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Egg-collar morphology and identity of nine species of Naticidae (Gastropoda)

in Taiwan, with an assessment of their phylogenetic relationships
Article in Journal of Molluscan Studies · November 2018

DOI: 10.1093/mollus/eyy041
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Journal of The Malacological Society of London
Molluscan Studies
Journal of Molluscan Studies (2018) 84: 354–378. doi:10.1093/mollus/eyy041
Advance Access publication date: 19 September 2018
Egg-collar morphology and identity of nine species of Naticidae (Gastropoda)

in Taiwan, with an assessment of their phylogenetic relationships
Downloaded from https://academic.oup.com/mollus/article-abstract/84/4/354/5102386 by guest on 01 November 2018

Dun-Ru Kang1, Koh Siang Tan2 and Li-Lian Liu1
1
Department of Oceanography, National Sun Yat-sen University, Kaohsiung, Taiwan 804, Republic of China; and
2
St John’s Island National Marine Laboratory, Tropical Marine Science Institute, National University of Singapore, 18 Kent Ridge Road, Singapore 119227
Correspondence: K.S. Tan; e-mail: [email protected] and L.-L. Liu; e-mail: [email protected]
(Received 28 August 2017; editorial decision 3 May 2018)
ABSTRACT
The gross morphology of egg (sand) collars laid by nine naticid species in six genera from Taiwan are
described and compared: Paratectonatica tigrina, Notocochlis cernica, N. gualtieriana, N. sp. aff. antoni, Naticarius
zonalis, Tectonatica bougei, Sinum haliotoideum (=planulatum), Mammilla melanostoma and M. melanostomoides. Egg
collars and adult animals were collected from Taiwan and its offshore islands, as well as from Four-Way
Closure Ridge (1,648 m depth) in the South China Sea. Mitochondrial COI and 16 S rRNA sequences
were obtained from both embryos and adults and the genetic distances of these sequences from maximum-
likelihood trees were then compared to determine the identity of the egg collars. Of the egg collars of nine
species identified using this procedure, seven are described here for the first time. Each species laid a con-
sistent, distinctive egg collar that differed in the overall shape and size, number of coils, capsule size and
packing density in different species. The largest collars, with basal diameters exceeding 8 cm, were laid by
M. melanostoma and S. haliotoideum, while the smallest belonged to Notocochlis spp., with basal diameters as
small as 1 cm. In the egg collars of all nine species examined, each embryo was enclosed within a single
capsule. COI and 16 S sequences from egg collars were analysed together with published sequences from
other studies. Seven groups, each with a unique egg-collar morphology, were defined, some of which were
coincident with supported phylogenetic clades. These results represent only a small fraction of some 47
species of Naticidae recorded to date from Taiwan, but the use of a combination of morphological and
molecular characters appears to be a viable method to identify egg collars. The egg-collar literature involv-
ing 52 naticid species worldwide is briefly reviewed in the light of our results.
INTRODUCTION
The adults generally appear to abandon the sand collars after laying
Egg (or sand) collars laid by naticids are often a conspicuous item them.
on intertidal soft shores around the world where these snails are In the tropics, where multiple naticid species can co-exist
present. In these egg masses, egg capsules, sediment (chiefly sand, together on the shore, matching egg collar to species is a difficult
mud; occasionally faecal pellets) and adhesive are mixed together task. While the process of identifying adult shells has benefited
(Giglioli, 1955; Kingsley-Smith, 2003) to form a marvellously func- from recent taxonomic treatments of some fossil and living mem-
tional, but ephemeral structure. Ventilation, camouflage and pro- bers (chiefly based on features of the shell, umbilicus and opercu-
tection (Aronowsky, 2003) are provided for the embryos for some lum; e.g. Marincovich, 1977; Majima, 1989), egg collar
weeks until they are ready to swim away as veligers or crawl away morphology is poorly elucidated in comparison. This is true for
as juveniles. In addition, the characteristic form of the collar pos- most naticid species living in the Indo-Pacific region, where diver-
sibly helps keep it in position on the substratum when subjected to sity is highest with more than 200 species (Kabat, 1990). Some 47
hydrodynamic forces (personal observation), while retaining its species of Naticidae have been recorded from Taiwan (Yeh &
shape and form in a fluid environment. However, despite being eas- Hsueh, 2009). Naticids continue to be economically important as
ily observed in the field during low tide, the specific identities of the food along the coastlines of the western Pacific including China
collars are difficult to ascertain. Spawning takes place under water (Zhang, 2016) and Southeast Asia (Poutiers, 1998). Recently, a
(Giglioli, 1955), possibly rapidly, and the act of laying capsules is number of authors notably Hülsken (2008) and Hülsken et al.
rarely observed in the field, although some species are able to pro- (2008, 2011), working in the Mediterranean, southern Australia
duce egg collars in captivity (Polinices lacteus: Bandel, 1976; Euspira and the Caribbean using molecular techniques, have revealed that
alderi [as Polinices]: Ansell, 1982; Euspira fusca: Rámon, 1994; most species lay egg collars with species-specific morphologies, so
Laguncula pulchella [as Euspira fortunei]: Okoshi & Sato-Okoshi, 2011). that in some cases they have provided additional information to
© The Author(s) 2018. Published by Oxford University Press on behalf of The Malacological Society of London, all rights reserved.
For Permissions, please email: [email protected]
EGG COLLARS OF NATICIDAE
clarify the taxonomy of disputed species complexes (Hülsken et al., Collar dimensions (Fig. 2) were measured in the laboratory fol-
2011). The sequences have also been useful in revealing phylogen- lowing Amio (1955) and Giglioli (1955), with modifications.
etic structure in the Naticidae (Hülsken et al., 2012), although Collars were initially divided into 11 morphotypes based on their
there are still large gaps in our knowledge of generic relationships. morphology (Fig. 3). Collar morphotype and locality were used to
Here we present the results of similar investigations carried out on annotate each sequence obtained in the study and in the resulting
common naticid species living on the shores of Taiwan and nearby molecular phylogenetic trees (Fig. 3). The shell heights (SH) of
islands, as well as on Dongsha (Pratas) Island atoll and Four-Way adult animals were measured, except for Sinum, where the shell
Closure Ridge (1,648 m depth) in the South China Sea. length (SL) is provided. Maximum length (L) and width (W) of
The main objectives of our study were to (1) characterize egg opercula were measured. Diameter (D) of egg capsules was
collars of naticids obtained in the field using molecular sequences recorded. Voucher specimens were deposited in the National
and morphology; (2) identify the egg collars by matching their Museum of Natural Science (NMNS), Taichung, Taiwan
sequences with those of adult specimens and (3) determine the (Table 1).
phylogenetic positions of the identified species in relation to other Examination of morphological characteristics of naticid speci-
naticids using available molecular sequences. mens and egg collars were made using stereo- and compound

microscopes. As egg collars are flexible, measurements given are
approximate, based on the shape assumed in seawater. Hand sec-
MATERIAL AND METHODS tions of living and preserved egg collars were made using a razor
Egg collars and adult animals were collected by hand on intertidal blade and the relative distribution of eggs with respect to sand
shores, by SCUBA diving in shallow water, or purchased from grains was observed under the microscope. Where possible, mito-
local fish markets, in Taiwan and adjacent islands including chondrial COI and 16 S rRNA from adults and collars were
Penghu, Kinmen and Dongsha, between 2011 and 2015 (Kang, sequenced. These were then compared with sequences from
2014; Fig. 1). In one case a museum specimen was used. Despite Hülsken (2008). Classification of naticids used in this study broadly
numerous visits to the field, we were unable to observe the depos- follows Kabat (1990), Hülsken (2008) and Torigoe & Inaba (2011).
ition of egg collars by animals during low tide. Adults from a Crude DNA was extracted with the Tissue & Cell Genomic
deep-sea trawl at Four-Way Closure Ridge (1,648 m depth) off the DNA Purification Kit (GeneMark, DP021-150), following the
southern coast of Taiwan were also included in the analysis manufacturer’s instructions, with modifications. Part of the foot
(Table 1). They were all either frozen at −20 ˚C or preserved in muscle of adults (1–25 mg) was put in extraction solution and
95% ethanol. ground with a tissue homogenizer. A piece of collar (10–40 mg)
Figure 1. Collection localities of naticid egg collars and adults in Taiwan. A. Mainland Taiwan, Penghu (Pescadores Is), Liuqiu I. and Green I. (see insets).
B. Kinmen I. off Xiamen, China. C. Dongsha (Pratas Atoll), South China Sea. Each locality is provided with a list of species for which the adult snail (S)
and/or egg collar (E) were either collected and sequenced, or photographed only (P). Generic abbreviations: E., Euspira; G., Glossaulax; L., Laguncula; M.,
Mammilla; N., Natica; Na., Naticarius; No., Notocochlis; P., Paratectonatica; Po., Polinices; Ps., Pseudopolinices; S., Sinum; T., Tanea; Te., Tectonatica. Except at Four-Way
Closure Ridge (depth 1,648 m), all specimens were collected either in the intertidal or shallow subtidal zones.
355
D.-R. KANG ET AL.
Table 1. Collection localities in Taiwan (see Fig. 1 for map) of sequenced specimens and egg collars, with museum accession numbers (NMNS-7708-0xx)
in parentheses.
Species Adult animal Egg collar Habitat; remarks
Paratectonatica tigrina (Röding, Taishi (04); Siang-shan (03); Kinmen Kinmen (31–33) Intertidal
1798) (02)
Notocochlis cernica (Jousseaume, Not available Kenting (49) Subtidal; collar photographed in field
1874) only; COI and 16S sequences of
adults from Hülsken (2008)
Notocochlis gualtieriana (Récluz, Dongsha (19, 20); Penghu (21–24) Dongsha (34,60); Penghu (35, 36, Intertidal to subtidal
1844) 40); Kenting (37–39); Kinmen (41)
Notocochlis sp. aff. antoni Kinmen (16–18) Kinmen (55–57) Intertidal

Notocochlis sp. 1 Not available Kenting (59) Subtidal; COI sequences of adults
from Hülsken (2008)
Naticarius alapapilionis (Röding, Dongshi market (15) Not available Subtidal trawl
1798)
Naticarius zonalis (Récluz, 1850) Penghu (14) Penghu (50, 51) Intertidal
Natica vitellus (L., 1758) Tungkang market (01) Not available Subtidal trawl
Tectonatica bougei Sowerby, 1908 Not available Kenting (58) Subtidal; egg collar photographed in
field, not sequenced; 16S
sequences of adults from Hülsken
(2008)
Sinum haliotoideum (L., 1758) Dongsha (museum spec. NMNS- Penghu (42–46)
(= planulatum Récluz in Chenu, 007263-00015)
1843)
Sinum sp. Daxi market (05) Not available Deep-sea trawl
Mammilla melanostoma (Gmelin, Not available Penghu (48); Dongsha (47) COI and 16S sequences of adults
1791) from Hülsken (2008)
Mammilla melanostomoides (Quoy Not available Penghu (53); Daxi (52) COI and 16S sequences of adults
& Gaimard, 1832) from Hülsken (2008)
Polinices flemingianus (Récluz, Kenting (09) Not available Subtidal
1844)
Polinices sp. 1 Dongshi (06) and Tungkang (07) Not available Commercial subtidal trawl
markets
Polinices sp. 2 Tungkang market (08) Not available Commercial subtidal trawl
Glossaulax didyma (Röding, 1798) Tungkang (12) and Dongshi (11) Not available Commercial subtidal trawl
markets
Glossaulax sp. Tungkang market (10) Not available Commercial subtidal trawl
Tanea lineata (Röding, 1798) Tungkang (27, 28) and Dongshi Not available Commercial subtidal trawl
(29, 30) markets
Euspira cf. plicispira (Kuroda, 1961) 1648 m, Four Way Closure Ridge, Not available Research deep-sea trawl
22˚ 5.01’N, 119˚ 48.04’E (25)
Pseudopolinices cf. nanus (Møller, 1,648 m, Four Way Closure Ridge, Not available Research deep-sea trawl
1842) 22˚ 5.01’N, 119˚ 48.04’E (26)
Laguncula pulchella Benson, 1842 Husia, Kinmen (13) Not available Intertidal
undetermined species Not available Kenting (54) Subtidal (SCUBA)
was placed in extraction solution with 0.5 g of glass beads (Sigma (elongation); after the last cycle, conditions were set to 95 °C for
G-9268, 425–600 μm) and homogenized with a benchtop Mini- 10 s (denaturation); 39.5 °C for 10 s (annealing) and 72 °C for
BeadBeater (BioSpec 3110BX). Samples were then lysed and cen- 2 min for final elongation. Conditions for 16 S were as follows:
trifuged several times to precipitate nuclei and debris. Finally, 95 °C, 5 min (denaturation); 49 °C, 45 s (annealing) and 72 °C,
50 μl double-distilled water was added to the resulting DNA pel- 1 min (elongation), followed by another 38 cycles: 95 °C for 25 s
lets, which were stored at –20 °C for later use. (denaturation); 49 °C for 25 s (annealing) and 72 °C for 35 s
The primers used for PCR amplification of COI and 16 S (elongation); after the last cycle, conditions were set at 95 °C for
rRNA genes are listed in Table 2. Amplification was conducted in 10 s (denaturation); 49 °C for 10 s (annealing); and 72 °C for
a thermal cycler (Applied Biosystems 2720). PCR conditions were 2 min for final elongation. The amplified DNA was directly
those used by Hülsken (2008). The COI setting was as follows: sequenced on an automated DNA sequencer (Applied Biosystems
95 °C, 5 min (denaturation); 39.5 °C, 45 s (annealing) and 72 °C, 3730xl DNA Analyzer). Additional sequences from Hülsken
1 min (elongation), followed by another 38 cycles: 95 °C for 20 s (2008) were also used. Sequences of Cypraea tigris (Cypraeidae),
(denaturation); 39.5 °C for 35 s (annealing) and 72 °C for 40 s Planaxis sulcatus (Planaxidae) and Strombus luhuanus (Strombidae)
356

Figure 2. Egg collar of Paratectonatica tigrina, illustrating terms used and measurements made on egg collars. Abbreviations: AD, apical diameter; AM, apical
margin; BD, basal diameter; BM, basal margin; CH, maximum collar height; CT, collar thickness; CW, crown width; TW, total width.
from Colgan et al. (2007) and Hülsken (2008) were used as out- tree. The COI sequence of Euspira cf. plicispira obtained from
group taxa. Four-Way Closure Ridge in the South China Sea was consistent
The sequences were aligned with those of other naticid species with those from the Philippines obtained by Hülsken (2008).
from Hülsken (2008) (Table 1) using CLUSTAL W in MEGA v. 6 The COI analysis suggested that a cryptic species may be involved
(Tamura et al., 2013), using the Kimura 2-parameter model and in Notocochlis cernica. Here, the sequence of an individual from the
maximum-likelihood (ML) method (Kimura, 1980). Bootstrap Philippines (sequence number 85211 of Hülsken, 2008) was assigned
probability (BS) estimates (1,000 replicates) (Felsenstein, 1985) to N. cernica s. s. together with an egg collar from Kenting (Fig. 5B,
were also made to indicate robustness of nodes in neighbour- C). This species was distinguished from another Notocochlis species,
joining trees. here assigned to Notocochlis sp. 1, based on sequences from Australia
and Indonesia (85112 and 91111 of Hülsken, 2008), which clustered
RESULTS with an undetermined egg collar, also from Kenting.
Resolution at the generic level and below was poor in both trees.
Molecular data analysis Only the genera Naticarius, Tanea and Conuber received significant BS
support. Relationships between the genera were not resolved.
For the COI gene, 52 sequences representing 21 species (Table 1,
The deep-sea circumboreal naticid Pseudopolinices cf. nanus clus-
Fig. 1) were obtained from adults and egg collars. These were
tered with Laguncula pulchella (=Euspira fortunei of authors; it is dis-
based on 29 adult animals and 23 collars, the latter representing
puted that L. pulchella may be a freshwater viviparid or assimineid)
eight naticid species, respectively. These sequences were analysed
in the 16 S tree (BS = 85%; Fig. 3B). No COI sequence was avail-
together with 51 naticid species sequenced by Hülsken (2008).
able for Pseudopolinices.
A total of 52 16 S sequences were examined, corresponding to
21 species (Table 1; Fig. 1). These sequences were based on 23
adult animals and 29 egg collars, of which collars represented ten Morphology
naticid species. They were analysed together with the sequences of
a further 40 naticid species from Hülsken (2008). A total of nine egg collars were matched with the corresponding
The resulting sequence lengths of COI and 16 S for naticid spe- adult species based on genetic distances that fell within the intra-
cies were approximately 410 and 400 bp, respectively (Supplementary specific range assessed for adults (Table 3). These comprised six
Material Tables S1, S2). In general, species sequence divergences of naticine, one sinine and two polinicine species in six genera. The
COI were greater than those of 16 S (see Table 3). For COI, the six naticines were Paratectonatica tigrina, Notocochlis cernica, N. gualtieriana,
within-species ranges were between 0.000 and 0.035 for Notocochlis gual- N. sp. aff. antoni, Naticarius zonalis, and Tectonatica bougei (based on an
tieriana and between 0.014 and 0.028 for N. sp. aff. antoni. For 16 S, the undetermined collar from Kenting); the two polinicines were
within-species range was between 0.000 and 0.018 for Sinum haliotoi- Mammilla melanostoma and M. melanostomoides., and the single sinine
deum, while Paratectonica tigrina, Naticarius zonalis and Mammilla melanosto- was Sinum haliotoideum. One egg collar was identified as Notocochlis sp.
moides showed no variation in genetic distance across individuals. 1 (based on an undetermined collar from Kenting; morphotype 9 in
Within genera, sequence divergences ranged from 0.046–0.079 Fig. 3) and another as a possible Euspira or Tanea sp. (also based on
(Mammilla) to 0.053–0.222 (Naticarius) for COI, and 0.013–0.166 an undetermined collar from Kenting; morphotype 8 in Fig. 3).
(Naticarius) to 0.063–0.093 (Sinum) for 16 S. Descriptions of the adult shell, operculum, animal (where avail-
COI and 16 S sequences provided by Hülsken (2008) and able) and egg collar (see Fig. 2 for terms and measurements used)
Colgan et al. (2007) were analysed together with those obtained in for each species are provided below.
this study. At the species level, COI and 16 S sequences of adults
and egg collars of the same species from different localities always Paratectonatica tigrina (Röding, 1798)
grouped together into the same clades with good BS support— (Fig. 4)
see, for example, M. melanostoma and N. zonalis (Fig. 3; both COI
and 16 S trees). However, there was one undetermined egg collar Shell (Fig. 4A, B): small, SH 21.5 mm; high-spired; suture narrow,
species (Morphotype 8 Kenting in Fig. 3), which clustered with the not deep; periostracum thin, yellow brown with fine axial lines.
Tanea clade in the COI tree but grouped with Euspira in the 16 S Several rows of dark brown to black, sometimes overlapping
357
D.-R. KANG ET AL.
dashes over yellowish white background. Surface of last whorl operculum. The two ribs separated by a narrow groove. Internal
smooth, with no axial undulations. Aperture wide, white; colu- surface smooth, greenish yellow, with thin corneous layer. L
mella narrow; umbilicus open, funicle relatively small. 13.0 mm, W 8.1 mm (for SH 21.5 mm).
Operculum (Fig. 4D, E): calcareous, white, external surface slightly Animal (Fig. 4C): foot uniformly translucent grey. Tentacles trans-
concave, with 2 narrow semicircular flattened ribs that terminate lucent grey, with anterior edges lined with black.
at inner edge adjacent to columella, where operculum is thickest
(1.5 mm). Outer rib at edge of operculum, where it is thinnest, Egg collar (Fig. 4F–H; n = 3): 1–1.5 whorls, basal margin wavy in
bears numerous fine, flattened granules along entire outer edge. horizontal plane. Apical margin smooth, not thickened. BD
Inner rib generally smooth, with some granules near edges of 42–58 mm; AD 15–24 mm, about half BD; TW 17–23 mm; CT
Figure 3. Continued on next page
358
Figure 3. Maximum-likelihood trees based on mitochondrial COI and 16 S sequences obtained from egg collars and adult Naticidae in Taiwan, combined
with additional data from Hülsken (2008). A. Mitochondrial COI tree. B. Mitochondrial 16 S rRNA tree. Roman numerals I–VII represent species groupings
based on the suggested egg-collar classification (Table 4). Species sequenced by Hülsken are marked (H), together with localities and original reference num-
bers of Hülsken (2008: appendix 6.1). Egg collars sequenced in this study are labelled according to a priori Morphotypes 1–11, with collection localities in
Taiwan; adults are labelled with collection localities only; reference numbers following localities correspond to voucher specimens deposited in National
Museum of Natural Science, Taichung (see Table 1 for accession numbers).
359
D.-R. KANG ET AL.
Table 2. Forward (F) and reverse (R) PCR primers used for PCR amplifi- lines; shell colour varies between having numerous (about 15) dis-
cation of COI and 16S rRNA genes, following Hülsken (2008). continuous brown axial lines interrupted by two whitish spiral
bands, to almost uniformly grey-brown with only faint evidence of
Gene Name Sequence 5′ to 3′
axial lines. Anterior region of shell usually white, as is area just
COI P388 (F) GCTTTTGTTATAATTTTYTT below suture defining the separation between penultimate and last
COI P390(R) CGATCAGTTAAARTATWGTAAT
whorls. Surface of last whorl with 18–20 narrow, raised axial
undulations that become flattened towards base of shell. Aperture
16S P744 (F) ATARRGAGTCGGACCTGCCCAGTGAT
wide, white; columella narrow; umbilicus open with medium-sized
16S P745(R) AGAATTTTAATGGTCGAACAGACCAACCYT
funicle.
The IUB code for mixed base is used: N = G, A, T, C; V = G, A, C; D = G, A, Operculum (Fig. 6E, F): calcareous, white, with greyish-brown con-
T; B = G, T, C; H = A, T, C; W = A, T; M = A, C; R = A, G; K = G, T; S = G, C;
cretion near anterior end. External surface slightly convex; single,
Y = C, T.
narrow, peripheral flattened rib present along its outer edge corre-
sponding to outer lip of shell. Operculum thickest (1.8 mm) along

inner edge that is in contact with shell columella. Entire rib sur-
0.6–1 mm; CH 18–23 mm; CW 4–9 mm, prominent, about 2/5 face smooth, narrow, flat and devoid of granules. Internal surface
CH. Capsules medium-sized, diameter (D) 170–245 μm, arranged smooth, greenish yellow, with thin corneous layer. L 14.3 mm, W
haphazardly within collar, density 2–3 capsules mm–2; each con- 8.9 mm (for SH 23.3 mm).
taining single embryo/larva.
Animal: foot translucent white, with very densely distributed brown
Remarks: better known by its junior synonym Natica maculosa Lamarck, to dark brown maculation on dorsal surface. Tentacles black-
1822 in older literature, it was placed in its own genus Paratectonatica tipped, otherwise white.
by Azuma (1961) in view of its radula, where two subequal cusps
flank the central cusp on either side, resembling the condition Egg collar (Fig. 6G–L; n = 9): 1.5–3.5 whorls, basal margin slightly
observed in members of the genus Sinum. Kabat (2000), however, wavy or flat, entire collar flattened with low profile. Apical margin
assigned tigrina to Notocochlis, which Kabat (1991) considered as a pos- well defined, smooth. BD 10–44 mm, AD 6–20 mm, about half
sible senior synonym of Paratectonatica. The species has a wide geo- BD; TW 4–18 mm; CT 0.3–1.2 mm; CH 2.5–11 mm; CW
graphical distribution, ranging from the Mascarene Islands in the 0.5–2.3 mm, narrow, about 1/5 CH. Capsules small, D
Indian Ocean to Queensland, Australia (Poutiers, 1998). 115–140 μm, arranged haphazardly inside collar, density 4–10
capsules mm–2, each containing single embryo/larva.
Notocochlis cernica (Jousseaume, 1874) Remarks: this and the next species are very similar in the morph-
(Fig. 5) ology of their shells, opercula as well as foot pigmentation. The
Shell (based on illustration by Saito, 2000: 261, pl. 130, fig. 48 and colour and pigmentation pattern of their shells appear to be highly
unnumbered figure showing living animal; see also Kabat, 2000; variable, resulting in a long list of synonymies that have not been
Hülsken, 2008): small, SH 20 mm, high-spired; suture weak, shallow; verified. The gross morphology of the egg collar agrees with the
surface with yellow-brown periostracum with 2 or 3 narrow light- illustration provided by Kay (1979), p. 242, fig. 82B. Notocochlis
coloured spiral bands connected by more numerous but thinner axial gualtieriana has a wide, Indo-Pacific distribution ranging from East
bands in shape of chevrons (Fig. 5A). Aperture wide, columella nar- Africa to Hawaii (Poutiers, 1998).
row, with open umbilicus; moderate-sized funicle partially occluding
umbilicus. Notocochlis sp. aff. antoni (Philippi, 1851)
(Fig. 7)
Operculum: calcareous, white; external surface bears two grooves on
outer margin (based on Hülsken, 2008). Shell (Fig. 7A, B): small, SH 15.8 mm; high-spired; suture narrow,
not deep; periostracum yellow-brown with fine axial lines; shell
Animal (Fig. 5A): foot translucent white with red spotting on dorsal colour almost uniformly grey-brown without markings. Anterior
surface (see also Saito, 2000). region of shell usually white, as is area just below suture defining
Egg collar (Fig. 5B, C; n = 1): 2.5 whorls, basal margin slightly the separation between penultimate and last whorls. Surface of
wavy along vertical axis. Apical margin well defined. BD 41 mm, last whorl has about 14 narrow axial raised undulations that
AD 13.7 mm, about 1/3 BD; TW 16 mm, CT 2 mm. CH become flattened towards base of shell. Aperture wide, white;
undetermined, CW 1.5 mm. Capsules small, D 140–155 μm, columella narrow; umbilicus nearly closed by large funicle.
arranged haphazardly inside collar; density 3.4 capsules mm–2, Operculum (Fig. 7C, D): calcareous, white, external surface slightly
each containing single embryo/larva. convex, with single, narrow peripheral flattened rib along outer
Remarks: adult shell resembles N. gualtieriana (see below) but the shell of edge, matching curvature of shell outer lip. Operculum thickest
N. cernica is thinner and sutures are weak (Torigoe & Inaba, 2011). (1.1 mm) along inner edge making contact with shell columella.
The shell of Naticarius zonalis could be confused with N. cernica, but the Entire rib surface smooth, narrow, flat, devoid of granules.
operculum in Naticarius species bears 5–8 curved grooves along the Internal surface smooth, greenish yellow, with thin corneous layer
outer edge, in contrast to only 1–2 curved opercular grooves in species and slightly protruding, brown nucleus. L 9.6 mm, W 6.0 mm (for
of Notocochlis (see also Kabat, 2000). We have also distinguished SH 15.0 mm).
N. cernica from a possibly cryptic Notocochlis species, based on their COI Animal: foot translucent white, with brown to dark brown spotting
and 16 S sequences (COI absolute genetic distance = 0.083 ± 0.000; on dorsal surface. Tentacles black-tipped, otherwise white.
16 S = 0.015 ± 0.003). The latter species is likely to be the ‘N. cernica’
sequenced by Hülsken (2008) from Australia and Indonesia. Egg collar (Fig. 7E–G; n = 3): 1–2 whorls, basal margin slightly
wavy or flat. Apical margin smooth, slightly recurved. BD
Notocochlis gualtieriana (Récluz, 1844) 31–47 mm; AD 14–20 mm, about half BD; TW 12–16 mm, CT
(Fig. 6) 0.6–1.2 mm. CH 9–16 mm, CW 1.1–3.3 mm, 1/5 to 1/10 CH.
Capsules small, D 90–130 μm, arranged haphazardly inside
Shell (Fig. 6A–D): small, SH 23.3 mm; high-spired; suture narrow, collar, density 9–13 capsules mm–2; each containing single
not deep; surface with yellow-brown periostracum with fine axial embryo/larva.
360
Table 3. Absolute genetic distances (±SD, range in parentheses, and sample size) between egg-collar sequences and their respective species examined for COI and 16S genes according to the phylogenetic trees
shown in Fig. 3. A. Within-species distances. B. Within-genera distances. ND, not determined.
COI 16 S
Species Within species Between egg collars Between adults Within species Between egg collars Between adults
Paratectonatica tigrina 0.001 ± 0.002 0.002 ± 0.002 0.000 ± 0.000 0.000 ± 0.000 0.000 ± 0.000 0.000
(0.000–0.003) (0.000–0.003) (0.000–0.000) (0.000–0.000) (0.000–0.000)
(n = 6) (n = 3) (n = 3) (n = 5) (n = 3) (n = 2)
Notocochlis cernica 0.000 ND ND 0.004 ND ND
(n = 2) (n = 1) (n = 1) (n = 2) (n = 1) (n = 1)
Notocochlis gualtieriana 0.019 ± 0.011 0.022 ± 0.010 0.018 ± 0.012 0.006 ± 0.004 0.005 ± 0.004 0.006 ± 0.006
(0.000–0.035) (0.000–0.035) (0.000–0.035) (0.000–0.013) (0.000–0.009) (0.000–0.013)
(n = 13) (n = 7) (n = 6) (n = 13) (n = 8) (n = 5)
Notocochlis sp. aff. antoni 0.019 ± 0.008 ND 0.019 ± 0.008 0.005 ± 0.005 0.009 ± 0.004 0.000
(0.014–0.028) (0.014–0.028) (0.000–0.013) (0.004–0.013)
(n = 3) (n = 0) (n = 3) (n = 5) (n = 3) (n = 2)
Notocochlis sp.1 0.000 ± 0.000 ND 0.000 ND ND ND

(0.000–0.000)
(n = 3) (n = 1) (n = 2) (n = 1) (n = 1) (n = 0)
Naticarius zonalis 0.009 ± 0.005 0.010 0.007 0.000 ± 0.000 0.00 0.000
(0.003–0.014) (0.000–0.000)
(n = 4) (n = 2) (n = 2) (n = 4) (n = 2) (n = 2)
Tectonatica bougei ND ND ND 0.004 ND ND
361
(n = 0) (n = 0) (n = 0) (n = 2) (n = 1) (n = 1)
Sinum haliotoideum 0.005 ± 0.003 0.006 ± 0.003 ND 0.007 ± 0.005 0.005 ± 0.005 ND
(0.000–0.010) (0.000–0.010) (0.000–0.018) (0.000–0.013)
(n = 5) (n = 4) (n = 1) (n = 6) (n = 5) (n = 1)
Mammilla melanostoma 0.005 ± 0.002 0.007 ND 0.006 ± 0.003 0.004 N.D
(0.003–0.007) (0.004–0.009)
(n = 3) (n = 2) (n = 1) (n = 3) (n = 2) (n = 1)
Mammilla melanostomoides 0.002 ± 0.002 0.000 ND 0.000 ± 0.000 0.000 ND
(0.000–0.003) (0.000–0.000)
(n = 3) (n = 2) (n = 1) (n = 3) (n = 2) (n = 1)
Genus COI 16S
Notocochlis 0.128 ± 0.025 0.063 ± 0.013

(0.072–0.198) (0.013–0.098)
(n = 24) (n = 22)
Naticarius 0.152 ± 0.043 0.099 ± 0.047
(0.053–0.222) (0.013–0.166)
(n = 13) (n = 12)
Mammilla 0.059 ± 0.010 0.049 ± 0.008
(0.046–0.079) (0.036–0.055)
(n = 8) (n = 7)
Sinum 0.133 ± 0.012 0.073 ± 0.008
(0.091–0.164) (0.064–0.084)
(n = 12) (n = 8)
D.-R. KANG ET AL.

Figure 4. Paratectonatica tigrina (Röding, 1798), Husia, Kinmen. A, B. Shell; SH 21.5 mm. C. Living animal. D, E. Operculum; L 13.5 mm. D. External
surface. E. Internal surface. F–H. Egg collar. F. Side view; CH 20 mm. G. Top view; BD 55 mm. H. Cross-section showing egg capsules (black spheres)
among sand grains; scale bar = 1 mm.
Remarks: our Taiwan specimens strongly resemble the shells of as opposed to the rather rough surface (due to the axial undula-
Natica ‘lurida’ illustrated by Philippi (1852: pl. 3, figs 12, 13), tions present) that is consistent across all specimens observed in
although the origin of these shells is unknown. In the same mono- this study; (2) the umbilicus of the shell of N. antoni is covered by a
graph, three other shells (plate 12, figs 2–4) were labelled as fol- callus, but in all of our specimens, a narrow but open umbilicus is
lows: fig. 2: N. lurida from Havana; fig. 3: N. lurida from Sandwich present; (3) the COI sequence of N. antoni (as identified by
Islands (Hawai’i); fig. 4: N. lurida from China (although it is stated Hülsken, 2008) appears to be different from our specimens from
as ‘Lieukieusinseln’, i.e. Ryukyu Islands, in the text; Ryukyu at Taiwan (see Fig. 3A). We are currently unable to confirm the
that time could have included Taiwan). Except for plate 12: fig. 2, taxonomic relationship between our specimens and N. antoni.
all the other illustrated shells also bear a strong resemblance to
our specimens collected in Taiwan. Philippi (1836) first referred to
N. lurida in his treatment of naticids from Sicily, when providing a Naticarius zonalis (Récluz, 1850)
correction to a species he described earlier, named Natica inter- (Fig. 8)
media. In the corrigendum, he referred intermedia to marochiensis
Gmelin in Lamarck, but noted that the shells which Chemnitz fig- Shell (Fig. 8A, B): small, SH 14.4 mm; high-spired; suture deep; peri-
ured in plate 188: figs 1907 and 1908 were similar to but not maro- ostracum yellow brown with fine axial lines; a white spiral band lies
chiensis, and hence named them as luridum. Chemnitz’s figures are between two dark brown bands that are unequal in width; broad
now referable to marochiensis Gmelin, 1791. Later, Philippi (1852) darker brown spiral band about twice width of the narrow dark
synonymized luridum and marochiensis under ‘maroccana Chemnitz’, brown band that lies anteriormost. Anterior to this narrow brown
which is marochiensis Gmelin (Coan & Kabat, 2017), a tropical band are about 10 brown dashes and, posterior to the broad dark
Atlantic species (see also Table 5). To add to the confusion, lurida brown band, there are larger irregular blotches of white and brown.
is a name that has been synonymized with N. gualtieriana and to a Surface of last whorl has about 13 narrow raised axial undulations
lesser extent, N. cernica (see e.g. Torigoe & Inaba, 2011), although that become flattened towards base of shell. Aperture wide, white;
they are clearly different as shown in our study. columella narrow; umbilicus open, funicle relatively small.
We have instead referred our specimens to another closely related Operculum (Fig. 8C, D): calcareous, white, external surface generally
species Notocochlis antoni Philippi, 1851 (also often synonymized flat, bearing 8 prominent semicircular ribs that terminate at inner
with N. gualtieriana; see e.g. Kilburn, 1976; Kabat, 2000; Torigoe edge adjacent to columella, forming 8 low teeth where operculum
& Inaba, 2011) originally described from the Ryukyu Islands, des- is thickest (1.2 mm). Outermost rib at edge of operculum where it
pite their differences: (1) N. antoni has a shiny, smooth shell surface, is thinnest bears numerous (>70) fine granules arranged in single
362

Figure 5. Notocochlis cernica (Jousseaume, 1874). A. Living animal (SH 18 mm), Green I., Taiwan. B. Egg collar (BD 41 mm), Kenting. C. Cross-section of
egg collar showing egg capsules (black spheres) among sand grains; scale bar = 1 mm.
row. Ribs separated by deep grooves about as wide to half width are clearly distinguished based on COI, 16 S, 18 S and H3
of the ribs. Grey concretion present near anterior region (orienta- sequences by Hülsken (2008). The shells of N. concinnus
tion when animal is withdrawn). Internal surface smooth, greenish (Dunker, 1860) and N. manceli (Jousseaume, 1874) also resemble
yellow, with thin corneous layer. L 7.9 mm, W 4.7 mm (for SH N. zonalis to different degrees. In the case of N. concinnus, the
14.4 mm). COI sequences from Hülsken (2008) for this species were suffi-
ciently different from those we obtained from adults and egg
Animal: foot translucent white with densely distributed pink or light masses in Taiwan (Fig. 3) to distinguish the two species, although
red dashes on dorsal surface. their egg masses are strikingly similar. As for N. manceli, there were
Egg collar (Fig. 8E–G; n = 2): 2–3.5 whorls, basal margin wavy along no sequences available for this species, but here we regard it suffi-
vertical axis; the waves traced by successive whorls are out of phase ciently different conchologically from N. zonalis.
with each other. BD 25–36 mm, AD 6–8 mm, about 1/5 BD; TW
12–15 mm; CT 0.9–1.5 mm; CH 8–11.6 mm; CW 1–2 mm, nar-
row, about 1/11 CH. Capsules large, D 470–550 μm, arranged in Tectonatica bougei Sowerby, 1908
single layer inside collar; density 0.5–1.1 capsules mm–2, each con- (Fig. 9)
taining single embryo/larva.
Shell: small, SH 4.5 mm; medium-spired; suture shallow; surface
Remarks: this species was synonymized with Naticarius sertatus with thin periostracum. Shell generally white with dark- or
(Menke, 1843) by Torigoe & Inaba (2011), but the two species reddish-brown irregular markings. Protoconch region also reddish
363
D.-R. KANG ET AL.
Figure 6. Notocochlis gualtieriana (Récluz, 1844). A–D. Shell. A. Dongsha; SH 15.6 mm. B. Dongsha, ventral view with operculum. C. Dongwei, dorsal
view; SH 16.5 mm. D. Dongwei, ventral view with operculum. E, F. Operculum, Tongliang; L 10.0 mm. E. External surface. F. Internal surface. G–L.
Egg collars. G, I, K. Dongsha, top, side view and cross section; BD 32 mm; CH 3 mm. H, J. Jingan, top and side view; BD 31 mm; CH 5 mm; L.
Dongwei, cross-section. Scale bars K, L = 1 mm.
brown. Aperture wide, white, with moderately broad columella Operculum: calcareous, white, external surface generally flat with
and closed umbilicus with large funicle (based on Hülsken, 2008: single, narrow peripheral flattened rib along outer edge, corre-
fig. A.27). sponding to outer lip of shell (based on Hülsken, 2008: fig. A.27).
364
Figure 7. Notocochlis sp. aff. antoni, Husia. A, B. Shell; SH 15.8 mm. C, D. Operculum, external and internal surfaces; L 10.0 mm. E–G. Egg collar, top, Downloaded from https://academic.oup.com/mollus/article-abstract/84/4/354/5102386 by guest on 01 November 2018
side view and cross-section; BD 42 mm; CH 10 mm. Scale bar G = 1 mm.
Animal: foot translucent white, with brown spots dashes on dorsal holotype of T. bougei examined by Hülsken (2008), bougei has a
surface of the frontal lobe (after Hülsken, 2008). thick shell and closed umbilicus, as opposed to the thin shell and
partially open umbilicus of venustula. Saito (2000, 2017) also
Egg collar (Fig. 9A, B; n = 1): 3 whorls, basal margin smooth along showed that the umbilicus of T. bougei (as Natica bougei) was almost
vertical axis. Apical margin distinct, slightly recurved. BD 8 mm, completely covered by callus. However, Kabat (2000) identified a
AD 2 mm, about ¼ BD; CW 3 mm, CT 0.5 mm; CH 3 mm, CW specimen from the Cook Islands showing an open umbilicus as
not measured. Capsules large, diameter 93–105 μm, arranged in a T. bougei. The taxonomic significance of the state of the umbilicus
single layer inside collar, density 0.02 capsule mm–2, each contain- appears to be played down by Torigoe & Inaba (2011). This may
ing single embryo/larva. have led them to synonymize the two species, as well as confound-
Remarks: this species was considered a synonym of Tectonatica venus- ing Notocochlis gualtieriana, whose shell has an open umbilicus, with
tula (Philippi, 1851) by Torigoe & Inaba (2011). Based on the N. sp. aff. antoni, which has a closed umbilicus. The latter two
365
D.-R. KANG ET AL.

Figure 8. Naticarius zonalis (Récluz, 1850), Tongliang. A, B. Shell; SH 14.4 mm. C, D. Operculum, external and internal surfaces; L 7.9 mm. E–G. Egg
collar, top, side view and cross-section; BD 28 mm; CH 11 mm. Scale bar G 1 mm.
species are clearly different based on our current study, reiterating Animal: Living specimen not observed. Foot large, white in pre-
the importance of the umbilicus as a taxonomic character at the served specimens (L 70 mm, W 20 mm, for SL 34 mm)
species level. Its significance at the genus level may be less certain,
given that species assigned to Tectonatica (all of which are charac- Egg collar (Fig. 10C–E; n = 5): 1–2.5 whorls, basal margin wavy
terized by a closed umbilicus) are generally grouped with other along horizontal axis, and in some parts the margin is folded
genera (e.g. Natica, Notocochlis) having an open or narrow umbilicus slightly inwards so that it is not visible from above. Apical margin
in both COI and 16 S trees (Fig. 3). smooth, upright. BD 62–87 mm, AD 33–47 mm, about half BD;
CW 26–43 mm, CT 0.7–0.9 mm; CH 24–39 mm, CW 4.6–10.6 mm,
about 1/4 to 1/5 CH. Capsules medium-sized, diameter
Sinum haliotoideum (Linnaeus, 1758) (D)170–210 μm, arranged haphazardly within collar, density 6–10
(Fig. 10) capsules mm–2; each containing single embryo/larva.
Shell (Fig. 10A, B): medium-sized, SL = 20–34 mm; low-spired and Remarks: the species names planatum Récluz in Chenu, 1843 and
flattened dorsoventrally. Suture narrow, not deep. Shell surface planulatum Récluz, 1843 have been used for this species but
with thin, yellow-brown periostracum with fine axial shell growth according to Kabat (1990), the name haliotoideum Linnaeus, 1758
lines visible beneath it; spiral lines on shell are not visible. Shell has priority, and the lectotype chosen by Kabat is closest to the
colour almost uniformly white. Aperture wide, white, with narrow species examined in this study. Members of the subgenus Sinum
columella; umbilicus small, shallow. have shells with prominent sutures and ribs in contrast to those
of Ectosinum (Iredale, 1931), which are lacking in spiral sculpture.
Operculum: small, triangular, corneous, thin and translucent; nor- Based on the sequence analyses of mt COI and 16 S rRNA
mally buried inside foot and not visible on living animal (Saito, (Fig. 3), Hülsken’s (2008) ‘haliotoideum’ may be another species of
2000). Sinum.
366
Figure 9. Tectonatica bougei (Sowerby, 1908), Kenting, egg collar. A. Field photograph; BD 8 mm. B. Cross-section. Scale bar B 1 mm.
Mammilla melanostoma (Gmelin, 1791) closed by swollen columella. Funicle absent. Both columella and
(Fig. 11) umbilicus stained dark brown.
Shell (Fig. 11A, B): large, SH 29 mm, high spired; suture shallow; Operculum (Fig. 11C): corneous, rusty brown. L 22 mm, W 10 mm
external surface mostly white with faint brown spiral bands of dif- (for SH 29.0 mm); size of operculum similar to shell aperture.
ferent widths and fine axial lines. Spiral sculpture absent.
Aperture wide, white, with narrow columella; umbilicus nearly Animal: foot large, white, no surface pigmentation.
367
D.-R. KANG ET AL.

Figure 10. Sinum haliotoideum (Linnaeus, 1758). A, B. Shell, dorsal and ventral views, Penghu (NMNS 003750-00030); SL 22.0 mm. C–E. Egg collar,
Tongliang, top, side view and cross-section; BD 74 mm; CH 25 mm. Scale bar E = 1 mm.
Figure 11. Mammilla melanostoma (Gmelin, 1791). A, B. Shell, Liuqiu (NMNS 003232-00039); SH 29 mm. C. Operculum, external surface; L 22 mm.
D–F. Egg collar, Tongliang, Penghu, top, side view and cross-section; BD 89 mm; CH 18 mm. Scale bar F = 1 mm.
368

Figure 12. Mammilla melanostomoides (Quoy & Gaimard, 1832), egg collar. A, B. Jingan, Penghu, top and side view; BD 48 mm; CH 14 mm. C. Daxi.
Scale bar C = 1 mm.
Figure 13. Egg collar of an undetermined naticid (possibly Tanea sp. or Euspira sp.), Kenting. A, B. Top and side view; BD 48 mm; CH 18 mm. C. Cross-
section. Scale bar C = 1 mm.
Egg collar (Fig. 11D–F; n = 2): 4 whorls, basal margin prominently Mammilla melanostomoides (Quoy & Gaimard, 1832)
sinuous in vertical plane, the waves traced by successive whorls (Fig. 12)
are in phase with each other. BD 89 mm, AD 25 mm; CW
33 mm, CT 1.4 mm. CH 18 mm, crown absent. Capsules Shell: large, SH 30 mm, high spired; suture shallow; external sur-
medium-sized, D 205–230 μm, arranged haphazardly within col- face mostly white with irregular brown spots. Spiral sculpture
lar, density 3–5 capsules mm–2; each containing single embryo/ absent. Aperture wide, white, with very narrow, concave colu-
larva. mella, umbilicus nearly closed by swollen columella. Funicle
absent. Both columella and umbilicus stained dark brown.
Remarks: This is a widespread and well known species in the Indo-
West Pacific (Poutiers, 1998). The living animal and egg collar are Operculum: corneous, rusty brown, substantially smaller than shell
illustrated in Chiu, Huang & Su (2011). aperture.
369
D.-R. KANG ET AL.
Animal: unknown. were supported as monophyletic. The following groups were dis-
cernible, here listed together with traditional taxonomic characters
Egg collar (Fig. 12A–C; n = 2): 2 and 4.5 whorls; basal margin wavy in of the shell and operculum:
vertical plane. BD 48 and 58 mm, AD 10 and 23 mm, CW 21 mm,
CT 1 and 2 mm; BD about three times AD; TW 14–22 mm, CW Group I: collars with tall crown and single whorl; medium-sized
1.9–2.6 mm. Crown is 1/8 height of upright egg mass. Capsules shell with distinct spire and open umbilicus; calcareous operculum
medium-sized, D 220–260 μm, arranged sparsely within collar, dens- with two ribs (e.g. Paractectonica tigrina; Fig. 4).
ity 3–6 capsules mm–2. Each capsule contains single embryo/larva. Group II: collars with moderate AD; small shell with distinct spire and
Remarks: this and the preceding species have similar shells and egg col- either open or nearly closed umbilicus; calcareous operculum with sin-
lars, but differ sufficiently in detail to be regarded as distinct species. gle rib (e.g. Notocochlis gualtieriana and N. sp. aff. antoni; Figs 6, 7).
Undetermined sp. Group III: collars with a small apical diameter, small shell with
(Fig. 13) distinct spire and open umbilicus, calcareous operculum with two
ribs (e.g. other Notocochlis spp.; Fig. 5).

Shell, operculum and animal: unknown. Group IV: small, pyramidal collars with very small apical diam-
Egg collar (Fig. 13A–C; n = 1): about 4 whorls; basal margin wavy eter; small shell with distinct spire and open umbilicus; calcareous
in vertical and horizontal planes. Apical margin smooth, slightly operculum with multiple ribs (e.g. Naticarius spp.; Fig. 8).
recurved. BD 50 mm, AD 18 mm, about 1/3 BD; TW 16 mm, Group V: collars with nearly perpendicular sides; shell with low,
CT 4 mm; CW 2 mm, about 1/8 CH. Capsules large, D indistinct spire and small, shallow umbilicus; small, corneous oper-
300–400 μm, arranged sparsely within collar, density 2–4 capsules culum (e.g. Sinum spp.; Fig. 10).
mm–2, each containing single embryo/larva.
Group VI: many-whorled collars with basal margin wavy in verti-
Remarks: the COI sequence obtained from this egg collar appeared cal plane; shell with distinct spire and almost closed umbilicus;
near Tanea spp. and Euspira spp. (Fig. 3), but without bootstrap sup- large, corneous operculum (e.g. Mammilla spp.; Figs 11, 12).
port. Unfortunately, none of the egg collars of Tanea species has been Group VII: many-whorled, flattened collars with extremely wavy
characterized to date (Table 5) and none of the known egg collars of basal margin; shell and opercular characters not determined
Euspira have a similar morphology to our specimen described above. (Fig. 13; possibly Tanea or Euspira spp.).
Egg-collar morphology and molecular phylogeny DISCUSSION

We examined the morphology of the egg collars in relation to the
phylogenetic trees (Fig. 3) obtained from sequence data and classi-
Molecular analyses
fied the 11 different egg collars into seven groups (I–VII; Table 4). Molecular tools applied to resolve phylogenies of various gastro-
While these bore some relationship to the molecular trees, not all pod groups have achieved varying degrees of success (see reviews
Table 4. Proposed classification of Taiwanese egg collars. A total of seven groups (I–VII) were recognized during this study based on their morphological
attributes.
Operculum/number of
Group Egg collar Shell Umbilicus ribs on surface Typical species
I Medium-sized, with tall crown; single Medium-sized, distinct Open C/2 Paratectonatica tigrina
whorl; basal diameter 2–3 times spire
apical diameter
II Small to medium-sized, low crown, Small-sized, distinct Open or nearly closed C/1 Notocochlis gualtieriana,
basal diameter twice apical spire N. sp. aff. antoni
diameter; 1.5–3.5 whorls;
III Small, with low crown; basal diameter Small-sized, distinct Open C/2 Notocochlis spp. and
4–5 times apical diameter; 1.5–2 spire Tectonatica bougei
whorls
IV Small pyramidal collars with very small Small-sized, distinct Open C/8 Naticarius zonalis
apical diameter; crown indistinct; spire
multiple whorls
V Generally large; high crown, basal Medium-sized, low Shallow NC, small Sinum haliotoideum
diameter <1.5 times apical spire
diameter; 1–2.5 whorls
VI Generally large; low or absent crown; Medium-sized, distinct Nearly closed NC, large Mammilla spp.
multiple whorled collars with wavy spire
margin along vertical axis; basal
diameter >2 times apical diameter
VII Generally large, low crown; multiple Medium- to large ? ? Unknown
whorled flattened collars with sized, distinct spire
extremely wavy basal margin; basal
diameter >3 times apical diameter
Shell and opercular characters corresponding to the groups are also provided.
Abbreviations: C, calcareous; NC, noncalcareous (i.e. corneous).
370
by Colgan et al., 2007; Aktipis et al., 2008; Ponder et al., 2008; Members of this genus have calcareous opercula that bear only
Zapata et al., 2014). Attempts to resolve genus-level relationships one or two peripheral ribs on their external surface (Kabat, 2000;
amongst the caenogastropods have also been somewhat uneven. Figs 6E, F, 7C, D). In Group II, members of which have one per-
Hülsken et al. (2008) using sequences based on H3, COI, 16 S and ipheral rib on their operculum and lay collars with a moderate
18 S genes, provided a first molecular phylogenetic tree for the apical diameter, N. gualtieriana (type species of Notocochlis) has very
Naticidae. The results, although based on a limited number of similar egg collar features to those of N. sp. aff. antoni. These two
taxa, recalled uncertainties raised by earlier researchers (e.g. species were synonymized by Kay (1979) and Torigoe & Inaba
Powell, 1933; Marincovich, 1977; Golikov & Sirenko, 1988; (2011), but are considered distinct species here, based on their
Majima, 1989; Bandel, 1999) who observed inconsistencies in genetic distances. Members of Group III, which form a well sup-
morphological characters such as the operculum and shell umbil- ported clade distinct from Group II (not monophyletic in either
icus used to delineate naticid subfamilies and genera. tree), have an operculum with two peripheral ribs and lay collars
In our study of Taiwanese Naticidae, we combined our new with a relatively small apical diameter. This suggests that further
COI and 16 S sequences with those from Hülsken et al. (2008). analysis could support the assignment of members of Group III to
The resulting dataset is still lacking some critical taxa (e.g. a different genus.

Globisinum, Amauropsis and deep-sea species) and the poorly sup- Egg collars with multiple whorls were also observed among mem-
ported basal clades in the phylogenetic trees reflect the very lim- bers of the subfamily Polinicinae, which is characterized by a corn-
ited number of taxa examined in this study. While we did not set eous operculum (Cernohorsky, 1971; Kilburn, 1976; Kabat, 2000;
out to examine the detailed relationships between naticid genera, Hülsken et al., 2012). Both Mammilla melanostoma and M. melanosto-
our trees do not contradict the topologies of those obtained by of moides lay distinctive collars having up to 4.5 whorls that are thrown
the Hülsken et al. (2008), which seem to suggest that the currently into sinuous folds in both the vertical and horizontal planes (Group
recognized Naticinae (i.e. taxa bearing calcareous opercula) are VI; Table 4, Figs 11D, E, 12A, B). These folds probably help to sta-
not a monophyletic clade. On the other hand, our results were bilize the collars on the substratum in moving water, while allowing
unable to corroborate the subsequent findings of Hülsken et al. oxygenated water to pass under and over the multilayered collar
(2012), which showed that species of Euspira, Mammilla and walls. This collar morphology is also shared by the conchologically
Polinices formed a clade, i.e. that the Polinicinae are a monophy- similar Indo-Pacific Polinices mammilla (see Gohar & Eisawy, 1967)
letic group. In both the COI and 16 S rRNA trees obtained here, and the Caribbean P. lacteus (see Bandel, 1976). However, most
the relationships between Euspira, Mammilla and Polinices were other polinicine species whose collars have been described produce
equivocal. Clearly there is a need for future work involving more single-whorled collars (Table 5), which can also be sinuous along
taxa, to clarify and stabilize the phylogenetic relationships between the basal edge. As noted above, our molecular results provided no
naticid genera. support for the traditional Polinicinae.
We also obtained an undetermined egg collar with multiple
whorls that was distinctly flattened, with sinuous folds, which we
Egg collars: identification, classification and review assigned as Group VII (Table 4; Figs 3, 13). Despite successfully
obtaining both COI and 16 S sequences from the egg collar, its
Egg collars were generally distinct among the nine species exam- identity remains unclear. In both trees the sequence grouped near
ined, differing in their overall shape, number of whorls, capsule Tanea and Euspira, but without statistical support.
size and arrangement of capsules within the collar, as previously A number of unrelated species lay egg collars with a distinct
detailed for other species (Thorson, 1940; Amio, 1955; Giglioli, crown. The naticine Paratectonatica tigrina lays a distinctive, single-
1955; Bandel, 1976; Baxter, 1983; Table 5). Our findings support whorled egg collar that has a tall crown (Group I; Fig. 4F, G).
the studies by Hülsken (2008) and Hülsken et al. (2008, 2011), This monotypic genus is otherwise distinguished by their unusual
showing that molecular analysis of egg masses allows the confirm- rachidian radula teeth, which resemble those observed in Sinum
ation of the identity of naticid species without having access to species. The egg collar of Natica marochiensis similarly has a tall
adult specimens, if suitable reference sequences from reliably iden- crown (Knudsen, 1950), as do those of the polinicines Glossaulax
tified adults are available. didyma, G. vesicalis and G. reiniana (Amio, 1955; Kim et al., 2007).
Among the six naticine species bearing calcareous opercula Most other naticid species appear to produce collars with a short
examined in this study, the small, tightly coiled egg collar of crown (e.g. Notocochlis, Naticarius and Sinum; Figs 6–8, 10), or a
Naticarius zonalis with a diminutive apical opening (Group IV; crown may be absent altogether (M. melanostoma; Fig. 11D).
Fig. 8E, F) appears to be similar to those of two congeners Unusually, the apical margin of the collars of N. sp. aff. antoni and
described by Hülsken et al. (2008, 2011; N. concinnus and N. onca; M. melanostomoides (Fig. 12A, B) is slightly but distinctly flared out-
Table 5). The egg collars of these three species all possess multiple wards (recurved).
whorls that are wound at an acute angle, such that the low-profile To date, the egg collars of only three species of the small sub-
collars have a small apical diameter relative to the base. In the family Sininae have been described (Table 5). Its members are
COI and 16 S trees (Fig. 3A, B), zonalis and onca belong to a clade characterized by reduced opercula and thin shells that are dorso-
of at least four Naticarius species. In contrast, two other larger- ventrally flattened, with large apertures. Their egg collars have up
shelled congeners, canrenus (see Bandel, 1976) and cruentatus (as heb- to 2.5 whorls (Group V; Table 4, Fig. 10C, D). Sinum japonicum is
raeus in Hülsken et al., 2008), lay collars with less than two whorls the only one of the three sinines with large egg capsules about
and the apical diameters are wider. Members of the genus 1 mm in diameter, each containing 10–12 larvae (Knudsen, 1994).
Naticarius possess calcareous opercula whose external surface bears The others have only a single larva in each capsule, which is
several sulci and spiral ridges or ribs (Fig. 8C, D; Cernohorsky, 170–290 μm in diameter. The egg collar of S. haliotoideum is dis-
1971; Kilburn, 1976; Kabat, 2000; Zhang, 2016). However, tinct in having its apical and basal diameters of comparable size
monophyly of the genus, as traditionally defined by shell and oper- (Fig. 10C, D), the walls having a nearly vertical orientation. This
cular characters, was not supported by our molecular analyses. is similar in shape to the egg collar identified as the same species
Three species of the genus Notocochlis also lay egg collars with by Hülsken (2008). However, the shell shown by Hülsken (2008:
multiple whorls (Groups II and III; Table 4, Figs 5B, 6G–J, 7E, fig A80A) is beige with stronger spiral striae, differing from the
F). However, the apical diameter of their collars is considerably white shell with very weak striae that is the lectotype of S. haliotoi-
wider than in Naticarius. The collars of Notocochlis cernica, N. gualtieri- deum. Both COI and 16 S genetic distances between our Sinum sp.
ana and N. sp. aff. antoni are consistent with those of N. isabelleana and Hülsken’s S. haliotoideum suggest that they are not conspecific,
from South America (Pastorino, Averbuj & Penchaszadeh, 2009). but share a similar egg collar morphology.
371
Table 5. Egg collar characteristics in the Naticidae, based on this study and available literature. Adult sizes, classification of subfamilies and genera, and geographical distributions of species follow Torigoe &
Inaba (2011), unless otherwise stated.
Basal Apical diameter,

Species (subfamily) Adult shell No. of Presence of crown; diameter or length × width Basal margin folds / Capsule size Capsule arrangement; No. of embryos in
(sample size) height (mm) whorls relative height (mm) (mm) direction of fold (μm) density (mm−2) capsule Distribution; reference
Paratectonatica tigrina 30 1–1.5 Yes; high 42–58 15–24 Sinuous / horizontal 170–245; Irregular; 2–3; 1 Indo-West Pacific; this study;
(Naticinae) (n = 9)
1 Yes; high 37–76 14–33 Sinuous / horizontal 220–300 Irregular; 3–5 Japan; Amio (1955; as Natica
maculosa);
1.5 Yes; high 56–79 ? Smooth or folded 251–284 Double layer 1 India; Natarajan (1957; as
Natica)
Notocochlis cernica 20 2.5 Yes; low 41 14 Slightly sinuous / vertical 140–155 3.4 1 W Pacific; this study
(Naticinae) (n = 1)
Notocochlis dillwyni 12 3–5 Yes; low 20–40 7–9 Flat or slightly sinuous ? ? ? Mediterranean to W. Africa
(Payraudeau, 1826) and Caribbean; Hülsken
(Naticinae) (n = 6) et al. (2008)
Notocochlis gualtieriana 20 1.5–3.5 Yes; low 10–44 6–20 Flat or slightly sinuous / 115–140 4–10 1 Indo-West Pacific; this study
(Naticinae) (n = 9) vertical (see also Kay, 1979:
fig. 82B)
15–27 1.5–2 Yes; low 35–50 ? Slightly sinuous 133 Single layer; 3–7 1 Pamban, India; Natarajan
D.-R. KANG ET AL.

capsules in spherical (1957; as ‘N.
egg space 183–367 μm marochiensis’)
in diameter
372
Notocochlis isabelleana 15 1–1.5 Yes; low 35–67 14–28 Sinuous / horizontal 154–235 14–16 1 Brazil to Argentina; Pastorino
(d’Orbigny, 1840) et al. (2009)
(Naticinae) (n = 23)
Notocochlis sp. aff. 20 1–2 Yes; low 31–47 14–20 Flat or slightly sinuous 90–130 9–13 1 Taiwan; this study
antoni (Naticinae)
(n = 14)
Naticarius canrenus 67 1–1.5 Yes; low 80–120 40–45 Slightly sinuous / 350–500 Single layer or double 1 Caribbean; Bandel (1976)
(L., 1758) (Naticinae) horizontal layer; 1
Naticarius concinnus 20 3 ?yes; low 25 2 Flat or slightly sinuous ? ? ? Japan–Australia; Hülsken
(Dunker, 1860) et al. (2011)
(Naticinae) (n = 1)
Naticarius hebraeus 30–40 1.25 Yes; low 75 40 Flat ? ? ? Mediterranean; Hülsken et al.
Martyn, 1784 (2008)
(Naticinae) (n = 1)
Naticarius onca (Röding, 25–30 3 Yes; low 30 0–5 Flat or slightly sinuous ? ? ? Indo-West Pacific; Hülsken
1798) (Naticinae) et al. (2011)
(n = 8)
Naticarius zonalis 20 2–3.5 Yes; low 25–36 6–8 Sinuous / vertical 470–550 0.5–1.1 1 Indo-West Pacific; this study (
(Naticinae) (n = 7) = sertatus Menke, 1843
according to Torigoe &
Inaba, 2011)
Natica livida Pfeiffer, 7–15 1.5 Yes; low 15–17 6–7 Slightly sinuous / 150 Irregular, generally single 2–5 Caribbean; Bandel (1976)
1840 (Naticinae) horizontal layer; 3
40 <1 Yes; low 70 28 Sinuous / horizontal 1,000 Single layer; 1 1
Natica vitellus (L., 1758) Indo-West Pacific; Thorson
(Naticinae) (1940; as Natica rufa)
Natica marochiensis 15–27 1 Yes; high 38 21 Sinuous / horizontal; egg 500 Single layer; 20–25 20–25 W Africa; Knudsen (1950)
(Gmelin, 1791) mass made of faecal embryos in each
(Naticinae) (n = 1) pellets capsule
Natica (Tectonatica)? 14 1.25 Yes; low 37–48 15 Flat ? ? ? Mediterranean Sea; Hülsken
rizzae (Philippi, 1844) et al. (2008; as
(Naticinae) (n = 4) Tectonatica)
Natica rubromaculata 20 ? ? (fragment only ? ? Slightly sinuous 750 Single layer; 1 1 W Africa; Knudsen (1950)
Smith, 1871 observed)
(Naticinae)
Natica sagraiana 10 1 Yes; low 24–40 20 Flat ?400 ? 1 Mediterranean Sea; Hülsken
(d’Orbigny, 1842) et al. (2008; as
(Naticinae) (n = 8) Tectonatica)
Natica (Tectonatica) 10 3 Yes; low 8 2 Flat 93–105 Single layer; 0.02 1 Tropical Pacific; this study
venustula Philippi,
1851 (Naticinae)
(n = 1)

Natica (Tectonatica) 20–41 leaf-like NA NA 30 (L) × 10 (W) Stalked base ? ? ? South Africa; Kilburn &
tecta Anton, 1839) attached to hard Rippey (1982)
(Naticinae) substratum
Cryptonatica adamsiana 30 1.5 ? 32–56 18–31 ? 290–340 Single layer; 1 12–13 Japan; Amio (1955; as
(Dunker, 1860) Natica)
(Naticinae)
373
Cryptonatica clausa 60 1 No 40 20 Flat 2,000–2,250 Single layer; 1 1 Arctic Ocean; Thorson (1935)
(Broderip & Sowerby,
1829) (Naticinae)
Cryptonatica janthostoma 50 1.2 Yes; low 88 37 Flat 770 Single layer; 1 18–26 Kamchatka to Korea;
(Deshayes, 1839) Kulikova, Kolbin, and
(Naticinae) Kolotukhina (2007)
Cryptonatica 40 1 Yes; low 73–105 30–46 ? 290–340 Single layer; 1 14–17 Japan; Amio (1955; as
janthostomoides Natica)
(Kuroda & Habe, 1949)
(Naticinae)
Sinum japonicum 15 1.2 Yes; low 60 25 Slightly sinuous 1,000 Single layer; 1 10–12 China and Japan; Knudsen
(Lischke, 1872) (1994)
(Sininae) (n = 1)
Sinum haliotoideum 20 1–2.5 Yes; low 62–87 33–47 Sinuous / vertical and 170–210 Irregular; 6–10 1 Indo-West Pacific; this study
(Sininae)(n = 4) horizontal
Eunaticina papilla 20 1.5 Yes, high 38–86 33–38 Flat or slightly sinuous 200–290 Irregular; 4–6 1 Africa to E Asia; Amio (1955;
(Gmelin, 1791) as Sinum)
(Sininae)
Conuber conicum 40 1 No 90 ? None 300 Diffuse in gelatinous 1–3 S Australia; Murray (1962)
(Lamarck, 1822) mass; no sand grains
(Polinicinae)
20–30 <1 Yes? ?40 ? None 350 1 S Australia; Murray (1966)
Continued
Table 5. Continued
Basal Apical diameter,

Species (subfamily) Adult shell No. of Presence of crown; diameter or length × width Basal margin folds / Capsule size Capsule arrangement; No. of embryos in
(sample size) height (mm) whorls relative height (mm) (mm) direction of fold (μm) density (mm−2) capsule Distribution; reference
Conuber incei (Philippi, Diffuse in gelatinous

1853) (Polinicinae) mass; no sand grains
Conuber sordidum 40–50 <1 No 70–100 10–20 None 625 Within 3 mm of outer 1–3 S Australia: Murray (1962);
(Swainson, 1821) surface in gelatinous Booth (1995)
(Polinicinae) mass; no sand grains
Mammilla melanostoma 40 4 No 89 25 Prominently sinuous / 205–230 Irregular; 3–5 1 Indo-West Pacific; this study
(Polinicinae) (n = 1) vertical
? 1 Yes; low 69 33 Flat 300 Double layer; 3–5 1 Indo-West Pacific: Gohar &
Eisawy (1967)
23 1 Yes; low 60 12 Flat 300? Double layer; 3–5 1 Hong Kong: Knudsen (1992;
as Polinices)
Mammilla 40 2, 4.5 Yes; medium 48, 58 10, 23 Sinuous 220–260 Irregular; 3–6 1 Indo-West Pacific; this study
melanostomoides
(Polinicinae) (n = 2)
Polinices lacteus 25 2.5–5.5 Yes; low 23–43 7–10 Sinuous / horizontal 200 Irregular; 10 1 Caribbean and W Africa;
(Guilding, 1834) Bandel (1976)
D.-R. KANG ET AL.

(Polinicinae)
Polinices hepaticus 50 0.75–1.5 Yes; low 40–45 20–23 Slightly sinuous / 300 Irregular; 5 1 Caribbean; Bandel (1976)
(Röding, 1798) horizontal
374
(Polinicinae)
Polinices mammilla L., 30–40 2–5 Yes; low 36–80 12–32 Sinuous / horizontal 135 Irregular; 25 1 Indo-West Pacific; Gohar &
1758 (Polinicinae) Eisawy (1967)
to 58 up to 5 Yes; low 30 17 Sinuous / horizontal? ? ? ? Kilburn & Rippey (1982; as
tumidus Swainson, 1840)
Neverita josephinia 40 1 Yes; low 51–59 38 Sinuous / horizontal 1,000 Single layer; 1 1 Mediterranean Sea; Giglioli
(Risso, 1826) (1955); Hülsken et al.
(Polinicinae) (n = 2) (2008)
Amauropsis islandica 20–30 ? No ? ? Sinuous / horizontal 1,500–1,750 Single layer; 1 1 N Atlantic; Thorson (1935);
(Gmelin, 1791) Giglioli (1955)
(Polinicinae)
Glossaulax reiniana 40 <1 Yes; high 52–64 21–27 Sinuous / horizontal 350–410 Irregular; 8–10 1 East China Sea to Japan;
(Dunker, 1877) Amio (1955; as Neverita)
(Polinicinae)
Glossaulax vesicalis 80 1 Yes; high 29–64 16–35 Sinuous / horizontal 1,300–1,450 Single layer; 1 1 Amio (1955; as Neverita)
(Philippi, 1849)
(Polinicinae)
Glossaulax didyma 40–90 1 Yes; high 80–95 60 Sinuous / horizontal 1,000 Single layer; 1 1 Indo-West Pacific; Thorson
(Röding, 1798) (1950; as Natica ampla)
(Polinicinae)
1 Yes; high 64–130 33–65 Sinuous / horizontal 520–580 Double layer; 2 1–3 Japan; Amio (1955; as
Neverita)
1 Yes; low 63–160 40–50 Sinuous / horizontal 240–260 ? 1
Monora I., Pakistan; Tirmizi &
Zehra (1983; as Neverita)
1.2 Yes; low 90 60 Flat 240–300 ? 1 China; Liu & Sun (2008; as
Neverita)
1 Yes; high 66 34 Flat 300 Double layer; 2 1 Hong Kong; Knudsen (1992;
as Polinices)
1 Yes; high ? ? Sinous / horizontal 530–570 Irregular; 1 Korea; Kim et al. (2007)
Laguncula pulchella 25–45 <1 Yes; low 69–96 27–43 Sinuous / horizontal ? ? 3–5 (about 100 nurse China, Korea, Japan;
Benson, 1842 cells present) Tomiyama (2013; as
(Polinicinae) Euspira fortunei)
? 1 Yes; low 50–120 40 Sinuous / horizontal 1,200–3,400 Single layer 2–8 (36–93 nurse Miyagi Pref., Japan; Sakai &
cells) Suto (2005; as Neverita
didyma)
Euspira catena (da 30 1 Yes; low 73 47 Flat 1,425×1,925 Single layer; 1 1–19 (up to 180 North Sea and
Costa, 1778) spheres nurse cells Mediterranean; Ankel
(Polinicinae) present) (1930); Hertling (1932; as
Lunatia); Giglioli (1949);
Fretter & Graham (1981)

Euspira fusca (Blainville, 30 <1 No 120 50 Flat 492×468 Irregular; 1 1 Mediterranean Sea; Ramón
1825) (Polinicinae) (1994; as Euspira)
Euspira montagui 10 1.2 No 40 10–13 Flat 230 ? ‘numerous’ Fretter & Graham (1981)
(Forbes, 1838) (?nurse eggs)
(Polinicinae)
Euspira heros (Say, 50–100 1.2 Yes; low 50–60 to 40 Flat 630×850 Single layer; 1 4–84 W Atlantic; Giglioli (1955)
375
1822) (Polinicinae) spheres

Euspira lewisi (Gould, to 125 1 Yes; low 120 50 Flat 200×250 Irregular; 4–5 1 E Pacific; Giglioli (1955)
Euspira nitida (Donovan, 10–20 1–3.5 No 17–40 8–18 Flat 208–254 Irregular; 1 North Sea; Ziegelmeier (1961;
1804) (Polinicinae) as Lunatia); Hülsken et al.
(2008; as Euspira)
Euspira pila (Pilsbry, 20–30 ? ? 79 38 ? 770 Single layer; 1 1 Yakovlev & Kolotukhina
1911) (1996)
Euspira triseriata (Say, 10 1.2 No 40 15 Flat 800×1,150 Single layer; 1 1–3 W Atlantic; Giglioli (1955)
Globisinum drewi 30 1.5 ? 100 45 Flat 6,500 Single layer; <1 ? New Zealand; Dell (1956)
(Murdoch, 1899)
(Globisininae)
Falsilunatia carcellesi 25 ? ? ? ? Flat 8,780–14,140 Single row; single layer, 1 Argentina, 200–2000 m;
(Dell, 1990) (? <1 Penchaszadeh et al. (2016,
Globisininae) as Bulbus carcellesi)
D.-R. KANG ET AL.
The largest egg collar observed in this study measured up to by Majima, 1989), differences in shell shape (Sun et al., 2012) and
90 mm in basal diameter, belonging to Mammilla melanostoma molecular diversity (Zhao et al., 2017).
(Group VI; Fig. 11D–F), which has an adult shell height of about As discussed above, we found some correspondence between
40 mm. The egg collar of the same species in Hong Kong molecular phylogeny and egg-collar morphology, but generally
reported by Knudsen (1992: pl. 4D) measured 60 mm in diameter. poor support for the monophyly of traditional genera. Genus-level
However, those of Glossaulax didyma (Indo-Pacific), Euspira fusca taxonomy is currently strongly dependent upon umbilical and
(Mediterranean) and E. lewisi (East Pacific) can be in excess of opercular characteristics (Kabat, 2000), but our results hint that
120 mm (Amio, 1955; Giglioli, 1955; Ramón, 1994). The egg col- egg-collar morphology may also prove to be a useful character
lar of the deep-sea Globisinum drewi was recorded by Dell (1956) to indicating phylogenetic affinity. As the identities of more egg col-
be about 100 mm in diameter. lars are revealed, the combination of morphological and molecu-
All egg collars examined in this study incorporated a single lar characters should result in a more natural and stable
embryo in each capsule, but there are a few species occurring else- classification of the Naticidae.
where whose capsules contain more than one embryo, including
Cryptonatica adamsiana and S. japonicum that have more than 10 eggs

in each capsule (Amio, 1955; Knudsen, 1994). Knudsen (1950) SUPPLEMENTARY MATERIAL
observed that each capsule in the egg collar of Natica marochiensis
from West Africa contained between 20 and 25 eggs, while those Supplementary material is available at Journal of Molluscan Studies
of Euspira heros from the Western Atlantic have up to 84 eggs in a online.
single capsule (Giglioli, 1955). Capsule sizes vary considerably
between species. The largest capsules recorded so far are those of
Falsilunatia carcellesi (Table 5; Penchaszadeh et al., 2016) with dia- ACKNOWLEDGEMENTS
meters of 8.8–14.1 mm. Each capsule accommodates a single larva
that undergoes direct development. The only globisinine whose We would like to express our appreciation to Ms Jingying Wu for
egg capsule has been described is Globisinum drewi from New field support, Ms Hsiao-yun Chuang for the photograph of a liv-
Zealand, which has a capsule diameter of 6.5 mm (Dell, 1956). ing specimen of Notocochlis cernica (Fig. 4A) and Mr Kun-Hsuan
Most species with planktotrophic larvae have capsule sizes ranging Lee (National Museum of Natural Science, Taichung) for photo-
between 100 and 300 μm, while those with larvae that undergo graphs of shells of Sinum haliotoideum and Mammilla melanostoma (Figs
some form of direct development have larger capsules, typically 10A, 11A). Specimens of Pseudopolinices cf. nanus and Euspira cf.
500–3,000 μm, in which some of the multiple eggs may act as plicispira were based on deep-sea trawl samples obtained from
nurse cells (e.g. Laguncula pulchella, Euspira catena and E. montagui; Four-Way Closure Ridge on the research vessel OR1 (CR1108).
Table 5). The size and arrangement of the capsules affect the Profs Ching-Nen Chen and Te-Yu Liao generously facilitated the
external appearance of the egg collar as well as its consistency. use of DNA extraction equipment for this project, while Prof.
Collars with large capsules feel more rigid than those incorporat- Keryea Soong (all National Sun Yat-sen University, Kaohsiung)
ing small capsules. Small capsules are usually arranged haphaz- provided funds for TKS to travel to Taiwan. We are also grateful
ardly in the sand matrix, although it is often difficult to see the to Dr T. Hülsken who generously made DNA sequences obtained
capsules from the surface even with the aid of a stereomicroscope. for his PhD thesis available on the internet. Dr Konstantin
In contrast, collars incorporating large capsules are at once recog- Lutaenko (Vladivostok) assisted with the translation of Russian
nizable due to the bulging collar surfaces created by the capsules text, and Dr Kazunori Hasegawa provided pertinent naticid litera-
within (e.g. Cryptonatica clausa, E. catena, E. pallida, E. heros and ture. Detailed, helpful comments by Drs Alan Kabat, Michael
E. triseriata; e.g. Thorson, 1935; Giglioli, 1955; Table 5). Hollmann and one other referee considerably improved the
Not all naticids lay egg collars. The naticine Natica tecta lays a manuscript. This study was supported by the Asia-Pacific Ocean
series of flattened leaflets made of sand, each attached to the hard Research Center at the National Sun Yat-sen University with
substratum by a stalk (Kilburn & Rippey, 1982: fig. 26). Members funding from the Ministry of Education, Taiwan, as well as the
of the polinicine genus Conuber lay gelatinous egg masses resem- Ministry of Science and Technology, Taiwan (MOST 105-3113-
bling thick sausages that are free of sand (Murray, 1962, 1966; M-005-001; MOST 106-3113-M-005-001) to LLL. This study
Booth, 1995; Table 5). was partially carried out by TKS at the St John’s Island National
The egg masses of some 52 species of Naticidae have been Marine Laboratory, a national research infrastructure under the
described (Table 5), since the first published description of an egg National Research Foundation, Singapore.
collar, that of Euspira catena, by Ankel (1930). This is less than 20%
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Legend to Supplementary Material.
Table S1. Mitochondrial COI sequences from naticid adults and egg collars used to construct
the COI maximum likelihood tree (Fig. 3A). Each sequence is accompanied by the species
name or morphotype of egg collar, with the corresponding locality in Taiwan. The numbers
after the locality indicate the individual sequence used.
Table S2. Mitochondrial 16S sequences from naticid adults and egg collars were used to
construct the 16S maximum likelihood tree (Fig. 3B). Each sequence is accompanied by the
species name or morphotype of egg collar, with the corresponding locality in Taiwan. The
numbers after the locality indicate the individual sequence used.
Appendix 1. COI sequences from naticid egg collars and snails from Taiwan.
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....|
10 20 30 40 50 60 70 80 90 100 110 120 130 140 150
Egg collar Morphotype 1 Husia 1 ------AATT GATTAGTTCC TTTAATATTA GGAGCTCCTG ATATAGCTTT TCCTCGGTTA AATAATATAA GTTTTTGGTT ATTACCTCCT GCACTATTAT TATTATTATC TTCAGCAGCT GTTGAAAGTG GAGTGGGAAC TGGATGGACA
Egg collar Morphotype 2 Kenting 2 -TTTGGTATT GATTAGTTCC TTTAATACTG GGGGCTCCTG ATATAGCTTT CCCTCGTCTG AATAATATAA GTTTTTGGTT ACTTCCACCT GCTTTATTAC TACTTTTATC TTCAGCTGCG GTCGAAAGAG GAGTCGGAAC AGGGTGAACT
Egg collar Morphotype 3 Jingan 4 --------TT GGTTAGTTCC ATTAATATTA GGAGCTCCTG ATATGGCATT TCCGCGATTA AATAATATGA GTTTCTGATT ATTACCTCCA GCTTTATTAT TATTGCTTTC TTCAGCAGCT GTAGAGAGTG GGGTTGGTAC TGGTTGAACA
Egg collar Morphotype 4 Tongliang 41 --------TT GATTAGTTCC TTTAATATTA GGAGCTCCTG ACATAGCTTT TCCTCGTTTA AATAATATAA GTTTCTGATT ACTTCCTCCA GCTTTATTAT TATTACTTTC TTCAGCTGCT GTAGAAAGTG GTGTTGGAAC AGGATGAACA
Egg collar Morphotype 5 Kenting 1 ---------- GATTAGTTCC TTTAATGCTT GGGGCTCCTG ATATAGCTTT TCCTCGACTA AATAATATAA GTTTCTGGTT ACTTCCTCCC GCTTTACTTT TATTATTATC TTCAGCTGCA GTAGAAAGAG GTGTTGGGAC TGGATGAACT
Egg collar Morphotype 6 Tongliang 54 --------TT GACTAGTACC ATTAATGCTA GGGGCCCCAG ACATAGCTTT TCCACGTTTA AATAATATAA GTTTTTGACT ACTGCCTCCA GCGCTTTTAT TACTGTTATC TTCAGCTGCA GTAGAAAGAG GTGTGGGAAC TGGATGAACT
Egg collar Morphotype 7 Daxi 1 --------TT GATTGGTTCC TTTAATGTTG GGAGCTCCTG ATATAGCTTT TCCTCGTTTA AATAATATAA GTTTTTGATT ACTTCCCCCA GCTTTATTAT TATTGCTTTC TTCAGCCGCT GTAGAAAGTG GTGTTGGAAC AGGATGAACA
Egg collar Morphotype 8 Kenting 23 --------CT GGTTAGTGCC TTTGATATTA GGTGCTCCAG ATATAGCTTT TCCTCGCTTA AATAATATAA GTTTTTGATT ATTACCTCCA GCTTTATTAC TATTACTATC ATCAGCTGCT GTAGAAAGCG GTGTAGGAAC TGGTTGAACT
Egg collar Morphotype 9 Kenting 3 -TTTGGAACT GACTGGTTCC CTTAATGTTA GGGGCTCCTG ATATGGCTTT TCCTCGTTTA AATAATATAA GTTTTTGATT ACTACCTCCT GCTTTACTTT TATTATTATC TTCAGCTGCA GTTGAAAGAG GTGTTGGAAC TGGATGAACT
Euspira cf. plicispira FC Ridge 15 ---------- ---------- ---------- --------AG ATATAGCTTT TCCTCGACGA AATAATATAA GTTTTTGACT TTTACCCCCA GCTTTATTGT TATTACTTTC TTCAGCTGCT GTAGAAAGTG GGGTGGGGAC AGGATGAACA
Glossaulax didyma Dongshi 1 --------CT GATTGGTTCC ACTAATGCTA GGAGCTCCAG ATATGGCTTT CCCGCGATTA AATAATATAA GTTTTTGACT TTTACCCCCA GCTTTATTAC TATTACTTTC ATCAGCTGCA GTAGAAAGGG GAGTTGGAAC AGGATGAACA
Glossaulax sp. Tungkang ---------- GATTGGTTCC ATTGATATTA GGGGCTCCAG ATATAGCTTT TCCACGATTA AATAATATAA GTTTCTGACT TCTTCCTCCA GCTTTGTTAC TATTACTTTC GTCGGCCGCG GTAGAGAGAG GAGTCGGGAC AGGATGAACA
Laguncula pulchella Husia 2 ---------- GATTAGTTCC TTTAATACTT GGAGCTCCAG ATATAGCTTT TCCTCGTTTA AATAATATAA GATTTTGACT ACTTCCACCT GCTTTATTAC TTCTACTCTC TTCAGCAGCA GTAGAAAGTG GTGTCGGAAC TGGATGAACA
Natica vitellus Tungkang 2 ---------- GATTGGTGCC TTTAATGTTG GGTGCTCCTG ACATAGCTTT TCCTCGTTTA AATAATATGA GTTTTTGATT ACTGCCTCCA GCATTATTAT TATTACTTTC TTCAGCAGCG GTAGAGAGAG GAGTTGGAAC TGGATGGACG
Naticarius alapapilionis Dongshi --------CT GATTAGTTCC CTTAATACTG GGAGCTCCTG ACATGGCTTT CCCTCGTTTA AATAATATAA GTTTTTGATT GCTACCACCA GCCTTACTAC TTTTACTCTC TTCAGCTGCA GTAGAAAGAG GTGTAGGAAC CGGATGAACT
Naticarius zonalis Tongliang ---------- ---------- ------GTTA GGGGCCCCAG ACATAGCTTT TCCACGTTTA AATAATATAA GTTTTTGACT ATTGCCTCCA GCGCTTTTAT TACTGTTATC TTCAGCTGCA GTAGAAAGAG GTGTGGGAAC TGGATGAACT
Notocochlis gualtieriana Dongwei 2 CCGGGTAATT GATTAGTTCC TTTAATGTTG GGGGCTCCTG ATATAGCTTT CCCTCGTCTG AATAATATAA GTTTTTGGTT ACTTCCACCT GCTCTATTAC TACTTTTATC TTCAGCTGCG GTTGAAAGAG GAGTTGGAAC AGGATGAACT
Notocochlis sp. aff. antoni Husia 2 ------AATT GATTAGTTCC TTTAATATTA GGGGCCCCTG ATATGGCTTT TCCCCGTTTA AATAATATAA GTTTTTGATT ACTTCCACCT GCTTTGTTAT TGCTATTATC TTCAGCTGCA GTTGAAAGAG GGGTTGGAAC AGGATGAACT
Paratectonatica tigrina Taishi 1 --------TT GATTAGTTCC TTTAATATTA GGAGCTCCTG ATATAGCTTT TCCTCGGTTA AATAATATAA GTTTTTGGTT ATTACCTCCT GCACTATTAT TATTATTATC TTCAGCAGCT GTTGAAAGTG GAGTGGGAAC TGGATGGACA
Polinices flemingianus Kenting --------TT GATTAGTTCC TTTAATATTA GGAGCTCCTG ATATAGCTTT TCCTCGTTTA AACAATATAA GTTTTTGGTT GCTTCCTCCA GCTTTATTAT TACTACTTTC TTCAGCAGCT GTAGAGAGTG GTGTTGGTAC AGGATGAACA
Polinices sp.1 Dongshi ---------- GATTAGTTCC TTTAATATTA GGGGCTCCTG ATATAGCTTT TCCTCGTTTA AATAATATAA GTTTTTGATT ATTGCCTCCT GCTTTACTGT TATTACTTTC TTCAGCGGCT GTGGAAAGCG GTGTTGGAAC AGGATGAACA
Polinices sp.2 Tungkang 3 ---------- --------CC TTTAATATTA GGAGCTCCTG ATATGGCTTT TCCCCGATTA AATAACATAA GTTTCTGGTT ATTACCTCCT GCCTTATTGT TATTACTTTC TTCAGCGGCT GTAGAAAGTG GTGTTGGAAC AGGATGGACA
Sinum haliotoideum Dongsha ---------- GGTTAGTTCC ATTAATATTA GGAGCTCCTG ATATGGCATT TCCGCGATTA AATAATATGA GTTTCTGATT ATTACCTCCA GCTTTATTAT TATTGCTTTC TTCAGCGGCT GTAGAGAGTG GGGTTGGTAC TGGTTGAACA
Sinum sp.1 Daxi ---------- ------TTCC TTTAATGTTA GGAGCTCCTG ACATAGCATT TCCTCGTTTG AATAATATAA GTTTTTGATT ATTACCTCCT GCTTTATTAT TATTACTTTC ATCTGCTGCA GTAGAAAGTG GTGTGGGAAC AGGATGAACA
Tanea lineata Dongshi 1 ---------- GATTAGTTCC TTTAATATTA GGAGCACCAG ACATAGCCTT TCCACGTCTT AATAATATAA GTTTTTGGCT ATTGCCTCCT GCTTTATTGT TACTTTTATC TTCAGCTGCT GTAGAAAGAG GTGTAGGAAC TGGATGAACA
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....|
160 170 180 190 200 210 220 230 240 250 260 270 280 290 300
Egg collar Morphotype 1 Husia 16 GTATACCCTC CATTATCTGG AAATTTAGCT CATGCAGGAG GATCTGTAGA TTTAGCAATT TTTTCTTTAC ACTTGGCTGG AGTTTCTTCC ATTTTGGGTG CTGTAAATTT TATTACCACG ATTATTAATA TACGATGACG TGGAATACAA
Egg collar Morphotype 2 Kenting 2 GTTTATCCTC CTTTATCTGG AAATTTGGCA CATGCCGGAG GCTCTGTAGA TTTAGCAATT TTTTCTTTAC ATTTAGCTGG TGTTTCATCT ATTTTAGGTG CCGTTAATTT TATTACAACG ATTATTAATA TGCGATGACG GGGTATGCAA
Egg collar Morphotype 3 Jingan 4 GTTTATCCTC CACTAGCTGG TAATTTAGCT CACGCCGGTG GTTCAGTAGA CTTGGCAATT TTTTCTTTAC ATTTAGCAGG TGCTTCATCT ATTTTAGGTG CTGTAAATTT TATTACTACT ATTATTAATA TACGATGACG AGGGATAGAA
Egg collar Morphotype 4 Tongliang 41 GTTTATCCTC CATTATCTGG TAATTTAGCT CATGCTGGTG GTTCTGTAGA CTTAGCAATT TTTTCTCTTC ATTTGGCTGG AGCTTCTTCA ATTTTAGGGG CTGCAAATTT TATTACTACT ATTATTAACA TACGATGACG AGGAATACAA
Egg collar Morphotype 5 Kenting 1 GTTTATCCTC CACTATCTGG AAATTTAGCA CATGCTGGTG GATCAGTTGA TTTAGCAATT TTCTCTTTAC ATTTAGCAGG TGTCTCATCT ATTTTAGGGG CCGCTAATTT TATTACAACA ATTATTAATA TGCGATGACG TGGTATACAA
Egg collar Morphotype 6 Tongliang 54 GTTTATCCTC CTTTATCTGG GAACTTAGCT CATGCAGGTG GCTCTGTAGA TCTAGCAATT TTTTCACTTC ATTTAGCAGG TGCTTCCTCG ATTTTAGGGG CCGTTAATTT TATTACGACT ATCATTAATA TACGGTGACG TGGTATACAA
Egg collar Morphotype 7 Daxi 1 GTTTATCCTC CATTGTCTGG TAATTTAGCT CATGCTGGTG GTTCTGTAGA CTTAGCAATC TTTTCTCTTC ATTTAGCTGG TGCTTCTTCA ATTTTAGGTG CTGCAAATTT TATTACTACC ATTATTAATA TGCGATGACG AGGAATACAA
Egg collar Morphotype 8 Kenting 23 GTATATCCAC CACTGTCTGG AAACTTGGCC CATGCCGGAG GTTCAGTTGA TTTAGCTATT TTTTCTCTTC ATTTAGCTGG TGCTTCTTCT ATTCTAGGTG CTGTAAACTT TATTACCACA ATTATTAATA TACGATGACG AGGAATAAAT
Egg collar Morphotype 9 Kenting 3 GTTTATCCTC CATTGTCCGG AAATTTAGCA CATGCTGGTG GATCAGTTGA TTTGGCAATT TTTTCTTTAC ATTTAGCAGG TGTTTCATCT ATTTTAGGGG CTGCCAATTT TATTACGACA ATTATTAATA TACGATGACG TGGTATGCAA
Euspira cf. plicispira FC Ridge 15 GTTTATCCCC CTTTATCTGG AAATTTAGCT CATGCTGGAG GCTCAGTAGA TTTAGCAATT TTTTCACTTC ATCTAGCTGG AGCTTCTTCT ATTTTAGGTG CCGTAAATTT TATTACTACG ATTATTAATA TACGATGACG AGGGATACAA
Glossaulax didyma Dongshi 1 GTCTATCCTC CTTTATCTGG TAATTTGGCT CATGCTGGTG GTTCTGTGGA TTTAGCAATT TTTTCTCTTC ATTTAGCTGG TGCTTCTTCT ATTTTAGGTG CTGTAAATTT TATTACTACG ATTATTAATA TACGATGACG GGGAATACAA
Glossaulax sp. Tungkang GTTTATCCTC CTTTATCCGG TAACCTGGCT CACGCCGGTG GTTCTGTAGA TTTAGCAATT TTCTCTCTTC ATTTAGCTGG TGCTTCTTCT ATTTTAGGTG CTGTAAATTT CATTACTACG ATTATTAACA TACGATGACG GGGGATACAA
Laguncula pulchella Husia 2 GTTTATCCTC CGCTATCAGG TAATCTAGCT CATGCTGGTG GATCTGTAGA TTTAGCAATT TTTTCTCTTC ATTTAGCTGG TGTTTCTTCA ATTCTAGGTG CTGTAAATTT TATTACTACA ATTATCAATA TACGATGACG GGGAATACAA
Natica vitellus Tungkang 2 GTTTATCCTC CGTTATCAGG GAATCTTGCT CATGCTGGGG GTTCGGTGGA TTTAGCTATT TTTTCTTTAC ATTTAGCTGG AGCTTCATCT ATTTTAGGTG CTGTAAATTT TATTACTACA ATTATTAATA TACGGTGACG TGGAATACAA
Naticarius alapapilionis Dongshi GTCTATCCTC CCTTATCTGG TAACCTGGCT CATGCCGGTG GATCTGTAGA TTTAGCAATT TTTTCTTTAC ACTTAGCTGG TGTATCTTCT ATTTTAGGAG CTGTTAATTT TATTACTACT ATTATTAATA TACGATGACG AGGTATACAG
Naticarius zonalis Tongliang GTTTATCCTC CTTTATCTGG AAACTTAGCT CATGCAGGTG GCTCTGTAGA TCTAGCAATT TTTTCACTTC ATTTAGCAGG TGCTTCTTCC ATTTTAGGGG CCGTTAATTT TATTACGACT ATCATTAATA TACGGTGACG TGGTATACAA
Notocochlis gualtieriana Dongwei 2 GTTTATCCTC CTTTATCTGG AAATTTGGCA CATGCCGGAG GTTCTGTAGA TTTAGCAATT TTTTCTTTAC ATTTAGCTGG TGTCTCATCT ATTTTAGGTG CCGTTAATTT TATTACAACG ATTATTAATA TGCGATGACG GGGTATGCAA
Notocochlis sp. aff. antoni Husia 2 GTCTACCCTC CTCTATCTGG AAACTTAGCG CATGCTGGAG GTTCAGTAGA TTTAGCAATT TTTTCTTTGC ATTTAGCTGG TGTTTCATCT ATTTTAGGTG CTGTCAATTT TATTACAACA ATTATTAATA TACGATGACG GGGTATGCAA
Paratectonatica tigrina Taishi 1 GTATACCCTC CATTATCTGG AAATTTAGCT CATGCAGGAG GATCTGTAGA TTTAGCAATT TTTTCTTTAC ACTTGGCTGG AGTTTCTTCC ATTTTGGGTG CTGTAAATTT TATTACCACG ATTATTAATA TACGATGACG TGGAATACAA
Polinices flemingianus Kenting GTCTATCCTC CCCTATCTGG TAATTTAGCT CATGCTGGTG GTTCTGTAGA TTTAGCAATT TTCTCTCTTC ATTTAGCAGG TGCTTCTTCA ATTTTAGGTG CTGCAAATTT TATCACTACT ATCATTAATA TACGATGACG AGGAATACAA
Polinices sp.1 Dongshi GTTTATCCTC CATTGTCAGG TAATTTGGCT CATGCTGGTG GTTCTGTAGA TTTGGCAATT TTTTCTCTCC ATTTAGCAGG TGTTTCTTCA ATTTTGGGTG CTGCTAACTT TATTACTACT ATTATTAATA TACGATGACG AGGTATGCAA
Polinices sp.2 Tungkang 3 GTCTATCCTC CATTGTCAGG TAATTTGGCT CATGCTGGTG GTTCTGTAGA TTTAGCAATT TTTTCTCTTC ATTTAGCAGG TGCTTCTTCA ATTTTAGGTG CTGCTAATTT CATTACTACT ATTATTAATA TACGATGACG AGGTATACAA
Sinum haliotodieum Dongsha GTTTATCCTC CACTAGCTGG TAATTTAGCT CATGCCGGTG GTTCAGTAGA CTTGGCAATT TTTTCTTTAC ATTTAGCAGG TGCTTCATCT ATTTTAGGTG CTGTAAATTT TATTACTACT ATTATTAATA TACGATGACG AGGGATAGAA
Sinum sp.1 Daxi GTTTATCCCC CATTAGCTGG TAATTTAGCT CATGCTGGGG GTTCAGTGGA TTTAGCAATT TTTTCTTTAC ATTTAGCTGG TGCTTCTTCT ATTTTAGGTG CTGTAAATTT TATTACTACT ATTATTAATA TACGGTGACG AGGAATGCAA
Tanea lineata Dongshi 1 GTGTATCCTC CTTTATCTGG AAACTTGGCT CACGCCGGTG GATCTGTTGA TTTAGCGATT TTTTCACTTC ATTTAGCTGG GGCTTCTTCG ATTTTAGGTG CTGTAAACTT TATTACTACA ATTATTAATA TACGATGACG AGGTATACAA
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....
310 320 330 340 350 360 370 380 390 400
Egg collar Morphotype 1 Husia 16 TTTGAACGAT TACCTTTATT CGTTTGATCT GTAAAAATTA CGGCCATCTT ACTACTTCTT TCCCTTCCTG TATTAGCTGG AGCTATTACT ATACTTTA-- ---------
Egg collar Morphotype 2 Kenting 2 TTTGAGCGAC TTCCTTTATT TGTCTGATCT GTAAAAATTA CAGCCATTTT ACTTTTACTT TCTTTACCAG TTTTAGCAGG TGCCATTACT ATA------- ---------
Egg collar Morphotype 3 Jingan 4 TTTGAACGAC TGCCGTTATT TGTATGATCT GTAAAAATTA CAGCAGTTTT GTTGCTTTTA TCTTTACCAG TCTTGGCTGG AGCTATTACT ATACTTTTAA CTGATCGA-
Egg collar Morphotype 4 Tongliang 41 TTTGAGCGAT TACCTTTATT TGTATGATCA GTAAAAATTA CTGCTATTTT ACTATTGTTG TCTCTACCTG TTTTAGCTGG AGCTATTACT ATACTTTT-- ---------
Egg collar Morphotype 5 Kenting 1 TTTGAACGTC TTCCATTATT TGTTTGATCA GTAAAAATTA CTGCTATTTT GCTTCTACTT TCTTTACCAG TTCTAGCTGG AGCTATTACT ATA------- ---------
Egg collar Morphotype 6 Tongliang 54 TTTGAACGAC TTCCTTTATT TGTTTGATCT GTAAAAATTA CAGCTGTTCT ACTTTTACTT TCTTTACCAG TTCTAGCAGG TGCTATTACT ATACTTTT-- ---------
Egg collar Morphotype 7 Daxi 1 TTTGAGCGAT TACCATTATT TGTATGATCA GTAAAAATTA CTGCTATTTT ATTATTGTTG TCTTTACCTG TTTTAGCTGG TGCTATTACT ATACTTTT-- ---------
Egg collar Morphotype 8 Kenting 23 TTTGAACGAT TGCCACTGTT CGTATGATCA GTAAAAATTA CAGCTGTATT ATTATTACTA TCTCTTCCTG TATTAGCTGG TGCTATTACT ATACTT---- ---------
Egg collar Morphotype 9 Kenting 3 TTTGAACGGC TTCCACTATT TGTTTGATCT GTAAAAATTA CTGCTATTTT ACTCTTACTT TCTTTACCAG TTTTGGCTGG AGCTATTACT ATACTAAAAC TTTATTCGA
Euspira cf. plicispira FC Ridge 15 TTTGAACGGC TACCTCTTTT TGTCTGATCT GTTAAGATTA CAGCTATTTT GTTACTTTTA TCTCTACCGG TATTAGCTGG AGCTATTACT ATA------- ---------
Glossaulax didyma Dongshi 1 TTTGAACGGT TACCATTATT TGTATGATCT GTAAAAATTA CAGCAATTTT ATTGTTATTA TCTTTGCCGG TGTTAGCAGG AGCTATTACT ATA------- ---------
Glossaulax sp. Tungkang TTTGAGCGGT TACCTTTATT TGTATGATCT GTAAAAATTA CAGCAATTTT ATTACTGTTA TCTTTACCAG TGTTAGCAGG AGCGATTACT ATA------- ---------
Laguncula pulchella Husia 2 TTTGAACGAC TACCTTTATT TGTTTGATCC GTTAAAATTA CGGCAATTTT ATTATTACTC TCTTTACCAG TTCTGGCAGG AGCTATTACT ATACTTTT-- ---------
Natica vitellus Tungkang 2 TTTGAGCGAT TGCCTTTATT TGTGTGGTCT GTAAAAATTA CGGCTATTTT GTTGCTTTTG TCGTTGCCTG TATTGGCGGG GGCTATTACT ATA------- ---------
Naticarius alapapilionis Dongshi TTTGAACGAC TTCCCTTATT CGTTTGATCA GTAAAAATTA CGGCAATTTT ACTTCTATTA TCTTTACCTG TTTTAGCAGG AGCTATTACT ATA------- ---------
Naticarius zonalis Tongliang TTTGAACGAC TTCCTTTATT TGTTTGATCT GTAAAAATTA CAGCTGTTCT ACTTTTACTT TCTTTACCAG TTCTAGCAGG TGCTATTACA ATACTTTT-- ---------
Notocochlis gualtieriana Dongwei 2 TTTGAGCGAC TTCCTTTGTT CGTTTGATCT GTAAAAATTA CAGCTATTTT ACTTCTACTT TCTTTACCAG TTTTAGCAGG TGCTATTACT ATACTTTT-- ---------
Notocochlis sp. aff. antoni Husia 2 TTTGAGCGAC TTCCTCTATT TGTTTGATCT GTAAAAATTA CAGCCATTTT ACTCTTGCTT TCTTTGCCAG TTTTGGCAGG TGCTATTACT ATACTTTT-- ---------
Paratectonatica tigrina Taishi 1 TTTGAACGAT TACCTTTATT CGTTTGATCT GTAAAAATTA CGGCCATCTT ACTACTTCTT TCCCTTCCTG TGTTAGCTGG AGCTATTACT ATACTTTT-- ---------
Polinices flemingianus Kenting TTTGAGCGGT TACCTTTATT CGTTTGATCT GTAAAAATTA CTGCTATCTT ATTGCTTTTG TCTTTGCCTG TCTTAGCTGG AGCCATTACT ATACTTTTAA CTGATCGA-
Polinices sp.1 Dongshi TTTGAACGGT TACCTTTATT TGTGTGATCT GTAAAAATTA CTGCTATTTT GTTGCTTCTG TCTCTACCTG TTTTAGCCGG TGCTATTACT ATACTTTT-- ---------
Polinices sp.2 Tungkang 3 TTTGAACGAT TACCTTTATT TGTGTGATCT GTGAAAATTA CTGCTATTTT GTTACTTCTG TCTTTGCCTG TTTTAGCTGG TGCTATTACT ATACTTTT-- ---------
Sinum haliotoideum Dongsha TTTGAACGAC TGCCGTTATT TGTATGATCT GTAAAAATTA CAGCAGTTTT GTTGCTTTTA TCTTTACCAG TCTTGGCTGG AGCTATTACT ATACTTT--- ---------
Sinum sp.1 Daxi TTTGAACGGT TACCATTATT TGTTTGATCT GTAAAAATTA CGGCAATTTT ATTGCTTTTA TCTCTTCCAG TTTTAGCTGG TGCTATTACT ATACTTTTA- ---------
Tanea lineata Dongshi 1 TTTGAACGGT TACCTCTCTT TGTATGATCT GTGAAAATTA CAGCTATTCT ATTACTTTTG TCACTTCCTG TATTAGCAGG AGCAATTACT ATACTTTTAA ACTGATCG-
Appendix 2. 16S rRNA sequences from naticid egg collars and snails from Taiwan.
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....|
10 20 30 40 50 60 70 80 90 100 110 120 130 140 150
Egg collar Morphotype 1 Husia 1605 AAAGACTTCT GCATCTTGTA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATATGAAC TCTCAAAATG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATTAAAA-TA A-TGGATCAT TATTTATAAA
Egg collar Morphotype 2 Husia 67 GAAGACTTCT GCATCTATCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATACGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAGG-AAT ---GGATCAA AATTTATAAA
Egg collar Morphotype 3 Jingan 4 GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATGAGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATTA-AAC-- A-TGGATCAC TATTTATAAA
Egg collar Morphotype 4 Tongliang 41 GAAGACTTCT GCATCTTTCA GGAAATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATATGAGC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACT--AAGTA A-TGGATCAT TATTCATAAA
Egg collar Morphotype 5 Kenting 1 GAAGACTTCT GCATCTATCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAT GGTAACTAAT TCTTTTAATC ACTAAG-AGT ---GGATCAA AATTTATAAA
Egg collar Morphotype 6 Tongliang 54 GAAGGCTTCT GCACCTGTCG GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATACGGAC TCTTAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAAGTAGT ---GGATCAA TATTTATATA
Egg collar Morphotype 7 Daxi 1 GAAGACTTCT GCATCTTTCA GGAAATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATATGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACT--AAATA G-TGGATCAC TACTCATAAA
Egg collar Morphotype 8 Kenting 23 GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATAAGAAC TCTTAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTCAATC ATTAAT--TA A-TGGATCAC TATTTATTTA
Egg collar Morphotype 9 Kenting 3 GAAGACTTCT GCATCTATCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAT GGTAACTAAT TCTTTTAATC ACTAAA-AAT ---GGATCAA AATTTATAAA
Egg collar Morphotype 10 Husia 23 GAAGACTTCT GCATCTGTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAAA-AAT ---GGATCAA GATTTATAAA
Egg collar Morphotype 11 Kenting 21 GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACTAAA-AGT ---GGATCAA AATTTATAGA
Glossaulax didyma Tungkang 1 GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATTAGAAC TCTCAAAAAA GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC GTTACGAT-A A-CGGATCAT TACTTATAAG
Glossaulax sp. Tungkang GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATTAGAAC TCTCAAAAAA GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC GTTATAAC-A A-CGGATCAT TACTTATAAG
Laguncula pulchella Husia 2 GAAGACTTCT GCATCTTTCA GGATATCCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATTAGAAC TCTCAAAAAA GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTCAATC ATTATAAGTA A-TGGATCAT TATTCATAAA
Natica vitellus Tungkang 2 GAAGACTTCT GCATCTATCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATAAGAAC TCTCAAAAAA GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACTAAAA--T AGTGGATCTA AATTTATAAA
Naticarius alapapilionis Dongshi GAAGACTTCT GCATCTGTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAAA-AAT ---GGATCAA TATTTATAAA
Naticarius zonalis Tongliang GAAGGCTTCT GCACCTGTCG GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATACGGAC TCTTAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAAGTAGT ---GGATCAA TATTTATATA
Notocochlis gualtieriana Dongwei 3 GAAGACTTCT GCATCTATCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAGG-AAT ---GGATCAA AATTTATAAA
Notocochlis sp. aff. antoni Husia 3 GAAGACTTCT GCATCTGTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACTCTTTTTT CGATATGAAC TCTCAAAAAG AATTATGCTG TTATCCCTAC GGTAACTAAT TTTTTTAATC ACTAAA-AAT ---GGATCAA GATTTATAAA
Paratectonatica tigrina Husia AAAGACTTCT GCATCTTGTA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATATGAAC TCTCAAAATG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATTAAAA-TA A-TGGATCAT TATTTATAAA
Polinices flemingianus Kenting GAAGACTTCT GCATCTTTCA GGAAATTCTA GTCCAACATC GAGGTCACAA ACCCTCTTTT CGATATGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACTATAAGTA G-TGGATCAC TAATTATAAA
Polinices sp.1 Tungkang 1 GAAGACTTCT GCATCTTTCA GGAAATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATAAGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATT--AAATA A-TGGATCAT TATTTATAAA
Polinices sp.2 Tungkang 3 GAAGACTTCT GCATCTTTCA GGAAATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATAAGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATT--AAATA A-TGGATCAT TATTTATAAA
Pseudopolinices cf. nanus FC Ridge 14 GAAGACTTCT GCATCTTTCA GGATATCCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATTAGAGC TCTTAAAAAA GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ACTATAATTA G-TGGATCAT TATTTATAAA
Sinum haliotoideum Dongsha GAAGACTTCT GCATCTTTCA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATGAGAAC TCTCAAAAAG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATTA-AAC-- A-TGGATCAT TATTTATAAA
Tanea lineata Dongshi 1 GAAGACTTCT GCATCTTACA GGATATTCTA GTCCAACATC GAGGTCACAA ACCCTTTTTT CGATATGAAC TCTCAAAATG GATTATGCTG TTATCCCTAC GGTAACTAAT TCTTTTAATC ATTAATAATA A-TGGATCAT TACTCATAAG
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....|
160 170 180 190 200 210 220 230 240 250 260 270 280 290 300
Egg collar Morphotype 1 Husia 1605 TTTT-AT-TT TTTTAAGAAG TTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTAC-A TAATTAAAAA TATT-T-TAA ATTTTTTTTA TTTTA-TATA TTAATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TAATTTAGGA
Egg collar Morphotype 2 Husia 67 TATGTCA--C TAATAAGAAG CTTTAATTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATCAAAAG TATTTT-TA- ATCTTTTTTA TTTTA-TAAA TGTTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GTATTTGAGT
Egg collar Morphotype 3 Jingan 4 TCTAGTTTAG TTGAAAGAAG CTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATCTTTT-A TAATCAAAAA TATAATAA-T ATTTTTTTTA TTTTA-TAAA TGTATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT AAATTTAGGT
Egg collar Morphotype 4 Tongliang 41 TTTTAAG-TT CATAAAGAAG CTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTTC-A TAACCAGAAA TATATC-TT- ATTTTCTTTA TTTTATCGAA TAGATTTAAA -GCTCGATAG GGTCTTCTTG TCTATTAATT TTATTTAGGT
Egg collar Morphotype 5 Kenting 1 TTTGTTG--T TTAAAAGAAG CTTTGTTTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATTAAAAG TATTTTATA- ATCTTTTTTA TTTTA-TAAA TGTTTTTAAA -GCTCAATAG GGTCTTCTTG TCTTTCAATA ATATTTGAGT
Egg collar Morphotype 6 Tongliang 54 TTCAATA--T TGTTAAGAAG CTTTAATTGT TCCTTAGTCG CCCCAACCAA AATTTTT-CT TAATTAAGAT TACCAATTG- ATTTCTTTTA TTTTA-AAAA TATTTTTAAA -GCTCGATAG GGTCTTCTTG TCTATCAATA ATATTTGAGT
Egg collar Morphotype 7 Daxi 1 TCTTAAA-TA TATAAAGAAG CTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTTC-A TAATTAGAAA TATACC-CT- ATTTTCTTTA TTTTATTAAA CGAATTTAAA -GCTCGATAG GGTCTTCTTG TCTATTAATT TTATTTAGGT
Egg collar Morphotype 8 Kenting 23 TTTT-AT-TT TTTCAAGAAG CTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTGTT-A TAATCGAAAG TATT-T-TAC ATCTTTTTTG TTTTA-TAAA TAAATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATTTAGGT
Egg collar Morphotype 9 Kenting 3 TCTATTA--T TTAAAAGAAG CTTTATTTGT TCCTTAGTCG CCCCAACCAA AATTTTT-AA TAATTAAAAG TATTTTATA- ATCTTTTTTA TTTTA-TAAA TATTTTTAAA -GCTCAATAG GGTCTTCTTG TCTTTCAATA ATATTTGAGT
Egg collar Morphotype 10 Husia 23 TTTGCTG--T TAGTAAGAAG CTTTAGTTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATTAAAAG TGTTTT-TA- ACCTTTTTTA TTTTA-TAAA TGTTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GTACTCGAGT
Egg collar Morphotype 11 Kenting 21 TCTATTA--T CAGTAAGAAG CTTTAACTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAAATAAGAG TATTCT-TA- ACCTCTTTTA TTTTA-TAAA TACTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GTATTTGGGT
Glossaulax didyma Tungkang 1 TTAA-GTTAT TAGAAAGAAG CTTTGTTTGT TCCTTAGTCG CCCCAACCAA AATTTTTT-A CAATTGAAAA TAT-TGAATT ATTTTTTTTA TCTTG-TAAA TATATTTCAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATCTAAGA
Glossaulax sp. Tungkang TTTA-GTTAT TGGAAAGAAG CTTTATTTGT TCCTTAGTCG CCCCAACCAA AATCTTTT-A CAATTGAAAA TAT-TAAATT ATTTTTTTTA TTTTG-TAAA TATATTTCAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TGATCTAAGA
Laguncula pulchella Husia 2 TCTAAATTTT -ATAAAGAAG CTTTATTTGT TCCTTAGTCG CCCCAACCAA AATTTTTT-A TGATTAAAAA TATTAAA-TA ATTTTTGTTA TTTCA-TAAA TGTATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT AAATTTAGGT
Natica vitellus Tungkang 2 TCTCAAT-CT TATAAAGAAG CTTTTCTTGT TCCTTAGTCG CCCCAACCAA AATATTAT-A TAACTAAAGA TATC-TATAT ACTCTTTTCA TCTTA-TATA TATATTTAAA AGCTCGATAG GGTCTTCTTG TCTTTTAATT AAATTAAGGC
Naticarius alapapilionis Dongshi TTCATTA--T TTATAAGAAG CTTTGCTTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATCAAGAG TATTC-ATA- ATCTCTTTCA TTTTA-TAAA TATTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GTATTTGAGT
Naticarius zonalis Tongliang TTCAATA--T TGTTAAGAAG CTTTAATTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TT TAATTAAGAT TACCAATTG- ATTTCTTTTA TTTTA-AAAA TATTTTTAAA -GCTCGATAG GGTCTTCTTG TCTATCAATA ATATTTGAGT
Notocochlis gualtieriana Dongwei 3 TATGTCA--C TAATAAGAAG CTTTAATTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATCAAAAG TATTTT-TA- ATCTTTTTTA TTTTA-TAAA TGTTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GCATTTGAGT
Notocochlis sp. aff. antoni Husia 3 TTTGCTG--C TAGTAAGAAG CTTTAGCTGT TCCTTAGTCG CCCCAACCAA AATTTTT-TA TAATTAAAAG TGTTTT-TA- ACCTTTTTTA TTTTA-TAAA TGTTTTTAAA -GCTCGATAG GGTCTTCTTG TCTTTCAATT GTACTCGAGT
Paratectonatica tigrina Husia TTTT-AT-TT TTTTAAGAAG TTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTAC-A TAATTAAAAA TATT-T-TAA ATTTTTTTTA TTTTA-TATA TTAATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TAATTTAGGA
Polinices flemingianus Kenting TCTTAAA-TT TATAAAGAAG CTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTCT-A TAATCAGAAA TAT-TT-GTA ATTTTCTTTA TTTTA-TAGA CAAACTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT CTATTTAGGA
Polinices sp.1 Tungkang 1 TTTTCAA-TT TAAAAAGAAG CTTTTTCTGT TCCTTAGTCG CCCCAACCAA AATTTTTC-A TAATTAGAAA TATCAA-TTG ATTTTCTTTA TCTTA-TGGA TAGATTTTAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATTTAGGT
Polinices sp.2 Tungkang 3 TTTTTAA-TT TAAAAAGAAG CTTTTTCTGT TCCTTAGTCG CCCCAACCAA AATTTTTC-A TAATTAGAAA TATTAA-TTA ATTTTCTTTA TCTTA-TAAA TGGATTTTAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATTTAGGT
Pseudopolinices cf. nanus FC Ridge 14 TCTAAGTTAT -ATGAAGAAG CTTTGTTTGT TCCTTAGTCG CCCCAACCAA AATTTTTT-A TGATTAAAAA TATCAAAATG ATTTTTGTCA TTTCA-TAAA TGGATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATTTAGGT
Sinum haliotoideum Dongsha TCTAGTTTAG TTGAAAGAAG CTTTTTTTGT TCTTTAGTCG CCCCAACCAA AATCTTTT-A TAATTAAAAA TATAATAA-T ATTTTTTTTA TTTTA-TAAA TGTATTTAAA -GCTCGATCG GGTCTTCTTG TCTTTTAATT AAATTTAGGT
Tanea lineata Dongshi 1 TCTTGAA-AT TATTAAGAAG TTTTTTTTGT TCCTTAGTCG CCCCAACCAA AATTTTTT-A TAACCTAAAA TATTAA-TAA ATTTTTTTTA TTTTA-TAAA TTTATTTAAA -GCTCGATAG GGTCTTCTTG TCTTTTAATT TTATTTAGGA
....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|....| ....|.
310 320 330 340 350 360 370 380 390 400
Egg collar Morphotype 1 Husia 1605 TTCTTCACCT AAAAAATAAG TTCTATAAAT TTTCGTAGAG ACAGTATTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Egg collar Morphotype 2 Husia 67 TTCTTCACTC AAAAAATAAA TTCTAAAAAT TTTTGTAGAG ACAGTGTAAT TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTACC
Egg collar Morphotype 3 Jingan 4 CTCTTCACCT AAAGGATAAA TTCTAAAAAT TTCTATGGAG ACAGTATTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTA--
Egg collar Morphotype 4 Tongliang 41 TTCTTCACCT AAAAAATAAT TTCTATCAAT TTTTATGGAG ACAGTATTGC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATG----- ------
Egg collar Morphotype 5 Kenting 1 TTCTTCACTC AAAAAATAAA TT-------- ---------- ---------- ---------- ---------- ---------- ---------- ---------- ------
Egg collar Morphotype 6 Tongliang 54 TTCTTCACTC AAAGAATAAA TTCTAGAAAT TTTTGTAGAG ACAGAATAAC TCTTGTCAAA CCATTCATAC TAGCCCTCCA ATTATAGGGC AAATG----- ------
Egg collar Morphotype 7 Daxi 1 TTCTTCACCT AAAAAATAAT TTTTATTAAT TTTTATAGAG ACAGTATTGC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTA--
Egg collar Morphotype 8 Kenting 23 TTCTTCACCT AAAAAATAGG TTCTAAGAAT TTTTATGGAG ACAGTACTAC TCTCGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTA--
Egg collar Morphotype 9 Kenting 3 TTCTTC---- ---------- ---------- ---------- ---------- ---------- ---------- ---------- ---------- ---------- ------
Egg collar Morphotype 10 Husia 23 TTCTTCACTC GAAAAATAAA TTCTGAAAAT TTTTGTAGAG ACAGTGTAAT TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTA--
Egg collar Morphotype 11 Kenting 21 TTCTTCACCC AAAGAATAAA TTCTAAAAAT TTCTGTAGAG ACAGTGTAAT TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTACC
Glossaulax didyma Tungkang 1 TTCTTCACTT AGAAAATAAA TTCTAGAAAT TTTTGCGGAG ACAGTATTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTACAGGGC AAATGATTAT GCTA--
Glossaulax sp. Tungkang TTCCTCACTT AGAAAATAAA TTCTAGAAAT TTTTGCGGAG ACAGTATTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTACAGGGC AAATGATTAT GCT---
Laguncula pulchella Husia 2 TTCTTCACCT AAAAAATAAA TTCTAAAAAT TTTTACGGAG ACAGTACTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATT-- ------
Natica vitellus Tungkang 2 TTCTTCACCT CAAAAATAAA TTCTAACAAA TTTTATAGAG ACAGCACTAT TCTCGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGAGC AAATGATTAT GCT---
Naticarius alapapilionis Dongshi TTCTTCACTC AAAAAATAAG TTCTAAGAAT TTCTATAGAG ACAGTATAAC TCATGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCTA--
Naticarius zonalis Tongliang TTCTTCACTC AAAGAATAAA TTCTAGAAAT TTTTGTAGAG ACAGAATAAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Notocochlis gualtieriana Dongwei 3 TTCTTCACTC AAAAAATAAA TTCTAAAAAT TTTTGTAGAG ACAGTGTAAT TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Notocochlis sp. aff. antoni Husia 3 TTCTTCACTC GAAAAATAAA TTCTGAAAAT TTTTGTAGAG ACAGTGTAAT TCTTGTCAAA CCATTCATAC TAGCCCTCTA ATTATAGGGC AAATGATTAT GCTACC
Paratectonatica tigrina Husia TTCTTCACCT AAAAAATAAG TTCTATAAAT TTTCGTAGAG ACAGTATTAC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Polinices flemingianus Kenting TTCTTCACCT AAAAAATAAT TTCTATTAAT TTCTATGGAG ACAGTATTGC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AA-TGATTAT GATA--
Polinices sp.1 Tungkang 1 TCCTTCACCT AAAAAATAAT TTCTATTAAT TTTTATGGAG ACAGTATTGC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Polinices sp.2 Tungkang 3 TCCTTCACCT AAAAAATAAT TTCTATTAAT TTTTATGGAG ACAGTATTGC TCTTGTCAAA CCATTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT GCT---
Pseudopolinices cf. nanus FC Ridge 14 TTCTTCACCT AAAAAATAAA TTCTTAAAAT TTTTGCGGAG ACAGTATTAC TCTTGTCGAT CCATTCATAC TAGCCCTC-A ATTATTGGGC AA-------- ------
Sinum haliotoideum Dongsha CTCTTCACCT AAAGGATAAA TTCTAATAAT TTCTATGGAG ACCGC-TTAC TCTTCTCAAC CCATTCCTAC TAGCCCTC-- ---------- ---------- ------
Tanea lineata Dongshi 1 TTCTTCACCT AAAAAATAAA TTCTATTTAG TCTCATAGAG ACAGCTTTAC TCTCGTCAAA CCGTTCATAC TAGCCCTC-A ATTATAGGGC AAATGATTAT G-----
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Egg-collar morphology and identity of nine species of Naticidae (Gastropoda)

Article in Journal of Molluscan Studies · November 2018

Dun-Ru Kang Koh Siang Tan

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Egg-collar morphology and identity of nine species of Naticidae (Gastropoda)

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Species Adult animal Egg collar Habitat; remarks

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Figure 3. Continued on next page

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EGG COLLARS OF NATICIDAE

Genus COI 16S

Notocochlis 0.128 ± 0.025 0.063 ± 0.013

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Egg-collar morphology and molecular phylogeny DISCUSSION

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Basal Apical diameter,

D.-R. KANG ET AL.

EGG COLLARS OF NATICIDAE

Basal Apical diameter,

Conuber incei (Philippi, Diffuse in gelatinous

D.-R. KANG ET AL.

EGG COLLARS OF NATICIDAE

1822) (Polinicinae) spheres

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