MICROSCOPY RESEARCH AND TECHNIQUE 33:2-11 (1996)

Alveus, gen. nov. (Bacillariaceae, Bacillariophyta), a Heavily

Silicified Diatom Found in Warm Water Oceans
IRENA KACZMARSKA AND GRETA A. FRYXELL
Department of Biology, Mount Allison University, Sackville, New Brunswick, EOA 3CO Canada; Department of Oceanography,
Texas A&M University, College Station, TX 778433146

KEY WORDS New genus, SEM, Systematics, Pattern of distribution, Silica export
ABSTRACT Scanning electron microscope images of complete and damaged frustules of Nitzs-
chia marina Grun. revealed an unusual chambered structure of the valve. This structure is not
present in any existing nitzschioid genus and warrants an independent generic status for this
taxon, which is therefore separated into Alueus gen. nov. Generic and emended specific definitions
are provided and presented in the context of their relationship to other genera within the family
Bacillariaceae. Deep relief of the inner surface of Alueus valves is similar to that of genera Fragi-
lariopsis, Denticulopsis, Neodenticula, and Crucidenticula. It differs from all these diatoms in the
alveolar striation. This heavily silicified diatom was found in samples from the US Joint Global
Oceanographic Flux Studies cruise "TO07 stations 2 and 4, (9" and 5"N respectively, 140"W)in the
equatorial Pacific. Presently, this uncommon species is restricted to warm waters. It is very com-
mon in the surface sediments of the equatorial Pacific and Indian Ocean and in the tropical and
subtropical Atlantic. The sedimentary record of Alueus marinus goes back to the Middle Late
Miocene (approximately 8 Ma years). o 1996 Wiley-Liss, Inc.
NOMENCLATURAL HISTORY OF THE TAXON Simonsen (1974)pointed out the unusual structure of
The taxon Nitzschia angustata var. marina was first the valve of Nitzschia marina detected through light
recognized by Grunow (Cleve and Moller, 1878) when microscope (LM) observations. He noted that the striae
identifying diatoms from the Balearic Islands, Medi- are chambered "much like in Pinnularia" (Simonsen,
terranean Sea (exsiccata, Cleve and Moller, 1878, 1974, p. 53) and occluded from the inside by a thin
slides 154 and 155), collected by Dr. F. Siiderlund. It is membrane. The membrane is perforated by small sub-
listed as one of 173 diatom species present on these marginal openings through which the chamber com-
slides (Cleve and Moller, 1878, pp. 6,7). Later, referring municates with the remainder of the interior of cell.
to the Balearic taxon as separate from the freshwater Our scanning electron microscope (SEMI observations
N . augustata, Grunow (Cleve and Grunow, 1880) made of frustules as well as two micrographs presented by
it a distinct species, Nitzschia marina, which he subse- Akiba and Yanagisava (1986) confirmed the accuracy
quently illustrated in Van Heurck (1881).The volume of Simonsen's observations. However, the taxonomic
of this publication housed in the Museum of Natural affinity of this diatom was outside the major thrust of
History in Vienna carries Grunow's handwritten name Akiba and Yanagisava's presentation (1986). Subse-
label on plate 57, Figures 26-27, which may be re- quently, we examined damaged and dissolving valves
garded as an iconotype. Unfortunately, Grunow's col- and expanded our understanding of the valve internal
lection which is also housed by this institution does not architecture.
possess a designated type of the species. Moreover, N . The goal of this paper is to document the unusual
marina is not listed in any of Grunow's slides (Riedl, structure of N . marina valves, which supports the sep-
personal communication, 1994). arate generic status. The diatom apparently has not
Independently, Heiden (Heiden and Kolbe, 1928) de- been examined in full detail using SEM, and conse-
scribed a species, Synedra gaussii, from equatorial and quently the taxonomic significance of the valve struc-
southern Atlantic waters, which was later put into syn- ture was not fully appreciated.
onymy with Nitzschia marina by Kolbe (1954) and re- MATERIALS AND METHODS
cently confirmed by Simonsen (1992) in his typification Water and net samples examined were collected by
of Heiden's taxa. In addition, Heiden described the va- R.V. Thompson Survey I Cruise TT007 in early spring
riety S . gaussii var. trigibba Heiden. Following Kolbe's 1992 at stations 2 and 4 (9ON-5"N and 140"W). Sam-
conclusion on the conspecificity of S. gaussii and N . pling methods were described in Murray et al. (1992)
marina, Simonsen (1992) proposed retaining the taxon
at the forma level by making this new combination, N .
marina forma trigibba (Heiden) Simonsen. W e agree
that it is useful to retain a separate form status for this Received January 26, 1994; accepted in revised form August 31, 1994.
diatom until more is known about the range of its mor- Address reprint requests to Irena Kaczmarska, Department of Biology, Mount
phological variation. Allison University, Sackville, New Brunswick, EOA 3CO Canada.

0 1996 WILEY-LISS. INC.

 GENUS ALVEUS
and Kaczmarska and Fryxell (in press). Cell densities
were enumerated by Utermohl technique (Kaczmarska
and Fryxell, in press), while biomass contribution was
calculated by cell volume method (Liu and Kaczmar-
ska, submitted).
Diatom frustules from net hauls were cleaned by
methods proposed by Hasle and Fryxell(1970) and Si-
monsen (1974), mounted in Hyrax, and examined in an
inverted ZEISS ICM-405 microscope (LM). Cleaned
frustules were mounted on aluminum stubs, each with
a glass coverslip affixed to its tops for SEM examina-
tion. The cleaned samples were sputter-coated with 20
nm gold and observed in a JEOL 6400 SEM (Texas
A&M University Electron Microscopy Center) a t an
accelerating voltage of 15 kV and a working distance of
8 mm. To demonstrate the maximum range of specific
variability, measurements of the specimens found in
the North Atlantic Warm Core Rings (Kaczmarska et
al., 1986) were also included in an emended species
description.
DESCRIPTION OF THE NEW GENUS AND
EXAMINED MATERIAL
ALueus Kaczmarska and G. Fryxell, gen. nov.
Frustula symmetria nitzschioidea. Raphe continua,
non elevata super superficiem valvae, moderate asym-
metrica. Copulae apertae. Striae alveolatae. Alveola
omnium striarum in limbo extendit.
Frustules with nitzschioid symmetry, elongate. Ra-
phe continuous, not elevated above the valve face sur-
face, moderately asymmetrical. Copulae open. Striae
alveolate, the chambered architecture of all striae con-
tinuous onto the mantle.
Diagnosis: Genus striis alveolatis a confamiliis di-
versa.
Etymology: Refers to the hollow (alveolar) structure
of striae.
Generitype: Alueus marinus (Grunow) Kaczmarska
et G. Fryxell, comb. nov.
Alueus marinus (Grunow) Kaczmarska
et G. Fryxell
Basionym: Nitzschiu marina Grunow (Cleve and
Grunow, 1880, p. 70). Nitzschiu angustutu var. marina
Grunow in Cleve and Moller (1878; page 6, slide 154,
1551, nomen nudum.
Synonym: Synedra gaussii Heiden (Heiden and
Kolbe, 1928).
Lectotype: Cleve and Moller (1878, (exsiccata) slide
154, Natural History Museum in Vienna).
Iconotype: Cleve and Grunow (1880, Plate 57, Fig.
26).
Collection Material and Earlier Descriptions
We have examined isosyntypic material (Cleve and
Moller, 1878, exsiccata) from the Balearic Islands
housed by the Natural History Museum in Vienna, the
Natural History Museum in London, and the Swedish
Museum of Natural History. The greatest range of
variation in size, shape, and ornamentation was found
on slide 154 from the B.M. Adams collection housed at
the Natural History Museum in London. This slide was
3
also best suitable for photomicrography. We found on
this slide a total of 11 complete or nearly complete
valves lying in the valve view, two apical fragments,
and one frustule with two girdle bands attached in gir-
dle view. These specimens (Figs. 1-19) were 98-160
pm long and 7.5-11 pm wide with 11-12 striae and
had 9-10 fibulae in 10 pm. Other key features can be
noted the eccentric raphe (Figs. 1,8), the rectangular
girdle view (Figs. 9, lo), the great range of size (Figs. 1,
7), and the coarse structure (Figs. 17, 18). Each stria
consisted of two rows of poroids which are barely re-
solvable in LM, with 18-19 poroids in 10 pm (Figs. 8,
11,13,14,16,19).Shorter valves were lanceolate (Figs.
1-3,5,12,13); longer valves were more linear (Figs. 6,
7). An apparently aberrant, lunate valve was also en-
countered (Figs. 4,111, contrasting with the linear lan-
ceolate shape (Fig. 12). Internal apertures of the cham-
bered striae noted by Simonsen (1974) were detectable
(Figs. 8, 13-16, 19). These Balearic specimens were
never formally described. Consequently, the name
Nitzschia angustata var. marina should be considered
nomen nudum.
Later, after examining his Atlantic material, Gru-
now in Cleve and Grunow (1880) recognized Nitzschia
angustata var. marina (erroneously spelled as N . an-
Figs. 1-19. (See page 4.) All specimens in Figs. 1-19 come from
the 1878 Cleve and Moller slide 155 representing collection material
from the Balearic Islands. The slide is in the B.M. Adams Collection
housed at the Natural History Museum, London.
Figs. 1-7. LM. Valves in the face view, variation of the shape and
size. Figs. 1, 2: Note curvature of raphe near poles. ~ 8 0 0 Fig.
. 3:
Slightly oblique view showing mantle adjacent to raphe. x 800. Fig. 4:
Curved, lunate valve with asymmetric margins. ~ 7 8 0 .Fig. 5:
Slightly curved valve. X 800. Fig. 6 Valve with almost parallel sides,
tapered at poles. x 580. Fig. 7: Oblique view of valve with rounded
poles. ~ 5 7 0 .
Fig. 8. LM. The position of the external apical raphe ending. Note
also poroids of striae. x 1,200.
Figs. 9,lO. LM. Valve and cingulum in girdle view. Fig. 9 focuses
on the mantle. Fig. 10 focuses on cingulum. x 800.
Fig. 11. LM. The same valve as in Fig. 4. Almost complete lunate
valve, arrowheads pointing to the row of internal apertures of the
chambered striae. x 1,240.
Figs. 12, 13. LM. The same valve as in Fig. 1. Fig. 12 shows a
continuous raphe system. Fig. 13 shows biseriate poroids of the ex-
ternal surface of striae. x 1,170.
Fig. 14. LM. Portion of the same valve as in Fig. 4, showing the
raphe (let%),biseriate striae, and internal apertures of chambered
striae (right). x 1,900.
Fig. 15. LM. Internal apertures of chambered striae (arrowheads).
x 2,000.
Figs. 16, 17. LM. Continuous raphe, biseriate striae (Fig. 17) and
the internal apertures of the chambered striae (Fig. 16, arrowheads).
x 1,210.
Figs. 18, 19. LM. Fig. 18 focuses on the raphe side of the valve
showing misalignment of striae on two sides of the raphe, Fig. 19 on
the internal apertures of the chambered striae (arrowhead) and bi-
seriate striae. x 1,160.
4 I. KACZMARSKA AND G.A. FRYXELL

Figs. 1-19. (Legend appears on page 3.)

 GENUS ALVEUS 5

Figs. 20-25. (Legend appears on page 6.)

6 I. KACZMARSKA AND G.A. FRYXELL
gusta var. marina, p. 70) as an independent species and
named it Nitzschia marina. His description gave the
following characteristics: valves 95-165 pm long and
8-9 pm wide, bearing 12 striae in 10 pm and 19 por-
oids in 10 pm within a striae. Although the original
description is very brief, the diatom is easily recogniz-
able in LM. Heiden and Kolbe (1928) collected the spe-
cies along an Atlantic transect spanning from 8"N to
35"s and from 12"W to 12"E. This material demon-
strates the greatest morphological variation thus far
observed. A new form, S. gaussii f. trigibba, was found
in this collection. The specimens conforming to the typ-
ical morphology were described as follows: 53-353 pm
long and 8-12 pm wide, with 10-12 striae in 10 p and
hstinctly decussate punctae of the striae. The trigib-
bous form differs from the typical form only in the out-
line of the valve margin.
Authors' Material
In the Pacific samples examined, the frustules were
almost always solitary. On only one occasion did we find
three cells joined by their valve faces. As the material
we examined was preserved, we restrain from comment
on the cellular content. The frustules were rectangular
in girdle view (Figs. 23, 25), 65-252 pm long and 6-9
pm wide, and with pervalval axis of 12-18 pm deep. In
four frustules examined in SEM the raphe systems were
located diagonally on the frustule, thus demonstrating
nitzschioid symmetry. Copulae were open at one apex
with a short and broad ligula a t the other end (Fig. 22,
arrowhead); no ornamentation was discernible on them
(Fig. 25). In the epicingulum there were three wide
copulae and possibly one narrow band a t the abvalvar
end (Fig. 25). The long valves were linear through most
of their length (Fig. 20); the short valves were lanceo-
late. Ends were shortly tapering and broadly rounded
(Figs. 21, 24). The mantle was deep, joining the valve
face a t an obtuse angle (Figs. 26, 35). The raphe was
continuous, not elevated above the valve face (Figs. 21,
24, 25), moderately eccentric, and closely paralleling
one margin (Figs. 20,21,24).The external apical raphe
Figs. 20-25. (See page 5.) All SEM figures represent specimens
from the US JGOFS TT007 station 2.
Fig. 20. SEM. Entire valve, outside view. x 1,100.
Fig. 21. SEM. Bottom apex of the valve from Fig. 20, showing the
external apical raphe ending and biseriate striae. Note shallow de-
pression running along the marginal ridge (arrowhead). x 4,200.
Fig. 22. SEM. A copula, open at one end and carrying a short stout
ligula at the other. Arrowhead points toward the ligula which is ob-
scured in this orientation. x 6,600.
Fig. 23. SEM. Girdle view of a frustule. x 6,600.
Fig. 24. SEM. Top valve apex of Fig. 20, showing external apical
raphe ending, with widened fissure and biseriate striae. Note contin-
uous shallow depression from Fig. 21 (arrowhead). x 4,300.
Fig. 25. SEM. Apical part of the frustule from Fig. 23, showing
raphe fissure (arrowhead)arrangement and morphology of three wide
copulae of the epicingulum and a putative fourth, narrow copula in
the abvalvar position (arrowhead). X 3,000.
endings were bent toward the middle of the valve apex
and terminated bluntly in widened fissures (Figs. 21,
24). Internally, the apical raphe endings terminated in
straight helictoglossae (Figs. 27,281. The valve face and
mantle bear biseriate striae on most of their external
surface (Figs. 20,21,24). These striae were interrupted
by a solid siliceous ridge running along the valve face
margin on the side opposite the raphe (Figs. 21,24,33,
34,351. The valve surface was partially depressed into
a shallow groove running close to this ridge (Figs. 20,
21,24).A low elevation corresponding to this depression
would likely be present in the sibling valve. Striae were
alveolate. The alveoles continued into the mantle (Figs.
30 and 21 arrowheads, 33-35). Alveoles opened exter-
nally through a row of large alternating poroids: 2-2.5
in 1 pm (Figs. 33-35). The number of alveolate striae
was unequal on the two sides of the raphe, being 8-10
striae in 10 pm on the narrower side, a number corre-
sponding to the number of fibulae (Figs. 26, 29-31).
These striae opened to the valve interior through por-
tulae (Figs. 29,32 arrowheads, 33). On the wider side of
the valve, striae were more numerous, 9-14 in 10 pm.
They opened to the valve interior by elliptical apertures
(Figs. 26 arrowhead, 30-32,33 arrow) along the valve's
margin.
Many features which are interpretable only through
an understanding of SEM micrographs can be seen in
the light microscope. The internal apertures of the
chambered striae on the wide side of the valve face are
visible in some of our specimens (Figs. 11, 15, 16) as
well as those examined by Simonsen (1974). A combi-
nation of the chambered striae (Fig. 29) and the asso-
ciated external longitudinal depression along with
margin ridge (Figs. 21,24) gives rise to the distinct line
on the specimens seen in LM (Figs. 1, 2 , 4 , 11-16, 18,
19).The biseriate nature of the striae is very difficult to
discern in the aged slides, although some micrographs
suggest this feature (Fig. 14). Although the raphe was
not elevated in the specimens examined in SEM (Figs.
21, 24, 251, light micrographs show that its position
generally corresponds with the longitudinal depression
on the sibling valve (Figs. 8, 11-16, 18, 19). A nonde-
pressed part of the valve (along with the raphe) may
therefore appear to be slightly elevated with respect to
the remaining valve surface.
TAXONOMIC AFFILIATION
The genus Nitzschia (Hustedt, 1930; Krammer and
Lange-Bertalot, 1988; VanLandingham, 1967-1979) is
one of the largest genera of diatoms and includes ap-
proximately 900 species (VanLandingham, 1967-1979)
whose frustule morphology demonstrates great diver-
sity. With thorough electron microscopical investiga-
tions, the intrageneric systematics of the genus Nitzs-
chiu has been recently under critical review (Hasle,
1993; Mann, 1986, personal communication). A new
concept of the genus based on the works of Mann
(19861, Round et al. (19901, and Hasle (1993) clearly
indicate the heterogeneity of this genus. Old genera
have been reinstated (e.g., Tryblionella W. Smith,
Fragilariopsis Hustedt, Pseudonitzschia H. Peragallo,
Gomphonitzschiu Grunow). New genera have been re-
cently established (e.g., Psammodictyon Mann, Gom-
 GENUS ALVEUS 7

Figs. 26-31. All SEM figures represent specimens from the US
JGOFS TT007 station 2.
Fig. 26. SEM. Inner surface of a complete valve shows fibulae
(arrow) and apertures of chambered striae (arrowhead). x 1,500.
Figs. 27,28. SEM. Internal apical raphe ending with helictoglossa
(arrows). x 15,000.
Fig. 29. SEM. Internal view of valve; raphe fissure located under-
neath fibulae between arrows. x 8,000.
Figs. 30, 31. SEM. Cross-section showing that the chambered
striae extends to the mantle which is proximal to the raphe (Fig. 30,
arrowheads) as well as to the mantle which is distal to the raphe (Fig.
31, arrowheads). Cross-section view of poroids (two small arrows point
to poroids detectable in this region) and fibulae (large arrow). Fig. 30:
x 6,000; Fig. 31: x 5,370.
8 I. KACZMARSKA AND G.A. FRYXELL
Figs. 32-35. All SEM figures represent specimens from the US
JGOFS n o 0 7 station 2.
Fig. 32. SEM. Incomplete valve with the external part missing.
Internal view with the siliceous plate forming the bottom of the cham-
bered striae. Arrowheads point to portulae of fibulae; the lower one is
partially damaged and exposes connection, via raphe canal, between
the chambered stria of the narrow part of the valve and the cell in-
terior. A row of apertures connecting chambered striae of the wide
part of the valve to the cell interior runs along the opposite margin.
~5,000.
Fig. 33. SEM. External view with the siliceousplate at the bottom
of chambered striae. The undamaged plate ascends laterally towards
the raphe and separates the chambered striae from it (black-and-
white arrowheads). The black arrowhead points to the openings of the
chambered striae into the raphe channel. x 8,000.
Fig. 34. SEM. External surface of the valve face showing a dis-
continuity of the areolation between valve face and mantle (arrow-
head) and an alternating biseriate pattern of areoles. x 10,000.
Fig. 35. SEM. Cross-section of a valve showing the position of the
mantle relative to the valve face and the depth of the transapical ribs
separating the chambered striae. x 10,000.
 GENUS ALVEUS
photheca Hendey and Sims, and Simonsenia Lange-
Bertalot) which would conform to the earlier concepts
of the genus Nitzschia (Hustedt, 1930; Krammer and
Lange-Bertalot, 1988). Alveus differs from all these
genera by having chambered striae.
Heavily silicified valves of several diatom species
comprising the genus Fragilariopsis (most notably
Fragilariopsis kerguelensis) bear superficial resem-
blance to the genus Alveus in their external ornamen-
tation, in particular in their biseriate areolation. How-
ever, their valves are unilayered. Thus the striae are
not alveolate, the mantle and transapical ribs are gen-
erally shallow, and when elevated (e.g., F. kerguelensis
(O'Meara) Hust.) the transapical ribs possess locking
projections, which facilitate the fit of the valvecopula
margin and the ribs similar to those present in the
denticuloid genera (Kaczmarska, unpublished data).
Denticuloid diatoms possess deep transapical ribs
(pseudosepta) between the striae. The marine genera
Denticulopsis (Simonsen) Akiba and Yanagisava, Neo-
denticula Akiba and Yanagisava, and Crucidenticula
Akiba and Yanagisava (all from family Bacillariaceae,
Akiba and Yanagisava (1986) implicitly) possess ex-
tensive systems of pseudosepta, crossbars, and mar-
ginal ribs projecting from the valve to the cell interior
(Akiba and Yanagisava, 1986; Simonsen and Kanaya,
1961).In addition, they differ from Alueus in the struc-
ture of areolae and the complex, septate structure of
the girdle.
The genus Alveus shares the nitzschioid symmetry
of the frustule with almost all members of the family
Bacillariaceae (Round et al., 1990) but differs from all
in the alveolar structure of the striae. We speculate
that the chambered striae could have developed from
the deep internal transapical ribs running between
every two rows of areolae. The free, distal ends of the
ribs were subsequently joined by a siliceous plate
parallel to the valve face. The plate enclosed the
chambers from the cell interior, as shown on the wide
side of the valve face. On the narrow side of the valve
face, the number of chambers was reduced to the
number of fibulae, while the basic alveolar structure of
the striae was preserved. None of the 15 genera
included in the Bacillariaceae (Round et al., 1990) and
two genera discovered by Akiba and Yanagisava
(1986) seen in SEM demonstrates a similar complexity
of the striae.
In the original description of the taxon, Grunow
(Cleve and Grunow, 1880) included Nitzschia marina
in the section Tryblionella within the genus Nitzschia.
The section was defined by a very eccentric raphe sys-
tem, valves with undulate surface, and indistinct fibu-
lae. Based on the fine structure of frustules as revealed
in SEM, Tryblionella has been recently reinstated as a
genus (Round et al., 1990). Using SEM these authors
documented structure of the generitype, Tryblionella
acuminata W. Smith. In T . acuminata, the valve is
monlayered and the striae are simple. Each stria con-
sists of a shallow groove perforated by a row of poroids.
The fibulae are distinct but very short. In addition, the
raphe is keeled and very eccentric in Tryblionella
(Round et al., 1990). Based on these observations,
Alveus rnarinus cannot belong to the genus Try-
9
blionella since it is lacking almost all of the diagnostic
characteristics.
Chambered striae similar to those in Alveus are
known from several families of diatoms. Long alveolar
striae, extending through the part of the valve face, are
present in Pinnulariaceae (Pinnularia Ehrenberg),
Scoliotropidaceae (Scoliopleura Grunow, Scoliotropis
Cleve, Biremis Mann, and Progonia H. Schrader),
Fragilariaceae (Catacombas Williams and Round, Pter-
oncola Holmes and Croll), Ardissoneaceae (Ardissonea
De Notaris), and Endopylaceae (Gephyria Arnott). The
first two families belong to the order Naviculales,
while the last three belong to the order Fragilariales
(Round et al., 1990), diatoms distinct from Alveus in
the frustule symmetry and the lack of raphe, respec-
tively. There are also numerous genera with striae con-
sisting of a series of loculate areolae which may be
viewed as an alveolar striae divided into numerous
smaller compartments-for example, Dictyoneis Cleve,
Diploneis Ehrenberg, Stauropsis Meunier, or Psam-
modictyon. Among these, only the last genus belongs to
the family Bacillariaceae. It differs from Alveus in that
it has a keeled raphe system and simple striae.
SPATIAL AND TEMPORAL DISTRIBUTION
Alveus rnarinus was found in several samples of the
US JGOFS Cruise TT007 Stations 2 and 4 a t depths of
60 and 3 m as well as in plankton net hauls. Its relative
frequency was 0.15% at depth 3 and 0.81% at depth 60
m of the micro-phytoplankton. Its biomass was esti-
mated to be 2.65 x 10T3-2.65 x pg CI1-I and
contributed 0.92-2,140 x of the total organic
carbon found at these depths. Thus, it was a minor
contributor to the diatom flora and to biomass in our
samples.
We also found this diatom in the Warm Core Rings of
the North Atlantic (Kaczmarska et al., 19861, where it
was rare in net hauls from the Sargasso Sea, Gulf
Stream waters, and Rings Center in the fall of 1981
and 1982, a t the locations between 35"41'-40"25'N and
63"25'-71"55'W. Equatorial and southern Atlantic as
well as Mediterranean records of occurrence are pro-
vided by Cleve and Grunow (1880), Heiden and Kolbe
(19281, and Hustedt (1958). Thus, the contemporary
distribution of the species spans from 40"N to 40"s in
the Atlantic Ocean. Simonsen (1974) and Cleve and
Grunow (1880)found the species in numerous locations
in the subtropical and tropical Indian Ocean and Ara-
bian Sea. In the equatorial Pacific Alueus marinus has
not been reported from the water column in earlier
quantitative studies (Fryxell et al., 1979; Hasle, 1959,
1960). However, it was reported from the southern Pa-
cific by Cleve and Grunow (1880). Interestingly, both
earliest reports (Cleve and Grunow, 1880; Heiden and
Kolbe, 1928) found both varieties of the species in the
gut of salps. A . marinus f. trigibba may outnumber the
type variety in the salp gut. In plankton samples A .
marinus is usually reported as rare or uncommon.
Alveus rnarinus is more frequently seen in the sedi-
ments. In the surface (bottom) and Quaternary sedi-
ments of the Equatorial Pacific, Atlantic and the In-
dian Ocean it is reported as one of the most common or
even dominant diatom (Jouse et al., 1971; Kolbe, 1954,
10 I. KACZMARSKA AND G.A. FRYXELL
1955, 1957). The abundant occurrence is confined to
the narrow equatorial band of siliceous bottom sedi-
ments (approximately 10°N-lO"S) in the Pacific and
Indian Oceans and in a broader band (30°N-18"S) in
the Atlantic. Other heavy silicified diatoms also re-
ported from the equatorial Pacific surface sediments
not far from the site of our collection, such as Azpeitia
nodulifer (Schmidt) G . Fryxell & P.A. Sims, Plank-
toniella sol (Wallich) Schutt, Thalassiosira oestrupii
(Ostenfeld) Proshkina-Lavrenko, and Thalassiothrh
spp. (JOUS~ et al., 1971), were found more frequently in
our samples than Alveus marinus. All these diatoms
are apparently established, although normally not nu-
merous, members of the phytoplankton community in
the equatorial Pacific. Heavily silicified diatom frus-
tules are more likely to escape dissolution after cell
death and during sinking to the sea bottom, where they
are often overrepresented in the surface sediments
when compared to their abundance in the water col-
umn (e.g., Fenner et al., 1976;Jouse et al., 1971). Other
diatoms that are common and frequent components of
regional phytoplankton communities may be found in
the sediments in such quantities that they impair drill-
ing operations, as in the case of microlayers contain-
ing valves of ThalassiothrhlThalassionemacomplex
(Kemp and Baldauf, 1993). Both Alveus and Thalassio-
thrixlThulassionema spp. may have been abundant in
the sediments as a result of selective preservation, as a
result of unusually favorable growth conditions or
physical accumulation such as occasionally reported
for other species (e.g., Rhizosolenk mats). In this con-
text Alveus, like other heavily silicified diatoms, may
contribute to the export of silica from the photic zone
and thus may play a significant role in the oceanic
silica budget. Accumulation of Alveus marinus in the
sediments has been well documented; however it is not
known what oceanographic and/or autecological condi-
tions are necessary or sufficient for it to occur. Better
understanding of the autecology of Alveus marinus, in
particular with respect to the growth requirements and
degree of silicification, may allow us to quantify its role
in the local and global oceanic cycling of silica.
Alveus marinus is known since the Middle Late Mi-
ocene (approximately 8 million (Ma) years ago). It has
been reported from the Pacific, Indian, and Atlantic
Oceans (Akiba and Yanagisava, 1986; Barron, 1985;
Burckle, 1972; Kaczmarska, 1990; Schrader, 1973,
1974). The Miocene valves of A . marinus retrieved
from the core RC12-65 (04"N, 144"W; Burckle, 1972)
and illustrated by Akiba and Yanagisava (1986) dem-
onstrate the same internal structure as observed in our
material. Occurrence in the Neogene sediments of the
northeastern Pacific (Akiba and Yanagisava, 1986;
Schrader, 1973) is of particular interest because there
it coexists with possibly related extinct taxa. Schrader
(1973) found it together with N . reinholdii Kanaya et
Koizumi, N . praereinholdii Schrader, N . heteropolica
Schrader, and N . pruefossilis Schrader (Schrader, 1973)
and stipulated a close relationship among the species.
Phylogenetic relationship among all but the first of
these species and Alveus marinus has been questioned
bv Simonsen (1974) and reauires further examination.
SLveral other 'Miocene and Pliocene species, such as N .
jouseana Burckle, N . mioceanica Burckle, and N . plio-
ceana (Brun) Mertz which occur mainly in low-latitude
sediments, also demonstrate affinity to A . marinus in
the LM and SEM morphology of the complete valves
(Akiba and Yanagisava, 1986). Planned detailed SEM
examination of the broken valves of these taxa may
elucidate their taxonomic affinity and phylogenetic re-
lationship.
ACKNOWLEDGMENTS
We thank L. Welling for taking samples on US
JGOFS cruise TT007 and Dr. J.W. Murray, the chief
scientist, and S. Kadar for excellent support. Ms. K.
Childs of the Natural History Museum in London and
Dr. H. Riedl of the Natural History Museum in Vienna
and the staff of the Swedish Museum of Natural His-
tory provided invaluable assistance in search for the
type material and with Cleve and Moller exsiccata. Dr.
M. Poulin of the Canadian Museum of Nature helped
with nomenclatural history.
Drs. H. vanderLeest and P. Fryxell kindly translated
diagnosis to Latin, and J. Ehrman expertly helped with
SEM operations. Liu Duan allowed us to use his un-
published estimations of the organic carbon and bio-
mass of Alveus marinus cells. Mr. B.P. Shaw provided
skillful laboratory assistance. We also thank Ms. L.
McIntyre for her darkroom work and assembling the
plates. Dr. J. Johansen's and two anonymous review-
er's comments were very helpful in improving the final
version of the manuscript. The support for this study
was provided by Marjorie Bell Faculty Sabbatical
Leave Fund (I.K.) and NSF grant OCE 91-15605
(G.A.F.).
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