stunningabout guide | about | species pages | icons | glossary seaweedsbeautiful

brownsa guide to the large brown seaweeds of Version 1, 2016

Kate Neill Wendy Nelson

with Michelle Kelly & Blayne Herr

1 about this guide | about seaweeds | species pages | icons | glossary about this guide

Seaweeds or marine macroalgae occur around the world and are found from the high shore to depths of around 200 m in some places. They are the marine equivalent of land plants and they vary in size from small filament- like species to huge . Large brown are common and obvious on most parts of New Zealand’s coastline and we hope this guide will help you identify them in the field.

This first part of STUNNING SEAWEEDS is a fully illustrated working e-guide to the most commonly encountered large seaweeds, the BEAUTIFUL BROWNS. 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 seaweeds and invertebrates that NIWA’s Coasts and Oceans group is presently developing.

The e-guide starts with a simple introduction to seaweeds, 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 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 plants 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. Outlying island groups are shown on the maps as a two-letter code: Ke = Kermadec Islands; Ch = ; Bo = Bounty Islands; An = ; Ak = ; Ca = Campbell Islands. 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.

Kate Neill is a marine biologist based in New Zealand at NIWA’s Wellington office. Kate has been involved in marine macroalgae research for 15 years and has worked on various aspects of their diversity, , distributions, aquaculture, and introduced species. Kate also works on starfish (Asteroidea) and is a co-author of the e-guide Extraordinary Echinoderms.

Wendy Nelson is a marine biologist/ phycologist based in New Zealand at NIWA’s Wellington office and the University of Auckland. Wendy is an authority on marine macroalgae, with a wide-ranging knowledge of all things seaweed, but for over 35 years has had a particular focus on macroalgal diversity and taxonomy. Wendy recently published an illustrated guide to New Zealand seaweeds.

For any advice on seaweeds you find, please email your photos and queries to Kate ([email protected]) or Wendy ([email protected])

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

Remember to check the websites for updated versions!

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 2 about this guide | about seaweeds | species pages | icons | glossary

a typical species page layout

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

species images inset images show variations and/or common name closeup detail of species body plan icon highlighting the basic shape, or a special characteristic, that defines a group of these organisms species classification life history icon see species index for highlighting arrangement geographic distribution

scale bar indicating relative size of organism in the main image

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

information details on external and internal distribution characters and section of coastline habitat or around offshore islands where species is most commonly found

make notes of where you encountered this species and let us know if you find it at a key taxonomic references it could also be ... new location some species are difficult to tell apart without more detailed information, so check the other species in the guide listed here to make sure that you have the correct species

3 about this guide | about seaweeds | species pages | icons | glossary

about seaweeds

Seaweeds (marine macroalgae) are found in a great number of habitats, from the shallow intertidal zone, to snorkelling and diving depths and deeper. So it is quite likely that whenever you are in or on the sea a seaweed won’t be far away. Seaweeds range in size from small filaments to huge plants more than 10 m high.

Like land plants, seaweeds photosynthesise and are grouped into three phyla, in two kingdoms, based on their photosynthetic pigments: brown (), green (Chlorophyta) and red (Rhodophyta) seaweeds.

This first version of the Stunning Seaweeds e-guide focuses on some beautiful brown seaweeds. All of the species in this version of the guide are in the class Phaeophyceae. The brown seaweeds (phylum Ochrophyta) include the largest species of seaweeds and are often an obvious sight on shorelines, forming forests in rocky areas. There are several commercial species of brown seaweeds, including those that produce alginates and several that are eaten e.g. ( pinnatifida).

The green seaweeds (phylum Chlorophyta) are the least diverse of the three phyla in New Zealand. Species of Ulva are known as sea lettuce and can form large blooms in some areas.

Phylum Rhodophyta (the red seaweeds) is the most diverse of the three phyla and includes some well- Chromista Plantae KINGDOM known species that produce agar (e.g. Gracilaria, Pterocladia) and the seaweed used in making PHYLUMOchrophyta PHYLUMChlorophyta PHYLUMRhodophyta () (green algae) (red algae) sushi (known as Nori in Japan and karengo in New Zealand, species of Pyropia and Porphyra).

The main body of a seaweed is called a thallus – seaweeds don’t have true leaves, or stems or roots like land plants. The main stem of a seaweed is usually called a stipe or axis, and the thallus is called a blade. Sometimes they have structures that look leaf-like. Seaweeds attach to rocky substrates with a variety of different holdfasts – sometimes very solid dome-like bases and other times made up of finger-like haptera that spread long the rock surface and attach to the rock. In this guide we use the terms stipes and blades throughout.

4 about this guide | about seaweeds | species pages | icons | glossary beautiful browns brown seaweed

Large brown seaweeds (the Ochrophyta) form forests in the coastal zone. Growing on rocky substrates, these seaweeds provide three-dimensional structure and habitat for many other coastal species - invertebrates, fishes and other seaweeds.

There are two main types of large browns – true kelps and fucoids. The true kelps belong to the order Laminariales and have a life history where the conspicuous phase - the kelp phase () that you see on the shore or when you are diving - alternates with a microscopic phase (). There are three genera of true kelps native to New Zealand – , and – and one introduced kelp, Undaria pinnatifida.

The fucoids, or members of the have a direct life history in which egg and sperm are produced and fuse to form a zygote. There are many genera and species of Fucales in New Zealand – e.g. , , Hormosira, Landsburgia, Marginariella, , Xiphophora. The species of that are known as bull kelp in New Zealand are actually fucoids and not true kelps.

Large brown seaweeds are very important along temperate coasts worldwide as they act as ecosystem engineers structuring the habitat and influencing the physical environment, affecting such features as light, nutrient concentrations and water movement. In addition to providing shelter and surfaces to settle on, seaweeds are also vitally important to local food chains as they contribute significant biomass.

Brown seaweeds are more closely related to various types of microscopic algae than they are to red or green seaweeds – they all share life in the sea and have some similarities in the ways they grow and attach to substrates, but they are actually very different from each other with respect to their origins.

Chromista Plantae

Ochrophyta Chlorophyta Rhodophyta

5 about this guide | about seaweeds | species pages | icons | glossary species index

Family Dictyotaceae Dictyota kunthii 8 ORDER

Family Durvillaeaceae 9 Durvillaea chathamensis 10 Durvillaea poha 11 Durvillaea willana 12 Family Hormosiraceae Hormosira banksii 13

Family Carpophyllum angustifolium 14 Carpophyllum flexuosum 15 Carpophyllum maschalocarpum 16 Carpophyllum plumosum 17 Cystophora platylobium 18 19

Fucales Cystophora scalaris 20 Cystophora torulosa 21 ORDER Landsburgia quercifolia 22 Phyllotricha verruculosa 23 Chromista

Ochrophyta Sargassum scabridum 24 CLASS Phaeophyceae Sargassum sinclairii 25 PHYLUM

KINGDOM Family Seirococcaceae Marginariella boryana 26 Marginariella urvilliana 27

Family Xiphophoraceae Xiphophora chondrophylla 28 Xiphophora gladiata 29

Family Alariaceae Undaria pinnatifida 30 Family 31 Family

Laminariales 32 ORDER Lessonia tholiformis 33 Lessonia variegata 34

6 about this guide | about seaweeds | species pages | icons | glossary species index

Family ligulata 35

Desmarestiales ORDER

Chromista Family Splachnidiaceae Ochrophyta

CLASS Splachnidium rugosum 36 Phaeophyceae PHYLUM Family Scytothamnaceae KINGDOM australis 37 ORDER

7 Class Phaeophyceae Order Dictyotales Family Dictyotaceae

classification Previously known as Dictyota kunthii Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Xiphophora gladiata It couldalsobe………. from Norfolk Island,ChileandPeru. three Kings, South, North, Stewart and Chatham Islands. Also known Found in the low to upper subtidal on intertidal open coasts around easily visible. not are but blades the over scattered are structures Fertile at bases. brown dark often brown, olive to Golden branches. prostrate look can tongues, fleshy shaggy small, when dense. Holdfast is hairy, in almost felted, and has lobed, covered sections, forked like, Plants are fan-like and up to 50 cm high. Blades divided into strap- Glossophora kunthii substrate (C.Agardh) Grev. habitat main image: Erasmo Macaya insetimage: Tracy Farr 10 cm 8 Class Phaeophyceae Order Fucales Family Durvillaeaceae

classification antarctica Durvillaea Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch,

morphology bull kelp, rimurapa Durvillaea willana Durvillaea poha Durvillaea It couldalsobe………. Atlanticislands. America andsouthern South Ocean, Southern the in widespread Also Islands. Campbell and Bounty, Auckland Chatham, Antipodes,Snares, including south, the in Island Stewart to down north the in Islands Kings Three the around coasts exposed of zone intertidal low the in rock on Found are noteasilyvisible. but blades the over scattered are structures Fertile green. olive to growsfrom large,browna dark to Golden holdfast. dome-shaped and unbranched, cylindrical, is stipe Short become cross-section. in oval or blades areas exposed narrower more and more whip-like, however, internal In they an always remain with rounded structure. rubbery, honeycomb and thick are portions, strap-like into to 5 m long, orsometimesup to 10 m long. Blades often divided massive, growing are and blades flattened of up made are Plants surface substrate (Cham.) Har. habitat main image: Kate Neillinsetmage: Erasmo Macaya 1 m 9 Class Phaeophyceae Order Fucales Family Durvillaeaceae

classification chathamensis Durvillaea Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Durvillaea poha Durvillaea antarctica Durvillaea It couldalsobe………. Endemic totheChathamIslands. Found in exposed areas from the low intertidal to about 2 m depth. are scatteredover thebladesbutarenoteasilyvisible. from a large, brown. dome-shaped holdfast. Dark structures Fertile honeycombno internal and grows Stipe is unbranched, structure. edges, wavy and sides parallel with flattened divided, not usually Plants grow massive blades, upto15mlongandaremadeof surface substrate C.H. Hay habitat main image: Wendy Nelsoninsetmage: Tracy Farr 1 m 10 Class Phaeophyceae Order Fucales Family Durvillaeaceae

classification poha Durvillaea Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Phycologia, 51(2),151–156. Fraser,Spencer,C.I., H.G.,Waters, J.M.(2012) morphology Durvillaea willana Durvillaea antarctica Durvillaea It couldalsobe………. D. antarcticaform’ of . D. pohawas previouslyknown andbroader asthe shorter ‘cape Islands. Auckland and Snares Stewart, Island, South southern above around band D.antarctica , a in often areas, exposed in intertidal the in Found visible. easily not are but blades the over golden. scattered are to structures orange-brown Fertile Blades clump. a of out growing stipes several with together join may Holdfasts tip. towards the flattened and broad flattened in cross-section. remain Short stipe is orange-yellow always and becomes but tips, the at whip-like become can Blades honeycomb structure. internal an with flattened, and broad massivePlants usually5–10mlong, blades, and are madeupof surface C.I. Fraser &J. Waters substrate sp. nov. (Fucales, Phaeophyceae): a buoyant southern bull-kelp species endemic to New Zealand. New to endemic species bull-kelp nov.buoyant sp.Phaeophyceae):(Fucales,southern a poha Durvillaea habitat main image: Kate Neillinsetmage: Erasmo Macaya 1 m 11 Class Phaeophyceae Order Fucales Family Durvillaeaceae

classification willana Durvillaea Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Durvillaea poha Durvillaea antarctica Durvillaea It couldalsobe………. Strait), South IslandandStewartCook Island. not (but Island North the of corner southeastern the of coasts exposed on subtidal upper to intertidal low the in Found are scattered over thebladesbutarenoteasilyvisible. structures Fertile brown. Dark disc. dome-shaped a Holdfast and produces cylindrical, side branches. Stipeisthick, structure. leathery, sometimesdividedintostraps, honeycomb withnointernal Plants grow massive blades, upto5mhighandconsistof toughand surface Lindauer substrate habitat main image: Wendy Nelsoninsetmage: Kate Neill 1 m 12 Class Phaeophyceae Order Fucales Family Hormosiraceae

classification banksii Hormosira Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch,

morphology Neptune’s necklace on o rcs r tns n h itria zn i setrd or sheltered Also known from . in zone Islands. Chatham intertidal and Stewart South, the North, around sites in exposed stones or rocks on Found over thebeadsandareeasilyvisibleassmallbumps. scattered are structures Fertile grey-brown. to Yellow-green disc. vary in size according to habitat and exposure. byAttaches a small hollow of ‘beads’ joined togetherchains by thin constrictions. Beads branched Plants upto30cm(sometimes50 cm) highand consist of substrate (Turner) Decne. habitat main image: Kate Neillinsetmage: Tracy Farr 5 cm 13 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification Carpophyllum angustifolium Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Carpophyllum flexuosum Carpophyllum maschalocarpum It couldalsobe………. Island. North around ThreeKingsIslandsandnorthern channels Found in the low to intertidal subtidal on exposed coasts or in surge stem andleaves. intheangle between ontheupper branches, the occur in bunches structures Fertile brown. Dark attachments. flattened peg-like with of branches up made is leaf-like Holdfast egg-shaped. of are cylindrical; bladders Consists are air stems high. branches, m of side 1.5 either on to carried blades up and shrub-like is Plant habitat J. Agardh main image: Kendall Clementsinsetmage: Wendy Nelson 70 cm 14 to 45m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification Carpophyllum flexuosum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Carpophyllum maschalocarpum Carpophyllum angustifolium It couldalsobe………. Chatham Islands. or channels, and pools deep deeper in sheltered areas around South, Stewart North, in and or subtidal, upper the in Found the anglebetween thestem andleaves. with smallleaves. are found in branches These specialised branches upper specialised on occur structures brown.Fertile dark brownto air midrib; creeping. Golden and flattened is rounded.Holdfast are bladders a have and broader are leaves Lower slightly. zags zig- and flattened is which stem, main a of side either branches on Leaf-likecarried high. are m blades 2 to up and shrub-like is Plant habitat (Esper) Grev. main image: Erasmo Macaya insetmage: Roberta D’Archino 10 cm 15 to 45m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification Carpophyllum maschalocarpum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Cystophora platylobium Carpophyllum flexuosum Carpophyllum angustifolum It couldalsobe………. exposed coastsaround SouthandChathamIslands. North, Found in the low intertidal to upper subtidal in sheltered areas or on andstem. in tightbundlesbetween theleaf brown. are has lateral Dark structures peg-like attachments. Fertile and flattened is Holdfast egg-shaped. are bladders air flattened; is broad and a stem, which either side of are carried onbranches and leaf-like are Blades high. m 1.5 to up and shrub-like is Plant habitat (Turner) Grev. main image: Erasmo Macaya insetmage: Kate Neill 10 cm 16 to 45m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification Carpophyllum plumosum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Landsbergia quercifolia It couldalsobe………. and theChathamIslands. exposedor Island aroundcoasts North Islands,Kings Three eastern Found in the low intertidal to upper subtidal in moderately sheltered the in branches angle between stemandleaf. long on are structures Fertile brown. greenish to and rounded. Holdfast is a prostrate extension the stem. Golden of stalked are bladders air flattened; is which stem, a of side either from oak leaf-shaped to very finely divided, and occur on branches Plant is shrub-like and up to 1.0 m high. Blades are divided, varying habitat (A. Rich), J.(A. Rich), Agardh 40 cm images: Kate Neill 17 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification platylobium Cystophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Carpophyllum maschalocarpum It couldalsobe………. known from Australia. deep in ususally water around coasts, southern North Island, South exposedand Stewart Islands. open, Also on subtidal the in Found two rows butaren’t onleaf-like structures easytosee. cone-shaped. Golden to brown.dark Reproductive are structures in to disc Holdfast oval. slightly to round are bladders Air flattened. is stalk Main leaf- branches. flat, side along are alternately Blades occuring high. lobes like m 2 to up and shrub-like are Plants habitat (Mertens) J.(Mertens) Agardh main image: LouHuntinsetmage: Roberta D’Archino 5 cm 18 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification retroflexa Cystophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Cystophora torulosa Cystophora scalaris It couldalsobe………. to ususally Islands, Chatham about 10mdepth.Alsoknown from Australia. and Stewart South, North, around coasts, exposed open, on subtidal to subtidal upper the in Found butaren’t easytosee.branchlets cylindrical the on rows two in are structures Reproductive brown. are stalks. on Holdfast short solid greyish-and conical. Medium to dark stems round to oval Main are bladders appearance. Air twisted. bushy sometimes and flattened a giving stem the around are arranged origin.Branchlets frombend back theirpointof narrow that andoccurin groups andcylindrical onsidebranches are Branchlets high. more or m 1.5 to up and shrub-like are Plants habitat (Labill.) J. Agardh main image: Erasmo Macaya insetmage: Kate Neill 20 cm 19 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification scalaris Cystophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Cystophora torulosa Cystophora retroflexa It couldalsobe………. Auckland and Chatham Stewart, Islands. South, Island, North around southern coasts exposed on subtidal to intertidal low the in Found butaren’t easytosee.branchlets cylindrical the on occur structures Reproductivegrey-brown. zig-zag Dark and tough are stalks strongly. Air bladders are oval to Main round. Holdfast solid and conical. branches. side on alternating groups flattened packed, narrow closely in are occur Branchlets and cylindrical high. and 1m to up and shrub-like are Plants J. Agardh habitat 25 cm image: MalcolmFrancis 20 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification torulosa Cystophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Cystophora scalaris Cystophora retroflexa It couldalsobe………. Australia. coasts, exposed on around North, South, subtidal Stewart and upper Chatham Islands. Also to known from intertidal low the in Found blades butaren’t easytosee. Golden conical. club-shaped and the rowson two in are brown.Reproductive structures solid flattened Holdfast but round. base, bladders the Air at above. cylindrical and thick are stems Main swollen are Branchlets and club-shaped high. and occur in m bunches on alternating side branches. 1 to up and shrub-like are Plants (R.Br.) J. Agardh habitat 10 cm images: Erasmo Macaya 21 to 40m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification quercifolia Landsburgia Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Carpophyllum plumosum It couldalsobe………. Kings, South,Stewart North, andChathamIslands. Three around coasts open on water deep to subtidal the in Found brown. Dark serrated marginsfound ontheupperbranches. conical. with leaves thicker and specialised, small, solid are structures Reproductive Holdfast branches. shed from scars have and cylindrical are andoccur stems Main branches. midrib, alternate on afaint with flattened are that leaves shaped Plants grow up to 1.5 m high and are shrub-like with small oakleaf- habitat Harv. main image: Roberta D’Archino insetmage:main image: Erasmo Roberta Macaya D’Archino 30 cm 22 to 40m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification Previously known as Phyllotricha verruculosa Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Sargassum sinclarii It couldalsobe………. century by whalingandsealingvessels. is considered to have been introduced to New Zealand in the19th species This Australia. from known Also Island. Stewart and Island South around harbours and bays sheltered in subtidal the in Found andnoteasilyvisible. branches upper in scattered structures Reproductive Mid-brown. Air branches. up. further developing thinner with disc-like becomes and conical but Holdfast rounded. bladders base at thick is stem Main that are leaf-like and variably shaped, carried on forked branches. blades with shrub-like are and high more or 1m to growup Plants Sargassum verruculosum habitat (C. Agardh) R.R.MDixon &Huisman images: Roberta D’Archino 10 cm 23 Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification scabridum Sargassum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, Sargassum sinclairii It couldalsobe………. New Zealand. Islandsandnorthern the Kermadec Found in the low intertidal to subtidal in pools on open coasts around swollenthe upperbranches, andwarty. in occur rounded.brown.Golden conical. structures Holdfast Fertile stems are which cylindrical and have spines; air bladders small and a midribandserrated margin.Bladesoccuronbushy, branched Plants grow to 1m high and are shrub-like with leaf-like blades with substrate Hook.f. &Harv. habitat 25 cm images: Wendy Nelson 24 to 40m Class Phaeophyceae Order Fucales Family Sargassaceae morphology classification sinclairii Sargassum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Carpophyllum flexuosum (lower leaves only) Sargassum scabridum It couldalsobe………. Stewart andChathamIslands. coasts, often in pools aroundor channels, Three Kings, South, North, Found in the low intertidal to upper subtidal on moderately exposed lobed ortoothedandarecarriedonstalks. and rounded. Holdfast domed. Golden brown. Fertile structures are stalked bladders air smaller; are branches upper large,on leaves are base at leaves flattened; slightly or rounded are which stems, blades leaf-like with with a midrib and toothed margin. Blades are on bushy branched shrub-like are and high 1m to grow Plants Hook.f. &Harv. habitat main image: Roberta D’Archino insetmage:main image: Erasmo Roberta Macaya D’Archino 30 cm 25 to 40m Class Phaeophyceae Order Fucales Family Seirococcaceae

classification boryana Marginariella Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Marginariella urvilliana It couldalsobe………. Island,South,Stewart, North southern Chatham andSnaresIslands. Foundfrom exposedon subtidal lowto the aroundcoasts intertidal blades ononesideonly. finger-likeare grow image) and edgesstructures (inset of from the manyGolden brown. conical andmadeupof thinbranches. Fertile and occur on margin of the blades, sometimes with a point. Holdfast egg-shaped bladders air below; bare is stem holdfast, the above on opposite edges the of stem, is which flattened and twists sharply alternate Blades edges. toothed with strap-like and long are that Plants grow blades to more than 2 m high and are comprised of substrate (A. Rich) Tandy.(A. Rich) habitat main image: insetmage: ChrisHepburn Erasmo Macaya 10 cm 26 to 40m Class Phaeophyceae Order Fucales Family Seirococcaceae

classification urvilliana Marginariella Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Marginariella boryana It couldalsobe………. and Snares Chatham, Stewart, South, Islands. Auckland Island, North around southern coasts exposed on subtidal to intertidal low the in Found the bladesononesideonly. brown. structures are Fertile finger-like and grow from the edges Golden of branches. thin many of up made conical Holdfast blades. the of margins growthe and roundon bladders below;air bare is is flattened and twists sharply above the holdfast, stem stem, which with toothededges. the onoppositeedges Bladesalternate of blades strap-like and long of consist and high m 2 to grow Plants substrate (A. Rich) Tandy.(A. Rich) habitat main image: Roberta D’Archino insetmage:main image: LouHunt Roberta D’Archino 10 cm 27 Class Phaeophyceae Order Fucales Family Xiphophoraceae

classification chondrophylla Xiphophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology he Kns sad ad oten ot Iln. lo nw from known Also Island. Australia. North northern and Islands Kings around Three coasts exposed on subtidal to intertidal low the in Found andnoteasilyvisible. aresunkenstructures intheupperbranches Reproductive Grey-brown. stem. lower the of extension flattened a Holdfast flattened. to rounded are branches base the towards blades thatbecomenarrower andmoreforked towards thetips, strap-likeand forked of consist and high cm grow30–50 Plants to substrate habitat (Turner) Mont.ex Harv. main image: insetmage: ChrisHepburn Kate Neill 10 cm 28 Class Phaeophyceae Order Fucales Family Xiphophoraceae

classification gladiata Xiphophora Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Dictyota kunthii It couldalsobe………. andCampbellIslands. AlsoknownAuckland from Australia. Antipodes,Snares,Stewart, Chatham, South, Island, aroundNorth southern coasts exposed on subtidal to intertidal low the in Found andnoteasilyvisible. aresunkenstructures intheupperbranches fan-shaped flat, forming plants. branches, Holdfast solid forked and disc-like. Grey-green-brown. on Reproductive blades elongate flattened, strap-like, of consist and high cm 50 to up grow Plants substrate (Labill.) Mont.ExKjellm. habitat main image: Roberta D’Archino insetmage:main image: JennDalen Roberta D’Archino 10 cm 29 Class Phaeophyceae Order Laminariales Family Alariaceae Undaria pinnatifida Undaria pinnatifida morphology classification Phaeophyceae) introduced genes populations. deducedfrom –origins andsuccession of Phycologia, mitochondria (Laminariales, 45,687–695. pinnatifida Undaria in diversity Genetic (2006) H. T.,Kawai,Kitayama,S.M., Boo,Wang,W.D., Aguilar-Rosas,L.E. K., W., Neill, Nelson, S., Uwai, Nelson, W. (2013)New Zealandseaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Wakame Ecklonia radiata Ecklonia It couldalsobe………. Argentina, Britain,France, Italy, Americaand Spain. North Islands. Native to Japan, Korea and China. Also invasive in Australia, found around the North Island, South, Stewart, Chatham and Snares including concrete and wood. Introduced to New Zealand. Presently Found in the low to intertidal subtidal growing on any hard surface, big appearstospiral around thestalk. often darker brown thantheblades, and deeply folded andwhen and thicker is It stalk. the of base the at found is and size in varies golden brown to dark brown. Reproductive structure (the ) continues into the stalk. Holdfast of forked finger-like branches. Pale deeply divided into lobes. Blades have a well defined midrib which Plants grow a large blade, upto1 m high and consist of often substrate (Harv.) Suringar. habitat main image: Kate Neillinsetmage: SherylMiller 30 cm 30 to 40m Class Phaeophyceae Order Laminariales Family Laminariaceae

classification Macrocystis pyrifera Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology temperate waters. Bounty, Antipodes, and Campbell Islands. Widespread Auckland in action. Found around southern North Island, South, Chatham, Stewart, Found in the subtidal on open coasts but sheltered from direct wave onsmoothbladesthathave noairbladders. patches darker as occur structures Reproductive brown. Golden structure. haptera cylindrical amassive forming dome-shaped intertwined of near the the top plant of where they sequentially.split off Holdfast edges and an oval bladder at the base. Blades grow from an area long of water. Blades Consist are long and high. leaf-like, wrinkled m with small teeth 20 on the to stipesthatcarrythebladesuptowardscylindrical the thesurface of up growing huge, be can Plants substrate (L.) C.Agardh. habitat main image: upperinsetmage: ChrisHepburn Erasmo Macaya lower insetmage: Kate Neill 1.5 m 31 to 100m Class Phaeophyceae Order Laminariales Family Lessoniaceae Ecklonia radiata Ecklonia radiata morphology classification Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate Undaria pinnatifida It couldalsobe………. and . ,Africa, South knownAustralia,from WestAlso Africa Islands but found on Bank, Mernoo 200 km east the of South Island. South, North, Stewart and Snares Islands. Absent from the Chatham Kings, Three around beds large forms sometimes coasts, exposed or exposed moderately on subtidal to intertidal low the in Found thebladeandarevisibleasdarker areas. the lobesof haptera. Golden brown. Reproductive structures occur as patches on finger-like Holdfast made up of unbranched. or cylindrial branches into lobes. Bladesmay and be smooth orwrinkled.Stipe is cylindrical Plants grow a large blade, to1 m high and consist of deeply divided (C. Agardh) J. Agardh. habitat main image: MalcomFrancis insetmage: Svenja Heesch 10 cm 32 Class Phaeophyceae Order Laminariales Family Lessoniaceae

classification Lessonia tholiformis Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology Found in the subtidal on open coasts. Endemic to the Chatham Islands. onthebladesandarevisibleasdarker areas. occur aspatches structures Reproductive brown. greenish to Golden branches. like finger- lobed and flattened with Holdfast flattened. to cylindrical are smoothand glossy, growing and stems thatare from branches Plants grow up to 1m high and consist of long, strap-like blades that substrate C.H. Hay habitat 15 cm image: Tracy F.r 33 Class Phaeophyceae Order Laminariales Family Lessoniaceae

classification Lessonia variegata Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, morphology North, SouthandStewartNorth, Islands. Found in the upper subtidal to deepwater on exposed coasts around and rigid Holdfast onthebladesandarevisibleasdarker areas. occur aspatches flattened. structures Reproductive and brown. light to tough brown Golden buttressed. are that stems and widely spaced marginal teeth.Bladesgrow from forked branches with strap-likeblades long of consist and high 1m to up are Plants substrate J. Agardh habitat main image: Roberta D’Archino insetmage:main image: Kate Roberta D’Archino Neill 20 cm 34 to 40m Class Phaeophyceae Order Family Desmarestiaceae morphology classification Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, substrate northern AfricaandEurope. northern America, North and South known Japan, Korea, Also Australia, southern Islands.from Campbell and Snares, Stewart, Auckland Antipodes, Chatham, Bounty, South, North, southern Islands, Kings aroundThree coasts open on subtidal to lowthe intertidal in Found rock. down andcanetch are not easily visible. Plants release acid when they are breaking Reproductive the blades and are buried in the surface structures of holdfast disc. Golden brown, or green and when drifting.detached short and cylindrical and expands into the blade. Attached by small Young blades may appear to be fringed with hairs. Stem is very with a midrib and similar secondary blades growing from theedges. blades ribbon-like long of consist and high m 1 to up grow Plants (Stackh.) J.V.(Stackh.) Lamour. habitat main image: Roberta D’Archino insetmage:main image: Kate Roberta D’Archino Neill 20 cm 35 Class Phaeophyceae Order Scytothamnales Family Splachnidiaceae

classification Splachnidium rugosum Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch,

morphology Dead man’s fingers ot, twr ad hta Ilns As kon rm Australia, Islands. fromSouth AfricaandJuanFernández known Also Islands. Chatham and Stewart South, Found in the intertidal on moderate to Reproductiveexposed coasts around brown.North, mid to Golden disc. thebranches. arevisibleassmallholesinthesurfaceof structures small material. jelly-like a clear, is with Holdfast filled are Branches swollen. than rather leathery look can branches Older branches. side and stems blunt-tipped swollen, of consist and high cm 20 to up grow Plants substrate (L.) Grev. habitat 8 cm images: Erasmo Macaya 36 Class Phaeophyceae Order Scytothamnales Family Scytothamnaceae Scytothamnus australis morphology classification Nelson, W. (2013) New Zealand seaweeds. Anillustrated guide. Te Papa Press, Wellington, 328pp. Adams, New 360 pp. N.M.(1994)Seaweeds Zealand: Anillustrated of guide. Canterbury University Press, Christchurch, ot, ot, twr ad hta Ilns As kon from known Also Islands. Chatham Australia andChile. and Stewart South, North, aroundexposure moderate of areas in mid-intertidal the in Found visible. the blades but are not easily are buried in the surface of structures small disc. Dull grey to khaki brown, or black when dry. Reproductive are branches flattened tocylindrical with pointed tips. Holdfast is a giving branched, a bushy and sometimes scruffy appearance. Main Plants grow to 30 cm high and are shrub-like, irregularly and densely substrate habitat (J. Agardh) Hook.f. &Harv. 10 cm images: Kate Neill 37 about this guide | about seaweeds | species pages | icons | glossary icons

marine macroalgae with photosynthetic pigments that result in their brown colour, and give rise brown to their common name (brown seaweeds, Phylum Ochrophyta). Includes the largest species on seaweeds shorelines, forming kelp forests in rocky areas body plan body

species recorded across the naturally occuring around New southern antipodean globe but restricted to the Zealand and Australia only hemisphere southern hemisphere only

species naturally ocurring outside of New Zealand waters introduced widespread species recorded globally and is found in New Zealand, invasive life history

naturally occuring around New native Zealand, endemic

bushy, composed of numerous inflated, hollow, or jelly-filled, bladders shrub-like branches and branchlets that flexible chamber emerge from a central stem

flattened or sheet-like thallus; composed of flattened, bladed may be simple, divided, or strap-like elongate, branches that are morphology include multiple blades longer than they are wide

thick skin, tough, flexible, leathery slightly elastic surface

anything man-made such as artificial indentation in rock filled with mooring blocks, mussel lines, rockpool substratum water, intertidal wharf piles

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

hard substrate such as rock mudstone, sandstone, basalt, compressed carbonates

underwater cliffs and slopes, coralline algae, seagrass or organisms exposed to wave algal beds wall algal beds surge and currents, and subdued illumination

exposed shoreline zone exposed shoreline zone between high and low tides, between high and low tides,

habitat including rock flats, pools, including rock flats, pools, intertidal overhangs, crevices, organisms subtidal overhangs, crevices, organisms exposed to wave action, exposed to wave action, temperature extremes, full temperature extremes, full illumination, and desiccation illumination, and desiccation

38 about this guide | about seaweeds | species pages | icons | glossary glossary agar a compound produced from red seaweeds and used in food and to make the agar plates commonly used in laboratories algal beds areas of seafloor with coralline algae, sea-grass or multiple seaweed species alginates compounds found in the cell walls of brown algae and used as emulsifiers or stabilisers in various industries artificial substratum anything man-made such as mooring blocks, mussel lines, wharf piles antipodean naturally occurring in New Zealand and Australia, and may include seamounts and ridges to the north blade seaweed blades range from a single sheet-like structure through to finely divided ‘leaves’ dendritic branching, tree-like diameter the distance across the widest point of a circle 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 gametophyte a phase in the life-cycle of those algae taht undergo alternation of generations; the stage that produces gametes gametes male and female cells involved in sexual reproduction gelatinous jelly-like, slippery habitat the environment and local situation in which an organism lives haptera finger-like outgrowths from the base of the stipe/stem that serve as attachment points to the substrate hard solid to the touch, not compressible, rigid 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 beyond New Zealand waters, now occurring in New Zealand and other locations, invasive, adventive leathery thick, tough, flexible, slightly elastic margins edge of a surface meandering wandering along and above substratum attached at intervals, repent midrib a raised rib or strengthened vein-like area along the mid line of leaf-like blades morphology form and structure, shape native naturally occurring in New Zealand, but may also occur naturally elsewhere, endemic range extension since first described in New Zealand, this species has been recorded elsewhere refuge safe place to hide from predators rock hard substratum such as mudstone, sandstone, basalt, compressed carbonates rock pool excavation in rock, filled with water, in the intertidal zone rough irregularly pitted and ridged surface, often tough sand small coarse grains of worn silica, rock, and shell seabed composed of a variety of sedimentary substrata including coarse gravels, shell hash and sands to finer sand, mud, and silts; associated organisms are susceptible to inundation and scouring from wave surge and currents, and subdued illumination sinuous wavy pattern smooth even, hairless, silky, can be slightly undulating soft easily compressible, elastic sporophyll a modified blade for spore-bearing stipe the stem-like region between the holdfast and blade of a seaweed subtidal zone below the low tide, including rock flats, slopes, walls, crevices, overhangs, boulder fields; associated organisms are exposed to wave surge, currents and subdued illumination thallus the entire structure of a seaweed; generally comprised of a holdfast, stipe/stem and blade/s wall underwater cliff or slope; associated organisms are exposed to wave surge, currents and subdued illumination warty bearing small flattened bumps or tubercles widespread species recorded globally zygote a cell that is formed when female and male gametes combine

39 about this guide | about seaweeds | species pages | icons | glossary acknowledgements

Thanks to the many talented photographers whose work was used with permission in our guide, including: Erasmo Macaya (Universidad de Concepción, ), Chris Hepburn (University of Otago), Lou Hunt (Ministry for the Environment), Kendall Clements (University of Auckland), Jenn Dalen (Te Papa), Roberta D’Archino, Tracy Farr, Malcolm Francis, Svenja Heesch, Sheryl Miller (all NIWA or ex-NIWA). 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 (2015/2016 SCI). further reading

Adams, N.M. (1994) Seaweeds of New Zealand. An illustrated guide. Canterbury University Press, Christchurch. 360 pp. Brown, M.T., Nyman, M.A., Keogh, J.A., Chin, N.K.M. (1997) Seasonal growth of the giant kelp Macrocystis pyrifera in New Zealand. Marine , 129, 417–424. Choat, J.H., Schiel, D.R. (1982) Patterns of distribution and abundance of large brown algae and invertebrate herbivores in subtidal regions of northern New Zealand. Journal of Experimental Marine Biology and Ecology, 60, 129–162. Cole, R.G., Babcock, R.C., Travers, V., Creese, R.G. (2001) Distributional expansion of Carpophyllum flexuosum onto wave- exposed reefs in north-eastern New Zealand. New Zealand Journal of Marine and Freshwater Research, 35, 17–32. Dean, P.R., Hurd, C.L. (2007) Seasonal growth, erosion rates, and and photosynthetic ecophysiology of Undaria pinnatifida (Heterokontophyta) in southern New Zealand. Journal of Phycology, 43, 1138–1148. Forrest, B.M., Brown, S.N., Taylor, M.D., Hurd, C.L., Hay, C.M. (2000). The role of natural dispersal mechanisms in the spread of Undaria pinnatifida (Laminariales, Phaeophyceae). Phycologia, 39, 547–553. Fraser, C.I., Spencer, H.G., Waters, J.M. (2012) Durvillaea poha sp. nov. (Fucales, Phaeophyceae): a buoyant southern bull- kelp species endemic to New Zealand. Phycologia, 51(2), 151–156. Gillanders, B.M., Brown, M.T. (1994) Seasonal variation in standing crop, reproduction and population structure of Xiphophora gladiata (Phaeophyceae: Fucales). Botanica Marina, 37, 35–41. Guiry, M.D., Guiry, G.M. (2015) AlgaeBase. World-wide electronic publication, National University of Ireland, Galway. http://www.algaebase.org; searched on 06 August 2015. Hay, C.H. (1979a) Some factors affecting the upper limit of the southern bull kelp Durvillaea antarctica (Chamisso) Hariot on two New Zealand shores. Journal of the Royal Society of New Zealand, 9, 279–289. Hay, C.H. (1979b) Growth, mortality, longevity and standing crop of Durvillaea antarctica (Phaeophyceae) in New Zealand. Proceedings of the International Seaweed Symposium 9, 97–103. Science Press, Princeton. Hay, C.H. (1989) Lessonia tholiformis sp. nov. (Phaeophyta: Laminariales) from the Chatham Islands, New Zealand. New Zealand Journal of Botany, 27, 461–469. Hay, C.H. (1990a) The dispersal of of Undaria pinnatifida by coastal shipping in New Zealand and the implication for further dispersal on Undaria in France. British Phycological Journal, 25, 301–313. Hay, C.H. (1990b) The distribution of Macrocystis (Phaeophyta: Laminariales) as a biological indicator of cool sea surface temperature, with special reference to New Zealand. Journal of the Royal Society of New Zealand, 20, 313–336. Hay, C.H., Luckens, P.A. (1987) The Asian kelp Undaria pinnatifida (Phaeophyta: Laminariales) found in a New Zealand harbour. New Zealand Journal of Botany, 25, 329–332. Hay, C.H., South, G.R. (1979) Experimental ecology with particular reference to proposed commercial harvesting of Durvillaea (Phaeophyta, Durvilleales) in New Zealand. Botanica Marina, 22, 431–436. Hay, C.H., Villouta, E. (1993) Seasonality of the adventive Asian kelp Undaria pinnatifida in New Zealand. Botanica Marina, 36, 461–476. Hurd, C.L., Nelson, W.A., Falshaw, R., Neill, K.F. (2004) History, current status and future of marine macroalgal research in New Zealand: Taxonomy, ecology, physiology and human uses. Phycological Research, 52, 80–106.

Miller, S.M., Hurd, C.L., Wing, S.R. (2011) Variations in growth, erosion, productivity, and morphology of Ecklonia radiata (Alariaceae; Laminariales) along a fjord in southern New Zealand. Journal of Phycology, 47(3), 505–516. Nelson, W. (2013) New Zealand seaweeds. An illustrated guide. Te Papa Press, Wellington, 328 pp. Hepburn, C.D., Hurd, C.L. (2005) Conditional mutualism between the giant kelp Macrocystis pyrifera and colonial epifauna. Marine Ecology Progress Series, 302, 37–48. Kain, J.M. (1982) Morphology and growth of the giant kelp Macrocystis pyrifera in New Zealand and . Marine Biology, 67, 143–157. Nelson, W.A. (1994) Distribution of macroalgae in New Zealand - an archipelago in space and time. Botanica Marina, 37, 221–233.

40 about this guide | about seaweeds | species pages | icons | glossary

Schiel, D.R., Foster, M.S. (2015). The biology and ecology of giant kelp forests. University of California Press, Oakland. 416 pp. Stevens, C.L., Hurd, C.L., Smith, M.J. (2002). Field measurements of the dynamics of the bull kelp Durvillaea antarctica (Chamisso) Heriot. Journal of Experimental Marine Biology and Ecology, 269, 147–171. Taylor, D.I. and Schiel D.R. (2010) Wave-related mortality in zygotes of habitat-forming algae from different exposures in southern New Zealand: the importance of ‘stickability’. Journal of Experimental Marine Biology and Ecology, 290, 229–245. Uwai, S., Nelson, W., Neill, K., Wang, W.D., Aguilar-Rosas, L.E. Boo, S.M., Kitayama, T., Kawai, H. (2006) Genetic diversity in Undaria pinnatifida (Laminariales, Phaeophyceae) deduced from mitochondria genes – origins and succession of introduced populations. Phycologia, 45, 687–695. Womersley, H.B.S. (1987) The marine benthic flora of southern Australia. Part II. Government Printer, South Australia, Adelaide. 484 pp.

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