments are linked exclusively or principally by cell
fusions. Genicula do not occur. The distinguishing features of genera with nonfossil species are summarized by
Harvey et al. (2006, p. 390, table 2). The status of disposition of most exclusively fossil genera referable to the Mastophoroideae requires re-assessment in a modern context.
Genus HYDROLITHON (Foslie) Foslie, 1909
Remarks. The genus Hydrolithon, as delimited by Harvey
et al. (2006), includes those species of Mastophoroideae
in which the basal layer of the thallus is not composed of
palisade cells and in which cells lining the pore canals of
tetrasporangial conceptacles are oriented more or less vertically (perpendicular to the thallus surface). Trichocytes
can occur singly or in horizontal rows or in vertical rows.
Although not diagnostic of the genus as a whole, the
occurrence of numerous horizontal rows of buried trichocytes, which is characteristic of certain species, provides a
useful clue to the generic affinities of specimens that possess them. Hydrolithon includes species that are exclusively fossil, or exclusively nonfossil, or both fossil and
nonfossil. Of the two species dealt with here, H. onkodes
is known from both fossil and nonfossil specimens while
H. taishakuensis is known only from fossil specimens.
Hydrolithon onkodes (Heydrich) Penrose and Woelkerling,
1992
Plate 1, figure 8; Text-figure 1D
Basionym. Lithothamnion onkodes Heydrich, 1897a, Bibliotheca
Botanica, 41, p. 6, pl. 1, fig. 11a–b.
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
1897a Lithothamnion onkodes Heydrich, p. 6, pl. 1,
fig. 11a–b.
1898a Goniolithon onkodes (Heydrich) Foslie, p. 8.
1901 Lithophyllum onkodes (Heydrich) Heydrich,
p. 533.
1909 Porolithon onkodes (Heydrich) Foslie, p. 57.
1988 Spongites onkodes (Heydrich) Penrose and Woelkerling; Penrose and Woelkerling, p. 173, figs 10–14.
1992 Hydrolithon onkodes (Heydrich) Penrose and Woelkerling; Penrose and Woelkerling, p. 83, figs 4–5.
1992 Porolithon onkodes (Heydrich) Foslie; Iryu, pp. 1167,
1171.
1994 Hydrolithon onkodes (Heydrich) Penrose and
Woelkerling; Verheij, p. 107, figs 31–35.
1996 Hydrolithon onkodes (Heydrich) Penrose and
Woelkerling; Penrose, pp. 261–263, fig. 119.
1998 Hydrolithon onkodes (Heydrich) Penrose and
Woelkerling; Baba, p. 561, pl. 3–22, figs A–D.
Heterotypic synonym. Porolithon hanzawai Ishijima, 1954, Privately Published, Y
uhodo, Tokyo, p. 52, pl. 3, fig. 5.
1954 Porolithon hanzawai Ishijima, p. 52, pl. 3, fig. 5.
Repository data. Porolithon hanzawai Ishijima, 1954; ‘IGPS coll.
cat. no. 79452’ (Ishijima 1954, p. 52).
Derivation of Ishijima name. No derivation of the specific epithet
hanzawai was provided in the protologues. The epithet, however,
almost certainly was chosen by Ishijima to honour Prof. Shoshiro Hanzawa (Tohoku University).
Lectotype of Hydrolithon onkodes. TRH A26-1494 (Woelkerling et al. 2005, p. 207). An account of the lectotype (from
Papua New Guinea) is provided by Penrose and Woelkerling
(1988, pp. 162–166, figs 10–14). Information on an isolectotype
in PC (cryptogamic herbarium at the Muséum National d’Histoire Naturelle, Département Systématique et evolution, Paris) is
provided by Woelkerling (1998, p. 357, figs 287–288).
Lectotype of Porolithon hanzawai. Ishijima (1954) did not designate a type. In accordance with ICBN Art. 9.13 (McNeill et al.
2006), we designate here as lectotype the material in the
thin section labelled ‘Archaeolithoth. Porolithon, Halimeda’
(Text-fig. 1D). The lectotype was illustrated in the protologue
(Ishijima 1954, pl. 3, fig. 5).
Age and locality. Pleistocene, Jun-k
o-shi (as Jin-k
o-shi), Takaosh
u, Taiwan. Jin-k
o-shi may be a misspelling of Jun-ko-shi. This
locality is located west of Hengchun, Pingtung County, southern
Taiwan. The sample should be from the Hengchun Limestone
on top of the West-Hengchun Terrace, or from blocks fallen
from the terrace.
Examination of the lectotype of Porolithon hanzawai Ishijima. The lectotype (Pl. 1, fig. 8; Text-fig. 1D) consists of a small
abraded and rounded fragment (1.2 mm long and 0.37 mm
thick) of a coralline plant transversely sectioned. The lectotype
409
almost certainly is monomerous, but monomerous construction
cannot be determined with certainty from transverse sections.
Both monomerous and dimerous construction is known in Porolithon onkodes (Penrose 1996, p. 261). Adjacent filaments are
connected by cell fusions. Cells are 15–20 lm in diameter and
20–40 lm long. Fifteen rows (up to 1.2 mm long and 0.02 mm
high) of horizontally aligned trichocytes are present, but no conceptacles or epithallial cells are evident.
Remarks. The lectotype specimen of Porolithon hanzawai
Ishijima belongs to the family Corallinaceae, subfamily
Mastophoroideae, genus Hydrolithon (Foslie) Foslie as circumscribed by Penrose and Chamberlain (1993), Harvey
et al. (2003a), Braga (2003) and Harvey et al. (2006). The
key feature is the occurrence of numerous buried horizontal rows of trichocytes. Although conceptacles are not
present in the type of P. hanzawai, buried rows of horizontal trichocytes have never been found in species of
Sporolithaceae or Hapalidiaceae or in subfamilies of Corallinaceae other than the Mastophoroideae. Within the
Mastophoroideae, the occurrence of rows of trichocytes is
not diagnostic of genera, but buried horizontal rows of
trichocytes occur abundantly only in some species of Hydrolithon (Woelkerling 1996c). In the protologue (Ishijima
1954, p. 52), Ishijima noted that hanzawai closely resembles Porolithon onkodes (now called Hydrolithon onkodes).
The examination of the lectotype of P. hanzawai has led
to the conclusion that it is morphologically and anatomically concordant with Hydrolithon onkodes and that therefore it is considered to be conspecific with and a
heterotypic synonym of that species (Table 1).
Hydrolithon taishakuensis (Ishijima) comb. nov.
Plate 2, figures 1–3; Text-figures 1E, 2A–B
Basionym. Lithothamnion taishakuensis Ishijima (as Lithothamnium), 1960a, St. Paul’s Review of Arts and Sciences, Natural
Science, 7, pp. 3–4, pl. 1, figs 6–7.
1960a Lithothamnium taishakuensis Ishijima, pp. 3–4, pl. 1,
figs 6–7.
Repository data. Not indicated in protologue.
Derivation of name. No derivation of the specific epithet taishakuensis was provided in the protologue, but it almost certainly is
derived from the name of the type locality, Taishaku-san.
Lectotype of Lithothamnion taishakuenesis. Ishijima (1960a)
did not designate a type. In accordance with ICBN Art. 9.13
(McNeill et al. 2006), we designate here as lectotype the material
in the thin section labelled ‘Archaeolithothamnium megamiensis
Ishi. (8)’ (Text-fig. 1E) that also was illustrated in the protologue
(Ishijima 1960a, pl. 1, figs 6–7).
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PALAEONTOLOGY, VOLUME 52
Age and locality. Late–Early to early–Middle Miocene (Nakamori
et al. 1991); Megami-yama (= Taishaku-san), Megami, Sagara
Town, Haibara County (currently in Makinohara City),
Shizuoka Prefecture, central Japan.
Examination of the lectotype. The lectotype material illustrated
and described by Ishijima (1960a) is easily recognized among
the several specimens present in the thin section (Text-fig. 1E).
This plant consists of an encrusting to warty undulated thallus
7 mm long and up to 2 mm thick. The thallus is monomerous
and has a non-coaxial, plumose, ventrally situated core from
which filaments curve outward to become perpendicular to the
thallus surface. Core region cells are 4 lm in diameter and 6 lm
long, while cells in the peripheral region are 5 lm in diameter
and 6 lm long. Cells of adjacent filaments are linked by cell
fusions. Trichocytes occur in vertical rows (Text-fig. 2A). Several
oblique tangential sections of uniporate conceptacles are present.
Conceptacles are uniporate, rounded in shape (120 lm in diameter and 70 lm in height), and buried in the thallus. Pore canals
of the conceptacles are short, conical in shape, 20 lm in diameter and 50 lm in height (Text-fig. 2B).
Remarks. The occurrence of vertically aligned trichocytes
excludes the assignment of Lithothamnion taishakuensis to
the Hapalidiaceae, subfamily Melobesioideae. The absence
of genicula, the co-occurrence of cell fusions and uniporate
conceptacles in the lectotype of taishakuensis indicate that
the plant belongs to the Corallinaceae, subfamily Mastophoroideae (Braga 2003; Harvey et al. 2003a; Braga et al.
2005). Trichocytes of the sort found in Ishijima’s type are
recorded so far for Neogoniolithon, Spongite and Hydrolithon, but not for other genera of Mastophoroideae (e.g.
Woelkerling 1996c). The occurrence of short pore canals
indicates that the pore canals were lined by cells orientated
more or less perpendicular to the conceptacle roof surface
and these cells did not protrude laterally into the canals.
Filaments of the conceptacle roofs are arranged more or
less parallel to the pore canals. This implies that the conceptacle roofs may have formed by filaments surrounding
and interspersed among sporangial initials. These characters strongly suggest that the plant belongs to Hydrolithon
rather than Neogoniolithon or Spongites (Table 1).
The relationships between Hydrolithon taishakuensis
Ishijima and H. murakoshii (Iryu and Matsuda 1996), a
Recent and fossil species from the Ryukyu Islands, require
further study to determine whether the two species are
conspecific. Both species produce long vertical rows of
trichocytes. The conceptacle chambers in H. murakoshii
appear to be slightly larger and more elongate than those
in H. taishakuensis, but further study may show that this
may not be diagnostically significant at species level.
Some but not all buried conceptacles of H. murakoshii
can become infilled by vegetative cells while no such infilling is evident in the type of H. taishakuensis. This difference also is of questionable taxonomic significance
because infilled conceptacles may occur in both but were
not preserved in Ishijima’s material of H. taishakuensis.
Until further data become available, the two species are
treated here as distinct. Should the two be shown to be
conspecific, the epithet murakoshii, which is based on a
nonfossil type, would have priority over the epithet taishakuensis, which is based on a fossil type.
Family HAPALIDIACEAE Gray, 1864
Remarks. The Hapalidiaceae contains those members of
the Corallinales that produce tetrasporangia in multiporate
conceptacles. The tetrasporangia possess apical plugs, and
each sporangium contains zonately arranged spores. Male
EXPLANATION OF PLATE 2
Figs 1–3. Hydrolithon taishakuensis (Ishijima) comb. nov. [= Lithothamnium taishakuensis Ishijima], lectotype, thin section
‘Archaeolithothamnium megamiensis Ishi. (8)’, Ishijima’s Collection; late–Early to early–Middle Miocene, Megami-yama, Shizuoka
Prefecture, central Japan. 1, encrusting to warty thallus bearing oblique tangential sections of uniporate conceptacles (arrows). 2–
3, encrusting part of the thallus showing several oblique tangential sections of uniporate conceptacles with characteristic short
pore canals.
Figs 4–5. Mesophyllum arakuraensis Ishijima, 1954, lectotype, thin section ‘498 ⁄ 3’, Ishijima’s Collection; Miocene, Arakura, Yamanashi
Prefecture, central Japan. 4, section of zoned peripheral region and buried multiporate conceptacles that protruded from
surrounding thallus surface at the time of reproduction. 5, coaxial ventral core with a single system of filaments with elongate
cells.
Figs 6–7. Mesophyllum contii Ishijima, 1954, lectotype, thin section ‘510’, Ishijima’s Collection; Miocene, Arakura, Yamanashi
Prefecture, central Japan. 6, longitudinal oblique section of a zoned fruticose protuberance. 7, detail of figure 6 showing
longitudinal-tangential section of conceptacles. Note poorly preserved sterile filaments in conceptacle chambers.
Fig. 8. Lithothamnion ishizuchiensis Ishijima, 1960b, lectotype, thin section ‘14’, Ishijima’s Collection; Eocene, Ishizuchi-yama, Ehime
Prefecture, south-western Japan. The very abraded fragment of a coralline thallus with several random sections of multiporate
conceptacles.
Scale bars represent 1 mm in 1, 0.50 mm in 5–6, 8, and 0.20 mm in 2–4, 7.
PLATE 2
1
2
3
4
5
6
7
8
IRYU et al., Corallinalean species
412
PALAEONTOLOGY, VOLUME 52
exclusively fossil genera referable to the Melobesioideae
requires re-assessment in a modern context.
A
d
c
B
T E X T - F I G . 2 . Hydrolithon taishakuensis (Ishijima) comb. nov.
[= Lithothamnium taishakuensis Ishijima], lectotype, thin section
labelled ‘Archaeolithothamnium megamiensis Ishi. (8)’. A,
Longitudinal section of the thallus of monomerous construction
showing the non-coaxial ventral core (c) and a peripheral dorsal
region (d) where trichocytes are aligned in vertical rows
(arrows). Scale bar represents 200 lm. B, uniporate conceptacle
buried in the thallus with a short, conical pore canal. Scale bar
represents 100 lm.
and female structures, however, are produced in uniporate
conceptacles. The family contains three subfamilies (see
Harvey et al. 2003a, p. 994 for details), but only one
(Melobesioideae) has been reported from the fossil record.
Subfamily MELOBESIOIDEAE Bizzozero, 1885
Remarks. The Melobesioideae contains those members of
the Hapalidiaceae in which cells of adjacent filaments are
linked by cell fusions and in which the roofs of the multiporate conceptacles are composed of cells (see Harvey
et al. 2003a, p. 994). The distinguishing features of genera
with nonfossil species are summarized by Harvey et al.
(2003b, p. 653, table 2). The status of disposition of most
Genus MESOPHYLLUM Lemoine, 1928
Remarks. The genus Mesophyllum, as delimited by Harvey
et al. (2003b, p. 653, table 2), includes species that are
exclusively fossil, or exclusively nonfossil, or both fossil
and nonfossil. Some difficulties attend the referral of fossil
specimens to Mesophyllum; these are explained below.
Lemoine (1928) established Mesophyllum for nongeniculate coralline red algae with multiporate conceptacles
and a thallus in which coaxial growth is evident in core
filaments. Lemoine did not designate a type species but
included both nonfossil and fossil species. Subsequently,
Ishijima (1942d, p. 174) designated the nonfossil species
M. lichenoides (Ellis) Lemoine as the type species. Woelkerling and Irvine (1986, 2007) provide information on
and a study of the type specimen of M. lichenoides.
Since 1928, Lemoine’s original concept of Mesophyllum
has undergone various changes, and a second nonfossil
genus (Synarthrophyton Townsend, 1979) containing
species with multiporate conceptacles and at least some
coaxial growth has been described. Both genera belong to
the Hapalidiaceae, subfamily Melobesioideae.
Based on present knowledge (Harvey et al. 2003b, p.
653, table 2), fossil specimens with multiporate conceptacles, cell fusions, and at least some coaxial growth almost
certainly belong to either Mesophyllum or to Synarthrophyton. Synarthrophyton, however, is distinguished from
Mesophyllum principally on a character that has never
been observed in fossils: Synarthrophyton has branched
spermatangial filaments in male conceptacles whilst Mesophyllum has unbranched spermatangial filaments in male
conceptacles (see Woelkerling 1996a; Harvey et al. 2003b,
2005).
Traditionally, palaeontologists have referred all fossil
specimens with multiporate conceptacles, cell fusions, and
at least some coaxial growth to Mesophyllum (e.g. Braga
et al. 1993; Aguirre and Braga 1998; Basso et al. 1998;
Bassi and Nebelsick 2000; Braga 2003), but it simply is
not possible at present to determine whether such fossil
specimens belong to Mesophyllum or Synarthrophyton.
Indeed, information on spermatiangial filaments in fossil
material is totally lacking, and no fossil specimens of Synarthrophyton have been reported in the literature. In these
circumstances, practical considerations have led us to
conclude that the traditional palaeontological practice of
referring all fossil specimens with multiporate conceptacles, cell fusions, and at least some coaxial growth to
Mesophyllum should be continued for the present. This
will consistently allow for the grouping of fossil material
with multiporate conceptacles and at least some coaxial
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
growth under a single generic name, thus facilitating both
cross comparisons between publications and the organization of taxonomic accounts within a publication.
Both species dealt with here under Mesophyllum are
known only from fossil material. We cannot determine
from available evidence whether either might belong to
Synarthrophyton instead of Mesophyllum.
Mesophyllum arakuraensis Ishijima, 1954
Plate 2, figures 4–7; Text-figure 1G–I
Basionym. Mesophyllum arakuraensis Ishijima, 1954, Privately
Published, Y
uhodo, Tokyo, p. 33, pl. 13, figs 3–4.
1954 Mesophyllum arakuraensis Ishijima, p. 33, pl. 13,
figs 3–4.
1961 Mesophyllum cf. M. arakuraensis Ishijima; Johnson,
p. 925, pl. 270, fig. 5; pl. 271, fig. 1.
Heterotypic synonym. Mesophyllum contii Ishijima, 1954, Privately
Published, Y
uhodo, Tokyo, pp. 33–34, pl. 19, figs 1–2.
1954 Mesophyllum Contii Ishijima, pp. 33–34, pl. 19,
figs 1–2.
? 1993 Mesophyllum cf. contii; Fravega et al., text-fig. 3.
Repository data. Mesophyllum arakuraensis, ‘IGPS coll. cat. no.
79434’ (Ishijima 1954, p. 33). Mesophyllum contii, ‘IGPS coll.
cat. no. 79435’ (Ishijima 1954, p. 34).
Derivation of name. No derivations of the specific epithets arakuraensis and contii were provided in the protologues. The epithet
arakuraensis, however, almost certainly was derived from the
name of the type locality, Arakura. The epithet contii almost certainly was chosen by Ishijima to honour Prof. Sergio Conti
(University of Genova, Italy).
413
Age and locality. Mesophyllum arakuraensis, Miocene; Arakura,
Mizuho Village, Minamitsuru County (currently in Fujiyoshida
City), Yamanashi Prefecture, central Japan. Mesophyllum contii,
Miocene; Arakura, Mizuho Village, Minamitsuru County (currently in Fujiyoshida City), Yamanashi Prefecture, central Japan.
Examination of the lectotype and isolectotype of Mesophyllum
arakuraensis. The lectotype is an abraded fragment 4.5 mm long
and 2.5 mm wide, and contains one transverse section and three
longitudinal oblique sections of the fruticose branches. The longitudinal sections show monomerous construction with a single
system of cell filaments that are coaxially arranged in rows in
the core and curve outward to become perpendicular to the
thallus surface. The thallus has evident growth bands resulting
from the interrupted growth of groups of filaments. Cells
(10 lm in diameter and 20 lm long) of adjacent filaments are
connected by cell fusions. Three tangential peripheral sections of
probable multiporate conceptacles are present (260 lm in diameter and 110 lm high; 270 lm in diameter and 180 lm high;
and 300 lm in diameter and 110 lm high). Poorly preserved,
infertile filaments occur in the conceptacle chambers. The conceptacles become buried within the thallus. The transverse section lacks a ventral core and only the dorsally located peripheral
region is preserved. Some growth bands are clearly discernible.
Cells of adjacent filaments are connected by cell fusions. No
conceptacles are present in the transverse section, and neither
epithallial cells nor subepithallial initials were identified.
The isolectotype (in thin section ‘499’) consists of a small
abraded fragment of an encrusting plant, 2 mm long and
0.8 mm thick. The thalli show coaxial ventral core with monomerous construction with a single system of filaments. The cells
(5–8 lm in diameter and 20–40 lm long) are connected by cell
fusions. Neither conceptacles nor epithallial cells nor subepithallial initials are present.
Isolectotype of Mesophyllum arakuraensis. Plant in thin section
labelled ‘499’ (Text-fig. 1H), illustrated by Ishijima (1954, pl. 13,
fig. 3).
Examination of the lectotype of Mesophyllum contii. The lectotype consists of a longitudinal oblique section of a fruticose
plant, 3 mm long and 1.8 mm wide, made up of several growth
stages. The thalli have a monomerous construction with a single
system of filaments that are coaxially arranged in the core and
tend to curve outward to become perpendicular to the thallus
surface. Cells (10–15 lm in diameter and 20–25 lm long) of
adjacent filaments are connected by cell fusions, which, together
with the micritization, make single filaments difficult to distinguish. Multiporate conceptacles are present. The sections of conceptacles are distally longitudinal-tangential and show only the
periphery of the chambers (up to 220 lm in diameter and up to
180 lm high). These conceptacles can be interpreted as the
remains of multiporate conceptacles. The conceptacles protrude
from surrounding thallus surface and later become buried in it.
No epithallial cells were identified.
Lectotype of Mesophyllum contii. Ishijima (1954) did not designate a type. In accordance with ICBN Art. 9.13 (McNeill
et al. 2006), we designate here as lectotype the material in the
thin section labelled ‘510’ (Text-fig. 1I). The lectotype was
illustrated in the protologue by Ishijima (1954) in pl. 19,
figs 1–2.
Remarks. The presence of cell fusions and possible multiporate conceptacles mean that the specimens of Mesophyllum arakuraensis belong to the Hapalidiaceae, subfamily
Melobesioideae (e.g. Braga 2003; Table 1). These features
combined with the occurrence of coaxial growth result in
placement in Mesophyllum as traditionally delimited by
Lectotype of Mesophyllum arakuraensis. Ishijima (1954) did not
designate a type, and there are two thin sections, respectively,
labelled ‘498 ⁄ 3’ and ‘499’. The material in thin section ‘498 ⁄ 3’
has conceptacles whereas the plant in thin section ‘499’ appears
sterile. In accordance with ICBN Art. 9.13 (McNeill et al. 2006),
we therefore designate here as lectotype the material in thin section ‘498 ⁄ 3’ (Text-fig. 1G). The lectotype was illustrated in the
protologue (Ishijima 1954, pl. 13, fig. 4).
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PALAEONTOLOGY, VOLUME 52
palaeontologists. Growth bands such those present in M.
arakuraensis have been illustrated by Woelkerling and
Harvey (1993, figs 7C and 28C) for several nonfossil
southern Australian species of Mesophyllum. As pointed
out by Foster (2001, p. 661; see also Basso 1995), groups
of primary bands that vary in cell size and wall thickness
can result in light-dark pairs of higher order bands. Cell
and growth band differences can be due to environmentally induced effects on growth and reproduction.
The relationships of M. arakuraensis to nonfossil species of Mesophyllum cannot be determined from features
evident in the lectotype or isolectotype.
Multiporate conceptacles and cell fusions in Mesophyllum contii are diagnostic of the Hapalidiaceae, subfamily
Melobesioideae (e.g. Braga et al. 1993; Aguirre and Braga
1998; Braga 2003). The occurrence of coaxial growth suggests an affinity with the genus Mesophyllum, but generic
placement cannot be determined in the absence of information on spermatangial conceptacles, as explained in the
remarks for the genus Mesophyllum above.
Both Mesophyllum contii and M. arakuraensis were
described in the same publication by Ishijima (1954), and
they come from the same type locality. In the present
study, conceptacles up to 200 lm in diameter and
180 lm high were found in the lectotype of M. contii,
but in the protologue, Ishijima (1954, p. 34) reported
conceptacles to be 150–300 lm in diameter and 100–
150 lm high. Ishijima’s dimensions markedly overlap
those found in the lectotype of M. arakuraensis (260–
300 lm in diameter and 110–180 lm high), and cell sizes
in the two taxa also largely overlap (M. arakuraensis,
10 lm in diameter and 20 lm long; M. contii 10–15 lm
in diameter and 20–25 lm long). These two taxa, therefore, are considered conspecific (Table 1). Although the
protologues of M. arakuraensis and M. contii were given
in the same publication (Ishijima 1954), the former is
described before the latter. Thus, M. arakuraenis has
nomenclatural priority over M. contii. Ishijima (1954,
p. 33) stated that M. arakuraensis most closely resembled
M. hasimotoi Ishijima (1943, p. 647), while M. contii
more or less resembled M. yabei Ishijima (1954, p. 34).
Ishijima (1954), however, made no direct comparisons of
M. arakuraensis and M. contii. Should all four species
prove to be conspecific, the oldest available name would
be Mesophyllum hashimotoi and not M. arakuraensis or
M. contii.
Melobesioid taxa of uncertain generic placement
The types of the following two species belong to the
Melobesioideae because they show cell fusions and multiporate conceptacles whose roofs are composed of cells.
There are insufficient data, however, to allow for
unequivocal generic placement in a modern context (see
discussion in Aguirre and Braga 1998; Rasser and Piller
1999; Bassi and Nebelsick 2000; Braga 2003; Table 1).
Ishijima’s original names (basionyms) are used in the
accounts below.
Lithothamnion ishizuchiensis Ishijima, 1960b
Plate 2, figure 8; Plate 3, figures 1–2; Text-figures 1F, 1J
Basionym. Lithothamnion ishizuchiensis Ishijima (as Lithothamnium), 1960b. Science Reports of the Tohoku University, second
series (Geology), special volume 4 (Hanzawa Memorial Volume),
p. 155, pl. 14, figs 1–2.
EXPLANATION OF PLATE 3
Figs 1–2. Lithothamnion ishizuchiensis Ishijima, 1960b, isolectotype, thin sections ‘4’ (Fig. 1) and lectotype, thin section ‘14’ (Fig. 2),
Ishijima’s Collection; Eocene, Ishizuchi-yama, Ehime Prefecture, south-western Japan. Details of oblique longitudinal sections of
multiporate conceptacles. Note that sterile filaments persist in the conceptacle chambers (arrows).
Figs 3–4. Lithothamnion kasedaensis Ishijima, 1954, lectotype, thin section ‘no. 154’, Ishijima’s Collection; Pliocene (?), Kaseda,
Kagoshima Prefecture, south-western Japan. 3, the encrusting plant with several sections of multiporate conceptacles. 4, detail of
figure 3 illustrating the multiporate conceptacles in which sterile filaments are preserved (arrows).
Figs 5–6. Sporolithon kobamazimensis (Ishijima) comb. nov. [= Archaeolithothamnion kobamazimensis Ishijima (as Archaeolithothamnium)],
Ishijima’s Collection; Eocene, Kohama-jima, Okinawa Prefecture, south-western Japan. 5–6, lectotype, thin section ‘Pellatispira A’. 5,
lumpy protuberance showing transverse and longitudinal sections of compartments. 6, detail of figure 5 showing some compartments
grouped into sori.
Figs 7–8. Sporolithon kobamazimensis (Ishijima) comb. nov. [= Archaeolithothamnion kobamazimensis Ishijima (as Archaeolithothamnium)],
Ishijima’s Collection; Eocene, Kohama-jima, Okinawa Prefecture, south-western Japan. 7–8, syntype, thin section ‘Pellatispira madraszi
douvillei’. 7, lumpy protuberance with compartments grouped in sori. 8, detail of figure 7 showing longitudinal sections of
compartments.
Scale bars represent 0.50 mm in 3, 5, 7, and 0.20 mm in 1–2, 4, 6, 8.
PLATE 3
1
2
3
4
5
6
7
8
IRYU et al., Lithothamnion, Sporolithon
416
PALAEONTOLOGY, VOLUME 52
1960b Lithothamnium ishizuchiensis Ishijima, p. 155, pl. 14,
figs 1–2.
Repository data. In the protologue, Ishijima (1960b, p. 159) indicated that the specimens were deposited in the Institute of Geology, Rikkyo University. The slides upon which the protologue
illustrations are based, however, are currently housed at the
Institute of Geology and Paleontology, Graduate School of Science, Tohoku University.
Derivation of name. No derivation of the specific epithet ishizuchiensis was provided in the protologue, but it almost certainly
is derived from the name of the type locality, Ishizuchi-yama.
Lectotype of Lithothamnion ishizukiensis. Ishijima (1960b) did
not designate a type. In accordance with ICBN Art. 9.13 (McNeill et al. 2006), we designate here as lectotype the material in the
thin section labelled ‘14’ (Text-fig. 1J). The lectotype was illustrated in the protologue (Ishijima 1960b, pl. 14, fig. 2). Material
in the designated lectotype is better preserved and shows more
features than in the isolectotype (see below).
Isolectotype of Lithothamnion ishizukiensis. Plants in the thin
section labelled ‘4’ (Text-fig. 1F). The isolectotype also was illustrated in the protologue (Ishijima 1960b, pl. 14, fig. 1).
Age and locality. Eocene; Ishizuchi-yama, Ehime Prefecture,
south-western Japan.
Examination of the lectotype and isolectotype. The lectotype (in
thin section ‘14’) is a very abraded fragment of a lumpy plant,
2.7 mm high and 2 mm long. The plant shows successive growth
stages with an encrusting ventral portion and a lumpy protuberance. The thallus is monomerous with non-coaxial (plumose)
ventral core; the thallus construction is monomerous in the protuberance. Epithallial cells were not preserved. The cells are very
small (c. 4 lm in diameter and 8 lm long), and cells of adjacent
filaments are connected by cell fusions. The numerous conceptacles, up to 300 lm in diameter and up to 80 lm high, are multiporate and ellipsoidal in shape, and are buried in the thallus.
Sterile filaments are preserved in conceptacle chambers.
The isolectotype (in thin section ‘4’) consists of an oblique
section of a small, abraded fragment, 2 mm long and 0.7 mm
thick. The thallus is monomerous. Epithallial cells are not preserved. The cells of adjacent filaments both in the ventral and in
the dorsal part of the thallus are connected by cell fusions. Cells
are 4–6 lm in diameter and 6–10 lm long. Three sections of
multiporate conceptacles buried in the thallus are present
(420 lm in diameter and 160 lm high; 230 lm in diameter and
130 lm high; 250 lm in diameter and 130 lm high). Sterile filaments persist in conceptacle chambers. None of these sections
clearly shows the pore canals.
Remarks. The occurrence of multiporate conceptacles and
cell fusions supports placement of this taxon in the Hapalidiaceae, subfamily Melobesioideae (e.g. Rasser and Piller
1999; Braga 2003). Lithothamnion (i.e. Lithothamnion
Heydrich 1897b nom. cons., non-Lithothamnium Philippi
1837 nom. rejec.; see Woelkerling 1983), the genus to
which Ishijima assigned his species, is characterized by
flattened epithallial cells with flared distal walls, and preservation of these cells is required for proper generic
placement of specimens (Aguirre and Braga 1998; Rasser
and Piller 1999; Braga 2003). The absence of preserved
epithallial cells, therefore, precludes any certain generic
assignment of this species (Table 1) and there are no
other features evident in Ishijima’s specimens that can be
used to ally his species to any particular genus or species
of Melobesioideae.
Lithothamnion kasedaensis Ishijima, 1954
Plate 3, figures 3–4; Text-figure 1K
Basionym. Lithothamnion kasedaensis Ishijima (as Lithothamnium), 1954, Privately Published, Y
uhodo, Tokyo, p. 28, pl. 7,
fig. 2a–c.
1954 Lithothamnium kasedaensis Ishijima, p. 28, pl. 7,
fig. 2a–c.
Repository data. In the protologue, Ishijima (1954, p. 28) indicated that the specimen was deposited in the Geological Institute
of the Tokyo University of Education (current Institute of Geoscience, Tsukuba University). The slide upon which the protologue illustrations are based, however, is currently housed at the
Institute of Geology and Paleontology, Graduate School of Science, Tohoku University.
Derivation of name. No derivation of the specific epithet kasedaensis has been provided in the protologue, but it almost certainly
is derived from the name of the type locality, Kaseda.
Lectotype of Lithothamnion kasedaensis. Ishijima (1954) did
not designate a type. In accordance with ICBN Art. 9.13
(McNeill et al. 2006), we designate here as lectotype the material
in thin section ‘no. 154’ (Text-fig. 1K). The lectotype was illustrated in the protologue (Ishijima 1954, pl. 7, fig. 2a–c).
Age and locality. Pliocene (?); Kaseda, Kawanabe County (currently in Minamisatsuma City), Kagoshima Prefecture, southwestern Japan.
Examination of the lectotype. The lectotype specimen is 7 mm
long and 1 mm thick and consists of a tangential oblique section
of an encrusting plant. The thallus is monomerous with a noncoaxial ventral core of filaments. The cells are small (5–15 lm in
diameter and 10–20 lm long), and cells of adjacent filaments are
connected by cell fusions. Epithallial cells were not preserved. Several multiporate conceptacles (up to 200 lm in diameter and up
to 60 lm high) are present. Some represent remains of early developmental stages of conceptacles. The conceptacles protrude from
the surrounding thallus surface and later become buried in it.
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
Remarks. The occurrence of multiporate conceptacles
and cell fusions supports placement of L. kasedaensis in
the Hapalidiaceae, subfamily Melobesioideae (e.g. Rasser
and Piller 1999; Braga 2003). The absence of preserved
epithallial cells, however, precludes any certain generic
assignment of this taxon (Table 1) for reasons already
outlined, and there are no other features evident in
Ishijima’s specimens that can be used to ally his
species to any particular genus or species of Melobesioideae.
Family SPOROLITHACEAE Verheij, 1993
Remarks. The Sporolithaceae contains those members of
the Corallinales that produce tetrasporangia individually
in calcified compartments (not conceptacles); commonly
the compartments are grouped into sori. The tetrasporangia possess apical plugs, and each sporangium
contains cruciately arranged spores. By contrast, male
and female structures are produced in uniporate
conceptacles similar to those of other coralline red
algae.
Verheij (1993) originally established the Sporolithaceae for the genus Sporolithon (see also Woelkerling
1988), which hitherto had been placed in the Corallinaceae. Another genus, Heydrichia, was described by
Townsend et al. (1994). Both genera are extant. Heydrichia differs from Sporolithon in having a pseudoparenchymatous wall (involucre) surrounding each
tetrasporangial compartment and in having more than
one stalk cell in each sporangial compartment (Harvey
et al. 2002). In Sporolithon, by contrast, tetrasporangial
compartments lack involucres and have only one stalk
cell in each sporangial compartment. Phylogenetic studies involving genera of Sporolithaceae have been published by Bailey and Chapman (1996) and Harvey et al.
(2002, 2003a). These analyses indicate that this family
is a distinct lineage separate from Corallinaceae and
Hapalidiaceae on molecular grounds, and this is supported by existing morphological ⁄ anatomical data
(Harvey et al. 2002, 2003a).
Of the above two extant genera, only Sporolithon is also
known with certainty from both fossil and nonfossil
material (e.g. Moussavian and Kuss 1990; Braga et al.
1993; Aguirre and Braga 1998; Bassi 1998; Braga and
Bassi 2006; Bassi et al. 2007; Tomás et al. 2007). Several
exclusively fossil genera, however, also are referable to the
Sporolithaceae. These include Axiophyllum Moussavian
(1989), Hemiphyllum Lemoine (1970), Kymalithon Lemoine and Emberger (1967), Parakymalithon Moussavian
(1987; see also Tomás et al. 2007), and Paraphyllum Lemoine (1970), but their status as distinct genera requires
further assessment.
417
Genus SPOROLITHON Heydrich (1897)
Remarks. The distinguishing features of Sporolithon have
been explained above. In the palaeontological literature,
many species referable to Sporolithon Heydrich (1897b)
traditionally have been placed in Archaeolithothamnion
Rothpletz ex Foslie (1898b, p. 3), commonly using the
orthographic variant spelling Archaeolithothamnium. The
name Archaeolithothamnion was validly published by Foslie (1898b, p. 3) whereas the name Archaeolithothamnium
was first coined by Rothpletz (1891) but was not validly
published (Woelkerling 1988; Moussavian and Kuss
1990). In the context of ICBN Art. 61.1 and 61.4 (McNeill et al. 2006), the validly published Archaeolithothamnion is the correct spelling, while the orthographic variant
Archaeolithothamnium is to be corrected to the validly
published form of the name whenever it appears in print.
Such corrections do not affect the authorship or priority
of names.
Over 140 species of Sporolithon (including Archaeolithothamnion) have been described. The vast majority
are known only from fossil specimens. While species
concepts have been clarified in a modern context for
some nonfossil species (e.g. Verheij 1993; Harvey et al.
2002), meaningful concepts for fossil species scarcely
exist. It is highly likely that a comprehensive reassessment of fossil species referable to Sporolithon will result
in most being reduced to synonymy or being treated as
dubious taxa. In the absence of such a study, which is
beyond the scope of the present paper, one can do little more than to report on types and update generic
names without commenting on possible relationships or
characters of diagnostic value. This approach has been
used in relation to the three Ishijima taxa dealt with
here.
Sporolithon kobamazimensis (Ishijima) comb. nov.
Plate 3, figures 5–8; Text-figure 1L–M
Basionym. Archaeolithothamnion kobamazimensis Ishijima (as Archaeolithothamnium), 1942b, Transactions of the Natural History
Society of Formosa, 32, pp. 354–355 (in Japanese), p. 357 (in
English), pl. 1, figs 1–3; 1 text-figure (unnumbered).
1942b Archaeolithothamnion kobamazimensis Ishijima (as
Archaeolithothamnium), pp. 354–355, pl. 1, figs 1–3;
1 text-fig (unnumbered).
1954 Archaeolithothamnion kobamajimensis Ishijima; Ishijima, pp. 18–19, pl. 1, fig. 3a–b; pl. 3, fig. 4 (as
Archaeolithothamnium).
Repository data. Institute of Geology and Paleontology, Graduate
School of Science, Tohoku University (Ishijima 1942b, p. 357, as
‘Geological Institute of the Tohoku Imperial University’).
418
PALAEONTOLOGY, VOLUME 52
Derivation of name. No derivation of the specific epithet
kobamazimensis was provided in the protologue, but it almost
certainly was derived from the name of the type locality,
Kobamazima (= currently called Kohama-jima).
Lectotype of Sporolithon kobamazimensis. Ishijima (1942b) did
not designate a type and there are two thin sections, respectively, labelled ‘Pellatispira A’ and ‘Pellatispira madraszi douvillei’. The material in thin section ‘Pellatispira A’ has several
well-preserved sporangial compartments cut at different angles,
whereas the plant in thin section ‘Pellatispira madraszi douvillei’
show mostly longitudinally oblique sectioned sporangial compartments. In accordance with ICBN Art. 9.13 (McNeill et al.
2006), we designate here as lectotype the material in the thin
section labelled ‘Pellatispira A’ (Text-fig. 1L). The two thin sections form part of a single specimen (as defined in ICBN Art.
8.2). The lectotype was illustrated in the protologue (Ishijima
1942b; unnumbered text-figure). In the protologue Ishijima
(1942c, p. 357) used the phrase ‘type slides’ which suggests that
he considered both slides to be part of the type. The second
thin section labelled as ‘Pellatispira madraszi douvillei’ represents therefore the syntype (as defined in ICBN Art. 9.4). The
specimen illustrated in pl. 1, figs 1–3 (Ishijima 1942b) was not
found.
longitudinally oblique sectioned. No layers of elongated cells
occur below the compartments.
Remarks. Calcified compartments are a diagnostic character of the family Sporolithaceae (Verheij 1993). The
reproductive features of the lectotype of ‘A. kobamazimensis’ are characteristic of sporangial plants of Sporolithon
(Table 1), to which we transfer the species here. Its status
as a distinct species requires further evaluation in the
context of a more comprehensive reassessment of fossil
species referable to Sporolithon.
Sporolithon kuboiensis (Ishijima) comb. nov.
Plate 4, figures 1–3; Text-figure 1N
Basionym. Archaeolithothamnion kuboiensis Ishijima (as Archaeolithothamnium), 1942b, Transactions of the Natural History Society of Formosa, 32, pp. 355–356 (in Japanese), p. 358 (in
English), pl. 2, figs 1–3; 1 text-figure (unnumbered).
Syntype of Sporolithon kobamazimensis. Plant in thin section
labelled ‘Pellatispira madraszi douvillei’ (Text-fig. 1M), illustrated
by Ishijima in 1954 (pl. 1, fig. 3a–b; pl. 3, fig. 4).
1942b Archaeolithothamnion kuboiensis Ishijima (as Archaeolithothamnium), pp. 355–356, pl. 2, figs 1–3; 1
text-figure (unnumbered).
1954 Archaeolithothamnion kuboiensis Ishijima; Ishijima,
pp. 19–20, pl. 2, fig. 4; pl. 3, figs 1–3 (as Archaeolithothamnium).
Age and locality. Eocene; Kohama-jima, Ryukyu Islands, Okinawa Prefecture, south-western Japan.
Repository data. Not indicated in protologue, but subsequently
(Ishijima 1954, p. 20) stated to be ‘IGPS coll. cat. no. 79424’.
Examination of the lectotype. The thin section labelled ‘Pellatispira A’ (Text-fig. 1L) contains the specimen illustrated by Ishijima (1942b) in an unnumbered text-figure.
The material in the thin section labelled ‘Pellatispira A’ (Textfig. 1L) consists of an abraded fragment of a protuberance
2 mm wide and 3 mm long that comes from a lumpy or fruticose plant (Pl. 3, figs 5–6). The fragment is monomerous in
construction, and filaments radiate from the core to the periphery. Each filament is composed of cells (30–40 lm long and 10–
20 lm in diameter) which are laterally connected to cells of
adjacent filaments by cell fusions. No epithallial cells were seen.
The protuberance has sporangial compartments that are cut at
different angles in the section. The compartments, 50–60 lm in
diameter and 70–80 lm high, are aggregated into irregularly
shaped sori. No layers of elongated cells are present below the
compartments.
Derivation of name. No derivation of the specific epithet kuboiensis was provided in the protologue, but it almost certainly was
derived from the name of the type locality, Kuboi.
Examination of the syntype. The thin section labelled ‘Pellatispira
madraszi douvillei’ (Text-fig. 1M) contains the specimen illustrated in 1954 (Ishijima 1954, pl. 1, fig. 3a–b; pl. 3, fig. 4). The
specimen, 2.4 mm wide and 2.8 mm long, is a fragment of a
protuberance (Pl. 3, figs 7–8). The thallus organization is radial
and consists of cells 30–40 lm long and 20 lm in diameter. Cell
fusions are present. No epithallial cells were recognized. Several
compartments grouped into four sori are present. The compartments, 50–60 lm in diameter and 70–100 lm high, are mostly
Lectotype of Sporolithon kuboiensis. Ishijima (1942b) did not
designate a type. In accordance with ICBN Art. 9.13 (McNeill
et al. 2006), we designate here as the lectotype the plant in the
thin section labelled ‘Archaeolithothamnium kuboiensis n. sp.
Ishijima’ (Text-fig. 1N). The lectotype was illustrated in the protologue by Ishijima (1942b, pl. 2, figs 1–3); and one unnumbered text-figure).
Age and locality. Miocene; Kuboi, Oishi Village, Minamitsuru
County, (currently in Town of Fujikawaguchiko), Yamanashi
Prefecture, central Japan.
Examination of the lectotype. The lectotype slide (Text-fig. 1N)
contains one large rounded section (19 · 13 mm) of a mono-specific rhodolith. The encrusting coralline thallus envelopes a coral
nucleus (about 4 mm in diameter). The thallus shows a dorsiventral monomerous organization with a poorly developed plumose
ventral region from which cell filaments curve towards the thallus
surface. Cells of core filaments are 15 lm long and 10 lm in
diameter. Cells (15 lm long and 10 lm in diameter) of contiguous filaments in the peripheral region are laterally aligned, giving
to the thallus the aspect of a uniform grid. Cells of adjacent
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
filaments are connected by cell fusions. No epithallial cells were
recognized. Several groups of calcified sporangial compartments
(sori) are present, mostly sectioned in a transverse oblique direction. The sori occur all along the thallus surface and are buried in
the thallus. The compartments, 50–60 lm in diameter and 70–
80 lm high, are elliptical to ovoid in shape. Ovoid stalk cells
occur at the base of some compartments (Pl. 4, fig. 2).
Remarks. Calcified compartments are diagnostic characters of Sporolithon, family Sporolithaceae (Verheij 1993).
The reproductive features of the lectotype of ‘A. kuboiensis’ are characteristic of sporangial plants of Sporolithon
(Table 1) to which we transfer the species here. Its status
as a distinct species requires further evaluation in the
context of a more comprehensive reassessment of fossil
species referable to Sporolithon.
Sporolithon taiwanensis (Ishijima) comb. nov.
Plate 4, figures 4–5; Text-figure 1O
Basionym. Archaeolithothamnion taiwanensis Ishijima (as Archaeolithothamnium), 1942c, Taiwan Tigaku Kizi, 13, pp. 2–3, 2 textfigures (unnumbered).
?
?
?
?
1942c Archaeolithothamnion taiwanensis Ishijima (as
Archaeolithothamnium), pp. 2–3, 2 text-figures
(unnumbered).
1954 Archaeolithothamnion taiwanensis Ishijima (as
Archaeolithothamnium), p. 18, pl. 1, fig. 5.
1957 Archaeolithothamnion taiwanensis Ishijima; Johnson,
pp. 218–219, pl. 46, figs 4–6 (as Archaeolithothamnium).
1961 Archaeolithothamnion cf. A. taiwanensis Ishijima;
Johnson, p. 919, pl. 267, fig. 7 (as Archaeolithothamnium).
1964 Archaeolithothamnion cf. A. taiwanensis Ishijima;
Johnson, pp. G10–11 (as Archaeolithothamnium).
1970 Archaeolithothamnion taiwanensis Ishijima; Vannucci, pp. 439–440, pl. 2, fig. 3 (as Archaeolithothamnium).
1975 A. taiwanensis Ishijima; Bellini and Mastrorilli, p. 42
(as Archaeolithothamnium).
1975 Archaeolithothamnion sp. cf. A. taiwanensis Ishijima;
Edgell and Basson, p. 170, pl. 1, fig. 5 (as Archaeolithothamnium).
1988 Archaeolithothamnion taiwanensis Ishijima; Fravega
et al., pp. 326–327, text-fig. 2 (as Archaeolithothamnium).
Repository data. Not indicated in protologue, but subsequently,
Ishijima (1954, p. 18) stated that the specimens deposited in the
Institute of Geology, Taihoku University. The material designated here as the lectotype, however, is stored at the Institute of
Geology and Paleontology, Graduate School of Science, Tohoku
University.
419
Derivation of name. No derivation of the specific epithet taiwanensis was provided in the protologue, but it almost certainly was
derived from the name of the type locality, Taiwan.
Lectotype of Sporolithon taiwanensis. Ishijima (1942c) did not
designate a type. In accordance with ICBN Art. 9.13 (McNeill
et al. 2006), we designate here as lectotype the material in the section of slide 21222 (Text-fig. 1O). The lectotype was illustrated by
Ishijima (1942c, p. 120) in the left-hand text-figure in the protologue. The same figure appears as pl. 1, fig. 5 in Ishijima (1954).
o (current Taitung
Age and locality. Pliocene; Payapaya, Tait
County), Taiwan.
Examination of the lectotype. The lectotype is up to 8 mm long
and up to 1.4 mm thick, abraded and attached to a large acervulinid foraminifer. The thallus is encrusting and monomerous
with core filaments curving upwards to form a peripheral
region. Cells (15 lm long and 10 lm in diameter) of adjacent
filaments are connected by cell fusions. No epithallial cells were
recognized. Several calcified compartments (40 lm in diameter
and 60–70 lm high) occur at the surface of the plant. They are
grouped into sori which are not raised above surrounding thallus surface. The compartment wall is very thick and dark in
colour. At high magnification under light microscope very small
cell filament-like structures appear to surround the inner
compartment wall.
Remarks. Archaeolithothamnion taiwanensis was first
published as a new species by Ishijima (1942c, as Archaeolithothamnium) and then published again as a ‘new
species’ by the same Ishijima (1954) without reference to
the earlier account. Both accounts are based on the same
material and thus the name dates from 1942 when the
first account was validly published. The plants identified
as A. taiwanensis by Bellini and Mastrorilli (1975) and
Edgell and Basson (1975) cannot be confidently attributed to this species as those authors did not illustrate
reproductive features. The specimen ascribed to A. taiwanensis and illustrated by Fravega et al. (1988, text-fig.
2) has a clear basal layer of elongated cells which is not
present in the type of taiwanensis.
The reproductive features evident in the lectotype of
Archaeolithothamnion taiwanensis are diagnostic of the
Sporolithaceae, and consequently the species is transferred
here to Sporolithon (Table 1). The presence of thick compartment walls, however, might suggest a possible fusion
between the original very small cells which surrounded
the compartments. If this is true, these cell filaments are
indicative of involucres. These features are considered
diagnostic characters of Heydrichia Townsend, Chamberlain and Keats (Harvey et al. 2002). However, the possible
affinity of taiwanensis to Heydrichia cannot be determined
with certainty until further material from the type locality
has to be collected and examined.
420
PALAEONTOLOGY, VOLUME 52
Johnson (1964) and Bellini and Mastrorilli (1975)
reported A. taiwanensis (as Archaeolithothamnium) but
did not illustrate the specimens (nor reported information about the occurrence of conceptacles) that they
attributed to this species. Edgell and Basson (1975) and
Fravega et al. (1988) illustrated the specimens attributed
to this species but the possible involucres visible in the
photos need to be confirmed in the original material.
Thus these records require confirmation.
TAXA OF UNCERTAIN FAMILY
PLACEMENT
The types of the following two taxa lack sufficient data to allow
for family or genus placement within the Corallinales (Table 1).
Consequently they are referred to by the basionym (original
name) although family and generic disposition are uncertain.
Lithophyllum nishiwadai Ishijima, 1944
Plate 4, figure 6; Text-figure 1P
1944 Lithophyllum Nishiwadai Ishijima, pp. 72–73,
text-fig. 7.
? 1975 Lithophyllum nishiwadai Ishijima; Bellini and
Mastrorilli, p. 55.
Repository data. Not indicated in protologue.
Derivation of name. No derivation of the specific epithet nishiwadai was provided in the protologue, but the epithet almost
certainly was chosen by Ishijima to honour Mr. Kyugaku
Nishiwada who reported the occurrence of ‘Lithothamnium
ramosissimum Ruess’ from Megami-yama, Shizuoka Prefecture,
central Japan (Nishiwada 1895; Ishijima 1944, p. 71).
Lectotype of Lithophyllum nishiwadai. Ishijima (1944) did not
designate a type. In accordance with ICBN Art. 9.13 (McNeill
et al. 2006), we designate here as lectotype material in the thin
section labelled ‘Lithophyllum nishiwadai n. sp.’. The lectotype
was illustrated by Ishijima (1944, text-fig. 7). The thin section is
illustrated here in Text-fig. 1P.
Ishijima (1944) did not provide a description of his material
but gave an indirect reference to a description validly published
by Nishiwada (1895) of plants that Nishiwada had identified as
Lithothamnium ramosissimum Reuss. Although Ishijima did not
explicitly cite Nishiwada (1895) in his paper, Ishijima’s direct
quote of information on cell dimensions from the Nishiwada
(1895) paper constitutes an indirect reference to valid publication, as indicated in ICBN Art. 32.1 (d), and 32.6 (McNeill et al.
2006).
Age and locality. Late–Early to early–Middle Miocene (Nakamori
et al. 1991); Megami-yama (= Taishaku-san), Megami, Sagara
Town, Haibara County (currently in Makinohara City), Shizuoka Prefecture, central Japan.
Examination of the lectotype. The lectotype thin section is from a
coral-coralline algal packstone that includes several bioclasts.
Two lamellate (flattened) branches of a possibly foliose thallus
constitute the lectotype and are evident in Ishijima (1944, textfig. 7). One branch is a small plant (2 mm long and up to
0.3 mm thick) with a monomerous organization showing coaxial
growth involving a single system of radiating filaments. Cells of
adjacent filaments are connected by cell fusions. The cells are
10–15 lm in diameter and 15–20 lm long. No conceptacles are
EXPLANATION OF PLATE 4
Figs 1–3. Sporolithon kuboiensis (Ishijima) comb. nov. [= Archaeolithothamnion kuboiensis Ishijima (as Archaeolithothamnium)],
lectotype, ‘Archaeolithothamnion kuboiensis n. sp. Ishijima’, Ishijima’s Collection; Miocene, Kuboi, Yamanashi Prefecture, central
Japan. 1, oblique section of the large specimen characterized by several sori composed of numerous compartments. 2, detail of a
sorus illustrating the stalk cells (arrows) at the base of the compartments. 3, plumose ventral core and peripheral filaments with
laterally well-aligned cells. vs, ventral surface; ds, dorsal surface.
Figs 4–5. Sporolithon taiwanensis (Ishijima) comb. nov. [= Archaeolithothamnion taiwanensis Ishijima (as Archaeolithothamnium)],
lectotype, thin section 21222, Ishijima’s Collection; Pliocene, Payapaya, Taitung County, Taiwan. 4, the encrusting thallus with
compartments grouped in a sorus which is located at the thallus surface. 5, detail of figure 4 illustrating the oblique longitudinal
sections of the compartments with thick dark walls.
Fig. 6. Lithophyllum nishiwadai Ishijima, 1944, lectotype, thin section ‘Lithophyllum nishiwadai n. sp.’, Ishijima’s Collection; late–Early
to early–Middle Miocene, Megami-yama, Shizuoka Prefecture, central Japan. Some ventral coaxial cell filaments and their
derivatives curve outwards to form a dorsal peripheral region where a possible aborted branching is present (b). Cell fusions
(arrows) are presents in the ventral core and in the peripheral regions.
Figs 7–8. Lithophyllum otsukiensis Ishijima, 1954, Ishijima’s Collection; Miocene, Otsuki,
Yamanashi Prefecture, central Japan. 7,
abraded lumpy protuberance with buried conceptacles (thin section ‘Lithoporella sp. nov.’, lectotype). 8, tangential section
through a conceptacle with surrounding vegetative thallus (thin section ‘no. 143’, syntype). Note conspicuous cell fusions
(arrows).
Scale bars represent 1 mm in 7, 0.50 mm in 1, 4, and 0.20 mm in 2–3, 6, 8, and 0.10 mm in 5.
PLATE 4
ds
ds
vs
1
vs
2
ds
vs
4
3
b
5
6
7
8
IRYU et al., Sporolithon, Lithophyllum
422
PALAEONTOLOGY, VOLUME 52
present. The second branch (Pl. 4, fig. 6) is a small fragment
(2.2 mm long and up to 0.6 mm thick) consisting of a ventral
core (about 0.25 mm thick) with coaxial filaments, some of
which curve outward to form a dorsal peripheral region
(c. 0.27 mm thick) where a possible aborted branching is present. In both the ventral core and in the peripheral regions the
cells in adjacent filaments are connected by cell fusions. The cells
are 30 lm length and 15 lm in diameter in the ventral core and
15 lm length and 20 lm in diameter in the peripheral region.
No conceptacles are present.
Remarks. The occurrence of cell fusions in the type of
nishiwadai excludes placement of the material in the
Corallinaceae, subfamily Lithophylloideae (which is
characterized by secondary pit-connections rather than
cell fusions), but suggests placement either in the
Hapalidiaceae, subfamily Melobesioideae or in the Corallinaceae, subfamily Mastophoroideae. The absence of
conceptacles, however, precludes any certain placement
at family or genus levels (Table 1). Bellini and Mastrorilli (1975, p. 55) reported Lithophyllum nishiwadai but
did not illustrate the specimens (nor reported information about the occurrence of conceptacles) that they
attributed to this species, and thus their record requires
confirmation.
Lithophyllum otsukiensis Ishijima, 1954
Plate 4, figures 7–8; Text-figures 1Q–R, 3
1954 Lithophyllum otsukiensis Ishijima, p. 44, pl. 27,
figs 1a–b, 4; pl. 28, fig. 1a–b; pl. 30, fig. 1.
Repository data. ‘IGPS coll. cat. no. 79447’ (Ishijima 1954, p.
44).
Age and locality. Miocene; Otsuki,
Kitatsuru County (currently
in City of Otsuki), Yamanashi Prefecture, central Japan.
Examination of the lectotype. The lectotype in the thin section
labelled ‘Lithoporella sp. nov.’ (Text-fig. 1Q; Pl. 4, fig. 7) contains two large coral fragments encrusted by coralline algae and
encrusting foraminifera. The thallus illustrated by Ishijima
(1954, pl. 28, fig. 1a–b) consists of an abraded lumpy protuberance (3 mm length and 2 mm width) belonging to an encrusting
undulated thallus about 35 mm long and 2 mm thick. The
abraded lumpy protuberance is encrusted by an acervulinid foraminifer. Due to the undulation of the plant, the orientation of
the section changes throughout the thallus. The thallus has a
dorsiventral monomerous construction with a well developed
coaxial ventral region (up to 300 lm thick) and a peripheral
region where portions of the ventral filaments curve outward
towards the thallus surface. The cells (8–10 lm in diameter and
20–25 lm long) of adjacent coaxial ventral filaments and those
of the peripheral region (8–10 lm in diameter and 20–25 lm
long) are joined by cell fusions. No trichocytes were seen. No
epithallial cells were recognized. Eight random tangential sections of uniporate conceptacles are present (Text-fig. 3). The
conceptacles protrude conspicuously from surrounding thallus
surface and later become buried; they are subellipsoidal in section with a flat floor, 530–620 lm in diameter and 200–240 lm
in height. The pore canal is conical with the major base 60 lm
in diameter and the minor base 40 lm in diameter; it is 120–
130 lm long. The pore canal is lined by multiple cells of the
roof filaments and may has been rimmed by those cells which
protruded into the canal.
Examination of the syntype. The syntype in the thin section
labelled ‘no. 143’ (Text-fig. 1R; Pl. 4, fig. 8) contains a large
coral fragment encrusted by coralline algal thalli. The plant illustrated by Ishijima (1954, pl. 27, figs 1a–b and 4) is an abraded
and bioeroded lumpy protuberance, about 6 mm long and
Derivation of name. No derivation of the specific epithet otsukiensis was provided in the protologue, but it almost certainly is
derived from the name of the type locality, Otsuki.
Lectotype of Lithophyllum otsukiensis. Ishijima (1954) did not
designate a type, and there are two thin sections, respectively,
labelled ‘Lithoporella sp. nov.’ and ‘no. 143’ with original material illustrated in the protologue (Text-fig. 1Q-R). The material
in thin section ‘Lithoporella sp. nov.’ has better preserved conceptacles with pore canals than the plant in thin section ‘no.
143’. The second thin section labelled as ‘no. 143’ represents the
syntype (as defined in ICBN Art. 9.4). In accordance with ICBN
Art. 9.13 (McNeill et al. 2006), we designate here as lectotype
the material in thin sections ‘Lithoporella sp. nov.’ (Textfig. 1Q). The lectotype was illustrated in the protologue by Ishijima (1954, pl. 28, fig. 1a–b).
A uniporate conceptacle of Lithophyllum
otsukiensis Ishijima, 1954, lectotype, thin section ‘Lithoporella sp.
nov.’. Note a conical pore canal lined by multiple cells of the
roof filaments. Scale bar represents 200 lm.
TEXT-FIG. 3.
Syntype of Lithophyllum otsukiensis. Plant in thin section
labelled ‘no. 143’ (Text-fig. 1R), illustrated by Ishijima in 1954
(pl. 27, figs 1a–b and 4).
IRYU ET AL.: RE-ASSESSMENT OF CORALLINALEAN SPECIES
4 mm thick, which belongs to a plant encrusting the large coral
fragment. This plant is c. 20 mm long and 1 mm thick. The
thallus is dorsiventral monomerous with a thick coaxial ventral
core (up to 280 lm thick) which passes into a peripheral region.
In the ventral part, the cells are 15–20 lm in diameter and 40–
60 lm long; in the peripheral region, the cells are 20 lm length
and 60 lm high. Cells of adjacent filaments are joined by cell
fusions. No epithallial cells are present. Nine uniporate conceptacles randomly sectioned are subellipsoidal in shape (530–
620 lm in diameter and 200–340 lm in height) and become
buried in the thallus surface. Only three tangential-oblique sections show short pore canals (50 lm in diameter and 120 lm in
height; Pl. 4, fig. 8).
Remarks. The occurrence of cell fusions in otsukiensis
excludes its assignment to the Corallinaceae, subfamily
Lithophylloideae and suggests a possible ascription either
to the Hapalidiaceae, subfamily Melobesioideae or to the
Corallinaceae, subfamily Mastophoroideae (e.g. Braga
2003; Harvey et al. 2003a; Braga et al. 2005) (Table 1).
Because the uniporate conceptacles have no contents, it is
not possible to determine whether the plants are tetrasporangial or gametangial. Gametangial conceptacles in both
the Hapalidiaceae (including the subfamily Melobesioideae) and the Corallinaceae (including the subfamily
Mastophoroideae) are uniporate. Genera in both subfamilies possess the features evident in the lectotype, and thus
it is not possible to suggest possible generic placement
other than to rule out those genera known only to have
dimerous construction (e.g. Melobesia and Epulo in the
Melobesioideae; Lithoporella, Mastophora, and Metamastophora in the Mastophoroideae).
CONCLUDING REMARKS
Ishijima’s collections deposited at the Institute of Geology
and Paleontology, Graduate School of Science, Tohoku
University contains the type material of 14 fossil coralline
algal species described by Ishijima from 1942 to 1960.
This collection is a part of the original fossil material of
Ishijima, who established at least 114 fossil coralline algal
species (65 nongeniculate and 49 geniculate coralline
algae) from Eocene to Pleistocene sedimentary successions
in the western Pacific (Iryu 2004).
We have typified 14 species in accordance with ICBN
rules (McNeill et al. 2006) selecting lectotypes from the
original type material. The types are re-described and
illustrated focusing on characters relevant to the modern
taxonomy of fossil corallines. This re-assessment of the
types has resulted in changes of genus and family disposition of seven species whilst one remains within the same
taxon to which was assigned by Ishijima (Table 1). The
species Hydrolithon taishakuensis (Ishijima) comb. nov
(= Lithothamnium taishakuensis), Sporolithon kobamazim-
423
ensis (Ishijima) comb. nov. (= Archaeolithothamnium kobamazimensis), Sporolithon kuboiensis (Ishijima) comb.
nov. (= Archaeolithothamnium kuboiensis) and Sporolithon
taiwanensis (Ishijima) comb. nov. (= Archaeolithothamnium taiwanensis) showed significant features justifying
their use as species names in coralline algal taxonomy.
Lithophyllum quadratum Ishijima is considered as a species of uncertain status within Lithophyllum pending further study. Lithoporella crassa Ishijima and Lithoporella
hayasakai Ishijima are considered heterotypic synonyms
of Lithophyllum prototypum (Foslie) Foslie, while Porolithon hanzawai Ishijima is a heterotypic synonym of
Hydrolithon onkodes (Heydrich) Penrose and Woelkerling.
No diagnostic characters occur in two types (Lithophyllum
nishiwadai, L. otsukiensis) and therefore the taxonomic
disposition of these taxa remains unknown.
The traditional palaeontological practice of referring all
fossil specimens with multiporate conceptacles, cell
fusions, and at least some coaxial growth to Mesophyllum
should be continued for the present. Mesophyllum arakuraensis Ishijima and M. contii Ishijima dealt with here
under Mesophyllum are known only from fossil material.
We cannot determine from available evidence whether
either might belong to Synarthrophyton instead of Mesophyllum. The relationships of M. arakuraensis Ishijima to
nonfossil species of Mesophyllum cannot be determined
from features evident in the lectotype or isolectotype.
Comparisons with the fossil species M. contii Ishijima are
provided in the account of that species. These two taxa
are considered conspecific and M. contii is a heterotypic
synonym of M. arakuraensis. The absence of preserved
epithallial cells precludes any certain generic assignment
of Lithothamnion ishizuchiensis Ishijima and L. kasedaensis
Ishijima. There are no other features evident in Ishijima’s
specimens that can be used to ally his species to any particular genus or species of Melobesioideae.
These results demonstrate that many fossil coralline
algal taxa were established based on features which are
not considered diagnostic in modern taxonomy. The
results also suggest the necessity of (1) re-assessing the
precise nature of other coralline algal taxa by revising
the original collections of Ishijima and of (2) striving to
interpret the taxonomy of this calcareous algal group
using the same classification system for both fossil and
nonfossil taxa.
Acknowledgements. We are grateful to K. Kurihara (Rikkyo University), H. Tanimura and K. Uemura (National Museum of
Nature and Science, Japan) for giving us the opportunity to
examine the coralline algal collections of W. Ishijima. Our investigation was financially supported in part by Grants-in-Aid for
Scientific Research, Japan Society for the Promotion of Science
(18340163 to Y. I.), by the 21st Century Center-of-Excellence
Program, ‘Advanced Science and Technology Center for the
424
PALAEONTOLOGY, VOLUME 52
Dynamic Earth’, at the Tohoku University, and by local research
funds at the University of Ferrara. Two anonymous reviewers
are thanked for their valuable comments and suggestions on the
paper. We also thank Chris Berry for his thoughtful editing of
the manuscript.
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