Extraordinary Echinoderms a Guide to the Echinoderms of New Zealand Version 1, 2014

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extraordinary echinoderms a guide to the echinoderms of New Zealand Version 1, 2014

outstanding ophiuroids amazingheavenly holothuriansasteroids exquisite echinoidscurious crinoids

Sadie Mills Kate Neill Owen Anderson Niki Davey with Michelle Kelly & Blayne Herr

about this guide

Echinoderms are found everywhere and are adapted to live in many habitats, from the intertidal zone down to the continental shelf, deep ocean trenches, and abyssal plains. They are an extraordinary and diverse group and we hope you will enjoy reading and using this guide to help identify them in the field.

EXTRAORDINARY ECHINODERMS is a fully illustrated working e-guide to the most commonly encountered shallow water species of sea stars, brittle stars, sea cucumbers, sea urchins and feather stars of New Zealand. It is designed for New Zealanders like you who live near the sea, dive and snorkel, explore our coasts, make a living from it, and for those who educate and are charged with kaitiakitanga, conservation and management of our marine realm. It is one in a series of e-guides on New Zealand marine invertebrates that NIWA’s Coasts and Oceans group is presently developing.

The e-guide starts with a simple introduction to living echinoderms, followed by a morphology (shape) index, species index, detailed individual species pages, and finally, icon explanations and a glossary of terms. As new species are discovered and described, new species pages will be added and an updated version of this e-guide will be made available.

Each echinoderm species page illustrates and describes features that enable you to differentiate the species from each other. Species are illustrated with high quality images of the animals in life. As far as possible, we have used characters that can be seen by eye or magnifying glass, and language that is non-technical. Information is provided in descriptive text or quick reference icons that convey information without words. Icons are fully explained at the end of this document and a glossary explains unfamiliar terms.

The contributors to this guide are:

Sadie Mills Ophiuroidea (brittle, basket and snake stars) Kate Neill Asteroidea (sea stars) Owen Anderson Crinoidea (feather stars and sea lilies) Echinoidea (sea urchins) Niki Davey Sadie Mills, Kate Neill, Owen Anderson, Niki Davey Holothuroidea (sea cucumbers)

For any ID advice on echinoderms that you encounter, please email your questions and images to [email protected]

Remember to check out

http://www.niwa.co.nz/coasts-and-oceans/marine-identification-guides-and-fact-sheets

for any updated versions!

a typical species page layout

taxonomic name of species taxonomic authority person(s) who first described this species common name of species

species images inset images show variations species classification see species index body plan icon for arrangement the basic shape of the animal, characteristic of certain groups

life history icon highlighting whether native or introduced, distribution beyond NZ, and typical size in life (scale bar)

depth range common depth range quick id icons around New Zealand highlighting shape, surface detail, habitat, and environment

abundance

information distribution details on external section of coastline and internal where species is most characters and commonly found habitat make notes of where you encountered this species and let us know if you find it at a new location

key taxonomic references it could also be ... some species are difficult to tell apart without careful identification, so check the species listed here to make sure that you have the correct species

about echinoderms

Echinoderms are found in a great number of habitats from the shallow intertidal zone, at snorkelling and diving depths, right through to the deepest trenches and abyssal plains of our oceans. So it is quite likely that whenever you are in or on the sea an echinoderm won’t be far away.

five Asteroidea Echinoderms There are classes of echinoderms, the (sea stars), Ophiuroidea (brittle, basket and snake stars), Crinoidea (feather stars and sea lilies), Echinoidea (sea urchins) and Holothuroidea (sea cucumbers).

The five classes are arranged throughout this guide according Crinoidea Echinoidea Asteroidea to their position in three subphyla Asterozoa (Asteroidea, Ophiuroidea Holothuroidea CLASS CLASS CLASS CLASS CLASS Ophiuroidea), Crinozoa (Crinoidea) and Echinozoa (Echinoidea, Holothuroidea).

Echinoderm means ‘spiny skin’ and this is seen in many of the echinoderm species, though not all of them have obvious spines. The basic body plan of an echinoderm is made up of a five-sided (pentaradial) symmetry and they don’t have a head or eyes. They all share a water vascular system, usually with tube feet, which helps them with breathing, feeding and movement in their habitat.

Within all of the groups there are species that are filter feeders, deposit feeders, scavengers and predators. With few exceptions the echinoderms have separate sexes and breed by fertilisation to create an embryo, but the sexes are usually very difficult or impossible to tell apart.

Echinoderms are fed on by fish, other echinoderms and humans. The most well-known species that make up part of the human diet are kina, Evechinus chloroticus (Valenciennes, 1846), and the sea cucumber Australostichopus mollis (Hutton, 1872). Some echinoderms have the remarkable ability of autonomous self-regeneration if they are about to be eaten or picked up. Sea cucumbers are able to eviscerate or spew out their internal organs and regrow a new set, while sea stars and brittle stars can drop an arm (or several) and regenerate it.

sea star feather star Asteroidea brittle star sea egg Crinoidea Echinoidea sea cucumber Ophiuroidea Holothuroidea

The Asteroidea are the group of echinoderms commonly known as starfish or sea stars. Many are star-shaped with a central disc, and five arms, however there are also many species with more than five arms. Arms may be long or short and the relationship between the sizes of the disc and the arms can be different between species. The bodies of starfish are made up of calcified plates which are either really obvious (like paving) or are partially or totally covered in skin, spines, and granules. Starfish can be distinguished from Ophiuroidea (brittle stars, etc) by the presence of a canal or furrow on the underside of their arms. These furrows contain the tube feet which are usually in rows of two or four.

There are seven orders of starfish; all seven orders are found in New Zealand waters and three are currently included in this guide.

Paxillosida This large group are predominantly found in soft sediment habitats. They lack an anus and their tube feet lack suckers.

Valvatida A very large order of many families, the Valvatida are dominated by species with five arms and two rows of tube feet. They include many biscuit-like starfish with obvious marginal plates but also some longer armed forms.

Forcipulatida The Forcipulatida are named after the forceps-like structure of the pedicellariae found in this group. This group contains several large, common predatory starfish that occur in the intertidal waters of New Zealand. Deep sea members are also relatively common.

Echinoderms Crinoidea Echinoidea Asteroidea Ophiuroidea Holothuroidea

outstanding ophiuroids brittle, basket & snake stars The Ophiuroidea are star-shaped echinoderms with a distinct central disc, and five slender arms, though there are some species with six, seven or eight arms.

Ophiuroids can be distinguished from Asteroidea (sea stars) by the underside of their arms, which lack a grooved canal. Their tube feet emerge from small pores in the underside of their arms, which are sometimes covered by bony scales. They are split into two orders, which look quite different from each other.

Euryalida (basket stars and snake stars)

The arms of this group coil vertically like a monkey tail and both sides of the disc and arms are covered in skin, sometimes with granules or small spines. The arms usually number five at the base, but some groups have more than five arms or arms that branch into a large number and form a coiled basket. These animals are quite often associated to corals or sponges that they cling to as a perch to reach up into the water column to catch food.

Ophiurida (brittle stars)

The arms are simple, unbranching and individual disc and arm plates are usually obvious with a few exceptions. The arm spines can be relatively spiny or smooth, very long or very short and inconspicuous, and similarly the disc plates can be unadorned forming a simple rosette pattern or can be covered in spines or granules obscuring some or all of the plates.

Echinoderms Crinoidea Echinoidea Asteroidea Ophiuroidea Holothuroidea

curious crinoids feather stars

Crinoids are the most ancient of the living classes of echinoderms and are split into two groups: stalked crinoids (sea lilies), which are usually permanently attached to rock and live exclusively in the deep sea (> 100 m); and the mobile and more diverse feather stars (comatulids), which occur in all depths.

The main body of the crinoid is called the crown which consists of a calyx (cup) housing the internal organs and numerous upward- pointing feather-like arms, composed of many small jointed segments, used to filter small organic particles from the passing currents.

The anus and mouth lie adjacent to each other on the upper surface of the calyx, leaving the underside free to bear the numerous slender jointed cirri used to grip the substrate.

Echinoderms Crinoidea Echinoidea Asteroidea Ophiuroidea Holothuroidea

exquisite echinoids sea eggs

The Echinoidea are generally circular or oval-shaped echinoderms, and come in two main forms: the ball-shaped “regular” echinoids such as the common kina, and the more variable “irregular” forms including heart urchins and sand dollars. Regular echinoids are usually found on rocky reefs while irregular species live almost exclusively on soft substrates. All New Zealand echinoids are native, but some are also found in Australia and others have a worldwide distribution. Although there are over 100 echinoid species recorded from New Zealand many either live deeper than 150 m or are tropical species found at the southern limit of their range only in the far north.

Regular echinoids

Apart from a few species not present in New Zealand, all species in the 12 living orders of regular echinoids have a round body- plan with pentameral symmetry based around 20 columns of calcium carbonate plates, and range from almost spherical to discus-shaped. They are covered in sharp spines used mainly for defence, and numerous tube feet used for movement, respiration, feeding, and defence. The five teeth visible on the lower surface are the sharp end of a complex feeding apparatus unique to this group called Aristotle’s Lantern.

Irregular echinoids

Irregular echinoids are also composed of 20 columns of calcium carbonate plates, but are distinguished from regular echinoids by their bilateral symmetry and usually live buried in soft sediment. They are mainly deposit feeders, either ingesting large amounts of sediment rich in organic material and excreting the inorganic component (cassiduloids), or selectively plucking organic particles and transporting them to the mouth with their tube feet via ciliated food-grooves (clypeasteroids) or a funnel to the surface (spatangoids). In most groups the ambulacral columns (those bearing the tube feet) form distinctive petals, and the spatangoids have bands of ciliated spines (fascioles) which create currents to aid respiration and waste management.

The strongly flattened clypeasteroids (sand dollars) have several specialisations including internal buttresses for added strength and much widened ambulacra to accommodate additional tube feet for food collecting and other tasks.

Echinoderms Crinoidea Echinoidea Asteroidea Ophiuroidea Holothuroidea

The Holothuroidea differ more than superficially from other echinoderms as they do not possess the conspicuous ‘spiny skeleton’; their body walls are soft or leathery. They are generally slow moving and/or sedentary often causing them to be targeted as food for fish and crustaceans, however, toxic compounds in body wall and a well known ability to eviscerate (discharge interior organs) and regenerate gives them quite a survival advantage.

On closer inspection the holothurians do contain remnants and features of the general echinoderm body plan. Their calcified skeleton is made up of microscopic ossicles embedded in the body wall. These ossicles display beautiful geometric shapes ranging from perfect wheels, to anchors, cups, tables and plates. These ossicles are often unique and are used to distinguish species.

They have a water vascular system which consists of anterior feeding tentacles which are actually modified tube feet and also true tube feet (Orders Molpadida and Synaptida do not have these). These tube feet vary in their arrangement on the body wall. Branching respiratory trees occupy the posterior end of the coelomic cavity and are aerated by the cloaca movements.

Holothurians are often common and conspicuous in New Zealand waters but mostly in offshore habitats. Australostichopus mollis is common in the intertidal zone, in rockpools and in the shallow subtidal zone.

There are six living orders of Holothuroidea: Synaptida, Elasipoda, Aspidochirotida, Dendrochirotida, Molpadida and Gephyrothuriida, of which the three represented in this guide are described below.

Aspidochirotida are deposit feeders with short pelate scooping tentacles. The ventral tube feet are locomotory and the dorsal are modified into papillae. Ossicles include tables, plates, rods, buttons and C-bodies. This order is the only one commercially exploited and in New Zealand waters contains the well-known conspicuous species Australostichopus mollis.

Dendrochirotida have a thick dense body wall and branching tentacles used for suspension feeding. A few have modified dendritic tentacles to enable deposit feeding. The tentacles are mucus covered catching small food items. Tube feet are usually over entire body or at least confined to the five radii. Ossicles include tables, perforated plates, baskets, cups and rods.

Molpadida have 15 modified digitate tentacles and have a clearly elongated tail. They are often smooth bodied as tube feet are absent. Ossicles contain tables, elongated rods, rocket shaped plates and caudinid cups.

Echinoderms Crinoidea Echinoidea Asteroidea Ophiuroidea Holothuroidea

21 24 25 Stegnaster inflatus Eurygonias hyalacanthus Pentagonaster pulchellus

20 23 45 22 Patiriella regularis Diplodontias dilatatus Asterodiscides truncatus

18 16 14 Astropecten polyacanthus Sclerasterias mollis Astrostole scabra

19 15 17 Luidia australiae Coscinasterias muricata Stichaster australis

35 33 26 Ophiomyxa brevirima Cryptopelta tarltoni Astrobrachion constrictum

27 28 36 Astroceras elegans Asteroporpa australiensis Ophionereis fasciata

34 29 30 Ophiopsammus maculata Ophiocentrus novaezelandiae Ophiactis resiliens

31 32 37 Clarkcoma bollonsi Ophiopteris antipodum Cenolia novaezealandiae

38 43 45 Oxycomanthus benhami Fellaster zelandiae Apatopygus recens

46 42 40 Echinocardium cordatum Pseudechinus novaezealandiae Pseudechinus albocinctus

39 41 44 Evechinus chloroticus Pseudechinus huttoni Centrostephanus rodgersii

52 48 49 Paracaudina chilensis Holothuria integra Amphicyclus thomsoni

51 47 50 Heterothyone alba Australostichopus mollis Squamocnus brevidentis

Family Asteriidae Astrostole scabra 14 Coscinasterias muricata 15 Sclerasterias mollis 16 Family Stichasteridae Forcipulatida Stichaster australis 17 ORDER

Family Astropectinidae Astropecten polyacanthus 18 Family Luidiidae Luidia australiae 19 Paxillosida ORDER Asteroidea

CLASS Family Asterinidae Patiriella regularis 20 Stegnaster inflatus 21 Family Asterodiscididae Asterodiscides truncatus 22 Family Odontasteridae

Valvatida Diplodontias dilatatus 23 Eurygonias hyalacanthus 24

ORDER Family Goniasteridae Pentagonaster pulchellus 25

Family Euryalidae Astrobrachion constrictum 26 Astroceras elegans 27 Family Gorgonocephalidae Euryalida Asteroporpa australiensis 28 Echinodermata PHYLUM ORDER

Family Amphiuridae Ophiocentrus novaezelandiae 29 Family Ophiactidae Ophiactis resiliens 30 Family Ophiocomidae Clarkcoma bollonsi 31 Ophiuroidea CLASS Ophiopteris antipodum 32 Family Ophiodermatidae Cryptopelta tarltoni

Ophiurida 33 Ophiopsammus maculata 34

ORDER Family Ophiomyxidae Ophiomyxa brevirima 35 Family Ophionereididae Ophionereis fasciata 36

Family Comasteridae Cenolia novaezealandiae 37 Oxycomanthus benhami 38 Comatulida Crinoidea CLASS ORDER

Family Echinometridae Evechinus chloroticus 39 Family Temnopleuridae Pseudechinus albocinctus 40 Pseudechinus huttoni 41

Camarodonta Pseudechinus novaezealandiae 42 ORDER

Family Clypeasteridae Fellaster zelandiae 43 Clypeasteroida ORDER

Family Diadematidae Centrostephanus rodgersii 44 Diadematoida Echinoidea ORDER CLASS

Family Apatopygidae Apatopygus recens 45 ORDER Neognathostomata

Family Loveniidae

Echinodermata Echinocardium cordatum 46 PHYLUM Spatangoida ORDER

Family Holothuriidae Australostichopus mollis 47 Holothuria integra 48 Aspidochirotida ORDER

Family Cucumariidae Amphicyclus thomsoni 49 Squamocnus brevidentis 50 Family Heterothyonidae Heterothyone alba 51 Dendrochirotida CLASS ORDER Holothuroidea

Family Caudinidae Paracaudina chilensis 52 Molpadida ORDER 13 Class Asteroidea Order Forcipulatida Family Asteriidae

classification scabra Astrostole morphology 15(1): 69 Research Freshwater Town, Marineand Asteroidea). J.C. New Zealand Journal of Astrostole (Echinodermata: (1981) scabra Prey anddietarycomposition inintertidal characteristics Paxillosida, Valvatida. NIWA Biodiversity Memoir 120:1 McKnight, D.G. New (2006) The Marine Fauna Zealand: Asteroidea of (sea-stars). 3. Orders Velatida, Spinulosida, Forcipulatida, Brisingida withaddenda to seven armed starfish It couldalsobe………. from Australia. the ChathamIslands, alsosouthwards totheSnaresIslands. Alsorecorded man-made structures, e.g.piles. Found throughout wharf New Zealandand tosubtidal.Thispredatoryspeciesiscommoninrocky areasandon Intertidal red,blue-grey.Dark surface fleshy with rows singlepointedspines, marginalplatesnotobvious. of Upper Seven thearms. arms, withdiscradius about1/10thelengthof Sclerasterias mollis Sclerasterias (11arms) Coscinasterias muricata surface –80. (Hutton, 1872) (5arms) –187. habitat environment inset image: Rachel Boschen main image: Pete Notman 5 cm 14 Class Asteroidea Order Forcipulatida Family Asteriidae

classification Coscinasterias muricata morphology food and fluctuating temperature. Marine Ecology Progress Series 233: 143–155. Barker,M., Skold, M.F., P.V.Mladenov star fissiparous sea the sexualasexualvariabilityin and reproductionSpatial of (2002) Paxillosida, Valvatida. NIWA Biodiversity Memoir 120:1 McKnight, D.G. New (2006) The Marine Fauna Zealand: Asteroidea of (sea-stars). 3. Orders Velatida, Spinulosida, Forcipulatida, Brisingida withaddenda to 11 armed starfish It couldalsobe………. throughout mainland New Zealandand Australia. Found farms. piles andundermussel aswharf such man-made structures rocky areaswhereprey asonrocky reefs, andon abundanceishigh,such tosubtidal.ThispredatoryspeciesiscommoninsandyIntertidal and obvious. Pale orange, grey/blue. rows singlepointedspines, spinesinsidelittlerosettes, of marginalplatesnot Uppersurfacewith Eleven thearms. arms, discradius ~1/6thelengthof Astrostole (7arms) scabra surface –187. Verrill, 1870 habitat : the role of : theroleCoscinasterias muricata of environment main image: ChrisWoods inset image: Pete Notman 10 cm 15 Class Asteroidea Order Forcipulatida Family Asteriidae

classification mollis Sclerasterias morphology Paxillosida, Valvatida. NIWA Biodiversity Memoir 120:1 McKnight, D.G. New (2006) The Marine Fauna Zealand: Asteroidea of (sea-stars). 3. Orders Velatida, Spinulosida, Forcipulatida, Brisingida withaddenda to Zoology 8(2):209 New ZealandJournal of Zealand. New of fiords P.southern J.Hill, some R., of Richardson, associations ShallowJ.,W.I., rock-wallbiological (1981) Main R. Grange, L. Singleton, de K.R., It couldalsobe………. Islands. Auckland tosubtidal.ThroughoutIntertidal New Zealand,from tothe the Kermadec marginal platesnotobvious. Apricottoorange orred. sitsinasmallrosette, platebearsasinglepointedspinewhich plates, each Five arms, sometimessix.Uppersurfacecovered inrows skin-covered of Astrostole (7arms) scabra surface –227. (Hutton, 1872) –187. habitat main image: Roberta D’Archino inset image: ChrisWoods environment 3 cm 16 Class Asteroidea Order Forcipulatida Family Stichasteridae

classification Stichaster australis morphology Ecology 41(3): 195–211. starfish NewZealand the of Barker,population a in M.F. recruitment and Breeding (1979) Paxillosida, Valvatida. NIWA Biodiversity Memoir 120:1 McKnight, D.G. New (2006) The Marine Fauna Zealand: Asteroidea of (sea-stars). 3. Orders Velatida, Spinulosida, Forcipulatida, Brisingida withaddenda to reef star It couldalsobe………. plus SnaresandChathamIslands. toshallowIntertidal subtidal.Commonthroughout theNew Zealandmainland Purple-pink andapricot. covered inorderedrows granules, multiple marginalplatesnotobvious. of Uppersurface Eleven thearms. arms, discradius ~1/3thelengthof Coscinasterias muricata surface (Verrill, 1871) –187. habitat Stichaster australis(Verrill).Stichaster Journal of ExperimentalMarine Biologyand Journal of inset image: Rachel Boschen environment main image: Kate Neill 5 cm 17 Class Asteroidea Order Paxillosida Family Astropectinidae

classification polyacanthus Astropecten Biodiversity Memoir116:1 H.E.S., McKnight,Clark, D.G. Asteroidea (Sea-stars) Orders Paxillosida New (2000) The Marine Fauna Zealand: Echinodermata: of and Notomyotida.

morphology comb star It couldalsobe………. Pacific. the south.Alsowidespreadinotherareasof with occasionalrecordsfurther Islands PlentyandBay theBay of New of Zealand,particularly northern Zealand region,found around Islandsandvery theKermadec commonin tosubtidal.Oftenoccursonsoftsubstrates,Intertidal e.g. sand.IntheNew orange, oftenbandedormottled. angle between anddisccarrytwo, arm uprightpointedspines. Biege, brown, with smallspines, marginalplatesobvious andcarrypointedspines. Platesin Uppersurfacecarpeted Five thearms. arms, discradius ~1/5thelengthof –196. surface Müller &Troschel, 1842 habitat environment main image: Roberta D’Archino main image: Roberta D’Archino inset image: Kate Neill 7 cm NIWA 18 Class Asteroidea Order Paxillosida Family Luidiidae

classification previously known asL.maculata Luidia australiae morphology Biodiversity Memoir116:1 H.E.S., McKnight,Clark, D.G. Asteroidea (Sea-stars) Orders Paxillosida New (2000) The Marine Fauna Zealand: Echinodermata: of and Notomyotida. It couldalsobe………. is mostcommonfrom IslandstoEastCape. theBay of Islandwhereit North Found Islandsandaround attheKermadec thenorthern tosubtidal.Oftenoccursonsoftsubstrates,Intertidal e.g. sandorgravel. cream orbeige andbrown. not obvious, butcarryenlarged grey spines. Bandedandmottleddark and small spines,surface covered marginalplates inclosely packed bundlesof Upper Seven thearms. arms, discradius about1/8or1/9thelengthof surface (7 arms) Astrostole (7arms) scabra –196. Döderlein, 1920 Döderlein,

habitat environment main image: Patrick Colin inset image: Rob Stewart 12 cm NIWA 19 Class Asteroidea Order Valvatida Family Asterinidae

classification regularis Patiriella morphology Asteroidea) based onmorphological and molecular data. P.M.,O’Loughlin, Waters, J.M., Roy, Patiriella anewM.S. species of (2002) Description of 1–269. H.E.S.,McKnight, D.G. Clark, Asteroidea (sea-stars) orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117: cushion star Stegnaster inflatus It couldalsobe………. AustraliaIslands. andTasmania. Hasspread tosouthern tosubtidal.EndemicNewIntertidal Zealand, includingtheChatham yellow, olive green,etc.Sometimes mottled. arecrescent-shaped. Colourvery thearms variable,of red,orange, pink, arranged,irregularly marginalplatesnotobvious. Platesalongthecentre or less. Uppersurfacecovered inoverlapping, scale-like plates, platesare Five arms, sometimesfour thearms orsix,discradius ~2/3thelengthof surface (Verrill, 1867) Journal of the Royal Society of New Zealand 32(4): 697 the Royal Society of Journal of habitat (Echinodermata, from New Patiriella Zealand,andreviewregularis (Echinodermata, of –711. inset images: Kate Neill&ChrisWoods environment main image: Pete Notman 6 cm 20 Class Asteroidea Order Valvatida Family Asterinidae

classification Stegnaster inflatus morphology Grace, R.V. Hutton(Class: Asteroidea, Stegnaster inflatus Family: Asterinidae).Tane behaviour of (1974)Feeding 20:163 1–269. H.E.S., McKnight, D.G. Clark, Asteroidea (sea-stars)orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117: ambush starfish It couldalsobe………. eating them.Widespreadthroughout New Zealandinshallow water. invertebrates. Thestarfishthenlowers ontothem,trapping themand itself thesubstrate, off species raises creatingarefuge for itself underneath small toshallowIntertidal subtidal.Known astheambush starasthispredatory mottled. Colour highlyvariable, red,orange, pink,yellow, olive green,etc.Sometimes covered inoverlapping skin-covered plates, marginalplatesnotobvious. Uppersurface Five thearms. arms, discradius about2/3thelengthof Patiriella regularis surface (Hutton, 1872) habitat environment –165. main andinsetimage: Andy Miller 3 cm 21 Class Asteroidea Order Valvatida Family Asterodiscididae

classification truncatus Asterodiscides morphology biogeography thegenus revisited.Zoosystema 31(3):419 of Lane, D.J.W., Rowe, F.W.E. Asterodiscides (2009) A new species of 1–269. H.E.S., McKnight, D.G. Clark, Asteroidea (sea-stars)orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117: fire brick star It couldalsobe………. Australia. andsoutheastern as southern New Islandsaswell ZealandandtheKermadec Subtidal. Found innorthern beige. they purple, areenlarged. Abrightlycolouredmixof orange, redand spaces inbetween. Marginalplatesnotobvious except tipswhere atarm surface covered withlarge andmediumgranules andtubercules, withsome Five arms, discradius approximately Upper thearms. 1/2thelengthof surface Pentagonaster pulchellus habitat (Coleman, 1911) –429. (Echinodermata, Asteroidea, Asterodiscididae) from the tropical southwest Pacific, and the and Pacific, southwest tropical the from Asterodiscididae) Asteroidea, (Echinodermata, environment main image: CrispinMiddleton inset image: Rob Stewart 4 cm 22 Class Asteroidea Order Valvatida Family Odontasteridae

classification Diplodontias dilatatus 1–269. H.E.S.,McKnight, D.G. Clark, Asteroidea (sea-stars) orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117: morphology It couldalsobe………. tosubtidal.FromIntertidal CookStrait southwards totheSnares Islands. orange. granules. Marginalplatestowards tipsareenlarged. arm Dulltobright plates covered ingranules, marginalplatesobvious andalsocovered in Uppersurfacewith Five thearms. arms, discradius ~1/2thelengthof Pentagonaster pulchellus surface (Perrier, 1875) habitat inset image: Roberta D’Archino main image: Rachel Boschen environment 2 cm 23 Class Asteroidea Order Valvatida Family Odontasteridae

classification hyalacanthus Eurygonias Fenwick, G.D.,Fenwick, theSnares islands, New southern Zealand. of Horning, D.S. Jr (1980) Echinodermata 1–269. H.E.S.,McKnight, D.G. Clark, Asteroidea (sea-stars) orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117: 437 morphology –445. It couldalsobe………. Snares Islands. toshallowIntertidal subtidal.Eastcoastfrom CookStrait southwards tothe cream, uppersurfaceisapricottoorange. and slightlyenlarged towards tips. Marginalplatesandpompomsare arm plates bearingrounded, pompom-like structures, marginalplatesobvious Uppersurfacewith Five thearms. arms, discradius ~2/3thelengthof Pentagonaster pulchellus surface Farquhar, 1913 habitat 14(4): Research Marine& Freshwater New ZealandJournal of environment Inset image: Roberta D’Archino main image: MalcolmFrancis 10 cm

24 Class Asteroidea Order Valvatida Family Goniasteridae

classification Pentagonaster pulchellus McKnight, D.G. (excluding Zoology 20(3):191 holothurians) from (1993) Records echinoderms theChatham Islands. New of Zealand Journal of 1–269. H.E.S.,McKnight, D.G. Clark, Asteroidea (sea-stars) orderValvatida. New (2001) The MarineFauna Zealand:Echinodermata: of NIWA Biodiversity Memoir117:

morphology biscuit star It couldalsobe………. around theSouthIsland. and easttotheChathamIslands. Particularly commoninCookStrait and IslandsouthtotheSnaresIslands, North tosubtidal.FromIntertidal northern tips areenlarged. Red, orange orpurple/brown. in hard,naked plates, marginalplatesvery obvious. Marginalplatesatarm UppersurfacecoveredFive thearms. arms, discradius ~2/3thelengthof Diplodontias dilatatus surface Gray, 1840 habitat environment inset image: Rachel Boschen main image: ChrisWoods –200. 4 cm 25 Class Ophiuroidea Order Euryalida Family Euryalidae

classification constrictum Astrobrachion 401–444. Mills, V.S., O’Hara, T.D. New Zealand.Zootaxa 3613(5): of biogenic Ophiuroidea) habitats onthecontinental shelf of (2013) Ophiuroids (Echinodermata: Memoir 115:1–79. McKnight, D.G. New Ophiuroidea: (2000) The Marine Fauna Zealand:Basket-stars Euryalinida). of and snake-stars (Echinodermata: NIWA Biodiversity morphology It couldalsobe………. Australia down to540m. scuba divers inFiordland.Alsoknown from theTasman Sea and south-eastern down to307 m inNew Zealand, and are well-known from being seen by thehostcoral. They of collectedinthemucus aredistributed planktonic particles as coral, such Lives onblack mutualistically Bodyarmspines. covered inskin,smoothtothetouch. solid morphs: colour Several red/brown,dark creamyarms. yellow, and white striped(pictured).Two orblack coiling long five mm, 15 diameter Disc Astroceras elegans Asteroporpa australiensis surface

habitat

(Farquhar, 1900) Antipathella fiordensis, feeding on environment image: Rob Stewart 5 cm 26 Class Ophiuroidea Order Euryalida Family Euryalidae

classification elegans Astroceras 401–444. Mills, V.S., O’Hara, T.D. New Zealand.Zootaxa 3613(5): of biogenic Ophiuroidea) habitats onthecontinental shelf of (2013) Ophiuroids (Echinodermata: Memoir 115:1–79. McKnight, D.G. New Ophiuroidea: (2000) The Marine Fauna Zealand:Basket-stars Euryalinida). of and snake-stars (Echinodermata: NIWA Biodiversity morphology It couldalsobe………. Caledonia. around Australia and between 140 m ineastern 350 and 700 m inNew Zealand, and are alsofound between 300 and 875 m intheTasman Sea, plankton from thewater column. They are distributed down to705 m inNew asplexaurid corals, seafans,Found onbranching such wherethey feedon Bodyarmspines. covered withskinandtubercles, slimy andrough tothetouch. tan and cream banded (see insetimages), both with whitespiny tubercles. two yellow, solid and morphs: colour Two arms. coiling five mm, 10 diameter Disc Astrobrachion constrictum Astrobrachion Asteroporpa australiensis surface

(Bell, 1917) habitat environment main andinsetimages: Rob Stewart 7 cm 27 Class Ophiuroidea Order Euryalida Family Gorgonocephalidae

classification australiensis Asteroporpa 401–444. Mills, V.S., O’Hara, T.D. New Zealand.Zootaxa 3613(5): of biogenic Ophiuroidea) habitats onthecontinental shelf of (2013) Ophiuroids (Echinodermata: Memoir 115:1–79. McKnight, D.G. New Ophiuroidea: (2000) The Marine Fauna Zealand:Basket-stars Euryalinida). of and snake-stars (Echinodermata: NIWA Biodiversity morphology It couldalsobe………. Seamount andarealsofound down Australia. toabout470minsouthern to 350 m atNorfolk IslandandWanganella Bank,about 140 m atGascoyne Newdistributed from Zealand,from about 30 to 510 m innorth-eastern 150 as primnoid and plexaurid corals,Lives sea fans. onbranching such They are coiltightly. arms the touch, Threetoseven withspinyand arms. armspines points. Body hard and rough to across tuberclesand hooks arranged disc.Microscopic rings of inbands on disc Disc diameter up to 10 mm, five arms. Black and white stripes around arms and Astroceras elegans constrictum Astrobrachion surface

H. L. Clark, 1909 H. L.Clark, habitat environment main image: Peter Marriott inset image: SadieMills 1 cm 28 Class Ophiuroidea Order Ophiurida Family Amphiuridae

classification Ophiocentrus novaezelandiae Forening 77:91–177. New Islands. T.fra DanskNaturhistorisk Ophiuroidea. II. Meddelelser Zealand andtheAuckland-Campbell Mortensen, Videnskabelige of (1924) Echinoderms 401–444. Mills, V.S., O’Hara, T.D. New Zealand.Zootaxa3613(5): of biogenic Ophiuroidea) habitatson the continentalshelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. Island andMarlborough Sounds. Found onsoftsandy ormuddy substrates, North predominantlywest coastof no covering plates. Discsoft,tearseasily, butwould feelspiny tothetouch. and ventral discplates. Large Sixtotenarmspines. opententacleporeswith Disc diameter10mm,five brittlespiny Even arms. spinesondorsal cover of Ophiactis resiliens surface habitat Gislén, 1926 environment main andinsetimage: Peter Marriott 2 cm 29 Class Ophiuroidea Order Euryalida Family Ophiactidae

classification Ophiactis resiliens Comparative Zoology,Comparative University Harvard 6:17–83. Lyman, T. (1879) Ophiuridae and Astrophytidae theexploring of voyage H.M.S. Challenger , underProf. of SirWyville Thomson, F.R.S. theMuseumof Partof 2.Bulletin Mills, V.S., O’Hara, T.D. New Zealand. of biogenic Ophiuroidea) habitats onthecontinentalshelf of (2013)Ophiuroids (Echinodermata: morphology It couldalsobe………. Tasman Australia down Seaandsouthern to520m. feet. Found around New Zealanddown toabout210m,butalsointhe withitstube holdfasts, tosuspensionfeed,capturingparticles raises itsarms Well camouflaged, irregular surfacesandalgae livesrock inintersticesof margin, four spinesby Singletentaclescale. Body midarm. ishard. margin, ondorsaldiscinjuveniles. Seven atdisc erect,bluntarmspines radial shields. Numerous atdistaltipof spines ondisc disc, whiteblotches Disc diameterto10mm,five brittlearms. onreddish Variable patterning Ophiocentrus novaezelandiae surface Lyman, 1879 habitat main andinsetimage: Peter Marriott environment Zootaxa 3613(5):401–444. 1 cm 30 Class Ophiuroidea Order Euryalida Family Ophiocomidae

classification bollonsi Clarkcoma Forening 77:91–177. New Islands. T.fra DanskNaturhistorisk Ophiuroidea. II. Meddelelser Zealand andtheAuckland-Campbell Mortensen, Videnskabelige of (1924) Echinoderms 401–444. Mills, V.S., O’Hara, T.D. New Zealand. of biogenic Ophiuroidea) habitats on the continental shelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. to about370mdeep. Alsofound Australia down inSouthern to630m. the ThreeKingsIslandsthroughout New ZealandsouthtoStewart Islanddown two withoutmicroscopic arehardtotellapart examination. Distributedfrom Lives inawiderange habitatssimilarto of Body hard. to oral (mouth)plates. Seven thin,flattenedarmspines. Two tentaclescales. granules ondorsaldisc,nospinesanddiscplatesvisibleventral discclose redcolour.Disc diameterto17mm,five Dark brittlearms. Densecovering of Ophiopteris antipodum surface

(Farquhar, 1908) habitat Ophiopteris antipodum , andthe environment main andinsetimage: AlanHart Zootaxa 3613(5): 2 cm 31 Class Ophiuroidea Order Ophiurida Family Ophiocomidae

classification Ophiopteris antipodum Forening 77:91–177. New Islands. T. II.Ophiuroidea. Zealand and the Auckland-Campbell Mortensen, of (1924) Echinoderms Mills,V.S., O’Hara, T.D. New Zealand. Zootaxa3613(5): 401–444. of biogenic Ophiuroidea) habitatsonthecontinentalshelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. m deep. Island,downIslands intheNorth toStewart Islandinthesouthdown to105 bryozoan beds, feedingonsmallinvertebrates. Distributedfrom theBay of Lives inarange rocky andsandy of habitatsfrom sponge gardens to spine modifiedintoasquareflatscale. Singletentaclescale. Body hard. spines cover ventral discandmargin.Five oar-like armspines, with upper smallgranules blunt covering ondorsaldiscandfirsttwo plates. Short of arm purple/brownDisc diameterto20mm,five Dark brittlearms. colour. Dense Clarkcoma bollonsi surface E. A.Smith,1877 habitat Videnskabelige Meddelelser fra Dansk Naturhistorisk fraDanskNaturhistorisk Meddelelser Videnskabelige environment main image: ChrisWoods inset image: CallumLilley 3 cm 32 Class Ophiuroidea Order Ophiurida Family Ophiodermatidae

classification tarltoni Cryptopelta Baker,Museum, Wellington theDominion Ophiuroidea). brittlestarsfrom Records(1974) New A.N. of New speciesof Zealand(Echinodermata: 401–444. Mills, V.S., O’Hara, T.D. New Zealand.Zootaxa3613(5): of biogenic Ophiuroidea) habitatson the continentalshelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. as CookStrait down to150mdeep. New Northland, Zealand,occasionallyfoundFound asfarsouth mainly off Found well camouflaged inamongstcoralline andrhodolithbeds. algae turf Body hard. margin. Seven Singlelarge armspines, pressedclosetoarm. tentaclescale. disc, except for oral shields, scatteredelongate granules ondorsaldisc smallgranules Densecovering ondorsalandventralcream bandedarms. of Disc diameter12–15mm,five Orange brittlearms. brown disc,dark and surface Baker, 1974 habitat environment inset image: Peter Marriott main image: Rob Stewart , 8:247–266. 2 cm 33 Class Ophiuroidea Order Ophiurida Family Ophiodermatidae

classification Ophiopsammus maculata Vail, L.L.,Rowe, F.W.E. the genera (1989) Status of Mills,V.S., O’Hara, T.D. New Zealand. Zootaxa3613(5): 401–444. of biogenic Ophiuroidea) habitatsonthecontinentalshelf of (2013) Ophiuroids (Echinodermata: Proceedings of the Linnean Society of New SouthWales 110:267–288. the LinneanSociety of species. Proceedings of morphology It couldalsobe………. Island. stars, they are agilepredators. Distributedfrom IslandstoStewart Bay of thelargestoccasionally befound rock pools. brittle Oneof indeepintertidal Lives onrocky shoresandbryozoan beds, commonsubtidallybutcan tentacle scales. Body hard. ventral discplates. Ten Two pressedclosetoarm. bluntarmspines to11short, entire dorsalsurfaceandventral disc.Rounded granules cover dorsaland discdiameter.size of crimsonwithdistinctive Dark smallredspotsover Large, common,discdiameterupto48mm,five upto snakyarms four times surface Ophiopeza and (Verrill, 1869) Ophiopsammus habitat (Echinodermata: Ophiuroidea) in Australian waters,(Echinodermata: a new withthe description of environment inset image: Rachel Boschen main image: ChrisWoods 6 cm 34 Class Ophiuroidea Order Ophiurida Family Ophiomyxidae

classification Ophiomyxa brevirima Harvard UniversityHarvard 25:165–376. Zoology, Comparative Comparative the Museum of Zoology. recent ophiurans: based on the collection of the Museum H.L. of (1915) Catalogue of Clark, Memoirs of Mills,V.S., O’Hara, T.D. New Zealand. Zootaxa3613(5): 401–444. of biogenic Ophiuroidea) habitatsonthecontinentalshelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. New Zealanddown toabout1100m. Found onsandy substrates inkelp holdfastsandinsponge gardens around glassy withserrated edge. Body issoft. 3–4persegment.Oral papillae alternating captured.Armspines easily if yellowish brown Thinskinondiscandarms, tears colourwithbandedarms. Disc diameter8–35mm,five brittlearms. Variable greenish,reddishor surface H. L. Clark, 1915 H. L.Clark, habitat environment main image: ChrisWoods inset image: CallumLilley 7 cm 35 Class Ophiuroidea Order Ophiurida Family Ophionereididae

classification Ophionereis fasciata 17 p. Hutton, F.W. (1872) Mills,V.S., O’Hara, T.D. New Zealand. Zootaxa3613(5): 401–444. of biogenic Ophiuroidea) habitatsonthecontinentalshelf of (2013) Ophiuroids (Echinodermata: morphology It couldalsobe………. about 300mdeep. with itstubefeet.Distributedonopenshoresaround New Zealanddown to thistothemouth through screensbetween andtransports mucus thearmspines deeper subtidalhabitats. by Feeds filteringcoarsesuspendeddetritus stonesintheshallowFound underneath subtidalrocky shoresandoutto bluntarmspines, singleoval Four short arms. tentaclescale. Body ishard. onlightergrey/brown. black of whiteandgrey-bandedblotches Black, with Disc diameterto16mm,fiverandom Discpatterned brittlearms. Catalogue of the Echinodermata of New Zealand, with diagnoses of thespecies,Colonial Museumand Geological Survey Wellington, Department, New Zealand,with diagnosesof of theEchinodermata of Catalogue Ophiactis resiliens surface Hutton, 1872 habitat environment inset image: CallumLilley main image: SadieMills 3 cm 36 Class Crinoidea Order Comatulida Family Comasteridae

classification Cenolia novaezealandiae morphology delineated by sequence andmicrosatellite markers. Molecular Phylogeneticsand Evolution 78:160–171. Naughton, K.M., O’Hara, T.D. Appleton, B., Gardner, M.G. Crinoidea: Comasteridae (2014) Sympatric cryptic species in thecrinoidgenus Cenolia(Echinodermata: McKnight, D.G. (1977)Additions totheNew Zealandcrinoid fauna. It couldalsobe………. Island, andwidespreadaround Australia andthesouthwest Pacific distancesby theNorth short of toswim. Found inthe north usingtheirarms This speciestypicallyclingstorock surfacesorotherorganisms andcanmove pinnules. (usually >20)anduniform colouredarm segments. Oxycomanthus plectrophorum isasimilarcolourbuthasmorearms comprising12–21individual theanimaltosubstrate), each to anchor yellowish pinnules. Thereare17–40cirri(appendages tipstothearm used food-gathering device. They reddish-purple, aremainlydark withdistinctive segmentedpinnules, themain branching segmented spinebearingdozensof starwithusually 10–20feather-like acentralFeather composedof arms habitat Oxycomanthus plectrophorum environment (A.H. Clark, 1918) (A.H. Clark, New ZealandOceanographic Records 3(11):93–112. main andinsetimage: CrispinMiddleton 7 cm 37 Class Crinoidea Order Comatulida Family Comasteridae

classification Oxycomanthus benhami morphology delineated by sequence andmicrosatellite markers. Molecular Phylogeneticsand Evolution 78:160–171. Naughton, K.M., O’Hara, T.D. Appleton, B., Gardner, M.G. Crinoidea: Comasteridae (2014) Sympatric cryptic species in thecrinoidgenus Cenolia(Echinodermata: McKnight, D.G. (1977)Additions totheNew Zealandcrinoid fauna. It couldalsobe………. elsewhere areusuallydeeperthan30m. the southwest Pacific. InFiordlandthey arecommonasshallow as8m,but Found allaround New Zealandandwidespreadaround Australia and fewer (usually<20)andhasyellowish arms pinnules. tipstothearm by isasimilarcolourbuthas toswim. Cenoliaspanoschistum usingtheirarms surfaces orotherorganisms, e.g. corals, black andcanmove distances short burgundy, chocolate-purple, deepbrown, They orblack. clingtorock comprising30–37 segments. Colourmainly animal tothesubstrate), each gathering the device. Thereare33–73cirri(appendages usedtoanchor segmentedpinnules, themainfood-segmented spinebearingdozensof starwithusually 31–44feather-like acentralFeather composedof arms habitat Cenolia spanoschistum environment (A.H. Clark, 1916) (A.H. Clark, New ZealandOceanographic Records 3(11):93–112. image: MalcolmFrancis 7 cm 38 Class Echinoidea Order Camarodonta Family Echinometridae

classification Evechinus chloroticus Freshwater Research 6(1–2): 48–68. Research Freshwater Dix, T.G. from different localities4.Age, Echinometridae) (Echinoidia: growth, chloroticus Marine and Evechinus (1972) Biology of and size. New ZealandJournal of 4(2):91–116. Research Marineand Freshwater T.G.Dix, from1. General. differentlocalities New ZealandJournal Echinometridae) of (Echinoidea: chloroticus Evechinus (1970a) Biology of kina morphology It couldalsobe………. recreational fisheries. barerock. Kinaareedible, significantcommercialand areas of supporting reefs. They oftenform large aggregations, algae toform strippingstandsof Kina arefound andunderledges nestledincracks orfullyexposed onopen areas. in southern sharp-tipped teeth.Canbevery large (upto180mmdiameter)especially but thetestgreenwhencleaned.Mouthcentral onthelower sidewithfive, the larger primaryspines, white. Tube feetandtheskinlayer reddish-brown, thesmallersecondaryspines,green spines. andoccasionally Spinetipsof flattened,withsolidsharp Round toslightlysub-pentagonal seaurchin, Centrostephanus rodgersii novaezelandiaePseudechinus surface (Valenciennes, 1846) habitat environment inset image: MalcolmFrancis main image: ChrisWoods 3 cm 39 Class Echinoidea Order Camarodonta Family Temnopleuridae

classification Pseudechinus albocinctus Fell, H.B. (1952) Echinoderms from New Southern Zealand. H.B.Fell, (1952) Echinoderms Oceanographic Institute Memoir47:1–91. McKnight, D.G. fauna. Benthos survey, theNew New Zealand shelf Zealand (1979) Anoutlinedistributionof theEchinoidea. station list,and distribution of It couldalsobe………. substrate,buried incoarseshell-rubble oronrocky reefs. Particularly abundantintheMarlborough Sounds, andoftenfound semi- apparatusFeeding (central onthelower side)withfive, sharp-tippedteeth. pattern. the pore-zones(tubefeet)producing asometimesdistinctive radial stripe test ispurplish-pinktogreyish-brown; thetuberclesalightershade, asare thetestmay beslightlyflattened.Thebare tips; thoseatthewidestpointof 10–12 mmlong, andgenerally red-brown orpurplishwithdistinctive white Ball-shaped, slightlyflatteneddorso-ventrally. andsolid,upto Spinesshort morphology Pseudechinus novaezealandiaePseudechinus huttoni Pseudechinus surface Zoology Publications of VictoriaUniversity 18:1–37. of Zoology Publications (Hutton, 1872) habitat environment image: Owen Anderson 1 cm 40 Class Echinoidea Order Camarodonta Family Temnopleuridae

classification Pseudechinus huttoni Fell, H.B. (1952) Echinoderms from New Southern Zealand. H.B.Fell, (1952) Echinoderms Oceanographic Institute Memoir47:1–91. McKnight, D.G. fauna. Benthos survey, theNew New Zealand shelf Zealand (1979) Anoutlinedistributionof theEchinoidea. station list,and distribution of It couldalsobe………. algae, andonrocky reefs. driftorattached a shelly-sandsubstrate wherethereisanabundanceof regions,common inFiordlandandothersouthern they areoftenfound on Mouth central onthelower sidewithfive, sharp-tippedteeth.Especially greenish insomeoradarker colouraround thebase. and spineswhitetopinkinlife, straw colouredwhendried;spines may be andsolid,toabout10–12mm.Test thetest.Spinesshort of the widestpart on theuppersurface. Onlarger specimenstuberclesform distinctive rows on Ball-shaped, varying inheightfrom moderately flattenedtoslightlyconical morphology Pseudechinus novaezelandiaePseudechinus albocinctus Pseudechinus surface Benham, 1908 Zoology Publications of VictoriaUniversity 18:1–37. of Zoology Publications habitat environment inset image: Owen Anderson image: MalcolmFrancis 4 cm 41 Class Echinoidea Order Camarodonta Family Temnopleuridae

classification Pseudechinus novaezealandiae Fell, H.B. (1952) Echinoderms from New Southern Zealand. H.B.Fell, (1952) Echinoderms Oceanographic Institute Memoir47:1–91. McKnight, D.G. fauna. Benthos survey, theNew New Zealand shelf Zealand (1979) Anoutlinedistributionof theEchinoidea. station list,and distribution of It couldalsobe………. least –innarrow crevicesbetween rocks oramongstrubble. They arefound mainlyonhardsubstrates, oftenhidden– inthedaytime at around theSouthIslandaswell asStewart, CampbellandMacquarieIslands. about50mmtestdiameter).Known of fromThis isasmallspecies(maximum apparatusFeeding (central onthelower side)withfive, sharp-tippedteeth. tubercles andaround thepores. may bewhiteorpurple-tipped.Thetestisadullgrey-green, paleronthe adense, forming short, coat;they uniform aregenerally greenish-grey, but Ball-shaped, moderately flatteneddorso-ventrally. Spinessolid and very morphology Pseudechinus albocinctus Pseudechinus huttoni Pseudechinus chloroticus Evechinus surface Zoology Publications of VictoriaUniversity 18:1–37. of Zoology Publications habitat (Mortensen, 1921) (Mortensen, environment image: Owen Anderson 2 cm 42 Class Echinoidea Order Clypeasteroida Family Clypeasteridae

classification Fellaster zelandiae Rowe, F.W.E., andGates, In Wells, Australia33:i–xiii, 1–510.CSIRO, J. (1995) Echinodermata. Melbourne. of A.(ed.). Zoological Catalogue Oceanographic Institute Memoir47:1–91. McKnight, D.G. fauna. Benthos survey, theNew New Zealand shelf Zealand (1979) Anoutlinedistributionof theEchinoidea. station list,and distribution of sand dollar morphology It couldalsobe………. AustraliaNew includingTasmania. Zealand and alsotheeastcoastof buried very shallow of inthesand,oftenhighdensities. Found around much Sand dollarsarehighlymobiledepositfeedersthat live onthesandorare the uppersurface, thelower themouthincentre of side, teethinternal. outwards from thecentreonbothsurfaces. Theanusisattheouteredge of Petal-shaped theuppersurfaceandfive regionsadorn food grooves radiate across thetestvisibleastenradiating stripes. Thebaretest isgreyish-white. light brown orgreyish-purple, spines. withvery Spinedifferentiation short Spherical, extremely flattenedseaeggwithasinuousperimeter; to dark surface (Gray, 1855) habitat environment main andinsetimage: Owen Anderson 3 cm 43 Class Echinoidea Order Diadematoida Family Diadematidae

classification rodgersii Centrostephanus Oceanographic InstituteMemoir 47:1–91. McKnight, D.G. fauna. Benthossurvey, theNew New Zealand shelf Zealand (1979) Anoutlinedistribution of theEchinoidea. station list,anddistribution of Miskelly, A.(2002) Rodgers sea urchin It couldalsobe………. Australia includingTasmania,coast of New Caledonia,andLordHowe Island. Island,theeast theNorth of come outtofeedatnight.Found inthenorth They areusuallyfound underbouldersandincrevicesduringdaylight, and tipped teeth.Canbequitelarge (toabout120mmdiameter). green ontheuppersurface. Mouthcentral onthelower sidewithfive, sharp- rough, serrated outersurface. Cleanedtestpale, purpleand withtraces of andslightlyiridescentgreenincolourwitha individuals; purplish-black inlargerabout equaltotestdiameterinsmallindividualsbutshorter quiteflattened.Spineshollow,Round seaurchin, brittle, andsharp;length morphology of Sea urchins Evechinus chloroticus Evechinus AustraliaandtheIndo-Pacific Publications,. Capricornia Sydney, Australia, 180p. surface habitat (A. Agassiz, 1863) environment image: MalcolmFrancis 2 cm 44 ` 45 2 cm image: Owen Anderson Owen image: environment Zoology Publications of 18: 1–37. Victoria University habitat (Milne-Edwards, 1836) (Milne-Edwards, surface Echinocardium cordatum It could also be ………. This sea urchin has an oval body shape which This sea urchin body is wider at the posterior end, has an oval The mouth (toothless in rounded on the upper side. and flat on the underside, The spines are of side. the centre of the lower adults) is located just forward In life the test and side. solid, short on the upper side and long on the lower the dead test is greyish-white. spines are reddish-brown, shelly sea bed, about 30 mm deep, buried typically in coarse, live They amounts of large out sediment and filtering ingesting feed by where they nutritious particles.

morphology

Fell, H.B. (1952) EchinodermsFell, H.B. Zealand. Southern New from Institute Oceanographic Zealand New EchinozoanThe (1962) Rise. Chatham the Island, and Macquarie of D.L. islands, Subantarctic Zealand Fauna New the Pawson, Memoir 42: 1–35.

Apatopygus recens recens Apatopygus classification Class Echinoidea Superorder Neognathostomata Family Apatopygidae Family Neognathostomata Superorder Echinoidea Class Class Echinoidea Order Spatangoida Family Loveniidae

classification Echinocardium cordatum Fell, H.B. (1958) Deep-Sea echinoderms of New Zealand. of H.B.Fell, (1958) Deep-Seaechinoderms Oceanographic Institute Memoir47:1–91. McKnight, D.G. fauna.Benthossurvey, theNew New Zealandshelf Zealand (1979)Anoutline distributionof theEchinoidea. stationlist,anddistributionof heart urchin morphology It couldalsobe………. Atlantic, andMediterranean, butdiscontinuously. Notfound intropical seas. Widespread around New andsouthPacific, ZealandandAustralia, north the through thesubstrate. from thatpassesthroughparticles themud theirdigestive tract when moving create arespiratory shaft tothesurface, andfeedby extracting minutefood orsand,theycentimetres deepinsoftmud usespecialisedtubefeetto golden togreenishgrey, flatagainst brushed thetest.Livinguptoseveral in adeepgroove continuesdown tothemouth.Spinesaglistening which The tubefeetform petalsontheuppersurface, theanteriorpetallying narrow end,themouth(toothless)onlower surfacenearthebluntend. Test convex heart-shaped, onuppersurface, flatterbelow. The anusatthe Apatopygus recens Apatopygus surface

habitat Zoology Publications from Victoria University of Wellington 24:1–40. from Victoria UniversityZoology Publications of (Pennant, 1777) environment inset image: Spencer, Mark AUSCAPE image: Owen Anderson 2 cm 46 Class Holothuroidea Order Aspidochirotida Family Holothuriidae

classification Australostichopus mollis Deuterostomia. Canterbury University Press, 371–400 pp. stars, seacucumbers, sea urchins, sealilies. InGordon, D.P. (Eds.), Mah, C.L., McKnight,D.G., Eagle, M.K.,Pawson, D.L., Ameziane, N.,Vance, D. J., Baker, H.E.S., Davey,Clark, A.N., seastars, brittle N.(2009) Phylum Echinodermata: 24(1):97–103. Research Freshwater mollis Stichopus Sewell, theecology of (1990) Aspects of M.A. morphology It couldalsobe………. New SouthWales coast around tosouthwestern Australia. Fiordland, Stewart IslandandtheMarlborough Sounds. Alsofound around bottoms. Itisdistributedallaround New Zealandwithhigherdensitiesin It lives in many habitatsfrom shallow rock poolstosubtidalsandy mud faecal coilemergingbehind. shield-shaped tentaclesshoveling mud sedimentintoitsmouthandathick tube feet,andasquishy texture. Itisregularyseenwithforward extended cm longandhasawarty dorsalsurface withnumerous radial rows ventral of bycharacterised itsmottledlightbrown body. orblack upto30 Itcanreach mollis Australostichopus Holothuria integra isNew Zealand’s mostconspicuousseacucumber surface (Hutton, 1872) New Zealand Journal of Marine & New‐eastern Holothuroidea) Zealand,New ZealandJournal innorth of (Echinodermata: New Zealand Inventory of Biodiversity, of New ZealandInventory volume Lophotrochozoa, Radiata, 1,Kingdom Animalia: habitat main andinsetimages: ChrisWoods environment 3 cm 47 Class Holothuroidea Order Aspidochirotida Family Holothuriidae

classification Holothuria integra Rowe, F.W.E., Gates, InWells,33: i–xiii,1–510.CSIRO, Australia J. (1995)Echinodermata. Melbourne. of A.(ed.). ZoologicalCatalogue morphology It couldalsobe………. from SouthAfrica andthePhilippines. New Zealandandalsoknown of part known tobedistributedinthenorthern Holothuria integra lives oncoarseshellandsand oramongstcobbles. Itis wall andpelate tentacles. and tubefeetventrally. yet leatherybody hasathickish Itissofttotouch indicate itextends to30cmlengthandhassmallpapillaeorbumpsdorsally itslifehistory.Little isknown of Specimenscollectedorphotographed Holothuria integra isgreyish-brown withwhitespots, and issausage-shaped; Australostichopus mollis Australostichopus Koehler &Vaney (1908) surface habitat main andinsetimage: NIWA environment 7 cm 48 Class Holothuroidea Order Dendrochirotida Family Cucumariidae

classification Amphicyclus thomsoni Pawson, Holothuroidea). Bulletin D.L. New (1970) The MarineFauna Zealand: Sea Cucumbers (Echinodermata: of 201:1–69. Research morphology It couldalsobe………. Island’s EastCapesouthtoStewart Island. depth butcanbeseenatdiveable depthsandisdistributedfrom theNorth debris. Itisalsofound underandbetween boulders. Itlives down to180m It lives crypticallyincoarsemuddy with shellysandwhereitcovers itself andcontractsconcentric rings. easily. Thisspeciesissquishy totouch numerous tubefeetradially and25dendritictentaclesarranged intwo upto7.5cminlengthandisyellowish-brown.wrinkles. Itreaches Ithas thomsonihasapentagon-shapedbodyAmphicyclus withnumerous transverse surface (Hutton, 1878) habitat New Zealand environment Department of

Scientific and image: NikiDavey 1.5 cm Industrial 49 Class Holothuroidea Order Dendrochirotida Family Cucumaridae

classification Squamocnus brevidentis O’Loughlin, P.M., MuseumVictoria 58(1):1–24. Holothuroidea). Memoirs N.K.(2000) TheNew of Alcock, ZealandCucumariidae (Echinodermata, morphology densities upto800perm This speciesisfound covering rock walls andplatforms inFiordland, surface soattimestiny canbefound a specimen. pinkspecks underneath into thecurrentswhenfeeding. Thefemales brood theiryoung ontheventral bumpy integumentandextensive areextended dendritictentacleswhich 5.5cm.Ithasatough lengthsof white bumps(tubefeetorpapillae)reaching Squamocnus brevidentis isastunningdeepredholothuriancovered insmall It couldalsobe………. rocks, inotherlocations. increvicesandamongstrubble 2 . Alternatvely itcanbefound. Alternatvely crypticallyunder surface (Hutton, 1872) habitat environment main image: Jason Moore inset image: NikiDavey 1.5 cm 50 Class Holothuroidea Order Dendrochirotida Family Heterothyonidae

classification alba Heterothyone Pawson, D.L. CookStrait, New (1963) HolothurianFauna of Zealand. morphology It couldalsobe………. round theentire New Zealandcoastandknown tobeeurybathic. It livesrubble. inasandy bottom,sometimesamongstfine Itis mud found after collection. like, tubefeet.Ittendstoholditsshape withnumerous andfirm stiff thick and theposteriorendtaperstoform aslendertail.Thebody wall isscale- retracted upto2.5cm.Thetentaclesarenormally and hasatotallengthof Heterothyone albaisavery commongrey towhiteholothurian.ItisU-shaped (Hutton, 1872) surface Zoology Publications from Victoria University of Wellington 36:1–38. from VictoriaUniversityZoology Publications of habitat 0.3 cm environment image: NIWA 51 Class Holothuroidea Order Molpadida Family Caudinidae

classification Paracaudina chilensis Memoirs of Museum Victoria 68: 37–65. Caudinidae). Memoirs of O’Loughlin, P.M.,Barmos, S., Van denSpiegel, Holothuroidea: D. Molpadida: Australia and New (2011) The paracaudinid sea cucumbers of Zealand (Echinodermata: 357–368. Davey,P.M. N.K.,O’Loughlin, Holothuroidea: Molpadida: Caudinidae). Zootaxa3613(4): New Zealand (Echinodermata: (2013) The caudinid sea cucumbers of morphology It couldalsobe………. widespread. Australia and further feeding andrespiration. They aredistributedthroughout New Zealand, It lives buriedinthesoftsedimentwithitsanusandtentacles exposed for integument. tothefirm attached and hasnotubefeet.Colouration ispalecreamtobrown, oftenwithgrit upto 15 cm.Thebodyand elongate wall tailreaching surfaceiswrinkled cast ontotheshorefollowing event. astorm Ithasadistinctbulbousbody Paracaudina chilensis is acommonholothurianthatfound in sediment or surface (Müller, 1850) habitat environment image: Peter Marriott 2.5 cm 52 about this guide | about echinoderms | morphology index | species index | species pages | icons | glossary icons

sea star asteroid feather star crinoid

brittle star ophiuroid sea cucumber holothurian

snake star ophiuroid with coiling arms sea egg echinoid body plan body

ophiuroid with branching basket star arms

naturally occuring around naturally occuring around native southwest pacific New Zealand, Australia New Zealand, endemic and other pacific locations

species first described since first described in from outside of New range NZ, this species has been introduced Zealand waters and is extention recorded elsewhere found in New Zealand and other locations, invasive life history naturally occuring around antipodean New Zealand and widespread species recorded globally Australia only

gherkin-shaped sea spherical, globular or ball gherkin cucumber with sharp or semi-spherical blunt protrusions

circular, distinctively sausage-shaped sea disk tail flattened, biscuit-shaped cucumber with thinner tail

coiling vertically, can wrap heart heart-shaped, cordate snake arms coral branches

intricate feeding can bend horizontally, no aristotle’s lantern brittle arms mechanism vertical coiling

an obvious series of flexible, stalked tentacles, margins marginal plates which may tube feet sometimes with suckers, or may not bear spines protruding from body wall morphology shield-shaped tentacles > 5 arms, may be 5+ arms pelate tentacles around sea cucumber between 6 and 12 mouth

5 long or short arms, branching, tree-like 5 arms sometimes 4 or 6 if dendritic tentacles tentacles around sea damaged cucumber mouth

finger-like tentacles very brittle, feather-like crinoid arms digitate tentacles around sea cucumber arms in multiples of 5 mouth

bearing small flattened even, hairless, silky, can be smooth warty bumps or tubercles, slightly undulating verrucose

irregularly pitted and surface covered in small to rough ridged surface, often granular medium sized granules tough

soft to the touch, easily hard to the touch, not soft hard compressible, elastic compressible, rigid

bearing irregularly spined surface covered in spines deeply wrinkled parallel ribs and grooves surface along the body wall

thick skin, tough, flexible, spikey surface covered in spikes leathery slightly elastic

bony units layered on the plates outer body wall

hard substrate such as very fine muddy and silty mudstone, sandstone, mud sediments derived from rock basalt, compressed terrigenous rocks, soils and carbonates clays

living or growing on shell, stone, and pebble the external surface of rubble epizoic/epiphytic rubble an animal (epizoic) or seaweed, (epiphytic)

habitat anything man-made such small coarse grains of sand artificial substratum as mooring blocks, mussel worn silica, rock, and shell lines, wharf piles

indentation in rock filled coralline algae, seagrass rockpool algal beds with water, intertidal or algal beds

exposed shoreline zone sand and rubble spread between high and low over underlying hard tides, including rock flats, substrate, organisms pools, overhangs, crevices, attached to basement rock intertidal covered rock organisms exposed to susceptible to inundation wave action, temperature and scouring from wave extremes, full illumination, surge and currents, and and desiccation subdued illumination

composed of a variety of sedimentary substrates zone below the low tide, including coarse gravels, including rock flats, slopes, shell hash and sands to walls, crevices, overhangs, finer sand, mud, and silts, subtidal boulder fields, organisms seabed organisms susceptible to exposed to wave surge inundation and scouring and currents, and subdued from wave surge and illumination currents, and subdued illumination

seabed raised into a bank of compacted rubbles and environment underwater cliffs and other carbonate materials slopes, organisms exposed including shell, kina and wall to wave surge and bank sealace hash, organisms currents, and subdued exposed to wave surge illumination and currents, and subdued illumination

underwater caves, shelves and overhangs, organisms may experience indents wave surge, subdued illumination, or near darkness

55 about this guide | about echinoderms | morphology index | species index | species pages | icons | glossary glossary algal beds areas of seafloor with coralline algae, sea-grass or multiple seaweed species amorphous without definable shape, often with lobed surface, potato or tuber-shaped, massive anterior towards the front aristotle’s lantern the intricate feeding apparatus (sometimes called ‘jaws’) unique to echinoids (sea urchins, sea eggs) artificial substratum anything man-made such as mooring blocks, mussel lines, wharf piles asteroid scientific name for a sea star or starfish antipodean naturally occurring in New Zealand and Australia, and may include seamounts and ridges to the north ball spherical, globular or semi-spherical banded stripes of two or more different colours bank seabed raised into a bank of compacted rubble and other carbonate materials including shell, kina and sea lace hash, organisms exposed to wave surge and currents, and subdued illumination blunt not sharp, rounded ends brittle fragile but rigid, breaks apart easily basket star popular name for a ophiuroid with branching arms brittle arms can bend horizontally, no vertical coiling, in brittle stars brittle star popular name for a ophiuroid bursal slit opening to the pouch or sac from whence juvenile ophiuroids or eggs are released calyx cup cirri appendages used to anchor crinoids (feather stars and sea lilies) to the seafloor cordate heart-shaped covered rock sand and rubble spread over underlying hard substrate, organisms attached to basement rock susceptible to inundation and scouring from wave surge and currents, and subdued illumination crinoid the scientific name for sea lilies and feather stars crinoid arms very brittle, feather like arms in multiples of 5 dendritic tentacles branching, tree-like tentacles around sea cucumber mouth deposit feeder an animal that feeds on particles of organic matter present in surface sediments diameter the distance across the widest point of a circle digitate tentacles finger-like tentacles around sea cucumber mouth disc circular, distinctively flattened, biscuit-shaped dorsal upper surface of the animal echinoid the scientific name for a sea urchin or sea egg endemic naturally occurring in New Zealand, but not elsewhere environment physical, chemical, ecological, behavioural, and other conditions experienced by an organism epiphytic living or growing on the external surface of a plant epizoic living or growing on the external surface of an animal eurybathic can live or be found at many depths feather star popular name for a form of crinoid lacking a stalk firm requires some pressure to compress fleshy feels like skin or edam cheese, dense food grooves channels in the test of sand dollars used for transporting food particles to the mouth gonad reproductive structure granular surface covered in small to medium sized rounded or square granules, giving a sand-papery texture due to calcareous or siliceous minerals in or on (echinoderms) the surface of the organism habitat the environment and local situation in which an organism lives hard solid to the touch, not compressible, rigid holothuroid the scientific name for a sea cucumber, holothurian indents underwater caves, shelves and overhangs, organisms that live there may experience wave surge, subdued illumination, or near darkness integument outer body wall or skin interstices the gaps and spaces between things e.g., rocks, sand-grains or seaweed holdfasts intertidal exposed shoreline zone between high and low tides, including rock flats, pools, overhangs, crevices, organisms that live there are exposed to wave action, temperature extremes, full illumination, and desiccation introduced species first described from outside of New Zealand waters and is found in New Zealand and other locations, invasive, adventive lateral side of an animal leathery thick, tough, flexible, slightly elastic margins an obvious series of marginal plates which may or may not bear spines morphology form and structure, shape mottled variable, blotchy, patterning of several colours

56 about this guide | about echinoderms | morphology index | species index | species pages | icons | glossary mud very fine silty sediments derived from terrigenous rocks, soils and clays naked surface unadorned by spines or granules, usually smooth native naturally occurring in New Zealand, but may also occur naturally elsewhere, endemic ophiuroid the scientific name for a brittle star, basket star or snake star oral related to the mouth of an animal ossicle a small mineral (calcium carbonate) element embedded in the body wall of an echinoderm papillae specialised dorsal tube feet lacking a suckered tip (holothurians), small bony scales that are attached to the jaw, mouth, disc, being free at one end (ophiuroids) peltate tentacles shield-shaped tentacles around sea cucumber mouth peristome parts surrounding the mouth of various invertebrates such as the echinoderms petals leaf-shaped concentrations of ambulacral pores on the tests of sand dollars and heart urchins pinnules small, segmented, food gathering appendages which give crinoids their feather-like appearance plastron in spatangoid echinoids, an enlarged area behind the mouth bearing locomotory spines plates bony units layered on the outer body wall of an echinoderm pom pom disc adornment resembling a fluffy ball posterior towards the rear of the organism primary spines long, large diameter sea urchin spines; may be a different colour to the secondary spines radius distance between the edge and centre of a circle range extension since first described in New Zealand, this species has been recorded elsewhere refuge safe place to hide from predators rock hard substrate such as mudstone, sandstone, basalt, compressed carbonates rockpool indentation in rock, filled with water, intertidal zone rosette the arrangement of small spines or plates in a radiating circle pattern rough irregularly pitted and ridged surface, often tough rubble shell, stone, and pebble rubble sand small coarse grains of worn silica, rock, and shell sea lily popular name for a stalked crinoid sea star popular name for an asteroid sea cucumber popular name for a holothurian, holothoroid sea urchin popular name for a echinoid sea egg popular name for a echinoid seabed composed of a variety of sedimentary substrates including coarse gravels, shell hash and sands to finer sand, mud, and silts, organisms susceptible to inundation and scouring from wave surge and currents, and subdued illumination secondary spines short, small diameter sea urchin spines; may be a different colour to the primary spines sinuous wavy pattern smooth even, hairless, silky, can be slightly undulating snake star popular name for an ophiuroid with coiled arms snake arms arms coiling vertically, can wrap around coral branches soft easily compressible, elastic starfish popular name for an asteroid spined surface covered with spines (echinoderms), or prickly bundles of very long spicules projecting from surface of the organism (sponges) subtidal zone below the low tide, including rock flats, slopes, walls, crevices, overhangs, boulder fields, organisms exposed to wave surge and currents, and subdued illumination surface patterning or ornamentation on the surface of the body of an animal tail sausage-shaped sea cucumber with thinner tail test calcium carbonate skeleton of a sea urchin, composed of twenty columns of individual plates translucent lets light through body wall or surface of organism, but not enough to perceive distinct details through it. transverse across the short axis of the body wall tube feet flexible, fluid-filled, stalked tentacles, sometimes with suckers, protruding from body wall tubercles hard, sometimes warty, calcified lumps that sit embedded in or on the body surface, in sea urchins they incorporate the basal articulation of the spines ventral lower surface or underside of the animal that sits on the seabed wall underwater cliffs and slopes, organisms exposed to wave surge and currents, and subdued illumination warty bearing small flattened bumps or tubercles widespread species recorded globally

Many of the specimens examined to produce this guide came from the NIWA Invertebrate Collection (NIC) and were collected under the following research programs: • Ocean Survey 20/20 Bay of Islands coastal biodiversity, sediment and seabed habitat project (voyages KAH0907 & TAN0906), funded by Land Information New Zealand (LINZ) • Ocean Survey 20/20 Chatham/Challenger biodiversity and seabed habitat project, jointly funded by the former New Zealand Ministry of Fisheries, LINZ, National Institute of Water & Atmospheric Research (NIWA), and Department of Conservation (DOC) • Biogenic habitats on the continental shelf project (voyages TAN1105 & TAN1108), funded by New Zealand Ministry for Primary Industries (MPI (Fisheries) (Biogenic habitats Project ZBD200801), New Zealand Foundation for Research, Science and Technology (CCM contract CO1X0907), NIWA Capability Fund (Project CF111358) and Oceans Survey 20/20 R/V Tangaroa days funded by LINZ • Rhodolith project funded by New Zealand MPI (Fisheries) (Project ZBD200903); Marine High Risk Site Surveillance programme (Project SOW12099) funded by MPI The preparation of this guide was funded by NIWA under Coasts and Oceans Research Programme 2 Marine Biological Resources: Discovery and definition of the marine biota of New Zealand (2013/2014 SCI).

Thanks to the many and talented photographers whose work was used with permission in our guide, including: Rachel Boschen, Victoria University Wellington & NIWA; Callum Lilley, DOC; Jason Moore, Wellington; Patrick Colin, Coral Reef Research Foundation, Palau, Micronesia; Crispin Middleton (including cover image of Diadema palmeri and introduction background images), Chris Woods, Malcolm Francis, Peter Marriott, Rob Stewart, Drew Lohrer, Roberta D’Archino, Pete Notman, Andy Miller, Sadie Mills, Owen Anderson, Niki Davey, Kate Neill, all from NIWA New Zealand. Line drawing in the introductory sections Heavenly Holothurians and Outstanding Ophiuroids are reproduced with permission from Charles Messing in Hendler et al. (1995). further reading

General Mah, C.L., McKnight, D.G., Eagle, M.K., Pawson, D.L., Ameziane, N., Vance, D. J., Baker, A.N., Clark, H.E.S., Davey, N. (2009) Phylum Echinodermata: sea stars, brittle stars, sea urchins, sea cucumbers, sea lilies. In Gordon, D.P. (Ed.), New Zealand Inventory of Biodiversity, Volume 1, Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia. Canterbury University Press, 371–400 pp. Hendler, G.L., Kier, M.P., Miller, J.E., Pawson, D.L. (1995) Sea stars, sea urchins and allies. Smithsonian Institution Press, Washington, 392 p. Miskelly, A. (2009) Sea urchins of the World: diversity, symmetry, and design [Ashley Miskelly, 2009, Blackheath, N.S.W. ISBN: 9780646518961 (hbk)], 103 p. Paine, R.T. (1971) A short-term experimental investigation of resource partitioning in a New Zealand rocky intertidal habitat. Ecology 52: 1096–1106. Waters, J.M., Roy, M.S. (2003) Marine biogeography of southern Australia: phylogeographical structure in a temperate sea-star. Journal of Biogeography 30: 1787–1796. Willan, R.C. (1981) Soft-bottom assemblages of Paterson Inlet, Stewart Island. New Zealand Journal of Zoology 8(2): 229–248. Wing, S. (2008) Subtidal invertebrates of New Zealand: a divers’ guide. Canterbury University Press, 148 p.

Ophiuroids Bell, F.J. (1917) Echinoderma. Part 1. Actinogonidiata. British Antarctic ‘Terra Nova’ Expedition 1910. Zoology 4: 1–10. Clark, H.L. (1909) Scientific results of the trawling expedition of H.M.C.S. Thetis. Echinodermata. Memoirs of the Australian Museum 4: 519–564. Farquhar, H. (1900) On a new species of Ophiuroidea. Transactions of the New Zealand Institute 32: 405–406. Gislén, T. (1926) On the generic types of the ophiuroid genus Ophiocentrus Ljungman (Amphiocnida Verrill). Göteborgs Kungliga Vetenskaps och Vitterhets Samhälles Handlingar 30: 3–16. Lyman, T. (1879) Ophiuridae and Astrophytidae of the exploring voyage of H.M.S. Challenger, under Prof. Sir Wyville Thomson, F.R.S. Part 2. Bulletin of the Museum of Comparative Zoology, Harvard University 6: 17–83.

Mortensen, T. (1925) Echinoderms from New Zealand and the Auckland-Campbell Islands. III–V. Asteroidea, Holothurioidea and Crinoidea. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 79: 263–420.

Echinoids Dix, T.G. (1970b): Biology of Evechinus chloroticus (Echinoidea: Echinometridae) from different localities 2. Movement. New Zealand Journal of Marine and Freshwater Research 4: 267–77. Dix, T.G. (1970c): Biology of Evechinus chloroticus (Echinoidea: Echinometridae) from different localities 3. Reproduction. New Zealand Journal of Marine and Freshwater Research 4: 385–405. McLary, D., Barker, M. (1998) Reproductive isolation? Interannual variability in the timing of reproduction in sympatric sea urchins, genus Pseudechinus. Invertebrate Biology 117(1): 75–95. Miskelly, A. (2009) Sea urchins of the world: diversity, symmetry, and design [Ashley Miskelly, 2009, Blackheath, N.S.W. ISBN: 9780646518961 (hbk)], 103 p.

Crinoids Hoggett, A.K., Rowe, F.W.E. (1986) A reappraisal of the family Comasteridae A.H. Clark, 1908 (Echinodermata: Crinoidea), with the description of a new subfamily and a new genus. Zoological Journal of the Linnean Society 88: 103–142.

Asteroids Byrne, M., Gonzalez-Bernat, M., Doo, S., Foo, S., Soars, N., Lamare, M. (2013) Effects of ocean warming and acidification on embryos and non-calcifying larvae of the invasive sea star Patiriella regularis. Marine Ecology Progress Series 473: 235–246. Crump, R.G. (1971) Annual reproductive cycles in three geographically separated populations of Patiriella regularis (Verrill), a common New Zealand asteroid. Journal of Experimental Marine Biology and Ecology 7(2): 137–162. Georgiades, E.T., Temara, A., Holdway D.A. (2006) The reproductive cycle of the asteroid Coscinasterias muricata in Port Phillip Bay, Victoria, Australia. Journal of Experimental Marine Biology and Ecology 332(2): 188–197. Grange, K.R., Singleton, R.I., Richardson, J.R., Hill, P.J., Main W. de L. (1981) Shallow rock-wall biological associations of some southern fiords of New Zealand. New Zealand Journal of Zoology 8(2): 209–227. Lane, D.J.W., Rowe, F.W.E. (2009) A new species of Asterodiscides (Echinodermata, Asteroidea, Asterodiscididae) from the tropical southwest Pacific, and the biogeography of the genus revisited. Zoosystema 31(3): 419–429. Town, J.C. (1980) Movement, morphology, reproductive periodicity, and some factors affecting gonad production in the seastar Astrostole scabra (Hutton). Journal of Experimental Marine Biology and Ecology 44(2): 111–132.

Produced for NIWA by TC Media Ltd www.tcmedia.co.nz July 2014 59