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Rare, deep-water and similar: revision of Sibogasyrinx
(Conoidea: Cochlespiridae)
Yuri Kantor, Nicolas Puillandre
To cite this version:
Yuri Kantor, Nicolas Puillandre. Rare, deep-water and similar: revision of Sibogasyrinx (Conoidea:
Cochlespiridae). European Journal of Taxonomy, Consortium of European Natural History Museums,
2021, �10.5852/ejt.2021.773.1509�. �hal-03360999�
19
European Journal of Taxonomy 773: 19–60 ISSN 2118-9773
https://doi.org/10.5852/ejt.2021.773.1509 www.europeanjournaloftaxonomy.eu
2021 · Kantor Yu.I. & Puillandre N.
This work is licensed under a Creative Commons Attribution License (CC BY 4.0).
Research article
urn:lsid:zoobank.org:pub:2A159A89-64D1-4352-8E0B-FA546983C99A
Rare, deep-water and similar: revision of Sibogasyrinx
(Conoidea: Cochlespiridae)
Yuri I. KANTOR 1,* & Nicolas PUILLANDRE 2
1 A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences,
Leninski prospect 33, 119071 Moscow, Russian Federation.
1,2 Institut Systématique Evolution Biodiversité (ISYEB), Muséum national d’histoire naturelle, CNRS,
Sorbonne Université, EPHE, Université des Antilles, 57 rue Cuvier, 75005 Paris, France.
* Corresponding author: kantor.yuri1956@gmail.com
2 Email: puillandre@mnhn.fr
1 urn:lsid:zoobank.org:author:48F89A50-4CAC-4143-9D8B-73BA82735EC9
2 urn:lsid:zoobank.org:author:00565F2A-C170-48A1-AAD9-16559C536E4F
Abstract. The genus Sibogasyrinx has to date included only four species of rare deep-water Conoidea,
each known from few specimens. In shell characters it strongly resembles three distantly-related
genera, two of which, Comitas and Leucosyrinx
A molecular phylogenetic analysis of a large amount of material of Conoidea has revealed the existence of
much additional undescribed diversity within Sibogasyrinx
cox1 gene and morphological characters
of 54 specimens, 10 species hypotheses are proposed, of which six are described as new species:
S. subula sp. nov., S. lolae sp. nov., S. maximei sp. nov., S. clausura sp. nov., S. pagodiformis sp. nov.
and S. elbakyanae Kantor, Puillandre & Bouchet sp. nov. One of the previously described species was
absent in our material. Most of the new species are very similar and are compared to Leucosyrinx spp.
Species of Sibogasyrinx are unique among Conoidea on account of the high intrageneric variability
in radular morphology. Three distinct radula types are found within Sibogasyrinx, two of which are
.
Keywords. Sibogasyrinx pyramidalis species complex, deep-water Conoidea, cox-1, integrative
taxonomy, new species.
Kantor Yu.I. & Puillandre N. 2021. Rare, deep-water and similar: revision of Sibogasyrinx (Conoidea: Cochle-
spiridae). European Journal of Taxonomy 773: 19–60. https://doi.org/10.5852/ejt.2021.773.1509
Introduction
The superfamily Conoidea is well known for high levels of homoplasy in shell form. Molecular studies
have revealed that very similar or nearly indistinguishable shell morphologies are found in distantly
related lineages (Puillandre et al. 2011; Abdelkrim et al. 2018), thus rendering the attribution of species
to genus and even family questionable, when based on shell characters alone.
European Journal of Taxonomy 773: 19–60 (2021)
20
Particularly striking examples of such homoplastic shells are found in the genera Comitas Finlay, 1926,
Leucosyrinx Dall, 1889, Sibogasyrinx Powell, 1969 and, to some extent, Comispira Kantor, Fedosov &
Puillandre, 2018, all of which possess rather similar fusiform shells with a long siphonal canal and axial
ribs or nodules on a more or less angular shoulder. Sibogasyrinx was originally established as a subgenus
of Leucosyrinx, at that time assigned to the Cochlespiridae (Powell 1966; Taylor et al. 1993). However,
molecular data demonstrate that Leucosyrinx constitutes a sister group to the Pseudomelatomidae, albeit
with low support, while Sibogasyrinx is clearly referable to the Cochlespiridae (Puillandre et al. 2011;
Abdelkrim et al. 2018). Furthermore, species of Sibogasyrinx are characterized by a radula typical of
Leucosyrinx spp.
which possess only marginal teeth (Bouchet et al. 2011). Both radula type and phylogenetic analyses
(Bouchet et al. 2011; Puillandre et al. 2011; Abdelkrim et alComitas in the
Pseudomelatomidae. Finally, the recently described Comispira is clearly referable to the Cochlespiridae,
both on anatomical and molecular grounds.
The taxonomy and morphology of these genera were treated in detail by Kantor et al. (2018); however,
although they can be reliably distinguished by the morphology of the radula, most of the species
have never been examined in this respect. Consequently, the generic placement of the anatomically
unknown species relies in most cases on shell characters only, even though generic diagnoses based
on conchological features are still lacking. For example, Sibogasyrinx originally included two species
with “peripheral angle right down at the lower suture” (Powell 1969: 343). Later it was found that this
character is inconsistent and in Sibogasyrinx sangeri Kantor et al., 2018 the angulation may be situated
at mid-whorl (Kantor et al. 2018). Finally, many species are known from a very limited number of
A consequence of this taxonomic imprecision is that the generic placement of species is often doubtful.
Most species are currently attributed to the pseudomelatomid genera Leucosyrinx and Comitas, with
26 and 53 valid Recent species, respectively (MolluscaBase), while the cochlespirid genera Comispira
and Sibogasyrinx only include two and three species, respectively. Furthermore, while DNA sequences
of only seven species of Leucosyrinx (four undescribed), three of Comitas (two undescribed), two of
Comispira and two of Sibogasyrinx
in the Muséum national d’histoire naturelle (MNHN) revealed the existence of numerous undescribed
species of Sibogasyrinx, including several that cluster with the type species, S. pyramidalis (Schepman,
Sibogasyrinx and a taxonomic
revision of the genus based on molecular and morphological data.
Material and methods
Sampling
Samples were collected during several expeditions of the MNHN (expeditions.mnhn.fr): DongSha 2014
and NanHai 2014 in the South China Sea, AURORA 2007 and PANGLAO 2005 in the Philippines,
SALOMON 2 and SALOMONBOA 3 in the Solomon Islands, BIOPAPUA, KAVIENG 2014, MADE
EP and PAPUA NIUGINI in Papua New Guinea, BOA1 and SANTO 2006 in Vanuatu, EXBODI,
Madagascar and GUYANE 2014 in French Guiana. Full details on the stations of these expeditions,
including dates, are available at the above-mentioned website. One additional sample was collected
in New South Wales, Australia, during the IN2017_V03 expedition on R/V Investigator. Specimens
processed before 2012 were anaesthetized using an isotonic solution of MgCl2
et al.
2014). The samples are recorded in the BOLD database and the sequences have been deposited in
GenBank (Table 1).
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
21
Table 1 (continued on next three pages). List of examined material included in molecular phylogenetic analysis. ABGD PSH = Automatic Barcode Gap
Discovery, primary species hypothesis; BOLD = Barcode of Life Datasystem; GB = Genbank. New BOLD and GB sequences are highlighted in red.
Reg. no. Expedition Stn Coordinates Depth (m) Family Genus ABGD PSHs species BOLD ID GB ID
MNHN-IM-
2007-42517 SALOMON 2 CP2276
814–980 Cochlespiridae Sibogasyrinx 3subula CONO792-08 MZ379777
MNHN-IM-
2007-42530 SALOMON 2 CP2187
482–604 Cochlespiridae Sibogasyrinx 3subula CONO806-08 MZ364512
MNHN-IM-
2007-42533 SALOMON 2 CP2194
440–521 Cochlespiridae Sibogasyrinx 3subula CONO809-08 MZ364501
MNHN-IM-
2009-13567 SALOMON 2 CP2184
464–523 Cochlespiridae Sibogasyrinx 3subula CONO3220-21 MZ364503
MNHN-IM-
2009-13568 SALOMON 2 CP2184
464–523 Cochlespiridae Sibogasyrinx 3subula CONO3214-21 MZ364481
MNHN-IM-
2009-17048 BIOPAPUA CP3750
654–660 Cochlespiridae Sibogasyrinx 3subula CONO3212-21 MZ364497
MNHN-IM-
2013-58409 KAVIENG 2014 CP4448
564–743 Cochlespiridae Sibogasyrinx 3subula CONO3229-21 MZ364494
MNHN-IM-
2007-17701 BOA1 CP2432
630–705 Cochlespiridae Sibogasyrinx 3subula CONO153-08 EU015646.1
MNHN-IM-
2007-17702 BOA1 CP2432
630–705 Cochlespiridae Sibogasyrinx 3subula CONO154-08 EU015647.1
MNHN-IM-
2007-17703 BOA1 CP2432
630–705 Cochlespiridae Sibogasyrinx 3subula CONO155-08 MN322452.1
MNHN-IM-
2007-17704 BOA1 CP2432
630–705 Cochlespiridae Sibogasyrinx 3subula CONO156-08 MZ364507
MNHN-IM-
2009-16972 BIOPAPUA CP3689
679–685 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3208-21 MZ364518
MNHN-IM-
2009-16973 BIOPAPUA CP3689
679–685 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3211-21 MZ364511
MNHN-IM-
2009-16988 BIOPAPUA CP3671
585–601 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3209-21 MZ364492
MNHN-IM-
2009-17000 BIOPAPUA CP3672
702–724 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3210-21 MZ364486
MNHN-IM-
2013-19716 PAPUA NIUGINI CP3981
688 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3224-21 MZ364517
MNHN-IM-
2013-19745 PAPUA NIUGINI CP3982
724 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3227-21 MZ364499
European Journal of Taxonomy 773: 19–60 (2021)
22
Table 1 (continued). List of examined material included in molecular phylogenetic analysis.
Reg. no. Expedition Stn Coordinates Depth (m) Family Genus ABGD PSHs species BOLD ID GB ID
MNHN-IM-
2013-19768 PAPUA NIUGINI DW3983
470–508 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3226-21 MZ364519
MNHN-IM-
2013-58845 KAVIENG 2014 CP4483
827–966 Cochlespiridae Sibogasyrinx 1cf. pyramidalis 2 CONO3231-21 MZ364516
MNHN-IM-
2009-13451 AURORA 2007 CP2729
593–600 Cochlespiridae Sibogasyrinx 2cf. pyramidalis 1 CONO2061-18 MG968492.1
MNHN-IM-
2013-44574 NanHai 2014 CP4118
700–723 Cochlespiridae Sibogasyrinx 2cf. pyramidalis 1 CONO2095-18 MG968491.1
MNHN-IM-
2013-44605 NanHai 2014 CP4118
700–723 Cochlespiridae Sibogasyrinx 2cf. pyramidalis 1 CONO2096-18 MG968493.1
MNHN-IM-
2013-50215 DongSha 2014 CP4129
590–633 Cochlespiridae Sibogasyrinx 2cf. pyramidalis 1 CONO2097-18 MG968494.1
MNHN-IM-
2009-11327 SANTO 2006 AT139
1262–1285 Cochlespiridae Sibogasyrinx 5pagodiformis CONO3206-21 MZ364502
MNHN-IM-
2009-16825 SALOMONBOA 3 CP2789
1250–1402 Cochlespiridae Sibogasyrinx 5pagodiformis CONO3221-21 MZ364496
MNHN-IM-
2009-16834 SALOMONBOA 3 CP2817
1136–1750 Cochlespiridae Sibogasyrinx 4elbakyanae CONO3219-21 MZ364498
MNHN-IM-
2013-19752 PAPUA NIUGINI CP3982
724 Cochlespiridae Sibogasyrinx 7sangeri CONO2080-18 MG968497.1
MNHN-IM-
2013-52052 BIOPAPUA CP3671
585–601 Cochlespiridae Sibogasyrinx 7sangeri CONO2088-18 MG968498.1
MNHN-IM-
2009-17021 BIOPAPUA CP3750
654–660 Cochlespiridae Sibogasyrinx 7sangeri CONO2058-18 MG968502.1
MNHN-IM-
2009-16989 BIOPAPUA CP3671
585–601 Cochlespiridae Sibogasyrinx 7sangeri CONO2079-18 MG968501.1
MNHN-IM-
2009-16766 SALOMON 2 CP2181
645–840 Cochlespiridae Sibogasyrinx 7sangeri CONO2062-18 MG968505.1
MNHN-IM-
2007-42523 SALOMON 2 CP2288
509–520 Cochlespiridae Sibogasyrinx 7sangeri CONO799-08 MG968466.1
MNHN-IM-
2009-16995 BIOPAPUA CP3674
788–805 Cochlespiridae Sibogasyrinx 7sangeri CONO2057-18 MG968495.1
MNHN-IM-
2009-17022 BIOPAPUA CP3729
575–655 Cochlespiridae Sibogasyrinx 7sangeri CONO2059-18 MG968496.1
MNHN-IM-
2009-17057 BIOPAPUA CP3742
448–470 Cochlespiridae Sibogasyrinx 7sangeri CONO2060-18 MG968499.1
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
23
Table 1 (continued). List of examined material included in molecular phylogenetic analysis.
Reg. no. Expedition Stn Coordinates Depth (m) Family Genus ABGD PSHs species BOLD ID GB ID
MNHN-IM-
2013-19961 PAPUA NIUGINI CP4014
630–870 Cochlespiridae Sibogasyrinx 7sangeri CONO2082-18 MG968504.1
MNHN-IM-
2009-13434 AURORA 2007 CP2663
562 Cochlespiridae Sibogasyrinx 7sangeri CONO2064-18 MG968500.1
MNHN-IM-
2009-16779 SALOMON 2 CP2288
509–520 Cochlespiridae Sibogasyrinx 7sangeri CONO2063-18 MG968503.1
MNHN-IM-
2007-42498 SALOMON 2 CP2227
508–522 Cochlespiridae Sibogasyrinx 6losa CONO772-08 MZ364480
MNHN-IM-
2013-59044 KAVIENG 2014 DW4500
150–170 Cochlespiridae Sibogasyrinx 6losa CONO3230-21 MZ364489
MNHN-IM-
2009-16831 SALOMONBOA 3 CP2767
416–425 Cochlespiridae Sibogasyrinx 6losa CONO3218-21 MZ364482
MNHN-IM-
2009-29230 EXBODI CP3864
460–708 Cochlespiridae Sibogasyrinx 8lolae CONO3222-21 MZ364514
MNHN-IM-
2009-29311 TERRASSES CP3098
511–1050 Cochlespiridae Sibogasyrinx 8lolae CONO3223-21 MZ364500
MNHN-IM-
2013-48156 KANADEEP CP4923
787 Cochlespiridae Sibogasyrinx 8lolae CONO3233-21 MZ364491
MNHN-IM-
2007-42537 SALOMONBOA 3 CP2839
506–567 Cochlespiridae Sibogasyrinx 8lolae CONO820-08 MZ364485
MNHN-IM-
2013-45883 MADEEP DW4323
720 Cochlespiridae Sibogasyrinx 9maximei CONO3228-21 MZ364490
MNHN-IM-
2009-16763 SALOMON 2 CP2182
762–1060 Cochlespiridae Sibogasyrinx 10 clausura CONO3215-21 MZ364484
MNHN-IM-
2013-48167 KANADEEP CP4966
975–980 Cochlespiridae Sibogasyrinx 10 clausura CONO3234-21 MZ364509
MNHN-IM-
2013-48169 KANADEEP CP4965
961 Cochlespiridae Sibogasyrinx 10 clausura CONO3235-21 MZ364487
MNHN-IM-
2013-48256 KANADEEP CP4964
963–970 Cochlespiridae Sibogasyrinx 10 clausura CONO3236-21 MZ364510
MNHN-IM-
2013-48258 KANADEEP CP4965
961 Cochlespiridae Sibogasyrinx 10 clausura CONO3237-21 MZ364504
MNHN-IM-
2013-48244 KANADEEP CP4963
978–1000 Cochlespiridae Sibogasyrinx 10 clausura CONO3238-21 MZ364495
MNHN-IM-
2013-48144 KANADEEP CP4965
961 Cochlespiridae Sibogasyrinx 10 clausura CONO3232-21 MZ364515
European Journal of Taxonomy 773: 19–60 (2021)
24
Table 1 (continued). List of examined material included in molecular phylogenetic analysis.
Reg. no. Expedition Stn Coordinates Depth (m) Family Genus ABGD PSHs species BOLD ID GB ID
AMS
C.519344 IN2017_V03 –
999–1013 Cochlespiridae Sibogasyrinx 10 clausura –MZ345688
MNHN-IM-
2013-19689 PAPUA NIUGINI CP3979
540–580 Pseudomelatomidae Leucosyrinx – G CONO3225-21 MZ364479
MNHN-IM-
2009-17089 BIOPAPUA DW3748 5°37’ S;
398–399 Pseudomelatomidae Leucosyrinx – G CONO3213-21 MZ364505
MNHN-IM-
2007-17846 EBISCO CP2600
603–630 Pseudomelatomidae Leucosyrinx – F CONO189-08 EU015655.1
MNHN-IM-
2009-16897 MIRIKY CP3250
46°0 493–662 Pseudomelatomidae Leucosyrinx – E CONO3207-21 MZ364493
MNHN-IM-
2007-42503 SALOMON 2 CP2219
650–836 Pseudomelatomidae Leucosyrinx – C CONO777-08 MZ364513
MNHN-IM-
2009-16769 SALOMON 2 CP2217
1045–1118 Pseudomelatomidae Leucosyrinx – D CONO3217-21 MZ364508
MNHN-IM-
2007-42445 PANGLAO 2005 CP2360
357–372 Pseudomelatomidae Leucosyrinx – A CONO716-08 MZ364488
MNHN-IM-
2009-16764 SALOMON 2 CP2182
762–1060 Pseudomelatomidae Leucosyrinx – B CONO3216-21 MZ364483
MNHN-IM-
2013-56288 GUYANE 2014 CP4367
351–354 Pseudomelatomidae Leucosyrinx –verrillii CONO2101-18 MG968474.1
MNHN-IM-
2013-56840 GUYANE 2015 CP4407
495–502 Pseudomelatomidae Leucosyrinx –verrillii CONO2109-18 MG968470.1
MNHN-IM-
2009-8188 CEAMARC 87EV524
397–411 Cochlespiridae Aforia – sp. CONO1120-10 KT448835.1
MNHN-IM-
2007-17919 PANGLAO 2005 CP2340
271–318 Cochlespiridae Cochlespira – sp. CONO294-08 EU015719.1
MNHN-IM-
2013-59407 ZhongSha 2015 CP4157
1205–1389 Cochlespiridae Comispira – sp. CONO2119-18 MG968463.1
MNHN-IM-
2007-42554 SANTO 2006 FR11 15°3652S,
167°1029E6–33 Pseudomelatomidae Crassispira – sp. CONO871-08 KX051344.1
MNHN-IM-
2007-17916 PANGLAO 2005 CP2385
982–989 Pseudomelatomidae Comitas – sp. CONO321-08 EU015731.1
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
25
DNA sequencing
DNA was extracted using the Epmotion 5075 robot (Eppendorf), following the manufacturer’s
recommendations. The barcode fragment (658 bp) of the mitochondrial cox1
the universal primers LCO1490/HCO2198 (Folmer et al. 1994). Polymerase chain reactions (PCRs)
were performed using a previously well-established protocol (Puillandre et al. 2017). The PCR products
Species delimitation
The cox1 sequences were aligned manually (no gaps were inferred). The Automatic Barcode Gap Discovery
(ABGD; Puillandre et al. 2012a) and Assemble Species by Automatic Partitioning (ASAP; Puillandre
et al. 2021) methods were used to propose primary species hypotheses (PSH), using the web versions
(https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb.html and https://bioinfo.mnhn.fr/abi/public/asap/)
with default parameters. Both methods use genetic distances only. ABDG automatically detects the gap
species hypotheses. ASAP screens all the genetic distances from the lowest to the highest and merges
sequences into ‘groups’ that are successively further merged until all sequences form a single group.
At each merging step, the partition is evaluated and given a score. At the end of the analyses, the 10
partitions with the lowest scores are provided (the lower the ASAP-score, the better the partition).
The robustness of the PSHs proposed by ABGD and ASAP was then evaluated by checking whether they
correspond to highly supported clades, whether they are conchologically and anatomically diagnosable,
and whether they are geographically and/or bathymetrically isolated, and in so doing they were converted
into secondary species hypotheses (SSH), following the methodology described in Puillandre et al.
(2012b).
Phylogenetic analyses
Phylogenetic trees were reconstructed using the cox1 alignment containing the Sibogasyrinx sequences,
plus Leucosyrinx, Comitas and Comispira sequences for comparison, and outgroups from the respective
families of the four genera (Cochlespiridae and Pseudomelatomidae) (Table 1). Maximum Likelihood
(ML) and Bayesian Approach (BA) were used to infer phylogenetic trees. ML was performed using the
IQ-Tree webserver (http://iqtree.cibiv.univie.ac.at/ et al. 2016), and robustness of the
tree was estimated using ultrafast bootstrapping (1000 replicates) (Minh et al. 2013). BA was performed
using MrBayes 3.2. (Ronquist et al. 2012), as implemented on the Cipres Science Gateway (MrBayes ver.
3.2.2 on XSEDE; http://www.phylo.org/portal2), with two parallel analyses consisting each of 8 Markov
chains for 30 000 000 generations with 5 swaps at each generation, a sampling frequency of one tree each
10 000 generations, a chain temperature of 0.02 and the parameters of the substitution model estimated
during the analysis. Convergence of each analysis was evaluated using Tracer ver. 1.4.1 (Rambaut
et al. 2014) by checking that ESS values all exceeded 200, and a consensus tree was reconstructed after
for each codon position was estimated during the analysis. For the BA, a substitution model with six
substitution categories, a gamma-distributed rate variation across sites approximated in four discrete
categories and a proportion of invariable sites was used for each codon position independently.
Morphological studies
Protoconchs were measured in standard position and the number of whorls counted according to
Bouchet & Kantor (2004). Foregut anatomy was examined by dissections. Radulae were prepared by
standard methods (Kantor & Puillandre 2012) and examined by scanning electron microscopy (TeScan
TS5130MM) at the Institute of Ecology and Evolution of the Russian Academy of Sciences (IEE RAS).
European Journal of Taxonomy 773: 19–60 (2021)
26
Abbreviations
AL = aperture length
IEE RAS = A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences,
Moscow, Russian Federation
I. = island
Is. = islands
lv = live collected specimen
MNHN = Muséum national d’histoire naturelle, Paris, France
PSH = primary species hypothesis
SL = shell length
SSH = secondary species hypothesis
stn = station
SW = shell width
USNM = National Museum of Natural History, Smithsonian Institution, Washington DC, USA
ZMA = Naturalis Biodiversity Center, Leiden, the Netherlands
Results
Phylogenetic analysis
ABGD and ASAP partitions
ABGD consistently returned a partition with 10 PSHs, numbered from PSH 1 to PSH 10 (Fig. 1). The best
ASAP partition includes 9 PSHs, with the ABGD PSHs 4 and 5 grouped into a single PSH. The second
best ASAP partition includes 10 PSHs, identical to the ABGD PSHs. The third best ASAP partition
includes 11 PSHs, splitting the ABGD PSH 2 into two PSHs, one including the single specimen from
the Philippines, the other including the three specimens from Taiwan. The remaining ASAP partitions
have a much higher score and will not be discussed further.
From PSHs to SSHs
All the ABGD PSHs are morphologically distinct, although PSHs 1 and 2 exhibit considerable similarity
(see below – Taxonomy). All the studied Sibogasyrinx have a protoconch suggestive of non-planktotrophic
development, thus suggesting limited dispersal abilities. All the ABGD PSHs including more than one
sample correspond to supported (Posterior Probabilities PP > 0.95 and Bootstraps BS > 90) or highly
supported (PP > 0.98 and BS > 95) clades. Three main lineages can be recognized, all highly supported.
is from Papua New Guinea, the latter from Taiwan and the Philippines. They are thus allopatric, but they
are not sister taxa, PSH 2 being sister to the morphologically distinct PSH 3 from Papua New Guinea
(together with the PSH 1, but not from the same station), but also occurring in the Solomon Islands and
Vanuatu. These three PSHs have overlapping bathymetric distributions, between 500 and 1000 m deep
(Fig. 2). Because of their cox-1
most closely related PSHs, we convert these 3 PSHs into 3 SSHs. The split of PSH 2 into two PSHs in
the third ASAP partition is interpreted as resulting from genetic distances between allopatric populations
within a single species, potentially linked to the limited dispersal abilities of the species.
The second main lineage includes PSH 4 and 5, represented by only one and two samples, respectively.
both are found in the Solomon Islands (PSH 5 also occurring in Vanuatu) (Fig. 2). They are easily
distinguished morphologically, and we convert these two PSHs into 2 SSHs, thus contradicting the
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
27
all these PSHs are found in at least two localities, and all of them have at least one representative from
the Solomon Islands, but never from the same station (although sometimes from geographically very
close stations). Three PSHs (6, 7 and 9) are found in Papua New Guinea, and again never at the same
found in relatively shallow water (between 150 and 500 m), whereas PSHs 7, 8 and 9 are found at
depths between 500 and 800 m, and PSH 10 is found even deeper, between 900 and 1000 m. In contrast
Fig. 1. Bayesian phylogenetic tree obtained with the cox1 dataset. Posterior probabilities (> 0.95) and
bootstraps (> 90) are shown for each node. The boxes in front of the lineages of Sibogasyrinx Powell,
1969 represent the ABGD PSHs, numbered from 1 to 10. Alternative PSH partitions obtained in the
second and third-best ASAP partitions are shown with dashed lines. The colors refer to the locality;
* = illustrated shells.
European Journal of Taxonomy 773: 19–60 (2021)
28
with the shallowest species sister to the four others, and the three intermediate ones in paraphyly and
including the deepest one. Finally, diagnostic morphological characters have been found to distinguish
cox-1 and morphological distinctiveness together with sympatric distributions
sometimes associated with bathymetric partitions, potentially correlated to ecological distinctiveness,
The three main lineages recovered (PSH 1–3 / PSH 4–5 / PSH 6–10), although not distinguishable by
Fig. 1). Furthermore, the cox-1 distances among these lineages are similar to the distances between
genera of Cochlespiridae. For these reasons, they could have been recognized as (sub)genera. However,
Fig. 2. Bathymetric ranges of species of Sibogasyrinx Powell, 1969. The dashed bar for PSH 4
(S. elbakyanae sp. nov.) marks the range of the trawl haul in which a single specimen was collected. The
grey bar for PSH 6 (S. losa Ardovini, 2021) indicates the depths of occurrence of the type material.
Stars indicate the depths of collection of the holotypes of non-sequenced species.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
29
because of their morphological similarity, and, more importantly, because of the lack of DNA sequence
data for the type species of the genus (see Taxonomy section below), attributing the name Sybogasyrinx
to any of the three lineages (and thus new names for the two others) would be untimely, and we thus
abstain from naming them.
From SSH to named species
Both PSH 1 and 2 resemble the type of S. pyramidalis and we refer to them as S. cf. pyramidalis 1 and
S. cf. pyramidalis 2 (see below – Taxonomy – for more discussion). The specimens of PSH 6 are very
similar to the type material of S. losa Ardovini, 2021 and we apply this name to it. PSH 7 contains the
holotype of S. sangeri, and this name has thus been applied to this PSH. We were not able either to link
the name archibenthalis (Powell, 1969) or any names proposed in other genera to any of the remaining
PSHs (PSHs 3–5 and 8–10), and they are thus described as new species: S. subula sp. nov. (= PSH 3),
S. elbakyanae sp. nov. (= PSH 4), S. pagodiformis sp. nov. (= PSH 5), S. lolae sp. nov. (= PSH 8),
S. maximei sp. nov. (= PSH 9) and S. clausura sp. nov. (= PSH 10). Sibogasyrinx archibenthalis is thus
the only species of the genus not represented in the molecular phylogeny.
For comparative purposes, we included several specimens of the distantly related genus Leucosyrinx
that bear a strong conchological resemblance to the species of Sibogasyrinx treated herein. All but
one, L. verrillii (Dall, 1881) (the type species of the genus), represent still unnamed species that are
marked on the tree as Leucosyrinx sp. A to Leucosyrinx sp. G (Fig. 1). The included specimens represent
a subsample of the sequenced material for the corresponding species and therefore we abstain from
describing the new species pending a general revision of the genus.
Taxonomy
Class Gastropoda Cuvier, 1795
Subclass Caenogastropoda Cox, 1960
Order Neogastropoda Wenz, 1938
Superfamily Conoidea Fleming, 1822
Family Cochlespiridae Powell, 1942
Genus Sibogasyrinx Powell, 1969
Leucosyrinx (Sibogasyrinx) Powell, 1969: 343.
Type species
Surcula pyramidalis Schepman, 1913 (original designation).
Diagnosis
Shell small to large, adult length from 27 to 65 mm, narrowly fusiform, rarely pagodiform, with concave
comprising narrow, close-set cords, often also on subsutural ramp, Shoulder with a row of strong
nodules, often obsolete on last whorl.
Venom gland does not pass through nerve ring and opens into oesophagus within or posterior to nerve
ring. Radula usually with central tooth, absent in one species, marginal tooth morphology variable,
folded longitudinally.
European Journal of Taxonomy 773: 19–60 (2021)
30
Sibogasyrinx pyramidalis (Schepman, 1913) complex
Sibogasyrinx pyramidalis (Schepman, 1913)
Fig. 3H–I
Surcula pyramidalis
Leucosyrinx (Sibogasyrinx) pyramidalis
Sibogasyrinx pyramidalis – Kantor et al.
Material examined
Holotype
Remarks
Kantor et al. (2018) provided illustrations of shells and radulae of sequenced specimens of what they
considered to be S. pyramidalis
and in the central South China Sea (= PSH 2, S. cf. pyramidalis 1). The shell outline of these specimens
of this generally variable species matched the illustration of the holotype, which was not examined.
The present molecular analysis revealed that there are two molecularly distinct but morphologically
cryptic species that match the description and illustrations of the holotype of S. pyramidalis (Schepman
in Kantor et al
(= PSH 1, S. cf. pyramidalis 2). The geographic ranges of these two PSHs do not include the type
locality of S. pyramidalis (Timor Sea, Pulau Rote Island). There are two possible explanations for this:
one of our species represents the true S. pyramidalis, or S. pyramidalis is a third species and both of
ours are new to science. Unfortunately, this can only be resolved by sequencing topotypic material
from the Timor Sea. In order to not add to the taxonomic ambiguity, we presently refrain from taking
any taxonomic decision, but for the sake of convenience refer here to our molecular species as S. cf.
pyramidalis 1 (= S. pyramidalis sensu Kantor et al. 2018) and S. cf. pyramidalis 2.
S. cf. pyramidalis 1 and S. cf. pyramidalis 2 (Kantor et al
Sibogasyrinx cf. pyramidalis (Schepman, 1913) 1 (= PSH 2)
Figs 3A–G, 4A–B
Material examined (all sequenced)
IM-2009-13451.
stn CP4129; MNHN-IM-2013-50215.
Description
largest specimen). SL 65.2 mm, AL (with canal) 35.1 mm, AL (without canal)
21.0 mm, SW 18.5 mm.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
31
Fig. 3. Sibogasyrinx pyramidalis (Schepman, 1913) complex. A–G. Sibogasyrinx cf. pyramidalis
1. A–C. South China Sea, MNHN-IM-2013-44605, SL 65.2 mm. D–F. Philippines, MNHN-
IM-2009-13451, SL 46.8 mm. G. South China Sea, MNHN-IM-2013-44574, SL 53.9 mm. H–I. Surcula
pyramidalis Schepman, 1913, holotype, ZMA.MOLL.136836, SL 45 mm (photos by K. Hasegawa,
copyright Naturalis, Leiden). J–Q. Sibogasyrinx cf. pyramidalis 2, Papua New Guinea. J–L. MNHN-
IM-2013-19745, SL 65.2 mm. M–O. MNHN-IM-2009-17000, SL 59.2 mm. P. MNHN-IM-2009-16973,
SL 41 mm. Q. MNHN-IM-2009-16988, SL 43 mm. R–T. Sibogasyrinx archibenthalis (Powell, 1969),
holotype, USNM 238773, SL 41.8 mm. All shells to same scale.
European Journal of Taxonomy 773: 19–60 (2021)
32
Moderately thick, strong except for fragile outer aperture lip, fusiform, with high spire and
long, narrow, straight siphonal canal. Protoconch small, globose, of about 1.5 strongly convex,
microshagreened whorls, eroded or missing in all specimens. Early teleoconch whorls weakly to
moderately angular at shoulder. Largest available specimen (SL 65.2 mm) of 10.3 teleoconch whorls.
a subsutural row of nodules, corresponding in shape to upper parts of thickened growth lines. Nodules
more distinct on upper whorls, increasing in number, up to 30 on penultimate whorl and 37 on last whorl
in largest specimen. Subsutural ramp may be completely smooth, or sometimes with very weak spiral
striation or dendritic lines. Shoulder with pronounced thickened and rounded nodules, reaching lower
lines on lower whorls. Nodules evanesce on last and even penultimate whorls at SL over 50 mm and
shoulder becomes evenly rounded; their number increase with size from 14–15 on upper whorls to
21–22 on lower whorls. Spiral sculpture of 3–6 moderately pronounced, closely spaced narrow cords
on shoulder, crossing shoulder nodules on spire whorls. Last whorl below shoulder with 40–45 cords
varying in width, their intervals 0.5–2 times width of cords, also distinct on canal. Shell base gradually
narrowing towards long, narrow, nearly straight siphonal canal. Aperture narrow, constricted posteriorly,
with narrow and thin parietal callus, outer lip with rounded angle at shoulder, weakly convex below
shoulder and weakly concave at transition to canal. Anal sinus moderately deep, subsutural, broadly
subsutural band, protoconch pale tan. Periostracum persists on part of shell, light yellowish.
Foregut similar to that of Sibogasyrinx cf. pyramidalis 2 (see below), except for presence of
a large oval accessory salivary gland, adjoining oesophagus, with rather thick duct, entering proboscis
and following along its wall.
n = 2; MNHN-IM-2009-13451, MNHN-IM-2013-50215). Comprising approximately 40
530
broadly concave, borders distinct and with narrow but rather long, sharply pointed cusp. Marginal teeth
elevated posterior edge, on developing part of radular tooth folding clearly visible (on Fig. 4A not fully
folded teeth are marked by white arrows), bringing posterior and anterior edges close together. During
tooth maturation the edges, particularly posterior one, progressively thicken, so that fully formed tooth
appears duplex (Fig. 4B).
Distribution
Sibogasyrinx cf. pyramidalis (Schepman, 1913) 2 (= PSH 1)
Figs 3J–Q, 4C–D, 5
Material examined (all sequenced)
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
33
Fig. 4. Radulae of species of Sibogasyrinx Powell, 1969. A–B. S. cf. pyramidalis (Schepman, 1913)
1, MNHN-IM-2009-13451 (shell seen on Fig. 3D–F). A. Part of radula with marginal teeth not fully
formed; white arrows indicate teeth that are not fully folded longitudinally (ae = anterior tooth edge;
pe = posterior tooth edge). B. Part of radula with fully formed marginal teeth. C. S. cf. pyramidalis 2,
MNHN-IM-2009-16972. D. S. cf. pyramidalis 2, MNHN-IM-2009-17000 (shell seen on Fig. 3M–O).
E. S. subula sp. nov., MNHN-IM-2007-17703. F. S. subula sp. nov., MNHN-IM-2009-13568 (shell seen
on Fig. 6H–I).
European Journal of Taxonomy 773: 19–60 (2021)
34
depth 827–966 m; KAVIENG 2014, stn CP4483; MNHN-IM-2013-58845.
Description
largest specimen). SL 65.2 mm, AL (with canal) 34.7 mm, AL without canal 22.9 mm,
SW 18.0 mm.
Extremely similar to that of Sibogasyrinx cf. pyramidalis
larger number of spiral cords crossing the shoulder nodules. Penultimate whorl of S. cf. pyramidalis 1
with 5–6 cords (increasing with SL), 6–8 in S. cf. pyramidalis 2.
n = 2, MNHN-IM-2009-16972, MNHN-IM-2009-17000). Head with long, cylindrical
tentacles, rounded on tips with closely spaced bases, eyes absent. One specimen (MNHN-IM-2009-17000)
male, with long penis with obliquely truncated tip. Seminal papilla occupies entire tip and is surrounded
by circular fold. Proboscis long, coiled, with broad conical base, occupied by large bulbous buccal mass,
spanning posterior ¼ of proboscis and protruding backwards from proboscis. Proboscis sheath with very
thin transparent walls. Remaining anterior part of proboscis narrow, gradually diminishing in diameter
edge of rhynchodeum. Radular sac and short oval odontophore situated on right side of buccal mass,
posterior to rear of proboscis. Oesophagus adjoining buccal mass slightly constricted and forming a
moderately long loop before passing through nerve ring. Salivary glands irregularly shaped, covering
dorsal portion of nerve ring. Accessory salivary gland not traced. Venom gland thick, long, strongly
convoluted, opens dorsally into oesophagus, immediately posterior to nerve ring (Fig. 5B). Muscular
bulb oval, of moderate size, situated to the right of oesophagus.
n = 2; MNHN-IM-2009-16972, MNHN-IM-2009-17000). Comprising approximately 40 rows
without canal). Radula width up to 300
to that of Sibogasyrinx cf. pyramidalis 1, central tooth has less rounded posterior margin and in one
specimen (MNHN-IM-2009-17000, Fig. 4D) the cusp is obtuse.
Fig. 5. Anterior foregut of Sibogasyrinx cf. pyramidalis (Schepman, 1913) 2, MNHN-IM-2009-16972.
A. Right latero-dorsal view. B. Dorsal view of the nerve ring and opening of the venom gland.
Abbreviations: mb = muscular bulb of the venom gland; nr = circumoesophageal nerve ring;
od = odontophore; oe = oesophagus; pr = proboscis; prr = proboscis retractors; rs = radular sac; sd =
salivary duct; sg = salivary gland; vg = venom gland.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
35
Remarks
Despite the similarities in shell and radular morphology, Sibogasyrinx cf. pyramidalis 1 and Sibogasyrinx
cf. pyramidalis 2 are clearly distinct from a molecular perspective. One morphological character of the
foregut that separates the two species is the presence of an accessory salivary gland in the former.
Distribution
Sibogasyrinx archibenthalis (Powell, 1969)
Fig. 3R–T
Leucosyrinx (Sibogasyrinx) archibenthalis
Material examined
Holotype
R/V Albatross, stn 5513; USNM 238773.
Description
holotype). SL 41.8 mm.
un-notched siphonal canal. Protoconch small, globose, of 1.5 smooth whorls. Teleoconch of 12 whorls.
carina, abutting lower suture. Last whorl distinctly angled by peripheral carina, and base rapidly
contracting toward moderately long anterior canal. Axial sculpture of rounded peripheral nodules,
about 13 per whorl, but these become obsolete on last two whorls; subsutural nodules or folds lacking.
Remarks
The species is known only from the holotype. In shell shape it is most similar to S. pyramidalis. The
last whorl is lower and the canal is proportionally much shorter in S. archibenthalis, while the spiral
cords on the subsutural ramp are more pronounced and are of the same strength as on the remaining
part of the last whorl. The last whorl is more angular in S. archibenthalis and the shell has a higher
number of teleoconch whorls; in the holotype of S. archibenthalis the number of teleoconch whorls is 12
(SL 43 mm), compared with 9.5 (SL 46.8 mm) in samples of S. cf. pyramidalis 1 from the Philippines of
similar shell size. Therefore, at present we consider S. archibenthalis to be a valid species.
Sibogasyrinx subula sp. nov. (= PSH 3)
urn:lsid:zoobank.org:act:6A992BBF-7C62-4F15-BFCC-AD9F347ED379
Figs 4E–F, 6
Sibogasyrinx sp. – Bouchet et al.
European Journal of Taxonomy 773: 19–60 (2021)
36
Diagnosis
Shell large, reaching 57 mm in length, narrowly fusiform, with concave, nearly smooth subsutural
ramp with darker subsutural band on posterior half of last whorl and strong nodules on shoulder. Spiral
sculpture of distinct, closely spaced cords below subsutural ramp. Radula with central tooth and duplex
marginal teeth.
Etymology
Latin ‘subula’ (noun in apposition), meaning ‘awl’, with reference to the general appearance of the spire.
Material examined
Holotype
stn CP2187; MNHN-IM-2007-42530.
Other material (all sequenced)
stn CP4448; MNHN-IM-2013-58409.
stn CP2184; MNHN-IM-2009-13567, MNHN-IM-2009-13568 • 1 lv; SW of Sta Isabel I.;
to IM-2007-17704.
Description
(holotype). SL 55 mm, AL (with canal) 31 mm, AL (without canal) 17.5 mm,
SW 15.6 mm.
(holotype). Moderately thick, strong except for fragile outer lip of aperture, fusiform, with high
spire and long, narrow, straight siphonal canal. Protoconch small, globose, of about 1.5 strongly convex,
microshagreened whorls. Protoconch/teleoconch transition marked by arcuate axial rib and appearance
shoulder. Teleoconch comprising just under 10 whorls in total. Suture shallowly impressed, subsutural
ramp broad, strongly concave. Subsutural region with a row of distinct sigmoidal nodules that correspond
32 on penultimate and 30 on last whorl. Subsutural ramp smooth except for subsutural row of nodules,
few thickened growth lines and 3–4 very weak spiral threads on last and penultimate whorls. Shoulder
with pronounced thickened and rounded nodules, weakly opisthocline on upper whorls, but more
whorls and 19 on last whorl. Spiral sculpture of 4–5 moderately pronounced, narrow cords on shoulder,
seen both on nodules and in their intervals, and rather distinct, weakly rounded cords below shoulder,
On last whorl cords occupy entire shell below shoulder, varying in width, their intervals 1–2 times width
of cords, in total 48 cords on last whorl, of which 28 on canal. Shell base gradually narrowing towards
long, narrow, nearly straight siphonal canal. Aperture narrow, constricted posteriorly, with very narrow
and thin parietal callus, outer lip partially broken, with rounded angle at shoulder, weakly convex below
shoulder and weakly concave at transition to canal. Anal sinus moderately deep, subsutural, broadly
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
37
subsutural band and darker subsutural band on posterior half of last whorl, protoconch light tan.
n = 1, MNHN-IM-2009-13568). Head with long cylindrical tentacles, rounded on tips with
closely spaced bases, eyes absent. Specimen male, with long penis with obliquely truncated tip. Seminal
Fig. 6. Sibogasyrinx subula sp. nov. A–C. Solomon Islands, holotype, MNHN-IM-2007-42530, SL
55 mm. D–F. Solomon Islands, MNHN-IM-2007-42533, SL 53.6 mm. G. Papua New Guinea, MNHN-
IM-2013-58409, SL 36.3 mm. H–I. Solomon Islands, MNHN-IM-2009-13568, SL 49.4 mm (radula seen
on Fig. 4F). J–L. Vanuatu, MNHN-IM-2007-17704, SL 53 mm. M. Vanuatu, MNHN-IM-2007-17701,
SL 46.0 mm. N. Solomon Islands, MNHN-IM-2009-13567, SL 57.4 mm. All shells to same scale.
European Journal of Taxonomy 773: 19–60 (2021)
38
papilla occupies entire tip and is surrounded by circular fold. Proboscis conical, with broad base, but
very narrow along most of its length. Proboscis retractors large, arranged in two bundles attached to
proboscis base. Buccal mass very large and broad, nearly as long as proboscis, oesophagus forming
short loop before nerve ring. Venom gland thick, medium-long and convoluted, opening dorsally into
oesophagus just posterior to nerve ring. Muscular bulb very large.
n = 3; MNHN-IM-2007-17701, MNHN-IM-2007-17703, MNHN-IM-2009-13568). All
specimens with similar radular morphology, radula comprising 38–40 rows of teeth, 11–17 nascent,
canal). Central tooth with broad, subrectangular basal plate, anteriorly overlapped by preceding tooth,
borders distinct, with narrow, sharply pointed cusp. Marginal teeth with strongly thickened posterior
edges, folded longitudinally (Fig. 4E–F), in all details similar to those of Sibogasyrinx cf. pyramidalis 1.
Remarks
Largest specimen attains an SL of 57.4 mm.
The other specimens are rather similar to the holotype in shell shape and sculpture. The entire subsutural
ramp may be pale yellowish on the last whorl, but the subsutural zone of the posterior part of last whorl
in larger specimens is always more darkly coloured than the rest of the shell. The subsutural ramp may
be completely smooth or with indistinct spiral threads, the shoulder angulation on the last whorl is
sometimes less distinct than in the holotype, as may be the cords below the shoulder on the penultimate
whorl (up to 4).
In shell outline, the new species resembles its congener Sibogasyrinx losa
much less distinct spiral sculpture on the subsutural ramp, as well as in its larger size (maximal SL
57.4 mm in S. subula sp. nov. and 52 mm in S. losa) and radular morphology. The shell shape of the new
species also strongly resembles that of some species of Leucosyrinx, particularly L. verrillii (Dall, 1881)
(Fig. 13A–C) and to a lesser extent the smaller specimens of Leucosyrinx sp. B and Leucosyrinx sp. G
(Fig. 13D and 13I, respectively). All these species of Leucosyrinx, however,
morphology, shorter siphonal canal and, according to the molecular tree (Fig. 1) and other published
Conoidea phylogenies (Puillandre et al. 2011; Abdelkrim et al
Distribution
This species is known from the Solomon Islands, Papua New Guinea and Vanuatu, and has a relatively
broad bathymetric range, from 440 to 814 m.
Sibogasyrinx sangeri Kantor, Fedosov & Puillandre, 2018 (= PSH 7)
Figs 7, 8A–B
Sibogasyrinx sangeri
Material examined
Holotype
PAPUA NEW GUINEA •
IM-2009-17022 (sequenced).
Other material (all sequenced)
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
39
724 m; PAPUA NIUGINI, stn CP3982; MNHN-IM-2013-19752 • 1 lv; Dampier Strait, E of Umboi I.;
IM-2009-13434.
SALOMON 2, stn CP2288; MNHN-IM-2007-42523, MNHN-IM-2009-16779.
Description
Holotype: SL 54.1 mm, AL (with canal) 31.3 mm, AL (without canal) 16.4 mm,
SW 14.1 mm. Largest available specimen attains SL 55.8 mm (MNHN-IM-2009-16779; Fig. 7D–E).
Thin, fragile, variable in shape, from narrowly fusiform to moderately broad (SW/SL ratio
from 0.22 to 0.27), with rather high spire and long, narrow, straight siphonal canal. Protoconch small,
globose, of 1.75 strongly convex, microshagreened whorls. Protoconch/teleoconch transition indistinct.
Protoconch diameter about 1.1 mm, height 1.2 mm. Early teleoconch whorls angular, usually in lower
part just above suture, but sometimes at mid-whorl; last whorl strongly to very weakly angled or evenly
rounded, even in specimens of the same size. Teleoconch comprising up to 10 whorls. Suture shallow,
becoming more broadly spaced and less discernible on later whorls, sometimes obsolete on last whorl of
largest specimens. Subsutural zone very weakly concave, nearly straight on penultimate and last whorls,
smooth except for a few irregularly spaced and indistinct spiral threads (sometimes oblique), or several
a row of larger, more pronounced nodules, just above suture in upper whorls or sometimes nearer mid-
whorl. In some specimens nodules absent on last whorl. Last whorl may bear low carina at periphery
(in specimens with nodules absent), giving it very weakly angled outline. Below periphery 2–3 distinct
spiral cords on penultimate whorl and about 30 cords below carina on last whorl and 20 on canal. Shell
base gradually narrowing towards long, nearly straight siphonal canal. Aperture narrow, constricted
posteriorly with broad, very thin parietal callus, outer lip partially broken, convex and weakly angled in
upper part and slightly convex below shoulder, and shallowly concave at transition to canal. Anal sinus
n = 1, MNHN-IM-2009-16995). Proboscis of moderate length in contracted stage, buccal
mass situated outside proboscis. Oesophagus very broad, forming short loop before passing through
nerve ring. Venom gland opening into oesophagus ventrally and immediately posterior to nerve ring.
n = 1, MNHN-IM-2009-16995) (Fig. 8A–B). Relatively short, comprising ca 40 rows of teeth,
365
trough-shaped during maturation, folded longitudinally when fully formed. On developing part of
radula, folding of teeth occurs abruptly, within one subsequent row (on Fig. 8B white arrow indicates
European Journal of Taxonomy 773: 19–60 (2021)
40
evident at 17th row in radula studied. Resulting folded tooth moderately broad, with sharp pointed tip,
formed by overlapping of both thickened margins (on Fig. 8A these overlapping parts are marked by
hollow white arrows).
Distribution
This species is known from the Solomon Islands, Papua New Guinea and the Philippines, over a
relatively broad bathymetric range, from 448 to 788 m.
Fig. 7. Sibogasyrinx sangeri Kantor, Fedosov & Puillandre, 2018. A–B. Holotype, Papua New Guinea,
MNHN-IM-2009-17022, SL 54.1 mm. C. Papua New Guinea, MNHN-IM-2009-16995, SL 47.7 mm
(radula seen on Fig. 8A–B). D–E. Solomon Islands, MNHN-IM-2009-16779, SL 55.8 mm. F. Papua
New Guinea, MNHN-IM-2009-17021, SL 53.7 mm. G–H. Papua New Guinea, MNHN-IM-2009-17057,
SL 38.8 mm. I–J. Philippines, MNHN-IM-2009-13434, SL 36.9 mm. K. Solomon Islands, MNHN-
IM-2009-16766, SL 26.3 mm. All shells to the same scale.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
41
Sibogasyrinx losa Ardovini, 2021 (= PSH 6)
Figs 8C, 9A–H
Sibogasyrinx losus
Material examined
Holotype
MNHN-IM-2000-37629 (originally R. Ardovini collection).
Paratype
PAPUA NEW GUINEA • 1 lv; same collection data as for holotype; R. Ardovini collection.
Other material (all sequenced)
stn DW4500; MNHN-IM-2013-59044.
Description
Largest available specimen (MNHN-IM-2007-42498): SL 49.9 mm, AL (with canal)
28 mm, AL (without canal) 16.7 mm, SW 13.5 mm. Holotype has SL 52 mm.
Moderately thick, strong except for fragile and often partially broken outer aperture lip, narrowly
fusiform, with high spire and long, narrow, straight siphonal canal. Protoconch small, globose, of just
over 1.5 strongly convex, microshagreened whorls (MNHN-IM-2009-16831). Protoconch/teleoconch
transition indistinct, marked by appearance of shoulder carina. Protoconch diameter 0.89 mm, height
0.72 mm. Spire whorls strongly angled at shoulder, last whorl with more rounded shoulder. Total
teleoconch whorls just under 9 in largest specimen. Suture shallowly impressed on last whorl and rather
deep on spire ones, subsutural ramp moderately broad, strongly concave. Subsutural region with a row
whorl and distinct on fourth whorl. Last, penultimate and antepenultimate whorls with 4–5 distinct
cords on subsutural ramp, their intervals equal to width of cords, followed below shoulder by 5–7 more
narrow, closely spaced cords. Shoulder with row of pronounced rounded nodules, more distinct on upper
teleoconch whorls and absent on last whorl, 15–17 on penultimate and antepenultimate whorls. Spiral
sculpture in addition to cords on subsutural ramp of distinct narrow cords, covering entire shell surface,
including shoulder nodules. On last whorl about 45 cords below shoulder, 25 of which on canal. Cords
even slightly wider than cords. Shell base gradually narrowing towards long, narrow, almost straight
siphonal canal. Aperture narrow, constricted posteriorly, with very narrow and thin parietal callus, outer
lip with rounded angle at shoulder, weakly convex below shoulder, weakly concave at transition to
canal.
with very light yellowish subsutural ramp and irregular darker blotches on subsutural ramp. Protoconch
light tan.
European Journal of Taxonomy 773: 19–60 (2021)
42
Fig. 8. Radulae of species of Sibogasyrinx Powell, 1969. A–B. Sibogasyrinx sangeri Kantor, Fedosov
& Puillandre, 2018, MNHN-IM-2009-16995 (shell seen on Fig. 7C). A. Part of radula with fully formed
marginal teeth; white hollow arrow indicates overlapping edges of the tooth at its tip. B. Part of radula
showing transition between unfolded (white arrow) and completely longitudinally folded (black arrow)
marginal teeth (ae = anterior tooth edge; pe = posterior tooth edge). C. Sibogasyrinx losa Ardovini,
2021, Solomon Islands, MNHN-IM-2009-16831 (shell seen Fig. 9E–F). D. Sibogasyrinx lolae sp. nov.,
MNHN-IM-2009-29311. E. Sibogasyrinx maximei sp. nov., holotype, MNHN-IM-2013-45883, anterior
end of radula. F. Sibogasyrinx pagodiformis sp. nov., MNHN-IM-2009-11327.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
43
n = 1; MNHN-IM-2009-16831). Head with moderately long, conical tentacles, rounded on
tips and with closely spaced bases, large eyes situated on small lobes at tentacle base. Proboscis conical,
posterior part of very thin proboscis sheath base weakly muscular. Buccal mass very large and broad,
posterior to proboscis base, constituting about half of proboscis length, oesophagus forming a very
short loop before nerve ring. Radular sac with small odontophore, opening dorso-laterally at right side
of buccal mass. Venom gland thick, moderately long and convoluted, opening into oesophagus within
nerve ring. Muscular bulb moderately large, elongated and folded in posterior part. Salivary glands
fused, relatively large, of irregular shape, acinous. Small ovate accessory salivary gland situated dorsally
at nerve ring.
n = 1; MNHN-IM-2009-16831) (Fig. 8C). Relatively short, consisting of 41 rows of teeth,
canal). Central tooth with subrectangular basal plate, having distinct borders and moderately long cusp.
Anterior margin overlapped by preceding row, posterior margin evenly and weakly rounded. Marginal
longitudinally when fully formed, with both margins overlapping at tooth tip. Resulting folded tooth
moderately broad, with sharp pointed tip, anterior edge with a narrow slit between folded tooth margins.
Remarks
Our specimens are very similar to the holotype and paratype (except lighter shells) and were collected
close to the type locality. Our specimens were collected at much shallower depths (150–508 m) as
compared to the holotype and paratype (1100 m). Nevertheless, the strong similarity of the shell shape
and sculpture, especially obvious when comparing the holotype with sequenced specimen of the same
himself considered the specimen MNHN-IM-2007-42498 as belonging to his newly described species
based on the photograph on the MNHN website. Smaller specimens (SL 34.4 and 29.9 mm) retain
This species is most similar to Sibogasyrinx subula
spiral cords on the subsutural ramp, a relatively narrower ramp resulting in a higher shoulder position on
some species of Leucosyrinx, particularly Leucosyrinx sp. A (Fig. 13H), but the shell is broader with
more numerous nodules on the shoulder. Compared to Leucosyrinx sp. F (Fig. 13K), the shell of S. losa
is larger, broader and has a coarser sculpture.
Distribution
150–1100 m.
Sibogasyrinx lolae sp. nov. (= PSH 8)
urn:lsid:zoobank.org:act:73404289-ED57-4ABD-A0D0-05939F095C40
Figs 8D, 9I–Q
Diagnosis
Shell medium-sized, reaching 43 mm in length, fusiform, with weakly concave, nearly smooth subsutural
ramp and nodules on shoulder of upper teleoconch whorls, absent on later whorls of large specimens.
Spiral sculpture weak, of closely spaced cords covering entire shell except subsutural ramp in some
specimens. Radula with central tooth and longitudinally folded marginal teeth with weakly thickened
edges.
European Journal of Taxonomy 773: 19–60 (2021)
44
Etymology
The species is named after Lola, the daughter of the second author.
Material examined
Holotype
3, stn CP2839; MNHN-IM-2007-42537.
Other material (all sequenced)
KANADEEP, stn CP4923; MNHN-IM-2013-48156.
Description
(holotype). SL 37.3 mm, AL (with canal) 22.9 mm, AL (without canal) 14.5 mm, SW
11.3 mm.
(holotype). Moderately thick, glossy, strong except for very fragile and partially broken outer
aperture lip, narrowly fusiform, with high spire and long, narrow, straight siphonal canal. Protoconch
small, globose, corroded, of about 1.5 whorls. Protoconch preserved in other juvenile specimen
(MNHN-IM-2013-48156), comprising just over 1.5 strongly convex, microshagreened whorls.
Protoconch/teleoconch transition indistinct, marked by appearance of axial ribs. Protoconch diameter
1.0 mm, height 0.74 mm. Spire whorls angled at shoulder, last whorl with less angular shoulder. Total
teleoconch whorls just under 8. Suture shallowly impressed on last whorl and rather deep on spire
whorls, subsutural ramp moderately broad, weakly concave on upper teleoconch whorls and nearly
whorl, and without spiral sculpture on upper four whorls and later with indistinct spiral cords, three
on last whorl. Shoulder of teleoconch whorls (except last one) with a row of distinct, opisthoclinely
on antepenultimate and penultimate whorls. Weak spiral cords on and below shoulder, starting from 3rd
teleoconch whorls, about twice as wide below shoulder. Penultimate whorl with four cords on shoulder
and four below shoulder. Shoulder smooth on last whorl, but with about 55 cords below shoulder, of
which 25 on canal. Cords weak, slightly wavy and closely spaced, their intervals about half the width
of cords. Shell base gradually narrowing towards long, narrow, nearly straight siphonal canal. Aperture
narrow, constricted posteriorly, with narrow, very thin parietal callus, outer lip badly broken, distinctly
impressed at shoulder, weakly convex below shoulder and shallowly concave at transition to canal. Anal
of outer lip. Shell very light yellowish, protoconch very light tan (in holotype) and light brown in
MNHN-IM-2013-48156.
n = 1; MNHN-IM-2009-29311). Male. Penis tip obliquely truncated, with long and very
narrow papilla, surrounded by circular fold, much larger in diameter than papilla itself. Proboscis
conical, moderately long, with broad base, anterior half rapidly narrowing towards tip. Proboscis
retractors distinct, arranged in two symmetrical lateral bundles, attached to inner proboscis walls at
border of its posterior third. Buccal mass with small radular sac, odontophore situated within proboscis
in its broader posterior part; elongate oval, occupying slightly less than half of proboscis length. Single
small accessory salivary gland present. Salivary glands separate, acinous, irregular in shape. Venom
gland long, moderately thick and convoluted, opening into oesophagus within the nerve ring.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
45
Fig. 9. A–H. Sibogasyrinx losa Ardovini, 2021. A–B. Holotype, Solomon Islands (MNHN-
IM-2000-37629), SL 52 mm. C–D. Solomon Islands, MNHN-IM-2007-42498, SL 49.9 mm.
E–F. Solomon Islands, MNHN-IM-2009-16831, SL 34.4 mm (radula seen on Fig. 8C). G–H. Papua
New Guinea, MNHN-IM-2013-59044, SL 29.9 mm. I–Q. Sibogasyrinx lolae sp. nov. I–J. Solomon
Islands, holotype, MNHN-IM-42537, SL 37.3 mm. K–L. New Caledonia, MNHN-IM-2009-29230,
SL 43.1 mm. M–N. New Caledonia, MNHN-IM-2009-29311, SL 29.1 mm. O–Q. New Caledonia,
MNHN-IM-2013-48156, SL 17.1 mm (O at the same scale as the other specimens, P–Q enlarged). All
shells (except P–Q) to same scale.
European Journal of Taxonomy 773: 19–60 (2021)
46
n = 1; MNHN-IM-2009-29311) (Fig. 8D). Short, comprising 28 rows of teeth, 13 nascent.
tooth length 160
distinct borders and long, narrow, sharp cusp. Anterior margin overlapped by preceding row, posterior
formed, becoming trough-shaped with weakly thickened edges during maturation, folded longitudinally
when fully formed, with both margins overlapping at tooth tip. Resulting folded tooth moderately broad,
with sharp pointed tip, anterior edge with a narrow slit between tooth margins. Tooth folding occurring
within 15th row of teeth.
Remarks
The largest specimen attains 43.1 mm.
Other specimens are similar to the holotype in shell shape, although some have much more pronounced
spiral sculpture, especially MNHN-IM-2009-29230, which also retains spiral cords on the shoulder of
the last whorl.
In shell shape and sculpture the new species is most similar to Sibogasyrinx losa, some specimens
being almost indistinguishable (e.g., holotype of S. losa and MNHN-IM-2009-29230). Sibogasyrinx
lolae sp. nov. can be distinguished from S. losa due to its generally less distinct spiral sculpture and less
S. cf. pyramidalis 1 and
2, S. lolae sp. nov., the nodules
on the shoulder become less pronounced or disappear on later whorls. Nevertheless, the nodules are still
pronounced on the shoulder of the last whorl in specimens of S. cf. pyramidalis 1 and 2 of the same size
as specimens of S. lolae sp. nov. which already lack nodules on the last whorl.
Distribution
This species is recorded from the Solomon Islands, the Coral Sea and southern New Caledonia, at depths
of 460–787 m.
Sibogasyrinx maximei sp. nov. (= PSH 9)
urn:lsid:zoobank.org:act:EEC38F7C-DD00-46B9-9D56-3915EB0B2CD2
Figs 8E, 10A–C
Diagnosis
Shell medium-sized, reaching 41.6 mm in length, fusiform, subsutural ramp weakly concave on early
whorls with nodules, absent on last whorl. Spiral sculpture weak, of indistinct closely spaced cords on
and below shoulder. Radula with central tooth and longitudinally folded marginal teeth with weakly
thickened edges.
Etymology
This species is named after Maxime, the son of the second author.
Material examined
Holotype
MNHN-IM-2013-45883.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
47
Description (holotype)
SL 41.6 mm, AL (with canal) 23.2 mm, AL (without canal) 15.7 mm, SW 12.4 mm.
Fig. 10. A–C. Sibogasyrinx maximei sp. nov., Solomon Sea, holotype, MNHN-IM-2013-45883,
SL 41.6 mm. D–M. Sibogasyrinx clausura sp. nov. D–F. Solomon Islands, holotype, MNHN-
IM-2013-48256, SL 41.4 mm. G–H. Coral Sea, Coriolis Bank, MNHN-IM-2013-48167, SL 37.9 mm.
I–J. Solomon Islands, MNHN-IM-2009-16763, SL 29.6 mm. K–L. Coral Sea, Coriolis Bank, MNHN-
IM-2013-48244, SL 35.6 mm. M. Coral Sea, Coriolis Bank, MNHN-IM-2013-48144, SL 31.7 mm. All
shells to same scale.
European Journal of Taxonomy 773: 19–60 (2021)
48
Moderately thick, strong except for fragile and partially broken outer aperture lip, fusiform,
with high, conical spire and moderately long, broad, straight siphonal canal. Protoconch small, globose,
of about 1.5 strongly convex, microshagreened whorls. Protoconch/teleoconch transition marked by
strongly arcuate axial rib, corresponding to shape of growth line. Protoconch diameter 1.0 mm, height
0.83 mm. Spire whorls weakly angled at shoulder, last whorl with hardly discernible shoulder. Total
with only traces of indistinct spiral striation. Shoulder of teleoconch whorls (except last one) with a
row of distinct elongated nodules, these nearly orthocline on upper whorls and weakly opisthocline
on penultimate whorl, 14–17 per whorl, reaching lower suture and intersected by spiral cords. Starting
from 2nd teleoconch whorl, spiral cords appearing on shoulder, these intersecting nodules and becoming
progressively stronger, from two on 2nd whorl to nine on penultimate one. Cords very closely spaced,
separated by narrow grooves. On last whorl cords cover entire shell surface below indistinct shoulder,
about 50 cords in total, of which 25 on canal. Cords weak, slightly wavy, very closely spaced just below
suture and with intervals 0.5–1.0 times width of cords on shell base and canal. Shell base gradually
narrowing towards moderately broad, almost straight siphonal canal. Aperture narrow, constricted
posteriorly, with narrow and very thin parietal callus, outer lip partially broken, evenly convex and
weakly concave at transition to canal.
with large forward extension of outer lip. Shell very light yellowish, protoconch light tan. Periostracum
smooth, tightly adhering.
n = 1; MNHN-IM-2013-45883) (Fig. 8E). Medium-long, comprising 40 rows of teeth, 12
canal). Central tooth with basal plate, having distinct borders and long, narrow, sharp cusp, anterior
margin overlapped by preceding row, posterior margin formed by two straight sections meeting at obtuse
during maturation, folded longitudinally when fully formed, with both margins overlapping at tooth tip.
Resulting folded tooth moderately broad with sharply pointed tip, border between both margins present
as a narrow slit at anterior edge. Tooth folding occurring within 14th row of teeth (counting from rear).
Remarks
The species is known from the holotype alone and is very similar to Sibogasyrinx clausura sp. nov.
(for comparison see remarks under the following species). In shell outline it is very similar to S. cf.
pyramidalis
on the penultimate whorl, whereas specimens of S. cf. pyramidalis 1 and 2 of similar size have up to
Distribution
Known only from the type locality.
Sibogasyrinx clausura sp. nov. (= PSH 10)
urn:lsid:zoobank.org:act:27ED0BF0-C78A-41E1-9064-E3EB0492864A
Figs 10D–M, 11A–B
Diagnosis
Shell medium-sized, reaching 41.5 mm in length, fusiform, subsutural ramp concave on upper teleoconch
whorls, weakly so on last whorl, nearly smooth (except for the axial riblets), shoulder bearing nodules on
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
49
spire whorls, absent on last whorl. Spiral sculpture of weak, closely spaced cords on and below shoulder.
Radula with central tooth and longitudinally folded marginal teeth with weakly thickened edges.
Etymology
Latin ‘clausura’ (noun in apposition), meaning ‘lockdown’, with reference to the sanitary restrictions
Material examined
Holotype
stn CP4964; MNHN-IM-2013-48256.
Other material (all sequenced)
R/V Investigator, cruise IN2017_V03_100; AMS C.519344.
KANADEEP, stn CP4966; MNHN-IM-2013-48167, MNHN-IM-2013-48235.
stn CP2182; MNHN-IM-2009-16763.
Description
(holotype, largest specimen). SL 41.4 mm, AL (with canal) 23.0 mm, AL (without canal)
13.7 mm, SW 12.4 mm.
(holotype). Moderately thick, slightly glossy, strong except for fragile and partially broken outer
aperture lip, narrowly fusiform, with high spire and moderately long, straight siphonal canal. Protoconch
small, globose, of about 1.5 strongly convex, eroded whorls. Protoconch/teleoconch transition marked
by strongly arcuate axial rib, corresponding to shape of growth lines, followed by 10 thinner and weaker
axial ribs and rather thickened growth lines. Protoconch diameter 1.0 mm, height 0.84 mm. Spire whorls
distinctly angled at shoulder, last whorl with scarcely discernible shoulder. Total teleoconch whorls just
over 8. Suture shallowly impressed, subsutural ramp broad, concave, on last whorl weakly concave.
Subsutural region with a row of distinct, dense, narrow, short, prosocline axial wrinkles, corresponding
(except last one) with a row of distinct opisthoclinely elongated nodules, extending to abapical suture
and penultimate whorls. Subsutural ramp smooth except for axial wrinkles mentioned above. Spiral
cords intersecting shoulder nodules beginning on 2rd teleoconch whorls (due to corrosion of shell their
number is unclear) and becoming progressively stronger, 7 on antepenultimate and 8 on penultimate
whorl. Cords closely spaced with intervals about half the width of cords. On last whorl cords covering
entire shell surface below indistinct shoulder, about 40 cords in total, of which 20 on canal. Cords weak,
slightly wavy, very closely spaced just below suture and with intervals 0.5–1.0 times cord width on shell
base and canal. Shell base gradually narrowing towards narrow, nearly straight siphonal canal. Aperture
narrow, constricted posteriorly, with narrow, moderately thick parietal callus, outer lip partially broken,
evenly convex and weakly concave at transition to canal. Anal sinus shallow, subsutural, broadly arcuate,
European Journal of Taxonomy 773: 19–60 (2021)
50
Fig. 11. Radulae of species of Sibogasyrinx Powell, 1969 and Leucosyrinx Dall, 1889. A–B. Sibogasyrinx
clausura sp. nov., MNHN-IM-2009-16763. A. Central part of radula with fully formed teeth.
B. Anteriormost part of radula at bending plane; marginal teeth at the right (black arrows) are unfolded
due to mechanical impact. C. Leucosyrinx verrillii (Dall, 1881), French Guiana, MNHN-IM-2013-56288
(shell seen on Fig. 13C). D. Leucosyrinx sp. F, Coral Sea, MNHN-IM-2007-17846 (shell seen on Fig.
13K). E. Leucosyrinx sp. G, Papua New Guinea, MNHN-IM-2009-17089 (shell seen on Fig. 13D).
F. Leucosyrinx sp. C, Solomon Islands, MNHN-IM-2007-42503 (shell seen on Fig. 13J).
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
51
slightly darker subsutural ramp, protoconch light tan. Periostracum smooth, retained between cords and
ribs.
n = 1; MNHN-IM-2009-16763). Male. Penis obliquely truncated at tip with short, large,
conical papilla occupying entire anterior part of penis, surrounded by circular fold. Eyes present. Proboscis
proboscis base muscular. Salivary glands small, not fused, with very long ducts that run within walls of
beyond proboscis base, radular sac lying outside proboscis. Venom gland very large, thick and strongly
convoluted, very constricted before opening into oesophagus in region of nerve ring. Muscular bulb
moderately large.
n = 1; MNHN-IM-2009-16763, AL 10.6 mm) (Fig. 11A–B). Relatively short, comprising
m
and long, narrow, sharply-pointed cusp. Anterior margin overlapped by preceding row, posterior margin
becoming trough-shaped with weakly thickened edges during maturation, folded longitudinally when
fully formed, with both margins overlapping at tooth tip. The resulting folded tooth is moderately broad
with a sharp pointed tip, border between margins appears as a narrow groove along anterior edge. Tooth
folding occurs within 15–16th row of teeth (counting from rear).
Remarks
The other specimens are very similar to the holotype in shell shape and sculpture. In some specimens
there is very indistinct spiral striation on the subsutural ramp and a single very weak spiral cord may
even be present.
This species is most similar to Sibogasyrinx maximei
narrower subsutural axial wrinkles (50 vs 43 on last whorl and 42 vs 34 on penultimate one) and less
numerous (40 vs 50) spiral cords on the last whorl, the holotypes of both species being of almost the
same size. Nevertheless, S. maximei sp. nov. is only represented in our material by a single specimen
Distribution
This species occurs in the Solomon Islands, New South Wales and on the Coriolis Bank in the Coral Sea,
at 762–1060 m. This bathymetric range corresponds to a single haul in the Solomon Islands, while all
other specimens were collected at 960–1000 m, indicating a narrower bathymetric range for the species.
Sibogasyrinx pagodiformis sp. nov. (= PSH 5)
urn:lsid:zoobank.org:act:DAA613D6-5AB6-4F89-A0AD-0C718A60307E
Figs 8F, 12A–D
Diagnosis
Shell small, reaching 29 mm in length, pagodiform, with strongly concave subsutural ramp bearing two
low spiral cords and distinct subsutural nodules. Axial sculpture of distinct opisthocline ribs extending
to shell base, 10–11 per whorl. Spiral sculpture of closely spaced cords on and below shoulder. Radula
lacking central tooth, marginal teeth trough-shaped with thickened edges.
European Journal of Taxonomy 773: 19–60 (2021)
52
Etymology
Material examined
Holotype
3, stn CP2789; MNHN-IM-2009-16825.
Other material (sequenced)
MNHN-IM-2009-11327.
Description
(holotype). SL 28.8 mm, AL (with canal) 15.2 mm, AL (without canal) 9.9 mm, SW
11.9 mm.
(holotype). Moderately thick, except for fragile, partially chipped outer aperture lip; pagodiform,
with moderately high spire and moderately long, narrow siphonal canal inclined to abaxially left.
Protoconch missing (rendering exact an whorl count and protoconch measurements impossible), upper
teleoconch whorls eroded. Teleoconch whorls rapidly increasing and strongly angled at shoulder, more
than 8 remaining. Suture shallowly impressed, subsutural ramp broad, concave. A row of distinct, sharp,
subsutural nodules on upper teleoconch whorls. Nodules less distinct on last three whorls and situated
on short, slightly raised, prosocline axial ribs. Number of ribs on uppermost whorls indeterminate due
22 on penultimate and 26 on last whorl. Subsutural ramp smooth except for nodules and two low, indistinct
subsutural cords. Shoulder with pronounced, thickened, opisthocline axial ribs on all teleoconch whorls.
Axial ribs on upper teleoconch whorls situated just above suture, extending to shell base on last whorl,
number of ribs constant, 11–12 per whorl. Ribs intersected by closely spaced, rounded spiral cords,
number increasing from 5 on uppermost remaining whorl to 10 on penultimate whorl. Last whorl with
total of 38 cords, including 18 on canal. Cords narrow, their intervals 0.5–1.5 times cord width. Shell
base rapidly narrowing towards narrow, moderately long siphonal canal. Aperture narrow, constricted
posteriorly, with thin, moderately broad parietal callus, outer lip partially broken, convex on subsutural
ramp, with rounded angle at shoulder, convex below shoulder and weakly concave at transition to canal.
darker base, glossy. Periostracum thin, persisting in some cord intervals.
n = 1; MNHN-IM-2009-11327). Female. Proboscis relatively short, straight, cylindrical.
Proboscis retractors large, in two bundles attached at proboscis base. Buccal mass within proboscis,
rather long, about half length of proboscis, radular sac opening on right. Venom gland very thick, not
long, opening with very short constriction just posterior to nerve ring. Venom bulb moderately large,
elongate.
n = 2; MNHN-IM-2009-16825, AL 9.9 mm, IM-2009-11327, AL ca 9 mm). Relatively short,
tooth length 150
when mature (Fig. 8F), with slightly thickened edges and sharp pointed tips. Teeth initially forming as a
folding of each tooth on 10–11th row, counting from rear.
Remarks
The second live collected specimen has a broken shell base. In sculpture pattern it is very similar to the
holotype.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
53
The species is distinct from all congeners in having a relatively broad, pagodiform shell with a shorter
canal. Conchologically the species is quite similar to Comitas paupera (Watson, 1881), a highly variable
Fig. 12. A–D. Sibogasyrinx pagodiformis sp. nov. A–B. Solomon Islands, holotype, MNHN-
IM-2009-16825, SL 28.8 mm. C–D. Vanuatu. MNHN-IM-2009-11327, SL 24.3 mm. E–H. Sibogasyrinx
elbakyanae Kantor, Puillandre & Bouchet sp. nov., holotype, Solomon Islands, MNHN-IM-2009-16834,
SL 27.3 mm. H. Enlarged fragment of penultimate and antepenultimate whorls. A–G to same scale.
European Journal of Taxonomy 773: 19–60 (2021)
54
the new species also strongly resembles a miniature copy of Leucosyrinx luzonica (Powell, 1969) (see
Kantor et al.
Another species of Leucosyrinx from the Solomon Islands, Leucosyrinx sp. D, is similar to the new
Distribution
This species was found in the Solomon Islands and Vanuatu at 1250–1262 m.
Sibogasyrinx elbakyanae Kantor, Puillandre & Bouchet sp. nov. (= PSH 4)
urn:lsid:zoobank.org:act:F9B671D9-4F84-4DF4-BAA0-3EF30AF06512
Fig. 12E–H
Diagnosis
Shell small, reaching 27.5 mm in length, fusiform, subsutural ramp strongly concave with distinct,
broadly-spaced spiral cords and dense subsutural nodules. Axial sculpture of distinct opisthocline ribs
extending to shell periphery. Spiral sculpture of distinct cords on and below shoulder.
Etymology
Philippe Bouchet joins us in dedicating this species to honor Alexandra Elbakyan, creator of the famous
sci-hub portal.
Material examined
Holotype
stn CP2817; MNHN-IM-2009-16834.
Description (holotype)
SL 27.3 mm, AL (with canal) 15.0 mm, AL (without canal) 9.6 mm, SW 8.9 mm.
Moderately thick, except for fragile outer aperture lip, partially chipped; fusiform, with high
spire and long, narrow, straight siphonal canal. Protoconch eroded and partially decollated, thus
rendering whorl count and measurements impossible. Teleoconch whorls strongly angled at shoulder,
about 8 whorls in total. Suture deeply impressed, subsutural ramp moderately broad, strongly concave.
Subsutural region with a row of dense, distinct, sigmoidal axial ribs that correspond in shape to upper
parts of thickened growth lines and form small rounded nodules at intersections with spiral cords. Rib
counts on uppermost whorls not possible due to shell surface erosion, about 24 on antepenultimate whorl,
preserved teleoconch whorl and 5 on last three whorls, one immediately subsutural followed by another
with interval about three times cord width, then three more closely spaced ones separated from second
axial ribs. Shoulder with pronounced, thickened, opisthocline axial ribs on all teleoconch whorls, about
12 on upper whorls, including penultimate one and 14 on last whorl. Ribs extending to abapical suture
on spire whorls, intersected by spiral cords, uppermost 2–3 very thin and closely spaced followed by
5–8 more broadly spaced. Last whorl with about 35 cords on shoulder and below, including 15 on canal.
Cords narrow and separated by intervals 1–3 times the width of cords on axial ribs on shoulder, more
broadly spaced on shell base and canal. Shell base gradually narrowing towards narrow, moderately
long, nearly straight siphonal canal. Aperture narrow, constricted posteriorly, with moderately broad,
thin parietal callus, outer lip partially broken, with rounded angle at shoulder, weakly convex below
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
55
shoulder and weakly concave at transition to canal. Anal sinus shallow, subsutural, broadly arcuate.
persisting in intervals between cords and ribs.
Adult male, penis long, gradually narrowing towards tip, obliquely truncated at tip with a
small but rather long conical papilla, surrounded by circular fold. Proboscis not long, conical, proboscis
Single oval accessory salivary gland present at level of anterior part of proboscis. Venom gland large,
thick and strongly convoluted, very constricted before opening into oesophagus within nerve ring.
Muscular bulb moderately large.
Minute, comprising about 15 rows of teeth. Radula length 650
width up to 100
trough-shaped. Due to poor preparation it was impossible to study tooth morphology in detail.
Remarks
This species is most similar to Sibogasyrinx losa
shorter canal, fewer, more distinct axial ribs on the shoulder (35 vs 46 on last whorl), and thinner, more
widely spaced spiral cords. In shell outline the new species resembles some species of Leucosyrinx,
particularly Leucosyrinx verrillii
shoulder nodules and radular morphology.
Distribution
Known only from the type locality.
Discussion
Conchologically, the species of Sibogasyrinx described herein are rather heterogeneous on one hand,
while on the other, some species are very similar to each other (e.g. S. cf. pyramidalis 1 and S. cf.
pyramidalis 2; S. losa and S. lolae sp. nov.). Furthermore, species of Sibogasyrinx
similar in shell shape and sculpture to species of Leucosyrinx (Fig. 13); thus, based on shell alone, it
is not possible to reliably attribute the species to either genus. Comparisons of species of Sibogasyrinx
with similar species of Leucosyrinx (still mostly undescribed) are provided in the remark sections. This
similarity in shell morphology between unrelated genera suggests that additional species of Sibogasyrinx
Leucosyrinx. Whether the
occurrence of similar shells in taxa that potentially diverged more than 130 mya (Abdelkrim et al. 2018)
is linked to the retention of ancestral state characters or to convergence remains to be determined.
Anatomically, species of Sibogasyrinx
radula characters. Three radula types were observed within the species of this genus, which is uncommon
for Conoidea, where intrageneric variation in radula form is generally limited. Two of these types are
Sibogasyrinx. The Type 1 of radula is found in S. cf.
pyramidalis 1 and 2 and S. subula sp. nov. (Fig. 4). Here, fully matured marginal teeth are similar to
including Leucosyrinx; Fig. 11C–F). The developing teeth in the younger, posterior part of the membrane
(Fig. 4A) are trough-shaped, with the anterior edge (Fig. 4A, ae) of the tooth poorly sclerotized and
attached to the membrane, while the posterior edge (Fig. 4A, pe) becomes progressively thicker along
the membrane, lifting from the membrane and drawing closer to the anterior edge, thus making the tooth
fold longitudinally. The fully formed and immature marginal teeth in the same radula membrane look
European Journal of Taxonomy 773: 19–60 (2021)
56
Fig. 13. Shells of species of Leucosyrinx Dall, 1889. A–B. Leucosyrinx verrillii (Dall, 1881), French
Guiana, MNHN-IM-2013-56840, SL 25.1 mm. C. L. verrillii, French Guiana, MNHN-IM-2013-56288,
SL 31.9 mm (radula seen on Fig. 11C). D–E. Leucosyrinx sp. G. D. Papua New Guinea, MNHN-
IM-2009-17089, SL 32.2 mm (radula seen on Fig. 11E). E. Bismarck Sea, MNHN-IM-2013-19689,
SL 47.3 mm. F. Leucosyrinx sp. D, Solomon Islands, MNHN-IM-2009-16769, SL 25 mm. G. Leucosyrinx
sp. E, Madagascar, MNHN-IM-2009-16897, SL 46.1 mm. H. Leucosyrinx sp. A, Philippines, MNHN-
IM-2007-42445, SL 34.1 mm. I. Leucosyrinx sp. B, Solomon Islands, MNHN IM-2009-16764, SL 29.9
mm. J. Leucosyrinx sp. C, Solomon Islands, MNHN-IM-2007-42503, SL 41.2 mm (radula seen on
Fig. 11F). K. Leucosyrinx
(radula seen on Fig. 11D). All shells to same scale.
KANTOR YU.I. & PUILLANDRE N., Revision of Sibogasyrinx (Conoidea: Cochlespiridae)
57
The Type 2 of radula is recorded in several species forming a well-supported clade: Sibogasyrinx sangeri,
S. losa, S. lolae sp. nov., S. maximei sp. nov. and S. clausura sp. nov. While the central teeth are well
thickened anterior edge (Fig. 8B, ae) and a not thickened posterior edge (Fig. 8B, pe). Then, mid-way
along the membrane, the teeth are abruptly folded longitudinally. The resulting folded tooth (Fig. 8A) is
moderately broad (half the width of the original unfolded plate) with a sharply pointed tip. The border
between both edges is evident as a narrow slit along the anterior margin of the tooth. Although this
folded pattern of the marginal teeth is rather stable and evidently remains unchanged in most of the
radulae studied, the teeth may become unfolded due to mechanical impact (on Fig. 11B such an unfolded
tooth on the working part of the membrane is marked by a black arrow).
Finally, the Type 3 of radula was found in Sibogasyrinx pagodiformis sp. nov. (Fig. 8F). Here the
marginal teeth are more similar to those of radula type 1, with a more thickened posterior edge, although
they retain a clear trough shape when fully formed. The central teeth are absent, which is unique for
Cochlespiridae in general and Sibogasyrinx in particular.
Sibogasyrinx pagodiformis sp. nov. has a broad, pagodiform shell, which is rather atypical for the genus.
Nevertheless, its position in Sibogasyrinx
anterior foregut. The venom gland in this species opens into the oesophagus just posterior to the nerve
ring, as in the other species of Sibogasyrinx examined (this character is a symplesiomorphy of the
Cochlespiridae; Abdelkrim et al. 2018), while in other families of Conoidea the venom gland passes
posterior to the radular sac.
The foregut anatomy is similar in the other species of Sibogasyrinx
bundles attached to the most posterior part of the proboscis sheath in S. pagodiformis sp. nov., S. lolae
sp. nov., S. subula sp. nov. and S. cf. pyramidalis, or absent as in S. elbakyanae sp. nov., S. clausura
sp. nov., S. losa). The buccal mass is usually situated posterior to the proboscis, but may be partially
(S. clausura sp. nov.) or completely (e.g., S. lolae sp. nov.) contained within the proboscis. The salivary
glands may be separate or fused. An accessory salivary gland was found in Sibogasyrinx cf. pyramidalis 1
and S. lolae
present in some other species as well.
Eyes may be present (S. losa, S. clausura sp. nov.) or absent (S. cf. pyramidalis, S. subula sp. nov.), and
this showed no relation to the bathymetric range of the species.
Presently, species of Sibogasyrinx
Sea and the Philippines to Indonesia (type species, S. pyramidalis) and south to Vanuatu, New Caledonia
and Australia. Despite a number of cruises led by MNHN at appropriate depths in the Indian Ocean, as
well as in Fiji, Tonga and French Polynesia, Sibogasyrinx was not recorded there, as well as in Japan
(8 species recorded) and Papua New Guinea (4 species), but it is not clear at present whether this is
a result of more intense sampling in these areas or a genuine distributional feature. The species are
lower bathyal (800–2000 m). Only one species (S. losa) was recorded as shallow as 150 m, but it
numerous samples, the bathymetric range is rather large, spanning several hundred meters, although, as
mentioned earlier, some species seem to have more limited bathymetric ranges.
European Journal of Taxonomy 773: 19–60 (2021)
58
Acknowledgments
Specimens were obtained during research cruises and expeditions organized by the MNHN and Pro-
Natura International as part of the Our Planet Reviewed program, and by the MNHN and the Institut de
Recherche pour le Développement as part of the Tropical Deep-Sea Benthos program (DongSha 2014
and NanHai 2014 in the South China Sea; AURORA 2007 and PANGLAO 2005 in the Philippines;
SALOMON 2 and SALOMONBOA 3 in the Solomon Islands; BIOPAPUA, KAVIENG 2014,
MADEEP and PAPUA NIUGINI in Papua New Guinea; BOA1 and SANTO 2006 in Vanuatu; EXBODI,
Madagascar; GUYANE 2014 in French Guiana), and we are grateful to numerous cruise leaders and
co-PIs: Wei-Jen Chen, Philippe Bouchet, Marivene Manuel Santos, Laure Corbari, Stephane Hourdez,
Ludivina Labe, Peter Kee Lin Ng, Karine Olu-Le Roy, Bertrand Richer de Forges and Sarah Samadi.
All expeditions operated under the regulations then in force in the countries in question and satisfy the
conditions set by the Nagoya Protocol for access to genetic resources. We thank Francesco Criscione and
Anders Halan for providing additional material from the Australian Museum. The authors thank Virginie
Héros, Barbara Buge, Philippe Maestrati, Julien Brisset (MNHN) and Alexander Fedosov (IEE RAS)
for their help in curating the vouchers and the associated data. Dr Roberto Ardovini kindly provided the
photos of the holotype of Sibogasyrinx losa. The photos of the holotype of Surcula pyramidalis were
provided by Jeroen Goud (Naturalis Biodiversity Center) and Dr Kazunori Hasegawa. We thank Elena
Mekhova from IEE RAS for assisting with shell photography. Our special thanks to Dr David Herbert,
who edited and corrected the manuscript, as well as Dr Philippe Bouchet and Dr Alexander Fedosov for
constructive reviews.
This study was conducted using Joint Usage Center “Instrumental methods in ecology” at the IEE RAS.
We thank A. Neretina and A. Nekrasov for their friendly and helpful assistance with the SEM facilities.
The project has received funding from the European Research Council (ERC) under the European
Union’s Horizon 2020 research and innovation programme (grant agreement no. 865101) to NP.
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Manuscript received: 6 April 2021
Manuscript accepted: 2 July 2021
Published on: 28 September 2021
Topic editors: Rudy C.A.M. Jocqué, Tony Robillard
Section editor: Thierry Backeljau
Desk editor: Danny Eibye-Jacobsen
Printed versions of all papers are also deposited in the libraries of the institutes that are members of
the EJT consortium: Muséum national d’histoire naturelle, Paris, France; Meise Botanic Garden,
Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural
Sciences, Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis
Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid,
Spain; Real Jardín Botánico de Madrid CSIC, Spain; Zoological Research Museum Alexander Koenig,
Bonn, Germany; National Museum, Prague, Czech Republic.