Revision of Euprotomus Gill, 1870. 5. A third putative hybrid in Euprotomus (Gastropoda, Caenogastropoda) with some additional remarks on hybridization in Euprotomus.

Abstract

A putative hybrid Euprotomus aratrum (Röding, 1798) x E. bulla (Röding, 1798) is reported upon. It is compared
with the two supposed parental species. Some additional remarks on putative Euprotomus hybrids are made.

Full text

JOURNAL OF THE NETHERLANDS MALACOLOGICAL SOCIETY
VOLUME 80 (1-3) | 8 OCTOBER 2016
Contents
BASTERA VOLUME 80 (1-3) | 8 OCTOBER 2016
Bank, R.A. 40 years of malacology in Amsterdam and Leiden: A Festschrift for Rob
Moolenbeek ..............................................................................................................................
Bank, R.A., Menkhorst, H.P.M.G. & Neubert, E. Descriptions of new and little-known
land snail taxa from Turkey, and establishment of a new genus (Gastropoda,
Pulmonata: Lauriidae, Enidae and Vitrinidae) ...................................................................
Dekkers, A.M. & Dekker, H. A new species of Turbo from the Red Sea (Gastropoda,
Turbinidae) ...............................................................................................................................
Winter, A.J. de, Leeuwen, S. van & Hovestadt, A. A new species of Glyphyalus
(Gastropoda, Pulmonata, Oxychilidae) from the Dutch Caribbean island of
St. Eustatius ..............................................................................................................................
Breure, A.S.H. Philippe Dautzenberg (1849–1935) and his time, towards the
reconstruction of an ancient science network .....................................................................
Janssen, A.W. & Goedert, J.L. Notes on the systematics, morphology and bio-stratigra-
phy of fossil holoplanktonic Mollusca, 24. First observation of a genuinely Late
Mesozoic thecosomatous pteropod ......................................................................................
Poorten, J.J. ter & Gemert, L.J. van. The genus Frigidocardium Habe, 1951 in the
Red Sea (Bivalvia, Cardiidae) ................................................................................................
Dekkers, A.M. A new tiny Conus species from the Philippines (Gastropoda, Conidae)................
Kronenberg, G.C. Revision of Euprotomus Gill, 1870. 5. A third putative hybrid in
Euprotomus (Gastropoda, Caenogastropoda) with some additional remarks on
hybridization in Euprotomus ..................................................................................................
Tenorio, M.J. Profundiconus robmoolenbeeki spec. nov.: A new deep water conoidean
gastropod from the Solomon Islands (Gastropoda, Conilithidae)...................................
Faber, M.J. & Gori, S. Infralittoral Rissoinidae (Gastropoda, Rissooidea) of Maldives
with the introduction of a new subfamily and one replacement name, the descrip-
tion of three new species, and a note on the identity of Rissoa rosea Deshayes, 1863....
Dijkstra, H.H. Annotations to the figured scallops (Mollusca, Bivalvia, Pectinidae) in
Gualtieri’s “Index Testarum Conchyliorum”, deposited in the Museo di Storia
Naturale e del Territorio at Calci (Pisa, Italy)......................................................................
Margry, C.J.P.J. Insulivitrina ingridae spec. nov., a fossil vitrinid from the Canary
Island of La Gomera (Gastropoda, Pulmonata)..................................................................
Gemert, L.J. van. Note on the publications of Edmond Saurin (1904-1977) on Recent Mol-
lusca; with a list of all his new pyramidellids (Gastropoda, Pyramidellidae) .....................
Maassen, W.J.M. Descriptions of two new carnivorous snail species from Thailand
(Pulmonata: Streptaxidae, Diapheridae)..............................................................................
Index to Volume 79 ........................................................................................................................
1
5
31
39
47
59
64
77
82
89
95
113
127
133
139
143
Robert Moolenbeek
at the
Zoological Museum Amsterdam
(2011)
Photograph by Ella Reitsma
Basteria_cover_80(1-3):Opmaak 1 9/15/2016 10:07 PM Page 1

Editorial board
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82
Basteria 80 (1-3): 82-88 (2016)
A putative hybrid Euprotomus aratrum (Röding, 1798)
xE. bulla (Röding, 1798) is reported upon. It is com-
pared with the two supposed parental species. Some
additional remarks on putative Euprotomus hybrids
are made.
Key words: Euprotomus aratrum,Euprotomus bulla, hybridization,
Indo-Pacific.
1For no. 4 in this series see: Kronenberg, G.C. & Harzhauser, M.,
2012. Europrotomus (Mollusca: Caenogastropoda: Strombidae): a
new Middle Miocene European strombid genus. – Paläontologis-
che Zeitschrift 86 (2): 147-159.
Introduction
Putative cases of hybridization in Indo-Pacific strom-
bids have been documented for a large number of
cases [for a preliminary overview see Kronenberg
(2008b) – but since then more cases have been re-
ported, for an overview see Liverani (2013: 5)]. So far,
three putative combinations of parental species form-
ing hybrids have been reported within the genus
Euprotomus Gill, 1870 (Kronenberg, 2008b). Two of
these combinations have been discussed, viz. E. vomer
(Röding, 1798) x E. bulla (Röding, 1798) (erroneously
described as Strombus hirasei Kuroda, 1942) by Kro-
nenberg (1999; 2002) and Dekkers (2012), and E. bulla x
E. chrysostomus Kuroda, 1942, by Liverani (2002). The
third combination, E. bulla xE. aratrum has been listed
by Kronenberg (2008b: Table 1) as a unique record.
This listing was based on a photograph made avail-
able by Jean-Pierre Barbier (Cebu, Philippines) of a
specimen collected at North Palawan (Coron Island),
but that specimen was not discussed in detail. Kronen-
berg (2008b) however, overlooked the specimen illus-
trated that same year in Kronenberg (2008a: pl. 230 fig.
6a, b); after re-examination of the image it turned out
that this specimen also appears to be a hybrid of
E. aratrum xE. bulla. A putative case of hybridization
involving a species of Euprotomus reported by Thach
(2007) has been dismissed by Kronenberg (2008b),
who, based on the illustration of the specimen (Thach,
2007: 18, fig. 39), concluded that the specimen was a
malformed E. aratrum (Röding, 1798).
In 2012 Dr. Nguyen Ngoc Thach (Nha Trang City,
Vietnam) made some images available of what at first
glance seemed to be a specimen of E. aratrum.
Dr. Thach however, already noted that there were
some remarkable differences with E. aratrum. And in-
deed, upon closer examination of the images, the spec-
imen could not be identified as that species with
certainty. Provisionally it was identified as E. aratrum
xE. bulla. Subsequently, Dr. Thach made the specimen
available for examination. The results are presented
below. I also take the opportunity to illustrate for the
first time the specimen mentioned in Table 1 by
Kronenberg (2008b).
Revision of Euprotomus Gill, 1870. 51.
A third putative hybrid in Euprotomus (Gastropoda,
Caenogastropoda) with some additional remarks
on hybridization in Euprotomus
Gijs C. Kronenberg
Associate, Naturalis Biodiversity Center, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands;
gijs.kronenberg@naturalis.nl
B2016-14-Kronenberg:Basteria-2015 9/7/2016 10:21 PM Page 82

Descriptive part
Class Gastropoda Cuvier, 1795
Subclass Caenogastropoda Cox, 1960
Order Littorinimorpha Golikov & Starobogatov, 1975
Superfamily Stromboidea Rafinesque, 1815
Family Strombidae Rafinesque, 1815
Euprotomus Gill, 1870
Euprotomus bulla xEuprotomus aratrum (Figs 1-10)
Euprotomus bulla (Röding, 1798) – Kronenberg 2008a: pl. 120 figs
6a-b.
Euprotomus bulla xE. aratrum – Kronenberg 2008b: 332 [Table 1].
Diagnosis. ― Adult shell with one digit at the adapi-
cal part of the outer lip, more or less parallel to the
shell’s axis, with columellar and parietal callus well
developed, slightly thicker at adbasal part of the last
whorl, apex and part of the spire covered with a dark
brown callus deposit.
Description. ― Adult shells with characters typi-
cal of species attributed to the genus Euprotomus Gill,
1870, measuring about 62-71 mm in length, with
about eight teleoconch whorls; the number of nuclear
whorls unknown. The first two to five teleoconch
whorls more or less extensively covered by the callus,
thereby obscuring the sculpture. First teleoconch
whorls with fine axial and spiral cords, resulting in a
reticulate sculpture. From the third whorl onwards
the axial cords gradually develop into ribs, overlaid
by the spiral sculpture. Simultaneously a subsutural
cord develops, broader than the other spiral cords.
On this cord an axially sculpture persists, but in other
spiral cords the axials gradually turn obsolete. From
the third or fourth whorl onwards a row of knobs on
the shoulder develops, gradually increasing in size.
Last whorl with an extensive callus on the adaper-
tural side, slightly thicker at the adbasal side, obscur-
ing about 45% of sculpture of the total of the last
whorl. On the last whorl there are about 11 knobs; on
the ventral side of the shell the knobs are covered by
the callus, more or less obscuring their presence.
Abapical of this row of knobs spiral cords are present,
with two cords being broader, bearing low knobs. A
third cord of intermediate broadness may be present
(Figs 6-8), but this third cord can also be absent (Figs
2-4). Suture of last whorl acutely curved adapically
and crossing the suture of the penultimate whorl.
Outer lip not broadly expanded, with a distinct,
deep, stromboid notch and a projection at the adapical
part, slightly diverging from the shell’s axis, reaching
beyond the apex. At early adult stage the rim of the
outer lip, abapically of the projection and adapically of
the stromboid notch, is slightly curved into adaper-
tural direction. When fully grown, the cavity thus
formed is filled with callus and subsequent thin layers
of callus cover the rim of the outer lip, resulting in a
subangled outer lip with a glaze. Anterior canal rather
long, curved addorsally, (almost) perpendicular to the
shell axis. Between the thickened outer lip and the pro-
jection, a low, very poorly developed secondary notch
is present. On the adapertural side of the outer lip, a
few poorly developed to very faint plicae are present
on the adapical part; at the point of attachment to the
spire there is a shallow canal.Adapertural side be-
tween strombid notch and anterior canal with a few
poorly developed plicae. Colour pattern very variable
[see illustrations herein and Kronenberg in Poppe
(2008a: pl. 120 fig. 6a-b) and discussion below].All
three specimens share a dark brown callus deposit on
the spire, the rim on the outer lip at the digit-like ex-
tension, and the part of the outer lip attachment to the
spire and near the stromboid notch.
Known records. ― So far only three specimens are
known, viz. (1) Philippines, near Palawan at about 3 m
depth; (2) Philippines, Olango Island at 10 m; and (3)
Vietnam, off Nha Trang, at unknown depth. These
specimens are kept in the collections of, respectively,
Jean-Pierre Barbier (Cebu) and Guido T. Poppe (Cebu),
and the author’s private collection (no. 6416). The latter
specimen will be deposited in Naturalis Biodiversity
Center (Leiden) at some point in the future.
Discussion
All three specimens appear to have a mixture of char-
acters of both E. bulla and E. aratrum, although they
differ remarkably from one another. A difference
with E. aratrum is the extensive callus deposit, reach-
ing the apex and covering a (large) part of the apex
and initial post nuclear whorls, which much more re-
sembles E. bulla.
The colour of the callus deposits on the ventral
side of the shells shows a great variability. The speci-
men in the Barbier collection is most reminiscent of E.
bulla. It is white except for (1) a patch with a light or-
ange hue near the adapical part of the columella, and
(2) the spire callus, which is dark brown (Figs 1-4).
The specimen in the Poppe collection is intermediate
in that character (see Kronenberg, 2008a: pl. 230 fig.
6a) and the specimen in the Kronenberg collection
(Fig. 5) is much closer to E. aratrum. Another differ-
ence with E. aratrum is the much paler colouration of
the extended columellar callus where the outer lip is
attached to the columella. This columellar callus is
whitish in the Kronenberg specimen, contrary to E.
aratrum where this patch is always dark, even in the
palest specimens. The Barbier and Poppe specimens
are less prominently coloured and do not show this
83
Kronenberg, G.C. – A third putative hybrid in Euprotomus
B2016-14-Kronenberg:Basteria-2015 9/7/2016 10:21 PM Page 83

character clearly. The colour of the aperture is much
more intense orange as is usually the case in E. ara-
trum, which usually is (much) paler. Only exception-
ally the colour of specimens of E. aratrum may reach
the same intensity. Also the sculpture and colour pat-
tern of the dorsal side of the specimen differ slightly
from that in E. aratrum.
As far as the colour pattern of the shell is con-
cerned, the pattern of the shell in the collection Kro-
nenberg (Figs. 5-8) is most reminiscent of E. aratrum,
with the irregular of light brown speckles and mot-
tling on a cream background. The specimen in the
Barbier collection is much more like E. bulla, especially
on the abapertural part of the outer lip, which has
some small white spots on a brownish background.
The specimen in the Poppe collection is much paler,
near white with a few brown blotches (Kronenberg,
2008a: pl. 230 fig. 6b). The dark brown blotch on the
centre of the dorsal side of that specimen has never
been encountered in E. aratrum to my knowledge, but
rarely occurs in E. bulla, see e.g. Kronenberg (2008a, pl.
230 fig. 2).
The main differences between the putative hybrids
and their probable parental species are summarized in
Table 1.
Unfortunately, like all other cases of presumed hy-
bridization in Indo-Pacific Strombidae (Kronenberg,
2008b), there is no definite proof of it. The animal of a
specimen that was considered to represent a putative
case of hybridization in a specimen of Lambis compa-
rable to specimens figured in De Turck et al. (1999: pl.
30 fig. 3) and Kronenberg (2008a: pl. 210 figs 1a, 1b)
has become available for examination. Preliminary re-
sults of the dissection of that specimen showed that its
anatomy differs from either of the putative parent
species and, surprisingly, not being intermediate (Si-
mone, unpublished data).
Some additional remarks on
Euprotomus
related to putative hybridization
Euprotomus bulla xE. aurisdianae
In his description of E. aurora, Kronenberg (2002b) al-
ready noticed that conchological characters of this
species are more or less intermediate between E. bulla
and E. aurisdianae. However, the possibility that E. au-
rora is a hybrid was rejected as none of the supposed
parental species lives in the (Western) Indian Ocean.
After Kronenberg’s (2002b) description, who
84
Basteria 80(1-3), 2016
Character Euprotomus aratrum E. aratrum xE. bulla Euprotomus bulla
Shell length 58 – 91 mm 62 – 71 mm 50 – 73 mm
Callus on spire
Only small part of the spire cov-
ered, near aachment of outer
lip, apex and first four to five
teleoconch whorls not covered
Dark chocolate brown, reaching
and partly covering apex
Reaching and partly covering
apex
Colour of callus Cream via orange to dark choco-
late brown, sometimes near black
Cream to dark chocolate brown,
with a paler, patch near aach-
ment of outer
White
Colour of aperture Cream via pale orange to bright
orange Bright orange Reddish
Colour rim of outer lip near digit
and strombid notch Dark brown to blackish Dark brown to blackish White
Dorsal sculpture below shoulder
knobs
Coarse spiral sculpture of close
set spiral cords, two or rarely
three cords broader, usually
bearing distinct knobs
Coarse spiral sculpture of close
set spiral cords, two cords
broader, usually bearing indis-
tinct knobs
Smooth to very low close set spi-
ral cords of nearly equal size
Plicae adapertural side outer lip Present, strong to moderately
strong
Present, moderately strong to
weak Absent
Table 1. Comparison of some of the shell characters of Euprotomus aratrum,E. bulla and the putative hybrid E. aratrum xE. bulla. Shell
measurements of E. aratrum and E. bulla round after Abbott (1960).
B2016-14-Kronenberg:Basteria-2015 9/7/2016 10:21 PM Page 84

85
Kronenberg, G.C. – A third putative hybrid in Euprotomus
Figs 1-4. Euprotomus aratrum XE. bulla: Philippines, North Palawan, Coron Island, by compressor diver, at about 3 meters in sand & sea-
weed bottom. Actual size: 62.2 mm in length. Coll. Jean-Pierre Barbier, unnumbered. Photographs by Jean-Pierre Barbier.
B2016-14-Kronenberg:Basteria-2015 9/7/2016 10:21 PM Page 85

86
Basteria 80(1-3), 2016
Figs 5-10. Euprotomus aratrum XE. bulla: Vietnam, off Nha Trang at unknown depth, by local fishermen. Actual size: 70.3 mm in length.
Coll. G.C. Kronenberg, 6416. Photographs by Eelco Kruidenier, Naturalis Biodiversity Center.
B2016-14-Kronenberg:Basteria-2015 9/7/2016 10:21 PM Page 86

excluded the Pacific from the distribution of E. aurora,
some records from the Pacific of E. aurora were pub-
lished (Monsecour, 2004; Vermeij, 2005 [as E. bulla,
but subsequently considered as E. aurora, Vermeij
pers. comm. to present author]; Kronenberg &
Dharma, 2005; Kronenberg, 2008a).
Recently E. aurora has also been reported from
American Samoa (Brown, 2011: 248). There was neither
description nor illustration. This record needs confir-
mation, as it is also possible that this concerns a rela-
tively heavy sculptured form of E. bulla. The possibility
of this specimen being a hybrid of E. aurisdianae xE.
bulla is provisionally excluded, because to my knowl-
edge E. aurisdianae does not live further east than the
Solomon Islands, where Abbott (1960: 127, pl. 103)
recorded it from Bougainville Island. Note however
that Abbott confused E. aurisdianae with E. aurora.Eu-
protomus bulla is known from Samoa; Abbott (1960:
129-130, pl. 105) recorded it from Tutuila Island.
More recently, Mr. David Monsecour (Aarschot,
Belgium) sent me images of a specimen of Euprotomus
sp. from the Solomon Islands. This appears to be a
genuine specimen of E. aurora. A discussion of this
specimen and a discussion on further records of E.
aurora from Pacific localities will be published else-
where (Monsecour & Delsaerdt, in prep.).
So far, no convincing hybrid E. aurisdianae xE.
bulla is known to me.
Euprotomus bulla xE. chrysostomus
In the original report on this presumed hybrid (Liv-
erani, 2002) two specimens, both from Sogod (Cebu,
Philippines) were illustrated (p. 59, figs 3-4), measur-
ing 74 and 66 mm, respectively. Liverani’s fig. 3 was
subsequently re-illustrated in Liverani, 2013 (pl. 162
fig. 2a, b). Although the same specimen is pho-
tographed, the colour of the dorsum in the images
(Liverani, 2002: fig. 3 right; Liverani 2013: fig. 2a) dif-
fers remarkably. On Liverani’s 2002 fig. 3 the dorsum
of the specimen appears to have a greyish pattern on
a white background, while in Liverani’s 2013 fig. 2a
the pattern is brown on a white background.
Both specimens are rather close to E. chrysostomus
(Kuroda, 1942), especially the specimen re-illustrated
in Liverani 2013. Both specimens illustrated by Liv-
erani have a callus deposit on the spire that reaches
further to the apex than in E. chrysostomus, but do not
reach the apex. In the text on Euprotomus, Liverani
(2002) stated that the interior of the outer lip is a rich
orange-red, being paler near the abapertural border.
Liverani further noted “A pair of light [sic] lirae are
present in the posterior canal.” These are not visible
on the accompanying images, so they must indeed be
very faint.
On http://www.stromboidea.de/?n=Species.Euproto-
musBullaXChrysostomus the specimens from the
Liverani collection are illustrated again, but the meas-
urements are indicated as 73 and 74.5 mm, contrary
to the caption on Liverani’s 2002 plate. The correct
measurements are 73,5 and 65,5 mm (pers. comm. V.
Liverani). On this website another putative E. bulla x
E. chrysostomus specimen from Panglao (Bohol,
Philippines) from the Börnke collection is illustrated.
It measures 65 mm and has the callus on the spire
more extended towards the apex –but not reaching it–
as in the Liverani specimens, but there are distinct
plicae in the adapical part of the adapertural side of
the outer lip. Another specimen (not illustrated) is
present in the Kronenberg collection. It was collected
at Bohol. In this specimen, the callus does reach the
apex, but plicae in the adapical part of the adaper-
tural side of the outer lip are not noticeable.
Euprotomus bulla xE. vomer
Bozzetti & Sargent (2011) described a new species in
Euprotomus, viz. E. kiwi. In their paper they also re-
ferred to E. hirasei (= E. vomer xE. bulla) as being “…
the smooth backed form that was described as Euproto-
mus hirasei Kuroda, 1942 [sic!; in the original descrip-
tion by Kuroda it is named Strombus hirasei]” (Bozzetti
& Sargent, 2011: 24). This observation, however, is not
correct; the holotype of S. hirasei has a distinct sculp-
ture of spiral cords on the dorsal side; see the descrip-
tion and figures of the holotype by Kuroda (1942: 8,
figs 3-4) and Kronenberg (1999: figs 2, 5, 8) and the
paratype by Habe (1964 and 1965: pl. 16 fig.1). Note
that the paratype illustrated by Habe is not indicated
as such in Habe’s works, but this information was ob-
tained from Dr. Kazunori Hasegawa and Dr. Hiroshi
Saito, both National Science Museum, Tokyo, Japan.
Kronenberg (2002a) subsequently reported on a
third specimen from Fiji and both Dekkers (2012) and
Liverani (2013: pl. 160 figs 3a-b) recorded a specimen
from New Caledonia. Dekkers (2012) discussed his
specimen, collected at Balabio Isl., (20°06’ S, 164°11’ E)
at 15-30 m, extensively and noted that there was some
variability in the shell characteristics of the putative
hybrid specimens known so far: the Fiji specimen
being rather close to E. vomer, the Japanese specimens
intermediate, and the Balabio Isl. specimen closer to E.
bulla. This agrees with the observations made by Kro-
nenberg on the putative Lambis hybrid, generally re-
ferred to as L. arachnoides (see Kronenberg, 1993).
In conclusion, there are at present three putative com-
binations of hybrid specimens known within Euproto-
mus, viz. E. bulla xE. aratrum (this paper); E. bulla xE.
chrysostomus (Liverani, 2002; Wieneke et al., 2016);
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88
Basteria 80(1-3), 2016
and E. bulla xE. vomer (Kronenberg, 1999; 2002a;
Dekkers, 2012; Liverani, 2013). It is remarkable that in
all these instances one of the supposed parent species
is E. bulla.
Acknowledgements
I would like to thank Dr. Nguyên Ngoc Thach (Nha
Trangh City, Vietnam) for making this specimen avail-
able for study. I also thank Jean-Pierre Barbier (Cebu,
Mactan, Philippines) for information on the hybrid
specimen in his collection. David Monsecour
(Aarschot, Belgium) kindly sent me images of the
Solomon specimen of E. aurora. Eelco Kruidenier (Nat-
uralis Biodiversity Center, Leiden) made the photo-
graphs. Ulrich Wieneke, Murnau, Germany read the
draft and made valuable suggestions for improvement.
This paper is dedicated to Mr. Robert G. Moolen-
beek on the occasion of his retirement from active
duty. Robert was the first to put forward to me that
Lambis wheelwrighti Greene, 1978, could be a junior
synonym of L. arachnoides Shikama, 1971, and the pos-
sibility of this being a hybrid (see Kronenberg, 1993).
References
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Indo Pacific Mollusca 1 (2): 33-146.
Bozzetti, L. & Sargent, D.M., 2011. A new species of Euprotomus
from New Zealand and the Kermadec Islands. – Visaya 3
(3): 23-27.
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