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Classification and Taxonomic Revision of Two Oyster Species from Peru: Ostrea
megodon (Hanley, 1846) and Crassostrea talonata (Li & Qi, 1994)

Article  in  Journal of Shellfish Research · August 2017

DOI: 10.2983/035.036.0208

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Journal of Shellfish Research, Vol. 36, No. 2, 359–364, 2017.

CLASSIFICATION AND TAXONOMIC REVISION OF TWO OYSTER SPECIES FROM PERU:

OSTREA MEGODON (HANLEY, 1846) AND CRASSOSTREA TALONATA (LI & QI, 1994)

CUI LI,1,2,3 HAIYAN WANG1 AND XIMING GUO2*

1
Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Road, Qingdao, Shandong 266071,
China; 2Haskin Shellfish Research Laboratory, Department of Marine and Coastal Sciences, Rutgers
University, 6959 Miller Avenue, Port Norris, NJ 08349; 3University of Chinese Academy of Sciences,
19 Yuquan Road, Beijing 100049, China

ABSTRACT Oyster classification is problematic because of variations in shell morphology, and the use of molecular data may
help to confirm species identity and clarify taxonomic confusion. Two ostreid oysters were collected from Peru and identified as
Undulostrea megodon (Hanley, 1846) and Talonostrea talonata (Li & Qi, 1994), based on shell morphology. To confirm their
taxonomic status, fragments of mitochondrial cytochrome oxidase I and 16S rRNA genes were sequenced. Phylogenetic analyses
of both sequences do not support the validity of the two genera that were solely founded on these two species. Oyster U. megodon
is clearly a member of Ostrea and T. talonata a member of Crassostrea. Thus, U. megodon should retake its original name of Ostrea
megodon (Hanley, 1846), and T. talonata should be renamed as Crassostrea talonata (Li & Qi, 1994). Oyster C. talonata is a
described species of China, and its occurrence in western Pacific is unusual and probably because of human facilitated
introductions.

KEY WORDS: Undulostrea megodon, Talonostrea talonata, Ostrea, Crassostrea, oyster phylogeny, Ostreidae, Ostrea megodon,
Crassostrea talonata

INTRODUCTION California to Panama, Hyotissa hyotis (Linnaeus, 1758) from

Oysters are marine bivalve molluscs that are widely distrib-
uted in world estuaries and oceans. As sessile filter feeders and
sometimes reef builders, oysters play important roles in the
ecology of some coastal systems (Beck et al. 2011). Many
oysters are important fishery and aquaculture species support-
ing significant industries worldwide (Carriker & Gaffney 1996,
Guo et al. 1999, Guo 2009). Despite the ecological and
economical importance of oysters, their diversity and distribu-
tion are not well understood. The exact number of oyster species
is unknown, and many of the described species may be
synonyms (Harry 1985). New species are still being discovered
(Wu et al. 2013, Xia et al. 2014). Oysters are difficult to identify
and classify because of the lack of reliable morphological
characters. Traditional classification based on shell morphol-
ogy that is highly susceptible to environmental influence has
resulted in misidentifications and taxonomic confusions. It is
only in the recent two decades that molecular techniques have
been applied to oyster classification (Banks et al. 1993, Foighil
et al. 1995, Wang et al. 2004, Lam & Morton 2006, Shilts et al.
2007, Reece et al. 2008, Polson et al. 2009, Wang et al. 2010,
Lazoski et al. 2011, Liu et al. 2011). Still, many species have not
been investigated with molecular tools.
Extent oysters are found in two families, Gryphaeidae and
Ostreidae, with the latter containing most of the species. Two
genera of Ostreidae, Ostrea and Crassostrea, are most speciose
and challenging to classify where cryptic species may exist (Liu
et al. 2011). The eastern Pacific Coast is rich in bivalve species
(Coan & Valentich-Scott 2012). More than 10 oyster species
have been previously reported along the Pacific Coast of
Americas including Crassostrea columbiensis (Hanley, 1846)
found from the Gulf of California to Ecuador (Lohan et al.
2015), Crassostrea corteziensis (Hertlein, 1951) from the Gulf of
*Corresponding author. E-mail: xguo@hsrl.rutgers.edu
DOI: 10.2983/035.036.0208
Mexico to Ecuador (Bieler et al. 2004), Hyotissa quercina
(Sowerby II, 1871) from Mexico to Peru, Ostrea angelica
(Rochebrune, 1895) from the Gulf of California to Ecuador
(Hertlein & Emerson 1956), Ostrea chilensis in Chile (Foighil
et al. 1999), Ostrea conchaphila (Carpenter, 1857) from Punta
Eugenia in Mexico to Panama, Ostrea lurida (Carpenter, 1864)
from Guerrero Negro in Mexico to British Columbia, Canada,
Saccostrea palmula from Mexico to Ecuador, Striostrea pris-
matica (Gray, 1825) from the Gulf of California to Peru (Loor
& Sonnenholzner 2016), and Undulostrea megodon from Mex-
ico to Peru (Harry 1985, Huber 2010, Coan & Valentich-Scott
2012). Not all of these species have been genetically confirmed,
and taxonomic status of some species remains uncertain (Salvi
et al. 2014, Raith et al. 2015). In addition to these native species,
several nonnative species have been introduced to or identified
along the Pacific Coast of Americas including Crassostrea gigas,
Crassostrea sikamea, and Crassostrea virginica, and Ostrea
equestris (Guo 2009, Raith et al. 2015).
Oyster specimens were collected from Peru and identified as
Undulostrea megodon (Hanley, 1846) and Talonostrea talonata
(Li & Qi 1994), based on shell morphology (Guo unpublished).
Oyster U. megodon is a common species found in Peru with
distribution extending to the Gulf of California in Mexico. It
was first identified as Ostrea megodon (Hanley, 1846) based the
type specimen from Peru (Coan & Kabat 2012) and later
assigned to the new genus Undulostrea by Harry (1985) as the
only species. It has not been characterized molecularly, and the
taxonomic status of genus Undulostrea needs genetic confirma-
tion. Oyster T. talonata is a described and common species from
China (Li & Qi 1994) that has not been reported anywhere else.
Its unexpected finding in Peru calls for genetic confirmation.
In this study, DNA sequences of two mitochondrial genes,
cytochrome oxidase subunit I (COI) and 16S rRNA, were
obtained from the two oysters from Peru. Phylogenetic analyses
suggest that Undulostrea megodon is a member of the eastern

359
360 LI ET AL.

Pacific Ostrea group and should retake its original name Ostrea TABLE 1.
megodon (Hanley, 1846), and Talonostrea talonata is a member Accession numbers of sequences used for phylogenetic analysis
of the western Pacific Crassostrea group and should be named including sequences obtained in this study (*) and those from
as Crassostrea talonata. the National Center for Biological Information database.

MATERIALS AND METHODS Accession numbers

Sample Collection Species 16S rRNA COI

Ostrea angelica KT317142.1 KT317451.1
On October 16, 2015,14 oysters were collected from Sechura Ostrea conchaphila KT317155.1 KT317465.1
Bay (–5.841279, –80.966087), Piura, in northern Peru. The Ostrea megodon KX364274* KX364276*
oysters were collected from a depth of about 8 m and found Ostrea chilensis AF052065.1 AF112289.1
attached to Peruvian scallops (Argopecten purpuratus). Whole Ostrea denselamellosa FJ743511.1 KP067908.1
oysters were fixed in 95% ethanol and transported to the Ostrea futamiensis LC051609.1 LC051627.1
Haskin Shellfish Research Laboratory, Rutgers University, Ostrea circumpicta AB898282.1 AB898294.1
for analysis. Ostrea edulis KJ818215.1 KJ818235.1
Ostrea stentina AF052074.1 DQ226522.1
Ostrea puelchana AF052073.1 DQ226518.1
DNA Extraction, Amplification, and Sequencing
Crassostrea angulate KC847117.1 KP216803.1
DNA was extracted from adductor muscle using the E.Z.N.A. Crassostrea gigas AF280611.1 KP099052.1
Mollusc DNA kit (OMEGA, #D3373–02). A fragment of Crassostrea hongkongensis KC847120.1 KP976208.1
Crassostrea zhanjiangensis JX899653.1 JX899646.1
COI was amplified with universal primers: LCO1490 (5#-
Crassostrea dianbaiensis AB972006.1 LC063808.1
GGTCAACAAATCATAAAGATATTGG-3#) and HCO2198
Crassostrea ariakensis FJ743507.1 KF272859.1
(5#-TAAACTTCAGGGTGACCAAAAAATCA-3#) Crassostrea nippona HM015198.1 HM015198.1
(Folmer et al. 1994), and a fragment of 16S rRNA was Crassostrea sikamea KC847116.1 AB904878.1
amplified by 16Sar (5#-CGCCTGTTTATCAAAAACAT-3#) Crassostrea belcheri AY160758.1 AY038077.1
and 16Sbr (5#-CCGGTCTGAACTCAGATCACGT-3#) Crassostrea madrasensis JF915518.1 GU591435.1
(Palumbi et al. 1991). Polymerase chain reaction (PCR) was Crassostrea brasiliana DQ839413.1 FJ717651.1
conducted in 25 ml with 12.5 ml Taq 2x master mix (New Crassostrea virginica KC429253.1 KF644145.1
England Biolabs, #M0270), 0.2 mM of each primer, and 10 ng Crassostrea columbiensis – KP455055.1
of template DNA. Crassostrea corteziensis KT317088.1 –
Crassostrea rhizophorae JN849107.1 KP455050.1
Polymerase chain reaction was performed on a Bio-Rad
Crassostrea sp. (China) KC847133.1 KC683515.1
iCycler with the following cycling profile: initial denature at
Crassostrea talonata (Peru) KX364275* KX364277*
95°C for 30 sec, 30 cycles of 95°C for 30 sec, 48°C (16S) or 51°C Hyotissa imbricate KC847136.1 AB076917.1
(COI ) for 1 min, and 68°C 1 min, and a final extension at 68°C Hyotissa hyotis AY376599.1 GQ166583.1
for 5 min.
Products of PCR were cleaned with rAPid alkaline phos-
phatase (Roche, Germany). Sequencing was performed in both
directions on an ABI 3500XL Genetic Analyzer (Applied HKY + I + G, respectively. Bayesian analysis for 16S rRNA was
Biosystems). carried out using the program MrBayes V.3.2.6 (Ronquist et al.
2012). The chain was run for 20 million generations for 16S
Phylogenetic Analyses rRNA and sampled parameters and trees every 10,000 genera-
tions, discarding 25% of the total sampled trees from the
Sequences from both directions obtained in this study were beginning of the chain as burnin. Bayesian posterior probabil-
aligned and manually merged. Sequences of other oyster species ities were obtained from a tree with clade credibility values. The
used for phylogenetic analysis were downloaded from the COI model was run for 10 million generations, logged tree every
National Center for Biological Information database (Table 5,000 generations with a burnin percentage of 25.
1). All sequences were aligned using CLUSTALW (Thompson
et al. 1994). The species Hyotissa imbricata (Lamarck, 1819) RESULTS
and Hyotissa hyotis (Linnaeus, 1758) of Gryphaeidae were
chosen as the outgroup, based on their close phylogenetic Shell Morphology
relationship with Ostreidae within the superfamily Ostreoidea
(Rafinesque, 1815) (Salvi et al. 2014). K2P distance model was The 14 oysters fell into two distinct groups in shell mor-
used to measure genetic divergences for pairwise comparisons phology. The first group consisted of 13 specimens whose shell
and average distance within genera, as part of MEGA6 morphology matched that of Undulostrea megodon (Hanley,
(Kimura 1980, Tamura et al. 2013). 1846), a common oyster of Peru (Harry 1985). These oysters are
Bayesian inference (BI) approach was used for separate flat with thin shells that are 45.0 ± 4.6 mm in height, 32.4 ±
phylogenetic analyses of 16S rRNA and COI sequences. The 3.6 mm in length, and 7.6 ± 0.7 mm in width. The shells are
best fitting models for BI analyses were selected using characterized by curving backward in the shape of an ‘‘L’’ or
jModelTest version 2.1 (Darriba et al. 2012) based on the a ‘‘C’’ with two to four large undulations along the anteroven-
Bayesian information criteria (Schwarz 1978). The selected tral margin (Fig. 1A). The right valve is purple, and the color of
models for 16S rRNA and COI genes were TPM2uf + G and the left valve is lighter than that of the right valve. The surface is
 CLASSIFICATION OF OYSTER SPECIES FROM PERU
Figure 1. Shell morphology of Ostrea megodon (A) and Crassostrea
talonata from Peru (B) compared with C. talonata from China (C). The
scale bar represents 10 mm.
smooth, without scales, but circular growth patterns are visible.
The interior of both valves is smooth and white, and some
chalky deposits are present at the margin of the shells. It has
4–10 chomata along the inner margin immediately next to the
hinge on either side. The hinge is short, without umbo cavity.
These characteristics matched shell morphology of U. megodon,
a common oyster of Peru, as described by Harry (1985).
The second group was represented by only one specimen,
which closely resembles Talonostrea talonata (Li & Qi, 1994)
(Fig. 1B, C). The shells are 22 mm in height, 16 mm in length,
and 5 mm in width. The left valve is convex and has seven strong
ribs radiating from hinge area to the valve edge, whereas the
right valve is flat and smooth. The left valve is reddish purple,
and the right valve is yellowish brown. There is no chomata
along the valves. These characteristics match description of T.
talonata as described by Li and Qi (1994).
Sequence and Phylogenetic Analysis
Three specimens representing Undulostrea megodon and the
only specimen of Talonostrea talonata were sequenced. These
sequences have been submitted to the National Center for
Biological Information GenBank under access numbers
KX364274–KX364277. The U. megodon sequences contained
one haplotype for COI and one haplotype for 16S. Searches in
GenBank indicate that these sequences are novel and do not
match that of any species listed in GenBank. This is expected as
there is no entry for U. megodon in GenBank.
361
The Talonostrea talonata sequences from Peru matched
that of an unnamed oyster Crassostrea sp. in GenBank
(KC847133 and KC683515). This unnamed oyster was col-
lected from Beibu Bay, China, in an unpublished study. Shell
morphology of this unnamed Crassostrea oyster (Fig. 1C)
matches that of T. talonata described by Li and Qi (1994)
as well as T. talonata from Peru as described previously. They
are similarly small in size and have five to seven strong ribs
radiating from the hinge area to shell margin on the left valve
(Fig. 1C). There is no sequence entry for T. talonata in
GenBank.
The final sequence set for phylogenetic analysis contained 28
taxa and was 437bp in length for 16S and 594bp for COI. The BI
tree constructed with COI (Fig. 2A) has a higher resolution than
that from 16S (Fig. 2B).
On both trees, Undulostrea megodon is found within a mono-
phylogenetic clade of Ostrea species (Fig. 2). Thus, U. megodon
belongs to the genus Ostrea and should retake its original name
Ostrea megodon (Hanley, 1946). On the COI tree that has higher
resolution, O. megodon is clustered with Ostrea lurida, Ostrea
conchaphila, Ostrea puelchana, Ostrea permollis, Ostrea sten-
tina, and Ostrea angelica, all Ostrea species from eastern Pacific,
adding phylogeographic support (Fig. 2A). Oyster O. megodon
is most closely related to O. puelchana as indicated by their K2P
genetic distance of 0.13 in COI (Table 2). This distance is
significantly higher than that between some of the other
recognized species.
On both phylogenetic trees, Talonostrea talonata from Peru
is clustered with Crassostrea sp. from China that also matches
T. talonata in morphology, and both are found within the clade
of Crassostrea (Fig. 2). Therefore, these T. talonata oysters
belong to Crassostrea and should be renamed as Crassostrea
talonata (Li & Qi, 1994). On both trees, C. talonata from Peru is
clustered with western Pacific species rather than eastern Pacific
species, suggesting that it is nonnative to Peru. The COI K2P
distance between C. talonata from China and Peru is 0.036
(Table 3) which is considerably lower than most interspecific
distances within Crassostrea, but similar to the distance between
Crassostrea gigas and Crassostrea angulata which have recently
been classified as two subspecies Crassostrea gigas gigas and
Crassostrea gigas angulata (Wang et al. 2010). The closest
relative of C. talonata is Crassostrea zhanjiangensis, with a
COI P2K distance of 0.174.
DISCUSSION
The two ostreid oysters from Peru were identified as
Undulostrea megodon (Hanley, 1846) and Talonostrea talonata
(Li & Qi, 1994), based on shell morphology. Although oyster
shells are highly variable, morphological identification of these
two species is relatively straightforward and unambiguous.
Oyster U. megodon is a common oyster of Peru that is readily
recognized by local biologists. It is characterized by two to four
undulations along its thin and flat shells (Harry 1985), which
matched specimens of this study. The specimens collected from
Peru, the type location, are clearly U. megodon. Oyster T.
talonata is a well-recognized species in China (Zhang & Lou
1956, Li & Qi 1994). It is characterized by its small size and five to
seven strong radiating ribs on its left valve. Although only one
specimen is obtained, the specimen is small and has the char-
acteristic ribs on its left valve. It was immediately recognized as
362 LI ET AL.

Figure 2. Bayesian inference trees of representative oysters based on COI (A) and 16S rRNA (B) sequences. Bayes posterior probability (BPP)
percentages greater than 50 are given for each node. In the Crassostrea clade, Asia–Pacific branches are in red and Atlantic species are in blue.

T. talonata at first sight, and its classification is confirmed by
DNA sequence analyses.
Oysters Undulostrea megodon and Talonostrea talonata were
sole founding members of their respective genera. Oyster U.
megodon was initially placed in the genus Ostrea as Ostrea
megodon by Hanley (1846). Harry (1985) moved it out of Ostrea
and established the new genus Undulostrea for its uniquely
shaped and undulated thin and flat shells. Phylogenetic analyses
with both COI and 16S sequences unambiguously place U.
megodon within a clade of Ostrea species, and therefore, U.
megodon should be move back into Ostrea and retake its
original name O. megodon (Hanley, 1846). The genetic distances
between O. megodon and its sister species are consistent with
within-genus distances and far below between-genus distances

TABLE 2.
K2P genetic distances in COI (below diagonal) and 16S (above diagonal) genes among representing species of Ostrea, Crassostrea,
and Hyotissa, showing Ostrea megodon is an independent species belonging to Ostrea.

Species 1 2 3 4 5 6 7 8 9 10 11 12
1 Crassostrea gigas – 0.210 0.214 0.200 0.207 0.214 0.187 0.201 0.218 0.204 0.217 0.360
2 Ostrea conchaphila 0.281 – 0.012 0.030 0.028 0.049 0.071 0.052 0.054 0.049 0.023 0.342
3 Ostrea angelica 0.265 0.098 – 0.028 0.028 0.043 0.065 0.052 0.049 0.051 0.023 0.342
4 Ostrea puelchana 0.285 0.122 0.141 – 0.038 0.046 0.070 0.036 0.044 0.060 0.036 0.358
5 Ostrea stentina 0.266 0.131 0.130 0.116 – 0.057 0.082 0.063 0.060 0.068 0.030 0.374
6 Ostrea circumpicta 0.308 0.212 0.205 0.186 0.200 – 0.068 0.065 0.062 0.059 0.057 0.365
7 Ostrea edulis 0.312 0.186 0.186 0.192 0.217 0.193 – 0.057 0.085 0.082 0.079 0.330
8 Ostrea chilensis 0.313 0.216 0.171 0.200 0.229 0.197 0.142 – 0.060 0.068 0.060 0.358
9 Ostrea denselamellosa 0.304 0.195 0.195 0.190 0.192 0.210 0.198 0.187 – 0.071 0.057 0.374
10 Ostrea futamiensis 0.302 0.195 0.199 0.223 0.195 0.206 0.190 0.207 0.208 – 0.068 0.358
11 Ostrea megodon 0.317 0.144 0.166 0.130 0.184 0.216 0.162 0.184 0.221 0.213 – 0.354
12 Hyotissa hyotis 0.544 0.462 0.520 0.449 0.461 0.512 0.459 0.502 0.528 0.496 0.471 –
 CLASSIFICATION OF OYSTER SPECIES FROM PERU 363

TABLE 3.
K2P genetic distances in COI (below diagonal) and 16S (above diagonal) genes among representing species of Crassostrea, Ostrea,
and Hyotissa, showing Crassostrea talonata belongs to Crassostrea.

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
1 C. brasiliana – 0.054 0.113 0.121 0.160 0.156 0.150 0.156 0.162 0.156 0.147 0.153 0.160 0.187 0.187 0.187 0.219 0.355
2 C. columbiensis 0.190 – 0.122 0.136 0.189 0.185 0.182 0.185 0.192 0.189 0.178 0.185 0.189 0.204 0.207 0.207 0.233 0.381
3 C. rhizophorae 0.232 0.265 – 0.030 0.157 0.154 0.157 0.160 0.179 0.172 0.176 0.169 0.170 0.194 0.211 0.208 0.197 0.355
4 C. virginica 0.253 0.289 0.191 – 0.157 0.154 0.154 0.157 0.164 0.166 0.173 0.163 0.167 0.182 0.188 0.185 0.203 0.339
5 C. angulata 0.252 0.313 0.253 0.248 – 0.005 0.012 0.020 0.041 0.049 0.052 0.043 0.046 0.104 0.107 0.110 0.187 0.360
6 C. gigas 0.252 0.330 0.268 0.248 0.031 – 0.018 0.020 0.041 0.049 0.052 0.043 0.052 0.104 0.107 0.110 0.187 0.360
7 C. sikamea 0.231 0.279 0.251 0.260 0.119 0.138 – 0.018 0.044 0.041 0.043 0.035 0.038 0.098 0.110 0.113 0.175 0.352
8 C. hongkongensis 0.253 0.294 0.289 0.275 0.130 0.146 0.153 – 0.036 0.033 0.041 0.035 0.036 0.087 0.095 0.098 0.171 0.364
9 C. nippona 0.259 0.286 0.266 0.257 0.169 0.162 0.174 0.138 – 0.065 0.062 0.057 0.057 0.098 0.101 0.104 0.200 0.359
10 C. ariakensis 0.267 0.281 0.279 0.294 0.149 0.155 0.156 0.144 0.148 – 0.059 0.054 0.051 0.092 0.098 0.101 0.165 0.348
11 C. dianbaiensis 0.260 0.273 0.286 0.290 0.167 0.187 0.185 0.173 0.193 0.162 – 0.020 0.049 0.081 0.087 0.090 0.172 0.357
12 C. madrasensis 0.272 0.274 0.301 0.263 0.170 0.170 0.205 0.168 0.175 0.185 0.110 – 0.038 0.087 0.092 0.095 0.175 0.357
13 C. belcheri 0.320 0.336 0.291 0.289 0.211 0.211 0.202 0.216 0.219 0.207 0.203 0.206 – 0.098 0.110 0.113 0.181 0.374
14 C. zhanjiangensis 0.295 0.308 0.281 0.292 0.248 0.259 0.211 0.234 0.223 0.278 0.233 0.240 0.275 – 0.023 0.020 0.184 0.356
15 C. sp. (China) 0.322 0.319 0.293 0.308 0.243 0.251 0.243 0.230 0.222 0.265 0.239 0.265 0.268 0.198 – 0.002 0.197 0.360
16 C. talonata (Peru) 0.294 0.319 0.293 0.292 0.240 0.254 0.214 0.233 0.218 0.265 0.229 0.257 0.253 0.174 0.036 – 0.194 0.356
17 O. edulis 0.345 0.342 0.334 0.330 0.317 0.312 0.289 0.307 0.284 0.297 0.258 0.278 0.312 0.260 0.311 0.311 – 0.330
18 H. hyotis 0.579 0.590 0.548 0.564 0.512 0.544 0.512 0.535 0.498 0.537 0.526 0.520 0.559 0.520 0.540 0.540 0.459 –

(Table 2). Thus, there is no justification of placing O. megodon
into a different genus. As U. megodon is the only species
supporting Undulostrea, moving it into Ostrea invalidates
Undulostrea as a genus and Undulostreini as a tribe (Harry
1985).
Similarly, phylogenetic analyses unambiguously place Talo-
nostrea talonata into Crassostrea, and T. talonata should be
renamed Crassostrea talonata. As T. talonata is the founding
species of the genus, moving it to Crassostrea also invalidates
Talonostrea as an independent genus. Three other genera,
Myrakeena, Ostreola, and Cryptoostrea, established by Harry
(1985) based on shell morphology, are also not supported by
DNA sequence data (Salvi et al. 2014), highlighting the need for
molecular reassessment of oyster taxonomy. Although it has
not been widely accepted, a recent study has proposed that
eastern and western Pacific Crassostrea species should belong
to two separate genera Crassostrea and Magallana (Salvi &
Mariottini 2017), respectively, based on the large genetic
distances between them (Salvi et al. 2014, Salvi & Mariottini
2017). If so, perhaps Talonostrea can be accepted as an
independent genus based on genetic distances (Table 2). Other
than genetic distances, Pacific and Atlantic species of Crassos-
trea are very similar in size, shell morphology, and biology.
Consensus is needed about whether genera should be estab-
lished by sequence data alone. Before genus Magallana is
accepted, T. talonata should be named as C. talonata as it is
well within the same clade on phylogenetic trees.
The observation of Crassostrea talonata in Peru, a described
species of China that has not been reported elsewhere, is
unexpected and intriguing. Oyster C. talonata is clearly a west-
ern Pacific species, as it is clustered with other western Pacific
species on the phylogenetic tree (Fig. 2). No oysters of the same
species or different species within such small genetic distances are
known to occur naturally on both sides of the Pacific. Therefore,
it is likely that C. talonata in Peru is a nonnative species from Asia
possibly introduced through human activities. Human activities
have facilitated the introduction of several oyster species, in-
cluding Crassostrea angulata from China to Europe, Crassostrea
gigas from Asia to many countries, Crassostrea sikamea and
Crassostrea ariakensis from Japan to the United States of
America, Ostrea equestris from Atlantic to Baja California
(Guo 2009, Raith et al. 2015).
It is worth noting that whereas the genetic distance between
species Crassostrea talonata from Peru and that from China is
below the interspecific distances among most known species, it is
comparable with that between Crassostrea gigas gigas and
Crassostrea gigas angulata that are considered as two subspecies
(Wang et al. 2010). Because only one sample from Peru and one
from China are available, genetic variation within the species
or species complex is unknown at this time. Further studies are
needed to sample C. talonata from the western and eastern Pacific
to determine if subspecies or species designation is warranted. It
is possible that C. talonata is widely distributed in Asian Pacific
and these in Peru may be from a different subspecies away from
southern China where Crassostrea sp. was collected.
In conclusion, two ostreid oysters from Peru were identified
as Undulostrea megodon and Talonostrea talonata, based on
shell morphology, but phylogenetic analysis with COI and 16S
sequences indicated that they should be renamed Ostrea
megodon and Crassostrea talonata, respectively. Oyster C.
talonata is a new record for Peru and likely a nonnative species
from the Asian Pacific. It would be interesting to study how and
where C. talonata originated in Asia.
ACKNOWLEDGMENTS
We thank Eric Hanschke for helping with sample collection.
This study is conducted at Rutgers University where CL was
a visiting student. We acknowledge partial support from
Rutgers University, USDA/NJAES project 1004475/NJ32920,
‘‘Taishan Overseas Scholar’’ project of Shandong, and China
Ocean Program (XDA10020303).
 364 LI ET AL.
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