Research Article |
Corresponding author: Suchana Chavanich ( suchana.c@chula.ac.th ) Academic editor: Nathalie Yonow
© 2021 Rahul Mehrotra, Manuel A. Caballer Gutiérrez, Chad M. Scott, Spencer Arnold, Coline Monchanin, Voranop Viyakarn, Suchana Chavanich.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Mehrotra R, Caballer Gutiérrez MA, Scott CM, Arnold S, Monchanin C, Viyakarn V, Chavanich S (2021) An updated inventory of sea slugs from Koh Tao, Thailand, with notes on their ecology and a dramatic biodiversity increase for Thai waters. ZooKeys 1042: 73-188. https://doi.org/10.3897/zookeys.1042.64474
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Improved access to field survey infrastructure throughout South-East Asia has allowed for a greater intensity of biodiversity surveys than ever before. The rocky bottoms and coral reef habitats across the region have been shown to support some of the highest sea slug biodiversity on the planet, with ever increasing records. During the past ten years, intensive SCUBA surveys have been carried out at Koh Tao, in the Gulf of Thailand, which have yielded remarkable findings in sea slug biology and ecology. In this work a brief history of sea slug biodiversity research from Thailand is covered and a complete inventory of sea slugs from Koh Tao, Gulf of Thailand is provided. This inventory is based on surveys from 2012 to 2020, with previously unreported findings since 2016. Habitat specificity and species-specific ecology are reported where available with a focused comparison of coral reef habitats and deeper soft-sediment habitats. The findings contribute 90 new species records for Thai waters (92 for the Gulf of Thailand) and report a remarkable consistency in the proportional diversity found to be exclusive to one habitat type or another. Additionally, taxonomic remarks are provided for species documented from Koh Tao that have not been discussed in past literature from Thailand, and a summary of previous records in the Indo-West Pacific is given.
Biodiversity exploration, coral reefs, Gulf of Thailand, Heterobranchia, soft sediment habitats
Contemporary sea slug research is largely dominated by investigations into biochemistry, taxonomy, and systematics of the vast diversity of species currently known. Broader aspects remain largely understudied such as development, trophic ecology, and biogeography. Recent years have seen a dramatic increase in the abundance of biodiversity inventories, particularly from regions where much of this work had been sparse before. The importance of documenting local and regional species ranges and diversity is often overlooked despite such studies contributing to our understanding of large-scale environmental issues such as increasing ocean temperatures (
Among the earliest records of sea slugs from Thailand were provided by
A dedicated survey effort was carried at the island of Koh Tao in the Gulf of Thailand which combined citizen science efforts with in-situ survey techniques and resulted in a dramatic increase in the documented diversity for the Gulf (
Benthic surveys were carried out using SCUBA at Koh Tao, Thailand, between January 2016 and February 2020. Roving-diver surveys were performed over both coral reef and soft sediment habitats. Belt transect surveys were also executed across both habitats following the Ecological Monitoring Protocol according to
Surveyed sites with location codes given in each species after specimen size.
Location | Code | Coordinates |
---|---|---|
Leuk Bay | LB | 10°4'11.65"N, 99°50'34.42"E |
Suan Olan Artificial Reef | SO | 10°4'6.70"N, 99°50'26.29"E |
Coral-Aid Artificial Reef | CA | 10°4'20.96"N, 99°50'31.84"E |
Shark Island | SI | 10°3'41.20"N, 99°50'40.54"E |
Sai Daeng | SD | 10°3'49.43"N, 99°50'23.80"E |
Shark Bay | SB | 10°3'39.75"N, 99°50'4.43"E |
Chalok Bay | CB | 10°3'44.77"N, 99°49'30.35"E |
Tao Tong | TT | 10°3'58.13"N, 99°49'4.76"E |
Sai Nuan and Three Rocks | SN | 10°4'45.02"N, 99°48'45.23"E |
Mae Haad | MH | 10°5'22.53"N, 99°49'14.07"E |
Sairee Beach | SRB | 10°6'0.99"N, 99°49'15.89"E |
Hin Pee Wee | HPW | 10°6'19.94"N, 99°48'47.73"E |
Sattakut Wreck | SW | 10°6'16.97"N, 99°48'47.52"E |
White Rock | WR | 10°6'27.94"N, 99°48'48.98"E |
Hin Fai Artificial Reef | HF | 10°6'43.42"N, 99°49'7.18"E |
Twins | TW | 10°7'1.93"N, 99°48'44.26"E |
Green Rock | GR | 10°7'31.24"N, 99°48'49.57"E |
Red Rock | RR | 10°7'19.92"N, 99°48'55.31"E |
Mango Bay | MB | 10°7'22.52"N, 99°50'5.06"E |
Hin Wong Pinnacle | HWP | 10°6'47.51"N, 99°51'1.95"E |
Hin Wong Bay | HWB | 10°6'12.30"N, 99°50'58.63"E |
Mao Bay North | AMN | 10°5'51.85"N, 99°51'7.69"E |
Mao Bay | AM | 10°5'32.95"N, 99°51'9.29"E |
Laem Thien | LT | 10°5'19.13"N, 99°51'17.64"E |
Tanote Bay | TB | 10°5'1.47"N, 99°50'57.50"E |
King Kong Rocks | KKR | 10°4'30.25"N, 99°50'46.46"E |
Chumphon Pinnacle | CP | 10°10'20.52"N, 99°46'44.49"E |
Southwest Pinnacle | SWP | 9°59'56.22"N, 99°46'44.28"E |
Sail Rock | SR | 9°56'42.47"N, 99°59'26.46"E |
The soft sediment habitats, located outside coral reefs, had a surface substrate composition of > 90% sand or silt particles. These are typically large areas lacking natural solid substrates and are therefore deficient in rugosity and stability over longer periods (discussed below). Areas of sandy substrate within or in close proximity to coral-dominated areas were included as coral reef (Fig.
Schematic outline of benthic zones classified in the surveys conducted at Koh Tao A fringing reef slope, usually near-shore B reef edge as determined by drastic reduction in reef-building scleractinian abundance, leading to zone of no stable substrate nor any Scleractinia C soft sediment habitats characterised by sand/silt dominated substrates and colonised by organisms absent/extremely rare in zones A and B. D Deepening of soft sediment slope resulting in a drastic reduction but not absence of soft-sediment colonisers. Illustrated by Pau Urgell Plaza.
For each documented species, a small number of specimens was examined closely for taxonomic purposes, with the vast majority of subsequent specimens recorded being noted for their ecology or simply their presence. Detailed specimen examination was carried out in-situ where possible or after sampling using high-magnification underwater photography. Ex-situ examination was carried externally on live specimens which were collected by hand and subsequently returned to their original habitats. All living specimens studied are here documented as ‘material examined’. Specimens were externally identified by the authors aided by in-situ photographs based on relevant literature and contrasted with known species prevalence in Thai waters (see Table
List of published literature between 1989–2020 in which sea slug diversity and distribution records in Thai waters are contributed.
References | Gulf of Thailand diversity | Total Thailand diversity |
---|---|---|
|
19 | 20 |
|
19 | 24 |
|
19 | 38 |
|
20 | 39 |
|
28 | 63 |
|
46 | 81 |
|
49 | 88 |
|
49 | 88 |
50 | 100 | |
|
51 | 101 |
|
52 | 102 |
|
111 | 203 |
|
111 | 204 |
|
154 | 239 |
|
156 | 241 |
|
160 | 245 |
Present study | 256 | 336 |
In total, 191 species of heterobranch sea slugs have been documented from Koh Tao to date across the orders Sacoglossa (14 species), Cephalaspidea (23 species), Aplysiida (5 species), Pleurobranchida (3 species), and Nudibranchia (146 species). These results more than double the known taxa from Koh Tao (new records for Koh Tao identified below by an asterisk *) and contribute a further 90 first records for Thai waters (96 for the Gulf of Thailand) from the island (not including the 32 species first documented in
Subclass Heterobranchia Burmeister, 1837
Superorder Panpulmonata Jörger, Stöger, Kano, Fukuda, Knebelsberger & Schrödl, 2010
Order Sacoglossa Ihering, 1876
Family Costasiellidae Clarke, 1984
Two specimens 3–6 mm, LB; two specimens 4–6 mm, SN.
In soft sediment habitats, beyond the coral reef where it feeds predominantly on Avrainvillea longicaulis (Kützing) G. Murray & Boodle, 1889 and less commonly on Vaucheria sp. Depth 10–18 m.
Costasiella kuroshimae is currently known from the Indo-Pacific including the Red Sea (
Due to the original description of the species being entirely based on external features, the identity of numerous similar species and the extent of the variability of the species has remained unclear for several years. Molecular work (
A Costasiella cf. kuroshimae 6 mm B Costasiella usagi 6 mm C Limapontiidae sp. 3 mm D Plakobranchus noctisstellatus 21 mm (photograph by Pau Urgell Plaza) E Plakobranchus ocellatus 32 mm F Plakobranchus papua 27 mm G Plakobranchus papua 21 mm on Holothuria edulis H Elysia aowthai 12 mm I Elysia asbecki 15 mm J Elysia cf. marginata 70 mm K Elysia mercieri 12 mm L Elysia obtusa 9 mm.
Three specimens 2–6 mm, LB.
In soft sediment habitats beyond the coral reef where it feeds predominantly on Avrainvillea longicaulis and less commonly on Vaucheria sp. An individual of the species has been observed to be naturally captured and ingested by the free-living coral Heteropsammia cochlea with subsequent investigations suggesting the species may represent viable prey for the coral in soft sediment habitats (
Across the Indo-Pacific including India (
One specimen 3 mm, location unknown.
Local ecology is unknown.
Unknown.
Similar to Ercolania translucens Jensen, 1993 or Stiliger sp. 7 in
Superfamily Plakobranchoidea Gray, 1840
Family Plakobranchidae Rang, 1829
One specimen 28 mm, SN; two specimens 26–31 mm, TT.
From deeper soft sediments outside coral reef habitats. Depth 15–25 m.
Vanuatu, Indonesia, Papua New Guinea (
Three specimens 25–32 mm, CB.
From shallow soft sediments to sandy areas along the reef edge. Rarely in deeper soft sediment habitats beyond the reef edge. Depth 0.5–11 m.
Plakobranchus ocellatus and P. cf. ocellatus are currently considered widespread across the Indo-Pacific including Kenya, Zanzibar, the Red Sea, Maldives, Seychelles, Réunion (
Specimens from Koh Tao were recently reviewed by
Three specimens 19–30 mm, SN.
Abundant in shallow soft sediment habitats and among the corals and soft sediments of the reef edge. Uncommon, but present in dense coral reef habitats. Rare in deeper soft sediment habitats outside the coral reef. Has been observed being ingested naturally by the scleractinian coral Pleuractis paumotensis (Stutchbury, 1833) but is mostly considered unpalatable by such corals (
Known only from the Philippines, Malaysia, Indonesia, and Papua New Guinea (
Specimens from Koh Tao were recently reviewed (
One specimen 14 mm, LB; one specimen 16 mm, TT.
From deeper soft sediments outside coral reef habitats. Depth 10–24 m.
Guam, Australia, and the Gulf of Thailand (
One specimen 15 mm, HF; one specimen 23 mm, TW.
In coral reef habitats throughout the island. Depth 3–18 m.
Australia, Samoa (
Three specimens 65–82 mm, CB.
Mostly recorded from specimens inhabiting a shallow, isolated patch of Halimeda macroloba Decaisne, 1841 in soft sediment habits, although presumably feeding on other nearby algae. Also observed from shallow coral reef habitats, rarely. Depth 0.5–6 m.
Elysia marginata is at present recorded from the Indo-Pacific including Myanmar (
Recent molecular investigations (
One specimen 12 mm, SO.
Found upon concrete artificial reefs in soft sediment habitats that formed part of coral restoration efforts. Depth 11–14 m.
Across the Indo-Pacific including the Red Sea (
Elysia mercieri is known to be predated upon by the nudibranch Gymnodoris okinawae Baba, 1936 (
One specimen 9 mm, CB; one specimen 12 mm, TW.
Among rubble, particularly found underneath the skeletons of dead Fungiidae corals, in shallow coral reef habitats. No association with prey was observed but is known to be part of a group of species feeding on the alga Bryopsis (
Across the Indo-Pacific including India (
Both individuals recorded were found adhering to the underside of dead fungiid skeletons. The species is locally rare and here included as a first record for the Gulf of Thailand and Thai waters in general.
Three specimens 3–7 mm, CB; one specimen 4 mm, SRB.
Feeds on Halimeda macroloba, on which it is highly cryptic. Host and prey found in soft sediment habitats near coral reefs between 0.5 and 9 m depth. Multiple individuals may be found feeding on a single prey item. Populations of the host algae H. macroloba have been found at only two locations at the island, 6–9 m depth at SRB and a small intertidal patch at site CB at 0.5–1.5 m. The abundance of E. pusilla has been found to be greater at CB than on the larger but less dense population of H. macroloba at site SRB.
Widespread across the Indo-Pacific including the Red Sea (
The status of Elysia pusilla and its taxonomic implications for the genus needs closer investigation (
One specimen 37 mm, CB.
In soft sediment habitats outside the coral reef. Observed feeding on a pinnate form of Caulerpa racemosa (Forsskål) J. Agardh, 1873 which is found chiefly in soft sediment habitats beyond the coral reef. A more lenticular/globular form of the algae can be abundant in some shallow, degraded reef habitats; however, specimens from Koh Tao not been observed associated with this variety. Depth 12–20 m.
At present Elysia tomentosa is considered widespread across the Indo-Pacific (but see remarks below) including Iran (
Recent molecular investigations have found that specimens recognised as Elysia tomentosa likely correspond to a complex of at least six species (
1 specimen 18 mm, HWB; 2 specimens 8–15 mm, LB.
Found in coral reef habitats throughout the island. Depth 2–25 m.
Thuridilla gracilis sensu lato is known from Maldives, Seychelles (
The taxonomic status of Thuridilla gracilis is at present unclear. Recent works (
Superfamily Cylichnoidea H. Adams & A. Adams, 1854
Family Colinatydidae Oskars, Bouchet & Malaquias, 2015
One specimen 3 mm, CB.
Endobenthic in sand in coral reef habitats. Depth 3–11 m.
The genus is currently recognised to be monospecific with Colinatys alayoi (Espinosa & Ortea Rato, 2004) known from the Bahamas, Cuba, Florida, and Martinique Island (
A Elysia pusilla 7 mm B Elysia cf. tomentosa 37 mm C Thuridilla cf. gracilis 18 mm D Colinatys sp. 3 mm E Aliculastrum cylindricum 30 mm F Aliculastrum debilis 28 mm G A. debilis (shell 12 mm) captured alive and predated by Haptosquilla cf. nefanda H Roxaniella multistriata 9 mm I Atys semistriatus 8 mm J Atys sp. 11 mm K Diniatys dentifer 5 mm L Diniatys dubius 6 mm.
Family Haminoeidae Pilsbry, 1895
Two specimens 25–30 mm, LB.
In soft sediment habitats outside the coral reef where it is found associated with cyanobacterial mats on the benthos. May also be found near the reef edge when cyanobacteria abundances increase, indicating a possible seasonal influence in abundance. Often found together with other Haminoeidae species from Koh Tao, apart from Atys sp. It may be a probable prey species for the mantis shrimp Haptosquilla cf. nefanda (Kemp, 1911) (see ecology of Aliculastrum debilis below). Depth 8–18 m.
Widespread across the Indo-Pacific including Mozambique (
Two specimens 30 mm, LB; one specimen 28 mm, TB.
Extremely similar to A. cylindricum (see above). Often found together with other Haminoeidae species from Koh Tao, apart from Atys sp. As part of the present surveys, observations were made of hunting and capture of specimens of A. debilis by the stomatopod Haptosquilla cf. nefanda (Fig.
Across the western Pacific including the Philippines, Guam, Tahiti, Fiji, and Hawaii (
Two specimens 5–9 mm, LB.
Very similar to those of Aliculastrum spp. though more regularly found immersed within/under cyanobacterial mats than on top. Often found together with other Haminoeidae species from Koh Tao, apart from Atys sp. Depth 8–18 m.
Known across the Indo-Pacific including Tanzania, the Philippines, Fiji (
Two specimens 8–12 mm, LB.
Very similar to R. multistriata, with which it is often found. A possible prey species for the mantis shrimp Haptosquilla cf. nefanda (see ecology of Aliculastrum debilis). Depth 8–18 m.
Across the Indo-Pacific including Japan (
One specimen 11 mm, AMB.
In soft sediment habitats outside the coral reef. Depth 23 m.
Atys sp. 6 (
Three individuals 2–5 mm, LB.
Very similar to the other soft-sediment associated Haminoeidae such as Aliculastrum spp., R. multistriata etc., which are often found together. Depth 8–18 m.
Diniatys dentifer is known from Madagascar, the Philippines, Japan, Indonesia, Papua New Guinea, Guam, Hawaii, French Polynesia (
Three individuals 3–6 mm, LB.
Very similar to D. dentifer. Depth 8–18 m.
Diniatys dubius is known from the Philippines, Indonesia, Papua New Guinea, Guam, Hawaii (
Three individuals 2–5 mm, LB.
While rarer than most other soft sediment associated Haminoeidae spp., from Koh Tao, the strong association with cyanobacterial mats is a shared feature across these species. Depth 8–18 m.
A Haloa sp. 5 mm B Lamprohaminoea ovalis 18 mm C L. ovalis 9 mm (photograph by Elouise Haskin) D L. ovalis 4 mm E Chelidonura cf. castanea 62 mm F Chelidonura punctata 32 and 34 mm G Niparaya sp. 4 mm H Philinopsis speciosa 18 mm I ‘Philinopsis’ coronata 35 mm (photograph by Phannee Mccarthy) J Tubulophilinopsis lineolata 38 mm K Tubulophilinopsis pilsbryi 35 mm L Tubulophilinopsis reticulata 30 mm (photograph by Kirsty Magson).
Unknown.
Six individuals 9–35 mm, SB; eight individuals 6–21 mm; LB.
White morphs (Fig.
Lamprohaminoea ovalis known from the Red Sea, Oman, Philippines, Vanuatu, Guam, French Polynesia, Hawaii (
With the recent comprehensive review of the genus Lamprohaminoea (
Family Aglajidae Pilsbry, 1895 (1847)
Two specimens 62–74 mm, TT.
In soft sediment habitats outside the coral reef. Depth 22–26 m.
Chelidonura castanea is currently known only from the Maldives (
Specimens from Koh Tao differ from those originally described by lacking orange spots across the dorsum, instead having only two tiny orange spots on the anterior portion of the head, on either side of the mouth. Additionally, the body is uniformly deep reddish brown with a thin white line on the upper margin of the cephalic shield. In the larger specimen (74 mm), both orange spots and the white line were markedly less distinct. The presence and absence of yellow/orange spots in such Aglajids has been shown to be an unreliable character for species delimitations (
Four specimens 32–39 mm, TT.
In soft sediment habitats outside the coral reef. Depth 22–26 m.
Chelidonura punctata is currently known from Kenya (
Two specimens 4 mm, CB; one specimen 3 mm, TT.
Ecology. Among rubble in coral reef habitats and soft sediments near the reef edge. Depth 4–8 m.
Niparaya sp. 3 is currently known only from eastern Malaysia (
Two specimens 18–26 mm, SB; one specimen 14 mm, TT; one specimen 11 mm, SN.
In soft sediment habitats outside the coral reef. Depth 14–26 m.
Widespread across the Indo-Pacific including Mozambique (
One specimen 35 mm, SRB.
In soft sediment habitats outside the coral reef. Depth 10 m.
‘Philinopsis’ coronata
is known from the Philippines (
The taxonomic validity of this species name remains unresolved after it was designated the type species for the recently erected genus Spinophallus by
One specimen 45 mm, LB; two specimens 32–38 mm, TT.
In soft sediment habitats outside the coral reef. Depth 12–18 m.
Currently known only from Australia (
Two specimens 35–39 mm, SN; two specimens 25–42 mm, TT; one specimen 19 mm, MB.
Abundant in soft sediment habitats outside the coral reef. A single individual of the species was observed being ingested by the scleractinian coral Heteropsammia cochlea (
Widespread across the Indo-Pacific including Mozambique (
One specimen 30 mm, SN.
In soft sediment habitats outside the coral reef. Depth 14–16 m.
Widespread across the Indo-Pacific including Mozambique (
One specimen 3 mm, LB.
In soft sediment habitats outside the coral reef. Depth 24 m.
Currently known only from the Gulf of Thailand, documented here for the first time.
A Migaya sp. 3 mm B Siphopteron makisig 3 mm C Siphopteron sp. 3 mm (photograph by Will Malsukum) D Philine orca 3 mm E Aplysia kurodai 30 mm (photograph by Geoffrey Chamayou) F Aplysia nigrocincta 9 mm G Bursatella cf. ocelligera 65 mm (photograph by Elouise Haskin) H Stylocheilus longicauda 45 mm (photograph by Kirsty Magson) I Stylocheilus striatus 29 mm J Berthella cf. caledonica 10 mm K Berthella martensi 60 mm (photograph by Paddy Steele) L Pleurobranchus forskalii 130 mm (photograph by Tine Kvamme).
Genus Siphopteron Gosliner, 1989
One individual 3 mm, SO.
In soft sediment habitats outside the coral reef. Depth 12–16 m.
Currently known only from the Philippines, Indonesia, and Australia (
One individual 3 mm, RR.
In soft sediment habitats outside the coral reef. Depth 20 m.
Unknown
Genus Philine Ascanius, 1772
Two individuals 2–4 mm, CB; one individual 3 mm, SB.
Observed exclusively under dead Fungiidae coral skeletons where it is extremely cryptic, although it may be abundant. The only cephalaspidean species recorded exclusively from the coral reef habitat at Koh Tao. Depth 3–8 m.
Widespread across the Indo-Pacific including Japan (
Superfamily Aplysioidea Lamarck, 1809
Family Aplysiidae Lamarck, 1809
Two specimens 60–81 mm, SN; one specimen 30 mm, MH.
In soft sediment habitats, occasionally found in aggregations, although more often observed as solitary. Depth 8–22 m.
Currently known only from China (
One specimen 9 mm, SB.
Under a dead Fungiidae coral among rubble in shallow coral reef habitats. Depth 6 m.
Across the Indo-Pacific including Mozambique, Mauritius, the Philippines, Indonesia, Papua New Guinea, Vanuatu (
One specimen 65 mm, SB.
In soft sediment habitats where it grazes on cyanobacterial mats on the benthos. Depth 18–25 m.
Bursatella ocelligera is known only from the Philippines (
One specimen 45 mm, SRB; one specimen 30 mm, TT.
In soft sediment habitats rarely and upon mooring ropes where it grazes on cyanobacteria. Far less common than S. striatus. The association with mooring ropes is believed to be driven by its pelagic lifestyle, as these ropes act as mid-water substrates for cyanobacterial growth. Depth 5–18 m.
Circumtropical including Brazil (
Recent work (
One specimen 25 mm, SRB; one specimen 22 mm, CB; one specimen 29 mm, TT.
From shallow and deep soft sediment habitats grazing of mats of cyanobacteria on the benthos. Depth 1–18 m.
Circumtropical including Brazil (
Order Pleurobranchida Deshayes, 1832
Superfamily Pleurobranchoidea Gray, 1827
Family Pleurobranchidae Gray, 1827
Three specimens 10–15 mm, TW.
Under coral rubble in shallow coral reef habitats. Depth 6–8 m.
Berthella caledonica is known from New Caledonia (
Overall colour variable from pink to light or dark brown, with numerous small, low tubercles across the dorsal surface, often surrounded by a brown ring, and with dark brown apices. A prominent brown mark surrounded by a diffuse ring of translucent white is located centrally on the dorsal surface. While specimens from Koh Tao resemble the description of Berthella caledonica (Risbec, 1928) rather well, records of Berthella africana (Pruvot-Fol, 1956) have also been made from Thailand, with an unclear locality (
One specimen 60 mm, LB.
Exclusively recorded from soft sediment habitats outside coral reefs. Depth 11–21 m.
Widespread throughout the Indo-Pacific including the Red Sea (
Individuals from Koh Tao have a dark, almost black mantle with numerous inconspicuous black spots. Berthella martensi was recorded by
One specimen 265 mm, SN; one specimen 55 mm, TT.
Exclusively recorded from soft sediment habitats outside coral reefs. Observed feeding on colonies of the tunicate Didemnum molle Herdmann, 1886. Depth 11–21 m.
Widespread throughout the Indo-Pacific including Mozambique (
Suborder Doridina Odhner, 1934
Superfamily Doridoidea Rafinesque, 1815
Family Actinocyclidae O’Donoghue, 1929
One specimen 10 mm, SO.
Among rubble in coral reef and reef edge habitats. Associated with an unidentified pale/creamy white sponge. Depth 4–12 m.
Across the Pacific including Australia (
A Hallaxa iju 10 mm B Hallaxa indecora 8 mm C Cadlinella ornatissima 30 mm (photograph by Mati Pauner) D Ceratosoma tenue 85 mm E Chromodoris mandapamensis 50 mm (photograph by Tine Kvamme) F Chromodoris cf. mandapamensis 7 mm G Chromodoris cf. balat 25 mm (photograph by Elouise Haskin) H Diversidoris aurantionodulosa 30 mm I Diversidoris crocea 8 mm (photograph by Pau Urgell Plaza) J Doriprismatica atromarginata 60 mm K Glossodoris cf. cincta 40 mm L Goniobranchus cf. albonares 5 mm.
One specimen 8 mm, SRB; one specimen 8 mm, CB.
Among rubble and under dead fungiid coral skeletons in coral reef habitats. Occasionally near isolated unattached colonies of sponge in deeper soft sediment habitats. Associated with an unidentified pale/creamy white sponge. Depth 4–20 m.
Red Sea (
Genus Cadlinella Thiele, 1931
Two specimens 12–30 mm, TT.
On rocks and among corals at offshore pinnacles. Depth 8–25 m.
Widespread in the Indo-Pacific including New Caledonia (
One specimen 85 mm, AM.
Soft sediment habitat. Depth 26 m.
Widespread throughout the Indo-Pacific including Japan (
One specimen 50 mm, CP.
Among corals and coral rubble at offshore pinnacle sites. 9–20 m.
Widespread throughout the Indo-Pacific including Mozambique (
While externally matching the original description of the species completely, the internal anatomy was not analysed to verify this, the importance of which is particular to this and other similar species (
Three specimens 5–10 mm, CB.
Among reef rubble, in particular under dead fungiid coral skeletons, in shallow coral reef areas. Depth 2–8 m.
Goniobranchus pruna (Gosliner, 1994) is known from Madagascar and South Africa (
Very similar to Chromodoris aff. mandapamensis (
One specimen 25 mm, CB.
Among reef rubble, in particular under dead fungiid coral skeletons, in shallow coral reef areas. Depth 4–8 m
Chromodoris balat is known only from the Philippines (
Similar to Chromodoris balat in having a striated dorsum with numerous large blotches and a broken yellow-orange marginal line. This species was differentiated from the similar Chromodoris striatella Bergh, 1877 based on these and other features (
One specimen 30 mm, SI.
Found upon its pink host sponge, Darwinella sp., at deeper reef and pinnacle sites, and in muck habitats. Depth 12–30 m.
Red Sea (
This species was mistakenly identified as Ardeadoris averni (Rudman, 1985) by
One specimen 8 mm, TT.
Ecology. Coral reef habitats. Usually cryptic on its sponge, a yellow Darwinella sp. (
Distribution. Widespread throughout the Indo-Pacific including Mozambique (
One specimen 60 mm, CP.
On rocks and among corals at offshore pinnacles. Depth 8–25 m.
Widespread throughout the Indo-Pacific including South Africa, Red Sea, French Polynesia, Solomon Islands, China, Philippines (
One specimen 40 mm, HWB.
Coral reefs throughout the island. Depth 5–15 m.
Papua New Guinea, the Philippines and Madagascar (
Previously recorded as Glossodoris cincta (
Three specimens 4–6 mm, CB.
Observed under dead fungiid coral skeletons and occasionally among rubble in shallow coral reef. Depth 3–8 m
Goniobranchus albonares is known from Australia (
Externally resembling both Goniobranchus albonares (Rudman, 1990) and Goniobranchus rubrocornutus (Rudman, 1985), the present species differs from the former by possessing a broken submarginal band of deep red and from the latter by the presence of completely white rhinophore clubs and gills as opposed to red. There is significant overlap in the range of both species, with G. rubrocornutus known from Australia, Hong Kong, and Japan (
One specimen 45 mm, SN.
Locally rare, known only from soft sediment habitats outside the coral reef. Depth 12–16 m.
Across the Indo-Pacific including Myanmar (
A Goniobranchus aureopurpureus 45 mm B Goniobranchus fidelis 12 mm C Goniobranchus geometricus 15 mm (photograph by Kirsty Magson) D, E Goniobranchus sinensis different morphs, 34 mm (D) and 55 mm (E) F Goniobranchus tumuliferus 10 mm G Goniobranchus verrieri 10 mm (photograph by Khumron Waipaka) H Goniobranchus sp. 55 mm (photograph by Phannee Mccarthy) I Hypselodoris cerisae 4 mm J Hypselodoris confetti 24 mm (photograph by Pau Urgell Plaza) K Hypselodoris decorata 18 mm L Hypselodoris infucata 12 mm.
Two specimens 6–18 mm CB; one specimen 15 mm, SB; one specimen 12 mm, SD.
Coral reef habitats throughout the region. Depth 2–25 m.
Widespread throughout the Indo-Pacific including Mozambique (
One specimen 15 mm, KKR.
Locally rare, known only from the reef edge and soft sediment habitats outside the coral reef. Depth 12–25 m.
Widespread throughout the Indo-Pacific including Myanmar (
None found presently.
Soft sediment habitats outside the coral reef. Depth 18–22 m.
Across the Indo-Pacific including Indonesia (
The species has historically been recorded from Koh Tao (
Three specimens 20–55 mm, CP.
Among rocks and corals at offshore rocky pinnacles. Depth 9–22 m.
Across the Indo-Pacific including the Gulf of Oman (
Incorrectly identified as Goniobranchus trimarginatus (Winckworth, 1946) by
Two specimens 10–15 mm, CP.
Predominantly found among coral and rock at an offshore submerged pinnacle site. Also sparsely recorded from the deeper soft sediment habitats near the island. Depth 11–25 m.
Across the western Pacific including Vietnam (
One specimen 10 mm, GR.
Coral reefs. Depth 5–10 m.
Widespread throughout the Indo-Pacific including Mozambique (
Locally known only from a single individual.
One specimen 55 mm, CP.
Among rocks and corals at offshore rocky pinnacles. Depth 18 m.
Goniobranchus sp. 5 (
Red reticulated specimens of Goniobranchus have been known to represent a complex of species often attributed to G. reticulatus or G. tinctorius. Recent molecular work by
One specimen 4 mm, CB.
Among reef rubble in shallow coral reef. Depth 4 m.
Japan, Malaysia, and Taiwan (
Though a small individual, it is identified as H. cerisae based on pink and purple pigmentation and dark brown lines with white spots. An absence of orange/burnt orange pigmentation separates it from H. krakatoa Gosliner & Johnson, 1999.
Two specimens 12–24 mm, SB.
Locally found exclusively from deeper soft sediment habitats of the island. Depth 14–25 m.
Philippines, Papua New Guinea, probably Indonesia and Hong Kong (
Two specimens 8–18 mm, CB.
Abundant among reef rubble, in particular under dead fungiid coral skeletons, in shallow coral reef areas. Rare in other habitats. Depth 2–12 m.
Widespread throughout the Indo-Pacific including Malaysia, Philippines, Indonesia, Papua New Guinea, New Caledonia, Vanuatu, and the Marshall Islands (
One specimen 25 mm, SRB; one specimen 12 mm, CB; one specimen 8 mm, SB.
Juveniles and smaller individuals common under dead fungiid corals and reef rubble in shallow coral reef areas, making up some of the most abundant nudibranch taxa in some areas. Larger individuals rarer. Throughout reef and deeper soft sediment habitats. Depth 2–25 m.
Widespread and abundant in the Indo-Pacific including Mozambique (
One specimen 6 mm, CB.
Observed under a dead fungiid coral skeleton in shallow coral reef. Depth 6 m.
Hypselodoris juniperae is currently known from Madagascar (
Externally similar to both Hypselodoris maculosa (Pease, 1871) and Hypselodoris juniperae Gosliner & Johnson, 2018 in bearing thin longitudinal white lines, dark purple spots, an opaque orange marginal band and white gills with red apices. Identified as the former species in
A Hypselodoris cf. juniperae 6 mm B Hypselodoris cf. kanga 40 mm C Hypselodoris katherinae 15 mm D Hypselodoris cf. lacuna 10 mm E, F Hypselodoris cf. maritima 20 mm (E) and 25 mm (F) G Hypselodoris tryoni, right specimen 35 mm (photograph by Kirsty Magson) H Mexichromis mariei 15 mm (photograph by Pau Urgell Plaza) I Mexichromis multituberculata 30 mm (photograph by Pau Urgell Plaza) J Mexichromis trilineata 8 mm K Verconia cf. hongkongiensis 6 mm L Asteronotus cespitosus 120 mm (photograph by Emily Palmer).
One specimen 40 mm, TW; two specimens 10–25 mm, SB.
Locally found exclusively from deeper soft sediment habitats of the island. Depth 14–25 m.
Misidentified by
Hypselodoris kanga bears blueish purple lines across its dorsum which are absent in this species, instead replaced by deep blue, almost black spots. These are more abundant towards the margin where they diffuse outwards turning into blue streaks closer to the edge. Given the historic confusion surrounding H. kanga (see
Two specimens 5–15 mm, CB.
Observed under dead fungiid coral skeletons and on rocks in shallow coral reef. Depth 3–15 m.
Indonesia, Eastern Malaysia, and the Philippines (
Recorded as undescribed from Koh Tao (
One specimen 10 mm, CB.
Observed under dead fungiid coral skeletons and on rocks in shallow coral reef. Depth 5 m.
Hypselodoris lacuna is known from the Philippines, Indonesia, Japan, Papua New Guinea, Vanuatu, and Aldabra Atoll (
Dorsum centrally translucent grey with a network of opaque white lines. Gills grey with light grey apices, rhinophore stalks translucent, clubs white basally turning red with white tips. Mantle edge pale yellow areas with alternating blue spots. Differentiated from H. lacuna by having a mostly translucent grey dorsal surface rather than just isolated circles and by the pale yellow areas between the marginal ring of blue spots. Similar to Hypselodoris sp. 8 in
Three specimens 14–30 mm, TT; two specimens 15–25 mm, SWP; one specimen 20 mm, SR.
On rocks and rubble within coral reef. Depth 5–25 m.
Hypselodoris maritima is recorded from Japan (
Initially recorded as H. maritima from Koh Tao (
Two specimens 30–35 mm, CB.
On rocks and rubble within coral reef. Depth 1–30 m.
Across the Indo-Pacific including Singapore (
One specimen 15 mm, SB.
In deep soft sediment habitats. Depth 20 m. Feeding on Dysidea sp. sponge.
Widespread throughout the Indo-Pacific including India (
Locally rare with only a single individual observed in the present surveys. Sharing the same habitat and prey preference as M. multituberculata.
One specimen 30 mm, SB, one specimen 8 mm, SN.
Observed in deep soft sediment habitats throughout the island, though uncommon. Often found associated with or actively feeding on Dysidea sp. sponge which grows unattached on the benthos. Depth 14–25 m.
Widespread throughout the Indo-Pacific including, India (
Three specimens 5–8 mm, CB; two specimens 5 mm, TW.
Usually found immersed in prey sponge Dysidea sp. under rubble and dead fungiid coral skeletons in shallow coral reef habitats. Uncommon, though multiple individuals may be observed together. Depth 3–8 m.
Across the western Pacific including Indonesia (
One specimen 6 mm, CB; one specimen 4 mm, TW.
Observed under dead fungiid coral skeleton and rubble in shallow coral reef. Depth 3–8 m.
Verconia hongkongiensis is known from Japan and Hong Kong (
Initially recorded as Hypselodoris bullockii (Collingwood, 1881) from a single small specimen by
Genus Asteronotus Ehrenberg, 1831
One specimen 120 mm, CB.
Locally found exclusively in soft sediment habitats. Depth 1–16 m.
Widespread Indo-Pacific including Australia, Indonesia, Mauritius (
One specimen 25 mm, CB.
Coral reefs. Depth 5–10 m.
Widespread Indo-Pacific including the Red Sea (
A Atagema intecta 25 mm (photograph by Nick Tringham) B Atagema spongiosa 95 mm C Atagema sp. 32 mm D Carminodoris cf. bifurcata 9 mm E Discodoris cebuensis 30 mm (photograph by Kirsty Magson) F Discodoris boholiensis 55 mm G Halgerda bacalusia (unknown size, photograph by Stephan Pelletier) H Jorunna funebris 87 mm I Jorunna sp. 10 mm J Peltodoris murrea 30 mm K, L Platydoris cf. formosa 35 mm, dorsal and ventral views (photographs by Kirsty Magson).
One specimen 95 mm, CB.
Observed upon prey sponge Dysidea sp. in deeper soft sediment habitats. Depth 14–24 m.
Gulf of Oman (
One specimen 32 mm, CB.
Observed among rubble in shallow coral reef habitats at night with no observed association with prey. Depth 4–6 m.
Atagema sp. 8 (
Externally distinct from Atagema spongiosa (Kelaart, 1858) in being completely translucent pale grey, including the circular pits, separated by ridges, along the mantle. Similar to Atagema sp. 8 (
Two specimens 9–17 mm, HF.
Among rubble in shallow coral reef habitats. Depth 3–8 m.
Carminodoris bifurcata is recorded across the Indo-Pacific including from Mozambique (
Specimens from Koh Tao resemble both Carminodoris bifurcata Baba 1993 and Carminodoris flammea (
One specimen 30 mm, SN.
Locally found exclusively in soft sediment habitats. Depth 14–18 m.
Across the Indo-Pacific including the Red Sea (
One specimen 55 mm, SD.
Found exclusively in soft sediment habitats. Depth 12–24 m.
Widespread across the Indo-Pacific including India (
No specimen collected.
Shallow coral reef habitats. Depth 5–10 m.
The range of this species appears to be very limited thus far including only Myanmar (
Recorded at Koh Tao from a single individual in 2011 and not recorded since. The included figure (Fig.
Two specimens 15–25 mm, LB; one specimen 30 mm, SI; one specimen 87 mm, CB.
Abundant throughout corals and rubble in both nearshore reefs and offshore pinnacles. Rarely observed in soft sediment habitats. Depth 2–35 m; preys on blue Xestospongia sp. (
Widespread in the Indo-Pacific including the Red Sea (
Two specimens 10–15 mm, SB.
Found exclusively in deeper soft sediment habitats. Depth 16–25 m.
Jorunna sp. 7 recorded in the Philippines (
An undescribed species covered in numerous long caryophyllidia, similar to Jorunna sp. 7 in
One specimen 30 mm, SR.
Documented here from a single record found upon submerged concrete artificial substrate at an offshore pinnacle site. Depth 30 m.
Widespread in the Indo-Pacific including the Red Sea (
One specimen 20 mm, CA; two specimens 35–40 mm, SI.
Among rubble at the coral reef edge. Depth 8–16 m.
Platydoris formosa is known from Australia (
Externally, the specimens bear similarity to Platydoris formosa, as detailed by
Two specimens 6–18 mm, CB.
Exclusively found under coral rubble and the skeletons of dead Fungiidae corals. Cryptic on its pink-red sponge. Depth 3–8 m.
Unknown.
The present species is only identified based on external morphology and as such has not been identified to species level.
A Rostanga sp. 18 mm B Sebadoris fragilis 70 mm C Thordisa sp. 12 mm D Doriopsis cf. granulosa 7 mm E Doriopsis pecten 6 mm F Doriopsis viridis 4 mm G Doris cf. immonda 5 mm H, I Dorididae sp. 15 mm (H photograph by Jeremy Coz) and 8 mm (I) J Goniodoridella sp. 1 6 mm K Goniodoridella sp. 2 3 mm (photograph by Khumron Waipaka) L Trapania cf. gibbera 5 mm.
One specimen 70 mm, SB.
Found under coral rubble and the skeletons of dead Fungiidae corals. Depth 3–8 m.
Across the Indo-Pacific including Madagascar (
One specimen 12 mm, TT.
Found in soft sediment habitats at 16 m depth.
Unknown.
Dorsal colour deep red, with white rhinophore clubs and gills. The most distinctive feature of the species appears to be a pair of distinct, elongated, white, conical papillae surrounded by 5–7 white extensions radiating out at the base of each papilla, on either side of the mid-dorsal ridge. Besides these are a few smaller but still elongated white papillae randomly distributed. Additionally, the base of the white gills and translucent red rhinophore stalks is surrounded by a thin white line which similarly can be found around the margin of the mantle. The most similar known species might be T. sanguinea Baba, 1955 which may be distinguished based on the ground colour, dorsal ornamentation (covered in papillae), and the colour of the rhinophores and gills.
Genus Doriopsis Pease, 1860
One specimen 7 mm, HF.
Under reef rubble and on rocks in coral reef habitats. Cryptic on its yellow prey sponge. Depth 6–12 m.
Doriopsis granulosa is found across the Indo-Pacific including India (
While clearly resembling Doriopsis granulosa (Pease, 1860) by an overall yellow dorsum, six gill leaves arranged horizontally in an almost transverse line, and numerous low, rounded tubercles, it bears differences from other descriptions (see
Two specimens 6 mm, CB.
Among rubble in coral reef habitats. Depth 3–6 m.
Across the Indo-Pacific including Taiwan (
Two specimens 4–6 mm, CB.
Among rubble in coral reef habitats. Depth: 3–6 m.
Known from China (
One specimen 5 mm, HF.
Under reef rubble and on rocks in coral reef habitats. Cryptic on its orange prey sponge. Depth 6–12 m.
Doris immonda is known across the Indo-Pacific including Japan (
Broadly matches the description and variations highlighted by
Three specimens 6–15 mm, SN.
In soft sediment habitats beyond the coral reef.
Unknown. Currently only documented from Koh Tao.
A small dorid with a dark mantle ranging from grey to dark brown, covered in numerous small, clearly separated pustules. Gills arranged circularly, pinnate, dark brown. The lamellate rhinophores are basally dark brown with translucent white clubs and reddish brown apices with white tips. A much more in-depth analysis of this species is needed to ascertain its placement.
Family Goniodorididae H. Adams & A. Adams, 1854
One specimen 5 mm, TT; two specimens 6–8 mm, CA.
Rare and cryptic within coral reef habitats. Depth 8–24 m.
Similar to Goniodoridella sp. 2 (
One specimen 3 mm, SI.
Rare and cryptic within coral reef habitats. Depth 14 m.
Similar to Goniodoridella sp. 10 (
One specimen 5 mm, CB.
Under rubble in shallow coral reefs. Depth 4–6 m.
Trapania gibbera is known from Indonesia, Japan, and Papua New Guinea (
Specimens from Koh Tao differ slightly from Trapania gibbera
Three specimens 8–25 mm, LB.
Among colonies of Didemnid tunicates in soft sediment habitats. Depth 12–15 m.
Known only from Indonesia (
While known to be predators of Entoprocta (
A, B Trapania miltabrancha 25 mm (A) two specimens on Didemnum molle (B) C Dendrodoris coronata 30 mm (photograph by Kirsty Magson) D Dendrodoris krusensternii 45 mm E Dendrodoris elongata 29 mm F Dendrodoris fumata 55 mm G Dendrodoris nigra 16 mm H Dendrodoris tuberculosa 88 mm I Phyllidia coelestis specimens 25–35 mm J Phyllidia elegans 45 mm (photograph by Pau Urgell Plaza) K Phyllidia exquisita 30 mm L Phyllidia ocellata 60 mm.
Family Dendrodorididae O’Donoghue, 1924 (1864)
One specimen 30 mm, LT.
On rocks and among corals in shallow reefs. Depth 2–6 m.
Widespread in the Indo-Pacific including the Red Sea (
Three specimens 8–45 mm, TT; one specimen 22 mm, SB.
Exclusively recorded from soft sediment habitats beyond the fringing coral reef. Depth 14–26 m.
Widespread in the Indo-Pacific including the Red Sea (
Three specimens 29–42 mm, CB.
Found under coral rubble and the skeletons of dead Fungiidae corals. Depth 3–8 m.
Across the Indo-Pacific including the Red Sea (
One specimen 7 mm, SB; one specimen 55 mm, CB.
Found under coral rubble and the skeletons of dead fungiid corals. Depth 3–8 m
Widespread in the Indo-Pacific including South Africa, Tanzania, Madagascar, Malaysia, Palau, Vanuatu, New Caledonia, Hawaii (
Two specimens 8–16 mm, LT; one specimen 25 mm, MH.
Found under coral rubble and the skeletons of dead Fungiidae corals. More abundant towards the edge of the reef, less abundant but present in soft sediment habitats outside of the coral reef. Depth 2–25 m.
Widespread in the Indo-Pacific including Mozambique (
Two specimens 24–88 mm, TT.
Found in soft sediment habitats outside the coral reef alongside D. krusensternii. Incorrectly classified as a coral reef-associated species from Koh Tao (
Widespread in the Indo-Pacific including the Red Sea (
Genus Phyllidia Cuvier, 1797
Three specimens 25–35 mm, GR.
Abundant in coral reef habitats. Depth 3–30 m
Widespread in the Indo-Pacific including Mozambique (
One specimen 45 mm, LT; one specimen 30 mm, RR; one specimen 45 mm, SP.
Abundant in coral reef habitats. Depth 3–30 m
Widespread in the western Pacific including), Indonesia, Taiwan, Australia, Guam, Solomon Islands (
One specimen 30 mm, SI.
Rare, found in coral reef habitats, Depth 15 m.
Known from the Maldives (
One specimen 30 mm, AMN; one specimen 65 mm, SP; three specimens 45–60 mm, HF.
Abundant in coral reef, reef edge, and soft sediment habitats. Depth 3–30 m.
Widespread in the Indo-Pacific including Mozambique (
Two specimens 28 mm, CB; one specimen 33 mm, TB.
Abundant in coral reef habitats. Depth 3–30 m
Known mostly from South-East Asia and the western Pacific including Malaysia, the Philippines, Indonesia, Hong Kong, Papua New Guinea, Australia, Japan, Fiji, Vanuatu, the Solomon Islands (
External identification of some specimens yielded the incorrect inclusion of Phyllidia marindica (Yonow & Hayward, 1991) by
A Phyllidia picta specimens 28 mm B Phyllidia varicosa 55 mm (photograph by Pau Urgell Plaza) C Phyllidiella nigra 50 mm D Phyllidiella cf. pustulosa 35 mm E Phyllidiopsis loricata 28 mm F Gymnodoris cf. alba 25 mm (photograph by Tine Kvamme) G Gymnodoris ceylonica 35 mm H Gymnodoris impudica 60 mm I Gymnodoris inornata 28 mm (photograph by Pau Urgell Plaza) J Gymnodoris sp. 1 25 mm (photograph by Pim Bontenbal) K, L Gymnodoris sp. 2 14 mm, same specimen feeding on Dendrodoris elongata.
One specimen 32 mm, HWB; one specimen 55 mm, SR.
Abundant in coral reef habitats. Depth 3–30 m.
Widespread in the Indo-Pacific including the Red Sea (
One specimen 50 mm, AMN; one specimen 50 mm, TB.
Abundant in coral reef habitats. Depth 3–30 m.
Widespread in the Indo-Pacific including the Philippines, Australia, Papua New Guinea (
Two specimens 35–40 mm, TT; one specimen 35 mm, CB.
Abundant in coral reef habitats. Depth 3–30 m.
‘Phyllidiella pustulosa’ is known from India (
Morphological and molecular work (
One specimen 28 mm, SB; one specimen 15 mm, CP.
Under and among rocks and coral rubble. Depth 3–30 m.
Across the Indo-Pacific including Australia, Guam, Marshall Islands, Tahiti (
Family Polyceridae Alder & Hancock, 1845
One specimen 25 mm, SI.
Among coral and rubble in coral reef habitats. Depth 4–18 m.
Gymnodoris alba is currently believed to be found across the Indo-Pacific, from the Red Sea (
Similar to Gymnodoris alba (Bergh, 1877) but more observations and material are needed to confidently ascertain an identification. A clarification of the G. alba species complex and indeed a revision of the genus is sorely needed to provide a biogeographical and taxonomic understanding of Gymnodoris in the Indo-Pacific. This species was first recorded as Gymnodoris sp. by
Two specimen 35 mm, TB.
Exclusively recorded from soft sediment habitats beyond the fringing coral reef. Depth 14 m.
Widespread in the Indo-Pacific including Mozambique (
One specimen 45 mm, TT; two specimens 65 mm, HF; one specimen 60 mm, TW.
In coral reef, rubble and soft sediment habitats throughout the island and nearby offshore pinnacles. Depth 5–25 m.
Widespread in the Indo-Pacific including the Red Sea (
One specimen 28 mm, HF.
Among coral and rubble in coral reef habitats. Depth 6–12 m.
Widespread in the Indo-Pacific including Mozambique (
One specimen 25 mm, TT.
On rocks and among coral and rubble in coral reef habitats. Depth 4–18 m.
First documented as Gymnodoris sp. from the Gulf of Thailand by
One specimen 14 mm, CB.
Among rubble in shallow coral reef habitats, observed feeding on Dendrodoris elongata by progressively feeding on the extended mantle around the animal. Depth 6 m.
Unknown.
Resembling multiple species considered undescribed according to
One specimen 12 mm, HF.
Among rubble in coral reef habitats. Depth 6–12 m.
Gymnodoris sp. 36 (
Smooth, translucent white dorsum with club shaped gill leaves. Observed feeding on an unknown nudibranch species (Fig.
A Gymnodoris sp. 3 12 mm B Gymnodoris nigricolor 5 mm C Gymnodoris cf. nigricolor 6 mm D Gymnodoris sp. 4 9 mm E Plocamopherus tilesii 90 mm F Polycera sp. 12 mm G Tambja amakusana 6 mm (photograph by Richard Brinke) H Tambja pulcherrima 40 mm (photograph by Liam Kelly) I Tambja sp. 4 mm (photo by Kaitlyn Harris) J Thecacera sp. 6 mm K Anteaeolidiella sp. 15 mm L Aeolidiopsis harrietae 17 mm.
Two specimens 5–7 mm, SN.
Recorded exclusively from deeper soft sediment habitats, crawling on the substrate. Depth 19–27 m.
Gymnodoris nigricolor is known from the Philippines, Japan, New Caledonia, and the Solomon Islands (
The present species closely resembles externally the description by
One specimen 6 mm, SB; one specimen 7 mm, LB.
Recorded exclusively from deeper soft sediment habitats, crawling on the substrate. Depth 19–27 m.
Unknown
Strikingly similar to Gymnodoris nigricolor from Koh Tao in sharing habitat, size, and much of their external morphology except the arc of gills. Both observed specimens, several months and kilometres apart, were found to possess white club-shaped gills instead of black. As such we treat this species as likely distinct.
One specimen 9 mm, TW.
In soft sediment habitats beyond fringing coral reefs. Depth 12–16 m.
So far only recorded in the Gulf of Thailand.
Incorrectly identified as Gymnodoris bicolor (Alder & Hancock, 1864) by
Two specimens 102 mm, TT.
Exclusively recorded from soft sediment habitats beyond the fringing coral reef. Depth 18–25 m.
Plocamopherus tilesii is known from Turkey (Yokeş et al. 2012), Vietnam (
One specimen 12 mm, SN.
Exclusively recorded from soft sediment habitats beyond the fringing coral reef where it feeds on Bugulidae spp. arborescent bryozoans. Depth 12–25 m.
Polycera sp. 1 (
One specimen 6 mm, CP.
Among rocks and corals on offshore pinnacle sites.
Widespread in the Indo-Pacific including Mozambique (
One specimen 40 mm, SWP.
Locally recorded exclusively in deep soft sediment habitats at an offshore pinnacle. Depth 25–30 m.
Known from South Korea, Japan, Taiwan, Malaysia, Papua New Guinea Australia, and New Zealand (
Externally, the present species matches the species recently described by
One specimen 4 mm, TT.
Recorded from soft sediment habitats beyond the fringing coral reef where it was found on Bugulidae spp. Depth 12–25 m.
Tambja sp. 2 (
Two specimens 6–16 mm, SB; one specimen 12 mm, KKR.
Exclusively recorded from soft sediment habitats beyond the fringing coral reef where it feeds on Bugulidae spp. arborescent bryozoans. Depth 12–25 m.
Similar species are known from Mozambique (
While similar to Thecacera cf. picta Baba, 1972 from Mozambique (
Superfamily Aeolidioidea Gray, 1827
Family Aeolidiidae Gray, 1827
One specimen 15 mm, CA.
Among corals and rubble in coral reef habitats. Depth 6–10 m.
Anteaeolidiella cacaotica (Stimpson, 1855) is known from Australia and Japan (
Two specimens 21–29 mm, LB; one specimen 17 mm, SRB.
Cryptic on their prey, Palythoa sp. which has thus far only been recorded growing on isolated pieces of rubble or artificial substrate (i.e., discarded plastic) in deeper soft sediment habitats outside the coral reef.
Known from Japan (
Specimens from Koh Tao have papillate rhinophores (similar to
Two specimens 25–30 mm, SB; one specimen 23 mm, CB.
Exclusively recorded on its prey species, the zoanthid Palythoa tuberculosa (Esper, 1805), on which it is extremely cryptic. Depth 1–18 m.
Across the Pacific including Japan (
Separated from Aeolidiopsis harrietae and A. palythoae (Gosliner, 1985) by the presence of smooth rhinophores and highly elongate body with between 17–23 pairs of cerata. Unlike specimens described by
A, B Aeolidiopsis ransoni 23 mm (A) and 25 mm (B) C Baeolidia salaamica 10 mm D Cerberilla ambonensis 20 mm E Cerberilla asamusiensis 8 mm (photograph by Pau Urgell Plaza) F, G Cerberilla cf. incola variants 12 mm (F) and 7 mm (G) H Cerberilla sp. 22 mm I Limenandra confusa 12 mm J Caloria indica 26 mm K Favorinus sp. 1 5 mm L Favorinus sp. 2 5 mm
One specimen 10 mm, CA.
Among corals and rubble in coral reef habitats. Depth 6–10 m.
Across the Indo-Pacific including Tanzania (
Rhinophores with numerous small white knobs leading to white apices, a faint white ring visible on the head, anterior to the rhinophores. Foot white and wide. Oral tentacles basally translucent with white tips, often held curled closer to the head (Fig.
One specimen 10 mm, TT; one specimen 20 mm, SN.
Exclusively found in deeper soft sediment habitats outside coral reef habitats, where it exhibits an endo-benthic substrate preference. Depth 14–20 m.
Across the Indo-Pacific including Mozambique (
One specimen 8 mm, TT.
Exclusively found in deeper soft sediment habitats outside coral reef habitats, where it exhibits an endo-benthic substrate preference. Depth 14–20 m.
Across the Indo-Pacific including Australia (
Two specimens 7–12 mm, TT.
Exclusively found in deeper soft sediment habitats outside coral reef habitats, where it exhibits an endo-benthic substrate preference. Depth 14–20 m.
Cerberilla incola is known from Australia (
Cerberilla incola as described by
One specimen 22 mm, SN.
Exclusively found in deeper soft sediment habitats outside coral reef habitats, where it exhibits an endo-benthic substrate preference. Depth 14–20 m.
Cerberilla sp. 4 (
One specimen 4 mm, LT; one specimen 7 mm, LB; two specimens 6–12 mm, CB.
On rocks and under rubble, including skeletons of dead fungiid corals, in coral reef habitats. Depth 2–14 m.
Until recently, known only from the Pacific including Costa Rica (
Genus Caloria Trinchese, 1888
One specimen 26 mm, TW; two specimens 31–39 mm, SWP; two specimens 18–25 mm, HF.
Abundant in coral reef habitats, particularly at the numerous coral reef restoration sites at Koh Tao and at rocky pinnacle sites nearshore and offshore. Depth 2–30 m.
Widespread across the Indo-Pacific including Mozambique (
One specimen 5 mm, CB; one specimen 5 mm, MH.
Documented in habitats ranging from shallow reef rubble and soft sediments to deeper soft sediment habitats. Depth 5–18 m.
Favorinus sp. 8 (
Similar to Favorinus sp. 8 in
Two specimens SB, 5 mm.
Documented from deeper soft sediment habitats only. Depth 18–25 m.
Favorinus sp. 4 and Favorinus sp. 12 (
One specimen GR, 9 mm.
Found exclusively on its prey hydroid Pennaria disticha (Goldfuss, 1820) uncommonly found in both coral reef and deeper soft sediment habitats. Depth 8–25 m.
Myja longicornis is known from Indonesia (
This genus was recently reviewed and expanded based on specimens from Japan and the Gulf of Thailand (
A Myja cf. longicornis 9 mm (photograph by Phannee Mccarthy) B, C Noumeaella sp. 1 5 mm D, E Noumeaella sp. 2 5 mm (photographs by Tony Myshlyaev) F Phidiana militaris 35 mm (photograph by Pau Urgell Plaza) G Phidiana sp. 7 mm H Phyllodesmium magnum 30 mm I Phyllodesmium cf. magnum 35 mm (photograph by Pau Urgell Plaza) J Phyllodesmium opalescens 35 mm, (photograph by Guillaume Gandoin) K Phyllodesmium sp. 55 mm (photograph by Pau Urgell Plaza) L Armina occulta 65 mm.
One specimen 5 mm, CB.
Found under rocks and coral rubble in shallow coral reef habitats. Depth 4–8 m.
Noumeaella sp. 4 is known from the Philippines (
One specimen 5 mm, SN.
Recorded from soft sediment habitats beyond the fringing coral reef. Depth 15–20 m.
Unknown. Here documented as a first record for Thai waters.
Generally translucent throughout with some patches of white or light grey, and cerata edged in white with white apices. Bears resemblance to a number of species within the genus and requires a larger sampling effort to identify further.
Two specimens 30–35 mm, SO; two specimens 30–40 mm, HF.
Occasionally found among rubble and corals in reef habitats. Most abundant at some artificial reef sites at the island, which were initially constructed from steel rebar or concrete to support coral restoration efforts. It is likely that these substrates, while promoting scleractinian conservation, also support different potential prey items than might be found in coral reef or soft sediment habitats. Depth 8–25 m.
Across the Indo-Pacific including the Gulf of Oman (
One specimen 7 mm, HWP.
Among rocks and coral rubble. Depth 5–16 m.
Unknown.
Bearing some similarities to Phidiana anulifera (Baba, 1949) and Phidiana semidecora (Pease, 1860).
Genus Phyllodesmium Ehrenberg, 1831
Two specimens 30–40 mm, TT.
In coral reef habitats where it feeds on the octocoral Sinularia sp. Depth 5–18 m.
Widespread across the Indo-Pacific including Mozambique (
One specimen 15 mm, TT; one specimen 35 mm, SB.
In soft sediment habitats outside coral reef habitats, found regularly on or in the immediate vicinity of Dendronephthya octocorals, though active feeding has not yet been confirmed. Depth 16–28 m.
Unknown
The present species externally appears to match Phyllodesmium magnum Rudman, 1991, though smaller with relatively short cerata and no brown pigment on the animal, having pale yellow tips to rhinophores and oral tentacles. All individuals recorded to date were separately found on or very close to Dendronephthya spp., distant from any colonies of Sinularia which at Koh Tao grow only on rocks in shallower coral reef habitats. While it is possible that some animals recorded at Koh Tao are indeed P. magnum with differences in external colouration reflecting a drastically different diet from that known for P. magnum, the present species is provisionally treated as distinct due to its unique ecology.
One specimen 35 mm, CP.
Among coral, rocks, and discarded fishing equipment at an offshore pinnacle site. Depth 16 m.
Known from the Philippines, Hong Kong, Korea, and Japan (
Two specimens 40–55 mm, HWB.
In shallow coral reef habitats where it observed feeding on a different species of Sinularia than P. magnum. Depth 2–8 m.
Phyllodesmium sp. 2 (
Similar to Phyllodesmium sp. 2 by
Family Arminidae Iredale & O’Donoghue, 1923 (1841)
Genus Armina Rafinesque, 1814
Two specimens 65–72 mm, SN.
Exclusively known from soft sediment habitats outside the coral reef where it feeds on the sea pen Virgularia sp. Depth 14–22 m.
Known from the western Pacific including Indonesia (
Two specimens 35–49 mm, SB; one specimen 41 mm, TT.
Exclusively known from soft sediment habitats outside the coral reef where it feeds on the sea pen Virgularia sp. One individual observed feeding on partially decomposed Pteroeides sp. (Octocorallia: Pennatulidae). Depth 12–29 m.
Known from the western Pacific including Japan (
Specimens identified as Armina semperi (Bergh, 1866) from coasts of Thailand correspond to this species (
A Armina scotti 41 mm B Dermatobranchus caeruleomaculatus 55 mm C Dermatobranchus dendronephthyphagus 40 mm D Dermatobranchus fortunatus 11 mm E Dermatobranchus semilunus 33 mm (photograph by Pau Urgell Plaza) F Dermatobranchus cf. striatus 14 mm G, H Dermatobranchus sp. 1 32 mm (G) and 28 mm (H) I, J Dermatobranchus sp. 2 27 mm K Bornella hermanni 12 mm L Bornella johnsonorum 35 mm.
Two specimens 55–67 mm, SN.
Exclusive to the soft sediment habitats outside the coral reef. Depth 14–24 m.
Known from the western Pacific including Malaysia, Indonesia, and the Philippines (
Two specimens 40–50 mm, SB.
Exclusive to the soft sediment habitats outside the coral reef where it may be found feeding on Dendronephthya sp. or partially buried in the nearby silt/sand. Depth 14–24 m.
Known from the West Pacific including Japan and Australia (
Recorded as Dermatobranchus sp. by
Two specimens 7–11 mm, CB; one specimen 9 mm, SB.
On rocks and under rubble, in particular under skeletons of dead Fungiidae corals, in shallow coral reef habitats. Depth 1–18 m.
Across the Indo-Pacific including the Indian Ocean of Java (
One specimen 33 mm, TB.
Exclusive to the soft sediment habitats outside the coral reef. Observed on Dendronephthya sp. octocoral, though active feeding was not observed. Depth 14–24 m.
Known from the West Pacific including Malaysia, the Philippines, Indonesia, and Papua New Guinea (
Specimens from Koh Tao are sometimes found with a pale yellow-pink margin to the oral veil. A similar trait is known from the closely related Dermatobranchus fasciatus
Two specimens 9–12 mm, SI; two specimens 11 mm, HPW; one specimen 14 mm, SW.
On rocks in coral reef habitats where it feeds on the octocoral Briareum stechei (Kükenthal, 1908). Occasionally observed among colonies of B. stechei growing on artificial substrates (i.e., discarded fishing nets) in deeper soft sediment habitats. Depth 6–22 m.
Dermatobranchus striatus is known from Indonesia, Papua New Guinea, and Japan (see
This species was recorded as Dermatobranchus striatus van Hasselt, 1824 by
Three specimens 28–40 mm, SB.
Exclusive to the soft sediment habitats outside the coral reef where it feeds on colonies of the octocoral Dendronephthya sp. Depth 14–24 m.
Unknown.
A species that vaguely resembles but is distinct from Dermatobranchus semilunus is regularly recorded from the same habitats and locations as other soft-sediment dwelling members of the genus. Specimens of Dermatobranchus sp. 1 externally appear to have characteristics of D. fasciatus and D. semilunus. All specimens have prominent longitudinal ridges on a generally white dorsum, with numerous black spots of varying sizes distributed along the ridges and margin of the oral veil. The oral veil always has patches of grey and the margin is sometimes pigmented with a yellow-pink band which is often pale or completely absent in some specimens. The dorsal surface usually has a single horizontal diffuse band approximately one third of the total animal length. The foot is pale pink to white, sometimes with numerous small black spots. The rhinophores have white tips, dark clubs with white lines along the lamellae, and white stalks with dark grey pigment along the inner edge of the stalks often forming a dark grey band in between and anterior to the rhinophores.
One specimen 27 mm, TT.
Exclusive to the soft sediment habitats outside the coral reef. Depth 21 m.
Unknown.
Dermatobranchus sp. 2 is characterised by prominent pale yellow marginal sacs that are very visible as conical papillae. The dorsal longitudinal ridges and grooves are white to pale brown and scattered sparsely with small brown spots of varying sizes. These spots extend to the oral veil, which is noticeably whiter than the dorsum. The rhinophore stalks are translucent white followed by a sharp black band at the base of the rhinophore club. This fades into brown, becoming paler apically with a translucent white apex. The anterior foot corners are blunt and short, with the foot being white. This species resembles D. fasciatus but differs in lacking any marginal pigmentation on either mantle or foot, and possessing prominent marginal papillae that are not seen in D. fasciatus.
Family Bornellidae Bergh, 1874
Genus Bornella Gray, 1850
Two specimens 12–18 mm, HWB.
Among corals, rocks and under rubble in shallow coral reef habitats. Depth 2–12 m.
Across the Indo-Pacific including Christmas Island in the Indian Ocean, Malaysia, the Marshall Islands (
One specimen 35 mm, CB.
On rocks and under rubble, in particular under skeletons of dead fungiid corals, in shallow coral reef habitats. Depth 2–8 m.
Bornella johnsonorum is known from the Marshall Islands in the Pacific (
The present species matches the external description of the species (
One specimen 31 mm, CB.
On rocks and under rubble, in particular under skeletons of dead Fungiidae corals, in shallow coral reef habitats. Depth 2–8 m.
Widespread across the Indo-Pacific including the Red Sea (
A Bornella stellifera 31 mm B Lomanotus vermiformis 25 mm C Scyllaea fulva 45 mm D Melibe viridis 97 mm E Melibe sp. 22 mm F Eubranchus ocellatus 22 mm G Eubranchus sp. 7 mm H Coryphellina exoptata 15 mm (photograph by Tine Kvamme) I, J Coryphellina cf. lotos variants 40 mm (I) and 35 mm (J) K Samla bicolor 12 mm L Phestilla fuscostriata 15 mm.
Genus Lomanotus Vérany, 1844
Two specimens 15–25 mm, SN; one specimen 17 mm, TW; one specimen 30 mm, SI.
Cryptic on its host hydroid Macrorhynchia sp., colonies of which are found exclusively in soft sediment habitats outside the coral reef. Depth 12–34 m.
Circumtropical, recorded from Florida, the Bahamas (
Genus Scyllaea Linnaeus, 1758
One specimen 45 mm, CB.
Recorded from a single individual on floating algae Sargassum oligocystum (Montagne, 1845).
Across the Indo-Pacific including Mozambique (
Genus Melibe Rang, 1829
One specimen 12 mm, TT; one specimen 122 mm, SB; one specimen 97 mm, SN.
Recorded from the soft sediment habitats outside the coral reef; however, individuals have rarely been observed swimming near the surface closer to shore. It is likely that these individuals were disturbed as no individuals have been recorded in shallower reef or sandy habitats after five years of survey. Depth 14–24 m.
Across the Indo-Pacific including Mozambique (
Two specimens 15–22 mm, TT.
In soft sediment habitats beyond the coral reef, grazing upon the substrate. Depth 17–22 m.
Melibe engeli Risbec, 1937 is known across the Indo-Pacific including Mozambique (
Externally similar to Melibe engeli and Melibe sp. 1 (
Family Eubranchidae Odhner, 1934
Genus Eubranchus Forbes, 1838
One specimen 22 mm, SN.
On its prey hydroid Idiellana pristis Lamouroux, 1816 rare in soft sediment habitats and absent from the coral reefs of Koh Tao. Depth 12–24 m.
Eubranchus ocellatus is known from the Red Sea (
It necessary here to clarify the brief historical records of Eubranchidae in Thai waters.
One specimen 7 mm, HF.
From an artificial reef structure at a reef restoration site in coral reef habitats. Depth 8–11 m.
Eubranchus sp. 2 (
The present species bears numerous dark brown to black spots throughout its body with bulbous transparent cerata with the digestive gland clearly visible. The dorsal colour and that of ceratal tips is a pale yellow-brown. The rhinophores, oral tentacles, and head are colourless with numerous small white patches spread throughout.
Genus Coryphellina O’Donoghue, 1929
One specimen 15 mm, GR.
Among rocks and corals in coral reef habitats. Depth 5–15 m.
Widespread across the Indo-Pacific including Mozambique (
This species was recently transferred to the genus Coryphellina in an extensive revision of the family Flabellinidae (
Two specimens 35–40 mm, HWP; two specimens 35 mm, KKR.
Often feeding on hydroids growing among rocks and corals in deeper coral reef habitats and on stable substrates such as discarded nets and the remains of large terrestrial plant matter in soft sediment habitats. It is likely that the currently unknown prey hydroid of this species is able to grow in greater abundance away from shallow coral reef habitats. Depth 10–35 m.
Coryphellina lotos is currently known only from Japan (
Specimens from Koh Tao strongly resemble Coryphellina lotos ; however, a few differences may indicate a possible cryptic species. While C. lotos is described as light violet with parts of the rhinophores, oral tentacles, and cerata apices as lilac to reddish lilac (
Genus Samla Bergh, 1900
Two specimens 9–12 mm, CB; one specimen 18 mm, SI; one specimen 16 mm, SN.
On rocks and under rubble, including skeletons of dead Fungiidae corals, in coral reef habitats. Depth 2–14 m.
Widespread across the Indo-Pacific including the Red Sea (
Genus Phestilla Bergh, 1874
Two specimens 11–15 mm, SRB; one specimen 12 mm, HF; one specimen 18 mm, LT.
Exclusively on its prey, the scleractinian coral Pavona decussata Dana, 1846, which is abundant throughout the depth range of coral reefs around Koh Tao. Depth 1–19 m.
Phestilla fuscostriata was previously known only from Hong Kong (
One specimen 30 mm, HWB; two specimens 25–30 mm, SO.
Exclusively on or in the immediate vicinity of its prey, the scleractinian coral Porites in coral reef habitats. Locally observed to be predating colonies of Porites lobata Dana, 1846, Porites lutea Milne Edwards, 1860, and Porites sp., all of which have been observed hosting the distinctive egg ribbons of the species. Depth 1–16 m.
Widespread across the Indo-Pacific including the Red Sea (
A Phestilla lugubris 30 mm B Phestilla melanobrachia 30 mm (photograph by Tom Jang) C Phestilla cf. minor 18 mm D Phestilla viei 20 mm E Phestilla subodiosa 2 mm F Phestilla sp. 1 8 mm G Phestilla sp. 2 4 mm H Phestilla sp. 3 3 mm I Trinchesia sp. 1 10 mm (photograph by Wanraya Kraikruan) J Trinchesia sp. 2 40 mm K Trinchesia sp. 3 45 mm (photograph by Pau Urgell Plaza) L Trinchesiidae sp. 6 mm.
Two specimens 15–30 mm, SI.
Exclusively on or in the immediate vicinity of its prey, the scleractinian corals Tubastraea spp., in coral reef habitats. Depth 6–32 m.
Widespread across the Indo-Pacific including Red Sea (
Four specimens 15–25 mm, CA.
Exclusively on or in the immediate vicinity of its prey, the scleractinian coral Porites in coral reef habitats. Locally observed to be predating colonies of Porites lobata and Porites lutea. Depth 6–14 m.
Across the Indo-Pacific including Tanzania, Australia, Hawaii (
Recent molecular analyses have revealed Phestilla minor to be a complex of up to six distinct species (
Two specimens 20–25 mm, AMN; one specimen 25 mm, TT; one specimen 33 mm, SB.
Exclusively on its prey, the scleractinian coral Pavona in coral reef habitats. Locally observed to be predating colonies of Pavona explanulata Lamarck, 1816. Depth 4–22 m.
Phestilla viei is known from Madagascar, Thailand, Philippines, Indonesia, and Papua New Guinea (
One specimen 2 mm, SB.
Exclusively on it its prey coral Montipora, in coral reef habitats. Depth 2–8 m.
Phestilla subodiosa is currently known from Thailand (type locality Koh Tao) and South Korea, and possibly Singapore (
Two specimens 8–14 mm, CB.
Exclusively on its prey, the scleractinian coral Goniopora in coral reef habitats. Locally observed to be predating colonies of Goniopora fruticosa Saville-Kent, 1891. Depth 2–8 m.
Phestilla sp. 3 (
Two specimens 4–7 mm, SB.
Exclusively on its prey, the scleractinian coral Acropora sp. in coral reef habitats. Depth 2–8 m.
Unknown. Here representing a first record for Thai waters.
One specimen 3 mm, HWB.
Recorded from a single individual found during sampling of the rare octocoral Nanipora (
Unknown.
Very similar to Phestilla subodiosa, which is considered an obligate feeder of Montipora sp. corals. While Montipora corals were observed in the vicinity, the present sample was observed upon Nanipora, although no feeding or egg masses were observed. The specimen has smooth rhinophores and oral tentacles with clear indications of a darker band approximately halfway on both. Cerata with clearly visible digestive glands and a distinct bulge followed by a subterminal black band (broken up into black spots in some cerata), terminating in translucent rounded apices.
One specimen 10 mm, TW.
Among rocks and corals in coral reef habitats. Depth 5–10 m.
Cuthona sp. 2 (
Incorrectly identified as Cuthona ornata Baba, 1937 by
Two specimens 35–40 mm, TT; one specimen 45 mm, TW.
Cryptic on its host hydroid Macrorhynchia sp., colonies of which are found exclusively in soft sediment habitats outside the coral reef. Depth 12–24 m.
Tenellia sp. 17 is known only from the United Arab Emirates (
Mistakenly identified as Cuthona yamasui Hamatani, 1993 by
Three specimens 12–45 mm, SN.
Cryptic on its host hydroid Macrorhynchia sp., colonies of which are found exclusively in soft sediment habitats outside the coral reef. Depth 12–24 m.
‘Cuthona yamasui’ (
Very similar to Trinchesia sp. 2, with which it shares a prey species (alongside Lomanotus vermiformis). Distinguished by the presence of a white body with brown rhinophores and oral tentacles, unlike the brown body with white markings around the rhinophores as seen in Trinchesia sp. 2. Cerata in the present species are basally transparent (with the blue digestive glands clearly visible) turning blue with a thin black band, a thick yellow band, and another thin black subapical band below translucent apices. The cerata of Trinchesia sp. 2 on the other hand are basally opaque white followed by a distinct large blue band and yellow apices with colourless tips. The present species is similar to Tenellia sp. 15 and Tenellia sp. 16 of
One specimen 6 mm, HF.
On artificial reef structures feeding on the same orange Corydendrium sp. hydroid as Unidentia aliciae, on which it is cryptic. Depth 8–12 m.
Cuthona sp. 19 (
Genus Unidentia Millen & Hermosillo, 2012
Three specimens 19–24 mm, HF.
On artificial reef structures feeding on the same orange hydroid Corydendrium sp. as Trinchesiidae sp. This hydroid is predominantly known from artificial reef structures at Koh Tao (see
Family Janolidae Pruvot-Fol, 1933
Two specimens 55–70 mm, LB; one specimen 38 mm, SW.
In soft sediment habitats outside the coral reef. Depth 18–28 m.
Across the Indo-Pacific including the Red Sea (
Family Tritoniidae Lamarck, 1809
One specimen 10 mm, CB.
Under rubble, among small hydroids, in coral reef habitats. Depth 5–8 m.
Across the Indo-Pacific including India (
The single specimen observed from Koh Tao appears paler than is typical for the species, though such colour differences are not unheard of (see
One specimen 35 mm, GR.
Known from a single specimen observed on the octocoral Echinogorgia sp. Depth 14 m.
Tritonia sp. 7 (
Prior to studies from Koh Tao, the documented diversity of sea slug taxa from the Gulf of Thailand numbered 111 species, with 204 species in total recorded for Thailand (Table
The first of these are those species that are documented preferentially from artificial substrates such as artificial reefs and debris. Nudibranch species such as Phidiana militaris, Trinchesiidae sp., and Unidentia aliciae were found in abundance over the years and were mostly or exclusively documented associated with prey growing on such substrates, with U. aliciae being described associated from these substrates (
The second specialist habitat type within the hard substrate habitats was the ventral surface of the skeletons of dead Fungiidae corals. These corals are known to be in high abundance around Koh Tao (
It is possible that many of these specialised organisms can colonise much of the benthic environment in the Gulf due to its particular characteristics. Apart from being among the western-most ecological regions attributed to the Pacific, the Gulf of Thailand differs greatly from the Andaman coast of Thailand, being a region inundated by heavy sedimentation due to the many rivers that flow into it and remained entirely frozen until the glacial retreat into the Holocene (
Beyond the habitat preferences, the present study further expands on the trophic dynamics of different sea slugs allowing insights into their place in the food webs of Koh Tao.
For example, predation upon sea slugs were documented from both habitat types, with predation upon Haminoeidae spp. in particular observed numerous times in the present study. In soft sediment habitats, predation by crustaceans (both decapod and stomatopod) appeared prominent (Fig.
The findings presented here highlight the need for a greater documentation and understanding of sea slug ecology in the Indo-Pacific as many questions remain regarding the habitat and prey preferences of the majority of species documented from the Gulf of Thailand and elsewhere. It is apparent that the diversity of sea slug taxa in the Gulf of Thailand has been greatly under-reported and that the marine habitats in the region support a high diversity of benthic species. With the increasing availability of SCUBA infrastructure around the Gulf, it is likely that a greater area within the region will be made accessible for the study of marine benthic environments. In conjunction, the expansion of surveys into deeper soft sediment habitats will likely further expand on the known diversity of a great number of species in these areas. Very little has been documented on the distinct biological and ecological characteristics of marine soft sediment habitats (
We would like to thank the survey team for their assistance, in particular Pau Urgell Plaza, Alyssa Allchurch, Elouise Haskin, Kirsty Magson, Genevieve Goulet, Kaitlyn Harris, Joel Rohrer, Raphael Danieau, Jan Koschorrek, Alexander Duseljee, Matthew Muncaster, Sriploy Chaisri, Rebecca Danielli, and Lena Schenke. We are grateful for comments by Vie Panyarachun, editor Nathalie Yonow and input from the reviewers which helped improve the manuscript. The first author received support via the Ratchadaphisek Somphot Endowment Fund for Postdoctoral Fellowship, Chulalongkorn University. This work was also supported by NRCT-JSPS Core to Core, UNESCO-IOC/WESTPAC, and UNESCO Japanese Funds-in-Trust and by Mubadala Petroleum (Thailand). Surveys and logistics were supported by Conservation Diver (Registered US Charity #20183007707; http://conservationdiver.com).