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Two new species of Caloglossa (Delesseriaceae, Rhodophyta) from the Americas,
C. confusa and C. fluviatilis spp. nov.
DAVID M. KRAYESKY1*, JAMES N. NORRIS
2, JOHN A. WEST3, MITSUNOBU KAMIYA
4, MATT VIGUERIE5, BRIAN S. WYSOR
6AND
SUZANNE FREDERICQ5
1Department of Biology, Slippery Rock University, Slippery Rock, PA 16057, USA2Department of Botany, Smithsonian Institution, Washington, DC 20013-7012, USA
3School of Botany, University of Melbourne, Parkville, Victoria 3010, Australia4Department of Marine Bioscience, Fukui Prefectural University, 1-1 Gakuencho, Obama, Fukui 917-0003, Japan
5Department of Biology, University of Louisiana at Lafayette, Lafayette, LA 70504-2451, USA6Department of Biology and Marine Biology, Roger Williams University, Bristol, R, 02809, USA
KRAYESKY D.M., NORRIS J.N., WEST J.A., KAMIYA M., VIGUERIE M., WYSOR B.S. AND FREDERICQ S. 2012. Two newspecies of Caloglossa (Delesseriaceae, Rhodophyta) from the Americas, C. confusa sp. nov. and C. fluviatilis sp. nov.Phycologia 51: 513–530. DOI: 10.2216/11-57.1
Although the systematics of Caloglossa has been intensely investigated, the species diversity of the genus is not yet fullyunderstood. Comparative chloroplast-encoded rbcL and nuclear LSU rDNA sequence analyses, combined withmorphological observations, reveal new records and two new species for the Americas. The species, previously identifiedas Caloglossa monosticha from the western Atlantic, is newly described as C. confusa sp. nov. It differs from PacificOcean C. monosticha primarily by its strongly constricted thallus nodes, and the sequence data presented here show thatit is most closely allied with other taxa of the C. continua species complex, but its relationship with these taxa is not fullyresolved. The freshwater C. fluviatilis sp. nov. from the Panama Canal is characterised by thalli arranged in tufts,adventitious branching, strong constrictions at the thallus nodes and lanceolate blades. Caloglossa fluviatilis forms amonophyletic group along with the species having only adventitious secondary branching. A dichotomous keyhighlights the diagnostic vegetative characters for separating the nine species of Caloglossa occurring in the Americas.
KEY WORDS: Atlantic Ocean, Caloglossa, Ceramiales, Delesseriaceae, LSU rDNA, Pacific Ocean, Panama Canal, rbcL,Rhodophyta, Systematics
INTRODUCTION
Caloglossa (Harvey) G. Martens (1869) is a euryhaline red
algal genus of shallow-water species that occur in man-
groves, saltwater/freshwater marshes, lagoons and rivers in
temperate and tropical regions of the world. The genus is
characterised by the isomorphic gametophytes and tetra-
sporophytes; thalli composed of thin, strap-shaped blades
that are monostromatic except for the midrib region; and
exogenous branching in which lateral branches are ini-
tiated through oblique division of subapical cells. The co-
occurrence of Caloglossa with members of other red
algae, such as Bostrychia Montagne, Catenella Greville
and Murrayella F. Schmitz, in mangroves was first noted
and extensively studied by Post. This ecological association
or assemblage of genera was referred to by Post (1936) as
the ‘Bostrychietum’, or the Bostrychia–Caloglossa associa-
tion. Within the polyphyletic Delesseriaceae Bory, Calo-
glossa is placed in the Caloglosseae M.J. Wynne, a tribe of
two genera that also includes Taenioma J. Agardh (Lin et
al. 2001; Wynne 2001; Choi et al. 2002).
Caloglossa is the most widespread genus of the Delesser-
iaceae (Papenfuss 1961), with most species being only
locally common within their reported range. Some species
distributions are more or less pantropical while others are
restricted. The genus has been the focus of considerable
taxonomic and phylogenetic research globally (Post 1936;
King & Puttock 1994; Kamiya 2004) and regionally (West
et al. 1994; Kamiya et al. 1997, 1999, 2003; Krayesky et al.
2011). These studies show that Caloglossa contains about
15 species. Taxonomically important morphological char-
acters have included nodal anatomy of the thallus, degree
of constriction and organization of cell row lineages, rhi-
zoidal anatomy/distribution, endogenous or adventitious
type of branching and internode morphology (for a review,
see King & Puttock 1994).
The taxonomic importance of rhizoidal anatomy/distri-
bution was further elucidated by Kamiya et al. (2003) with
the description of rhizoidal position types. Rhizoidal types
in the Americas include type C (rhizoids are derived from
ventral pericentral cells in the nodal region two to three
pericentral cells above and below the node, in addition to
an occasional ventral wing cells that directly flanks one
of these pericentral cells), type E (rhizoids derived from
ventral pericentral cells around the region of the node in-
cluding ventral node pericentral cell and the lateral peri-
central cells that flank it), type F (rhizoids derived from
ventral pericentral cells around the region of the node
above the ventral node of a pericentral cell, in addition to
ventral wing cells located directly around the nodal region),
and type G (rhizoids derived from ventral pericentral cells* Corresponding author ([email protected])
Phycologia (2012) Volume 51 (5), 513–530 Published 4 September 2012
513
Table
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So
urc
eo
fsp
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su
sed
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isst
ud
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occ
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Sp
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Co
llec
tio
nn
um
ber
(Her
bari
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acc
essi
on
nu
mb
er)
Str
ain
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nD
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tor
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tio
nn
um
ber
(Her
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acc
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on
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mb
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Str
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mb
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Co
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C.
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tC
.‘o
gasa
wara
ensi
s’K
28
JW4130
Ch
arl
ott
eH
arb
or,
Flo
rid
a,
US
A19
Sep
.2000
J.A
.W
est
rbcL
:EU
349107
LS
U:
JN845507
C.
‘ogasa
wara
ensi
s’K
30
JW3415
Mia
mi,
Flo
rid
a,
US
A14
Jun
.1994
J.A
.W
est
rbcL
:JN
845518
C.
‘ogasa
wara
ensi
s’K
31
JW3374
Pu
eblo
Vie
jo,
Gu
ate
mala
22
Mar.
1993
J.A
.W
est
rbcL
:JN
845519
LS
U:
JN845508
C.
‘ogasa
wara
ensi
s’K
32
JW3408
Jam
esIs
lan
d,
So
uth
Caro
lin
a,
US
A17
Jun
.1994
J.A
.W
est
rbcL
:JN
845520
C.
‘ogasa
wara
ensi
s’K
37
JW3018
I.d
oC
ard
oso
,S
ao
Pau
lo,
Bra
zil
24
No
v.
1989
J.A
.W
est
rbcL
:JN
845521
LS
U:
JN845509
C.
‘ogasa
wara
ensi
s’K
39
JW3412
Fo
rtP
ula
ski,
Geo
rgia
,U
SA
18
Jun
.1994
J.A
.W
est
rbcL
:JN
845522
C.
‘ogasa
wara
ensi
s’K
76
—D
ayto
na
Bea
ch,
Flo
rid
a,
US
A1
Feb
.2004
T.O
.C
ho
C.
rotu
ndata
Kam
iya
K36
JW3375
Lik
in,
Gu
ate
mala
22
Mar.
1993
J.A
.W
est
rbcL
:JN
845523
C.
‘rotu
ndata
’K
72
—B
ay
of
Pan
am
a,
Pan
am
a9
Sep
.1999
B.
Wyso
rrb
cL:
JN845524
LS
U:
JN845510
C.
ruet
zle
rii
Kra
yes
ky,
Fre
der
icq
&J.
N.
No
rris
K146
—C
ud
joe
Key
,F
lori
da,
US
A23
No
v.
2005
T.O
.C
ho
rbcL
:H
M775452
LS
U:
HM
775478
516 Phycologia, Vol. 51 (5), 2012
below first and second axial cells of the main and lateral
axis, in addition to several ventral wing cells located directly
above the node). In some cases these are not exclusive to a
single species, as four rhizoid position types occur in the
Americas and more than four species are recognized.
Presently, there are nine species of Caloglossa in the
Americas (Pedroche et al. 1995; West & Zuccarello 1995;
Kamiya et al. 2000, 2003; Lin et al. 2001; Krayesky et al.
2011): C. apicula (Durant) Krayesky, Fredericq & J.N.
Norris, C. apomeiotica J.A. West & Zuccarello, C. beccarii
(Zanardini) De Toni, C. intermedia Kamiya & J.A. West, C.
leprieurii (Montagne) G. Martens, C. monosticha Kamiya,
C. ogasawaraensis Okamura, C. ruetzleri Krayesky, Freder-
icq & J.N. Norris, and C. ‘stipitata’ E. Post reassessed by
Kamiya et al. (2003) as C. rotundata Kamiya. Recent stu-
dies on C. leprieurii support the view that genus is more
diverse than previously reported and Caloglossa species
may not be as widespread as formerly thought (Krayesky
et al. 2011). Past investigations of Caloglossa (Kamiya &
West 2008; Kamiya et al. 2003, 2011) are often based on a
broad species concept that includes considerable phenotyp-
ic plasticity; C. monosticha and C. leprieurii are two such
examples.
Our goals are threefold. First, the characters used to
delimit species of Caloglossa as defined by King & Puttock
(1994) and Kamiya et al. (2003) will be evaluated. Second,
the status of C. monosticha, a species in the C. continua
complex, as a single disjunctly distributed species sensu
Kamiya et al. (2003) will be assessed. To further clarify, the
C. continua complex is a group of species that were at one
time all thought to be a single taxon, namely, C. continua
(Okamura) R.J. King & Puttock, and this species complex,
as described by Kamiya (2003), encompasses C. continua
spp. continua, C. continua spp. axillaris R.J. King & Puttock,
C. monosticha, C. postiae Kamiya & R.J. King and
C. saigonensis T. Tanaka & P.H. Ho. The distinctiveness
of taxa of the C. continua complex has been the focus of
considerable taxonomic studies (King & Puttock 1994;
Kamiya et al. 1997, 1999, 2003; Wynne & De Clerck 1999).
To address C. continua complex species in the Americas,
C. ‘monosticha’ from North Carolina to Brazil is evaluated
as the second goal of this study. The final objective of
our study is to assess the validity of a singular report of
C. beccarii from Panama (Lin et al. 2001).
MATERIAL AND METHODS
Molecular data
Silica gel-dried, alcohol-preserved specimens and extracted
DNA samples were deposited at the University of
Louisiana at Lafayette (LAF). Plastid-encoded rbcL and
nuclear-encoded LSU rDNA was selected to infer a
phylogeny of Caloglossa from the Americans (Lin et al.
2001). Protocols for DNA extraction followed the proce-
dure in Gavio & Fredericq (2002). Gene amplification and
sequence protocol of rbcL and LSU rDNA followed Gavio
& Fredericq (2002) and Lin et al. (2001), respectively.
Primers used for gene amplification and sequencing of the
rbcL were listed in Freshwater & Rueness (1994), andTable
1.
Co
nti
nu
ed.
Sp
ecie
s
Co
llec
tio
nn
um
ber
(Her
bari
um
acc
essi
on
nu
mb
er)
Str
ain
nu
mb
erL
oca
tio
nD
ate
Co
llec
tor
Gen
Ban
kacc
essi
on
nu
mb
er
C.
ruetz
leri
iK
169
—T
win
Cays,
Bel
ize
18
May
2006
S.
Fre
der
icq
rbcL
:H
M775453
LS
U:
HM
775479
C.
ruetz
leri
iK
170
—T
win
Cays,
Bel
ize
12
May
2006
S.
Fre
der
icq
rbcL
:H
M775454
LS
U:
HM
775480
C.
ruetz
leri
iK
74
—S
um
mer
lan
dK
ey,
Flo
rid
a,
US
A4
Feb
.2004
T.O
.C
ho
rbcL
:H
M775451
LS
U:
HM
775477
C.
ruetz
leri
i—
JW3431
Isla
Mo
rad
a,
Flo
rid
a,
US
A15
Jun
.1994
J.A
.W
est
rbcL
:A
Y150316
LS
U:
AF
522225
C.
ruetz
leri
iK
240
—Is
laS
ola
rte,
Bo
cas
del
To
ro,
Pan
am
a17
Jan
.2007
B.
Wyso
r&
D.W
.F
resh
wate
rrb
cL:
HM
775455
C.
saig
onen
sis
Tan
ak
a&
Ph
am
-Ho
an
gH
o—
JW3846
Po
rtD
ick
son
,S
elan
go
r,M
ala
ysi
a14
May
1998
?L
SU
:A
F522243
C.
stip
itata
E.
Po
stK
33
JW3837
Mo
rib
,S
elan
go
r,W
est
Mala
ysi
a12
May
1998
J.A
.W
est
rbcL
:JN
845525
C.
vie
illa
rdii
(Ku
tzin
g)
Set
chel
l—
JW3328
Gard
enIs
lan
d,
Ad
elaid
eA
ust
rali
a22
Jan
.1993
J.A
.W
est
rbcL
:A
Y150329
LS
U:
AF
522205
Taen
iom
aper
pusi
llum
J.A
gard
hK
75
—F
low
erG
ard
enB
an
ks,
Gu
lfo
fM
exic
o?
S.
Fre
der
icq
rbcL
:JN
845526
T.
perp
usi
llum
—M
K1268
Yak
om
o,
Ok
ino
erab
uIs
lan
d,
Jap
an
20
Feb
.2000
M.
Kam
iya
LS
U:
AF
522249
Krayesky et al.: Two new Caloglossa species from the Americas 517
additional primers used were presented in Gavio &
Fredericq (2002) and Lin et al. (2001). The gene amplifi-
cation and sequencing primers for the middle seg-
ment of LSU rDNA used were listed in Freshwater et al.
(1999).
Alignment
The sequenced samples analyzed are listed in Table 1, and
the sequences deposited in GenBank. RbcL and LSU rDNA
sequence data sets were compiled with Sequencher 4.1
(Gene Codes Corp., Ann Arbor, MI, USA), then imported
into MacClade v4.0 (Maddison & Maddison 2000). The
LSU rDNA sequence data were first aligned by the use of
ClustalX 1.8 (Thompson et al. 1997) before being imported
into MacClade v4.0 and PAUP* for added manual align-
ment. A concatenated data set was not generated, as there
were not both rbcL and LSU rDNA sequences for all sam-
ples analyzed.
Phylogenetic analysis
Phylogenetic analyses were conducted using maximum
parsimony (MP) and maximum likelihood (ML) algorithms
as implemented in PAUP* 4.0b10 (Swofford 2003) and
PHYML (Guindon & Gascuel 2003), respectively. Bayesian
inference was performed in MrBayes 3.0 (Huelsenbeck &
Ronquist 2001). The rbcL and LSU rDNA sequence data
Fig. 1. Maximum likelihood tree for rbcL sequence data (2ln L 8160.22765) of Caloglossa species in the Americas showing the interspecificrelationship. Branch lengths are proportional to sequence change. Species names highlighted in red represent species new to science. Threetiers of numbers at each node; top numbers are MP bootstrap values, middle numbers are ML bootstrap values and bottom numbers areBayesian posterior probabilities (in %).
518 Phycologia, Vol. 51 (5), 2012
utilized Taenioma perpusillum J. Agardh as a single out-
group following Kamiya et al. (2003). Taenioma perpusillum
was also selected as the outgroup based on the view of
Papenfuss (1961) that Caloglossa and Taenioma are closely
related genera.
Parsimony trees obtained under the Fitch criterion of
equal weights for all substitutions (Fitch 1971) were
inferred from a heuristic search, excluding uninformative
characters, with 1000 random sequence additions holding
10 trees at each step and the tree-bisection-reconnection
(TBR) swapping algorithm. Support for nodes in the MP
analysis were assessed by calculating bootstrap proportion
values (Felsenstein 1985) as implemented in PAUP* by
generating 1000 bootstrap data sets, from resampled data,
with 1000 random sequence additions.
Optimal models of sequence evolution to fit the data
alignment estimated by hierarchical likelihood ratio tests
were performed by Modeltest v.3.6 (Posada & Crandall
1998). The model of sequence evolution chosen for both data
files was the GTR+I+G (general time reversible model with
variable base frequencies, symmetrical substitution matrix).
An ML phylogram was generated for each of the two data
sets, using the substitution model, gamma distribution and
proportion of invariable sites determined by the model. For
Fig. 2. Maximum likelihood tree for LSU sequence data (2ln L 4803.02285) of Caloglossa species in the Americas showing the interspecificrelationships. Branch lengths are proportional to sequence change. Species names highlighted in red represent species new to science. Threetiers of numbers at each node; top numbers are MP bootstrap values, middle numbers are ML bootstrap values and bottom numbers areBayesian posterior probabilities (in %).
Krayesky et al.: Two new Caloglossa species from the Americas 519
each data file, the ML tree and ML bootstrap values
(generated from 1000 bootstrap trees) were inferred by
PhyML 3.0, using the nearest neighbor interchange branch
swapping method. An ML phylogram was also generated for
each of the two data sets in PAUP*. The ML phylogram
generated in PAUP* was used as the backbone of the ML
tree for both the rbcL and the LSU data sets.
For the Bayesian analysis, the optimal model of sequence
evolution obtained for each data set was used to set up the
Markov chain Monte Carlo (MCMC) search for the
Bayesian analyses. Four chains of the MCMC were run,
sampling one tree every 100 generations for 2 3 106
generations starting with a random tree for each of the two
data files. The analyses’ phylogenetic inferences were based
Table 2. Morphological differences amongst species of Caloglossa in the Americas, including C. monosticha. The definition of rhizoiddistribution, FLA, NA and FMA follows Kamyia et al. (2003).
C. apicula C. apomeiotica C. intermedia C. fluviatilis sp. nov. C. leprieurii
Endogenousbranching
present present present present present
Adventitiousbranching
present,occasionally
absent absent present absent
Number of rhizoidsper cell
multiple multiple multiple multiple multiple
Rhizoid distribution type F type F type F type F type FRhizoids tightly
adhering at base(cortical pad)
absent present present present present
Adaxial cell rowderived from theFLA1
absent absent present present absent
Number of cell rowsderived from theNA2
1–2 1–2 1(2) 1 1–2
Number of cell rowsderived from theFMA3
1–3 (1–)2–5 1–3 2–4 (2–)3–7
Constriction at thenode
slightly slightly slightly strongly(short interval)
slightly
Blade shape strap-like strap-like strap-like lanceolate strap-likeMiddle blade width
(mm)0.1–1.0 0.9–2.5(–4) 0.9–1.2 0.8–1.1 0.5–1.4
Blade length (mm) 1.0–3.0 1.5–4.0 2.0–3.0 3.0–6.0(–7.0) 1.5–4.0Thallus length (cm) 0.3–1.2 0.3–1.7 0.7–1.2 (0.5–)0.7–1.3 0.8–2.0
C. monosticha C. confusa sp. nov. C. ogasawaraensis C. rotundata C. ruetzleri
Endogenousbranching
present present absent present present
Adventitiousbranching
absent absent present present,occasionally
absent
Number of rhizoidsper cell
single single single single multiple
Rhizoid distribution type G type G type E type C type FRhizoids tightly
adhering at baseabsent absent absent absent present
Adaxial cell rowderived from theFLA
present present present present absent
Number of cell rowsderived from theNA
1–3 1–3 1 1 1
Number of cell rowsderived from theFMA
1 1–2 1 1–2 1
Constriction at thenode
slightly strongly(long interval)
slightly strongly(long interval)
strongly(long interval)
Blade shape strap-like ellipticalto ovate
strap-like ellipticalto rotundate
ellipticalto ovate
Middle blade width(mm)
0.5–1.5 1.0–1.5 0.1–0.5 1.7–2.2 1.1–2.0
Blade length (mm) 1.5–3.0 (2.0–)2.5–5.5 1.5–4.5 3.0–5.0 3.0–6.0Thallus length (cm) 0.5–1.2 0.5–1.1 0.7–1.7 0.5–1.0 0.5–1.0
1 FLA, first axial cell from lateral axis.2 NA, nodal axial cell opposite lateral branch.3 FMA, first axial cell at main axis opposite lateral branch.
520 Phycologia, Vol. 51 (5), 2012
on the trees sampled after ‘burn-in’. A 50% majority rule
consensus, as implemented by PAUP*, was computed from
those saved trees. This frequency corresponded to the
posterior probability of the clades.
Morphological studies
Herbarium specimens as well as live collections were used in
this study. Information on taxa studied, collection sites,
date of collection and collectors are listed in Table 1. For
microscope observations, specimens were stained with 1%
aqueous aniline blue acidified with 0.1% HCl. Voucher spe-
cimens and materials for morphological studies were fixed
and stored in 5% formalin in seawater and/or pressed as
herbarium sheets deposited in LAF and the Algal Collec-
tion of the U.S. National Herbarium (US). Abbreviations
for herbaria follow the online Index Herbariorum (http://
sweetgum.nybg.org/ih).
RESULTS
Molecular analysis
The rbcL and LSU rDNA sequences included in this study
represent a wide sampling of Caloglossa, with a strong em-
phasis on the America species. The rbcL data set consisted of
37 samples (399 parsimony informative sites out of 1467
base pairs); MP generated 36 trees (tree length 5 1207 steps)
and the topology of the MP trees did not differ significantly
from that of the ML and Bayesian trees. The LSU data set
consisted of 30 samples (253 parsimony informative sites out
of 1017 aligned base pairs); MP generated five trees (tree
length 5 788 steps), the topology of which did not differ
significantly from that of the ML and Bayesian trees. As the
MP, ML and Bayesian trees did not differ significantly, only
the ML trees are presented (Figs 1, 2).
Both the rbcL and the LSU ML trees (Figs 1, 2)
demonstrate the genus to be monophyletic but without
strong support. The rbcL-based phylogeny (Fig. 1) shows
Caloglossa to be composed of two major clades: clade 1
containing C. apomeiotica, C. intermedia, C. leprieurii, C.
apicula, C. ruetzleri, C. vieillardii (Kutzing) Setchell, C.
continua, C. monosticha and C. confusa sp. nov. and clade 2
including C. ogasawaraensis, C. stipitata, C. bengalensis (G.
Martens) R.J. King & Puttock, C. fluviatilis sp. nov. and C.
rotundata. These clades are well supported by moderate to
strong bootstrapping values and posterior probabilities.
The species of Caloglossa are separated into three clades
based on LSU sequence analysis (Fig. 2). Clade 1 has full
support for Bayesian posterior probability with moderate
support for the MP and ML analyses. The specific rela-
tionships in the clade 1 of the LSU tree is nearly identical to
that of rbcL tree in Figure 1 except for C. saigonensis and a
collection of C. monosticha from Monkey Mia, Australia,
which are represented by only a single LSU sequence, and
C. intermedia is represented by a single rbcL sequence in
this study. Clade 2 is well supported (Fig. 2) and likewise
is similar to that of Figure 1 apart from the position of
C. beccarii, a taxon represented by two LSU sequences.
Caloglossa fluviatilis and C. stipitata do not form a mono-
phyletic clade, and C. bengalensis is not closely allied with
C. ogasawaraensis. The change in the relationship of these
species to one another is not surprising given the low to
nonexistent MP and ML support shown in Figure 2 for
these monophyletic assemblages. In addition, within the
LSU tree, C. rotundata is not nested within clade 2 but
forms a separate clade, and clade 3 receives low MP sup-
port and Bayesian posterior probabilities.
The separation of species into the major clades appears to
reflect characteristics of secondary branching observed in
these taxa. Clade 1 in the rbcL tree (Fig. 1) contains the
species bearing endogenous bladelets and with some species
generating the occasional adventitious bladelets, namely,
C. apicula. Clade 2 (Fig. 2) encompasses Caloglossa species
with exclusive adventitious branching that is common to
abundant in individuals, except for C. rotundata, which bears
occasional adventitious branching and common endoge-
nous branching. The LSU data (Fig. 2) differ from the rbcL
data in that species with exclusive adventitious branching
form a clade separate from C. rotundata, whose secondary
branching is primarily endogenous and occasionally adven-
titious.
All species sampled from the rbcL sequence data are
strongly supported (Fig. 1), and the LSU sequence data
(Fig. 2) further corroborate that the species reported here
are molecularly distinct. These well-supported monophy-
letic groups that compose the species of Caloglossa from the
Americas (Figs 1, 2) are likewise morphologically distinct
and are discernible from one another (Table 2). The rbcL
sequence data further indicate that the molecular distinc-
tion of the species sampled as interspecies sequence diver-
gence ranges between 2.7% and 14.1% and that intraspecies
sequence divergence ranges between 0.1% and 1.5% for
Caloglossa species.
The rbcL and LSU sequence data presented here suggest
that both C. ogasawaraensis and C. rotundata as currently
understood encompass additional species. An intraspecific
rbcL sequence divergence ranging from 5.0% to 5.4% exists
between C. ogasawaraensis of the western Pacific and the
Americas, with both the western Pacific clade and the clade
from the Americas being well supported (Figs 1, 2). At
present, limited samples exist for C. rotundata, a species
known only from its type locality and from a newly reported
collection from Panama. The intraspecific rbcL sequence
divergence among a cultured individual that is part of the C.
rotundata type collection and a Panama collection is 6.4%.
Comparative morphological and systematic study
The research presented by Kamiya et al. (2003) suggests
that species of Caloglossa reported from the Americas, such
as C. monosticha and C. beccarii, may actually be restricted
to other regions of the world. Molecular data (Figs 1, 2)
corroborate the morphological differences based on subtle
but distinct differences in thallus node constriction, blade
length and rhizoid morphology (Table 2), thus indicating
that these taxa are undescribed. A dichotomous key is here
provided to highlight the diagnostic features amongst all
the species of Caloglossa recognised for the Americas.
Krayesky et al.: Two new Caloglossa species from the Americas 521
Species’ authorities are listed in the sections following the
key.
Key to the species of the Caloglossa in the Americas
1a. Adaxial cell row derived from first axial cell of
lateral axis absent . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Adaxial cell row derived from first axial cell of
lateral axis present . . . . . . . . . . . . . . . . . . . . . . . 5
2a. Basal rhizoidal cells loosely arranged, not forming a
cortical pad . . . . . . . . . . . . . . . . . . . . . . C. apicula
2b. Basal rhizoidal cells tightly adherent forming a
cortical pad . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3a. Only one cell row derived from the first axial cell of
the main axis opposite the lateral branch; thallus
highly constricted at nodes over long interval, with
internodes forming elliptical ‘blades’ . . . . C. ruetzleri
3b. Usually more than one cell row derived from the
first axial cell of the main axis opposite the lateral
branch; thallus not highly constricted at nodes,
strap-like throughout . . . . . . . . . . . . . . . . . . . . . 4
4a. Number of cell rows derived from the first axial cell of
the main axis opposite the lateral branch (1–)2–5; thal-
lus width at median internode region 0.9–2.5(–4.0) mm.
. . . . . . . . . . . . . . . . . . . . . . . . C. apomeiotica
4b. Number of cell rows derived from the first axial
cell of the main axis opposite the lateral branch
(2–)3–7; thallus width at median internode region
0.5–1.4 mm . . . . . . . . . . . . . . . . . . . . . C. leprieurii
5a. Endogenous branching absent . . . . . . . . . . . . . . . 6
5b. Endogenous branching present . . . . . . . . . . . . . . . 7
6a. Thallus blades lanceolate and thallus nodes strongly
constricted; median blade width (0.8–1.1 mm). . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . C. fluviatilis
6b. Thallus blades strap-like and thallus nodes slightly
constricted; median blade width (0.1–0.5 mm). . . . .
. . . . . . . . . . . . . . . . . . . . . . C. ogasawaraensis
7a. Rhizoids clustered at the node between the main and
lateral axis, type G rhizoid arrangement . . . . C. confusa
7b. Rhizoids clustered in another arrangement other
than type G. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8a. Thallus blades elliptical to rotundate; rhizoids not
forming a cortical pad; adventitious branching present
(occasionally) . . . . . . . . . . . . . . . . . . . . C. rotundata
8b. Thallus blades strap-like; rhizoids forming a cortical
pad; adventitious branching absent. . . . . .C. intermedia
New species of Caloglossa
Caloglossa confusa Krayesky, J.A. West et Kamiya, sp. nov.
(Figs 3–4, 6–13)
Thalli plani, materia herbarii pallidarosea ad pallidam
violaceorubram, subdichotomi ramosi, usque ad 0.5–1.1 cm
long, constantes ex costae regione alis monostromaticis
alteruter latere. Regio costae constans ex duo cellulis
transversalibusque duo cellulis lateralibus periaxialibus,
serie axiali cellulis. Laminae constrictae valde ad nodos,
(2.0–)2.5–5.5 mm longaeque 1.0–1.5 mm latae internodi
medianam regionem, afiliformes ad nodum. Internodi aut
laminae ovati. Ramificatio endogenea praesens. Ramifica-
tio adventitia absens. Fila rhizoidea fasciculata nodum inter
axem principalem lateralemque. Series cellularum adaxialis
oriunda e prima axiali cellula lateralis axis praesens.
Numerus serierum cellularum oriundarum e cellula axiali
nodalis opposita ramum laterale 1–3. Numerus serierum
cellularum oriundarum e prima cellula axiali in axem
principalem opposita ramum laterale 1–2. Tetrasporangia
(48–)52–68 mm latis, 56–68 mm longis. Diversae structurae
reproductivae non visae.
Thalli flat, herbarium material light pink to pale violet-
red, subdichotomously branched, 0.5–1.1 cm long (Figs 3,
4), consisting of a midrib region with monostromatic wings
to either side (Fig. 6). Midrib region composed of two
transverse and two lateral periaxial cells, and one axial cell
series per segment. Blades strongly constricted at nodes
(Fig. 7), (2.0–)2.5–5.5 mm long and 1.0–1.5 mm wide at
median node region. Internodal regions or ‘blades’ ovate
(Fig. 3). Endogenous branching present (Fig. 10). Adven-
titious branching absent. Rhizoids clustered at the node
between the main and lateral axis (Figs 11, 12). Adaxial cell
row derived from first axial cell from lateral axis present
(Fig. 8). Number of cell rows derived from nodal axial cell
opposite lateral branch 1–3 and from the first axial cell of
the main axis opposite the lateral branch 1–2 (Fig. 9).
Tetrasporangia 56–68 mm tall, (48–)52–68 mm in diameter
(Fig. 13). Other reproductive structures not observed.
ETYMOLOGY: The species epithet, confusa, reflects the
confusion or misidentification of this taxon in the past as
another species of the Caloglossa continua complex, namely,
C. monosticha.
HOLOTYPE: Plantation Key, Florida, USA [24u599230N,
80u329350W]; 15 June 1994, coll. J.A. West, West 3426
(SLRO 19072).
DISTRIBUTION: Western Atlantic: North Carolina to Bra-
zil. Caloglossa confusa appears to be uncommon in the
tropical coastlines of the western Atlantic.
HABITAT: Growing associated and often entangled with
Bostrychia.
ADDITIONAL SPECIMENS STUDIED: Paratypes: North Car-
olina, USA: Fort Fisher (J.A. West, 26 June 2003); Mexico:
Isla Chinchorro, Laguna de Terminos. Campeche (L.
Huerta, s. dat.); Brazil: I. do Cardoso, Sao Paulo (J.A.
West, 4 April 1990).
ADDITIONAL SPECIES STUDIED: Caloglossa monosticha
Kamiya; Australia: Derby [M. Kamiya, 2 October 1991,
holotype (TNS-AL-42365)].
REMARKS: In the Americas, Caloglossa confusa sp. nov.
was previously identified as ‘C. monosticha’ based on
morphology (Kamiya et al. 2000, 2003, 2011). Kamiya et al.
(2003) already recognised the strains isolated from Florida
and Brazil had stronger constrictions at each node than the
western Pacific strains and also demonstrated their distant
relationship on the basis of the LSU analysis. Hybridization
studies further demonstrated that the strain from Florida
was reproductively isolated from three culture collections
from the Pacific (Kamiya et al. 2003). In the present study,
the rbcL analysis that was not carried out in the Kamiya et
al. (2003) corroborates that the western Atlantic specimens
522 Phycologia, Vol. 51 (5), 2012
are genetically distinct from the western Pacific ones.
Furthermore, the specimens from North Carolina and
Mexico were newly obtained for our study. Consequently,
we concluded to distinguish the western Atlantic entity as a
new species, C. confusa.
Wynne & De Clerck (1999) have suggested that Calo-
glossa monosticha sensu Kamiya et al. (1997) was not a
distinct species and placed C. monosticha in synonymy with
C. saigonensis based on a lack of distinctive morphological
features between the two taxa. Kamiya et al. (1999) in a
subsequent work resurrected C. monosticha based on mor-
phological characteristics, namely, the presence of a single
row of cells derived from a nodal cell, and blade shape
as evidence that C. monosticha is a distinct species from
C. saigonensis.
Studies of the type material of C. monosticha reveal
vegetative differences distinguishing it from C. confusa. The
most striking difference between these two species is the
strong constriction at the node in C. confusa (Fig. 3) versus
a slight constriction in C. monosticha (see Kamiya et al.
1997, figs 3–5), along with different blade shapes. Under
similar culture conditions, C. confusa tends to generate
ovate blades (Fig. 4); whereas, C. monosticha does not, thus
demonstrating the stability of the character (Fig. 5). Blade
length in mature blades is generally greater in C. confusa
(2.5–5.5 mm) than in C. monosticha (1.5–3.0 mm), with
some degree of overlap within each species range. Although
these differences are not overwhelming, they are sufficiently
different to identify each species from the other. In addition
to vegetative characters, rbcL and LSU rDNA sequence
data (Figs 1, 2) indicate that the two taxa form separate
lineages, and when additional taxa of the C. continua
complex are included in the analysis, C. confusa and C.
monosticha do not form a monophyletic group. The mole-
cular evidence (Fig. 2) also supports the view of Kamiya
et al. (1997, 1999) that C. monosticha is distinct from
C. saigonensis.
Caloglossa confusa is easily differentiated from the other
species of the genus reported from the Americas as it is
the only taxon reported to have the type G rhizoidal
arrangement as defined by Kamiya et al. (2003). Superfi-
cially, this species could be confused only with C. rotundata
and C. ruetzleri, which share a similar blade morphology
with C. confusa, elliptical to rotundate and ovate to sub-
ovate, respectively. Both C. rotundata and C. ruetzleri have
a different rhizoid arrangement from C. confusa (Table 2).
In addition, C. rotundata generates adventitious branches,
and C. ruetzleri lacks an adaxial cell row derived from the
first axial cell of a lateral axis.
Caloglossa fluviatilis Krayesky, Fredericq et J.N. Norris, sp.
nov.
(Figs 14–25)
Thalli caespitosi, plani, materia herbarii pallida fusca ad
pallidam violaceorubram, subdichotomi ramosi, usque ad
(0.5–)0.7–1.3 cm longis, constantes ex costae regione alis
monostromaticis alteruter latere. Regio costae constans
ex duo cellulis transversalibusque duo cellulis lateralibus
periaxiaibus, serie axiali cellulis. Laminae lanceolatae, me-
dianam regionem 6.0(–7.0) mm longaeque 0.8–1.1 mm latae
constrictae valde ad nodos. Ramificatio endogenea absens.
Ramificatio adventitia praesens. Fila rhizoidea formantia
cellulis periaxialibus nodi alatocellulis marginalibusque
vicinis. Cellulae basales rhizoideae non arcte appressae for-
mantes tumulum corticalem similem stipitis. Series cellu-
larum adaxialis oriunda e prima axiali cellula lateralis axis
praesens. Numerus serierum cellularum oriundarum e cel-
lula axiali nodalis opposita ramum laterale unus. Numerus
serierum cellularum oriundarum e prima cellula axiali in
axem principalem opposita ramum laterale 2–4. Reproduc-
tio ignota.
Thalli arranged in tufts (Fig. 14), flat, herbarium
material pale brown to pale violet-red, subdichotomously
branched (Figs 15, 16), and up to (0.5–)0.7–1.3 cm long,
consisting of a midrib region with monostromatic wings on
either side. Midrib region composed of two transverse and
two lateral periaxial cells, and one axial cell series. Blades
lanceolate 3.0–6.0(–7.0) mm long and 0.8–1.1 mm wide at
median node region and strongly constricted at nodes.
Endogenous branching absent. Adventitious branching
present (Figs 22–25). Rhizoids forming from nodal periax-
ial cells in addition to adjacent marginal wing cells (Figs 20,
21). Basal rhizoidal cells not tightly adherent to form a
stipe-like cortical pad. Adaxial cell row derived from first
axial cell from lateral axis present (Fig. 19). Number of cell
rows derived from nodal axial cell opposite lateral branch is
one; two- to four-cell rows derived from the first axial cell
of main axis opposite lateral branch (Figs. 17, 18).
Reproductive structures were not found.
ETYMOLOGY: This species epithet, fluviatilis (from the
Latin ‘of the stream’), is named for the freshwater system
the species inhabits.
HOLOTYPE: Pedro Miguel Lock, Pedro Miguel, Republic
of Panama [08u59946.420N, 79u35930.490W]; on cement of
western wall of upper western lock (freshwater), 5 May
1999, coll. B.S. Wysor, 421 (US Alg. Type Coll. 217746);
isotype LAF.
DISTRIBUTION: Central America: Known only from the
type locality.
HABITAT: Occurring in freshwater stream or rivers;
attached to cement.
REMARKS: Caloglossa fluviatilis is newly reported from
tropical Central America. It was previously reported from
Panama as C. beccarii by Lin et al. (2001). The rhizoidal
arrangement of this species is of the type F arrangement of
Kamiya et al. (2003) and not the type D arrangement as
seen in C. beccarii and C. stipitata reported by Kamiya
et al. (2003). Thallus nodes in C. beccarii are only mod-
erately constricted; whereas, nodes in C. fluviatilis are
strongly constricted. Furthermore, LSU sequence data
indicate that C. fluviatilis forms a lineage that is separate
from C. beccarii. The clarification presented herein
effectively removes C. beccarii from the marine flora of
the western Atlantic.
Caloglossa fluviatilis is easily distinguished from other
Caloglossa species from the region. Although C. apomeio-
tica, C. leprieurii and C. fluviatilis have a type F rhizoidal
arrangement as described in Kamyia et al. (2003), thalli of
Krayesky et al.: Two new Caloglossa species from the Americas 523
524 Phycologia, Vol. 51 (5), 2012
C. fluviatilis are more constricted at the nodes, possess
adaxial cell rows derived from the first axial cell of the
lateral axis, lack a corticated pad at the base of the rhizoid
filaments and produce adventitious branching. Caloglossa
ogasawaraensis likewise can easily be differentiated from C.
fluviatilis in that the former has a type E rhizoid
arrangement (Kamiya et al. 2003), a median thallus blade
width of 0.1–0.5 mm and a range that is smaller and does
not overlap with that of C. fluviatilis (0.8–1.1 mm).
Furthermore, unlike the new species, C. ogasawaraensis is
not strongly constricted at the nodes.
The only species from the region that C. fluviatilis may
superficially be mistaken for are C. confusa, C. rotundata
and C. ruetzleri due to the thallus blades not being strap-
like. Although the nodes of C. fluviatilis are strongly
constricted as in these three species, the blades are
lanceolate in C. fluviatilis (Table 2). Caloglossa fluviatilis
also generates proliferous adventitious branches and
appears to lack endogenous branching; however, endoge-
nous branching is present in C. confusa, C. rotundata and C.
ruetzleri, and adventitious branching is lacking in all three
except for C. rotundata, where it is occurs rarely. This
species further differs from C. confusa, C. rotundata and C.
ruetzleri in rhizoid anatomy; however, whereas C. ruetzleri
shares with C. fluviatilis a type F rhizoid arrangement, the
rhizoidal filaments in the former are organized in a
corticated pad at their base.
Other species of Caloglossa reported from the Americas
Caloglossa leprieurii (Montagne) G. Martens
(1869, pp. 234, 237)
BASIONYM: Delesseria leprieurii Montagne (1840, 196–197,
pl. 5, fig. 1).
TYPE LOCALITY: Sinnamary, NW of Cayenne, French
Guiana.
DISTRIBUTION: Caloglossa leprieurii appears widespread in
tropical waters of western Atlantic, eastern Pacific and
Indian Oceans. Western Atlantic Ocean: Bermuda, Puerto
Rico, Guadeloupe, French Guiana, Suriname, Guyana and
Venezuela. Eastern Pacific: Costa Rica. Indian Ocean:
India, Madagascar, South Africa (Krayesky et al. 2011).
HABITAT: Epiphytic on mangrove pneumatophores and prop
roots; intertidal to very shallow subtidal (Krayesky et al. 2011).
REMARKS: The generitype C. leprieurii is differentiated
from most of the species reported from the Americas,
except for C. apicula, C. apomeiotica and C. ruetzleri, based
on the absence of an adaxial cell row derived from the first
axial cell from the lateral axis. The species can be confused
with the somewhat similar C. apomeiotica (see remarks
below for C. apomeiotica), but it is easily differentiated
from C. ruetzleri in blade morphology and in cell row
number from the first axial cell at the main axis opposite
the lateral axis (Table 2). Caloglossa leprieurii can likewise
be readily distinguished from C. apicula as rhizoid filaments
are organized in a cortical pad in C. leprieurii and not C.
apicula; it is also tropical in its distribution unlike the
temperate C. apicula.
Caloglossa leprieurii has been wrongly applied to western
Atlantic specimens presently distinguished as other spe-
cies (e.g. C. apicula, C. apomeiotica, C. intermedia and C.
ruetzleri) by investigators for nearly 150 years (Krayesky
et al. 2011). In a recent study of Caloglossa, namely,
Kamiya et al. (2011), many strains from this area were still
assigned to C. leprieurii because that report paper was
published prior to the appearance of Krayesky et al. (2011).
A combination of rbcL, LSU and morphological data has
elucidated part of the C. leprieurii complex occurring in
the Americas, and at present this species is no longer
verified from western Atlantic habitats in the United States
(Krayesky et al. 2011).
Caloglossa apomeiotica J.A. West & Zuccarello (1994,
p. 383, figs 1–15)
TYPE LOCALITY: On mangrove prop roots, Puerto San
Carlos, Bahıa Magdalena, Baja California Sur, Pacific
Mexico.
DISTRIBUTION: Eastern Pacific: Baja California Sur, Mex-
ico to Central America. Western Atlantic: South Carolina,
USA to Brazil (Krayesky et al. 2011).
HABITAT: Associated with mangroves, growing on pneu-
matophores of Avicennia germinans (Linnaeus) Linnaeus
and Laguncularia racemosa (Linnaeus) Gaerth and prop
r
Figs 3–13. Caloglossa confusa sp. nov. (Figs 3, 6, 7, 9, 11, 13: Isla Chinchorro, Campeche, Mexico; Figs 4, 8, 10, 12: R. Pereque, I. doCardoso, S.P. Brazil.; Fig. 5: Holotype of Caloglossa monosticha; M. Kamiya, 2 October 1991, TNS-AL-42365).
Fig. 3. Habit. Scale bar 5 2.0 mm.Fig. 4. Cultured specimen of C. confusa. Scale bar 5 1.0 mm.Fig. 5. Thallus of cultured material of C. monosticha. Scale bar 5 1.0 mm.Fig. 6. Juvenile node. Short straight arrow points to the nodal axial cell; long straight arrow points to the first axial cell of main axis;arrowhead points to the first axial cell of the lateral axis. Scale bar 5 100 mm.Fig. 7. Mature node showing strong constriction at thallus node. Scale bar 5 0.5 mm.Fig. 8. Nodal region with an adaxial cell row derived from the first axial cell of lateral axis. Scale bar 5 100 mm.Fig. 9. Main axis of a mature node. Arrowheads point to the cell row lineages derived from the first axial cell of the main axis opposite thelateral branch. Scale bar 5 100 mm.Fig. 10. Endogenous branching. Scale bar 5 0.25 mm.Fig. 11. Rhizoid cell initials; cell initials located apically of midrib cells demonstrating type G arrangement of rhizoids of Kamiya et al.(2003). Scale bar 5 100 mm.Fig. 12. Mature rhizoids of the type G arrangement showing rhizoidal filament located apically of midrib cells. Scale bar 5 100 mm.Fig. 13. Blade with tetrasporangia. Scale bar 5 0.5 mm.
Krayesky et al.: Two new Caloglossa species from the Americas 525
Figs 14–25. Caloglossa fluviatilis sp. nov. (Figs 14–25: Pedro Miguel Lock, Pedro Miguel, Panama).Fig. 14. Habit. Scale bar 5 2.0 mm.Fig. 15. Thallus blades showing strong constriction at node. Scale bar 5 1.0 mm.Fig. 16. Lateral axis (arrow) demonstrating significantly less growth than the main axis. Scale bar 5 1.0 mm.Fig. 17. Juvenile node. Short straight arrow points to the nodal axial cell; long straight arrow points to the first axial cell of main axis;arrowhead points to the first axial cell of the lateral axis. Scale bar 5 100 mm.Fig. 18. Higher magnification of the juvenile node in Fig. 17. Arrowheads point to the cell row lineages derived from the first axial cell ofthe main axis opposite the lateral branch. Scale bar 5 100 mm.Fig. 19. Nodal region of thallus illustrating the presence of an adaxial cell row derived from the first axial cell of the lateral axis(arrowhead). Scale bar 5 50 mm.
526 Phycologia, Vol. 51 (5), 2012
roots of Rhizophora mangle Linnaeus (Krayesky et al.
2011).
REMARKS: Caloglossa apomeiotica is closely allied with C.
leprieurii, C. apicula and C. ruetzleri (Krayesky et al. 2011).
It can be readily distinguished from C. apicula by the
presence of a cortical pad associated at the base of the
rhizoid filaments and by ovate blades in that of C. ruetzleri.
Caloglossa apomeiotica can be separated vegetatively from
C. leprieurii by the number of cell rows derived from the
first axial cell of the main axis opposite to the lateral
branch; however, there is overlap in the cell row range
between these species [i.e. two to five in C. apomeiotica vs
three to seven in C. leprieurii]. Therefore, material in which
the nonoverlapping parts of the cell row range can be
observed is required to make a definite determination be-
tween these two species. Caloglossa apomeiotica has some-
what more robust thalli than C. leprieurii (Table 2), and
material of the latter whose median blade width is closer to
the larger end of the range in C. apomeiotica will further aid
in distinguishing C. apomeiotica from C. leprieurii.
Caloglossa apicula (Durant) Krayesky, Fredericq & J.N.
Norris (2011, figs 4 A–L)
BASIONYM: Apiarium apicula Durant (1850, p. 18).
TYPE LOCALITY: On Fucus vesiculous Linneaus, northwest
side of Upper New York Bay, Jersey City, New Jersey,
USA.
DISTRIBUTION: Northern Gulf of Mexico: Texas to middle
Gulf Florida. Eastern United States: Connecticut to middle
Atlantic Florida (Krayesky et al. 2011). Caloglossa apicula
appears widespread along the coast of the northern Gulf of
Mexico and the eastern United States and is often common
where found. It represents much of what in the past was
identified by workers as C. leprieurii.
HABITAT: Growing on rock, Spartina alterniflora Loise-
leur-Deslongchamps, and epiphytic on pneumatophores of
A. germinans (Linnaeus) Linnaeus in salt marshes and
mangroves; also occasional in freshwater streams and rivers
on sediments and grasses (Krayesky et al. 2011).
REMARKS: Caloglossa apicula is a temperate species. It can
be easily distinguished from other species of Caloglossa
with similar distribution. Unlike C. ogasawaraensis, this
species lacks abundant adventitious branching, does not
possess an adaxial cell row derived from the first axial cell
of the lateral axis and has a different rhizoidal arrangement
(Table 2). Two other species, C. apomeiotica and C. inter-
media, both produce a cortical pad in which the base of
rhizoid filaments are attached in contrast to C. apicula,
which lacks this structure.
Caloglossa ruetzleri Krayesky, Fredericq & J.N. Norris
(2011, figs 6 A–G)
HOLOTYPE: Epiphytic on mangrove pneumatophores and
prop roots, Twin Cays, Belize, [16u489N; 88u059W], 18 May
2006, intertidal, coll. S. Fredericq, J.N. Norris, C.F.D.
Gurgel & R.H. Sims, K169 (US Alg. Type Coll.- 0210477);
isotypes: LAF.
DISTRIBUTION: Caribbean Sea: Panama and Belize. Gulf of
Mexico: Florida Keys (Krayesky et al. 2011). Caloglossa
ruetzleri appears thus far to be restricted to the neotropics.
HABITAT: Intertidal; growing on red mangrove pneumat-
ophores and prop roots (Krayesky et al. 2011).
REMARKS: Caloglossa ruetzleri is distinct from C. apomeio-
tica and C. leprieurii in being highly constricted at the nodes
and in generating an ovate to subovate blade morphology.
In addition, C. ruetzleri generates only a single cell row that
is derived from the first axial cell of the main axis opposite
the lateral axis unlike in C. apomeiotica (two to five) and
C. leprieurii (three to seven). The only other taxa that
C. ruetzleri could superficially be confused with based on
blade morphology and constriction at the nodes are
C. confusa, C. rotundata and perhaps C. fluviatilis. The
presence of a cortical pad in which the bases of the rhizoid
filaments are joined, in addition to the absence of an adaxial
cell row derived from first axial cell from lateral axis, easily
distinguish C. ruetzleri from the former three species.
Caloglossa intermedia Kamiya & J.A. West (2000, p. 419,
figs. 1–5)
TYPE LOCALITY: James Island, South Carolina, USA.
DISTRIBUTION: Northwestern Atlantic: New Jersey to Florida.
HABITAT: Growing on Salicornia perennis (P. Miller) A.J.
Scott and on Spartina alterniflora in salt marshes.
SPECIMENS STUDIED: South Carolina: James Is. (topotype:
D.R. Wiseman, 10 July 2003); Georgia: Sapelo Is. (M.
Darley, 24 October 1990); Florida: St. Augustine (G.A. Hall,
s. dat., US-00296142).
REMARKS: Caloglossa intermedia is here newly reported
from the eastern Florida coast. It is easily distingui-
shed from the other species reported from the region, C.
apomeiotica, C. apicula and C. ogasawaraensis. Caloglossa
ogasawaraensis is different from C. intermedia by its smaller
median blade width and production of abundant adventi-
r
Figs 20, 21. Rhizoids at node displaying the type F arrangement of Kamiya et al. (2003). Mound of corticated cells at the base of therhizoid filaments absent. Scale bars 5 100 mm.Fig. 22. Thallus node with multiple adventitious branches. Scale bar 5 1.0 mm.Figs 23, 24. Higher magnification of the thallus node in Fig. 22. Arrow points to a juvenile adventitious branch. Scale bars 5 100 mm.Fig. 25. Thallus node with an adventitious branch at vertical arrow. Horizontal arrow points to the lateral axis showing significantly lessgrowth than the main axis. Scale bar 5 100 mm.
Krayesky et al.: Two new Caloglossa species from the Americas 527
tious branches (Table 2). Caloglossa apicula and C. oga-
sawaraensis do not develop rhizoids that are tightly
adhering at the base to form a mound of cells (cortical
pad), as seen in C. intermedia. Furthermore, C. intermedia
does generate an adaxial cell row derived from the first
axial cell of the lateral axis; whereas, both C. apomeiotica
and C. apicula lack this condition. Caloglossa intermedia is
further distinct based on rbcL sequence data.
Caloglossa ogasawaraensis Okamura (1897, pp. 12–13, figs
A–D)
TYPE LOCALITY: Ogasawara-gunto (Bonin Islands), Japan.
DISTRIBUTION: Western Pacific: Japan to Australia. Indian
Ocean: India, Bombay. Eastern Atlantic: Tanzania, Zanzi-
bar. Western Atlantic: South Carolina, USA to Brazil.
HABITAT: Associated with mangrove pneumatophores.
SPECIMENS STUDIED: Western Atlantic: Florida: Charlotte
Harbor (J.A. West, 19 September 2000); Miami, (J.A. West,
14 June 1994); Daytona Beach (T.O. Cho, 1 February
2004); Georgia: Fort Pulaski (J.A. West, 18 June 1994);
South Carolina: James Is. (J.A. West, 17 June 1994);
Guatemala: (J.A. West, 22 March 1993); Brazil: I. do
Cardoso, Sao Paulo, (J.A. West, 24 November 1989).
WESTERN PACIFIC: Bali: Nusa Lembongan (J.A. West,
25 April 1999); Japan: Bonin Islands (C. Wright, s. dat.,
US-00296138).
REMARKS: This is the first published report of C. oga-
sawaraensis from the Gulf of Mexico. Caloglossa ogasawar-
aensis is morphologically distinct from other species
primarily based on its fine stature of which the median
blade width is 0.1–0.5 mm, and the possession of a type E
rhizoid arrangement (Kamiya et al. 2003). It differs from
all other reported species from the Americas, except
C. fluviatilis and C. rotundata, in its prolific generation of
adventitious branches. The rbcL and LSU rDNA sequence
data suggest that the western Atlantic populations repre-
sent a separate species from that of the western Pacific
populations. Despite a disjunct distribution, they are
treated here for the present as a single species until a more
detailed investigation can determine if there are morpho-
logical characteristics that distinguish between these two
groups.
Caloglossa rotundata Kamiya (2003, pp. 493–494, figs 17–20)
TYPE LOCALITY: Buena Vista, Likin, Guatemala.
DISTRIBUTION: Eastern Pacific: Guatemala to Panama.
Caloglossa rotundata appears to be uncommon in its range.
HABITAT: Growing on prop roots and trunks of the
mangrove Rhizophora mangle L. and a marine mudflat.
SPECIMENS STUDIED: Guatemala: Likin (J.A. West, 22
March 1993); Panama: Bay of Panama (B. Wysor, 9
September 1999).
REMARKS: Appearing narrowly distributed, C. rotundata is
known only from the type collection and our newly reported
material collected from the Bay of Panama, Pacific Panama.
This species is easily differentiated from other species of
Caloglossa reported from the tropics of the Americas (see
Table 2) by its rhizoidal morphology, as it is the only species
known to have the type C rhizoidal arrangement presented
in Kamiya et al. (2003). As C. rotundata possesses an axial
cell row derived from the first axial cell of the lateral axis, it
can easily be distinguished from species that lack it, such as
C. apomeiotica, C. leprieurii and C. ruetzleri. In addition to
rhizoidal morphology, C. rotundata is distinguished from
C. ogasawaraensis and C. confusa in that it produces both
endogenous and adventitious branching and because the
median blade width is much finer in C. ogasawaraensis
(Table 2). Caloglossa fluviatilis can further be differentiated
from C. rotundata based on the prolific generation of
adventitious branching, lack of endogenous branching and
the lanceolate blade morphology of the thallus in the
former.
The rbcL and LSU sequence data demonstrate that the
type material from Guatemala and the Panama collection
form a monophyletic group. However, there seems to be a
significant amount of genetic divergence between these two
populations, suggesting that they are two separate spe-
cies. The arrangement of rhizoids in the Panama collection
appears to approach more of a type B arrangement of
Kamiya et al. (2003) except for the occasional formation
of rhizoid filaments from dorsal periaxial cells and the
common appearance of adventitious branching. At this
time, there is not enough Panamanian material to properly
evaluate or make a substantial type collection; thus, it
remains herein identified as ‘C. rotundata’ until additional
material can be obtained.
DISCUSSION
Nine species of Caloglossa are now reported for the Amer-
icas: C. apicula, C. apomeiotica, C. confusa, C. fluviatilis,
C. intermedia, C. leprieurii, C. ogasawaraensis, C. rotundata
and C. ruetzleri. Caloglossa monosticha as reported by
Kamiya et al. (2003) is excluded from the marine flora of
the Americas. The taxon going under the name ‘C. mono-
sticha’ from the western Atlantic is herein identified as a
new species, C. confusa. Caloglossa confusa and C. mono-
sticha do not represent a monophyletic group when
analyzed with other members of the C. continua complex
based on rbcL and LSU rDNA sequence data, and their
blade shape and length also demonstrate that these are two
different species. Rhizoidal arrangement and sequence data
likewise support the premise that C. confusa is most closely
related to the members of the C. continua complex. The
collection from Panama that was putatively identified as
C. beccarii by Lin et al. (2001) is described as a new strictly
freshwater species, C. fluviatilis. Although these two taxa
contain adventitious branching and according to Sato &
Akiyama (2001) C. beccarii can also be found in freshwater
localities, these species differ from one another in rhizoidal
arrangement and degree of constriction at the thallus nodes.
Both rbcL and LSU rDNA sequence data further support
the conclusion that they are separate species (Figs 1, 2). In
addition, C. fluviatilis is part of the monophyletic group
528 Phycologia, Vol. 51 (5), 2012
that contain Caloglossa species, whose only mode of
secondary branching is adventitious.
The nine species reported here are distinct based on
molecular sequence data and vegetative characters. All nine
species are well supported by rbcL and LSU rDNA data,
except for C. fluviatilis, as we have included only one
sequence for both markers since it is known only from the
type locality (Figs 1, 2). Additionally, interspecific sequence
divergence for rbcL of the nine species ranges between 2.7%
and 14.1%. It has been shown that a sequence divergence,
for rbcL, of at least 2.7% between Caloglossa taxa indicates
they are separate species (Krayesky et al. 2011). These
species can furthermore be differentiated from one another
based on vegetative characters that include rhizoidal
morphology, degree of constriction at the thallus nodes,
secondary branching pattern, blade morphology, blade
width, number of cell rows cut off from the first axial cell
of the main axis, presence or absence of an adaxial cell
row derived from first axial cell of lateral axis and, when
present, the position of the cystocarp on the blades
(terminal vs terminal and subterminal).
Caloglossa in all likelihood is more diverse in Central
America than has been reported. For example, the collection
from the Bay of Panama, Pacific Panama, is most closely
allied with C. rotundata based on its morphological affinities
and rbcL sequence data. Unfortunately, lack of sufficient
material limits comparative morphological studies to
determine if it is a new or previously described species;
hence, it is tentatively referred to as ‘C. rotundata’.
Of all the species reported in this study, C. ogasawaraensis
has the broadest distribution, being nearly pantropical and
also found in subtropical to temperate localities. Other
Caloglossa species appear more restricted in their range than
C. ogasawaraensis, suggesting it may encompass more than
one taxon. A worldwide revision of C. ogasawaraensis is
needed to better understand the intraspecific relationships
within this species and to assess if it actually encompasses
more than one species. More specimens from regions
outside the Americas, specifically the region of the type
locality, are needed to accurately understand the intraspe-
cific relationships within C. ogasawaraensis.
The present study also demonstrates that, according to
rbcL and LSU sequence data, Caloglossa is monophyletic
based on the ML phylogenies. This is likewise supported by
the studies of Kamiya et al. (2003), as their MP and ML
phylogenies for their LSU data set of the genus were
likewise monophyletic. When we used an outgroup of both
T. perpusillum and Centroceras gasparrinii (Meneghini)
Kutzing (data not shown) the MP, ML and Bayesian
phylogenies demonstrated that Caloglossa was found to be
monophyletic and well supported.
ACKNOWLEDGEMENTS
The National Science Foundation Grants DEB PEET
0328491, DEB 091905, DEB 0936216 and OISE 0819205
who supported parts of this study. We would like to thank
all collectors cited in Table 1 and National Museum of
Natural History (US) for sending material used in this
study. In addition, we would also like to thank Giuseppe C.
Zuccarello, Craig, W. Schneider and Jerry G. Chmielewski
for their helpful remarks on this manuscript.
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Received 3 June 2011; accepted 20 January 2012
Associate Editor: Craig Schneider
530 Phycologia, Vol. 51 (5), 2012