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![](https://binary.pensoft.net/fig/207136/singlefigmini/file.jpg)
Phylogenetic tree based on mitochondrial COI sequences. Numbers by branches show bootstrap values (Maximum Likelihood analysis) and posterior probabilities (Bayesian analysis). Only significant numbers (>80% and >0.9) are indicated. Onchidella, Peronia, Peronina, Platevindex, Onchidium, Onchidina, and Melayonchis sequences serve as outgroups. Numbers for each individual correspond to unique identifiers used for DNA extraction. All sequences of Wallaconchis specimens are new, except two sequences obtained from GenBank (from specimens from China). Information on individually-identified specimens can be found in the lists of material examined and in Table
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![](https://binary.pensoft.net/fig/207138/singlefigmini/file.jpg)
Phylogenetic tree based on concatenated mitochondrial COI, 16S, and 12S sequences. Numbers by branches show bootstrap values (Maximum Likelihood analysis) and posterior probabilities (Bayesian analysis). Only significant numbers (>80% and >0.9) are indicated. All sequences of Wallaconchis specimens are new. Information on individually-identified specimens can be found in the lists of material examined and in Table
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![](https://binary.pensoft.net/fig/207139/singlefigmini/file.jpg)
Maximum parsimony bootstrap consensus tree based on concatenated nuclear ITS1, ITS2, and 28S sequences. Only bootstrap values >70% are indicated. All sequences of Wallaconchis specimens are new. Information on individually-identified specimens can be found in the lists of material examined and in Table
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![](https://binary.pensoft.net/fig/207140/singlefigmini/file.jpg)
Maximum Likelihood phylogenetic tree based on concatenated nuclear ITS1, ITS2, and 28S sequences. Numbers by branches show bootstrap values (only numbers >80% are indicated). All sequences of Wallaconchis specimens are new. Information on individually-identified specimens can be found in the lists of material examined and in Table
![Download image](https://arpha.pensoft.net//i/download-icon-30.png)
![](https://binary.pensoft.net/fig/207141/singlefigmini/file.jpg)
Maximum parsimony bootstrap consensus tree based on concatenated nuclear ITS1 and ITS2 sequences. Only bootstrap values >70% are indicated. All sequences of Wallaconchis specimens are new. Information on individually-identified specimens can be found in the lists of material examined and in Table
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![](https://binary.pensoft.net/fig/207142/singlefigmini/file.jpg)
Geographic distribution of Wallaconchis species. The colors used for each species are the same as those used in phylogenetic trees (Figs
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![](https://binary.pensoft.net/fig/207143/singlefigmini/file.jpg)
Habitats and live specimens, Wallaconchis sinanui, Indonesia, Ambon, Lateri. A Type locality, mudflat adjacent to a shallow river through a mangrove (st 128) B Close up of mud with a few individuals of W. sinanui (same locality as A) C Wider view of the mud with many individuals of W. sinanui (same locality as A) D Dorsal view, 9 mm long [2746] (UMIZ 00059) E Holotype, dorsal view, 9 mm long [2737] (UMIZ 00058) F Dorsal view, 9 mm long [2740] (UMIZ 00059).
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![](https://binary.pensoft.net/fig/207144/singlefigmini/file.jpg)
External morphology and digestive system, Wallaconchis sinanui, Indonesia, Ambon (A–C) holotype [2737] (UMIZ 00058) (D–F) [2738] (UMIZ 00059). A Dorsal notum B Anterior region, ventral view C Posterior region, ventral view D Digestive system, dorsal view E Digestive system, dorsal view (digestive gland removed) F Digestive system, ventral view (digestive gland removed). Abbreviations: a anus ddg dorsal digestive gland f foot fo female opening hn hyponotum i intestine mo male opening oddg opening of the dorsal lobe of the digestive gland oldg opening of the lateral lobe of the digestive gland opdg opening of the posterior lobe of the digestive gland ol oral lobe ot ocular tentacle pn pneumostome pdg posterior lobe of the digestive gland ppg peripodial groove st1 first stomach chamber st2 second stomach chamber st3 third stomach chamber st4 fourth stomach chamber. Scale bars: 2 mm (A, B, C); 1 mm (D, E, F).
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![](https://binary.pensoft.net/fig/207145/singlefigmini/file.jpg)
Radula, Wallaconchis sinanui, Indonesia, Ambon [5845] (UMIZ 00059). A Rachidian and innermost lateral teeth B Rachidian and innermost lateral teethC Transition between inner lateral teeth and outer lateral teeth D Outermost lateral teeth E Right lateral teeth with basal lateral spine F Outermost lateral teeth. Abbreviations: rt rachidian tooth mc median cusp lc lateral cusp 1llt first left lateral tooth 1rlt first right lateral tooth 2llt second left lateral tooth 2rlt second right lateral tooth bls basal lateral spine (of lateral tooth) nls no lateral spine hlt hook of lateral tooth ollt outermost left lateral tooth. Scale bars: 10 μm (A); 20 μm (B, D, E); 40 μm (C), 10 μm (F).
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![](https://binary.pensoft.net/fig/207146/singlefigmini/file.jpg)
Nervous system, Wallaconchis sinanui, holotype, Indonesia, Ambon [2737] (UMIZ 00058) scale bar 0.5 mm. Abbreviations: lcg left cerebral ganglion lpg left pedal ganglion lplg left pleural ganglion rcg right cerebral ganglion rpg right pedal ganglion rplg right pleural ganglion vg visceral ganglion.
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Reproductive system, Wallaconchis sinanui, holotype, Indonesia, Ambon [2737] (UMIZ 00058). A Hermaphroditic (female), posterior parts, scale bar 0.8 mm B Anterior, male copulatory parts, scale bar 2 mm. Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca v vestibule.
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![](https://binary.pensoft.net/fig/207148/singlefigmini/file.jpg)
Penis, Wallaconchis sinanui, Indonesia, Ambon. A Penis evaginated outside the male opening, scale bar 200 μm [5846] (UMIZ 00059) B Penis removed from vestibule, scale bar 20 μm [5844] (UMIZ 00059).
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![](https://binary.pensoft.net/fig/207149/singlefigmini/file.jpg)
Habitats, Wallaconchis uncinus, Indonesia. A, B Sulawesi, Wori, tall mangrove forest of Sonneratia and some Avicennia with a rocky area and dead logs (st 90) C Ambon, Lateri, muddy mangrove with Rhizophora and Avicennia (st 131) D Type locality, Ambon, Lateri, mudflat adjacent to a shallow river through a mangrove (st 128) E Halmahera, Sofifi, mangrove with Sonneratia trees, dense roots and hard mud (st 205) F Ternate, Bastiong, muddy rocks nearby a mangrove patch (st 203).
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![](https://binary.pensoft.net/fig/207150/singlefigmini/file.jpg)
Live specimens, Wallaconchis uncinus, Indonesia. A Dorsal view, 25 mm long [5070], Halmahera (UMIZ 00010) B Holotype, dorsal view, 22 mm long [2751], Ambon (UMIZ 00004) C Dorsal view, 30 mm long [2752], Ambon (UMIZ 00006) D Dorsal view, 28 mm long [2843], Ambon (UMIZ 00008) E Dorsal view, 15 mm long [3138], Bali (UMIZ 00005) F Dorsal view, 25 mm long [5080], Halmahera (UMIZ 00011) G Ventral view, same as F H Ventral view, same as B.
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![](https://binary.pensoft.net/fig/207151/singlefigmini/file.jpg)
External morphology and digestive system, Wallaconchis uncinus, Indonesia. A Anterior region, ventral view, Sulawesi, scale bar 2 mm [2268] (UMIZ 00005) B Posterior region, ventral view, Sulawesi, scale bar 2 mm [2268] (UMIZ 00005) C Dorsal notum (papillae with dorsal eyes), Halmahera, scale bar 2.6 mm [5056] (UMIZ 00009) D Digestive system, dorsal view, Sulawesi, scale bar 1 mm [2268] (UMIZ 00005). Abbreviations: a anus ddg dorsal lobe of digestive gland f foot fo female opening hn hyponotum i intestine mo male opening ol oral lobe ot ocular tentacle pdg posterior lobe of the digestive gland pn pneumostome ppg peripodial groove st2 stomach chamber 2.
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![](https://binary.pensoft.net/fig/207152/singlefigmini/file.jpg)
Radula, Wallaconchis uncinus, Indonesia. A Rachidian and innermost lateral teeth, Bali, scale bar 10 μm [3138] (UMIZ 00008) B Rachidian and innermost lateral teeth, Halmahera, scale bar 20 μm [5079] (UMIZ 00011) C Transition from inner to outer lateral teeth, scale bar 20 μm, same as B D Outer lateral teeth, scale bar 40 μm, same as B E Outermost lateral teeth, scale bar 20 μm, same as B F Outermost lateral teeth, Ambon, scale bar 20 μm [2843] (UMIZ 00007). Abbreviations: bls basal lateral spine (of lateral tooth) nls no lateral spine.
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![](https://binary.pensoft.net/fig/207153/singlefigmini/file.jpg)
Reproductive system, Wallaconchis uncinus, Indonesia. A Hermaphroditic (female), posterior parts, Sulawesi, scale bar 2 mm [2268] (UMIZ 00005) B Anterior, male copulatory parts, Halmahera, scale bar 3 mm [5079] (UMIZ 00011) C Anterior, male copulatory parts, Bali, scale bar 1 mm [3138] (UMIZ 00008) D Penis (fully evaginated), Halmahera, scale bar 0.5 mm [5079] (UMIZ 00011). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca v vestibule.
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![](https://binary.pensoft.net/fig/207154/singlefigmini/file.jpg)
Penis, Wallaconchis uncinus, Indonesia. A Two most distal loops of fully evaginated penis (the loop on the right is proximal to the loop on the left), Halmahera, scale bar 0.5 mm [5079] (UMIZ 00011) B Two most distal loops of (nearly fully evaginated) penis (the loop on the right is proximal to the loop on the left), Ambon, scale bar 300 μm [2843] (UMIZ 00007) C Proximal region of penis, detail view of curved hooks, Sulawesi, scale bar 100 μm [2268] (UMIZ 00005) D Distal region of penis which is not fully evaginated, scale bar 300 μm, same as C. Abbreviations: ch curved hooks ih immature hooks rh rounded hooks.
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![](https://binary.pensoft.net/fig/207155/singlefigmini/file.jpg)
Habitats, Wallaconchis buetschlii. A Indonesia, Sulawesi, Wori, tall mangrove forest of Sonneratia and some Avicennia with a rocky area and dead logs (st 84) B Australia, Queensland, Yule Point, sandy area with a few Rhizophora and Avicennia trees (st 100) C Indonesia, Ambon, Wai, narrow band of old Avicennia trees on sandy mud, with old logs on ground (st 132) D Indonesia, Lombok, Don Don, old Avicennia forest, with coral rubble (st 149) E Indonesia, Sulawesi, Mantehage Island, rocks behind a mangrove of Sonneratia and Rhizophora (st 91) F Indonesia, Bali, Gilimanuk, sandy mudflat outside Rhizophora and Avicennia mangrove (st 156).
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![](https://binary.pensoft.net/fig/207156/singlefigmini/file.jpg)
Live specimens, Wallaconchis buetschlii. A Dorsal view, 46 mm long [2722], Indonesia, Ambon (UMIZ 00026) B Dorsal view, 31 mm long [2853], Indonesia, Ambon (UMIZ 00027) C Dorsal view, 23 mm long [2249], Indonesia, Sulawesi (UMIZ 00024) D Dorsal view, 30 mm long [2928], Indonesia, Kei Islands (UMIZ 00029) E Dorsal view, 52 mm long [3159], Philippines, Luzon (PNM 041209) F Dorsal view, 36 mm long, [3409], Philippines, Bohol (PNM 041212) G Ventral view, 44 mm long [5898], Indonesia, Timor (UMIZ 00072) H Ventral view, same as F I Ventral view, 43 mm long [2982], Indonesia, Lombok (UMIZ 00030).
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![](https://binary.pensoft.net/fig/207157/singlefigmini/file.jpg)
Radula, Wallaconchis buetschlii (A–D) Indonesia, Sulawesi [2120] (UMIZ 00021) (E) Australia, Queensland [2554] (MTQ st100) (F) Australia, Queensland [2555] (MTQ st100). A Rachidian and innermost lateral teeth, scale bar 10 μm B Rachidian and innermost lateral teeth, scale bar 40 μm C Inner lateral teeth, scale bar 30 μm D Outermost lateral teeth, scale bar 20 μm E Outermost lateral teeth, scale bar 10 μm F Transition between inner lateral teeth and outer lateral teeth, with the basal lateral spine revealed on the underside of the inner lateral teeth at the bottom, scale bar 30 μm.
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![](https://binary.pensoft.net/fig/207158/singlefigmini/file.jpg)
Reproductive system, Wallaconchis buetschlii. A Hermaphroditic (female), posterior parts, Indonesia, Sulawesi, scale bar 2 mm [2120] (UMIZ 00021) B Anterior, male copulatory parts, Australia, Queensland, scale bar 1 mm [2555] (MTQ st100) C Anterior, male copulatory parts, Indonesia, Kei, scale bar 1 mm [2933] (UMIZ 00029). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland of oviduct fibers attaching to the inner wall of the visceral cavity ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca.
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![](https://binary.pensoft.net/fig/207159/singlefigmini/file.jpg)
Penis, Wallaconchis buetschlii, Indonesia. A Longitudinal ridge on the internal surface of the penis, Kei Islands, scale bar 60 μm [2933] (UMIZ 00029) B Close-up of a groove between two longitudinal ridges showing smooth internal surface (no hooks), Sulawesi, scale bar 20 μm [2120] (UMIZ 00021).
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![](https://binary.pensoft.net/fig/207160/singlefigmini/file.jpg)
Habitats, Wallaconchis gracile, Indonesia. A–B Bali, Gilimanuk, muddy-sandy beach near a mangrove (st 155) C–D Ternate, sandy mudflat adjacent to a rocky beach (st 220) E–F Timor, sandy area of mangrove with ripples in the sand and near crab pellets (st254).
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![](https://binary.pensoft.net/fig/207161/singlefigmini/file.jpg)
Live specimens, Wallaconchis gracile. A Dorsal view, 19 mm long [3107], Indonesia, Bali (UMIZ 00056) B Dorsal view, 15 mm long [5158], Indonesia, Halmahera (UMIZ 00057) C Dorsal view, 21 mm long [3648], Philippines, Bohol (PNM 041231) D Dorsal view, 19 mm long [2252], Indonesia, Sulawesi (UMIZ 00055) E Dorsal view, 15 mm long [5159], Indonesia, Halmahera (UMIZ 00057) F Dorsal view, 27 mm long [5932], Indonesia, Timor (UMIZ 00074) G Dorsal view, 25 mm long [3652], Philippines, Bohol (PNM 041231) H Frontal view, same as F I Ventral view, 25 mm long [3653], Philippines, Bohol (PNM 041231) J Ventral view, same as F.
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![](https://binary.pensoft.net/fig/207162/singlefigmini/file.jpg)
Radula, Wallaconchis gracile, Indonesia, Bali (A–C) [3106] (UMIZ 00056) (D) [spm #1] (UMIZ 00056). A Rachidian and innermost lateral teeth, scale bar 6 μm B Rachidian and innermost lateral teeth, scale bar 20 μm C Transition between inner lateral teeth and outer lateral teeth, scale bar 30 μm D Left lateral teeth, scale bar 30 μm E Left lateral teeth, gradually increasing in length towards the left, scale bar 40 μm F Outermost lateral teeth, scale bar 20 μm.
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![](https://binary.pensoft.net/fig/207163/singlefigmini/file.jpg)
Reproductive system, Wallaconchis gracile, Indonesia. A Hermaphroditic (female), posterior parts, Sulawesi, scale bar 2 mm [2280] (UMIZ 00055) B Anterior, male copulatory parts, Bali, scale bar 2 mm [3106] (UMIZ 00056). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland of oviduct fibers attaching to the inner wall of the visceral cavity ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca.
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![](https://binary.pensoft.net/fig/207164/singlefigmini/file.jpg)
Penis, distal part free in the vestibule, Wallaconchis gracile, Indonesia, Bali, scale bar 40 μm [3106] (UMIZ 00056).
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![](https://binary.pensoft.net/fig/207165/singlefigmini/file.jpg)
Habitats, Wallaconchis nangkauriense. A Indonesia, Sulawesi, sand, rocks, pieces of wood behind narrow coastal mangrove (st 85) B India, Andaman Islands, Burman Nala, slugs were on the stilt roots (st 53) C Indonesia, Ambon, Haruku Island, narrow mangrove of Sonneratia trees on rocky beach (st 127) D Close up (same locality as C) E Indonesia, Bali, Pemuteran, Labuhan Lalang Harbor, coral rubble, rocks and a few Avicennia (st 157) F Philippines, Bohol, Maribojoc, coral rubble of uplifted coral reef (st 191).
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![](https://binary.pensoft.net/fig/207166/singlefigmini/file.jpg)
Live specimens, Wallaconchis nangkauriense. A Dorsal view, 28 mm long [3162], Philippines, Luzon (PNM 041204) B Dorsal view, 23 mm long [1786], Indonesia, Sumatra (UMIZ 00012) C Dorsal view, 18 mm long [2156], Indonesia, Sulawesi (UMIZ 00013) D Dorsal view, 23 mm long [1788], Indonesia, Sumatra (UMIZ 00012) E Dorsal view, 34 mm long [3269], Philippines, Bohol (PNM 041206) F Dorsal view, 24 mm long [2161], Indonesia, Sulawesi (UMIZ 00013) G Dorsal view, 32 mm long [3268], Philippines, Bohol (PNM 041206) H Dorsal view, 28 mm long [1787], Indonesia, Sumatra (UMIZ 00012) I Dorsal view, 24 mm long [3217], Philippines, Luzon (PNM 041205) J Ventral view, same as G K Ventral view, 28 mm long [3427], Philippines, Bohol (PNM 041208) L Ventral view, 22 mm long [2731], Indonesia, Ambon (UMIZ 00017).
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![](https://binary.pensoft.net/fig/207167/singlefigmini/file.jpg)
Radula, Wallaconchis nangkauriense, Indonesia, Bali [3129] (UMIZ 00020). A Rachidian and innermost lateral teeth, scale bar 20 μm B Innermost lateral teeth, scale bar 20 μm C Outer lateral teeth, scale bar 20 μm D Outermost lateral teeth, scale bar 20 μm.
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![](https://binary.pensoft.net/fig/207168/singlefigmini/file.jpg)
Reproductive system, Wallaconchis nangkauriense. A Hermaphroditic (female), posterior parts, Philippines, Bohol, scale bar 4 mm [3276] (PNM 041206) B Anterior, male copulatory parts, Indonesia, Sulawesi, scale bar 5 mm [2192] (UMIZ 00014) C Anterior, male copulatory parts (vestibule removed around the penis), Indonesia, Sumatra, scale bar 2 mm [1788] (UMIZ 00012). Abbreviations: bc body cavity bw lateral left body wall dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct p penis ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca.
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![](https://binary.pensoft.net/fig/207169/singlefigmini/file.jpg)
Penis, Wallaconchis nangkauriense, Philippines. A Luzon, scale bar 200 μm [3208] (PNM 041205) B Bohol, scale bar 200 μm [3268] (PNM 041206).
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![](https://binary.pensoft.net/fig/207170/singlefigmini/file.jpg)
Mangrove habitats, Wallaconchis ater. A–B Indonesia, Bali, Pemuteran, Labuhan Lalang Harbor, coral rubble, rocks and a few Avicennia (st 157) C–D Indonesia, Lombok, Don Don, old Avicennia forest, with coral rubble (st 149) E–F Philippines, Bohol, Mabini, open Avicennia and Sonneratia mangrove with sand and coral rubble (st 196).
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![](https://binary.pensoft.net/fig/207171/singlefigmini/file.jpg)
Coral rubble habitats, Wallaconchis ater. A Indonesia, Sulawesi, Bahoi, sand and small rocks outside a mangrove (st 88) B Philippines, Bohol, Maribojoc, uplifted, dead, coral flat covered with sand and algae adjacent to a few mangrove trees (st 202) C Philippines, Bohol, Maribojoc, coral rubble of uplifted coral reef (st 191) D Indonesia, Halmahera, Foli, rocky shore by beach (st 217).
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![](https://binary.pensoft.net/fig/207172/singlefigmini/file.jpg)
Live specimens, dorsal view, Wallaconchis ater. A 33 mm long [3265], Philippines, Bohol (PNM 041214) B Dorsal view, 34 mm long [3271], Philippines, Bohol (PNM 041222) C 26 mm long [3210], Philippines, Luzon (PNM 041213) D 26 mm long [3405], Philippines, Bohol (PNM 041219) E 26 mm long [2164], Indonesia, Sulawesi (UMIZ 00036) F 38 mm long [5057], Indonesia, Halmahera (UMIZ 00046) G Dorsal view, 22 mm long [3137], Indonesia, Bali (UMIZ 00053) H 25 mm long [2157], Indonesia, Sulawesi (UMIZ 00036).
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![](https://binary.pensoft.net/fig/207173/singlefigmini/file.jpg)
Live specimens, Wallaconchis ater. A Dorsal view, 43 mm long [2970], Indonesia, Lombok (UMIZ 00043) B Dorsal view, 37 mm long [2170], Indonesia, Sulawesi (UMIZ 00036) C Ventral view, 35 mm long [2974], Indonesia, Lombok (UMIZ 00043) D Ventral view, 36 mm long [3266], Philippines, Bohol (PNM 041214) E Ventral view, 26 mm long [3405], Philippines, Bohol (PNM 041219) F Ventral view, 33 mm long [3272], Philippines, Bohol (PNM 041214) G Ventral view, 32 mm long [3277], Philippines, Bohol (PNM 041222) H Ventral view, 37 mm long [3420], Philippines, Bohol (PNM 0412224).
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![](https://binary.pensoft.net/fig/207174/singlefigmini/file.jpg)
Radula, Wallaconchis ater, Indonesia (A–B) Bali [3137] (UMIZ 00053) (D) Sulawesi [2330] (UMIZ 00039) (C, E–F) Sulawesi [2220] (UMIZ 00050). A Rachidian and innermost lateral teeth, scale bar 20 μm B Rachidian and left innermost lateral teeth, scale bar 30 μm C Rachidian and innermost lateral teeth, scale bar 50 μm D Rachidian and right innermost lateral teeth, scale bar 50 μm E Outermost lateral teeth, scale bar 10 μm F Transition between inner lateral teeth and outer lateral teeth, scale bar 40 μm.
![Download image](https://arpha.pensoft.net//i/download-icon-30.png)
![](https://binary.pensoft.net/fig/207175/singlefigmini/file.jpg)
Reproductive system, hermaphroditic (female), posterior parts, Wallaconchis ater, Indonesia A Lombok, scale bar 2 mm [2970] (UMIZ 00043) B Sulawesi, scale bar 3 mm [2220] (UMIZ 00050). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct rs receptaculum seminis sp spermatheca.
![Download image](https://arpha.pensoft.net//i/download-icon-30.png)
![](https://binary.pensoft.net/fig/207176/singlefigmini/file.jpg)
Reproductive system, Wallaconchis ater, Indonesia. A Anterior, male copulatory parts (vestibule removed, showing the coiled and U-shaped penis), Halmahera, scale bar 2 mm [5078] (UMIZ 00047) B Penis (coil of loops removed from vestibule), Sulawesi, scale bar 2 mm [2283] (UMIZ 00038) C Anterior, male copulatory parts, scale bar 5 mm, same as B D Anterior, male copulatory parts, Kei Islands, scale bar 1 mm [2939] (UMIZ 00041). Abbreviations: dd deferent duct dpp distal penial projections p penis ps penial sheath rm retractor muscle v vestibule.
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![](https://binary.pensoft.net/fig/207177/singlefigmini/file.jpg)
Reproductive system, anterior, male copulatory parts, Wallaconchis ater. A Vestibule removed, showing the coiled and U-shaped penis, Indonesia, Sulawesi, scale bar 3 mm [2220] (UMIZ 00050) B Vestibule with coiled penis inside, Philippines, Bohol, scale bar 3 mm [3270] (PNM 041220). Abbreviations: dd deferent duct dpp distal penial projections p penis ps penial sheath rm retractor muscle v vestibule.
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![](https://binary.pensoft.net/fig/207178/singlefigmini/file.jpg)
Penis, Wallaconchis ater, Indonesia. A Detail of distal portion when fully relaxed (with distal penial projections) and partial view of the coil, Indonesia, Halmahera, scale bar 500 μm [3393] (PNM 041218) B Detail of distal portion when fully relaxed (with distal penial projections) and partial view of the coil, Halmahera, scale bar 500 μm [5125] (UMIZ 00054) C Partial view of the coil (with six loops), Philippines, Bohol, scale bar 500 μm [3266] (PNM 041214) D Cross-section showing the longitudinal fold, Bali, scale bar 300 μm [3137] (UMIZ 00053).
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Habitats, Wallaconchis graniferum. A Philippines, Bohol, Loay, mostly sand, very few rocks, and a few small Avicennia trees (st 198) B Close up of sand with shells and Avicennia roots (same locality as A) C Indonesia, Timor, Oesapa, sandy area of mangrove with Avicennia and Sonneratia trees spread out D Close up of sediment and onchidiid tracks (same locality as C).
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Live specimens, Wallaconchis graniferum, Philippines. A Dorsal view, 46 mm long [3163], Luzon (PNM 041227) B Dorsal view, 30 mm long [5902], Indonesia, Timor (UMIZ 00073) C Dorsal view, 28 mm long [3636], Bohol (PNM 041228) D Dorsal view, 26 mm long [3635], Bohol (PNM 041228) E Ventral view, same as A, F Ventral view, same as B.
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Radula, Wallaconchis graniferum, Philippines. A Rachidian and innermost lateral teeth, Bohol, scale bar 20 μm [3636] (PNM 041228) B Rachidian and innermost lateral teeth, Luzon, scale bar 30 μm [3163] (PNM 041227) C Transition between inner lateral teeth and outer lateral teeth, Bohol, scale bar 30 μm [3635] (PNM 041228) D Outermost lateral teeth, Bohol, scale bar 30 μm [3636] (PNM 041228).
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Reproductive system, Hermaphroditic (female), posterior parts, Wallaconchis graniferum, Philippines. A Luzon, scale bar 5 mm [3163] (PNM 041227) B Bohol, scale bar 2 mm [3638] (PNM 041225). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct rs receptaculum seminis sp spermatheca.
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Reproductive system, anterior, male copulatory parts, Wallaconchis graniferum, Philippines. A Penis with two proximal regions of hooks separated from a third distal region of hooks by a gap with no hooks (the hooks of the first, most proximal region are on the other side of the penis and cannot be seen here), Luzon, scale bar 0.7 mm [3163] (PNM 041227) B Anterior male copulatory parts scale bar 3.6 mm same as A. C Penis partially evaginated inside the vestibule (removed) Bohol scale bar 1 mm [3638] (PNM 041225). Abbreviations: dd deferent duct pr1 first most proximal penial region (with hooks on the other side) pr2 second proximal penial region (with hooks) pr3 third distal penial region (with hooks) ps penial sheath rm retractor muscle sr smooth region of the penis (with no hooks) v vestibule.
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Evaginated penis, Wallaconchis graniferum, Philippines (A–C) Luzon [3163] (PNM 041227) (D) Bohol [3636] (PNM 041228) (E) Bohol [3638] (PNM 041225). A Proximal region of evaginated penis with hooks, scale bar 400 μm B Middle region of evaginated penis with long slender hooks on one side, and shorter hooks on the other side, scale bar 400 μm C Distal region of evaginated penis with distal hooks, scale bar 300 μm D Partially evaginated penis with proximal hooks (more distal hooks are inside the rest of the invaginated penis), scale bar 200 μm E Partially evaginated penis with most proximal hooks (more distal hooks are invaginated inside), scale bar 200 μm. Abbreviations (from proximal to distal): ph proximal hooks (first penial region) lh long slender hooks (second penial region) sh short hooks (second penial region) nh no hooks (gap) dh distal hooks (third penial region).
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Penis, Wallaconchis graniferum, not evaginated and opened, Philippines, Bohol (A–B) [5762] (PNM 041226) (C) [5761] (PNM 041228). A Penis invaginated inside the penial sheath opened to reveal regions of penial hooks, scale bar 200 μm B Most proximal region of penis invaginated inside the penial sheath opened to reveal penial hooks, scale bar 200 μm C Penis invaginated inside penial sheath (both opened to reveal the penial hooks), scale bar 0.5 mm. Abbreviations (from proximal to distal): ph proximal hooks (first penial region) lh long slender hooks (second penial region) sh short hooks (second penial region) nh no hooks (gap) dh distal hooks (third penial region).
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Habitats and live specimens, Wallaconchis achleitneri, Australia, Queensland, Bowen, Doughty Creek. A–B Type locality, sandy area with Avicennia and Rhizophora trees in narrow band by the creek (st 117) C–D Dorsal view, approximately 14 mm long (MTQ st117) E Ventral view, approximately 14 mm long (MTQ st 117).
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Radula, Wallaconchis achleitneri, Australia, Queensland (MTQ st117). A Rachidian and innermost lateral teeth, scale bar 6 μm [#1] B Rachidian and innermost lateral teeth, holotype, scale bar 20 μm [3534] C Transition between inner lateral teeth and outer lateral teeth, holotype, scale bar 30 μm [3534] D Outermost lateral teeth, holotype, scale bar 10 μm [3534] E Lateral teeth with basal lateral spine decreasing in size from right to left, scale bar 10 μm [#1] F Left lateral teeth, scale bar 100 μm [#2].
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Reproductive system, Wallaconchis achleitneri, Australia, Queensland (MTQ st117). A Hermaphroditic (female), posterior parts, scale bar 1 mm [#4] B Anterior, male copulatory parts, scale bar 1 mm [#1]. Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca v vestibule.
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Penis, Wallaconchis achleitneri, Australia, Queensland (MTQ st117). A Scale bar 100 μm [2645] B Holotype, scale bar 100 μm [3534].
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Habitats, Wallaconchis comendadori. A Type locality, Philippines, Bohol, Guindulman, rocks and coral rubble near a few Avicennia trees (st 197) B View of the shore (same locality as A) C Indonesia, Bali, Pemuteran, Labuhan Lalang Harbor, coral rubble, rocks and a few Avicennia trees (st 157) D Indonesia, Lombok, Don Don, old Avicennia forest, with coral rubble (st 149).
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Live specimens, Wallaconchis comendadori. A Dorsal view, 16 mm long [3133], Indonesia, Bali (UMIZ 00064) B Dorsal view, 18 mm long [2985], Indonesia, Lombok (UMIZ 00063) C Dorsal view, 13 mm long [3131], Indonesia, Bali (UMIZ 00064) D Dorsal view, 12 mm long [2725], Indonesia, Ambon (UMIZ 00061) E Dorsal view, 7 mm long [2983], Indonesia, Lombok (UMIZ 00063) F Dorsal view, holotype, 12 mm long [3626], Philippines, Bohol (PNM 041232) G Ventral view, same as B H Ventral view, 15 mm long [2315], Indonesia, Sulawesi (UMIZ 00060) I Ventral view, 22 mm long [3385], Philippines, Bohol (PNM 041234).
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Radula, Wallaconchis comendadori. A Rachidian and innermost lateral teeth, Philippines, Bohol, scale bar 20 μm [3385] (PNM 041234) B Rachidian and innermost lateral teeth, Indonesia, Lombok, scale bar 20 μm [2985] (UMIZ 00063) C Rachidian and inner lateral teeth, scale bar 30 μm, same as B D Transition between inner lateral teeth and outer lateral teeth, arrow shows basal lateral spine, scale bar 30 μm, same as B E Transition between inner lateral teeth and outer lateral teeth, scale bar 30 μm, same as B F Outermost lateral teeth, Indonesia, Ambon, scale bar 20 μm [3539] (UMIZ 00061).
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Reproductive system, Wallaconchis comendadori. A Hermaphroditic (female), posterior parts, Indonesia, Lombok, scale bar 3 mm [2985] (UMIZ 00063) B Anterior, copulatory parts, Philippines, Bohol, scale bar 2 mm [3385] (PNM 041234). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct ps penial sheath rm retractor muscle rs receptaculum seminis sp spermatheca.
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Penis, Wallaconchis comendadori, Philippines, Bohol [3385] (PNM 041234). A View of entire penis, scale bar 0.5 mm B Distal tip of penis, scale bar 50 μm.
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Habitats, Wallaconchis melanesiensis, Indonesia. A Halmahera, Foli, large rocks with algae high in intertidal (st 217) B Close-up of boulder showing the algae where the onchidiid was found (same locality as A) C Lombok, Seriwe Bay, narrow rocky beach between a cliff and the sea water D Halmahera, Sofifi, muddy, rocky intertidal (st 204).
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Live specimens, Wallaconchis melanesiensis, Indonesia. A Dorsal view, 25 mm long [5131], Halmahera (UMIZ 00069) B Dorsal view, 27 mm long [5133], Halmahera (UMIZ 00069) C Holotype, dorsal view, 15 mm long [5065], Halmahera (UMIZ 00065) D Dorsal view, 23 mm long [5132], Halmahera (UMIZ 00069) E Dorsal view, 27 mm long [2963], Lombok (UMIZ 00070) F Dorsal view, 43 mm long [2735], Ambon (UMIZ 00068) G Ventral view, same as B H Ventral view, same as C.
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Radula, Wallaconchis melanesiensis, Indonesia (A–C) Halmahera [5132] (UMIZ 00069) (D) Ambon [2733] (UMIZ 00068). A Rachidian and innermost lateral teeth, scale bar 20 μm B Rachidian and inner left lateral teeth, scale bar 20 μm C Transition between inner lateral teeth and outer lateral teeth, scale bar 30 μm D Outermost lateral teeth, scale bar 20 μm.
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Radula, Wallaconchis melanesiensis, Vanuatu [5484] (IM-2013-62406). A Rachidian and innermost lateral teeth, scale bar 20 μm B Rachidian and inner left lateral teeth, scale bar 50 μm C Transition between inner lateral teeth and outer lateral teeth, scale bar 100 μm D Outermost lateral teeth, scale bar 20 μm E Hook of left lateral teeth from underneath, without basal lateral spine, scale bar 30 μm F Hook of left lateral teeth from underneath, without basal lateral spine, scale bar 20 μm.
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Reproductive system, hermaphroditic (female), posterior parts, Wallaconchis melanesiensis. A Indonesia, Halmahera, scale bar 2 mm [5131] (UMIZ 00069) B Vanuatu, scale bar 1 mm [5484] (IM-2013-62406) C Indonesia, Lombok, scale bar 2.4 mm [2963] (UMIZ 00070). Abbreviations: dd deferent duct fgm female gland mass hd hermaphroditic duct hg hermaphroditic gland ov oviduct rs receptaculum seminis sp spermatheca.
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Reproductive system, anterior, male copulatory parts, Wallaconchis melanesiensis. A Vanuatu, scale bar 1 mm [5484] (IM-2013-62406) B Indonesia, Halmahera, scale bar 2 mm [5132] (UMIZ 00069) C Indonesia, Lombok, scale bar 2 mm [2963] (UMIZ 00070). Abbreviations: dd deferent duct p penis ps penial sheath rm retractor muscle v vestibule.
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Penis, Wallaconchis melanesiensis, Papua New Guinea, Madang, scale bar 10 μm [5421] (IM-2013-14039).
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Phylogenetic relationships of Wallaconchis species, with shifts in habitat and character state mapped (for the details on node support and numbers of specimens included in each species, see Figs