# Mitogenomic phylogeny of mud snails of the mostly Atlantic/ Mediterranean genus Tritia (Gastropoda: Nassariidae)

Yi Yang, Samuel Abalde, Carlos Afonso, Manuel J Tenorio, Nicolas

Puillandre, Jose Templado, Rafael Zardoya

► To cite this version:

Yi Yang, Samuel Abalde, Carlos Afonso, Manuel J Tenorio, Nicolas Puillandre, et al.. Mitogenomic phylogeny of mud snails of the mostly Atlantic/ Mediterranean genus Tritia (Gastropoda: Nassariidae). Zoologica Scripta, 2021, 50 (5), pp.571-591. 10.1111/zsc.12489. hal-03172830

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Distributed under a Creative Commons Attribution 4.0 International License
1 Submitted to:

2 *Zoologica Scripta*

3 Second revised version: March 8th, 2020

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Mitogenomic phylogeny of mud snails of the mostly Atlantic/Mediterranean
genus *Tritia* (Gastropoda: Nassariidae)

Yi Yang¹, Samuel Abalde¹, Carlos L. M. Afonso², Manuel J. Tenorio³, Nicolas Puillandre⁴, José
Templado¹, and Rafael Zardoya¹

¹Museo Nacional de Ciencias Naturales (MNCN-CSIC), José Gutiérrez Abascal 2, 28006, Madrid,
Spain;

²Centre of Marine Sciences (CCMAR), Universidade do Algarve, Campus de Gambelas, 8005-139
Faro, Portugal

³Departamento CMIM y Q. Inorgánica-INBIO, Facultad de Ciencias, Universidad de Cadiz; 11510
Puerto Real, Cádiz, Spain;

⁴Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum National d'Histoire Naturelle,
CNRS, Sorbonne Université, EPHE, Université des Antilles, 57 rue Cuvier, CP 26, 75005 Paris,
France.

<footnote>*Corresponding author ([rafaz@mncn.csic.es](mailto:rafaz@mncn.csic.es))</footnote>
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**Abstract**

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**KEYWORDS**

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Nassariinae, *Tritia*, *Ilyanassa*, mitochondrial genome, phylogeny, chronogram

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# 1. INTRODUCTION

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Mud snails or dog whelks (family Nassariidae) are small- to medium-sized snails, readily recognized by their fusiform, slender to broadly ovate shells of moderately sculptured surface (from smooth to cancellate or reticulate) with more or less prominent axial and spiral ribs and cords (Cernohorsky, 1984). The base of the shell aperture has a notch for extension of their long siphon. The columella is short with a pronounced siphonal fold (Cossmann, 1901). The reduced operculum is thin and corneous with a terminal nucleus and often marginally serrated. Most nassariids species are facultative scavengers or non-selective predators (Morton, 2011). Their characteristic long inhalant siphon (as long as their body) together with a well-developed osphradium allows acute chemoreception and efficiently leads to carrion or prey. Their often very wide and flattened foot enables these sea snails to glide quickly and easily over soft substrata.

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The nassariids constitute an important component of the biodiversity of tropical and temperate seas and attain their greatest diversity in the tropical Indo-Pacific (Cernohorsky, 1984). A few species have reached the polar seas (Nekhaev, 2014). Mud snails inhabit predominantly in shallow water soft bottoms, but some live in deep water, and many characterize certain types of sediments where they are dominant (Harasewych, 1998; Hayashi, 2004). For instance, the species of the genus *Bullia* are the most conspicuous surfing organisms of the swash zone of many sandy beaches in the southern hemisphere, and some species of *Nassarius* and *Tritia* display dense populations in estuaries and coastal lagoons. Several species have adapted to brackish water and a few have successfully conquered freshwater habitats in Southeast Asia (Strong, Galindo, & Kantor, 2017). Due to their abundance, nassariids have been well studied from different points of view. For instance, many nassarid species are known to accumulate toxins and as such constitute a health problem especially in Asia (Hwang et al., 2005); other species have been used as model organisms, such as *Tritia obsoleta* (as *Ilyanassa obsoleta*) in developmental biology (Goulding & Lindberg, 2016) or *Bullia digitata* in ecological and behavioral studies (Brown, 1980). Furthermore, nassariids have a rich fossil record (Gili & Martinell, 1993, Tracey, Todd, & Erwin, 1993; Haasl, 2000; Harzhauser & Kowalke, 2004; Landau, da Silva, & Gili, 2009) since the lower Cretaceous (Taylor, Morris, & Taylor, 1980) that should help understanding major evolutionary trends within the family when interpreted within a phylogenetic framework.

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The first thorough classification of nassariids was based on the shape of the shell columella (Cossmann, 1901), and is still in use with slight modifications (Cernohorsky, 1984). The family was traditionally divided into three subfamilies: Nassarinae, Dorsaniinae and Cylleninae, until Allmon (1990) transferred several genera previously included in Dorsaniinae to establish the new subfamily Bulliinae. However, the taxonomy of these subfamilies, especially the status of genera and
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subgenera, has remained far from resolved because of rampant convergence in shell morphology and other phenotypic characters due to adaptations to similar environmental or selective pressures. Moreover, most nassariids species were assigned by default to the genus *Nassarius* and many are still provisionally included in this genus, which demands thorough revision.

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Recently, a molecular phylogeny including more than 200 putative nassariid species was reconstructed based on partial sequences of five (mitochondrial and nuclear) genes (Galindo et al., 2016). This molecular phylogeny showed that the morphology-based taxonomy of the family (Cernohorsky, 1984; Allmon, 1990; Haasl, 2000; Kantor, 2003) was dominated by homoplasy and had little phylogenetic significance. Genera were redefined, the new subfamily Buccinanopsinae was created, and the taxon Photinae was reestablished at subfamily level (Galindo et al., 2016).

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The reconstructed phylogeny of Galindo et al. (2016) showed that the cosmopolitan subfamily *Nassariinae* comprises about 80% of nassariid species diversity (MolluscaBase, 2020a). Up to five strongly supported clades were recovered within this subfamily and were assigned to genera with relatively distinct geographical distributions: the Indo-Pacific *Nassarius* and *Reticunassa* (the “pauper-complex” sensu Kool & Dekker, 2006, 2007, Galindo, Kool, & Dekker, 2017); the mostly Atlantic-Mediterranean *Tritia*; the (East and West) African *Naytia*; and the Caribbean-Panamanian *Phrontis*. Furthermore, these genera showed different types of protoconch. The multispiral protoconchs of species assigned to *Nassarius* and *Reticunassa* have one or two keels, whereas those of species included within *Tritia*, *Naytia*, and *Phrontis* lack a keel (Galindo et al., 2016). Transitions to a paucispiral protoconch have occurred independently within different genera (Galindo et al., 2016).

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Phylogenetic relationships among species within the different genera remained largely unresolved in several cases (Galindo et al., 2016). After Galindo et al. (2016) around 60 species were reclassified into genera other than *Nassarius* but more than 300 species remained tentatively in this genus (MolluscaBase, 2020b) awaiting further phylogenetic studies. In this regard, two subsequent mitogenomic studies that focused on the genus *Nassarius* improved the resolution of phylogenetic relationships among species restricted to the China seas (Yang, Li, Kong, & Yu, 2019; Yang et al., 2020), compared with previous attempts based on partial gene sequences (e.g., Zhang & You, 2009; Chen & Zhang, 2012; Pu et al., 2017). As a result of these mitogenomic studies, the composition of the genus *Nassarius* was refined, with *Nassarius jacksonianus* and *Nassarius* sp. (incorrectly identified as *N. acuticostus*; see Supporting Information Figure S1) placed with strong statistical support as sister to *Reticunassa* (Yang, Li, Kong, & Yu, 2019). Furthermore, another mitogenomic phylogenetic study focused on genus *Reticunassa* revealed cryptic species (Yang, Li, Kong, & Yu, 2018). These studies highlight the need of studying in depth the species composition
and phylogenetic relationships within the five recognized Nassariinae genera. They also confirm the
high levels of resolution achieved by complete mitochondrial genomes when inferring phylogenetic
relationships of gastropods below the family level (Uribe, Puillandre, & Zardoya, 2017).

The traditional taxonomy of the Atlantic/Mediterranean nassariids was mainly based on shell
characters, with species classified into different genera (Hinia, Nassarius, Sphaeronassa, Naytiopsis,
Cyclope and Ilyanassa). Even though species of Cyclope possess rather unique flattened shell
geometry, and Ilyanassa is restricted to the Northwest Atlantic, the molecular phylogeny of
Nassariidae found no evidence to support these genera, which were both synonymized with Tritia
(Galindo et al., 2016). At present, a total of 40 species are considered valid within Tritia
(MolluscaBase, 2020c), including four species endemic to the Gulf of Gabès that were recently
recognized as valid (Aissaoui, Galindo, Puillandre, & Bouchet, 2017). Several Tritia species extend
their range distribution to West African coasts living in sympatry with Naytia species.

In order to clarify the taxonomy of the genus *Tritia* in the Atlantic/ Mediterranean region, a
robust phylogeny of the group is needed. Thus far, only two complete mitochondrial genomes of
*Tritia* species are available i.e., *T. obsoleta* (DQ238598; Simison, Lindberg, & Boore, 2006, as
*Ilyanassa obsoleta*) and *T. reticulata* (EU827201; Cunha, Grande, & Zardoya, 2009). In the present
study, we sequenced the nearly complete (without the control region or partial gene sequences)
mitochondrial genomes of 21 specimens representing 20 species of *Tritia* (i.e., half of the diversity
of the genus) and of one specimen of the freshwater nassariid *Anentome* sp. as outgroup. Our aims
were: (1) to test the monophyly of the genus *Tritia* as currently defined (MolluscaBase, 2020c); (2)
to reconstruct robust phylogenetic relationships within the genus; (3) to date the origin and main
cladogenetic events during the diversification of this genus; and (4) to test the validity of shell
morphology in *Tritia* taxonomy.

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## 2. MATERIALS AND METHODS

### 2.1. Samples and DNA extraction

The complete list of specimens analyzed in the present study, including their sampling sites,
collectors, and museum vouchers is shown in Table 1. Samples were stored in 100% ethanol at 4°C
or -20°C, and total genomic DNA was isolated from foot muscle using the DNeasy Blood & Tissue
Kit (Qiagen, Germany).

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### 2.2. PCR amplification and sequencing
Nearly complete (without the control region or partial gene sequences) mt genomes were amplified in two main steps. First, fragments of *cox1*, *cox3*, *rrnS*, *rrnL*, and *cob* genes were amplified through standard PCRs, which were carried out in a total volume of 25 µl with 1 µl of template DNA (25–40 ng/µl), 2.5 µl of 10× buffer (Mg²⁺ plus), 0.5 µl of dNTPs (2.5 mM), 0.5 µl of each primers (10 µM), 0.2 µl of Taq DNA polymerase (5 U/µl), 0.25µl of BSA (10 mg/ml) and 19.55 µl of DEPC water.

The PCR conditions were as follows: an initial denaturing step at 94 °C for 5 min; 40 cycles of denaturing at 94 °C for 60 s, annealing at 39–48°C for 60 s, and extension at 72 °C for 90 s; and a final extension step at 72 °C for 10 min. Standard-PCR products were purified by ethanol precipitation, and Sanger sequenced at Macrogen (Seoul, Korea). Information on standard PCR primers is shown in Supporting Information Table S1.

In the second step, the obtained sequences were used to design specific primer pairs for long PCR amplification of the remaining mitogenome in four fragments (see Supporting Information Table S1). The long PCR reactions contained 2.5 µl of 10× buffer (Mg²⁺ plus), 3 µl of dNTPs (2.5 mM), 0.5 µl of each primers (10 µM), 0.8 µl of template DNA (25–40 ng/µl), 0.2 µl of TaKaRa LA Taq DNA polymerase (5 U/µl), 0.2 µl of BSA (10 mg/ml) and DEPC water up to 25 µl. The following PCR conditions were used: initial denaturing step at 94 °C for 60 s; 45 cycles of denaturing at 98 °C for 10 s, annealing at 53-56 °C for 30 s, and extension at 68 °C for 60 s per kb; final extension step at 68 °C for 10 min. Long PCR products were purified by ethanol precipitation, and fragments from the same mitogenome were pooled together in equimolar concentrations for high-throughput sequencing. For each mitogenome of *Tritia*, a separate indexed library was constructed using the NEXTERA XT DNA library prep Kit (Illumina, San Diego, CA, USA) and run in an Illumina MiSeq platform (2 × 150 paired-end) at AllGenetics (A Coruña, Spain) and NIMGenetics (Madrid, Spain). Among the 20 libraries, five contained also the mitogenomes of cones or buccinoids from different projects (see Supporting Information Table S1).

Due to the degraded condition of the source DNA of *T. ovoidea* and *T. tingitana*, the partial mt genomes of these species were mostly amplified through standard PCRs, as only few long PCRs worked (see Supporting Information Table S1). Those amplified fragments with sizes ≤ 1,000 bp were directly Sanger sequenced with the corresponding PCR primers, whereas the longer PCR products were Sanger sequenced using a primer walking strategy. Sequencing was performed in automated sequencers (ABI PRISM 3700) using the BigDye® Terminator v3.1 cycle-sequencing kit
(Applied Biosystems, Foster City, CA, USA), and following the manufacturer's instructions.

For some species with tentative identification, namely *N. jacksonianus* and *N. acusticostus*, or
of unexpected phylogenetic position, namely *T. ephamilla* and *T. vaucheri*, the barcoding *cox1*
fragment was PCR amplified and Sanger sequenced to double check species identity (see
*Supporting Information Figure S1*).

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## 2.3. Genome assembly and annotation

The Sanger sequences were assembled using Sequencher 5.0.1. For the specimens from the MNHN
collections, some *cox1* sequences were downloaded from Genbank (see Supporting Information
Table S1). For Illumina sequence data, the reads corresponding to different individuals were sorted
by the corresponding library indices. Raw sequences were analyzed for quality control using
FastQC v.0.11.9 (Andrews, 2010), and trimmed and filtered out according to their quality scores
using PRINSEQ v.0.20.4 (Schmieder & Edwards, 2011). Assembly of the mitogenomes was
performed using Geneious Prime 2019.0.3 (Kearse et al., 2012). The mitogenomes were constructed
by repeatedly mapping the original reads (setting a minimum identity of 99%) to the Sanger
fragments from the same mitogenome.

The newly determined mt genomes were annotated with Geneious Prime 2019.0.3. Annotations
of protein-coding genes (PCGs) were defined by setting a limit of 75% nucleotide identity to
previously published nassariid mitogenomes (Table 1) in Geneious Prime 2019.0.3, and
corroborated using the MITOS Webserver (Bernt et al., 2013) with the invertebrate mitochondrial
genetic code. The transfer RNA (tRNA) genes were further identified using tRNA scan-SE 1.21
(Schattner, Brooks, & Lowe, 2005) and ARWEN (Laslett & Canbäck, 2008). The ribosomal RNA
(rRNA) genes were first identified using the MITOS Webserver (Bernt et al., 2013). The software
rendered shorter rRNA genes (due to sequence variability in the ends), and their boundaries were
assumed to be between the adjacent genes by comparison with other nassariid mt genomes.

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## 2.4. Sequence alignment

The newly sequenced mitogenomes were aligned along with those of other nassariids available in
Genbank (Table 1). Representatives of different genera of Nassariinae were included in the ingroup
to test the monophyly of *Tritia*. The freshwater nassariid *Anentome* sp. (subfamily Anentominae)
215 was used as outgroup to root the tree (Galindo et al., 2016). A dataset concatenating the nucleotide
216 sequences of the 13 PCGs and two rRNA genes was constructed and analyzed.

217 The 13 PCGs were aligned separately guided by amino acid translations (according to the
218 invertebrate mitochondrial genetic code) using Translator X (Abascal, Zardoya, & Telford, 2010), .
219 Nucleotide sequences of the two rRNA genes were aligned separately using MAFFT v7 (Katoh &
220 Standley, 2013) with default parameters. Ambiguously aligned positions were removed using
221 Gblocks v0.91b (Castresana, 2000) with the following settings: minimum sequence for flanking
222 positions: 85%; maximum contiguous non-conserved positions: 8; minimum block length: 10; gaps
223 in final blocks: no. Finally, the different single alignments were concatenated into a single data set
224 in Geneious Prime 2019.0.3. Sequences were converted into different formats for downstream
225 analyses using DAMBE5 (Xia, 2013).

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227 **2.5. Phylogenetic analyses**

228 Pairwise uncorrected *p* sequence distances to delimit species were estimated based on the alignment
229 including the nucleotide sequences of the 13 PCGs and the two rRNA genes. Phylogenetic trees
230 were reconstructed using maximum likelihood (ML; Felsenstein, 1981) and Bayesian inference (BI;
231 Huelsenbeck & Ronquist, 2001). ML analyses were carried out using IQtree 1.6.10 (Nguyen,
232 Schmidt, Von Haeseler, & Minh, 2015) in the CIPRES gateway (Miller, Pfeiffer, & Schwartz, 2010),
233 allowing partitions to have different evolutionary rates (-spp option) and with 10,000 ultrafast
234 bootstrap pseudoreplications (-bb option). BI analyses were performed with MrBayes
235 v.3.1.2 (Ronquist & Huelsenbeck, 2003), running four simultaneous Monte Carlo Markov chains
236 (MCMC) for 10,000,000 generations, sampling every 1000 generations, and discarding the first 25%
237 generations as burn-in. Two independent runs were performed to increase the chance of adequate
238 mixing of the Markov chains and to increase the chance of detecting failure to converge, as
239 determined by using Tracer v1.6. The effective sample size (ESS) of all parameters was higher than
240 200. The resulting phylogenetic trees were visualized in FigTree v1.4.2.

241 The best partition schemes and best-fit substitution models for BI analysis of the data set were
242 determined (see Supporting Information Table S2 for selected best fit partitions and models) using
243 PartitionFinder 2 (Lanfear et al., 2017), under the Bayesian Information Criterion (BIC; Schwarz,
244 1978). For the 13 PCGs, the partitions tested were: all genes combined; all genes separated (except
245 *atp6-atp8* and *nad4-nad4L*); and genes grouped by subunits (*atp*, *cob*, *cox* and *nad*). Additionally, these three partition schemes were tested considering separately the three codon positions. The two 246 rRNA genes were analyzed with two different schemes (genes grouped or separated). The best-fit 247 substitution models for ML analysis were calculated with ModelFinder (Kalyaanamoorthy, Minh, 248 Wong, Haeseler, & Jermiin, 2017) as implemented in IQ-TREE 1.6.10. 249

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251 **2.6. Ancestral Character State Reconstruction**

252 The type of larval shell or protoconch with a single (paucispiral) or multiple whorls (multispiral)
253 was accessed based on literature (Galindo et al., 2016; Aissaoui, Galindo, Puillandre, & Bouchet,
254 2017) except for *T. vaucheri*, which was determined by direct examination of the shells (the number
255 of whorls was counted to the nearest quarter whorl; Modica et al., 2020). Both types of protoconch
256 were used as a proxy of the lecithotrophic (direct development using yolk reserves) or
257 planktotrophic (with a feeding veliger phase) modes of larval development (Strathmann, 1978). The
258 evolution of larval development was analyzed by performing an ancestral character state
259 reconstruction in Mesquite v3.6.1 (Maddison & Maddison, 2018) using the Tracing Character
260 History option under ML and mapping onto the identical topology inferred by the ML and BI
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263 **2.7. Estimation of divergence times**

264 The program BEAST v.1.10.4 (Drummond & Rambaut, 2007) was used to perform a Bayesian 265 estimation of divergence times among main nassariid lineages based on PCGs only. An uncorrelated 266 relaxed molecular clock was used to infer branch lengths and node ages. The tree topology was 267 fixed using the identical topology recovered by the ML and BI analyses. For the clock model, the 268 lognormal relaxed-clock model was selected, which allows rates to vary among branches without 269 any *a priori* assumption of autocorrelation between adjacent branches. For the tree prior, a Yule 270 process of speciation was employed. The partitions selected by PartitionFinder 2 (see above) were 271 applied. The final Markov chain was run twice for 100 million generations, sampling every 10,000 272 generations and the first 10 million were discarded as burn-in, according to the convergence of 273 chains checked with Tracer v.1.6. The effective sample size of all the parameters was above 200. 274

The posterior distribution of the estimated divergence times was obtained by specifying three
calibration points based on fossil data as priors for divergence times of the corresponding splits. The prior distributions of the three calibration nodes were selected following Ho & Phillips (2009). For the origin of the lineage leading to *T. neritea* and *T. pellucida*, a calibration point was set at the minimum limit of 3.6 million years ago (Mya; exponential distribution, offset: 3.6; mean: 1.0) based on the fossils of *Cyclope migliorinii* that are the oldest for the stem group (Gili & Martinell, 1999; Landau, da Silva, & Grigis, 2009) and dated from the Pliocene (Placenzian; 3.6 - 2.6 Mya) of Spain according to the Paleobiology Database (https://paleobiodb.org). A second calibration point was set at the origin of *Tritia*. A lognormal distribution was applied, with the minimum of 23 Mya and a 95% upper limit of 28.4 Mya (offset: 23; mean: 1.9; standard deviation: 1.9) based on *Nassarius (Hinia)* *pygmaea* from Germany (Albright et al., 2019) and *Nassarius (Hinia)* *schlotheimi* from Hungary (Báldi, 1973), both from the Oligocene (Chattian; 28.4 - 23.0 Mya), the oldest Nassariinae fossils in Europe (Lozouet, 1999). A third calibration point was set at the root of the tree. A lognormal distribution was applied, with the minimum of 55.8 Mya and a 95% upper limit of 58.7 Mya (offset: 55.8; mean: 1.0; standard deviation: 1.0) based on the oldest known Nassariinae fossil, which is *Buccitriton sagenum* from the Paleocene (Thanetian; 58.7 - 55.8 Mya) of Louisiana, USA (Glawe, Anderson, & Bell, 2014).

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## 3. RESULTS

### 3.1. Sequencing and assembly

All newly sequenced mt genomes in the present study lacked the sequences of the *trnF* gene, the control region, and the start of the *cox3* gene because the corresponding fragment could not be PCR amplified. The number of reads, mean coverage, length, and accession number in Genbank of each mitogenome are provided in Table 1. The mt genomes of *T. incrassata*-SP and *T. varicosa* received the minimum (53x; 30,800 reads) and maximum (23,350x; 2,765,768 reads) coverage, respectively (Table 1).

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### 3.2. Genome organization

The newly determined *Tritia* mitogenomes encode for 13 PCGs, two rRNA and 22 tRNA genes (but note that the *trnF* gene could not be determined in all mt genomes; see annotation of each mt genome in Supporting Information Table S3). They all share the same genome organization
described for Caenogastropoda (Osca, Templado, & Zardoya, 2015), with most genes encoded by
the major strand and a cluster of tRNA genes (*trnM*, *trnY*, *trnC*, *trnW*, *trnQ*, *trnG*, *trnE*) and the
*trnT* gene encoded by the minor strand (Figure 1). All PCGs start with the conventional initiation
codon ATG (but note that the start of the *cox3* gene could not be determined in all mt genomes; see
Supporting Information Table S3). As for the termination codons, two genes (*cox2* and *atp8*)
consistently ended with TAA and two (*nad4L* and *nad5*) stopped with TAG. The incomplete stop
codon (TA-), which becomes functional after polyadenylation (Ojala et al., 1981), was observed in
all *nad2* genes but that of *Tritia grana*, which ended with TAA. The remaining genes varied in their
stop codons depending on the species (Supporting Information Table S3).

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315 **3.3. Phylogenetic relationships**

Phylogenetic relationships of *Tritia* were reconstructed based on the nucleotide sequences of the
concatenated 13 PCGs and two rRNA genes using probabilistic methods (Figure 2). The final
matrix was 13,101 positions in length. According to the BIC, the best partition scheme for the PCGs
was the one combining genes by subunits but analyzing each codon position separately (Supporting
Information Table S2). For the rRNA genes, the best partition scheme was the one combining
together *rrnL* and *rrnS* genes. Both ML (−*lnL* = 133,593.77) and BI (−*lnL* = 133,905.39 for run 1;
−*lnL* = 133,908.66 for run 2) analyses arrived at identical topologies (Figure 2).

According to the reconstructed phylogeny, *N. jascksonianus* and *Nassarius* sp were recovered
closer to *Reticunassa* than to other *Nassarius* species (Figure 2). *Tritia vaucheri* was placed sister to
the clade including (*N. jascksonianus* and *Nassarius* sp.) + *Reticunassa*, and apart from the
remaining *Tritia* species, which formed a natural group. Within this latter group, the Northwest
Atlantic species *T. obsoleta* was placed as sister to three mostly Northeast Atlantic/ Mediterranean
clades (I-III). Clade I included ten species distributed into five lineages: (1) *T. ephamilla* (endemic
to New Zealand and southern Australia) and *T. ovoidea* + *T. elata*, (2) *T. denticulata*, (3) *T. tingitana*,
(4) *T. reticulata* and *T. nitida*, and (5) the “*T. cuvierii* complex” including *T. unifasciata* and *T.
cuvierii* + *T. tenuicosta*. This clade was sister to clades II and III (Figure 2). Clade II comprised *T.
varicosa* and *T. incrassata* (including one specimen from a Spanish Mediterranean population and
one from Norway). Clade III included *T. elongata* + (*T. pallaryana* and *T. corniculum*) sister to a
group formed by *T. grana* + (*T. neritea* and *T. pellucida*) and *T. pfeifferi* + *T. mutabilis* (Figure 2).
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**3.4. Ancestral character state reconstruction**

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The ancestral character state reconstruction analysis inferred that the Atlantic/Mediterranean nassariids could have originated from an ancestor with a multispiral protoconch, which indicates a planktotrophic mode of larval development (Figure 3). The “*T. cuvierii* complex”, clade III, and *T. tingitana* were inferred to represent independent evolutionary transitions to a paucispiral protoconch, suggesting a secondary loss of the planktotrophic mode of larval development to lecitotrophy.

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**3.5. Divergence times**

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Main cladogenetic events within *Tritia* were dated using an uncorrelated relaxed molecular clock model, which was calibrated using three Atlantic fossils. The origin of the stem lineage leading to genus *Tritia* (including *T. obsoleta* and without *T. vaucheri*) was dated 24.0 (26.2–23.1, 95% highest posterior density interval, HPD) million years ago (Mya). The first event of diversification within the crown group of *Tritia* was estimated at a mean of 20.4 (23.5–17.1) Mya, separating the Northwest Atlantic *T. obsoleta* from *T. ephamilla* and East Atlantic/ Mediterranean *Tritia* species.

The branching of the three main East Atlantic/ Mediterranean clades was estimated to have occurred around 18.9–17.6 (22.0–14.4) Mya. Finally, main diversification events leading to extant *Tritia* species were estimated to have occurred between 16.9–3.9 (20.1–2.0) Mya, whereas divergences of some closely-related species or populations were dated about 0.6–0.1 Mya (Figure 4).

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# 4. DISCUSSION

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**4.1. Taxonomy of the non-*Tritia* Nassariinae**

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363

364

The often-ornamented shell of nassariids, which could in principle provide numerous characters for taxonomic classification, and had been widely used in the past for this purpose, was found to be prone to convergence at different taxonomic levels (particularly within Nassariinae) when a rather comprehensive molecular phylogeny of the family was inferred (Galindo et al., 2016). This phylogeny provided the long needed scaffold to continue resolving the systematics of the family and opened the way to further phylogenetic studies focusing on relationships within particular clades. Here, we focused on one of these clades, which included nassariinae mostly from the Atlantic/Mediterranean region and was ascribed to genus *Tritia* (Galindo et al., 2016). We
365 sequenced the near-complete (without the control region or partial gene sequences) mitogenomes of
366 20 species belonging to *Tritia* and reconstructed a highly resolved phylogeny, which included more
367 than half of the species diversity of this genus (MolluscaBase, 2020c) as well as other nassariinae
368 genera and *Anentome* sp. (subfamily Anetominae) as outgroup.

369 Within the first offshoot of the tree, *N. jascksonianus* and *Nassarius* sp. were recovered closer
370 to *Reticunassa* than to other *Nassarius*, as previously suggested (Yang, Li, Kong, & Yu, 2019). This
371 is not surprising considering the geographic disparity of the samples and that not all species
372 ascribed to *Nassarius* were included in the molecular phylogeny of the family, impeding proper
373 assignment to clades (Galindo et al., 2016). Given the relative phylogenetic position of the two
374 species and the large uncorrected *p* distance separating them from other clades (16–18%), they
375 could be allocated in a new genus, whose formal description would need a larger taxon sampling
376 and thorough morphological revision beyond the scope of this study.

377 According to the reconstructed phylogeny, the ascription of *T. vaucheri* to genus *Tritia* was
378 strongly rejected. This taxon was originally described as *Nassa* (*Himia*) *vaucheri* (Pallary, 1906).
379 After examining the syntype of this species, Cernohorsky (1975), transferred it to genus *Chauvetia*
380 (in family Buccinidae). Later, Hoenselaar and Moolenbeek (1988) transferred the species to genus
381 *Nassarius* and designated a lectotype. Finally, the new combination *T. vaucheri* (currently accepted
382 in MolluscaBase, 2020c) was established in a checklist of marine mollusks recorded in Spanish
383 marine waters (Gofas et al., 2017). *Tritia vaucheri* was recovered as an early divergent lineage in
384 the phylogeny, sister to the Indo-Pacific clade (*N. jascksonianus* and *Nassarius* sp.) + *Reticunassa*,
385 although with low statistical support. Hence, this species does not belong to genus *Tritia*, but to
386 another Atlantic genus that is only distantly related and still needs to be determined (*Genus incertae
387 sedis*, *Gen. inc. sed.*). Therefore, the general shell morphology resemblance of this taxon to other
388 *Tritia* species would represent yet another case of convergence within Nassariidae (Galindo et al.,
389 2016). *Genus incertae sedis vaucheri* is found in the southwestern Mediterranean coasts of Spain
390 and in the nearby Atlantic coasts from south Portugal to Morocco being replaced by *Nassarius
391 argenteus* from Senegal down to the Gulf of Guinea (Gofas, 2011). This latter species was not
392 included in our study but may also belong to the same genus as *Gen. inc. sed. vaucheri*. The
393 above-mentioned distribution is typical of several species of *Tritia* but also of several sympatric
394 species of the African genus *Naytia*. Galindo et al. (2016) provisionally included in this latter genus
395 *N. glabrata* (type species), *N. granulosa*, *N. johni*, from West African coasts, and *N. priscardi* from Madagascar, and they highlighted that the members of this genus were characterized by high morphological disparity. The type species and *N. granulosa* exhibit a smooth shell, similar to that of the species of the *Bullia* group, which led Simone & Pastorino (2014) to erroneously include *N. granulosa* in the genus *Bullia*. Conversely, *N. johni* and *N. priscardi*, exhibit a reticulate shell sculpture. *Gen. inc. sed. vaucheri* is quite similar to these two latter species, so it could be possible that it belongs to *Naytia* (the position of this genus in the phylogeny of Galindo et al. 2016 is sister to *Reticunassa*, although with low support). It will be necessary to include mitogenomes of species from other genera (especially from West Africa) into the nassariid phylogeny in order to fully resolve the phylogenetic position of *Gen. inc. sed. vaucheri* and its generic placement.

405 The first split among the species attributed to the genus *Tritia* separated the Northwest Atlantic 406 species *T. obsoleta* from the East Atlantic/Mediterranean species. This node is inconsistent with the 407 results reported for the nassariid phylogeny (Galindo et al., 2016), in which *T. obsoleta* emerged in 408 an internal position within *Tritia*, although with low bootstrap support. In the past, *T. obsoleta* was 409 assigned as the type species to genus *Ilyanassa* (Stimpson, 1865), which was a taxon that included 410 nassariids from North America (Cernohorsky, 1984). Although *Ilyanassa* had been widely used in 411 previous literature, the internal position of *T. obsoleta* within *Tritia* in Galindo et al. (2016), led 412 these authors to synonymize *Ilyanassa* with *Tritia*. In contrast, our results reveal a correlation 413 between the phylogenetic position of *T. obsoleta* and its geographical distribution, which together 414 with a significant level of genetic divergence (13.7%) support the reinstatement of the genus 415 *Ilyanassa*. To further resolve this part of the tree, it would be also necessary to include *I. trivittata* 416 and other Northwest Atlantic nassariids (Rosenberg, 2009) into the phylogeny, as well as *T*. 417 *buchardi* from southern Australia and New Zealand, which is close to *Ilyanassa* according to the 418 phylogeny of Galindo et al. (2016). 419

420 **4.2. Taxonomy of *Tritia***

421 Within the mostly East Atlantic/Mediterranean group, Clade I comprised 10 species, which are 422 characterized by the presence of a shell with a more or less marked sculpture. The first diverging 423 lineage of Clade I included *T. ephamilla* and *T. elata* + *T. ovoidea*. The former species is endemic to 424 New Zealand and southern Australia whereas the latter two are found in the Mediterranean Sea and
along the West African coasts down to Angola. The inclusion of *T. ephamilla* within *Tritia* was also recovered by Galindo et al (2016), although its exact sister group was not resolved. In addition to the geographic disparity, there is also a prominent morphological difference between *T. ephamilla* (globose with nodulose shell sculpture) and *T. elata* and *T. ovoidea* (slender-ovoid with slight spiral striations). The presence of *T. ephamilla* in this part of the tree could reflect a Thetyan origin of the Northeast Atlantic/Mediterranean mud snails and a subsequent southwards expansion towards tropical waters in the West Africa coasts (Galindo et al., 2016). A striking example of a similar disjoint distribution is constituted by the seagrass genus *Posidonia*, which displays the same contrasting biogeographic pattern. One species, *P. oceanica*, is distributed in the Mediterranean Sea whereas the other species of the genus are restricted to subtropical and temperate Australian waters (Aires et al., 2011). Other Northeast Atlantic/Mediterranean gastropods with a similar disjoint distribution and suspected of having Thetyan origin are the top shell snails of the subfamily Cantharidinae (Uribe et al., 2017b). In this case, the endemics to Oceania were recovered as sister to all Atlantic/ Mediterranean taxa. *Tritia elata* is a circalittoral species typical of soft bottoms of the continental shelf from the southern Iberian Peninsula to Angola. It shows a quite variable shell sculpture along the West African coast where it has been sometimes recorded as *Nassarius semistriatus*, a very similar fossil species extinct during the Pleistocene (Adam & Knudsen, 1984). This last species, as well as the fossil species *Nassarius martinelli* (described by Gili, 1992 from the West Mediterranean Pliocene and differing from *T. elata* by its paucispiral protoconch) likely belong to the stem group of *Tritia*.

The other lineage within Clade I included *T. denticulata* sister to a rather diversified clade including *T. tingitana* sister to the “*T. cuvierii* complex” plus *T. reticulata* and *T. nitida*. The relative position of *T. denticulata* within clade I is in agreement with the Nassariidae molecular phylogeny (Galindo et al., 2016) but here showing high statistical support. *Tritia denticulata* is another circalittoral species distributed in the Alboran Sea and along the West African coasts down to Angola, including the Canary, Madeira and Cabo Verde islands (Rolán & Hernández, 2005). This species is morphologically similar to a group of West African species, namely *N. turbineus*, *N. webbei*, *N. desmoulioides*, and *N. arcadioi* (Rolán & Hernández, 2005), whose ascription to *Tritia* remains to be tested by their incorporation into the molecular phylogeny. In addition, the fossil species *N. clathratus* (Rolán & Hernández, 2005) could belong to the stem lineage of *Tritia*. The
phylogenetic position of *T. tingitana* (endemic of the Strait of Gibraltar), as the second diverging
lineage within Clade I, is in clear disagreement with Galindo et al. (2016), who recovered *T.
tingitana* as sister to *T. corniculum* with maximal support in a clade equivalent to our Clade III (see
below). The two species are morphologically very different, and difficult to misidentify, suggesting
a possible mislabeling error.

The last lineage of Clade I included three species corresponding to the “*T. cuvierii* complex”
plus *T. reticulata* and *T. nitida* (Galindo et al., 2016; Aissaoui, Galindo, Puillandre, & Bouchet,
2017). Within the “*T. cuvierii* complex”, *T. cuvierii* and *T. unifasciata* are sympatric along the
Mediterranean coasts, and differ in the size of their shell, protoconch and egg capsules (Moreno &
Templado, 1994). *Tritia unifasciata* has faint axial ribs only in the first whorls of the shell. The third
species of the complex here studied was *T. tenuicosta*, endemic to the Gulf of Gabès, whose relative
position was in agreement to that recovered by Aissaoui, Galindo, Puillandre, & Bouchet (2017).
Another two endemic species of the Gulf of Gabès, *T. lanceolata* and *T. djerbaensis* also belong to
the “*T. cuvierii* complex”, and *T. cuvierii* may also include additional local cryptic species (Aissaoui,
Galindo, Puillandre, & Bouchet, 2017). According to Galindo et al. (2016), the West African *T.
miga* would be closely related to the “*T. cuvierii* complex”. Given its distribution and morphological
similarity, the endemic species from Cabo Verde islands, *T. caboverdensis* (Rolán, 1984) may also
belong to this complex (pending molecular confirmation).

The closely related and well-known species *T. reticulata* and *T. nitida* have been widely studied
from different viewpoints (see Rolán & Luque, 1994; Albaina et al., 2012; Couceiro et al., 2012,
and references therein). Both species, sometimes misidentified, share similar life histories and
dispersal capacities, but differ in their environmental affinities. The netted dog-welk *T. reticulata*
inhabits a wide range of marine soft bottoms and is mainly an East Atlantic species with a more or
less continuous distribution from the British Isles to Morocco and the Canary Islands that enters
into western Mediterranean. The estuarine dog-welk *T. nitida* is restricted to sheltered brackish
habitats with a patchy distribution in the Mediterranean Sea and Black Sea coasts as well as in the
East Atlantic Ocean from Morocco to Norway (Albaina et al., 2012). Therefore, this latter species
probably diverged from *T. reticulata* by a habitat shift from marine to brackish waters.

Clade II in our phylogeny (also recovered in Galindo et al., 2016) included *T. varicosa* (often
previously recorded as *T. pygmaea*) and *T. incrassata*. Both species are sympatric in the
485 Mediterranean Sea and along the Northeast Atlantic coast from Sweden to the Iberian Peninsula.

486 They are similar in terms of shell size (about 10 mm) and ornament (little nodules on shell formed by costae and spiral ridges), but the nodules of *T. varicosa* are much smaller, squarer and more upstanding than those of *T. incrassata* (Fretter, Graham & Andrews, 1986). Divergence between

489 both species could have been caused by adaptation to dissimilar habitats: mainly rocky bottoms in *T. incrassata* and soft bottoms with beds of *Zostera* and *Cymodocea* seagrasses in *T. varicosa* (JT, pers. obs.). According to Galindo et al. (2016), *T. senegalensis* and *T. goreensis*, and several other species endemic to Senegal, would be sister to the northern species within this clade.

493 With a planktotrophic larval stage, *T. incrassata* is widely distributed along the Mediterranean and Northeast Atlantic region. The pairwise uncorrected *p* distance between the two studied individuals of *T. incrassata* from two distant locations (Norway and the Spanish Mediterranean) reached 3.8%, which was even higher than the value obtained for sibling species such as *T. neritea* and *T. pellucida* (Supporting Information Table S4). This result indicates that the diversity of *T. incrassata* was previously underestimated and that it may include putative cryptic species. This hypothesis will need to be further confirmed by the inclusion of data from nuclear genes to discard potential events of incomplete lineage sorting and hybridization (Alexander et al., 2017). A previous phylogeographic study on *T. nitida* suggested the existence of two glacial refugia during the Pleistocene, one in the Atlantic (around the Iberian Peninsula) and the other in the Paleo-Mediterranean Sea (Albaina et al., 2012). It may well be the case that the observed genetic divergence between the two populations of *T. incrassata*-NW and -SP is a result of isolation in the two Pleistocene refugia. After the last glaciation, the Strait of Gibraltar and the Almeria-Oran oceanographic front would have acted as well documented barriers to gene flow between the Atlantic and Mediterranean populations, as reported for *T. nitida* (Albaina et al., 2012) and other marine organisms (Patarnello, Volckaert, & Castilho, 2007). Our results show the need for further studies with dense sampling at the population level along the distribution range of *T. incrassata* in order to determine its diversity and elucidate the evolutionary processes driving its diversification.

511 Finally, Clade III in the reconstructed tree is composed of eight species with smooth (absence of sculpture) shells as shared derived character. The absence of sculpture may have occurred independently in several lineages of Nassariinae, some of which were already defined in the Early-Middle Miocene (Manganelli, Spadini & Martini, 2010). Phylogenetic relationships within
Clade III are in agreement with those previously reported (Galindo et al., 2016; Aissaoui, Galindo, 2015); here is moderately supported as sister to *T. pfeifferi* and was not resolved in previous studies. In Galindo et al. (2016), *T. conspersa* was recovered as sister to *T. pfeifferi* within this clade. The earliest divergent lineage within our Clade III includes species of the "*T. corniculum* complex". The pairwise uncorrected *p* distance obtained between *T. pallaryana* (a recently described species from the Gulf of Gabès; Aissaoui, Galindo, Puillandre, & Bouchet, 2017) and *T. corniculum* showed the least value of all comparisons (0.6%; Supporting Information Table S4), and could cast doubts on the taxonomic validity of the former. However, we conservatively maintain *T. pallaryana* as valid given significant ecological and morphological differences (Aissaoui, Galindo, Puillandre, & Bouchet, 2017) as well as because the sequence divergence value falls within the so-called "grey zone" of speciation (Roux et al., 2016). *Tritia elongata*, also endemic of the Gulf of Gabès, was also recovered within the "*T. corniculum* complex", in agreement with Aissaoui, Galindo, Puillandre, & Bouchet (2017). *Tritia corniculum* inhabits sandy-muddy bottoms in shallow, sheltered waters of brackish and marine environments, typically in coastal lagoons (Iannotta, Toscano, & Patti, 2009) with a patchy distribution along the Mediterranean and Northeast Atlantic coasts. Its direct development without a planktonic larval phase, together with the discontinuous distribution of its habitat, may promote genetic isolation among populations and incipient speciation processes, which are reflected in great intra-specific morphological variability (Iannotta, Toscano, & Patti, 2009; Gofas, 2011).

The remaining species of Clade III were *T. mutabilis* (type species of the currently unaccepted genus *Sphaeronassa*) and *T. pfeifferi* sister to a lineage comprising *T. grana* (type species of the currently unaccepted genus *Naytiopsis*) and the flattened-shell sibling species *T. neritea* and *T. pellucida* (previously assigned to the currently unaccepted genus *Cyclope*). While *T. mutabilis* inhabits infralittoral fine sands and muddy sand banks along the Mediterranean coasts, *T. pfeifferi* is an East Atlantic species (from northeastern Spain to Mauritania) adapted to shallow and intertidal sheltered soft bottoms that is replaced by *T. conspersa* in Canary Islands (Gofas, 2011). On the other hand, while *T. grana* inhabits clean, mobile, fine infralittoral sands in open waters, *T. neritea* and *T. pellucida* are characteristic of sandy-muddy shallow bottoms in estuaries, sheltered bays and marine lagoons, often subject to variable salinity (even brackish) waters (Gili & Martinell, 2000;
545 Gofas, 2011).

546

547 **4.3. Protoconch evolution**

548 The robust phylogeny of *Tritia* prompts some considerations about the evolution of the mode of
549 larval development within the genus. Since Thorson (1950), a large body of studies has focused on
550 the evolution of the mode of larval development in marine invertebrates. Transitions between modes
551 of development are considered highly asymmetrical from species with feeding larval stages
552 (planktotrophic) to species with non-feeding larval development (lecithotrophic), with the
553 evolutionary reversal of this transition considered very rare or impossible (Strathmann, 1978;
554 Modica et al., 2020). In both extant and fossil gastropods, the presence of a paucispiral or a
555 multispiral protoconch is used as proxy for the lecithotrophic and planktotrophic alternative modes
556 of development, respectively (Shuto, 1974; Jablonski & Lutz, 1980). The shift in mode of larval
557 development has important evolutionary consequences, as planktotrophic larvae are able to survive
558 longer in the water column and thus have higher dispersal capabilities than lecithotrophic larvae,
559 which barely move from the places where eggs were laid (Shank, 2009). Hence, species with
560 planktotrophic larvae usually are widely distributed and maintain high levels of gene flow between
561 populations, whereas species with lecithotrophic larvae show narrower distribution ranges and are
562 prone to allopatric speciation processes (Collin, 2001; Cunha et al., 2005; Abalde et al. 2017;
563 Modica, 2020).

564 The ancestor of the genus *Tritia* was inferred to have had planktotrophic veliger larvae, as is
565 widely assumed for caenogastropods (Ponder & Lindberg, 1997, 2019; Modica et al., 2020). The
566 lecithotrophic state is a derived condition, which evolved independently in at least three terminal
567 lineages: species within Clade III as well as species of the “*T. cuvierii* complex”, and *T. tingitana*
568 within Clade I. In addition, some but not all of the species of the “*T. denticulata* complex” have
569 planktotrophic development, and the potentially closely related *Nassarius turbineus* from the Gulf
570 of Guinea (not included in our phylogeny) exhibits paucispiral protoconch indicative of another
571 independent shift to lecithotrophy within Clade I. The loss of planktotrophy has occurred several
572 other times independently within Nassariinae including some *Reticunassa* species (Galindo et al.,
573 2016). Transitions from planktotrophic to lecithotrophic larval development have been recurrent in
574 diverse caenogastropod lineages (e.g. Lieberman, Allmon, & Eldredge, 1993; Reid, Rumbak, &
Thomas, 1996; Duda & Palumbi, 1999; Collin, 2004) and the fossil record from the Northeast Atlantic and Mediterranean shows this trend towards the loss of planktotrophy in several taxa as recently as the Pliocene and Pleistocene periods (Oliverio, 1996). During this evolutionary transition, intermediate forms of development (facultative feeding larvae, short-lived planktotrophs, species with dispersal polymorphisms, and species showing poecilogony, i.e., the presence of the two types of development) should appear (Collin, 2012). However, these intermediate stages are generally ephemeral in evolutionary terms, and either go extinct or evolve into direct developers. In rare cases, the phenotypically plastic intermediates might persist if they show a selective advantage under heterogeneous environmental conditions compared to the two extremes (Collin, 2012). In gastropods, the known cases of poecilogony are prevalent among sea slugs in the suborder Sacoglossa, short-lived planktotrophs are found among calyptraeids, and facultative lecithotrophy has been reported in the nudibranch genus *Phestilla* (Collin, 2012).

The frequent and asymmetric transition to lecithotrophy may have key macroevolutionary consequences because it implies shifts in rates of speciation and extinction. On the one hand, species with lecithotrophic development have higher rates of speciation, and thus they are assumed to progressively increase through time compared to widely dispersing species. For instance, Harzhauser & Kowalke (2004) studied the nassariid gastropods from the Neogene Paratethys and observed a trend in the fossil record towards the local appearance of species showing paucispiral protoconch in rather short time periods. On the other hand, regional environmental perturbations could cause the extinction of narrowly ranging species, whereas more broadly dispersed species could survive elsewhere (e.g., Jablonski, 1986). Therefore, higher extinction rates would lower the net diversity of lecithotrophs despite the fact that this condition originates frequently, and this evolutionary pattern could explain the long-term maintenance of planktotrophy (Collin & Moran, 2018; Modica et al., 2020). In this regard, several authors have highlighted that planktotrophic species are more resistant to extinction and persist longer on geological time scales relative to lecithotrophic species (e.g. Hansen, 1980; Jablonski, 1986; Crampton et al., 2010). For instance, planktotrophic species of the family Nassariidae from the Neogene fossil deposits of the Atlantic/Mediterranean area had a mean duration of 9.8 million years, whereas the average duration of several non-planktotrophic species was 2.8 million years (Gili & Martinell, 1994).

Given the high frequency of the transition to lecithotrophy (together with its enhancing effects
on extinction rates), it is likely that even if most living species of a taxon are included in a
molecular phylogeny, the total number of independent transitions to direct development will be
underestimated due to ignorance of what may have occurred in extinct lineages (Collin & Moran,
2018). Hence, fossil data appears necessary to testify and supplement the results of molecular data
to improve the knowledge of the evolution of this important trait in the life cycle in the East Atlantic
and Mediterranean nassariid gastropods (Lozouet & Galindo, 2015; Van Dingenen, Ceuleman,
Landau, & da Silva, 2015). For instance, the West Mediterranean Pliocene species *Nassarius
martinelli* is distinguished from the extant planktotrophic *T. elata* by its paucispiral protoconch,
indicative of lecithotrophic development (Gili, 1992). *Nassarius martinelli* was apparently quite
local in South France, inhabiting circallittoral bottoms, and became extinct at the end of the
Pliocene or during the first glacial period of the Pleistocene, probably as a consequence of
glacio-eustatic processes producing a sharp drop in sea level (Gili, 1992). *Tritia neritea* and *T.
pellucida* are the only extant species among Nassariinae gastropods with a depressed, discoidal shell,
dorsally convex and flatter in the ventral side. The early Pliocene fossil *Tritia migliorinii* (as
*Cyclope migliorinii*) shows the above-mentioned peculiar shell morphology (Gili & Martinell, 1999,
2000) and is considered a member of the stem group. It has a multispiral protoconch (i.e,
planktotrophic larval development) implying a putative independent transition to direct
development within this lineage of Clade III. Another instance of shift in mode of larval
development could be that of some fossil forms closely related to *T. mutabilis* from the
Mediterranean and Atlantic Pliocene that may have had a rather short planktotrophic phase (Gili &
Martinell, 1990) suggesting yet another transition from plankotrophy to lecithotrophy within Clade
III.

627

4.4. Tempo of diversification

Our reconstructed time tree dates the origin of Nassarinae genera throughout the Oligocene, which
is consistent with the fossil record (Lozouet & Galindo, 2015). A trend towards cooler conditions
during the Oligocene (Zachos et al., 2001) is likely to have favored temperate taxa (Uribe et al,
2017b). During this period, the onset of deep-water circulation between the Arctic and North
Atlantic Oceans (Davies et al., 2001) may have constituted a strong barrier to marine taxa dispersal
between North America and Eurasia. In this regard, the separation of *Ilyanassa* from *Tritia* at both
sides of the Atlantic and the subsequent diversification of the main eastern lineages occurred right after the Oligocene-Miocene transition. The late Oligocene is thought to have been a time of relatively warm global temperatures while in the early Miocene there was a rapid expansion of the Antarctica ice sheet and large-scale climate fluctuations attributed to an Earth orbital anomaly (Zachos et al., 2001). These global climate oscillations might have triggered diversification events leading to the extant genera within Nassariinae. In parallel, continental drift produced the collision of Laurasia and Gondwana about 19 Mya and the final closure of the Eastern Tethys Seaway (the so-called Terminal Tethyan event) around 14 Mya (Harzhauser et al., 2007; Okay, Zattin, & Cavazza, 2010; Hou & Li, 2018). The closure definitively ended the previous flow of tropical marine gastropods between the Indian Ocean and the proto-Mediterranean Basin (Hamon et al., 2013). In fact, fragmentation of the Tethys Ocean has been suggested as the main driver of the disjunct, endemic and relict distributions of many Tethyan lineages (i.e. Ozawa et al., 2009; Rodríguez-Flores et al., 2020). Accordingly, the puzzling phylogenetic position of the Australian and New Zealand endemic *T. ephamilla*, which diverged about 16 Mya from East Atlantic/ Mediterranean taxa (as well as the sister relationship of the Australian *T. buchardi* and the Northwest Atlantic *I. obsoleta*; Galindo et al. 2016), would represent one of the latest split events prior to the closure.

The origin of extant species started at the end of the Miocene, when there was a sustained global cooling event that started about 12 Mya and culminated about 5.4 Mya (Herbert et al., 2016), causing an eustatic sea level drop of -10 to -30 m (Hodell et al., 2001). Simultaneously, and because of tectonic movements, the connection between the East Atlantic Ocean and the Mediterranean Sea at the Strait of Gibraltar was closed, resulting in the desiccation of the Mediterranean basin, the well-known Messinian Salinity Crisis (MSC, 5.96 to 5.33 Mya; Krijgsman et al., 1999). Altogether, these climatic, eustatic, and tectonic changes resulted in drastic shifts in the paleoecosystems of the East Atlantic/Mediterranean region that likely triggered speciation, as has been shown for other groups (Davis et al., 2016, Abalde et al., 2017; Uribe et al., 2017b). The ecological adaptation to the variable salinity waters of this period likely promoted the divergence between *T. nitida* and *T. reticulata*, between *T. mutabilis* and *T. pfeifferi*, and between *T. grana* and *T. neritea* / *T. pellucida*, which were dated in the Miocene according to the chronogram. The independent transitions to the direct mode of larval development were also dated in this period. The transition from the Pliocene
to the Pleistocene was characterized by a sudden cooling (about 2.6 Mya) and subsequent
665
666
glacial-interglacial periods with extreme climate oscillations and drastic eustatic sea level changes
667
(Filippelli & Flores, 2009), which may explain the divergence between sibling taxa such as *T.*
668
*pallaryana* and *T. corniculum*, *T. neritea* and *T. pellucida*, and the two distant populations of *T.*
669
*incrassata*.

Our estimates of major divergence events within Nassariinae are about 11-24 Mya younger
670
than those previously reported in Galindo et al. (2016), which dated the origin of the lineage leading
671
to *Tritia* and *Ilyanassa* to 48 Mya (Eocene), separation of both genera at 32 Mya (Oligocene), and
672
the diversification of main lineages within *Tritia* during the Miocene. The main difference between
673
the two inferences relies on the calibration points, which in Galindo et al. (2016) were chosen to
674
date the much older Nassariidae origin within Buccinoidea (note that two calibration points were
675
the same in both studies) and the fewer representatives of *Tritia* in their chronogram. Given that our
676
calibration points were chosen to fit better in time the question at hand, and that there is congruence
677
of our estimated dates with climatic, eustatic, and tectonic changes, as well as the rich fossil record
678
of East Atlantic/Mediterranean Nassariinae during the Miocene and Pliocene (Cernohorsky, 1984),
679
we propose the present time tree as our preferred working hypothesis for the timing of the origin
680
and diversification of genus *Tritia*.
681

682

5. CONCLUSIONS
***

The reconstructed mitogenomic tree provided high resolution of *Tritia* phylogenetic relationships,
684
and allowed determining that *Gen. inc. sed. vaucheri* does not belong to the genus. Given its
685
distribution, this result calls for a morphological and ecological revision of the West African genus
686
*Naytia*. The strongly supported sister group relationship between the Northwest Atlantic *I. obsoleta*
687
and the three East Atlantic/Mediterranean clades (I-III), justify the reinstatement of the genus
688
*Ilyanassa*, and shows the need for a careful revision of the two genera. Within *Tritia*, the additional
689
described species endemic to West Africa need to be incorporated into the mitogenomic phylogeny.
690
In addition, the notable pairwise divergence between the two *T. incrassata* specimens from Norway
691
and the Spanish Mediterranean strongly suggests the possibility of the existence of cryptic species
692
that needs to be further confirmed with nuclear data and a larger sampling. The robust phylogeny
693
allowed inferring that the ancestor of Nassariinae had planktotrophic larvae and that several
694
695 independent transitions to lecithotrophic larvae occurred during the evolutionary history of the
group. Hence, it would be interesting to compare morphological and gene expression changes
696 during development of Nassariinae lecithotrophic larvae in search for footprints of convergence.
697 The inferred divergence dates suggest an important role of climatic, eustatic, and tectonic changes
698 during the Oligocene, Miocene, and Pleistocene influencing local Atlantic/Mediterranean
699 paleoecosystems and promoting the origin, main lineage diversification, and recent speciation
700 events within *Tritia*, respectively. Several recent divergences between some pairs of close species
701 could be due to a habitat shift from open marine to sheltered brackish waters. Our study indicates
702 that mt genomes could render similar robust results when applied to other Nassariinae genera and
703 would be highly useful in reconstructing a robust phylogeny of the entire Nassariidae family, hence
704 providing the long-needed framework to understand diversification and adaptation processes
705 underpinning the evolutionary success of this family.
706

707

708 ACKNOWLEDGEMENTS

709 We are grateful to the colleagues who helped us collect in the Gulf of Gabès or sent us specimens
710 from other Mediterranean, Atlantic, Australian and New-Zealand localities: Manuel Caballer, Serge
711 Gofas, Philippe Maestrati, Javier Martin, Jean Pierre Miquel, Leopoldo Moro, Marco Oliverio,
712 Gianni Spada, Jacques Pelorce, Emmanuel Vassard, and Bruce Marshall. Barbara Buge and Sadie
713 Mills are also thanked for their help in curating the MNHN and NIWA samples, respectively. We
714 thank two anonymous reviewers for detailed and insightful comments on a previous version of the
715 manuscript. This work was funded by the Spanish Ministry of Economy, Industry and
716 Competitiveness (CGL2016-75255-C2-1-P [AEI/FEDER, UE] to R.Z.; BES-2014-069575 to S.A).
717 Yi Yang was supported by a scholarship from the China Scholarship Council (CSC-201806330034)
718 for studying and living abroad. The project has received funding from the European Research
719 Council (ERC) under the European Union's Horizon 2020 research and innovation programme
720 (grant agreement No. 865101) to NP.
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1062 **Figure captions**

1063

1064 Figure 1. Gene order of *Tritia* mitogenomes. All newly sequenced mt genomes shared identical genome organization. The genes encoded in the major and minor strands are shown in the top and bottom rows, respectively. Genes encoding for proteins (blue), rRNAs (pink), and tRNAs (red) are depicted with different colors. The relative positions of the primers for long PCR are shown.

1069 Figure 2. Phylogenetic relationships within Nassariinae based on the nucleotide sequences of 13 protein-coding genes plus two rRNA genes. The Bayesian inference (BI) phylogram is shown. The non-nassariinae *Anentome* sp. was used as outgroup. The three Northeast clades are indicated. Numbers at nodes are statistical support values for ML (bootstrap proportions in percentage)/BI (posterior probabilities). Solid red circles represent nodes with maximum support values. The bar indicates substitutions per site. Additional *Tritia* species whose relative phylogenetic positions were resolved in Galindo et al. (2016) but were not included in this study are shown with a dashed line. Ventral images of the voucher specimens are shown.

1078 Figure 3. Ancestral character state reconstruction of *Tritia* protoconch (paucispiral *versus* multispiral) using the ML model Mk1 in Mesquite. Pie charts represent the statistical support at every node.

1081 Figure 4. Chronogram with age estimates of major divergence events within Nassariinae based on the nucleotide sequences of the concatenated mitochondrial protein coding genes, and using Bayesian relaxed dating methods (BEAST). Horizontal bars represent 95% credible intervals of relevant nodes, and calibration constraints are indicated with an asterisk on the corresponding nodes. Dates (and highest posterior density intervals, HPD) are in millions of years. Geological epochs are indicated.

1087
<figure><img src="image_2.png" /><figcaption>Tritia</figcaption></figure>

cox1

cox2

D

atp8

atp6

rrnS

V

rrnL

L1

L2

nad1

P

atp6

cob

S1

nad4L

nad4

H

nad5

F

CR

cox3

K

A

R

nad1

nad3

nad2
<figure><img src="image_3.png" /></figure>
<figure><img src="image_4.png" /></figure>

Tritia neritea

Tritia pellucida

Tritia grana

Tritia pfeifferi

Tritia mutabilis

Tritia pallaryana

Tritia corniculum

Tritia elongata

Tritia incrassata-SP

Tritia incrassata-NW

Tritia varicosa

Tritia cuvierii

Tritia tenuicosta

Tritia unifasciata

Tritia reticulata

Tritia nitida

Tritia tingitana

Tritia denticulata

Tritia ovoidea

Tritia elata

Tritia ephamilla

Ilyanassa obsoleta

Nassarius succinctus

Nassarius sinarus

Nassarius javanus

Nassarius foveolatus

Nassarius conoidalis

Nassarius nodifer

Nassarius pullus

Nassarius variciferus

“Tritia” vaucherii

Reticunassa hiradoensis

Reticunassa fratercula

Nassarius jacksonianus

Nassarius sp.

<figure><img src="image_5.png" /></figure>

<figure><img src="image_6.png" /></figure>

Paucispiral protoconch

<figure><img src="image_7.png" /></figure>

Multispiral protoconch
<figure><img src="image_8.png" /></figure>

Paleocene

Eocene

Oligocene

Miocene

Pli

Ple
<table><caption>Table 1. Mitochondrial (mt) genomes analyzed in this study.</caption><thead><tr><th>Species</th><th>Location</th><th>Collector</th><th>No. reads</th><th>Coverage (x)</th><th>Length (bp)</th><th>Genbank Acc. No.</th><th>Voucher</th></tr></thead><tbody><tr><td>Tritia corniculum</td><td>Ilha da Armona, Portugal</td><td>CLMA</td><td>34 502</td><td>75</td><td>15 114</td><td>MT755650</td><td>MNCN:ADN: 118970</td></tr><tr><td>Tritia cuvierii</td><td>Cabo de Palos, Spain</td><td>JT</td><td>36 582</td><td>306</td><td>15 185</td><td>MT755651</td><td>MNCN:ADN: 118971</td></tr><tr><td>Tritia grana</td><td>Cabo de Palos, Spain</td><td>JT</td><td>40 296</td><td>131</td><td>15 198</td><td>MT762151</td><td>MNCN:ADN: 118972</td></tr><tr><td>Tritia incrassata-SP</td><td>Cabo de Palos, Spain</td><td>JT</td><td>30 800</td><td>53</td><td>15 159</td><td>MT762153</td><td>MNCN:ADN: 118973</td></tr><tr><td>Tritia incrassata-NW</td><td>Bergen, Norway</td><td>David Osca</td><td>38 716</td><td>151</td><td>15 170</td><td>MT762152</td><td>MNCN:ADN: 118974</td></tr><tr><td>Tritia mutabilis</td><td>Market, Madrid, Spain</td><td>YY</td><td>40 182</td><td>250</td><td>15 148</td><td>MT762154</td><td>MNCN:ADN: 118975</td></tr><tr><td>Tritia neritea</td><td>Vigo, Spain</td><td>SA</td><td>35 082</td><td>216</td><td>15 179</td><td>MT762155</td><td>MNCN:ADN: 118976</td></tr><tr><td>Tritia pellucida</td><td>Mar Menor, Spain</td><td>JT</td><td>40 234</td><td>317</td><td>15 175</td><td>MT768039</td><td>MNCN:ADN: 118977</td></tr><tr><td>Tritia pfeifferi</td><td>Praia dos Cavacos, Portugal</td><td>CLMA</td><td>43 804</td><td>76</td><td>15 142</td><td>MT768040</td><td>MNCN:ADN: 118978</td></tr><tr><td>Tritia unifasciata</td><td>Mar Menor, Spain</td><td>JT</td><td>43 288</td><td>322</td><td>15 168</td><td>MT768043</td><td>MNCN:ADN: 118979</td></tr><tr><td>Tritia vaucherii</td><td>Isla Canela, Spain</td><td>CLMA</td><td>58 170</td><td>83</td><td>15 153</td><td>MT768044</td><td>MNCN:ADN: 118980</td></tr><tr><td>Tritia denticulata</td><td>Candelaria, Spain</td><td>—</td><td>2 142 656</td><td>19 059</td><td>15 167</td><td>MT755652</td><td>MNHN-IM-2009-21546</td></tr><tr><td>Tritia ephamilla</td><td>Kermadec Ridge, New Zealand</td><td>—</td><td>1 849 562</td><td>14 802</td><td>15 157</td><td>MT755654</td><td>MNHN-IM-2009-24014</td></tr><tr><td>Tritia pallaryana</td><td>Golfe de Gabès, Tunisie</td><td>Chrifa Aissaoui</td><td>2 515 502</td><td>21 974</td><td>15 144</td><td>MT762157</td><td>MNHN-IM-2009-30939</td></tr><tr><td>Tritia tenuicosta</td><td>Golfe de Gabès, Tunisie</td><td>Chrifa Aissaoui</td><td>2 494 246</td><td>19 378</td><td>15 169</td><td>MT768042</td><td>MNHN-IM-2013-31865</td></tr><tr><td>Tritia elongata</td><td>Errimel, Tunisie</td><td>Chrifa Aissaoui</td><td>2 133 232</td><td>18 698</td><td>15 145</td><td>MT755653</td><td>MNHN-IM-2013-32570</td></tr><tr><td>Tritia nitida</td><td>Vigo, Spain</td><td>Samuel Abalde</td><td>2 415 740</td><td>21 498</td><td>15 157</td><td>MT762156</td><td>MNCN:ADN: 118981</td></tr><tr><td>Tritia varicosa</td><td>Ponta da Piedade, Portugal</td><td>CLMA</td><td>2 765 768</td><td>23 350</td><td>15 142</td><td>MT768041</td><td>MNCN:ADN: 118982</td></tr><tr><td>Tritia elata</td><td>Ponta da Piedade, Portugal</td><td>CLMA</td><td>2 493 798</td><td>21 897</td><td>15 167</td><td>MT768045</td><td>MNCN:ADN: 118983</td></tr><tr><td>Tritia tingitana</td><td>Isla de Tarifa, Spain</td><td>Serge Gofas</td><td>—</td><td>—</td><td>13 337</td><td>MW194096</td><td>MNCN:ADN: 118984</td></tr><tr><td>Tritia ovoidea</td><td>Cañón de Carchuna, Spain</td><td>Serge Gofas</td><td>—</td><td>—</td><td>14 889</td><td>MW194095</td><td>MNCN:ADN: 118985</td></tr><tr><td>Anentome sp.</td><td>Aquarium market, Madrid, Spain</td><td>YY</td><td>2 528 802</td><td>19 514</td><td>15 069</td><td>MT755649</td><td>MNCN:ADN: 118986</td></tr></tbody></table>
Genbank mt genomes

<table><thead><tr><td>Species</td><td>Length (bp)</td><td>Accession No.</td><td>Reference</td></tr></thead><tbody><tr><td>Tritia obsoleta</td><td>15 263</td><td>DQ238598</td><td>Simison, Lindberg, &amp; Boore, 2006</td></tr><tr><td>Tritia reticulata</td><td>15 271</td><td>EU827201</td><td>Cunha, Grande, &amp; Zardoya, 2009</td></tr><tr><td>Nassarius variciferus</td><td>15 269</td><td>KM603509</td><td>Yang, Li, Kong, &amp; Yu, 2018</td></tr><tr><td>Nassarius succinctus</td><td>15 329</td><td>KT768016</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius nodifer</td><td>15 337</td><td>KT818617</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius conoidalis</td><td>15 332</td><td>KT826694</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius pullus</td><td>15 278</td><td>KT900947</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius sinarus</td><td>15 325</td><td>MH346208</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius foveolatus</td><td>15 343</td><td>MH346209</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius javanus</td><td>15 325</td><td>MH346210</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Reticunassa fratercula</td><td>15 174</td><td>KT826695</td><td>Yang, Li, Kong, &amp; Yu, 2018</td></tr><tr><td>Reticunassa hiradoensis</td><td>15 194</td><td>MG744569</td><td>Yang, Li, Kong, &amp; Yu, 2018</td></tr><tr><td>Nassarius jacksonianus</td><td>15 234</td><td>MH346212</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr><tr><td>Nassarius sp.</td><td>15 240</td><td>MH346211</td><td>Yang, Li, Kong, &amp; Yu, 2019</td></tr></tbody></table>
<table><caption>Table S1. Standard and Long PCR primers for *Tritia* mitogenomes.</caption><tbody><tr><td><i>Tritia corniculum</i></td><td colspan="3">Sequenced by Allgenetics, with a buccinoid species in the same library</td></tr><tr><td><strong>Long PCR</strong></td><td></td><td></td><td></td></tr><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td><td></td></tr><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td><td></td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td><td></td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td><td></td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td><td></td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td><td></td></tr><tr><td><strong>Primer Link cox1</strong></td><td></td><td></td><td></td></tr><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td><td></td></tr><tr><td>COI-tritiaF</td><td>ACWGCYCAYGCTTTYGTAATAAT</td><td>cox1 (400)</td><td></td></tr><tr><td>COI-tritiaR</td><td>TGYATNCCTCGTCATCGYATRTT</td><td></td><td></td></tr><tr><td><i>Tritia cuvierii</i></td><td colspan="3">Sequenced by Allgenetics, in a whole library</td></tr><tr><td><strong>Long PCR</strong></td><td></td><td></td><td></td></tr><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td><td></td></tr><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td><td></td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td><td></td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td><td></td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td><td></td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td><td></td></tr><tr><td><strong>Primer Link cox1</strong></td><td></td><td></td><td></td></tr><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td><td></td></tr><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td><td></td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td><td></td></tr><tr><td><i>Tritia grana</i></td><td colspan="3">Sequenced by Allgenetics, with a cone species in the same library</td></tr><tr><td><strong>Long PCR</strong></td><td></td><td></td><td></td></tr><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td><td></td></tr><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td><td></td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td><td></td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td><td></td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td><td></td></tr></tbody></table>
<table><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td></tr></tbody></table>

*Tritia incrassata-sp* Sequenced by Allgenetics, with a cone species in the same library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>

*Tritia incrassata-nw* Sequenced by Allgenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>

*Tritia mutabilis* Sequenced by Allgenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr></tbody></table>
<table><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>

*Tritia neritea*

Sequenced by Allgenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAAATCA</td><td></td></tr></tbody></table>

*Tritia pellucida*

Sequenced by Allgenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAAATCA</td><td></td></tr></tbody></table>

*Tritia pfeifferi*

Sequenced by Allgenetics, with a buccinoid species in the same library
## Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

## Primer Link cox1

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>

*Tritia unifascita* Sequenced by Allgenetics, in a whole library

## Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

## Primer Link cox1

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td></tr></tbody></table>

*Tritia vaucheri* Sequenced by Allgenetics, with a buccinoid species in the same library

## Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

## Primer Link cox1

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead></table>
<table><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></table>

*Tritia denticulata* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-trnF (4.0K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

cox1 Genbank Acc. No: KY451292

**Primer Link cytb**

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cytbF</td><td>GGWTAYGTWYTWCCWTGRGGWCARAT</td><td>cytb (0.5K)</td></tr><tr><td>cytbR</td><td>GCRTAWGCRAAWARRAARTAYCAYTCWGG</td><td></td></tr></tbody></table>

*Tritia ephamilla* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.4K)</td></tr><tr><td>coblaR</td><td>GCYCCTCARAARGATATYGACCTCARGG</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-trnF (4.0K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

cox1 Genbank Acc. No: KY451298

**Primer Link cytb**

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cytbF</td><td>GGWTAYGTWYTWCCWTGRGGWCARAT</td><td>cytb (0.5K)</td></tr><tr><td>cytbR</td><td>GCRTAWGCRAAWARRAARTAYCAYTCWGG</td><td></td></tr></tbody></table>

*Tritia pallaryana* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr></tbody></table>
<table><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></table>

**Primer Link cox1**

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COI-tritiaF</td><td>ACWGCYCAYGCTTTYGTAATAAT</td><td>cox1 (400)</td></tr><tr><td>COI-tritiaR</td><td>TGYATNCCTCGTCATCGYATRTT</td><td></td></tr></tbody></table>

*Tritia tenuicosta* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.4K)</td></tr><tr><td>coblaR</td><td>GCYCCTCARAARGATATYGACCTCARGG</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-trnF (4.0K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

cox1 Genbank Acc. No: KY489424

**Primer Link cytb**

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cytbF</td><td>GGWTAYGTWYTWCCWTGRGGWCARAT</td><td>cytb (0.5K)</td></tr><tr><td>cytbR</td><td>GCRTAWGCRAAWARRAARTAYCAYTCWGG</td><td></td></tr></tbody></table>

*Tritia elongata* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

cox1 Genbank Acc. No: KY489440

*Tritia nitida* Sequenced by NIMGenetics, in a whole library

**Long PCR**

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr></tbody></table>
<table><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td></tr></tbody></table>

*Tritia varicosa*

Sequenced by NIMGenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>16slaF</td><td>GTGCAAAGGTAGCATAATCATTTGCCT</td><td>rrnL-trnF (6.1K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>

*Tritia elata*

Sequenced by NIMGenetics, in a whole library

### Long PCR

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>cox1laF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>16slaF</td><td>GTGCAAAGGTAGCATAATCATTTGCCT</td><td>rrnL-trnF (6.1K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

### Primer Link cox1

<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>COINassaF</td><td>TCTACAAATCATAAAGACATTGG</td><td>cox1 (700)</td></tr><tr><td>COINassaR</td><td>ACYTCAGGATGACCAAARAATCA</td><td></td></tr></tbody></table>
Tritia ovoidea Sequenced by Sanger

## Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox11aF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (5.7K)</td></tr><tr><td>16SlaR</td><td>ATGCTGTTATCCCTACGGTAACTAATT</td><td></td></tr><tr><td>16SZouF</td><td>GCGGTACTCTGACCGTGCAA</td><td>rrnL-cob (2.5K)</td></tr><tr><td>coblaR</td><td>GCYCCTCARAARGATATYGACCTCARGG</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox11aR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr><tr><td>15F</td><td>TCAGCYTGTGGATTTGRARAT</td><td>trnH-trnF (1.6K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr></tbody></table>

## Walking Primers

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>1F</td><td>TGATTTCTTARGAGAGGTTCA</td><td>TrnQ-rrnS (1K)</td></tr><tr><td>1R</td><td>TGCTATGTTACGACTTATCTC</td><td></td></tr><tr><td>2F</td><td>AAGCTRTTGGGTTCATACCCC</td><td>TrnM-TrnE (0.5K)</td></tr><tr><td>2R</td><td>TAGATTTTCATTCTAAARGRATA</td><td></td></tr><tr><td>3F</td><td>ATGCTTGTAGACATTTTTTCTTC</td><td>atp6-trnC (0.9K)</td></tr><tr><td>3R</td><td>TCGATTTTGCARRTCGATGTT</td><td></td></tr><tr><td>4F</td><td>TTTTAYGGTCARTGTTCYGAAAT</td><td>cox2-atp6 (0.8K)</td></tr><tr><td>4R</td><td>ACAATATGYCCRGCYCTTATRTT</td><td></td></tr><tr><td>5F</td><td>ATGAGCCTATGAGGACAAYTAGGATT</td><td>cox2-trnD (0.7K)</td></tr><tr><td>5R</td><td>GYRACRTAAGRCTGACAAYCT</td><td></td></tr><tr><td>6F</td><td>GAYATTATRCTYCAYGAYACHTAYTA</td><td>cox1-cox2 (0.7K)</td></tr><tr><td>6R</td><td>TCATAGCTTCARTATCATTGRTG</td><td></td></tr><tr><td>7F</td><td>AAYATRCGATGACGAGGRATRCA</td><td>cox1 (0.9K)</td></tr><tr><td>7R</td><td>GCYTCYCABACAATTACYATRAA</td><td></td></tr><tr><td>9F</td><td>AGGGCTGCTAACTTTGTTTTG</td><td>trnS-cox1 (1.1K)</td></tr><tr><td>9R</td><td>ACRTTRTAAAGTTGRTCRCACC</td><td></td></tr><tr><td>10F</td><td>CTAGCTGTTAATTAGRAGAAT</td><td>trnN-nd2 (0.75K)</td></tr><tr><td>10R</td><td>ATRAARTAYTTWACRGCYGAYTG</td><td></td></tr><tr><td>11F</td><td>TTYTTYTGAGCTTAYTTTCAYAG</td><td>cox3-trnI (0.87K)</td></tr><tr><td>11R</td><td>ACATCAATGATTYCCGTTCAT</td><td></td></tr><tr><td>15F</td><td>TCAGCYTGTGGATTTGRARAT</td><td>nd5 (0.9K)</td></tr><tr><td>15R</td><td>TTRTGRATAAATATHCCDGCRA</td><td></td></tr><tr><td>16F</td><td>ATTGGNGCHAAYTAYGAAAAYGA</td><td>nd5-trnF (0.9K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr></tbody></table>

## Standard PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>12F</td><td>ATGGCRCGTAATCCTTTTCATTT</td><td>cox3-trnA (0.8K)</td></tr><tr><td>12R</td><td>CTRAATGCAAATCARRTCTTT</td><td></td></tr><tr><td>13F</td><td>TTTGAAGCTTCTTTAATTCCWAC</td><td>nd4-trnH (0.8K)</td></tr><tr><td>13R</td><td>ATYTYCAAATCCACARGCTGA</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-nd4 (0.9K)</td></tr></tbody></table>
<table><tr><td>14R</td><td>TTKACYCTATTTTGMCTYGCTAA</td><td></td></tr><tr><td>18F</td><td>DCATTCTGCCTGTTTAKCAAA</td><td>rrnL-trnL2 (0.8K)</td></tr><tr><td>18R</td><td>GGGCTTAAACCTAATGCACTT</td><td></td></tr><tr><td>19F</td><td>AATGCAATTGACTTAGGMTCA</td><td>trnL1-trnP (1K)</td></tr><tr><td>19R</td><td>ACCCAAGCTYTAGCTTCCAAT</td><td></td></tr><tr><td>20F</td><td>TTGGCAGTCGCTTTTTTTACTCT</td><td>nd1-nd6 (1K)</td></tr><tr><td>20R</td><td>CTAAARAYAATTARYGCTCTCAT</td><td></td></tr><tr><td>21F</td><td>ATTAGCATTGGAAGCTARAGC</td><td>trnP-cob (0.6K)</td></tr><tr><td>21R</td><td>AARATRGGRTGHACTTTTCGAAT</td><td></td></tr><tr><td>22F</td><td>ATGCGAAGTCCYATTCGAAAAGT</td><td>cob (1K)</td></tr><tr><td>22R</td><td>ACMGGACAAGCYCCAATTCAWGT</td><td></td></tr><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCAGGT</td><td>rrnL-rrnS (1.2K)</td></tr><tr><td>16sZouR</td><td>TCACGTAGAATTTTAATGGTCG</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-nd4 (1.2K)</td></tr><tr><td>nd4lnd4R</td><td>GGTAATGAAGCTGCTACAGTGTA</td><td></td></tr></table>

*Tritia tingitana* Sequenced by Sanger

## Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>16SZouF</td><td>GCGGTACTCTGACCGTGCAA</td><td>rrnL-cob (2.5K)</td></tr><tr><td>coblaR</td><td>GCYCCTCARAARGATATYGACCTCARGG</td><td></td></tr><tr><td>12F</td><td>ATGGCRCGTAATCCTTTTCATTT</td><td>cox3-cox1 (3K)</td></tr><tr><td>cox1laR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr><tr><td>5F</td><td>ATGAGCCTATGAGGACAAYTAGGATT</td><td>cox2-rrnS (3K)</td></tr><tr><td>12slaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>15F</td><td>TCAGCYTGTGGATTTGRARAT</td><td>trnH-trnF (1.6K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-trnH (2.5K)</td></tr><tr><td>13R</td><td>ATYTYCAAATCCACARGCTGA</td><td></td></tr></tbody></table>

## Walking Primers

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>1F</td><td>TGATTTCTTARGAGAGGTTCA</td><td>TrnQ-rrnS (1K)</td></tr><tr><td>1R</td><td>TGCTATGTTACGACTTATCTC</td><td></td></tr><tr><td>2F</td><td>AAGCTRTTGGGTTCATACCCC</td><td>TrnM-TrnE (0.5K)</td></tr><tr><td>2R</td><td>TAGATTTTCATTCTAAARGRATA</td><td></td></tr><tr><td>3F</td><td>ATGCTTGTAGACATTTTTTCTTC</td><td>atp6-trnC (0.9K)</td></tr><tr><td>3R</td><td>TCGATTTTCARRTCGATGTT</td><td></td></tr><tr><td>4F</td><td>TTTTAYGGTCARTGTTCYGAAAT</td><td>cox2-atp6 (0.8K)</td></tr><tr><td>4R</td><td>ACAATATGYCCRCGCYCTTATRTT</td><td></td></tr><tr><td>5F</td><td>ATGAGCCTATGAGGACAAYTAGGATT</td><td>cox2-trnD (0.7K)</td></tr><tr><td>5R</td><td>GYRACRTAAGRCTGACAAYCT</td><td></td></tr><tr><td>9F</td><td>AGGGCTGCTAACTTTGTTTTG</td><td>trnS-cox1 (1.1K)</td></tr></tbody></table>
<table><tr><td>9R</td><td>ACRTTRTAAAGTTGRTRCRTCACC</td><td></td></tr><tr><td>10F</td><td>CTAGCTGTTAATTAGRAGAAT</td><td>trnN-nd2 (0.75K)</td></tr><tr><td>10R</td><td>ATRAARTAYTTWACRGCYGAYTG</td><td></td></tr><tr><td>11F</td><td>TTYTTYTGAGCTTAYTTTCAYAG</td><td>cox3-trnI (0.87K)</td></tr><tr><td>11R</td><td>ACATCAATGATTYCCGTTCAT</td><td></td></tr><tr><td>12F</td><td>ATGGCRCGTAATCCTTTTCATTT</td><td>cox3-trnA (0.8K)</td></tr><tr><td>12R</td><td>CTRAATGCAAATCARRTCTTT</td><td></td></tr><tr><td>13F</td><td>TTTGAAGCTTCTTTAATTCCWAC</td><td>nd4-trnH (0.8K)</td></tr><tr><td>13R</td><td>ATYTYCAAATCCACARGCTGA</td><td></td></tr><tr><td>coblaF</td><td>GAYCCWGARAAYTTTATYCCTGCTAAYCC</td><td>cob-nd4 (0.9K)</td></tr><tr><td>14R</td><td>TTKACYCTATTTTGMCTYGCTAA</td><td></td></tr><tr><td>15F</td><td>TCAGCYTGTGGATTTGRARAT</td><td>nd5 (0.9K)</td></tr><tr><td>15R</td><td>TTRTGRATAAATATHCCDGCRCA</td><td></td></tr><tr><td>16F</td><td>ATTGGNGCHAAYTAYGAAAAYGA</td><td>nd5-trnF (0.9K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>18F</td><td>DCATTCTGCCTGTTTAKCAAA</td><td>rrnL-trnL2 (0.8K)</td></tr><tr><td>18R</td><td>GGGCTTAAACCTAATGCACTT</td><td></td></tr><tr><td>19F</td><td>AATGCAATTGACTTAGGMTCA</td><td>trnL1-trnP (1K)</td></tr><tr><td>19R</td><td>ACCCAAGCTYTAGCTTCCAAT</td><td></td></tr><tr><td>20F</td><td>TTGGCAGTCGCTTTTTTTACTCT</td><td>nd1-nd6 (1K)</td></tr><tr><td>20R</td><td>CTAAARAYAATTARYGCTCTCAT</td><td></td></tr><tr><td>21F</td><td>ATTAGCATTGGAAGCTARAGC</td><td>trnP-cob (0.6K)</td></tr><tr><td>21R</td><td>AARATRGGRTGHACTTTTCGAAT</td><td></td></tr><tr><td>22F</td><td>ATGCGAAGTCCYATTCGAAAAGT</td><td>cob (1K)</td></tr><tr><td>22R</td><td>ACMGGACAAGCYCCAATTCAWGT</td><td></td></tr></table>

Standard PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnL-rrnS (1.2K)</td></tr><tr><td>16sZouR</td><td>TCACGTAGAATTTTAATGGTCG</td><td></td></tr><tr><td>cob-12870F</td><td>GGGTATGTCTTACCCTGAGGTCAGAT</td><td>cob (0.4K)</td></tr><tr><td>cob-12870R</td><td>GCATACGCAAAGAGGAAGTATCATTCTGG</td><td></td></tr></tbody></table>

Anentome sp. Sequenced by NIMGenetics, in a whole library

Long PCR

<table><thead><tr><th>Primer</th><th>Sequence 5'-3'</th><th>Fragment (bp)</th></tr></thead><tbody><tr><td>cox11aF</td><td>AAYATRCGATGACGAGGRATRCARTTTGA</td><td>cox1-rrnS (4K)</td></tr><tr><td>12SlaR</td><td>ATAGGCTGCACCTTGATCTGACGTYTT</td><td></td></tr><tr><td>12SlaF</td><td>CAGTTTGTATACCGTCGTCGTCAGGT</td><td>rrnS-cob (4.5K)</td></tr><tr><td>cobla2R</td><td>GGRTTAGCWGGRATAAARTTYTCWGGRTC</td><td></td></tr><tr><td>cobla2F</td><td>CCYTGAGGTCARATATCYTTYTGAGGRGC</td><td>cob-trnF (4.3K)</td></tr><tr><td>trnF-R</td><td>CACCCTAGCATCTTCAGTGCTATGCTCT</td><td></td></tr><tr><td>cox3la2F</td><td>ATGGCRCGNAATCCTTTTCATTTAGTWGARTT</td><td>cox3-cox1 (2.5K)</td></tr><tr><td>cox11aR</td><td>CGAGGAAAAGCYATRTCAGGAGC</td><td></td></tr></tbody></table>

Primer Link cox1
<table><thead><tr><td>Primer</td><td>Sequence 5'-3'</td><td>Fragment (bp)</td></tr></thead><tbody><tr><td>LCO1490 F</td><td>GGTCAACAAATCATAAAGATATTGG</td><td>cox1 (709)</td></tr><tr><td>HCO2198 R</td><td>TTAACTTCAGGGTGACCAAAAAATCA</td><td></td></tr></tbody></table>
<table><caption>Table S2. Best fit partitions and substitution models.</caption><thead><tr><th></th><th>Set Partition<br>(Conducted by<br>PartitionFinder 2)</th><th>Best Model for BI<br>(Selected by<br>PartitionFinder 2)</th><th>Best Model for<br>ML (Selected by<br>IQtree 1.6.10)</th></tr></thead><tbody><tr><td rowspan="10">Best Partition CDS genes<br>(BIC= 254577.82)</td><td>atp6-8 1th</td><td>TRN+G</td><td>TN+F+G4</td></tr><tr><td>atp6-8 2th</td><td>K81UF+I</td><td>K3Pu+F+I</td></tr><tr><td>atp6-8 3th</td><td>TVM+I+G</td><td>K3Pu+F+I+G4</td></tr><tr><td>cox1-2-3 1th</td><td>TRN+I+G</td><td>TN+F+I+G4</td></tr><tr><td>cox1-2-3 2th</td><td>TVM+I+G</td><td>TVM+F+I+G4</td></tr><tr><td>cox1-2-3 3th</td><td>GTR+I+G</td><td>TVM+F+I+G4</td></tr><tr><td>cob 1th</td><td>TRN+I+G</td><td>TIM2+F+I+G4</td></tr><tr><td>cob 2th</td><td>TVM+I+G</td><td>TVM+F+I+G4</td></tr><tr><td>cob 3th</td><td>K81UF+I+G</td><td>K3Pu+F+I+G4</td></tr><tr><td>nad1-2-3-4-4L-5-6 1th</td><td>GTR+I+G</td><td>TIM3+F+I+G4</td></tr><tr><td rowspan="3">Best Partition to rRNA genes<br>(BIC= 26625.61)</td><td>nad1-2-3-4-4L-5-6 2th</td><td>GTR+I+G</td><td>GTR+F+I+G4</td></tr><tr><td>nad1-2-3-4-4L-5-6 3th</td><td>GTR+I+G</td><td>TVM+F+I+G4</td></tr><tr><td>rrnS-L</td><td>GTR+I+G</td><td>GTR+F+I+G4</td></tr></tbody></table>
<caption>Table S3. Mitochondrial genome features</caption>

*Tritia corniculum* 15,114 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-747</td><td>incomplete</td><td>—</td><td>TAG</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>763-827</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>829-896</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>906-974</td><td>69</td><td>—</td><td>—</td><td>5</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>980-1047</td><td>68</td><td>—</td><td>—</td><td>15</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1063-1130</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1134-1487</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1488-1555</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1556-2610</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2611-4146</td><td>1536</td><td>ATG</td><td>TAA</td><td>29</td></tr><tr><td>Cox2</td><td>H</td><td>4176-4862</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-ASp</td><td>H</td><td>4861-4928</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4930-5088</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5096-5791</td><td>696</td><td>ATG</td><td>TAA</td><td>38</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5830-5896</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5900-5967</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5969-6033</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6034-6099</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6101-6162</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6164-6231</td><td>68</td><td>—</td><td>—</td><td>11</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6243-6307</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6308-7265</td><td>958</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7266-7333</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7334-8685</td><td>1352</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8686-8757</td><td>72</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8760-8828</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8829-9770</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9777-9844</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9846-10346</td><td>501</td><td>ATG</td><td>TAG</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10352-11491</td><td>1140</td><td>ATG</td><td>TAA</td><td>9</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11502-11565</td><td>64</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11566-11630</td><td>65</td><td>—</td><td>—</td><td>20</td></tr><tr><td>Nad4L</td><td>H</td><td>11651-11947</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11959-13314</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13314-13378</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad5</td><td>H</td><td>13379-15100</td><td>1722</td><td>ATG</td><td>TAG</td><td>14</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia cuvierii 15,185 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-776</td><td>incomplete</td><td>—</td><td>TAA</td><td>12</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>789-853</td><td>65</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>856-922</td><td>67</td><td>—</td><td>—</td><td>10</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>933-1001</td><td>69</td><td>—</td><td>—</td><td>8</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1010-1077</td><td>68</td><td>—</td><td>—</td><td>17</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1095-1162</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1166-1519</td><td>354</td><td>ATG</td><td>TAG</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1520-1587</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1588-2642</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2643-4178</td><td>1536</td><td>ATG</td><td>TAA</td><td>26</td></tr><tr><td>Cox2</td><td>H</td><td>4205-4891</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-ASp</td><td>H</td><td>4890-4957</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4959-5117</td><td>159</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Atp6</td><td>H</td><td>5124-5819</td><td>696</td><td>ATG</td><td>TAA</td><td>42</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5862-5928</td><td>67</td><td>—</td><td>—</td><td>4</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5933-5999</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6001-6066</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6067-6132</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6133-6196</td><td>64</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6198-6264</td><td>67</td><td>—</td><td>—</td><td>14</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6279-6343</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6344-7302</td><td>959</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7303-7370</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7371-8723</td><td>1353</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8724-8792</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8795-8863</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8864-9805</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9811-9878</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9880-10380</td><td>501</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10386-11525</td><td>1140</td><td>ATG</td><td>TAA</td><td>9</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11535-11599</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11600-11667</td><td>68</td><td>—</td><td>—</td><td>25</td></tr><tr><td>Nad4L</td><td>H</td><td>11693-11989</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>12001-13356</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13356-13421</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13423-15144</td><td>1722</td><td>ATG</td><td>TAG</td><td>41</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia denticulata 15,167 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-771</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>787-852</td><td>66</td><td></td><td></td><td>3</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>856-923</td><td>68</td><td></td><td></td><td>8</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>932-1002</td><td>71</td><td></td><td></td><td>7</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1010-1076</td><td>67</td><td></td><td></td><td>17</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1094-1161</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1165-1518</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1519-1586</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1587-2641</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2642-4177</td><td>1536</td><td>ATG</td><td>TAA</td><td>22</td></tr><tr><td>Cox2</td><td>H</td><td>4200-4886</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4885-4952</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4954-5112</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5120-5815</td><td>696</td><td>ATG</td><td>TAA</td><td>42</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5858-5924</td><td>67</td><td></td><td></td><td>6</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5931-5998</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6000-6063</td><td>64</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6064-6129</td><td>66</td><td></td><td></td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6127-6193</td><td>67</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6193-6261</td><td>69</td><td></td><td></td><td>12</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6274-6338</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6339-7296</td><td>958</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7297-7364</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7365-8716</td><td>1352</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8717-8785</td><td>69</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8788-8856</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8857-9798</td><td>942</td><td>ATG</td><td>TAG</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9805-9872</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9874-10374</td><td>501</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Cytb</td><td>H</td><td>10381-11520</td><td>1140</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11531-11595</td><td>65</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11595-11664</td><td>70</td><td></td><td></td><td>18</td></tr><tr><td>Nad4L</td><td>H</td><td>11683-11979</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11991-13346</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13346-13412</td><td>67</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13414-15135</td><td>1722</td><td>ATG</td><td>TAG</td><td>32</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia elongata 15,145 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-752</td><td>incomplete</td><td>—</td><td>TAA</td><td>12</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>765-831</td><td>67</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>831-900</td><td>70</td><td>—</td><td>—</td><td>7</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>908-978</td><td>71</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>981-1047</td><td>67</td><td>—</td><td>—</td><td>14</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1062-1129</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1133-1486</td><td>354</td><td>ATG</td><td>TAG</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1487-1554</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1555-2609</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2610-4145</td><td>1536</td><td>ATG</td><td>TAA</td><td>29</td></tr><tr><td>Cox2</td><td>H</td><td>4175-4861</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-ASp</td><td>H</td><td>4860-4927</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4929-5087</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5095-5790</td><td>696</td><td>ATG</td><td>TAA</td><td>38</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5829-5895</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5897-5968</td><td>72</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5968-6032</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6033-6098</td><td>66</td><td>—</td><td>—</td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6096-6162</td><td>67</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6163-6231</td><td>69</td><td>—</td><td>—</td><td>12</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6244-6308</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6309-7267</td><td>959</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7268-7335</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7336-8686</td><td>1351</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8687-8756</td><td>70</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8758-8828</td><td>71</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad1</td><td>H</td><td>8828-9769</td><td>942</td><td>ATG</td><td>TAG</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9776-9843</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9845-10345</td><td>501</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10351-11490</td><td>1140</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11499-11569</td><td>71</td><td>—</td><td>—</td><td>-2</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11568-11635</td><td>68</td><td>—</td><td>—</td><td>19</td></tr><tr><td>Nad4L</td><td>H</td><td>11655-11951</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11963-13318</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13318-13382</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad5</td><td>H</td><td>13383-15104</td><td>1722</td><td>ATG</td><td>TAG</td><td>41</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia ephamilla 15,186 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-776</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>792-856</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>858-925</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>935-1003</td><td>69</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1013-1080</td><td>68</td><td>—</td><td>—</td><td>18</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1099-1166</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1170-1523</td><td>354</td><td>ATG</td><td>TAA</td><td>-1</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1523-1592</td><td>70</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad2</td><td>H</td><td>1592-2646</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2647-4182</td><td>1536</td><td>ATG</td><td>TAA</td><td>27</td></tr><tr><td>Cox2</td><td>H</td><td>4210-4896</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4895-4962</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4964-5122</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5130-5825</td><td>696</td><td>ATG</td><td>TAA</td><td>43</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5869-5935</td><td>67</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5938-6003</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6005-6069</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6070-6135</td><td>66</td><td>—</td><td>—</td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6133-6199</td><td>67</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6199-6267</td><td>69</td><td>—</td><td>—</td><td>13</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6281-6345</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6346-7301</td><td>956</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7302-7369</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7370-8719</td><td>1350</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8720-8788</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8791-8861</td><td>71</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad1</td><td>H</td><td>8861-9802</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9809-9876</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9878-10378</td><td>501</td><td>ATG</td><td>TAA</td><td>4</td></tr><tr><td>Cytb</td><td>H</td><td>10383-11522</td><td>1140</td><td>ATG</td><td>TAA</td><td>16</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11539-11603</td><td>65</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11603-11673</td><td>71</td><td>—</td><td>—</td><td>20</td></tr><tr><td>Nad4L</td><td>H</td><td>11694-11990</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>12002-13357</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13357-13422</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13424-15145</td><td>1722</td><td>ATG</td><td>TAG</td><td>41</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia grana 15,198 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-770</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>786-850</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>851-918</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>928-996</td><td>69</td><td>—</td><td>—</td><td>16</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1013-1080</td><td>68</td><td>—</td><td>—</td><td>12</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1093-1160</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1164-1517</td><td>354</td><td>ATG</td><td>TAG</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1518-1585</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1586-2641</td><td>1056</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2642-4177</td><td>1536</td><td>ATG</td><td>TAA</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4206-4892</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4891-4958</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4960-5118</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5126-5821</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5863-5929</td><td>67</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5932-5999</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6001-6066</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6067-6132</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6134-6195</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6197-6264</td><td>68</td><td>—</td><td>—</td><td>12</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6277-6341</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6342-7302</td><td>961</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7303-7370</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7371-8720</td><td>1350</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8721-8789</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8792-8860</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8861-9802</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9809-9876</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9878-10378</td><td>501</td><td>ATG</td><td>TAA</td><td>18</td></tr><tr><td>Cytb</td><td>H</td><td>10397-11536</td><td>1140</td><td>ATG</td><td>TAA</td><td>12</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11549-11614</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11615-11682</td><td>68</td><td>—</td><td>—</td><td>23</td></tr><tr><td>Nad4L</td><td>H</td><td>11706-12002</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>12014-13369</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13369-13434</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13436-15157</td><td>1722</td><td>ATG</td><td>TAG</td><td>41</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia incrassata-Norway 15,170 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-772</td><td>incomplete</td><td>—</td><td>TAA</td><td>14</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>787-851</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>853-920</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>930-998</td><td>69</td><td>—</td><td>—</td><td>7</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1006-1072</td><td>67</td><td>—</td><td>—</td><td>15</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1088-1154</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1158-1511</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1512-1579</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1580-2634</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2635-4170</td><td>1536</td><td>ATG</td><td>TAG</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4199-4885</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4884-4951</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4953-5111</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5119-5814</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5856-5922</td><td>67</td><td>—</td><td>—</td><td>5</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5928-5994</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5996-6061</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6062-6127</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6129-6190</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6192-6258</td><td>67</td><td>—</td><td>—</td><td>13</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6272-6336</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6337-7292</td><td>956</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7293-7360</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7361-8711</td><td>1351</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8712-8780</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8783-8851</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8852-9793</td><td>942</td><td>ATG</td><td>TAG</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9800-9867</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9869-10369</td><td>501</td><td>ATG</td><td>TAA</td><td>4</td></tr><tr><td>Cytb</td><td>H</td><td>10374-11513</td><td>1140</td><td>ATG</td><td>TAA</td><td>11</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11525-11590</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11591-11658</td><td>68</td><td>—</td><td>—</td><td>19</td></tr><tr><td>Nad4L</td><td>H</td><td>11678-11974</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11986-13341</td><td>1356</td><td>ATA</td><td>TAA</td><td>1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13343-13407</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad5</td><td>H</td><td>13408-15129</td><td>1722</td><td>ATG</td><td>TAG</td><td>41</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia incrassata-Spain 15,159 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-775</td><td>incomplete</td><td>—</td><td>TAA</td><td>14</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>790-854</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>856-923</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>933-1001</td><td>69</td><td>—</td><td>—</td><td>7</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1009-1075</td><td>67</td><td>—</td><td>—</td><td>15</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1091-1157</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1161-1514</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1515-1582</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1583-2637</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2638-4173</td><td>1536</td><td>ATG</td><td>TAG</td><td>27</td></tr><tr><td>Cox2</td><td>H</td><td>4201-4887</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4886-4953</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4955-5113</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5121-5816</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5858-5924</td><td>67</td><td>—</td><td>—</td><td>5</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5930-5996</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5998-6062</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6063-6128</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6130-6191</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6193-6259</td><td>67</td><td>—</td><td>—</td><td>13</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6273-6337</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6338-7293</td><td>956</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7294-7361</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7362-8712</td><td>1351</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8713-8781</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8784-8852</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8853-9794</td><td>942</td><td>ATG</td><td>TAG</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9801-9868</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9870-10370</td><td>501</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10376-11515</td><td>1140</td><td>ATG</td><td>TAA</td><td>11</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11527-11591</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11592-11659</td><td>68</td><td>—</td><td>—</td><td>19</td></tr><tr><td>Nad4L</td><td>H</td><td>11679-11975</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11987-13342</td><td>1356</td><td>ATA</td><td>TAA</td><td>1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13344-13408</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad5</td><td>H</td><td>13409-15130</td><td>1722</td><td>ATG</td><td>TAG</td><td>29</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia mutabilis 15,148 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-750</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>766-830</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>832-899</td><td>68</td><td>—</td><td>—</td><td>7</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>907-975</td><td>69</td><td>—</td><td>—</td><td>10</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>986-1053</td><td>68</td><td>—</td><td>—</td><td>16</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1070-1137</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1141-1494</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1495-1562</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1563-2617</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2618-4153</td><td>1536</td><td>ATG</td><td>TAA</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4182-4868</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4867-4934</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4936-5094</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5102-5797</td><td>696</td><td>ATG</td><td>TAG</td><td>42</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5840-5906</td><td>67</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5909-5975</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5977-6042</td><td>66</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6043-6108</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6110-6171</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6173-6239</td><td>67</td><td>—</td><td>—</td><td>14</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6254-6318</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6319-7278</td><td>960</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7279-7346</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7347-8699</td><td>1353</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8700-8768</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8771-8839</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8840-9781</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9788-9855</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9857-10357</td><td>501</td><td>ATG</td><td>TAG</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10363-11502</td><td>1140</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11513-11577</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11578-11645</td><td>68</td><td>—</td><td>—</td><td>21</td></tr><tr><td>Nad4L</td><td>H</td><td>11667-11963</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11975-13330</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13330-13395</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13397-15118</td><td>1722</td><td>ATG</td><td>TAG</td><td>30</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia neritea 15,179 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-771</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>787-851</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>852-919</td><td>68</td><td></td><td></td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>929-997</td><td>69</td><td></td><td></td><td>16</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1014-1081</td><td>68</td><td></td><td></td><td>18</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1100-1167</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1171-1524</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1525-1592</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1593-2647</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2648-4183</td><td>1536</td><td>ATG</td><td>TAA</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4212-4898</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4897-4964</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4966-5124</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5132-5827</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5869-5935</td><td>67</td><td></td><td></td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5939-6006</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6008-6071</td><td>64</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6072-6137</td><td>66</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6139-6200</td><td>62</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6202-6268</td><td>67</td><td></td><td></td><td>11</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6280-6344</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6345-7300</td><td>956</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7301-7368</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7369-8720</td><td>1352</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8721-8789</td><td>69</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8792-8860</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8861-9802</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9808-9875</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9877-10377</td><td>501</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>Cytb</td><td>H</td><td>10388-11527</td><td>1140</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11536-11600</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11601-11667</td><td>67</td><td></td><td></td><td>22</td></tr><tr><td>Nad4L</td><td>H</td><td>11690-11986</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11998-13353</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13353-13421</td><td>69</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13423-15144</td><td>1722</td><td>ATG</td><td>TAG</td><td>35</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia nitida 15,157 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-750</td><td>incomplete</td><td>—</td><td>TAA</td><td>12</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>763-827</td><td>65</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>829-898</td><td>70</td><td></td><td></td><td>6</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>905-975</td><td>71</td><td></td><td></td><td>8</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>984-1051</td><td>68</td><td></td><td></td><td>20</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1072-1139</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1143-1496</td><td>354</td><td>ATG</td><td>TAG</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1497-1564</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1565-2619</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2620-4155</td><td>1536</td><td>ATG</td><td>TAA</td><td>26</td></tr><tr><td>Cox2</td><td>H</td><td>4182-4868</td><td>687</td><td>ATG</td><td>TAA</td><td>-1</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4868-4935</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4937-5095</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5103-5798</td><td>696</td><td>ATG</td><td>TAG</td><td>43</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5842-5908</td><td>67</td><td></td><td></td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5912-5982</td><td>71</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5982-6045</td><td>64</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6046-6111</td><td>66</td><td></td><td></td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6109-6175</td><td>67</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6175-6243</td><td>69</td><td></td><td></td><td>14</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6258-6322</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6323-7282</td><td>960</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7283-7349</td><td>67</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7350-8701</td><td>1352</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8702-8770</td><td>69</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8772-8840</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8841-9782</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9788-9855</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9857-10357</td><td>501</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Cytb</td><td>H</td><td>10364-11503</td><td>1140</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11512-11576</td><td>65</td><td></td><td></td><td>3</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11580-11649</td><td>70</td><td></td><td></td><td>18</td></tr><tr><td>Nad4L</td><td>H</td><td>11668-11964</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11976-13331</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13331-13396</td><td>66</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13398-15119</td><td>1722</td><td>ATG</td><td>TAG</td><td>38</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia pallaryana 15,144 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-753</td><td>incomplete</td><td>—</td><td>TAG</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>769-833</td><td>65</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>835-902</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>912-980</td><td>69</td><td>—</td><td>—</td><td>5</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>986-1053</td><td>68</td><td>—</td><td>—</td><td>15</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1069-1136</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1140-1493</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1494-1561</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1562-2616</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2617-4152</td><td>1536</td><td>ATG</td><td>TAA</td><td>29</td></tr><tr><td>Cox2</td><td>H</td><td>4182-4868</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4867-4934</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4936-5094</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5102-5797</td><td>696</td><td>ATG</td><td>TAA</td><td>38</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5836-5902</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5906-5973</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5975-6039</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6040-6105</td><td>66</td><td>—</td><td>—</td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6103-6169</td><td>67</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6169-6238</td><td>70</td><td>—</td><td>—</td><td>10</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6249-6313</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6314-7271</td><td>958</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7272-7339</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7340-8690</td><td>1351</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8691-8762</td><td>72</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8764-8834</td><td>71</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad1</td><td>H</td><td>8834-9775</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9782-9849</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9851-10351</td><td>501</td><td>ATG</td><td>TAG</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10357-11496</td><td>1140</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11507-11570</td><td>64</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11570-11636</td><td>67</td><td>—</td><td>—</td><td>19</td></tr><tr><td>Nad4L</td><td>H</td><td>11656-11952</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11964-13319</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13319-13383</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad5</td><td>H</td><td>13384-15105</td><td>1722</td><td>ATG</td><td>TAG</td><td>39</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia pellucida 15,175 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-771</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>787-851</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>852-919</td><td>68</td><td>—</td><td>—</td><td>9</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>929-997</td><td>69</td><td>—</td><td>—</td><td>16</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1014-1081</td><td>68</td><td>—</td><td>—</td><td>18</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1100-1167</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1171-1524</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1525-1592</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1593-2647</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2648-4183</td><td>1536</td><td>ATG</td><td>TAA</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4212-4898</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4897-4964</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4966-5124</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5132-5827</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5869-5935</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5939-6006</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>6008-6072</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6073-6138</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6140-6201</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6203-6269</td><td>67</td><td>—</td><td>—</td><td>11</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6281-6345</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6346-7301</td><td>956</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7202-7369</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7370-8718</td><td>1349</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8719-8787</td><td>69</td><td>—</td><td>—</td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8790-8858</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8859-9800</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9806-9873</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9875-10375</td><td>501</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>Cytb</td><td>H</td><td>10386-11525</td><td>1140</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11534-11598</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11599-11665</td><td>67</td><td>—</td><td>—</td><td>22</td></tr><tr><td>Nad4L</td><td>H</td><td>11688-11984</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11996-13351</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13351-13417</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13419-15140</td><td>1722</td><td>ATG</td><td>TAG</td><td>35</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia pfeifferi 15,142 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-747</td><td>incomplete</td><td>—</td><td>TAA</td><td>15</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>763-827</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>828-895</td><td>68</td><td></td><td></td><td>5</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>901-969</td><td>69</td><td></td><td></td><td>16</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>986-1053</td><td>68</td><td></td><td></td><td>14</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1068-1135</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1139-1492</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1493-1560</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1561-2615</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2616-4151</td><td>1536</td><td>ATG</td><td>TAA</td><td>28</td></tr><tr><td>Cox2</td><td>H</td><td>4180-4866</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4865-4932</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4934-5092</td><td>159</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>Atp6</td><td>H</td><td>5101-5796</td><td>696</td><td>ATG</td><td>TAA</td><td>42</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5839-5905</td><td>67</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5907-5974</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5976-6041</td><td>66</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6042-6107</td><td>66</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6109-6170</td><td>62</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6172-6238</td><td>67</td><td></td><td></td><td>12</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6251-6315</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6316-7271</td><td>956</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7272-7339</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7340-8691</td><td>1352</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8692-8760</td><td>69</td><td></td><td></td><td>3</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8764-8832</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8833-9774</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9781-9848</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9850-10350</td><td>501</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>Cytb</td><td>H</td><td>10356-11495</td><td>1140</td><td>ATG</td><td>TAA</td><td>10</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11506-11570</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11571-11638</td><td>68</td><td></td><td></td><td>21</td></tr><tr><td>Nad4L</td><td>H</td><td>11660-11956</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11968-13323</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13323-13388</td><td>66</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13390-15111</td><td>1722</td><td>ATG</td><td>TAG</td><td>31</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia varicosa 15,142 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-762</td><td>incomplete</td><td>—</td><td>TAA</td><td>18</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>781-847</td><td>67</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>847-916</td><td>70</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>917-985</td><td>69</td><td></td><td></td><td>20</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1006-1072</td><td>67</td><td></td><td></td><td>14</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1087-1154</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1158-1511</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1512-1579</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1580-2634</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2635-4170</td><td>1536</td><td>ATG</td><td>TAG</td><td>22</td></tr><tr><td>Cox2</td><td>H</td><td>4193-4879</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4878-4945</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4947-5105</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5113-5808</td><td>696</td><td>ATG</td><td>TAA</td><td>41</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5850-5916</td><td>67</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5919-5985</td><td>67</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5987-6051</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6052-6117</td><td>66</td><td></td><td></td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6115-6181</td><td>67</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6182-6248</td><td>67</td><td></td><td></td><td>13</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6262-6326</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6327-7283</td><td>957</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7284-7351</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7352-8701</td><td>1350</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8702-8770</td><td>69</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8773-8841</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8842-9783</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9789-9856</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9858-10358</td><td>501</td><td>ATG</td><td>TAA</td><td>2</td></tr><tr><td>Cytb</td><td>H</td><td>10361-11500</td><td>1140</td><td>ATG</td><td>TAA</td><td>8</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11509-11573</td><td>65</td><td></td><td></td><td>-1</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11573-11641</td><td>69</td><td></td><td></td><td>20</td></tr><tr><td>Nad4L</td><td>H</td><td>11662-11958</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11970-13325</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13325-13389</td><td>65</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13390-15111</td><td>1722</td><td>ATG</td><td>TAG</td><td>31</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia tenuicosta 15,169 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-770</td><td>incomplete</td><td>—</td><td>TAA</td><td>12</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>783-848</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>850-919</td><td>70</td><td>—</td><td>—</td><td>8</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>928-996</td><td>69</td><td>—</td><td>—</td><td>11</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1008-1074</td><td>67</td><td>—</td><td>—</td><td>15</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1090-1157</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1161-1514</td><td>354</td><td>ATG</td><td>TAA</td><td>-1</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1514-1583</td><td>70</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad2</td><td>H</td><td>1583-2637</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2638-4173</td><td>1536</td><td>ATG</td><td>TAA</td><td>25</td></tr><tr><td>Cox2</td><td>H</td><td>4199-4885</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4884-4951</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4953-5111</td><td>159</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Atp6</td><td>H</td><td>5118-5813</td><td>696</td><td>ATG</td><td>TAA</td><td>42</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5856-5922</td><td>67</td><td>—</td><td>—</td><td>3</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5926-5996</td><td>71</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5996-6060</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6061-6126</td><td>66</td><td>—</td><td>—</td><td>-3</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6124-6190</td><td>67</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6191-6257</td><td>67</td><td>—</td><td>—</td><td>12</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6270-6335</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6336-7296</td><td>961</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7297-7364</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7365-8716</td><td>1352</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8717-8785</td><td>69</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8787-8857</td><td>71</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>Nad1</td><td>H</td><td>8857-9798</td><td>942</td><td>ATG</td><td>TAA</td><td>4</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9803-9870</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9872-10372</td><td>501</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Cytb</td><td>H</td><td>10379-11518</td><td>1140</td><td>ATG</td><td>TAA</td><td>11</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11530-11593</td><td>64</td><td>—</td><td>—</td><td>-1</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11593-11662</td><td>70</td><td>—</td><td>—</td><td>24</td></tr><tr><td>Nad4L</td><td>H</td><td>11687-11983</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11995-13350</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13350-13415</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13417-15138</td><td>1722</td><td>ATG</td><td>TAG</td><td>31</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>
Tritia unifasciata 15,168 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-771</td><td>incomplete</td><td>—</td><td>TAG</td><td>12</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>784-848</td><td>65</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>851-918</td><td>68</td><td></td><td></td><td>10</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>929-997</td><td>69</td><td></td><td></td><td>9</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>1007-1074</td><td>68</td><td></td><td></td><td>17</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1092-1159</td><td>68</td><td></td><td></td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1163-1516</td><td>354</td><td>ATG</td><td>TAG</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1517-1584</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1585-2639</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2640-4175</td><td>1536</td><td>ATG</td><td>TAA</td><td>25</td></tr><tr><td>Cox2</td><td>H</td><td>4201-4887</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4886-4953</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4955-5113</td><td>159</td><td>ATG</td><td>TAA</td><td>7</td></tr><tr><td>Atp6</td><td>H</td><td>5121-5816</td><td>696</td><td>ATG</td><td>TAA</td><td>40</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5857-5923</td><td>67</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5926-5992</td><td>67</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5994-6057</td><td>64</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6058-6123</td><td>66</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6125-6186</td><td>62</td><td></td><td></td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6188-6254</td><td>67</td><td></td><td></td><td>14</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6269-6333</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6334-7292</td><td>959</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7293-7360</td><td>68</td><td></td><td></td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7361-8712</td><td>1352</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8713-8781</td><td>69</td><td></td><td></td><td>2</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8784-8852</td><td>69</td><td></td><td></td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8853-9794</td><td>942</td><td>ATG</td><td>TAA</td><td>5</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9800-9867</td><td>68</td><td></td><td></td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9869-10369</td><td>501</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Cytb</td><td>H</td><td>10376-11515</td><td>1140</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11527-11591</td><td>65</td><td></td><td></td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11592-11659</td><td>68</td><td></td><td></td><td>21</td></tr><tr><td>Nad4L</td><td>H</td><td>11681-11977</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11989-13344</td><td>1356</td><td>ATA</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13344-13408</td><td>65</td><td></td><td></td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13410-15131</td><td>1722</td><td>ATG</td><td>TAG</td><td>37</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td></td><td></td><td>—</td></tr></tbody></table>
Tritia vaucheri 15,153 bp

<table><thead><tr><th>Gene</th><th>Strand</th><th>Location</th><th>Size (bp)</th><th>Start Codon</th><th>Stop codon</th><th>Intergenic nucleotides</th></tr></thead><tbody><tr><td>Cox3</td><td>H</td><td>1-750</td><td>incomplete</td><td>—</td><td>TAA</td><td>13</td></tr><tr><td>tRNA-Lys</td><td>H</td><td>764-829</td><td>66</td><td>—</td><td>—</td><td>4</td></tr><tr><td>tRNA-Ala</td><td>H</td><td>834-901</td><td>68</td><td>—</td><td>—</td><td>8</td></tr><tr><td>tRNA-Arg</td><td>H</td><td>910-978</td><td>69</td><td>—</td><td>—</td><td>5</td></tr><tr><td>tRNA-Asn</td><td>H</td><td>984-1052</td><td>69</td><td>—</td><td>—</td><td>8</td></tr><tr><td>tRNA-Ile</td><td>H</td><td>1061-1128</td><td>68</td><td>—</td><td>—</td><td>3</td></tr><tr><td>Nad3</td><td>H</td><td>1132-1485</td><td>354</td><td>ATG</td><td>TAA</td><td>0</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>1486-1553</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad2</td><td>H</td><td>1554-2608</td><td>1055</td><td>ATG</td><td>TA</td><td>0</td></tr><tr><td>Cox1</td><td>H</td><td>2609-4144</td><td>1536</td><td>ATG</td><td>TAA</td><td>29</td></tr><tr><td>Cox2</td><td>H</td><td>4174-4860</td><td>687</td><td>ATG</td><td>TAA</td><td>-2</td></tr><tr><td>tRNA-Asp</td><td>H</td><td>4859-4926</td><td>68</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Atp8</td><td>H</td><td>4928-5086</td><td>159</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>Atp6</td><td>H</td><td>5093-5788</td><td>696</td><td>ATG</td><td>TAG</td><td>40</td></tr><tr><td>tRNA-Met</td><td>L</td><td>5829-5895</td><td>67</td><td>—</td><td>—</td><td>8</td></tr><tr><td>tRNA-Tyr</td><td>L</td><td>5904-5970</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Cys</td><td>L</td><td>5972-6035</td><td>64</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Trp</td><td>L</td><td>6036-6102</td><td>67</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gln</td><td>L</td><td>6104-6165</td><td>62</td><td>—</td><td>—</td><td>1</td></tr><tr><td>tRNA-Gly</td><td>L</td><td>6167-6233</td><td>67</td><td>—</td><td>—</td><td>13</td></tr><tr><td>tRNA-Glu</td><td>L</td><td>6247-6311</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>12s</td><td>H</td><td>6312-7264</td><td>953</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Val</td><td>H</td><td>7265-7332</td><td>68</td><td>—</td><td>—</td><td>0</td></tr><tr><td>16s</td><td>H</td><td>7333-8687</td><td>1355</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8688-8756</td><td>69</td><td>—</td><td>—</td><td>4</td></tr><tr><td>tRNA-Leu</td><td>H</td><td>8761-8829</td><td>69</td><td>—</td><td>—</td><td>0</td></tr><tr><td>Nad1</td><td>H</td><td>8830-9771</td><td>942</td><td>ATG</td><td>TAA</td><td>6</td></tr><tr><td>tRNA-Pro</td><td>H</td><td>9778-9846</td><td>69</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad6</td><td>H</td><td>9848-10348</td><td>501</td><td>ATG</td><td>TAA</td><td>12</td></tr><tr><td>Cytb</td><td>H</td><td>10361-11500</td><td>1140</td><td>ATG</td><td>TAA</td><td>13</td></tr><tr><td>tRNA-Ser</td><td>H</td><td>11514-11578</td><td>65</td><td>—</td><td>—</td><td>0</td></tr><tr><td>tRNA-Thr</td><td>L</td><td>11579-11644</td><td>66</td><td>—</td><td>—</td><td>20</td></tr><tr><td>Nad4L</td><td>H</td><td>11665-11961</td><td>297</td><td>ATG</td><td>TAG</td><td>11</td></tr><tr><td>Nad4</td><td>H</td><td>11973-13328</td><td>1356</td><td>CTT</td><td>TAG</td><td>-1</td></tr><tr><td>tRNA-His</td><td>H</td><td>13328-13393</td><td>66</td><td>—</td><td>—</td><td>1</td></tr><tr><td>Nad5</td><td>H</td><td>13395-15116</td><td>1722</td><td>ATG</td><td>TAG</td><td>37</td></tr><tr><td>tRNA-Phe</td><td>H</td><td>missing</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>