# Snails in depth: Integrative taxonomy of Famelica, Glaciotomella and Rimosodaphnella (Conoidea: Raphitomidae) from the deep sea of temperate Australia

Francesco Criscione, Anders Hallan, Nicolas Puillandre, Alexander Fedosov

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Francesco Criscione, Anders Hallan, Nicolas Puillandre, Alexander Fedosov. Snails in depth: Integrative taxonomy of Famelica, Glaciotomella and Rimosodaphnella (Conoidea: Raphitomidae) from the deep sea of temperate Australia. Invertebrate Systematics, 2021, 35 (8), pp.940-962. 10.1071/IS21008. hal-03477619

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Snails in depth: Integrative taxonomy of Famelica, Glaciotomella and Rimosodaphnella (Conoidea: Raphitomidae) from the deep sea of temperate Australia

<table><tr><td>Journal:</td><td><i>Invertebrate Systematics</i></td></tr><tr><td>Manuscript ID</td><td>IS21008.R2</td></tr><tr><td>Manuscript Type:</td><td>Research paper</td></tr><tr><td>Date Submitted by the Author:</td><td>n/a</td></tr><tr><td>Complete List of Authors:</td><td>Criscione, Francesco; Australian Museum Research Institute, Life and Geoscience<br>Hallan, Anders; Australian Museum Research Institute, Life and Geoscience<br>Puillandre, Nicolas; Museum National d'Histoire Naturelle, Systematique et Evolution<br>Fedosov, Alexander; Russian Academy of Sciences</td></tr><tr><td>Keyword:</td><td>Australia, mitochondrial DNA, molecular systematics, Gastropoda, morphology, species delineation</td></tr></table>

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## Short summary

Distribution patterns of deep-sea benthic gastropods remain poorly known, yet such knowledge is crucial to approaching their systematics and understanding their diversity. With focus on the raphitomid genera *Famelica*, *Glaciotomella* and *Rimosodaphnella*, we combine genetic, shell, anatomy and distribution data to identify four new species from Australia and beyond, which we here describe. We identify the set of morphological characters shared by congeneric species and highlight patterns of rarity and endemicity in some species. Our findings suggest that radula and protoconch features are good proxies for genus-level phylogenetic relationships. We discuss the significance of ecological factors (such as choice and availability of preys) as potential drivers in the evolution of this hyper-diverse group of gastropods.
1 Snails in depth: Integrative taxonomy of *Famelica*, *Glaciotomella* and *Rimosodaphnella*

2 (Conoidea: Raphitomidae) from the deep sea of temperate Australia

3

4

5 Francesco Criscione A*, Anders Hallan A, Nicolas Puillandre B, Alexander Fedosov C

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7

8

9 A - Australian Museum Research Institute, 1 William Street, Sydney NSW 2010, Australia

10 B - Institut Systématique Evolution Biodiversité (ISYEB), Muséum National d'Histoire

11 Naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles. 57 rue Cuvier, CP 26,
12 75005 Paris, France

13 C - A.N. Severtsov Institute of Ecology and Evolution of Russian Academy of Sciences,
14 Leninski Prospect 33, Moscow 119071, Russia

15 * Corresponding author: francesco.criscione@austmus.gov.au

16

17

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22

23

24

25

26 Running Head: Integrative taxonomy of three raphitomid genera

27

28

29

30 Keywords: shell, radula, deep sea, mtDNA, species delimitation, biodiversity

31
32 Abstract

33 The deep sea of temperate south-eastern Australia appears to be a 'hotspot' for diversity
34 and endemism of conoidean neogastropods of the family Raphitomidae. Following a series
35 of expeditions in the region, a considerable amount of relevant DNA-suitable material has
36 become available. A molecular phylogeny based on this material has facilitated the
37 identification of diagnostic morphological characters, allowing the circumscription of
38 monophyletic genera and the introduction of a number of new genus-level taxa. Both
39 named and new genera are presently being investigated through integrative taxonomy, with
40 the discovery of a significant amount of undescribed species. As part of this ongoing
41 investigation, this study focuses on the genera *Famelica* Bouchet & Warén, 1980,
42 *Glaciotomella* Criscione, Hallan, Fedosov & Puillandre, 2020 and *Rimosodaphnella*
43 Cossmann, 1914. We subjected a comprehensive mitochondrial DNA dataset of
44 representative deep-sea raphitomids to the species delimitation methods ABGD and ASAP,
45 which recognised 18 and 15 primary species hypotheses (PSHs) respectively. Following
46 additional evaluation of shell and radular features, as well as examination of geographic and
47 bathymetric ranges, nine of these PSHs were converted to secondary species hypotheses
48 (SSHs). Four SSHs (two in *Famelica* and two in *Rimosodaphnella*) were recognised as new to
49 science and their formal descriptions are provided herein.
50 Introduction

51 Turriform Conoidea is the most diverse group of marine molluscs (Horton et al. 2019), with
52 considerable species diversity occurring in the deep sea, where it is often coupled with low
53 abundance (Sysoev 1997; Kantor et al. 2008; Bouchet et al. 2009). One of the total fifteen
54 families of the group, Raphitomidae Bellardi, 1875, is among the most poorly studied
55 (Bouchet et al. 2011), notably in the deep sea (Criscione et al. 2021). Because of their high
56 diversity, low abundance and morphological complexity combined, the systematics of many
57 deep-sea genera and species remains tentative, with a large proportion of taxa awaiting
58 description (Bouchet et al. 2009). To further complicate matters, most of the taxonomic
59 accounts available for groups below family level are primarily based on shell morphology
60 (Sysoev and Ivanov 1985; Sysoev and Kantor 1986, 1987; Sysoev 1988, 1990, 1996a, b;
61 Bouchet and Sysoev 1997; Sysoev 1997; Bouchet and Sysoev 2001; Sysoev and Bouchet
62 2001; Morassi and Bonfitto 2006; Kantor et al. 2012; Stahlschmidt and Chino 2012; Bonfitto
63 and Morassi 2013; Morassi and Bonfitto 2015; Kantor et al. 2016) and only three studies do
64 include molecular data (Fassio et al. 2019; Russini et al. 2020; Criscione et al. 2021). This is
65 partly due to a lack of study material, particularly that suitable for molecular analysis
66 (Criscione et al. 2021). Due to widespread shell homoplasy, some of the earlier studies have
67 incorrectly attributed a considerable number of unrelated species to very few raphitomid
68 genera, colloquially termed 'dumpsters' (such as *Pleurotomella* Verril, 1872 and *Gymnobela*
69 Verrill, 1884) (Criscione et al. 2021). Based on their support as clades and on diagnostic
70 morphological characters, Criscione et al. (2021) demonstrated the non-monophyly of such
71 genus-level groups and, in constraining them, introduced a number of new genus-level taxa
72 and described their type species. The same study also revealed a multitude of putatively
73 undescribed species among deep-sea raphitomids of a temperate region roughly
74 corresponding to the central and eastern part of the South Australia realm (#26 of Costello
75 et al. 2017) and the southern part of the Tropical Australia and Coral Sea realm (#16 of
76 Costello et al. 2017) (Fig. 1). With nearly half of the currently accepted deep-sea genera (24
77 out of 50) and nearly a third (7) endemic, this area is a potential 'hot spot' for raphitomid
78 diversity (Criscione et al. 2021). Thanks to recent sampling efforts in the area (MacIntosh et
79 al. 2018; Williams 2018; O'Hara et al. 2020), relatively abundant raphitomid samples,
80 suitable for genetic analysis, are available. This material has allowed studies whose
81 methodological design for species delimitation transcends the mere conchological approach
of earlier taxonomical accounts and capitalises on the combination of morpho-anatomical
and molecular data (i.e. 'integrative taxonomy', Dayrat 2005; Will et al. 2005). As a result of
these studies on the Australian Raphitomidae, new species have been described for the
genera *Gladiobela* Criscione, Hallan, Puillandre & Fedosov, 2020 (5 species) and *Pagodibela*
Criscione, Hallan, Puillandre & Fedosov, 2020 (2) (Hallan et al. 2021) and of *Austrobela*
Criscione, Hallan, Puillandre & Fedosov, 2020 (4), *Austrotheta* Criscione, Hallan, Puillandre &
Fedosov 2020 (1), *Spergo* Dall, 1895 (4) and *Theta* Clarke, 1959 (2) (Criscione et al., in press).
Based on a larger sampling size, including specimens from beyond Australian waters, this
study aims to revisit species delimitation and taxonomy in three genera: *Famelica* Bouchet
& Warén, 1980, *Glaciotomella* Criscione, Hallan, Puillandre & Fedosov 2020 and
*Rimosodaphnella* Cossmann, 1916. In the analysis of Criscione et al. (2021), these genera
(along with *Veprecula* Melvill, 1917) constituted a monophyletic group (clade 'A' of fig. 2 in
Criscione et al. 2021). Formal descriptions are here presented for newly recognised species.
Furthermore, revised genus diagnoses and new anatomical and morphological data are
introduced for both established and new taxa, which are discussed in terms of their
diagnostic utility at the genus level. Finally, geographic and bathymetric distributions are
presented and discussed for the taxa treated herein.
99 Materials and methods

100

101 Taxon sampling

102 The samples studied herein were selected among all Raphitomidae ethanol-preserved material from the malacological collections of the Australian Museum, Sydney (AMS), the South Australian Museum, Adelaide (SAMA) and the Muséum national d'Histoire naturelle, Paris (MNHN). All studied material has been collected off Australia during the expeditions IN2015_C01, IN2015_C02 (GAB) and IN2017_V03 (Tasman and Coral Seas), targeting (among other sites) a number of Commonwealth Marine Reserves (CMR), and other localities (mainly of the tropical and temperate Indian and Pacific Oceans), during a number of voyages that formed part of the Tropical Deep-sea Benthos programme of MNHN (expeditions.mnhn.fr; Fig. 1, Table S1).

110 111 As a result of ongoing systematic research on the Conoidea at the AMS and MNHN, several hundreds of (mostly unpublished) sequences of the mitochondrial gene cytochrome oxidase subunit 1 (*cox1*) were obtained (see methodology below) from a considerable number of largely undescribed raphitomid taxa. In order to assist with the selection of the study material, a pilot analysis was performed on a dataset including all raphitomid *cox1* sequences, using the neighbour-joining method (NJ) (Saitou and Nei 1987) implemented in MEGA 7 (Kumar *et al.* 2016). In particular, the dataset included *cox1* sequences of the holotypes for the type species of several deep-sea raphitomid genera, including *Glaciotomella investigator* Criscione, Hallan, Puillandre and Fedosov *et al.* 2020 (GenBank AN: MN983178). The dataset also encompassed *cox1* sequences of well-recognisable species, such as *Famelica tasmanica* Sysoev & Kantor, 1987, as well as of several Australian undescribed species included in the phylogeny of Criscione *et al.* (2021), such as those placed in the genus *Rimosodaphnella* therein. All *cox1* sequences of *Glaciotomella*, *Famelica* or *Rimosodaphnella* used in (Criscione *et al.* 2021) and all sequences that (in the results of the NJ analysis; Fig. S1) were more closely related to the species of these three genera than to any other raphitomid genus, were selected as ingroup and used in the molecular analysis described below (Table S1). Sequences representing 13 raphitomid species from 13 genera were selected as outgroups (Table S1). Their choice was based on the phylogeny of Criscione *et al.* (2021), containing many southern and south-eastern Australian Raphitomidae.
Among the ingroup specimens, morphological examination was only conducted on those collected in Australian waters. However, for samples outside Australia, examination of shell photographs was possible and was utilised when necessary and appropriate. Geographic and bathymetric data were available for all ingroup specimens. Geographic distributions were assessed with reference to marine biogeographic realms as delimited in Costello *et al.* (2017). According to Bouchet *et al.* (2008), when inferring bathymetric distributions of species from sampling depth intervals, only shallower depth values were considered, as there is no evidence that the species collected occurs beyond that value.

Molecular methods

Molecular work was performed in laboratories at two different institutions (AMS and MNHN). Unless otherwise stated, the same methodology was followed by both laboratories. DNA was extracted from small pieces of foot muscle by use of a Bioline Isolate II Genomic DNA extraction kit for animal tissue, following the standard procedure of the manual (AMS) or using the Epmotion 5075 robot (Eppendorf), following the recommendations by the manufacturer (MNHN). A fragment of *cox1* was amplified using the primer pairs LCO1490/HCO2198 for *cox1* (Folmer *et al.* 1994). PCR reactions were performed in volumes of 25 µl, containing 3 ng DNA, 1X Qiagen CoralLoad PCR Buffer, 2.5mM MgCl2, 0.25mM dNTP, 0.5mM of each primer, 0.5 µg/µl of BSA and 0.2 µl of Bioline MyTaq DNA polymerase. Amplification consisted of an initial denaturation step at 94°C for 4 min, followed by 37 cycles of denaturation at 94°C for 30 s, annealing at 50°C for 30 s, followed by extension at 72°C for 1 min. The final extension was at 72°C for 5 min. PCR products were purified and sequenced by the Macrogen (AMS) and Eurofins (MNHN) sequencing facilities. When necessary, chromatograms were manually corrected for misreads and forward and reverse strands were merged into one sequence file using CodonCode Aligner v. 9.0.1 (CodonCode Corporation, Dedham, MA). Sequence alignments were generated using MUSCLE as implemented in MEGA7 (Kumar *et al.* 2016). Sequences were deposited in GenBank (Table S1). Phylogenetic trees were generated using Maximum Likelihood (ML) and Bayesian inference (BI) methods. ML was performed using the program IQ-Tree v. 2.1.1 (Minh *et al.* 2020), including the implemented model-finder function (Kalyaanamoorthy *et al.* 2017) and ultrafast bootstrapping (BS, 1000 replicates) (Hoang *et al.* 2017). The BI analysis was performed in MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003) and included 2 runs of 10⁷
162 generations, with 4 chains each and a sampling frequency of one tree per 1000 generations.
163 Other parameters were set to default. After checking for convergence (ESS>200) with Tracer
164 (Rambaut et al. 2018), a consensus tree was calculated after discarding the first 25% trees as
165 burn-in. According to MrBayes manual (p. 94), a priori model testing was not performed,
166 and the GTR+G+I model was applied to the BI analysis. Nodal support was assessed as
167 Bayesian posterior clade probabilities (BPP). Pairwise genetic distances were calculated for
168 *cox1* sequences using the Kimura-2 parameter model (K2p, Kimura 1980) as implemented in
169 MEGA7 with the option 'pair-wise deletion of gaps'.

170

171 Morphological examinations

172 All studied samples consisted of animals and shells, from which they had been extracted
173 following the methodology described in Criscione et al. (2021). We studied shell morphology
174 and (when possible) internal anatomy, including radular morphology. Shells of sequenced
175 specimens were affixed to plasticine and positioned with their vertical axis parallel to the
176 observation plane. Each shell was then photographed from above using a digital SLR
177 camera. Maximum shell length (SL) and width (SW) were measured on digitised images
178 using the calibrated ruler tool in Adobe Photoshop CC v.20.0.6. Measurements were
179 rounded to the nearest 0.1 mm. The number of shell whorls was counted under a Leica MZ8
180 stereomicroscope, according to Bouchet and Kantor (2004). While it was possible to count
181 the number of teleoconch whorls for almost all studied specimens, protoconch whorls could
182 only be counted occasionally due to widespread erosion of the apex. Anatomical studies
183 were conducted on animals removed from ethanol and briefly rehydrated in distilled water.
184 Using standard dissection tools, the venom apparatus was excised and the radular sac
185 isolated and placed on a glass slide; during this dissection process, head-foot, mantle,
186 genital and (non-radula) foregut characters were examined where possible. After dissolution
187 in dilute commercial bleach, clusters of hypodermic teeth were rinsed repeatedly in distilled
188 water, then separated into individuals and ligament-connected pairs/smaller clusters.
189 Subsequently, the glass stub was affixed to a carbon adhesive placed on a 12 mm diameter
190 aluminium mount. All samples were imaged at Macquarie University, Sydney, using a
191 Phenom XL Scanning Electron Microscope.

192

193 Species delimitation
194 Firstly, the Automatic Barcode Gap Discovery (ABGD, Puillandre *et al.* 2012a;

195 https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb.html) was used with a p-distance model,

196 the relative gap width (X) set to 1 and the other parameters left to default.

197 Secondly, the Assemble Species by Automatic Partitioning (ASAP, Puillandre *et al.* 2020;

198 https://bioinfo.mnhn.fr/abi/public/asap/) was applied. Similar to ABGD, ASAP proposes

199 species partitions based on genetic distances only; however, ASAP screens all the genetic

200 distances from the lowest to the highest and merges sequences into "groups" that are
201 successively further merged until all sequences form a single group. At each merging step,

202 the partition is evaluated and given a score. At the end of the analyses, the 10 partitions

203 with the lowest scores are provided (the lower the ASAP-score, the better the partition).

204 Resulting ABGD and ASAP groups were considered primary species hypotheses, henceforth

205 referred to as PSHs. Following Puillandre *et al.* (2012b), conversion of PSHs to secondary

206 species hypotheses (SSHs) was conducted through comparative examination of

207 morphological characters as well as through evaluation of geographic and bathymetric data

208 and phylogenetic reconstruction based on the DNA sequences.

209 In particular, when converting an individual PSH to SSH, the occurrence of the following

210 conditions was assessed: (i) the PSH is a highly supported clade (BPP>0.98 and BS>90%), (ii)

211 all its constituent specimens share at least one distinctive morphological feature deemed

212 not to be polymorphic or ecophenotypic and without exhibiting intermediate forms, (iii) the

213 PSH maintains genetic or morphological divergence and/or bathymetric partitioning when

214 occurring in sympatry with another PSH. When available, species names were assigned to

215 SSHs based on the current taxonomy. New species names were introduced when no names

216 were available. Formal descriptions for these taxa are given in the systematics section

217 below.
218 Results

219 Molecular studies

220 Molecular analyses were based on a total of 46 *cox1* sequences (28 newly produced and 18 GenBank-sourced). Of the total sequences employed, 33 constituted the ingroup and the remaining 13 were used as outgroups.

223 In the vicinity of the barcode gap (i.e. between 0.8 and 3.3%, the highest intra-PSH genetic distance and the lowest inter-PSH genetic, respectively), the ABGD analysis of the *cox1* ingroup dataset consistently returned a primary partition with eighteen groups of sequences (PSHs). Among all PSHs, six (F1, F3, G1, G2, R1, R2) contained exclusively Australian samples, one (F2) included samples from Australian seas and beyond, while the remaining eleven (F4, FA, GA, R3-R5, RA-RE) encompassed sequences from outside Australian waters.

230 The best ASAP partition (i.e. with the lowest ASAP-score) returned 15 PSHs, 12 of which were identical to those resulting from ABGD and the remaining three corresponding to the combinations F1+F3, G1+G2 and R4+R5. The second-best ASAP partition included 14 PSHs, with RD and RE grouped (when compared to the 15-species ASAP partition). The ASAP-score of the alternative partitions were much higher and are thus ignored.

235 The BI and ML analyses generated trees (Fig. 2, S2) with similar topologies and no supported incongruences. Clades representing PSH-level relationships and above were generally well-supported, with very few exceptions. In both analyses, three major genus-level clades were retrieved, namely *Famelica* (BPP=1, BS=95%), *Glaciotomella* (BPP=1 and BS=95%) and *Rimosodaphnella* (BPP=0.98 and BS=94%). These three clades included five, three and ten ABGD PSHs respectively, all forming highly supported (in terms of nodal support) and well-differentiated (in terms of branch lengths) clades. The only exception was R1, which received no support from the BI analysis and moderate support from the ML analysis (BS=95%). ASAP PSHs R4+R5 and G1+G2 were highly supported by both analyses (Fig. 2, S2), while F1+F3 received no support.

245 Within the genus-level clades, there was no overlap between intra- and inter-ABGD-PSH distances. In the *Famelica* clade, the intra-PSH pairwise distance in *cox1* measured for the only PSH with more than one sample (F2) was 0.8%, with inter-PSH distances ranging from 5.2 to 18.2% (average=14.5%) (Table 1). The lowest inter-PSH distances were observed between F1 and F3 (the two ABGD-PSH grouped by ASAP). No intra-ABGD-PSH pairwise
distances could be measured for the *Glaciotomella* clade and the inter-PSH distances ranged from 3.3 to 16.0 % (average=9.7 %), with the lowest value recorded between G1 and G2 (the two ABGD-PSH grouped by ASAP). In the *Rimosodaphnella* clade, the intra-PSH pairwise distances in *cox1* ranged from 0 to 0.3 % (average=0.2 %) with inter-PSH distances ranging from 4.6 to 15.6 % (average=11.7 %) (Table 2). The lowest inter-PSH distances were observed between R4 and R5 (the two ABGD-PSH grouped by ASAP) and the highest intra-PSH distances were found within R2. The genetic distance between RD and RE, the two ABGD-PSH grouped in the second-best ASAP partition, is 6.7%.

## 259 Morphological studies

Shell morphology was examined for all sequenced material and internal anatomy, including radular morphology (when possible) for at least two sequenced specimens per PSH.

Detailed morpho-anatomical observations refer to PSHs examined herein, and do not include PSHs assigned with a letter suffix (i.e., FA, GA, etc.). Protoconchs of many specimens were considerably eroded and could therefore not be studied. For the remaining specimens, some sculptural detail could be inferred from heavily eroded protoconchs by careful examination using a microscope. However, owing to their very poor quality, these protoconchs are not figured herein. When observed, protoconch sculpture was highly conserved within each genus-level clade and could not be used to differentiate congeneric PSHs. While *Rimosodaphnella* PSHs possessed a protoconch with a typical raphitomid diagonally cancellate sculpture (Fig. 3C), *Famelica* and *Glaciotomella* PSHs exhibited a protoconch with median keel as the only spiral element (Fig. 3A-B). In these two genera, axial elements ('pillars' in Bouchet *et al.* 2011, p. 283) were present, that were restricted to the area below the keel in *Famelica* (Fig. 3A) and extended also above the keel in *Glaciotomella* (Fig. 3B).

All PSHs in the *Famelica* clade exhibited a gross shell morphology and whorl profile that were comparatively similar (Figs 4-5), with only one (F4) differing in its sculptural elements (prominent spiral keels). The radula was absent in all *Famelica* specimens studied.

Very similar shells were exhibited by PSHs in the *Glaciotomella* clade (Fig. 6), with shells of G1 and G2 being remarkably larger than GA and sharing the same sculptural pattern. While the radula of GA was not studied, G1 and G2 shared nearly identical hypodermic teeth (Fig. 6).
282 *Rimosodaphnella* PSHs varied notably in shell morphology (Figs 7-8), from comparatively large and broad (R2, Fig. 7D-F), to large and elongate (R1, Fig. 7A-C), to very small (e.g. R4 and R5, Fig. 8B-C). Generally, axial sculpture was more prominent than spiral sculpture in all shells of all PSHs, with two exceptions: RA, where axials were absent (Fig. 5C) and R2, where spiral and axial elements were equally prominent (Fig. 7G). The radula of both R1 and R2 (Fig. 10) consisted of similar straight, unbarbed hypodermic teeth that in R2 were shorter and broader.

289

290 Geographic and bathymetric distributions

291 The *Famelica* clade is recorded from at least four marine realms (sensu Costello *et al.* 2017) (Fig. 1A), the *Glaciotomella* clade from three (Fig. 1A) and the *Rimosodaphnella* clade from at least six marine realms (Fig. 1B). *Famelica* and *Rimosodaphnella* both exhibit a wide bathymetric range (respectively 398-4144 m and 350-4750 m), while the range of *Glaciotomella* (600-2474 m) is narrower (Fig. 11). In *Famelica*, the sister PSHs F1 and F3 are known only from off Australia, where they are found respectively at bathyal and abyssal depths of the east coast (Fig. 1A). F2 is known from two widely separated localities, both geographically (Fig. 1A) and bathymetrically (Fig. 11), namely in the temperate GAB (3389 m) and off the tropical New Caledonia (815 m). The only samples of the remaining *Famelica* PSHs, F4 (Papua New Guinea, PNG – 398 m; Figs 1A, 8) and FA (Taiwan – 999 m; Figs 1A, 11), were collected outside the seas of Australia.

302 Two of the three ABGD PSHs of *Glaciotomella*, G1 and G2, are found only off temperate Australia (albeit G1 at the temperate-tropical boundary) at comparable depths (2474 and 2007 m). The remaining PSH, GA is only known from a single record in shallower waters (600 m) off the Solomon Islands.

306 For *Rimosodaphnella*, virtually all records of the only two Australian PSHs, R1 and R2, are limited to a relatively restricted area off the east coast of Tasmania (Fig. 1B), where they occupy fairly disjunct bathymetric ranges (bathyal vs. abyssal; Fig. 11). The only sample of RA is found off the south coast of Chile (Fig. 1B) in much shallower waters (766 m, Fig. 11). All other *Rimosodaphnella* ABGD PSHs occur at lower latitudes in the warmer and shallower waters of the Philippines (R4 and R5; Figs 1B, 11), Melanesia (R3, RB, RC; Figs 1B, 11) and Polynesia (RD and RE; Figs 1B, 11).

313
314 PSH to SSH conversion

315 Comparative examination of the morphological, geographic and bathymetric data available
316 was employed to attempt the conversion of PSHs to SSHs. As generating morphological data
317 for most species with distribution outside Australian waters was beyond the scope of this
318 study, testing of seven PSHs (i.e. FA, GA, RA-RE), out of the total retrieved by ABGD and
319 ASAP, was not attempted and these are pending further sampling and taxonomic
320 investigation. As detailed below, nine of the PSHs retrieved by ABGD and ten of those
321 retrieved by ASAP satisfied at least one of the conditions described in the methodological
322 section. It must be noted that some of the PSHs consisted of one single sequence and, for
323 those, support value and intra-PSH genetic distance were not available. Also, the
324 information available on their geographic and bathymetric distribution was limited to a
325 single collection event. The evidence for PSHs to SSHs conversion is detailed below, where
326 congeneric PSHs are compared with each other according to their relationships as resolved
327 by the molecular analysis (Figs 2 and S2).

328 Two haplotypes of the *Famelica* clade represent two distinct ABGD PSHs (F1 and F3) but are
329 combined into one (F1+F3) by ASAP. While the comparatively low genetic distance
330 separating them (5.2%) could be interpreted as an indication of genetic connectivity
331 between two allopatric conspecific populations (Fig. 1A), this is not sufficient to convert
332 F1+F3 to SSH, as criteria (i) and (ii) are not met. In fact, F1+F3 is not statistically supported
333 (Figs 2, S2) and there is considerable difference between F1 and F3 in shell features (with F1
334 possessing a much more slender and darker shell; Fig. 4A, D-F). For these reasons F1 and F3
335 (as delimited by ABGD) are converted into two separate SSHs.

336 The only *Famelica* PSH including more than one sample, F2, exhibited a value of intra-PSH
337 genetic distance (0.8%) considerably lower than the distance value (13.8%) separating it
338 from its closest relative, F4. In addition, F2 and F4 differ significantly in shell features (Fig.
339 5C, F-G) and occur at different depths in distinct geographic areas (Figs 1, 11), which clearly
340 suggests that they should be treated as separate SSHs.

341 Two sequences in the *Glaciotomella* clade are treated as distinct PSHs by ABGD (G1 and G2)
342 but are combined into one highly supported PSH (G1+G2; BPP=1, BS=100%; Figs 2, S2) by
343 ASAP. G1 and G2 are found in allopatry, they are separated by comparatively low genetic
344 distance (3.3%), have very similar shells (Fig. 6A-B), and in terms of observed anatomical
345 characters differ only in G2 exhibiting a peculiar squamose osphradium. Whether the latter
346 character is of any taxonomic value or a result of a developmental anomaly is not clear,
347 therefore the evidence available is interpreted as supporting combination of G1 and G2 (as
348 suggested by ASAP) and subsequent conversion to a single SSH.

349 In the *Rimosodaphnella* clade, R1 was separated from its most closely related PSH (RA) by
350 comparatively large genetic distance (6.4%; Table 2) and could be differentiated by its more
351 pronounced cancellate shell sculpture (Fig. 7A-C). R1 occurs in a very distant realm (Fig. 1B)
352 at a different depth (Fig. 11). For these reasons, and despite the comparatively low support
353 (BPP=0.71, BS=96%) received in the molecular analysis (Figs 2, S2), R1 is converted into SSH.
354 R2 fulfils all criteria necessary to be converted to SSH. This PSH is separated from its closest
355 relatives (R1 and RA) by high genetic distance (9.3 and 6.4%; Table 2), far higher than its
356 intra-PSH genetic distance (0.3%). It has a notably different shell (Fig. 7D-F) and it is found in
357 sympatry with R1 in the South Australian realm (Fig. 1B) at deeper sites (Fig. 11).

358 R3, sister to the R1+RA+R2 clade, is separated from each of these by high genetic distance
359 (Table 1) and differs considerably in shell shape and sculpture (Fig. 8E). It is allopatric to RA,
360 R1 and R2 (Fig. 1B) and is found at much shallower depth than the two latter PSHs (Fig. 11).
361 Due to its marked genetic and morphological distinctiveness and distinct bathymetric
362 distribution, R3 is converted to SSH.

363 R4 and R5 were recognised by ABGD only, while ASAP combined them into a single PSH
364 (R4+R5). Their members exhibit very similar shells (Fig. 8B-C) and are separated by the
365 lowest inter-PSHs distance (4.6%) measured for PSHs of *Rimosodaphnella*. They also occur
366 sympatrically (Fig. 1B) and at comparable depth (Fig. 11). As no genetic, morphological,
367 geographic or bathymetric criterion can be applied to reliably distinguish them, they are
368 here combined (as indicated by ASAP) into a single SSH.

369

370 Assigning names to SSHs

371 A search was conducted for all names available and potentially applicable to the nine SSHs
372 resulting from the conversion process described above. By consulting the relevant literature
373 on Raphitomidae (Dall 1881; Watson 1881; Verrill 1884; Dautzenberg and Fischer 1896;
374 Schepman 1913; Barnard 1963; McLean and Poorman 1971; Bouchet and Warén 1980;
375 Sysoev and Kantor 1987; Sysoev 1990, 1996b; Figueira and Absalão 2012; Bonfitto and
376 Morassi 2013; Criscione et al. 2021) and by comparison of molecular and morphological
377 data available on type specimens with the data generated on sequenced specimens, we
378 found five names applicable to five SSHs. One SSH, G1+G2, including the holotype of G.

379 *investigator*, could be readily assigned to this species. As the constituent specimens of F3,

380 F4, R3 and R4+R5 shared shells that closely resembled the holotypes of respectively

381 *Famelica tasmanica* Sysoev & Kantor 1987 (Fig. 4D), *Veprecula polyacantha* Stahlschmidt,

382 Chino & Kilburn 2012 (Fig. 5G), *Rimosodaphnella solomonensis* Bonfitto & Morassi 2013 (Fig.

383 8D) and *R. brunneolineata* Bonfitto & Morassi 2013 (Fig. 8A), these SSH were attributed to

384 these taxa. This required the formal transfer (as hereby proposed) of the latter species to

385 *Famelica* as *Famelica polyacantha* (Stahlschmidt, Chino & Kilburn, 2012) n. comb. As no

386 available names could be found for the remaining four SSHs, these were assigned to new

387 taxa: namely *Famelica turritelloides* n. sp. (F1), *F. acus* n. sp. (F2), *Rimosodaphnella*

388 *guraradara* n. sp. (R1) and *R. truvana* n. sp. (R2). Formal taxonomic descriptions of these

389 newly recognised species are provided below.
390 Systematics

391 General remarks

392 If not stated otherwise, holotypes are dissected ethanol-preserved specimens, and all
393 systematic descriptions are based on the holotype. Shell whorls counts (approximated to
394 one decimal unit) are reported with reference to intact whorls only. When applicable, the
395 expression 'at least' is used in combination with the whorl count to indicate potential
396 additional missing whorls that could not be counted. Shell and head-foot colouration
397 reported in the descriptions is based on observations performed prior to fixation and thus
398 may not be fully reflected in the illustrations provided (Figs 3-8).

399 Measurements of radular features, mainly the length of the adapical opening and the dorsal
400 blade, are given as ratios of the length of the shaft. The 'shaft' is here defined as the entire
401 length of the tooth minus the base, where there is a notable swelling and angulation that
402 clearly differentiates it from the comparatively thin-walled hypodermic, rolled structure.
403 This is done to ensure consistency with the terminology used in Criscione et al. (2021).

404

405 Superfamily Conoidea Fleming, 1822

406 Family Raphitomidae Bellardi, 1875

407 Genus *Famelica* Bouchet & Waren, 1980

408

409 Type species: *Pleurotomella catharinae* Verrill & S. Smith [in Verrill], 1884 by subsequent
410 designation by Bouchet and Warén (1980).

411

412 Diagnosis

413 Shell narrowly fusiform to turriform with very high spire, thin- or very thin-walled, often
414 translucent. Protoconch pagodiform, multispiral, of four to five dark orange whorls, three to
415 four of which bearing a median keel, with orthocline axial pillars below the keel and no
416 sculpture above. Teleoconch of up to seven evenly convex, uniformly dark orange, cream to
417 white transparent whorls. Subsutural ramp indistinct, or well-pronounced, steep to very
418 steep, narrow to moderately wide. Shell base convex, often clearly demarcated from long,
419 slender siphonal canal with deep to moderate concavity. Spiral sculpture of densely set
420 rounded subequal cords, or (rarely) of a pair of sharp prominent median keels. Axial
421 sculpture of very fine collabral growth lines. Aperture elongate, a half to less than one third
422 of shell length. Outer lip thin, finely sculptured; inner lip smooth, with or without very thin
callus. Anal sinus moderately deep, U-shaped. Radula, and venom apparatus absent. Animal
423 with broad to long, narrow funnel; cephalic tentacles moderately short, cylindrical to weakly
424 tapering, with small eyes situated at their lower portion not recorded. Rhynchocoel
425 capacious.
426

427

428 Remarks

429 *Famelica* is characterised by the combination of: (a) a multispiral keeled protoconch with
430 axial pillars below the keel (this study, Fig. 3A; Bouchet and Warén 1980, p. 88-90, figs 277-
431 281; Figueira and Absalão 2012, fig. 4), (b) a very slender, thin-walled and spirally sculptured
432 teleoconch (this study, Figs 4-5; Dall 1889, p. 117, pl. 10, fig. 12; Barnard 1963, fig. 1b;
433 Bouchet and Warén 1980, figs 186-189; Sysoev and Kantor 1987, figs a-d; Sysoev 1990, pl. 2,
434 fig. 8; 1996b, figs 28-29; Figueira and Absalão 2012, fig. 3) and (c) absence of radula,
435 proboscis and venom gland (Criscione et al. 2021). This combination of morpho-anatomical
436 characters is unique among the Raphitomidae (Criscione et al. 2021). The distinctive
437 protoconch morphology of *Famelica* is shared by only one other deep-sea raphitomid genus,
438 the monotypic *Aliceia* Dautzenberg and Fischer, 1897 (Dautzenberg and Fischer 1897, p.
439 182, pl. 4, fig. 16; Bouchet and Warén 1980, p. 90, fig. 230; Figueira and Absalão 2012, p. 5,
440 fig. 2). Although its type species, *A. aenigmatica* Dautzenberg and Fischer, 1897, exhibits a
441 very distinctive shell, with a labial process and a deep umbilicus (Dautzenberg and Fischer
442 1897, p. 182, pl. 4, fig. 15, 17-18; Bouchet and Warén 1980, p. 90, fig. 190; Figueira and
443 Absalão 2012, p. 5, fig. 2), its overall shape and whorl profile are not too dissimilar from
444 those of some species of *Famelica* (e.g. *F. bitrudis* (Barnard, 1963). No anatomical details are
445 available for *Aliceia*, including information on the presence of a radula. Their remarkable
446 similarity may suggest a close phylogenetic relationship between *Aliceia* and *Famelica*.
447 However, testing of this hypothesis depends on the availability of live-collected material of
448 *A. aenigmatica* for molecular and anatomical investigation.

449 Prior to this study, *Famelica* included ten Recent species: the type species *F. catharinae*
450 (Verrill & S. Smith [in Verrill], 1884) (Fig. 4B), *F. monoceros* (R. B. Watson, 1881), *F.*
451 *mirmidina* (Dautzenberg & H. Fischer, 1896), *F. monotropis* (Dautzenberg & H. Fischer, 1896)
452 (Bouchet and Warén 1980 figs 187, 188 and 189 respectively) and *F. scipio* (Dall, 1889) (Dall
453 1889 pl. 10, fig. 12) from the Atlantic; *F. bitrudis* (Barnard, 1963) (Barnard 1963, fig. 1b;
Sysoev 1996b, figs 28-29) and *F. tajourensis* Sysoev & Kantor, 1987 (Fig. 5B) from the Indian Ocean as well as *F. pacifica* Sysoev & Kantor, 1987 (Fig. 4C), *F. nitida* Sysoev, 1990 (Fig. 5A) and *F. tasmanica* Sysoev & Kantor, 1987 (Fig. 4D) from the Pacific Ocean. Of these species, only the latter was included in the molecular analysis of this study (Fig. 2, S2) and no molecular data is available for the other species, including the type species. With the inclusion of the two new species described below (*F. turritelloides* n. sp. and *F. acus* n. sp.) as well as the transfer of a further species (*F. polyacantha* n. comb.), thirteen species now comprise the genus, occurring from bathyal to abyssal depth in all of the world's oceans.
Most species have very similar shell morphology and sculpture, with the exception of *F. tajourensis*, possessing a protoconch with axial pillars extending above the keel (Sysoev and Kantor 1987, fig. 1). This species exhibits an unusual pagodiform teleoconch whorl profile (Fig. 5B), which is shared with *F. monotropis* (Bouchet and Warén 1980 fig. 189). The notably shorter siphonal canal and absence of spiral sculpture of *F. nitida* (Fig. 5A) and the prominent keels of *F. polyacantha* (Fig. 5F-G), represent a further departure from the typical genus shell morphology. While our results (Fig. 2, S2) show that the latter species belongs to *Famelica*, further study is required to confirm the generic attribution of the remaining taxa, clarify status of putative new species (e.g. FA) and elucidate the phylogenetic relationships within *Famelica*.

472

473

*Famelica turritelloides* n. sp. (PSH F1)

474

(Fig. 4A)

475

Material examined

476

Holotype: Australia, Queensland, off Moreton Bay, (-26.95, 153.94), IN2017_V03_101, 2520

477

m (AMS C.482253). COI: MN983170.

478

479

Distribution

480

Known only from the type locality.

481

482

Etymology

483

For the turreted shape of its shell, vaguely resembling that of the unrelated

484

caenogastropods of the genus *Turritella*.

485
486 ZooBank registration

487 http://zoobank.org/urn:lsid:zoobank.org:act:69A31671-72A6-45E1-9048-1A04352BB83E

488

489 Description

490 Shell (SL=23.3, SW=6.4 mm) thin-walled, elongate-fusiform to turriform with strongly
491 convex whorls and long, slender siphonal canal. Protoconch pagodiform, multispiral, of at
492 least 3 whorls, all bearing a median keel, with axial pillars below keel and no sculpture
493 above. Teleoconch of 6.5 strongly convex, uniformly dark cream whorls. Suture adpressed;
494 subsutural ramp steep, moderately wide. Spiral sculpture of fine, regularly set rounded
495 cords (14 on penultimate whorl, 32 on last whorl). Axial sculpture of very fine collabral
496 growth lines, most distinct on subsutural ramp, and below - in interspaces between spiral
497 cords. Shell base convex, with pronounced concavity at its transition to long, undulating
498 siphonal canal. Aperture moderately wide; aperture length less than one third of shell
499 length. Outer lip thin, unsculptured, inner lip smooth, with very thin callus. Anal sinus
500 moderately deep, U-shaped. Shell colouration light to dark orange, with dark orange
501 protoconch. Animal with long, cylindrical rhynchostome funnel; cephalic tentacles
502 moderately short, cylindrical, eyes not detected. Rhynchocoel long, capacious.

503

504 Remarks

505 This species exhibits a very similar sculpture to that of four other *Famelica* species: *F. tasmanica* (Fig. 4D-F), *F. pacifica* (Fig. 4C), *F. catharinae* (Fig. 4B) and *F. monoceros*, from
506 which it can be distinguished by possessing a notably higher spire. Among the high-spired
507 *Famelica* species, *F. turritelloides* can be recognised by its strongly convex whorl profile and
508 orange shell colour. While eyes have not been observed in studied material, it cannot be
509 ruled out that they occur.

511

512

*Famelica acus* n. sp. (PSH F2)

513

(Figs 3A, 5C-D)

514

515 Material examined

516 Holotype (Fig. 5C): New Caledonia, (-22.33, 167.37), EXBODI, CP3844, 815-970 m (MNHN

517 IM-2009-24922). COI: MW459351.
518 Paratype (Fig. 5D): Australia, GAB, (-34.77, 130.71), IN2017_C01_207, 3389 m, 1 wet (= 519 ethanol-preserved specimen) (SAMA D49339).

520

521 Distribution

522 New Caledonia and GAB.

523

524 Etymology

525 In reference to its very elongate shell, derived from ‘acus’ (Latin = needle). Noun in 526 apposition.

527

528 ZooBank registration

529 http://zoobank.org/urn:lsid:zoobank.org:act:03D9ECD1-37D1-439D-9DF9-4DF189D8F801

530

531 Description. Shell (SL=19.1, SW=3.4 mm) very thin-walled, fragile, lanceolate-fusiform to 532 turriform. Protoconch pagodiform, multispiral, of four dark orange whorls, three of which 533 bearing a median keel, with axial pillars below the keel and no sculpture above. Teleoconch 534 of seven evenly convex, uniformly white semitransparent whorls. Subsutural ramp indistinct 535 on early whorls; more clearly demarcated after onset of spiral sculpture, steep, moderately 536 wide. Sculpture of very thin cords (about 36 on last whorl) and very fine collabral growth 537 lines. Last adult whorl convex, clearly demarcated from long, tapering siphonal canal by 538 smooth, shallow, concave transition. Aperture elongate, one third of shell length. Outer lip 539 thin, unsculptured, attenuated towards tip of siphonal canal in its lower portion; inner lip 540 smooth, with no callus. Anal sinus rather shallow, U-shaped. Animal with short, broad 541 funnel (possibly contracted upon fixation); cephalic tentacles moderately short, somewhat 542 tapering, small eyes situated at their lower portion, adjacent to base. Rhynchocoel 543 capacious.

544

545 Remarks

546 The very elongate, weakly sculptured shell (Fig. 5C-D) differentiates this species from most 547 of its congeners. *F. bitrudis* possesses a very similar, but broader and shorter shell, which 548 lacks spiral sculpture (Barnard 1963, fig. 1b; Sysoev 1996b, figs 28-29). Remains of setae of 549 syllid polychaetes were found in the digestive tract of the paratype specimen. The
550 rhynchostome funnel in this species is most likely long as in *F. turritelloides* n. sp., as
551 indicated by dense latitudinal folds in the rhynchodeum floor, suggesting it has become
552 fixed in a retracted state.

553

554 *Famelica tasmanica* Sysoev & Kantor, 1987 (PSH F3)

555 (Figs 4D-F)

556 *Famelica tasmanica* - Sysoev & Kantor, p. 1256, figs б-в

557

558 Material examined

559 Other material: Australia, Tasmania, Flinders CMR, (-40.473, 149.397), IN2017_V03_015,
560 4114-4139 m, 1 wet (AMS C.571629); Australia, NSW, off Bermagui, (-36.35, 150.91),
561 IN2017_V03_043, 4763-4750 m, 1 wet (AMS C.519370).

562

563 Remarks

564 Both specimens studied here (Fig. 4E-F) are significantly larger than the holotype (Fig. 4D),
565 this owing to difference in age.

566

567 Genus *Glaciotomella* Criscione, Hallan, Puillandre & Fedosov, 2020

568

569 Type species: *Glaciotomella investigator* Criscione, Hallan, Puillandre & Fedosov, 2020 by
570 original designation

571

572 Diagnosis

573 Shell with cyrtoconoid spire, chalky, semitranslucent to opaque. Protoconch whorls 3 to 4
574 with dense, thin axial riblets and prominent median carina. Protoconch-teleoconch
575 boundary sharp. Teleoconch of about five whitish whorls; whorl profile with weakly
576 pronounced subsutural ramp, broadly convex below. Suture deep. Sculpture below
577 subsutural ramp of low collabral axial ribs. Spiral sculpture of evenly distanced primary
578 cords, slightly weaker than axial ribs, and 1-3 weaker secondary cords in intersections
579 between primary ones. Last adult whorl globose, siphonal canal long, clearly differentiated
580 from shortly constricted base. Aperture broadly pyriform, about half of shell length. Anal
581 sinus moderately deep, J-shaped. Cephalic tentacles small; eyes minute. Rhynchostome and
582

rhynchostomal sphincter extremely large; rhynchocoel short, rhynchostome funnel

583

moderately developed. Radula of long, straight, cylindrical hypodermic teeth with no

584

distinct barbs or blades.

585

586

Remarks

587

*Glaciotomella* is defined by the combination of: (a) a multispiral, axially ribbed and keeled

588

protoconch (Fig. 3B), (b) strongly convex whorls, with cancellate sculpture (Fig. 6C-D, but

589

also Criscione *et al.* 2021, fig. 3g) and (c) straight hypodermic teeth with no barbs or blade

590

(Fig. 6). The protoconch description contained in the diagnosis (above) is based on the type

591

species (SAMA D44120 – not figured due to erosion) and on the PSH GA (MNHN IM-2009-

592

19042; Fig. 3B). Such protoconch morphology is uncommon among raphitomids and

593

(besides *Glaciotomella*) is possessed only by species of *Neopleurotomoides* Shuto, 1971,

594

such as the type species *N. rufoapicatus* (Schepman, 1913) (Schepman 1913, p. 75, pl. 29,

595

fig. 6c; Shuto 1971p. 6, pl. 1, fig. 2), *N. distincta* (Bouchet & Warén, 1980) (Bouchet and

596

Warén 1980, p. 43, fig. 233), *N. callembryon* (Dautzenberg and Fischer, 1896) (Bouchet and

597

Warén 1980, p. 43, fig. 232) and *N. aembe* (Figueira & Absalão, 2012) (Figueira and Absalão

598

2012, p. 8, fig. 7). General shell morphology and sculpture are very similar in species of

599

*Glaciotomella* (this study, Fig. 6A-C; Criscione *et al.* 2021, fig. 3g) and *Neopleurotomoides*

600

(Fig. 6D and Bouchet and Warén 1980, figs 100-103; Figueira and Absalão 2012, figs 5-6),

601

with species of the former differing mainly by their considerably larger shells. In addition,

602

species of the two genera share nearly identical, unbarbed hypodermic teeth (Fig. 6 for

603

*Glaciotomella*; see Bouchet and Warén 1980, fig. 45 for *Neopleurotomoides*). The

604

remarkable morphological similarity between *Glaciotomella* and *Neopleurotomoides* may be

605

an indication of a close phylogenetic relationship. However, the extent of this relationship

606

and its resulting taxonomic implications are still to be determined, pending availability of

607

molecular data on the type species of *Neopleurotomoides*.

608

*Glaciotomella* species are also similar to species of *Pleurotomella s.s.* (see Criscione *et al.*

609

2021) in possessing a shell with prominent sculpture, strongly convex whorls with an

610

impressed suture and a long siphonal canal and a comparatively straight, awl-shaped radula.

611

However, they differ from the latter genus in having a distinctly broader, more convex and

612

less shouldered whorl profile. While further study is required on the radula in both genera,

613

differences based on examinations thus far are that *Glaciotomella* does not possess a barb
or blade, is very straight and relatively cylindrical, and has a comparatively small ligament, whereas *Pleurotomella* may possess a very small dorsal blade, is more tapering and slightly curved, and generally has a larger ligament. Furthermore, our molecular analyses (Fig. 2, S2) are in agreement with that of Criscione *et al.* (2021) and suggest that the two genera are in fact not closely related within Raphitomidae.

619

620

*Glaciotomella investigator* Criscione et al., 2020 (PSH G1+G2)

621

(Figs 6A-B, 9A-B)

622

623

Material examined

624

Holotype (Fig. 6A): Australia, New South Wales (NSW), Hunter CMR, (-32.575, 153.162),

625

IN2017_V03_070, 2595-2474 m (AMS C571621).

626

Other material: Australia, GAB, (-34.62, 130.28), IN2015_C02_449, 2007-2067 m, (SAMA

627

D44120).

628

629

Distribution

630

Southern and Eastern Coast of Australia

631

632

Remarks

633

This genus was previously known only for the holotype of its type species *G. investigator*,

634

described without mention of its heavily eroded protoconch (Criscione *et al.* 2021, p. 25).

635

The additional specimen studied here for the species, SAMA D44120, was collected with an

636

intact protoconch that is described as follows: protoconch whorls three with dense, thin

637

axial riblets and a prominent median carina; protoconch-teleoconch boundary sharp.

638

Unfortunately, the apex was accidentally damaged while the specimen was being handled

639

for DNA extraction and cannot be illustrated. A nearly identical protoconch is possessed by

640

MNHN IM-2009-19042 (GA; Fig. 3B).

641

642

Genus *Rimosodaphnella* Cossman, 1916

643

644

Type species: *Murex textilis* Brocchi, 1814by subsequent designation by Cossmann (1916)

645
646 Diagnosis

647 Shell elongate-fusiform, moderately thick, with high spire. Protoconch multispiral, dark
648 orange, with diagonally cancellate sculpture. Teleoconch white to cream to pale purple;
649 whorl profile evenly convex (with angulation) to subcylindrical or pagodiform. Suture deeply
650 impressed. Subsutural ramp wide, concave, rather steep, sculptured by dense, raised
651 arcuate growth lines, indicating shape of the anal sinus. Below subsutural ramp, axial
652 sculpture of regular, opisthocline ribs, rarely absent. Spiral sculpture of rather densely set
653 cords, overriding axials, and typically forming nodules at intersections. Last adult whorl
654 evenly convex below subsutural ramp, clearly demarcated from straight, moderately long
655 siphonal canal. Aperture elongate, outer lip thin, inner lip with thin callus, gently recurved
656 toward left. Anal sinus wide, deep, u-shaped.

657 Animal uniform cream; head broad, blunt. Cephalic tentacles tapering evenly toward blunt
658 tip; eyes very small. Proboscis relatively long (broad and short when retracted); venom
659 gland, convoluted; muscular bulb pearlescent, kidney-shaped to ovate.

660 Radular teeth of hypodermic type, straight, tightly rolled, often rugose; barbs absent;
661 adapical opening elongate; ventral blade sharp; base moderately broad; lateral process
662 present; basal opening subcircular. Ligament moderately large, broad.

663

664 Remarks

665 Two main combined diagnostic features define *Rimosodaphnella*, namely: (a) a high-spired
666 shell with cancellate sculpture (Brocchi 1814, p. 283, pl. 8, fig. 14; McLean and Poorman
667 1971, p. 111, fig. 50; Bonfitto and Morassi 2013, figs. 1A-C, G-I, P, 2A-B) and (b) straight,
668 tightly rolled, unbarbed hypodermic teeth with a sharp dorsal blade (Fig. 10). An exhaustive
669 comparison of the shell of *Rimosodaphnella* with the shell of other raphitomid genera can
670 be found in Bonfitto and Morassi (2013). Bonfitto and Morassi (2013) critically reviewed the
671 Recent and fossil species attributed to *Rimosodaphnella*, restricting this genus to its fossil
type species *R. textilis* (Brocchi, 1814) (Fig. 7l) and to four additional Recent species: *R.*
672 *morra* (Dall, 1881) (Fig. 7J, off Cuba, 822 m), *R. brunneolineata* Bonfitto & Morassi, 2013
673 (Fig. 8A) and *R. tenuipurpurata* Bonfitto & Morassi, 2013 (Fig. 8J) and (both from the
674 Philippines, 180-250 m) as well as *R. solomonensis* Bonfitto & Morassi, 2013 (Fig. 8D) (from
675 Solomon Islands and Vanuatu, 456-551 m). Bonfitto and Morassi (2013) removed from the
676 genus *R. deroyae* McLean & Poorman 1971 (off Galapagos Islands, 200 m). However, the
677
678 observation of the shell of the holotype (Fig. 7H) is consistent with its inclusion in
679 *Rimosodaphnella*. With the addition of the two species named herein, the total number of
680 species of *Rimosodaphnella* (both fossil and Recent) is increased to eight. While generic
681 attribution is molecularly confirmed for four out of seven Recent *Rimosodaphnella* species
682 (Figs 2 and S2), genetic data is pending for the remaining three species.

683

684

*Rimosodaphnella guraradara* n. sp. (PSH R1)

685

(Figs 7A-C, 10A)

686

Material examined

687

Holotype (Fig. 7A). Australia, NSW, Jervis CMR, (-35.33, 151.26), IN2017_V03_056, 2650-
2636 m, (AMS C.571613). COI: MN983199

688

Paratypes. As per holotype, 1 wet (AMS C.571684), 1 wet (AMS C.571688); Tasmania (TAS),
Freycinet CMR, (-41.73, 149.12), IN2017_V03_004, 2820-2751 m, 1 wet (AMS C.519332), 1
wet (AMS C.571770); TAS/Victoria (VIC), Bass Strait, (-39.46, 149.28), IN2017_V03_022,
2760-2692 m, 1 wet (AMS C.482194).

693

694

Etymology

695

In reference to the long hypodermic teeth of its radula, derived from the combination of
'gurara' (Aboriginal Australian language Tharawal = long) and 'dara' (Tharawal = tooth),
noun in apposition.

698

699

ZooBank registration

700

http://zoobank.org/urn:lsid:zoobank.org:act:3D4ED0A0-FF1E-4E76-9D5E-7CDD91062F73

701

702

Distribution

703

Tasman Sea.

704

705

Description. Shell elongate-fusiform, (SL=22.8 mm, SW=8.8 mm), apex pointed but
protoconch eroded. Teleoconch uniformly whitish, 5.5 whorls; whorl profile convex, with
moderate angulation in earlier whorls. Subsutural ramp wide, concave, rather steep; suture
deployed. Sculpture of regular rounded, opisthocline ribs (about 20 on penultimate
whorl) on whorl periphery, intersected by uneven flattened cords, separated by deep and

706

707

708

709
narrow striae. Ribs more elevated on early teleoconch whorls and becoming lower, with
shallower interspaces on last adult whorl. Spiral cords forming weak, horizontally elongated
nodules at intersections with axial ribs. Subsutural ramp sculptured by dense, raised arcuate
growth lines, indicating shape of the anal sinus. Last adult whorl evenly convex below
subsutural ramp, clearly demarcated from straight, moderately long siphonal canal. Last
whorl periphery with predominant spiral sculpture; axial ribs becoming obsolete toward
base. Aperture elongate, approximately 40% of shell length; outer lip thin, unsculptured;
inner lip whitish, with thin callus, gently recurved toward left. Anal sinus wide, deep, U-
shaped. Animal uniform cream; head broad, blunt. Cephalic tentacles of medium length,
broad at base, tapering evenly toward blunt tip; weak longitudinal furrow observed in right
tentacle of one specimen; eyes very small. Proboscis relatively long (broad and short when
retracted); venom gland of medium length, convoluted; muscular bulb pearlescent, bean-
shaped to ovate, of medium size. Radular tooth hypodermic, straight, tightly rolled,
attaining approximately 175 µm in length; barb absent; adapical opening elongate; ventral
blade sharp, approximately 1/5 of length of tooth; basal third of tooth somewhat rugose, in
some teeth distinctly rugose throughout, in places with deep indentations; base moderately
broad, rugose; lateral process present; basal opening subcircular, unrestricted. Ligament
relatively large and broad.

728

729 Remarks

The shell of this species (Fig. 7A-C) can be differentiated from most congeners by the
combination of larger size and convex whorl profile with weakly-defined shoulder. R.
*truvana* is comparable in size and sculpture but possesses distinctly shouldered whorls.
Most other congeners, particularly R. *deroyae* (Fig. 7H) and the PSHs RC, RD and RE (Fig. 8G-
I), possess fewer but more prominent sculptural elements. Finally, the PSH RA (Fig. 7G) can
be differentiated from R. *guraradara* by a complete lack of axial elements in the former. The
radula of R. *guraradara* is considerably longer than that of R. *truvana* based on the material
examined.

738

739

Rimosodaphnella truvana n. sp. (PSH R2)

740

(Figs 7D-F, 10B)

741
742 Material examined

743 Holotype (Fig. 7D). Australia, TAS/VIC, Bass Strait, (-39.55, 149.55), IN2017_V03_030, 4197-4133 m, (AMS C.571685). COI: MN983200.

745 Paratypes. As per holotype, 1 wet (AMS C.519315), 1 wet (AMS C.519316), 1 wet (AMS C.571615), 1 wet (AMS C.571686), 1 wet (AMS C.571689); TAS, Flinders CMR, (-40.47, 149.40), IN2017_V03_015, 4114-4139 m, 1 wet (AMS C.571687).

748

749 Etymology

750 In reference to the closest land mass to the type locality, derived from 'Truvana' (Aboriginal Australian language Palawa Kani = Cape Barren Island), name in apposition.

752

753 Zoobank registration number

754 http://zoobank.org/urn:lsid:zoobank.org:act:614EE2C7-1A95-4049-9179-08DA467FD60D

755

756 Distribution

757 Tasman Sea.

758

759 Description. Shell solid, broadly fusiform, (SL=24.5 mm, SW=11.2 mm), protoconch eroded.

760 Teleoconch uniformly whitish, of at least six whorls. Early teleoconch whorls convex, subsequent whorls distinctly shouldered with nearly subcylindrical profile, giving spire somewhat stepped appearance. Suture impressed. Subsutural ramp concave, sculptured with fine, arcuate collabral growth lines. Sculpture of rounded, slightly opisthocline axial ribs (about 22 on penultimate and last whorl) intersected by dense, uneven cords. Axials more elevated, with deeper interstices on early teleoconch whorls and lower, densely set on last adult whorl. Spiral cords rounded, becoming somewhat nodulose at intersections with axial elements. Last adult whorl strongly convex, sculptured predominantly by spiral elements, clearly demarcated from long and slender siphonal canal. Aperture wide, elongate-pyriform, approximately half of shell length; outer lip thin, simple; inner lip whitish, with well-developed callus, gently twisting toward left. Anal sinus wide, moderately deep, U-shaped.

761
762
763
764
765
766
767
768
769
770
771
772
773
Anatomy (based on holotype and paratypes AMS C.571687, C.571686, C.571689): animal uniform whitish to cream; head broad, blunt. Penis large, somewhat flattened, muscular, coiling clockwise, ¼ distal end tapering. Cephalic tentacles broad, of medium length,
774 cylindrical, tip blunt; eyes very small. Osphradium and ctenidium large. Venom apparatus:

775 Proboscis far retracted into rhynchocoel in specimens examined; venom gland moderately

776 long, convoluted; muscular bulb large, pearlescent, kidney shaped.

777 Radular teeth of hypodermic type, straight, tightly rolled, attaining approximately 125 µm in

778 length; barb absent; terminal pore elongate; ventral blade sharp, approximately 1/5 of

779 length of tooth; some teeth rugose, in places with deep indentations; base moderately

780 broad, of coarse texture; lateral process present; basal opening subcircular. Ligament

781 comparatively broad and large.

782

783 Remarks

784 *Rimosodaphnella truvana* n. sp. can be distinguished from all other congeners by its much

785 broader and larger shell with a subcylindrical rather than convex whorl outline (Fig. 7D-F). It

786 differs from the shell of *R. guraradara* (Fig. 7A-C) by its far more angular shoulder at the

787 whorl periphery, the length of its aperture which is approximately equal to the length of the

788 spire (compared to the relatively shorter aperture in the former), and its more pronounced

789 cancellate sculpture. The radula of *R. truvana* is shorter than that of *R. guraradara*.
790 Discussion

791 The five-gene phylogeny of Criscione et al. (2020) established the phylogenetic framework
792 upon which the new genus *Glaciotomella* was recognised and described, and it is shown
793 herein that there is strong support in both BI and ML analyses for its monophyly, as is the
794 case for *Famelica* and *Rimosodaphnella* (Figs 2, S2).

795 The level of genetic distinctiveness reported for *cox1* sequences of the three genera (Tables
796 1, 2) was generally comparable with those reported for the other deep-sea raphitomid
797 genera (Hallan *et al.* 2021; Criscione et al., in press) at the intra-specific level, but overall
798 higher (up to more than three times; *e.g.* average 4.1% in *Theta* vs. 14.2% in *Glaciotomella*)
799 at the inter-specific level. It remains unclear whether this is a reflection of a comparatively
800 reduced species sampling (most species of these genera have low abundance) or indication
801 of a higher diversification rate in these lineages when compared with other raphitomid
genus-level lineages. Other than being supported by molecular evidence, the integrity of the
802 genera studied herein is also corroborated by morpho-anatomical features diagnostic for
803 each genus. Among those features, the sculpture of the protoconch and the radula were
804 particularly conserved among congeneric species and therefore represented solid diagnostic
805 genus-level characters.
806

807 *Rimosodaphnella* possessed a multispiral larval shell, with protoconch II exhibiting a
808 diagonally cancellate sculpture (Fig. 3C). This configuration is the most commonly recorded
809 in Raphitomidae, where it may display a range of variations. Conversely, protoconchs with a
810 keel and axial ‘pillars’, such as those observed here for *Famelica* and *Glaciotomella* (Fig. 3A-
811 B) are considered a departure from the prevailing raphitomid pattern (Bouchet *et al.* 2011).
812 Among deep-sea raphitomid genera, keeled protoconchs are shared by *Famelica*, *Aliceia*,
813 *Neopleurotomoides* and *Glaciotomella*, which can be separated into pairs by the two former
814 genera having no axial elements above the keel, as opposed to the two latter ones where
815 axial elements persist above the keel. This, and other obvious morpho-anatomical affinities,
816 exhibited in pairs by these genera (see above Remarks to genera), coupled with the close
817 phylogenetic relationship between *Famelica* and *Glaciotomella* (this study, Figs 2, S2;
818 Criscione *et al.* 2021, fig. 2), might suggest that all deep-sea raphitomid genera with keeled
819 protoconchs are part of the same lineage (clade A of fig. 2 of Criscione *et al.* 2021), although
820 this requires testing through molecular data. Nevertheless, owing to the presence in that
821 lineage of at least one genus with a typical diagonally cancellate protoconch
822 (Rimosodaphnella: this study, Figs 2, S1; Criscione et al. 2021, fig. 2), it is unlikely that a
823 'keeled-pillared' protoconch is the plesiomorphic state for this lineage (Criscione et al.
824 2021).

825 Three main types of radular morphology have been reported for Australian deep-sea
826 raphitomid genera (Criscione et al. 2021): (1) double-barbed, (2) awl-shaped with dorsal
827 blade and (3) awl-shaped with no distinct blade. The latter two were recorded for two
828 genera in this study: (2) in *Rimosodaphnella* (Fig. 10) and (3) in *Glaciotomella* (Fig. 9). The
829 hypodermic teeth lacking barbs or blades, or reduced in size are commonly found in
830 raphitomids, and are thought to be a result of secondary simplification of morphology
831 (Kantor and Taylor 2002, see also below)(Kantor & Taylor 2002, also see discussion below).

832 Unfortunately, diagnostic value of simplified morphology is questionable as it is virtually
833 impossible to extract a phylogenetic signal associated with loss of a structure and to
834 ascertain, whether loss of certain feature has happened once or several times in the
835 evolution.

836 None of the species of *Famelica* studied here possess a radula or a venom apparatus, in
837 agreement with that reported by Criscione *et al.* (2021) for the genus. As hypodermic teeth
838 as well as a well-developed intraembolic proboscis and venom gland are plesiomorphic in
839 Raphitomidae (Kantor and Sysoev 1989; Bouchet *et al.* 2011), the lack or reduction of these
840 structures in some lineages has been interpreted as a derived state, which has occurred
841 independently in multiple unrelated deep-sea lineages (Kantor and Taylor 2002; Criscione *et
842 al.* 2021). The loss of such structures (involved in predation by stabbing and injecting
843 venom) has in some studies been associated with adaptation to alternative feeding
844 strategies (Kantor and Sysoev 1989; Taylor *et al.* 1993; Kantor and Taylor 2002). In some
845 raphitomid groups, the loss of radula and venom apparatus has been accompanied by
846 substantial anatomical modifications, such as the development of a prominent rhynchodeal
847 introvert (e.g. some *Raphitoma* spp.), the expansion of the cavity between rhynchodeum
848 and body walls (e.g. *Abyssobela* Kantor & Sysoev, 1986 and *Teretiopsis* Kantor & Sysoev,
849 1989) or the evolution of accessory rhynchodeal organs (e.g. Tritonoturris subrissoides
850 (Hervier, 1897); Fedosov 2007). No obvious anatomical rearrangements have occurred in
851 some other groups (e.g. *Taranis* Jeffreys, 1870). The well-developed rhynchodeal introvert
852 was also found to be a common feature in Terebridae, another lineage of Conoidea
853 remarkable for tendency toward complete loss of the venom apparatus (Taylor 1990; Taylor
et al. 1993; Castelin et al. 2012). Furthermore, terebrid species with a well-developed proboscis and hypodermic radulae appeared to have smaller introverts compared to those with reduced or lacking radula and/or proboscis (Taylor 1990). This observation had corroborated earlier observation on the feeding of terebrids, which demonstrated that the rhynchodeal introvert plays a primary role in feeding for some terebrids (Miller 1975). In Raphitomidae it was also showed that a well-developed introvert can occur in taxa possessing a prominent venom apparatus and radula: *Hemilienardia*, *Austrobela* and *Pueridaphne* (Kantor and Taylor 2002; Criscione et al. 2021). One could hypothesize that development of a functional rhynchodeal introvert is a prerequisite to simplification of the organs related to envenomation (which has occurred in most living Raphitomidae), often leading to their complete reduction. It is possible that initially, the role of the introvert was limited to retention of prey during envenomation, as for cone-snails of the subgenus *Conus* (*Gastridium*). Conids of this group first engulf fish-preys into an enormous rhynchostome funnel, and then stab them with a radular harpoon (Olivera et al. 2015) Species of *Conus* (*Gastridium*) are remarkable for their simplified radular teeth, lacking elaborated barbs and serrations, that are structures that keep prey 'hooked' preventing its escape. Indeed, once prey is enclosed in a rhynchocoel, it cannot escape, and as the retention of prey is taken over by rhynchostome funnel, elaborated radular weaponry becomes unnecessary, and undergoes reduction. The widespread morphological simplification of the hypodermic teeth in Raphitomidae can possibly be explained by the same causal link (i.e. by partial transfer of their functions to the rhynchodeal introvert). Progressively, the introvert might enable alternative feeding mechanisms, which do not require envenomation (similar to the mechanisms observed in Terebridae) and which eventually led to the complete loss of radula, venom gland and proboscis. Some occasional prey findings in radula-less raphitomids (e.g. polychaete setae identified as belonging to a member of the family Syllidae, recovered from *Famelica acus* n. sp. SAMA D49339) suggest that these species retain a predatory lifestyle.

Shells are heterogeneous for most species of all three genera, but generally exhibit similar gross morphology, with very few exceptions. The overall strongly elongate shell shape of *Famelica* (Fig. 4-5), the fusiform shell with strongly convex whorls of *Glaciotomella* (Fig. 6) and the cancellate teleoconch sculpture of *Rimosodaphnella* (Fig. 7-8) are the main recognisable characters in their respective composite taxa. However, while most congeneric
886 species are similar in shell morphology [e.g. *F. tasmanica* (Fig. 4D-F), *F. acus* (Fig. 5D-E) and
887 *F. turritelloides* (Fig. 4A), *R. guraradara* (Fig. 7A-C) and *R. truvana* (Fig. 7D-F) or *R.

888 brunneolineata (Fig. 8A-C) and *R. solomonensis* (Fig. 8D-F)], other species or PSHs [e.g. *F.
889 polyacantha* (Fig. 5F-G) or *Rimosodaphnella* RA (Fig. 7G)] exhibit shell features that are
890 somewhat divergent from those shared by their congeners. Some species of
891 *Rimosodaphnella* and *Glaciotomella* studied here have much larger shells than others.

892 Although the small sample size did not allow thorough statistical testing, the size of their
893 shells (and bodies) appears to be positively correlated with depth, with species from the
894 lower bathyal (i.e. *R. guraradara* and *R. truvana* for the former genus and *G. investigator* for
895 the latter) larger than species (or PSHs) of shallower waters (i.e. *R. brunneolineata* and
896 *Glaciotomella* GA for each genus respectively). A size-depth clinal pattern is well-known for
897 bathyal neogastropods (Rex and Etter 1998; Harasewych and Kantor 2004; McClain *et al.*

898 2005) and for turriform conoideans in particular ('turrids' in Rex *et al.* 1999; McClain and Rex
899 2001) and has been interpreted as an adaptive response to an environmental gradient in the
900 deep sea. The increase in maximum size attained with depth would provide these snails
901 with increasing metabolic rates and competitive advantages as food resources diminish (Rex
902 and Etter 1998; Rex *et al.* 1999). For instance, as their polychaete prey becomes scarcer with
903 depth (Thistle *et al.* 1985), larger raphitomids may benefit from increased mobility as well as
904 ability to feed on a wider range of prey sizes (Levinton 1982, 1987). As is often the case for
905 deep-sea studies, it is difficult to establish the extent to which inevitably limited
906 observations reflect such adaptive mechanisms, and to which they are confounded by
907 sampling biases (see Criscione *et al.* 2021).

908
909 Biogeographic and bathymetric patterns

910 Most species treated herein occur within an area corresponding approximately to the South
911 Australia marine realm of Costello *et al.* (2017). Unsurprisingly, the records are
912 concentrated in the areas of sampling: a relatively restricted portion of the GAB and on a
913 section of the temperate eastern Australian coast between south-eastern Tasmania and
914 Moreton Bay (Fig. 1). Samples included in this study that were collected outside this region
915 tend to cluster around the areas targeted during MNHN-led voyages in the tropical Indo-
916 Pacific and Pacific (Table S1).
Of the three genera studied here (restricted to the sequenced samples), one (*Glaciotomella*) seems to be relatively restricted, recorded only in Australia and Melanesia (Fig. 1A) whereas the other two have ranges that from South Australia extend respectively to the Central Pacific (*Famelica*, Fig 1A) and to the Central and Eastern Pacific (*Rimosodaphnella*, Fig. 1B). When *Famelica* is regarded in its entirety (i.e. including all species herein attributed to it, irrespective of the availability of molecular data for them – see Remarks under this genus), it displays a much broader, transoceanic range, encompassing all of the world's Oceans.

Geographic ranges of comparable size, purportedly stretching over two or more Oceans, have been reported for other raphitomid genera present in Australia, such as *Austrobela* and *Theta* (Criscione et al., in press) as well as *Pagodibela* (Hallan *et al.* 2021). If confirmed by molecular data, a hypothetical *Glaciotomella* + *Neopleurotomoides* genus-level group (see above), including species from three different Oceans, would also feature a similarly broad range. When *Rimosodaphnella* is regarded in its entirety (i.e. including species whose attribution is pending molecular testing), it exhibits a prevalent South Pacific distribution.

However, further sampling outside this area (such as the Atlantic, where *R. morra* is found) may reveal the presence of further species. Among the species reported here for Australia, none occur outside the two realms encompassing Australian waters (Fig. 1). However, future deep-sea sampling may reveal records beyond these realms, particularly for species occurring further north, such as *F. acus* or *F. turritelloides* n. spp. Species of other deep-sea raphitomid genera, recorded for the Tropical Australia & Coral Sea realm (#16 of Costello *et al.* 2017), commonly have ranges extending into neighbouring tropical realms [Hallan *et al.* (2021) for *Pagodibela baruna* (Sysoev, 1997); Criscione et al., in press for *Austrobela procera* (Sysoev & Bouchet, 2001) and *Spergo fusiformis* (Habe, 1962)].

The comparatively wide geographic distributions of *F. acus* and *G. investigator* are not entirely unexpected given the mounting evidence of wide distributions in some species of other deep-sea raphitomid genera, such as *Austrobela*, *Spergo* and *Theta* (Criscione et al., in press) as well as *Gladiobela* and *Pagodibela* (Hallan *et al.* 2021).

As neighbouring regions with comparable environmental conditions remain virtually unexplored, it is plausible that disjunct distributions, like that observed for *F. acus* and *G. investigator* (Fig. 1A), are the result of sampling bias, rather than a reflection of any underlying biological process. Low mean *cox1* genetic distance between two populations, when compared with the range of distances calculated within one of the populations of the
raphitomid *Gladiobela angulata* Criscione, Hallan, Puillandre & Fedosov, 2020 has been interpreted as indication of high genetic connectivity (Hallan *et al.* 2021). In order to perform similar comparisons for *F. acus* and *G. investigator*, more than one sequence from each of the two disjunct populations would be necessary. In addition, due to the aforementioned sampling bias, it remains unclear whether genetic connectivity in these species is realised through a stepping-stone process, involving additional geographically intervening populations, or through long-range dispersal of the planktotrophic larva (or both).

*Rimosodaphnella guraradara* and *R. truvana* are present in sympatry off the coast of East Tasmania (Fig. 1B), where there is no overlap in their bathymetric ranges, with a gap of more than 1000 m (Fig. 11), despite some sampling coverage within this gap at corresponding areas. It has been suggested that partitioning into separate bathymetric niches is the driver of speciation of sister species of some other Australian deep-sea raphitomid genera, such as *Gladiobela* (Hallan *et al.* 2021) and *Austrobela* (Criscione et al., in press), albeit the ecological factors involved remain unknown. As *R. guraradara* and *R. truvana* are not recovered as sister taxa based on our results, speciation via bathymetric partitioning appears unlikely; rather, a scenario of a secondary contact between the two species appears more plausible.

Due to the limited material of *F. turritelloides*, we cannot infer much about its biogeography. While it cannot be ruled out that its rarity may be an artefact of sampling, it is possible that this species may indeed be relatively rare, as the regions in which it has been collected are comparatively well-sampled (MacIntosh *et al.* 2018; O’Hara *et al.* 2020) and have revealed the presence of a diverse raphitomid fauna (Criscione et al., 2020). For species of other raphitomid genera from the same regions, such as *Gladiobela* (Hallan *et al.* 2021), *Theta and Austrotheta* (Criscione et al., in press), scarce records have been interpreted as evidence of their intrinsic rarity. Conversely, some *Rimosodaphnella* species studied here, notably *R. guraradara* and *R. truvana*, can be considered relatively common, as observed for most species of *Austrobela* (Criscione et al., in press) and some of *Gladiobela* (Hallan *et al.* 2021). Estimates of bathymetric ranges of deep-sea turriform conoideans genera have traditionally relied on morphology-based genus attribution (Bouchet 1990; Kantor *et al.* 2016). For raphitomid genera, Criscione *et al.* (2021) provided the first available depth range size, based on Australian species whose generic attribution was confirmed by molecular data. By
complementing molecular data on further species from beyond Australia, this and other
studies suggest range size expansions for a number of these genera. Among these expanded
depth ranges, those of *Spergo* Dall, 1895 (4432 m; Criscione et al., in press), *Famelica* (4352
m; this study), *Gladiobela* Criscione, Hallan, Puillandre and Fedosov, 2020 (4276 m; (Hallan
et al. 2021) and *Rimosodaphnella* (3874 m; this study) are the widest four reported for the
family to date. The bathymetric range of *F. acus* (at least 2574 m; Fig. 11) constitutes one of
the widest ever reported for a molecularly confirmed conoidean species, second only to that
of *Gladiobela acris* Hallan, Criscione, Fedosov and Puillandre, 2021 (Hallan et al. 2021). Due
to the scarcity of material for most other species studied here, little inference can be made
with regards to their bathymetric zonation.
991 Conflict of interest

992 The authors declare no conflicts of interest.

993

994 Acknowledgments

995 This work has been made possible through financial support from the Australian

996 Government (ABRS grant RF217-57, principal investigator FC). The participation of AF was

997 also supported by the Russian Science Foundation (grant 19-74-10020). The participation of

998 NP was also supported by the European Research Council (ERC) under the European Union's

999 Horizon 2020 research and innovation programme (grant agreement No. 865101).

1000 Voyages in the GAB were part of: (a) the GAB Research Program [GABRP – a collaboration

1001 between BP, CSIRO, the South Australian Research and Development Institute (SARDI), the

1002 University of Adelaide and Flinders University] and (b) the GAB Deepwater Marine Program

1003 (GABDMP – a CSIRO led research program sponsored by Chevron Australia]. Funding for the

1004 ‘Eastern Abyss’ voyage (IN2017_V03) was provided by the Marine Biodiversity Hub (MBH),

1005 supported through the Australian Government’s National Environmental Science Program

1006 (NESP). The authors wish to thank the CSIRO MNF for its support in the form of sea time

1007 onboard, support personnel, scientific equipment and data management. We also thank the

1008 scientific staff and crew who participated in all voyages generating the samples studied

1009 herein. The MNHN samples used in this study originates from shore-based expeditions

1010 (PANGLAO 2004, PAPUA NIUGINI; PI Philippe Bouchet) and deep-sea cruises (AURORA 2007,

1011 BIOPAPUA, BOA 1, EXBODI, KANACONO, KAVIENG 2014, MAINBAZA, PANGLAO 2005,

1012 SALOMON 2, SALOMON BOA 3, TARASOC, ZHONGSHA 2015; PIs Philippe Bouchet, Tin-Yam

1013 Chan, Laure Corbari, Nicolas Puillandre, Sarah Samadi, Wei-Jen Chen, Bertrand Richer de

1014 Forges) conducted by MNHN, Pro-Natura International (PNI) and Institut de Recherche pour

1015 le Développement as part of the Our Planet Reviewed and the Tropical Deep-Sea Benthos

1016 programs. Funders and sponsors included a bilateral cooperation research funding from the

1017 Taiwan Ministry of Science and Technology (MOST 102-2923-B-002-001-MY3, PI Wei-Jen

1018 Chen) and the French National Research Agency (ANR 12-ISV7-0005-01, PI Sarah Samadi),

1019 the Total Foundation, Prince Albert II of Monaco Foundation, Stavros Niarchos Foundation,

1020 and Richard Lounsbery Foundation. One sample was collected during the INSPIRE cruise

1021 (Chile) onboard R/V Melville (SCRIPPS institutions). All expeditions operated under the

1022 regulations then in force in the countries in question and satisfy the conditions set by the
1023 Nagoya Protocol for access to genetic resources (expeditions.mnhn.fr). We would like to
1024 express our gratitude to Mandy Reid, Alison Miller and Jennifer Caiza (AMS) for assistance
1025 with registration and databasing of material, to Virginie Héros, Philippe Maestrati and
1026 Barbara Buge (MNHN) for the sample preparation and databasing and to Andrea Crowther
1027 (SAMA, Adelaide. Thanks are also due to Sue Lindsay and Chao Shen (Macquarie University,
1028 Sydney) for facilitating SEM work. We are indebted to several people providing photos of
types and other relevant specimens: Adam Baldinger and Alana Rivera (MCZ, Cambridge,
1029 USA), Antonio Bonfitto (Bologna University, Italy), Jeroen Goud and Bram van der Bijl
1030 (NMNL, Leiden), Kazunori Hasegawa (NSMT, Tokyo, Japan), Lindsey Groves (LACM, Los
1031 Angeles), Peter Stahlschmidt (Universität Koblenz-Landau, Koblenz, Germany), Ellen Strong
1032 (NMNH, Washington), Alexander Sysoev (ZMMU, Moscow), Giorgio Teruzzi and Anna
1033 Alessandrello (MSNMi, Milan).
1034
1035 References

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<table>
  <tr>
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</td>
  </tr>
</table>
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1313

1314
1315 Figure captions

1316

1317 Figure 1. Distribution of taxa studied herein with focus on the South West Pacific. (A)

1318 Records of sequenced specimens of *Famelica* (circles) and *Glaciotomella* (triangles). (B)

1319 Records of sequenced specimens of *Rimosodaphnella* (squares). The insets show records of
1320 *Rimosodaphnella* in off the coast of Chile (upper righolotype) and in the Tuamotu

1321 Archipelago (lower left). Numbers and letters in shapes indicate PSHs/species of: *Famelica*

1322 (circles: 1 – F1/F. *turritelloides*, 2 – F2/F. *acus*, 3 – F3/F. *tasmanica*, 4 – F4/F. *polyacantha*, A – FA); *Glaciotomella* (1 – G1/G. *investigator*, 2 – G2/G. *investigator*, A – GA) and

1323 *Rimosodaphnella* (squares: 1 – R1/R. *guraradara*, 2 - R2/R. *truvana*, 3 – R3/R. *solomonensis*, 4 – R4/R. *brunneolineata*, A – RA, B – RB, C – RC, D – RD, E – RE). Thin lines mark limits

1324
1325
1326 among marine realms (numbered as in Costello et al., 2017). Main scalebars = 1000 km,
1327 insets scalebars = 200 Km.

1328

1329 Figure 2. Bayesian consensus phylogram (BI) based on analyses of *cox1* sequences. Numbers

1330 above branches indicate nodal support by Bayesian posterior probabilities (BPP). Numbers

1331 below PSH nodes indicate nodal support (%) by bootstrap (BS) resulting from the ML

1332 analysis of Fig. S1, with 100% values indicated by asterisks. Names of species described

1333 herein and sequences of Australian samples are in bold. Vertical bars mark distinct PSHs as
1334 delimited by ABGD on the corresponding *cox1* dataset. Dashed rectangles mark alternative

1335 PSH groupings as delimited by the ASAP method. Symbols and numbers next to taxon

1336 names correspond to those used in Figure 1. Samples whose shells are figured are

1337 underlined. Scale bar (right) = 7.5 mm (shells of R3-R5, RB-RE, GA), 10 mm (other shells).

1338

1339 Figure 3. Protoconchs of species/PSHs studied here. (A) F2/F. *acus* n. sp. holotype MNHN

1340 IM-2009-249221; (B) GA/G. sp. MNHN IM-2009-19042; (C) R5/R. brunneolineata, MNHN

1341 IM-2009-18977.

1342

1343 Figure 4. Shells of *Famelica* PSHs/species. (A) F1/F. *turritelloides* n. sp. holotype AMS

1344 C.482253; (B) *F. catharinae* (Verrill & S. Smith [in Verrill], 1884) holotype USMN 37871; (C) *F.*

1345 *pacifica* Sysoev & Kantor, 1987 holotype ZMMU LC 5369; (D) *F. tasmanica* Sysoev & Kantor,
1346

1987 holotype ZMMU LC 5370; (E) F3/F. tasmanica AMS C.519370; (F) F3/F. tasmanica AMS

1347

C.571629. Scale bar = 10 mm.

1348

1349

Figure 5. Shells of *Famelica* PSHs/species. (A) *F. nitida* Sysoev, 1990 holotype ZMMU LC

1350

5737; (B) *F. tajourensis* Sysoev & Kantor, 1987 holotype ZMMU LC 5371; (C) F2/F. acus n. sp.

1351

holotype MNHN IM- 2009-249221; (D) F2/F. acus n. sp. paratype SAMA D49339; (E) FA/F. sp.

1352

MNHN IM-2013-61627; (F) F4/F. *polyacantha*, MNHN IM-2009-17104; (G) *F. polyacantha*

1353

(Stahlschmidt, Chino & Kilburn 2012) holotype NSMT Mo 78456. Scale bar = 10 mm (A-D, F-

1354

G); 12.5 mm (E).

1355

1356

Figure 6. Shells of *Glaciotomella* and *Neopleurotomoides* PSHs/species. (A) G1/G.

1357

*investigator* Criscione et al. 2020 holotype AMS C.571621; (B) G2/G. *investigator* Criscione

1358

et al. 2020, SAMA D44120; (C) GA/G. sp. MNHN IM-2009-19042, (D) *N. rufoapicatus*

1359

(Schepman, 1913) holotype NMNL MOLL.136877. Scale bar = 10 mm (A-B), 5 mm (C), 4 mm

1360

(D).

1361

1362

Figure 7. Shells of *Rimosodaphnella* PSHs/species. (A) R1/R. *guraradara* n. sp., holotype AMS

1363

C.571613; (B) R1/R. *guraradara* n. sp., paratype AMS C.571684; (C) R1/R. *guraradara* n. sp.,

1364

paratype AMS C.519332; (D) R2/R. *truvana* n. sp., holotype AMS C.571685; (E) R2/R. *truvana*

1365

n. sp., paratype AMS C.571615; (F) R2/R. *truvana* n. sp., paratype AMS C.571686; (G) RA/R.

1366

sp., MNHN IM-2013-57410; (H) *R. deroyae* n. sp., holotype LACM 1543; (I) *R. textilis* (Brocchi,

1367

1814), holotype MSNMi 5340; (J) *R. morra* (Dall, 1881) holotype MCZ 7107. Scale bar = 10

1368

mm (A-H), 7.5 mm (I), 2.5 mm (J).

1369

1370

Figure 8. Shells of *Rimosodaphnella* PSHs/species. (A) *R. brunneolineata* Bonfitto & Morassi,

1371

2013, holotype MZB 49760; (B) R5/R. brunneolineata, MNHN IM-2009-18977; (C) R4/R.

1372

*brunneolineata*, MNHN IM-2007-42460; (D) *R. solomonensis* Bonfitto & Morassi, 2013,

1373

holotype MNHN 25803; (E) R3/R. solomonensis Bonfitto & Morassi, 2013, MNHN IM-2007-

1374

42524; (F) RB/R. sp., MNHN IM-2007-17836; (G) RC/R. sp., MNHN IM-2013-58315; (H) RD/R.

1375

sp., MNHN IM-2007-38690; (I) RE/R. sp., MNHN IM-2007-39365; (J) *R. tenuipurpurata*

1376

Bonfitto & Morassi, 2013, holotype MZB 49758. Scale bar = 10 mm.

1377
1378 Figure 9. Hypodermic radular teeth of *Glaciotomella* PSHs/species studied herein. (A)

1379 G1/*Glaciotomella investigator* Criscione et al., 2020 holotype AMS C.571621; (B)

1380 R2/*Glaciotomella investigator* Criscione et al. 2020, SAMA D44120. Scale bar = 100 µm.

1381

1382 Figure 10. Hypodermic radular teeth of *Rimosodaphnella* PSHs/species studied herein. (A)

1383 R1/*Rimosodaphnella guraradara* n. sp., paratype AMS C.519274; (B) R2/*Rimosodaphnella truvana* n. sp., paratype AMS C.571689. Scale bar = 50 µm.

1384

1385

1386 Figure 11. Bathymetric ranges of PSHs/taxa studied herein as inferred from records of

1387 sequenced specimens. Species represented by a single record are indicated by a circle.

1388
1389 Tables captions

1390

1391 Table 1. Intra- and inter-PSHs/specific genetic differentiation of *cox1* sequences in *Famelica*

1392 by means of K2p distances. Intra-PSHs/specific distances shaded. Maximum and minimum

1393 values of inter-PSHs/specific distance in bold. Inset: minimum, maximum and average intra-

1394 and inter-PSHs/specific p-distances within *Famelica*. Species codes: acu, *acus*; pol,

1395 *polyacantha*; tas, *tasmanica*; tur, *turritelloides*. Codes of species described herein in bold.

1396

1397 Table 2. Intra- and inter-PSHs/specific genetic differentiation of *cox1* sequences in

1398 *Rimosodaphnella* by means of K2p distances. Intra-PSHs/specific distances shaded.

1399 Maximum and minimum values of inter-PSHs/specific distance in bold. Inset: minimum,

1400 maximum and average intra- and inter-PSHs/specific p-distances within *Rimosodaphnella*.

1401 Species codes: bru, *brunneolineata*; gur, *guraradara*; sol, *solomonensis*; tru, *truvana*. Codes

1402 of species described herein in bold.

1403
<figure><img src="image_5.png" /><figcaption>Distribution of taxa studied herein with focus on the South West Pacific. (A) Records of sequenced specimens of Famelica (circles) and Glaciotomella (triangles). (B) Records of sequenced specimens of Rimosodaphnella (squares). The insets show records of Rimosodaphnella in off the coast of Chile (upper righolotype) and in the Tuamotu Archipelago (lower left). Numbers and letters in shapes indicate PSHs/species of: Famelica (circles: 1 - F1/F. turritelloides, 2 - F2/F. acus, 3 - F3/F. tasmanica, 4 - F4/F. polyacantha, A - FA); Glaciotomella (1 - G1/G. investigator, 2 - G2/G. investigator, A - GA) and Rimosodaphnella (squares: 1 - R1/R. guraradara, 2 - R2/R. truvana, 3 - R3/R. solomonensis, 4 - R4/R. brunneolineata, A - RA, B - RB, C - RC, D - RD, E - RE). Thin lines mark limits among marine realms (numbered as in Costello et al., 2017). Main scalebars = 1000 km, insets scalebars = 200 Km.</figcaption></figure>

170x126mm (600 x 600 DPI)
<figure><img src="image_6.png" /></figure>

Bayesian consensus phylogram (BI) based on analyses of cox1 sequences. Numbers above branches indicate nodal support by Bayesian posterior probabilities (BPP). Numbers below PSH nodes indicate nodal support (%) by bootstrap (BS) resulting from the ML analysis of Fig. S1, with 100% values indicated by asterisks. Names of species described herein and sequences of Australian samples are in bold. Vertical bars mark distinct PSHs as delimited by ABGD on the corresponding cox1 dataset. Dashed rectangles mark alternative PSH groupings as delimited by the ASAP method. Samples whose shells are figured are underlined. Scale bar (right) = 7.5 mm (shells of R3-R5, RB-RE, GA), 10 mm (other shells).

168x171mm (600 x 600 DPI)
<figure><img src="image_7.png" /><figcaption>Figure 3. Protoconchs of species/PSHs studied here. (A) F2/F. acus n. sp. holotype MNHN IM-2009-249221; (B) GA/G. sp. MNHN IM-2009-19042; (C) R5/R. brunneolineata, MNHN IM-2009-18977.</figcaption></figure>

199x65mm (300 x 300 DPI)
<figure><img src="image_8.png" /><figcaption>Figure 4. Shells of Famelica PSHs/species. (A) F1/F. turritelloides n. sp. holotype AMS C.482253; (B) F. catharinae (Verrill & S. Smith [in Verrill], 1884) holotype USMN 37871; (C) F. pacifica Sysoev & Kantor, 1987 holotype ZMMU LC 5369; (D) F. tasmanica Sysoev & Kantor, 1987 holotype ZMMU LC 5370; (E) F3/F. tasmanica AMS C.519370; (F) F3/F. tasmanica AMS C.571629. Scale bar = 10 mm.</figcaption></figure>
<figure><img src="image_9.png" /><figcaption>Figure 5. Shells of Famelica PSHs/species. (A) F. nitida Sysoev, 1990 holotype ZMMU LC 5737; (B) F. tajourensis Sysoev & Kantor, 1987 holotype ZMMU LC 5371; (C) F2/F. acus n. sp. holotype MNHN IM-2009-249221; (D) F2/F. acus n. sp. paratype SAMA D49339; (E) FA/F. sp. MNHN IM-2013-61627; (F) F4/F. polyacantha, MNHN IM-2009-17104; (G) F. polyacantha (Stahlschmidt, Chino & Kilburn 2012) holotype NSMT Mo 78456. Scale bar = 10 mm (A-D, F-G); 12.5 mm (E).</figcaption></figure>
<figure><img src="image_10.png" /><figcaption>Figure 6. Shells of Glaciotomella and Neopleurotomoides PSHs/species. (A) G1/G. investigator Criscione et al. 2020 holotype AMS C.571621; (B) G2/G. investigator Criscione et al. 2020, SAMA D44120; (C) GA/G. sp. MNHN IM-2009-19042, (D) N. rufoapicatus (Schepman, 1913) holotype NMNL MOLL.136877. Scale bar = 10 mm (A-B), 5 mm (C), 4 mm (D).</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>Figure 7. Shells of Rimosodaphnella PSHs/species. (A) R1/R. guraradara n. sp., holotype AMS C.571613; (B) R1/R. guraradara n. sp., paratype AMS C.571684; (C) R1/R. guraradara n. sp., paratype AMS C.519332; (D) R2/R. truvana n. sp., holotype AMS C.571685; (E) R2/R. truvana n. sp., paratype AMS C.571615; (F) R2/R. truvana n. sp., paratype AMS C.571686; (G) RA/R. sp., MNHN IM-2013-57410; (H) R. deroyae n. sp., holotype LACM 1543; (I) R. textilis (Brocchi, 1814), holotype MSNMi 5340; (J) R. morra (Dall, 1881) holotype MCZ 7107. Scale bar = 10 mm (A-H), 7.5 mm (I), 2.5 mm (J).</figcaption></figure>
<figure><img src="image_12.png" /><figcaption>Figure 8. Shells of Rimosodaphnella PSHs/species. (A) R. brunneolineata Bonfitto & Morassi, 2013, holotype MZB 49760; (B) R5/R. brunneolineata, MNHN IM-2009-18977; (C) R4/ R. brunneolineata, MNHN IM-2007-42460; (D) R. solomonensis Bonfitto & Morassi, 2013, holotype MNHN 25803; (E) R3/R. solomonensis Bonfitto & Morassi, 2013, MNHN IM-2007-42524; (F) RB/R. sp., MNHN IM-2007-17836; (G) RC/R. sp., MNHN IM-2013-58315; (H) RD/R. sp., MNHN IM-2007-38690; (I) RE/R. sp., MNHN IM-2007-39365; (J) R. tenuipurpurata Bonfitto & Morassi, 2013, holotype MZB 49758. Scale bar = 10 mm.</figcaption></figure>
<figure><img src="image_13.png" /><figcaption>Figure 9. Hypodermic radular teeth of Glaciotomella PSHs/species studied herein. (A) G1/Glaciotomella investigator Criscione et al., 2020 holotype AMS C.571621; (B) R2/Glaciotomella investigator Criscione et al. 2020, SAMA D44120. Scale bar = 100 µm.</figcaption></figure>
<figure><img src="image_14.png" /><figcaption>Figure 10. Hypodermic radular teeth of Rimosodaphnella PSHs/species studied herein. (A) R1/Rimosodaphnella guraradara n. sp., paratype AMS C.519274; (B) R2/Rimosodaphnella truvana n. sp., paratype AMS C.571689. Scale bar = 50 µm.</figcaption></figure>
<figure><img src="image_15.png" /><figcaption>Figure 11. Bathymetric ranges of PSHs/taxa studied herein as inferred from records of sequenced specimens. Species represented by a single record are indicated by a circle.</figcaption></figure>

83x84mm (600 x 600 DPI)
<table><thead><tr><th></th><th>FA</th><th>F1/tur</th><th>F2/acu</th><th>F3/tas</th><th>F4/pol</th><th></th><th></th><th></th></tr></thead><tbody><tr><th>FA</th><td>-</td><td></td><td></td><td></td><td></td><td></td><td>min</td><td>max</td><td>mean</td></tr><tr><th>F1/tur</th><td>0.130</td><td>-</td><td></td><td></td><td></td><td>within</td><td>0.008</td><td>0.008</td><td>0.008</td></tr><tr><th>F2/acu</th><td>0.171</td><td>0.169</td><td>0.008</td><td></td><td></td><td>between</td><td>0.052</td><td>0.182</td><td>0.145</td></tr><tr><th>F3/tas</th><td>0.129</td><td>0.052</td><td>0.154</td><td>-</td><td></td><td></td><td></td><td></td><td></td></tr><tr><th>F4/pol</th><td>0.182</td><td>0.174</td><td>0.138</td><td>0.147</td><td>-</td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><thead><tr><th></th><th>R1/gur</th><th>R2/tru</th><th>RA</th><th>R3/sol</th><th>R4/bru</th><th>R5/bru</th><th>RB</th><th>RC</th><th>RD</th><th>RE</th></tr></thead><tbody><tr><td>R1/gur</td><td>0.002</td><td></td><td></td><td></td><td></td><td></td><td></td><td>min</td><td>max</td><td>mean</td></tr><tr><td>R2/tru</td><td>0.093</td><td>0.003</td><td></td><td></td><td></td><td></td><td></td><td>0.000</td><td>0.003</td><td>0.002</td></tr><tr><td>RA</td><td>0.064</td><td>0.122</td><td>-</td><td>-</td><td></td><td></td><td>within</td><td>0.046</td><td>0.156</td><td>0.117</td></tr><tr><td>R3/sol</td><td>0.092</td><td>0.098</td><td>0.117</td><td>-</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>R4/bru</td><td>0.125</td><td>0.125</td><td>0.132</td><td>0.113</td><td>-</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>R5/bru</td><td>0.128</td><td>0.104</td><td>0.135</td><td>0.123</td><td>0.046</td><td>-</td><td></td><td></td><td></td><td></td></tr><tr><td>RB</td><td>0.127</td><td>0.138</td><td>0.133</td><td>0.115</td><td>0.107</td><td>0.128</td><td>-</td><td></td><td></td><td></td></tr><tr><td>RC</td><td>0.141</td><td>0.136</td><td>0.156</td><td>0.126</td><td>0.122</td><td>0.132</td><td>0.102</td><td>0.003</td><td></td><td></td></tr><tr><td>RD</td><td>0.147</td><td>0.141</td><td>0.141</td><td>0.131</td><td>0.129</td><td>0.127</td><td>0.092</td><td>0.098</td><td>0.000</td><td></td></tr><tr><td>RE</td><td>0.119</td><td>0.129</td><td>0.124</td><td>0.115</td><td>0.111</td><td>0.116</td><td>0.091</td><td>0.099</td><td>0.067</td><td>-</td></tr></tbody></table>
<caption>Figure 51 - NJ tree generated based on a dataset including all available AMS and MNNH COI sequences of Raphitomidae. Inset: sequences of Raphitomidae in the dataset. The tree is a single tree, with the criteria described in the text.</caption>

The criteria described in the text.

<figure><img src="image_16.png" /></figure>

Invertebrate Systematics

<figure><img src="image_17.png" /></figure>
<figure><img src="image_18.png" /></figure>

Figure S1. Maximum likelihood (ML) tree based on analyses of COI sequences. Numbers near branches indicate nodal support (%) by ML bootstrap (BS). Support values for clades below the PSH/species level are omitted. Names of species described herein, and sequences of Australian samples are in bold. Vertical bars mark distinct primary species hypotheses (PSHs) as delimited by the ABGD method. Dashed rectangles mark alternative PSH groupings as delimited by the ASAP method.

168x225mm (600 x 600 DPI)