Research Article |
Corresponding author: Gi-Sik Min ( mingisik@inha.ac.kr ) Academic editor: Spyros Sfenthourakis
© 2018 Ji-Hun Song, Niel L. Bruce, Gi-Sik Min.
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:
Song J-H, Bruce NL, Min G-S (2018) The first records of Stenobermuda Schultz, 1982 and Tenupedunculus Schultz, 1979 from Australia, with description of two new species from the Great Barrier Reef (Isopoda, Asellota, Stenetriidae). ZooKeys 733: 1-24. https://doi.org/10.3897/zookeys.733.20474
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The genera Tenupedunculus Schultz, 1982 and Stenobermuda Schultz, 1979 are recorded for the first time from beyond the Southern Ocean, at the Great Barrier Reef, Australia. Tenupedunculus serrulatus sp. n. and Stenobermuda warooga sp. n. are described from Heron Island and Lizard Island respectively, both in the Great Barrier Reef. The genus Tenupedunculus is revised and a new diagnosis presented, with Tenupedunculus virginale Schultz, 1982, T. pulchrum (Schultz, 1982), and T. serrulatus sp. n. being retained within the genus, and the remaining species here regarded as Stenetriidaeincertae sedis (eight species).
Asellota , Australia, Great Barrier Reef, new species, Stenetriidae , Stenobermuda , Tenupedunculus
The family Stenetriidae Hansen, 1905 comprises 12 accepted genera (
In this paper, two new species of Stenetriidae are described from Great Barrier Reef, Australia: Tenupedunculus serrulatus sp. n. and Stenobermuda warooga sp. n. These are the first records of these genera from Australian waters. The genus Stenobermuda is known to occur on coral reefs in East Africa (
Including the present genera the Great Barrier Reef is now known to have four genera of Stenetriidae. The rich collections made during the Census of Marine Life’s (CoML) Census of Coral Reef Ecosystems (CReefs) Program, housed at the Museum of Topical Queensland, hold abundant specimens of the genera Liocoryphe Serov & Wilson, 1995, Tristenium Serov & Wilson, 1995, Mizothenar Serov & Wilson, 1995 and Stenetrium Haswell, 1881 (from Ningaloo Reef, Western Australia).
Sampling. See
Descriptions. See
Digital inking. Pencil illustrations were scanned and electronically inked using a graphics tablet (Wacom Intuos4, Düsseldorf, Germany) and Adobe Illustrator CS5. A dorsal view of the pleotelson for each species was drawn with the aspect specifically positioned to allow for descriptive measurements. In habitus drawings, specimen curvature of these animals often distorts the true length of the pleotelson.
Permits. Specimens were collected under permits: Great Barrier Reef Maine Park Authority GBRMPA G08/26156.1, G09/32313.1; Queensland Fisheries Service QFS 95152.
Abbreviations.MTQ – Museum of Tropical Queensland; RS – robust seta/e.
Tenupedunculus
Schultz, 1982: 77.–
Tenupedunculus elongatus Schultz, 1982; by original designation and monotypy.
Tenupedunculus elongatus (type species), south-eastern Argentine Basin, 4696 m; T. virginale (Schultz, 1982), Scotia Sea, Antarctica, 567 m; T. pulchrum (Schultz, 1982), southern Argentina, 1911 m; T. serrulatus sp. n., Great Barrier Reef, Australia, 25 m.
Species here excluded from Tenupedunculuss. str., are regarded as Stenetriidaeincertae sedis: Tenupedunculus acutum (Vanhöffen, 1914); T. beddardi (Kussakin, 1967); T. dentimanum (Kussakin, 1967); T. drakensis (Schultz, 1982); T. inflectofrons (Schultz, 1982); T. serraticaudum (Kussakin & Vasina, 1984); T. smirnovi (Vasina, 1982); and T. haswelli (Beddard, 1886).
(male). Cephalon frontal margin antennal spines small; lateral spines moderate, acute, slightly longer than antennal spines or sub-equal length. Pseudorostrum quadrate to trapezoid, wider than long. Eyes small, round. Male pereopod 1 ischium–carpus superodistal margin produced with acute process, inferodistal margins not produced; propodus moderate, length 1.2–1.5 times maximum width, 1.9–2.1 times carpus length, propodal palm transversely truncate or distally inflected; dactylus length similar to propodus distal width. Male pleopod 2 appendix masculina bluntly rounded apically, without apical setae.
(male).Body dorsal surface smooth or sparsely setose, widest at pereonites 6 and 7; pereonite 1 length greater than 0.9 times pereonite 2 length; pereonites 2–4 lateral margins convex, anteriorly acute. Pleotelson length subequal to width; lateral margins or finely serrate, sub-parallel, posterolateral spines prominent, margin posterior to spines rounded with weak or no apical lobe; dorsal surface smooth, or sparsely setose. Cephalon lateral margins smooth or finely serrate. Antennae length equal or longer than total body length, article 1 lateral spine absent. Pereopod 1 basis superior margin with irregularly spaced setae along length; propodal palm with teeth along palm margin; dactylus length subequal to propodal palm length. Pleopod 1 protopod rectangular, lateral margin setae present; rami lateral margins evenly convex. Pleopod 2 protopod longer than wide, distal apex blunt, transversely truncate; appendix masculina lateral margin groove absent. Pleopod 5 distal apex with 3–5 plumose setae.
The genus Tenupedunculus Schultz, 1982 was established as a monotypic genus based on a single male specimen lacking legs.
The characters that serve best to identify the genus Tenupedunculus are: small round eyes; anterior margin of head with only lateral spines prominent; pseudorostrum wider than long, quadrate or trapezoid; inferodistal margin of ischium–carpus in male pereopod 1 without process, and superodistal margin of ischium–carpus usually produced and acute (strongly produced as a process in T. serrulatus sp. n.). The principle differentiating and diagnostic characters for Tenupedunculussensu stricto are presented in Table
Principle differentiating characters for Tenupedunculussensu stricto (male).
Tenupedunculus sensu stricto | ‘dentimanum group’ | |
---|---|---|
1. Eyes | small, round | large, reniform |
2. Cephalon, lateral spines | moderate | large |
3. Pseudorostrum | quadrate to trapezoid, wider than long | round to acute, longer than wide |
4. Pereopod 1 ischium–carpus superodistal margin | produced, with acute process | produced, with acute process(except carpus) |
5. Appendix masculina | without apical setae | with apical setae |
6. Appendix masculina | bluntly rounded apically | excavate rounded apically |
All species of the genus, with the exception of the new species, are from the Antarctic and the sub-Antarctic region—off Argentina, Patagonian Shelf and also Scotia Sea (all Atlantic sector); at depths 500 to 4696 metres. Tenupedunculus serrulatus sp. n., from the southern Great Barrier Reef, is the first record of the genus from beyond the Southern Ocean and from depths less than 500 metres.
The species listed here are retained in Tenupedunculus Schultz, 1982, but excluded from the genus sensu stricto as they either lack the diagnostic characters of Tenupedunculus or possess unique characters that also preclude their inclusion in other stenetriid genera. The here termed ‘dentimanum group’ of species, particularly when considered in conjunction with their shared characters, potentially warrants a new genus.
‘dentimanum group’
All species share the following characters: cephalon antennal spines small, lateral spines large. Pseudorostrum anteriorly round to acute, as long as or longer than wide. Eyes large, reniform. Male pereopod 1 ischium and merus superodistal margins weakly to strongly forming an acute process, inferodistal margins not produced. Male pleopod 2 appendix masculina excavate, apically rounded, with apical setae.
Tenupedunculus beddardi (Kussakin, 1967). Southern Argentina; 680 m; similar to T. dentimanum with the following characters common to the group: pseudorostrum approximately as long as wide, lateral spines large, eyes reniform and male pereopod 1 carpus with distinct acute superodistal process. The pseudorostrum uniquely converges to a narrowly rounded apex. The appendix masculina is acute (not excavate), differing significantly from that of others in this group, but on balance the species otherwise agrees well with and is best placed within the ‘dentimanum group’ at present.
Tenupedunculus dentimanum (Kussakin, 1967). Southern Argentina; 680 m; pseudorostrum as long as posterior width, anteriorly broadly rounded.
Tenupedunculus inflectofrons (Schultz, 1982). Scotia Sea, Antarctica; 588 m; pseudorostrum rounded; male pereopod 1 not known; appendix masculina with terminal process.
Tenupedunculus smirnovi (Vasina, 1982). Patagonian Shelf; 500 m. Female only; seems to have reniform eyes, pseudorostrum stepped, acute; female pereopod 1 with ischium and merus with acute superodistal margin but not carpus; pleotelson with distinct caudomedial lobe.
Ungrouped species.
Tenupedunculus acutum (Vanhöffen, 1914). Gauss Station, Davis Sea; 3397 m; pseudorostrum longer than wide, anteriorly rounded with median point; moderate lateral spines on cephalon; eyes moderate in size (more than six ommatidia) round (eye shape is not entirely clear in the original figures); appendix masculina blunt (excavate), with apical setae; male pereopod 1 ischium and merus with acute processes but carpus without process. Eye size and shape precludes inclusion of T. acutum in the ‘dentimanum group’.
Tenupedunculus haswelli (Beddard, 1886). Rio del la Plata; 1097 m; eyes reniform; male pereopod 1 with superodistal process on carpus (i.e. pereopod 1 similar to Tenupedunculus serrulatus sp. n.); not evident if there is a rostrum or pseudorostrum; eyes reniform; appendix masculina not known.
Tenupedunculus drakensis (Schultz, 1982). Tierra del Fuego, Argentina; 548 m; pseudorostrum rounded to acute; weak lateral spines on cephalon; reniform eyes; appendix masculina not known; male pereopod 1 not known. Originally placed in Protallocoxa Schultz, 1978 this species was later transferred to Tenupedunculus by
Tenupedunculus serraticaudum (Kussakin & Vasina, 1984). South Atlantic; 500 m; pseudorostrum anteriorly round to acute; large lateral spines on cephalon; reniform eyes; appendix masculina blunt (excavate) with apical setae; male pereopod 1 carpus with superodistal process.
1 | Lateral margins of the body (from cephalon to pleotelson) with serrations; shallow-water species, found at depths less than 50 m | T. serrulatus sp. n. |
– | Lateral margins of the body (from cephalon to pleotelson) without serrations; deep-water species, found at depths greater than 500 m | 2 |
2 | Posterior margin of pleotelson distinctly produced | T. virginale |
– | Posterior margin of pleotelson not produced, obtusely or evenly rounded | 3 |
3 | Posterior margin of pleotelson obtusely rounded, with indications of uropodal bases | T. pulchrum |
– | Posterior margin of pleotelson evenly rounded, without indications of uropodal bases | T. elongatus |
All material from Capricorn Group, southern Great Barrier Reef.
♂ (4.2 mm), ‘Harry’s Bommie, Heron Island, 23.46053°S, 151.9293°E, 13 November 2010, reef slope, dead Acropora, 9 m, CReefs stn. HI10-002C, coll. C. Buxton (MTQ W33638).
2 ♂ (4.5, 4.1 mm [dissected]), same sample as holotype, (MTQ W52903). ♀ (5.8 mm [pereopod 1 dissected]), same data as holotype (MTQ W33654). ♂ (5.1 mm [pereopod 7 dissected]), Sykes Reef west, 23.4316°S, 152.0493°E, 14 November 2010, reef slope, 27 m, CReefs stn. HI10-009F, coll. J. Reimer (MTQ W33694). 5 ♀ (3.2–5.0 mm), 1 juv. (1.5 mm), Sykes Reef west, 23.4316°S, 152.0493°E, 14 November 2010, reef slope, 27 m, CReefs stn. HI10-009F, coll. J. Reimer (MTQ W33695). 20 ♂ and ♀, same data as holotype, coll. C. Buxton, stn. HI10-002B (MTQ W33673, W33644) and HI10-002C (MTQ W33636, W33642). ♀ (3.5 mm), Lamont Reef, 23.5932°S, 152.0655°E, 16 November 2010, reef slope, dead Acropora, 9 m, CReefs stn. HI10-019B/1, coll. M. Blazewicz (MTQ W33753). 5 ♂ (3.8–5.8 mm), Heron Island, southern side ‘Twin Peaks’, 23°28.357'S, 151°57.593'E, 28 November 2009, small rubble, 13–17 m CReefs stn. HI09-125F, coll. N.L. Bruce & K. Schnabel. (MTQ W52904). ♀ (5.8 mm), Heron Island 23.43238°S, 152.03375°E, 14 November 2009, CReefs stn. 018, no other data (MTQ W52905). ♂ (3.5 mm), 2 ♀ (2.8, 5.1 mm), Lamont Reef, southern side, 23°36.125'S, 152°03.152'E, 19 November 2009, coarse sand and small rubble, 9.7 m, CReefs stn. HI09-058D, coll. K. Schnabel & N.L. Bruce. (MTQ exW31591). MTQ W52906). 8 ♂ and ♀, Harry’s Canyons, Heron Reef, 23°28.389'S, 151°57.835'E, 18 November 2009, reef slope, small rubble and coarse sand, 6 m, CReefs stn HI09-045D, coll. N.L. Bruce & K. Schnabel (MTQ W52907). 2 ♂, 2 ♀, 2 imm., Sykes Reef, 23°25.929'S, 152°02.924'E, 18 November 2009, 26 m, coll. S. Smith & A. Anderson (MTQ W52908).
All Heron Island: north-eastern side, 20 November 2009, small rubble and sand at base of large bommies 7 m, CReefs stn HI09-064D (MTQ W31595). “The Patches” (=Mystery Bommie), 28 November 2009, rubble, mid-channel, 18 m, CReefs stn HI09-123C (MTQ W31604). “Harrys Bommie”, 13 November 2010, dead coral on sandy bottom, CReefs stn HI10-002B, 10 m (MTQ W33669).
From combining the Latin words ‘serrula’ (serrated) and the ending of ‘marginatus’, alluding to the serrated body margins of this species.
(male). Body (Fig.
Tenupedunculus serrulatus sp. n., male holotype. A body, dorsal view B body, lateral view, sternal keel C cephalon, dorsal view D antennula E antenna F enlargement of peduncular articles 1–4 of antenna G enlargement of antennal flagellum articles H uropod. Scale bars: 1 mm (A, B), 0.5 mm (C), 0.2 mm (D, F, G, H), 0.4 mm (E).
(male).Body (Fig.
Antennula (Fig.
Antenna (Fig.
Mandible (Fig.
Pereopod 1 (Fig.
Pereopod 1 ischium length 1.6 times width; inferior margin with one short seta; superodistal margin produced, with eight long setae, apex acute.
Pereopod 1 merus angular; merus length 1.1 times width, 0.8 times carpus length, 0.8 times ischium length; inferior margin with one short seta, two long setae; inferior submargin with a dense patch of long setae; distal margin with no setae; superodistal margin produced, apex acute, densely setose with long setae and two short setae.
Pereopod 1 carpus triangular; length 1.5 times width, 1.1 times ischium length; distal margin convex, with two short setae; inferior margin clearly defined, and densely setose along distal two-thirds only; inferior submargin with a dense patch of short setae. Superior carpal process long, bladelike; length 3.3 times width, 1.7 times carpal width; extending distally approximately half length of propodus; apex acute, densely setose; inferior margin smooth, straight, densely setose along distal two-thirds only, with several rows of setae; superior margin slightly convex, setose along full length.
Pereopod 1 propodus robust superiorly with inferior side of article flattened; length 1.5 times maximum width, 3.6 times proximal width, 2.2 times ischium length; inferior margin clearly defined, long, 0.7 times propodus length, 0.6 times superior margin length, densely covered with rows of long and serrate setae and with submarginal row of short setae; superior margin setose, setae regularly spaced. Propodal palm (Fig.
Pereopod 1 dactylus robust; length 4.8 times width, 1.4 times propodal palm width, 0.9 times propodus distal width (not including process), 0.6 times propodus length; superior margin distal third setose, with regularly spaced setae; distal margin setae regularly spaced along entire length; and more sparsely distributed long setae; mesial surface sparsely setose.
Pereopod 2 (Fig.
Pereopod 7 (Fig.
Pleopod 1 (Fig.
Uropod (Fig.
(female).Pereopod 1 (Fig.
Pleopod 2 (Fig.
Males 2.6–5.1 mm (mean 3.9 mm, n = 6); females 2.3–5.8 mm (mean 3.8 mm, n = 24).
Tenupedunculus serrulatus sp. n. is distinguished from other species of Tenupedunculus by the following unique combination characters: merus superodistal margin with strongly produced distally rounded process, more than 3.0 times as long as wide; male pleopod 2 appendix masculina bluntly rounded apically, without apical setae; and the lateral margins of the body (from cephalon to pleotelson) are serrated.
Tenupedunculus serrulatus is most similar to the deep-water species T. pulchrum (Schultz, 1982) with regard to general external appearance, but is distinguished by the following: small body size (< 6 mm vs. 9 mm for the holotype of T. pulchrum), serrations on lateral margin of the whole body (vs. smooth); the very strongly produced process on pereopod 1 carpus superodistal margin, approximately 2.5 times as long as width of carpus proximal margin (vs. weak process, approximately 0.6 times as long as width of carpus proximal margin); and appendix masculina without any acute part (vs. with small acute part). Tenupedunculus serrulatus is the only shallow-water tropical species in the genus.
Heron Island and reefs of the Capricorn Group, southern Great Barrier Reef, Australia; at depths of 6–27 m.
Stenobermuda
Schultz, 1979: 905.–
Stenetrigus Schultz, 1982: 58.
Stenobermuda acutirostrata Schultz, 1979; by original designation and monotypy.
Stenobermuda acutirostrata Schultz, 1979, Bermuda (type species); S. brucei Kensley & Schotte, 2002, Zanzibar, Tanzania; S. iliffei Kensley, 1994, Bermuda; S. mergens Botosaneanu & Iliffe, 1999, Bahamas; S. syzygus (Barnard, 1940), South Africa.
Stenobermuda is a small genus with both marine and stygobiont species. Recent diagnoses do not require modification in light of other recent revisions, and the genus can be readily recognised by the prominent, narrow, and acute rostral process, small or absent eyes, and pereopod 1 articles without prominent processes or an expanded propodus. Sexual dimorphism is weak in the genus.
A diagnostic character of the genus Stenobermuda Schultz, 1979 is the acute and distinct rostrum, but one species, S. iliffei Kensley, 1994 is described as having a rostrum but figured with a pseudorostrum. The presence or absence of a rostrum of S. iliffei therefore does need to be confirmed to assess its status within the genus. Other than the apparent difference is rostrum the species agrees entirely with Stenobermuda.
Cave species are indicated by brackets.
1 | Eyes (ommatidia) absent | [S. mergens] |
– | Eyes (ommatidia) present | 2 |
2 | Dorsal coxal plates absent | 3 |
– | Dorsal coxal plates present | 4 |
3 | Body length < 3 mm; pereopod 1 propodus narrow, length 1.9 times maximum width | [S. iliffei] |
– | Body length > 6 mm; pereopod 1 propodus expanded, length 1.1 times maximum width | S. syzygus |
4 | Eyes with five ommatidia; dorsal coxal plates large | S. acutirostrata |
– | Eyes with four ommatidia; dorsal coxal plates small | 5 |
5 | Pereopod 1 propodus expanded, length 1.1 times maximum width; rostrum proximal lateral margin straight | S. brucei |
– | Pereopod 1 propodus narrow, length 1.9 times maximum width; rostrum proximal lateral margin convex | S. warooga sp. n. |
Holotype. ♂ (1.6 mm), Yonge Reef, northern Great Barrier Reef, 14.57302°S, 145.6189°E, 10 September 2010, outer reef slope, coarse sand, 25 m, CReefs stn. LI10-126B (MTQ W32968).
3 ♂ (1.7 [all appendage dissected], 1.2 [antennula and pleopod 2 dissected], 1.5 mm [pleopod 1 dissected]), same data as holotype (MTQ W52909). 3 ♂ (1.4, 1.5 [dissected], 1.7 mm), High Rock (between Direction Islands and Ribbon Reef No. 10), northern Great Barrier Reef, 14.82462°S,145.552°E, 6 September 2010, coral rubble, 8 m, CReefs stn. LI10-092A, coll. C. Buxton (MTQ W32917). ♂ (1.2 mm), 14.57302°S, 145.61980°E, Yonge Reef, 10 November 2010, outside; small coral rubble in spur, 20 m CReefs stn LI10-126A (MTQ W52910). ♂ (1.4 mm), Yonge Reef, 14.60681°S, 145.6311°E, 20 February 2009, outer reef front., dead coral, 30 m, coll. CReefs stn LI09-15B Shawn Smith & Julian Caley (MTQ W52911).
The epithet ‘warooga’ is an Aboriginal word meaning small child, in reference to the small size of this species; noun in apposition.
(male). Body (Fig.
(male).Body (Fig.
Antennula (Fig.
Antenna (Fig.
Mandible (Fig.
Pereopod 1 (Fig.
Pereopod 1 ischium length 1.9 times width; inferior margin with one short seta; distal margin with one short setae; superodistal margin not produced, apex rounded.
Pereopod 1 merus rectangular; merus length 1.4 times width, 0.8 times carpus length, 0.9 times ischium length; inferior margin with two short setae, one long seta; distal margin with no setae; superodistal margin not produced, apex rounded, with two short setae.
Pereopod 1 carpus rectangular; length 2.0 times width, 1.3 times ischium length; distal margin convex; inferior margin clearly defined, and with four stout setae and four long setae; superodistal margin not produced, apex obliquely truncate.
Pereopod 1 propodus robust and narrow; length 1.9 times maximum width, 3.2 times proximal width, 1.3 times ischium length; inferior margin clearly defined, long, 0.7 times propodus length, 0.7 times superior margin length, lightly setose, regularly spaced setae along entire length; superior margin setae absent. Propodal palm (Fig.
Pereopod 1 dactylus convex in mid-section; length 1.9 times width, 1.3 times propodal palm width, 0.8 times propodus distal width (not including process), 0.4 times propodus length; superior margin distal third with four long setae. Distal margin with three serrate setae. Mesial surface not setose.
Pereopod 2 (Fig.
Pereopod 7 (Fig.
Pleopod 1 (Fig.
Uropod (Fig.
Not known.
Males 1.2–1.7 mm (mean 1.5 mm, n = 9).
Stenobermuda warooga sp. n. can be identified by the following unique combination characters: small body size of the adult male (< 2 mm); rostrum proximal lateral margin with convex margin; pereonite 5 distolateral margin not produced; pereopod 1 propodus narrow, length 1.9 times maximum width (Table
Comparison of diagnostic characters between S. warooga sp. n. and other species of Stenobermuda (male).
Total length (mm) of adult male | Rostrum (Pseudorostrum), proximal lateral margin | Rostrum/ Pseudorostrum | Eyes (ommatidia) | Dorsally visible coxae (size) | Pereopod 1 propodus length | |
S. warooga sp. n. | 1.6 | Convex | Rostrum | Small rounded (4 ommatidia) | Pereonites 4–6 (small) | Narrow, 1.9 times maximum width |
S. acutirostrata | 4.8 | Straight | Rostrum | Small rounded (5 ommatidia) | Pereonites 1, 4–6 (large) | Normal, 1.6 times maximum width |
S. brucei | 3.1 | Straight | Rostrum | Small rounded (4 ommatidia) | Pereonites 4–6 (small) | Expanded, 1.1 times maximum width |
S. iliffei | 2.9 | Straight | Pseudorostrum (?) | Small rounded (4 ommatidia) | Not visible | Narrow, 1.9 times maximum width |
S. mergens | 3.2 | Straight | Rostrum | Absent | Pereonites 1–6 (small) | Normal, 1.6 times maximum width |
S. syzygus | 6.5 | Straight | Rostrum | Small rounded (4 ommatidia) | Not visible | Expanded, 1.1 times maximum width |
Yonge Reef and High Rock, Lizard Island region, northern Great Barrier Reef both outer barrier reefs; at depths of 8–30 m.
This work was supported by a grant from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR201524202, NIBR201624202). J.-H. Song thanks the Museum of Tropical Queensland, Queensland Museum for provision of facilities during his visit.
The authors gratefully acknowledge Australian Biological Resources Study for providing support to NLB for this study (ABRS grant 2009-30). Material was collected under the auspices of the CReefs project organised by the Australian Institute of Marine Science (AIMS), Townsville. The CReefs Australia Project was generously sponsored by BHP Billiton in partnership with The Great Barrier Reef Foundation, the Australian Institute of Marine Science and the Alfred P. Sloan Foundation; CReefs is a field program of the Census of Marine Life. The authors thank Julian Caley and Shawn Smith (AIMS) for their excellent organisation and field support; Magda Błażewicz-Paszkowycz (University of Lodz, Poland) and Kareen Schnabel (NIWA, Wellington, New Zealand), Slava Ivanenko (Moscow State University, Russia), and James Reimer (University of the Ryukyus, Japan) for their excellent help while collecting at Lizard and Heron Islands. This is contribution number 204 of the NWU Water Research Group.