Diseases, pathogens and parasites of Undaria pinnatifida

Diseases, pathogens and parasites 

of Undaria pinnatifida 

MAF Biosecurity New Zealand Technical Paper No: 2009/44 

Authors: 

Neill, K., Heesch, S., Nelson, W. 

National Institute of Water and Atmospheric Research, Private 

Bag 14-901, Wellington 

Prepared for BNZ Post-clearance Directorate 

By National Institute of Water and Atmospheric Research 

As contract No: ZBS2005-01 

ISSN 1176-838X (Print) 

ISSN 1177-6412 (Online) 

ISBN 978-0-478-35752-3(Print) 

ISBN 978-0-478-35753-0(Online 

April 2008

Disclaimer 

While every effort has been made to ensure the information in this publication is accurate, the 

Ministry of Agriculture and Forestry does not accept any responsibility or liability for error or 

fact omission, interpretation or opinion which may be present, nor for the consequences of 

any decisions based on this information. 

Any view or opinions expressed do not necessarily represent the official view of the Ministry 

of Agriculture and Forestry. 

The information in this report and any accompanying documentation is accurate to the best of 

the knowledge and belief of the National Institute of Water & Atmospheric Research Ltd 

(NIWA) acting on behalf of the Ministry of Agriculture and Forestry. While NIWA has 

exercised all reasonable skill and care in preparation of information in this report, neither 

NIWA nor the Ministry of Agriculture and Forestry accept any liability in contract, tort or 

otherwise for any loss, damage, injury, or expense, whether direct, indirect or consequential, 

arising out of the provision of information in this report. 

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Telephone: (04) 474 4100 

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© Crown Copyright - Ministry of Agriculture and Forestry

Contents Page 

Executive Summary 1 

Overall objective 2 

Specific objectives 2 

1. Introduction 3 

2. Methods and Materials 5 

2.1. Definitions 5 

2.2. Data Sources 6 

2.3. Database 7 

2.4. Mapping 8 

3. Results 8 

3.1. General Comments 8 

3.2. description of known pathogen-host relationships 9 

3.3. Laminariales 13 

3.4. Brown algae other than Laminariales 16 

3.5. Red algae 20 

3.6. Green algae 27 

3.7. Xanthophyceae 28 

4. Discussion 30 

I: Assessment of information available on seaweed diseases worldwide and in New Zealand 30 

II: Assessment of threats by pathogens of Undaria to New Zealand native marine flora 31 

III: Future strategy for screening populations and increasing knowledge of risk posed by 

diseases/parasites/pathogens to New Zealand macroalgae and coastal communities 32 

5. Conclusions 33 

6. Acknowledgements 33 

7. References 34 

i

Executive Summary 

A detailed desk study was carried out on diseases, pathogens and parasites of Undaria and 

other macroalgae. In additional to published literature, data sources included specimens 

housed in New Zealand herbaria, and information obtained through email and personal 

contacts. An Access database was established to enter data from the relevant papers and to 

record details of the diseases, parasites and pathogens. The database contains a complete 

listing of all papers considered (927 references) of which 549 pertinent papers are included in 

the reference list in this report. 

The information on diseases of seaweeds is very patchy and the emphasis of published work 

lies in two main areas: diseases occurring in monocultures of farmed species, mainly in East 

and Southeast Asia (particularly affecting the key economic genera Porphyra, Laminaria, 

Undaria, Gracilaria, Eucheuma and Kappaphycus), and observations of certain groups of 

pathogens in particular geographic regions as a consequence of the research interests of a 

particular team or research group, leading to “pockets of information”. The amount of 

information contained in the references we investigated varied greatly between articles, 

ranging from reports of the occurrence of pathogens to multi-paper treatments of certain 

diseases. The latter are especially numerous for farmed macroalgae e.g. Pythium porphyrae, 

the agent causing the red rot disease in Porphyra species (Porphyra cultivation is a billion 

dollar industry in Asian countries). Other agents, in contrast, have only been observed once 

and often only incidentally in the course of other research. 

The only disease reported in Undaria from its introduced range is the infection of thalli with 

the pigmented endophytic brown alga Laminariocolax aecidioides, both in Spain (Veiga et al. 

1997) and in Argentina (Gauna et al. personal communication). It is not clear whether this 

endophyte originates from Japanese populations introduced with the host or from European or 

Argentinian populations respectively. Laminariocolax aecidioides is known from other, 

native European kelps such as Laminaria hyperborea in the German Bight and Norway, and 

Saccharina latissima in the Western Baltic Sea (Lein et al. 1991; Ellerstdottir & Peters 1995, 

1997; Peters & Schaffelke 1996), but it has not been reported from southern Europe. It also 

occurs in the native range of U. pinnatifida, in Japan (Yoshida & Akiyama 1978). Genetic 

studies may determine the origin of the Spanish and Argentinean populations and thus shed 

some light on whether endophytes were or can be transmitted with host sporophytes (or other 

disease agents). 

None of the known pathogens of Undaria have so far been observed in/on U. pinnatifida in 

New Zealand, however, populations of U. pinnatifida around New Zealand have not been 

screened for the presence of diseases, pathogens and parasites. Given that there is evidence 

that New Zealand has received at least 10 separate introduction events of Undaria pinnatifida 

(Uwai et al. 2006), it would be important to construct a sampling regime that reflected this 

known genetic diversity within New Zealand populations of Undaria. 

Seaweeds that are diseased are under-collected in New Zealand and, as a consequence, the 

status of knowledge about biotic diseases, pathogens and parasites is deficient: it is not 

possible to evaluate risk posed by introduced diseases, pathogens and parasites on the basis of 

current understanding of the native biota. Whilst experts in the field of algal diseases such as 

Correa (1997) stress the need for studies on the mechanisms of infection and the spread of the 

pathogens within and among host individuals, as well as on the genetics of the host-pathogen 

interaction, the basic underpinning surveys and research are required in New Zealand to 

MAF Biosecurity New Zealand Diseases, pathogens and parasites of Undaria pinnatifida • 1

document the biodiversity and distribution of diseases, pathogens and parasites within 

macroalgae. 

OVERALL OBJECTIVE: 

To determine and assess the threats that known diseases, pathogens and parasites of Undaria 

pinnatifida pose to native New Zealand macroalgae 

SPECIFIC OBJECTIVES: 

1. To undertake a review (literature, email, telephone) and map the known distribution of the 

diseases, pathogens and parasites which have been recorded to affect Undaria pinnatifida 

(and/or closely related members of the Laminariales) 

a. in its native range (Japan, Korea and the Kamchatka Peninsula of Russia), and, 

b. in its introduced range (Australia, United Kingdom, France, USA, Argentina, New 

Zealand). 

2. To undertake a review (literature, email, telephone) and map the geographic distribution of 

the status of known diseases, pathogens and parasites in macroalgae. 

3. To determine if any of the diseases, pathogens and parasites identified in specific objective 

1 are present in New Zealand. 

4. To determine if any of the diseases, pathogens and parasites identified in specific objective 

2 are present in native New Zealand macroalgae. 

2 • Diseases, pathogens and parasites of Undaria pinnatifida MAF Biosecurity New Zealand

1. Introduction 

Undaria pinnatifida is a large kelp (Laminariales, Phaeophyceae) native to the north western 

Pacific (Japan, Korea, China and the Kamchatka Peninsula of Russia) (Akiyama & Kurogi 

1982; Silva et al. 2002; Guiry & Guiry 2007). It was introduced to Europe in the 1970s 

associated with the transport of oysters from Asia (Perez et al. 1981; Bourdouresque et al. 

1985; Castric-Fay et al. 1993; Fletcher & Farrell 1998). In the 1980s Undaria was recorded in 

New Zealand (Hay & Luckens 1987; Hay 1990), Tasmania, Australia (Sanderson 1990), in 

the 1990s in Argentina (Casas & Piriz 1996), Victoria, Australia (Campbell & Burridge 

1998), and in the 2000s from California, USA (Silva et al. 2002) and Baja California, Mexico 

(Aguilar-Rosas et al. 2004). 

Since its detection in New Zealand, Undaria has spread primarily by human-mediated vectors 

such as vessel hulls and marine farming equipment. This species has the potential to displace 

native macroalgae (environmental impact), alter habitat for commercial species 

(environmental and economic impact), disrupt aquaculture activities (economic impact) and 

may affect the cultural values of particular sites. 

At present, Undaria in New Zealand has been reported from Great Barrier Island, Auckland 

(Waitemata Harbour), Coromandel, Tauranga, Gisborne, Napier, Port Taranaki, Wellington 

and the Wellington region of Cook Strait in the North Island, in the Marlborough Sounds, 

Nelson, Golden Bay, Kaikoura, Lyttelton, Akaroa, Timaru, Oamaru, Dunedin Harbour, Bluff 

in the South Island and also from Stewart Island and the Snares Islands. The potential exists 

for this species to be spread further to regions regarded as having high conservation values 

such as the other sub-Antarctic islands, Stewart Island (apart from Paterson Inlet and Oban), 

the Chatham Islands, Fiordland, Abel Tasman National Park, as well as the islands of the 

Hauraki Gulf, and offshore islands of north eastern New Zealand. A genetic study of Undaria 

populations in New Zealand has revealed multiple introductions have occurred with 10 

distinct haplotypes present in New Zealand, although only a single haplotype was found in 

current North Island populations (Uwai et al. 2006). 

As well as the direct impact this species has on indigenous biota, it also poses a potential risk 

to native New Zealand macroalgae through diseases, pathogens and parasites. Infectious 

diseases in macroalgae are caused by a wide variety of organisms ranging from viruses, 

bacteria, cyanobacteria, fungi (phycomycetes, ascomycetes, fungi imperfecti), 

heterokontophytes (oomycetes, labyrinthulids), nematodes, protozoans, through to endophytic 

and parasitic macroalgae (Chlorophyta, Phaeophyceae, Rhodophyta) (Andrews 1976; Goff 

1982; Apt 1988b; Correa 1994; Bouarab et al. 2001a; Van Etten et al. 2002). In addition, the 

grazing of macroalgae by herbivores results in vulnerability to disease/access for pathogenic 

organisms. Within its native distribution Undaria has a number of known diseases, pathogens 

and parasites (e.g. Yoshida & Akiyama 1978; Rho et al. 1993; Jiang et al. 1997). Undaria, as 

a member of the Laminariales, has a heteromorphic life history with the 2 life stages having 

entirely different morphologies – the sporophyte grows up to several metres in length whereas 

the gametophyte is microscopic growing to only several hundred microns in size. Any 

consideration of diseases in macroalgae needs to consider the vulnerability of different life 

history stages to diseases/pathogens/parasites. 


Biological invasions are understood to pose a significant threat to biodiversity, and parasites 

are considered to play a role in determining the outcomes of invasions. Transmission of 

parasites to native species from the invading species can influence the fitness of native taxa, 

mediating competitive interactions. Introduced diseases may have catastrophic impacts or 

may result in persistent and sub-lethal effects on natives and consequent impacts on 

community structure (Prenter et al. 2004). Introduced hosts may also play a role as reservoirs 

for native diseases, pathogens and parasites from which potentially deleterious “spillback” of 

infection to native hosts may occur (Prenter et al. 2004; Tompkins & Poulin 2006). Tompkins 

& Poulin (2006) observe that although many parasites are apparently lost from hosts when 

they are introduced to a new environment, the introduced hosts tend to acquire generalist 

parasites from the native biota. Impacts of disease are often dependent on the context, with 

multiple abiotic and biotic factors implicated in the emergence of parasites, invasion 

processes, and the impacts experienced by native biota (Blaustein & Kiesecker 2002). Factors 

which increase host susceptibility to infection, including a range of stressors such as habitat 

alteration and degradation, may make them more prone to introduced parasites. Artificial 

rearing and aquaculture increase the potential for disease transmission as well as increasing 

potential host susceptibility in crowded or sub-optimal growth conditions. Correa (1997) 

considers that long term strategies for disease control in macroalgal farms will only succeed if 

the genetics of disease resistance in the host and virulence in the pathogen are understood. 

The risks that diseases, pathogens and parasites of Undaria pose to the New Zealand marine 

environment have yet to be quantified. To fully understand Undaria’s impacts and to 

effectively implement control or management options, diseases, pathogens and parasites 

associated with this species as well as with other macroalgae need to be documented, both 

internationally and nationally. 

In this study the status of known diseases, pathogens and parasites of Undaria was 

determined (Objective 1) and literature was reviewed for reports of these diseases, pathogens 

and parasites in Undaria populations in New Zealand (Objective 3). Diseases, pathogens and 

parasites of other macroalgae known internationally were summarised (Objective 2), as was 

information relating to those present in macroalgal populations in New Zealand (Objective 4). 

Within a wider consideration of diseases, pathogens and parasites of macroalgae there are 

difficulties in confirming a causal role of specific organisms that have been implicated in 

disease or infection. The confirmation of Koch’s postulates is the exception rather than the rule. 

Thus, the database developed in this proposal has considered all organisms that have been 

associated with infection/disease/pathology with a clear indication of the evidence linking 

specific organisms to disease states/symptomology. 


2. Methods and Materials 

2.1. DEFINITIONS 

A disease is defined by various authors as: 

• either “… a continuing disturbance to the plant’s normal structure or function such that it 

is altered in growth rate, appearance or economic importance” (Andrews 1976), 

• “... the abnormal, injurious and continuous interference with physiological activities of the 

host" (Andrews 1979a, page 429; 1979b, page 448), 

• or a "... disturbance of the normal appearance and function of a plant” (Correa 1994) 

Diseases can have a variety of causes. This study only deals with infectious diseases, i.e. 

diseases caused by another organism (i.e. viruses, bacteria, protozoa, animals, fungi, other 

algae). This excludes diseases due to adverse abiotic conditions, i.e. physiological diseases 

caused by factors such as UV light, high or low temperature, or by dehydration (Gäumann 

1951; Andrews 1976). Exceptions are diseases caused by organisms affecting hosts weakened 

by adverse abiotic conditions. 

A pathogen is an organism that causes a disease. 

In the literature, an agent is often called a pathogen when it is found to be associated with a 

disease or aberrant appearance. However, for true pathogenicity, causality has to be 

demonstrated according to Koch’s postulates (Andrews & Goff 1984): 

1. the agent must be associated in every case with the disease under natural conditions, and 

the disease must not appear in the absence of the agent 

2. the agent must be isolated in pure culture and characterised. 

3. typical symptoms must develop when the host is inoculated with the agent under suitable 

conditions, and the appropriate control inoculations must be made concurrently. 

4. the causal agent must be re-isolated and demonstrated to be identical to the agent isolated 

originally. 

An endophyte is an "organism living within a host plant" (Greek: éndon = inside; 

phytón = plant; Womersley 1987) 

An epiphyte is an organism living on the surface of a plant (its basiphyte). Epiphyte species 

can be opportunists (i.e. grow on all available surfaces), generalist epiphytes (i.e. grow on a 

variety of algal substrates), or specialists (i.e. grow on one or a few algal surfaces). Obligate 

specialist epiphytes are restricted to the epiphyte habit and to particular hosts. In the database 

we primarily focused on obligate or specialist epiphytes. In some cases of obligate 

epiphytism, such as in Polysiphonia lanosa growing on Ascophyllum nodosum, a directional 

exchange of nutrients from basi- to epiphyte has been experimentally demonstrated (Citharel 

1972), however, such information is lacking in the majority of cases. Organisms that were 

isolated from the surface of macroalgae but can also grow on other substrates are not 

considered in this study. 

A parasite is an organism which benefits to the detriment of its host organism. Usually, this 

means a physiological dependence, e.g. parasitic algae are unpigmented and thus, as 

heterotrophic organisms, rely at least to some extent on their host for nutrition, especially 

carbohydrates (Goff 1983; Correa 1994, 1997). 


Especially in older literature the terms pathogen and parasite, or parasite and endophyte, tend 

to be used interchangeably. This should be avoided. For example, endophytic algae may but 

need not be parasitic, while not all parasites live inside the tissue of their host. Likewise, the 

term symbiosis is often used as opposite to parasitism. However, in its original definition, i.e. 

sensu de Bary (1879), a symbiosis is "...a phenomenon in which dissimilar organisms live 

together..." (de Bary 1879, cited in Paracer & Ahmadjian 2000; Goff 1983; Correa 1994), thus 

including parasitism. 

Classification system: We have based the hierarchical classification used in this study on the 

work of Cavalier-Smith (1998) with modifications adopted for the New Zealand Species 2000 

project (pers. comm. D. Gordon, NIWA). The hierarchy is provided as Appendix 1. In this 

study the term “fungus” comprises true fungi, such as Ascomycetes, but also taxa that are 

traditionally treated as fungi, but really belong to the Ochrophyta/Chromista, i.e. oomycetes, 

Labyrinthula sp., etc. 

The organisms are treated as follows: 

Section in report Kingdoms/phyla included 

Viruses Viruses, Virus-like particles (VLPs) 

Bacteria Bacteria including phyla Eubacteria, Cyanobacteria, Proteobacteria, & Mycoplasmalike 

Organisms 

Fungi Fungi, Chromista (phyla Bigyra, Sagenista) 

Animals Animalia, Protozoa 

Other algae Plantae, Chromista (phylum Ochrophyta) 

2.2. DATA SOURCES 

2.2.1. Literature Review: 

A detailed literature search was carried out by NIWA information management staff to locate 

literature on diseases, pathogens and parasites of Undaria and other macroalgae. 

The literature searching strategy and terms were 1+3 and 2+3, where 1= seaweeds, 

macroalgae, Undaria, Laminaria, Macrocystis, Laminariales, Phaeophyceae, Phaeophyta; 2= 

Rhodophyta, Chlorophyta, Phaeophyceae, Phaeophyta; 3= disease, pathogen, parasite, 

endophyte, symbiosis, ascomycetes, bacteria, fungi, cyanobacteria, bluegreen/blue-green/blue 

green alga*e, chytrid*iomycetes, labrinthulids, virus*es, nematodes, copepod*s. The 

databases searched included standard marine bibliographic sources (e.g. SCOPUS, Web of 

Science, ASFA, Google Scholar) and also web sites of marine research organisations were 

explored. The references obtained were entered into an EndNote database. 

Titles and abstracts of literature were scrutinised to determine relevance to the review and 

papers were scored (immediate acquisition, later acquisition, possible inclusion, no 

relevance). The scoring of literature was carried out by 2 people and cross–checked by a third 

to check for consistent treatment. Relevant literature was obtained, and there was an iterative 

review of key words. Additional papers to be scored and entered into the database were 

located through scrutiny of reference lists and earlier review articles. 

Translations were made of key papers in Chinese, Spanish, French and German. Generally 

papers in Japanese (and some in Chinese) included English abstracts/ summaries as well as 

captions in English for tables, and graphs. 


2.2.2. E-mail and personal contacts: 

A message about the project requesting literature and general information was sent to Algae- 

L, a bulletin-board-type forum for people interested in any aspect of algae (terrestrial, 

freshwater and marine). In addition the archives of Algae-L were searched (May 1995 - 

October 2007) for any messages containing the words “disease” (37 hits) “pathogen” (19 

hits), “parasite” (14 hits). The majority of these were found to be references to books, 

microalgae, or to be otherwise irrelevant. 

Personal contacts and/or email messages were sent to key researchers in this field including 

Professor Juan Correa (Universidad Catolica de Chile, Santiago, Chile), Mr Smith (Australian 

Centre for International Agricultural Research, Australia), Dr M. Polne-Fuller (University of 

California, Santa Barbara, USA), Dr Bruce Harger (Sunshine Marine Farms, USA), Professor 

Ma & Dr Bin Sun (Shanghai Fisheries University, China), Professor Sung Min Boo 

(Chungnam National University, Korea), Dr M. Gauna (Universidad Nacional del Sur, Bahia 

Blanca, Argentina), Dr Danilo Largo (University of San Carlos, Philippines). 

2.2.3. Herbaria: 

The collections of the herbaria holding the majority of macroalgal specimens in New Zealand 

were examined (Auckland Museum – including the Lindauer and ex-Auckland University 

collections [AK/AKU], Museum of New Zealand Te Papa Tongarewa [WELT], Landcare 

Manaaki Whenua [CHR]). In addition the NZFungi database of Landcare was searched for 

specimens of algal parasitic taxa known to be reported from New Zealand. The data are 

presented in Appendix 2. 

2.3. DATABASE 

An Access database was established to enter data from the relevant papers and to record 

details of the diseases, parasites and pathogens. The following fields were included: 

• bibliographic data (including author, year, title, book/journal/publication details, abstract, 

keywords, comments, language); 

• characteristics of the agent (including classification [Kingdom, Phylum, Class, Order, 

Family, Genus, original genus and species name, current genus and species name, species 

authority], common name, agent type, associated species/community, secondary agent); 

• characteristics of the host (including classification [Kingdom, Phylum, Class, Order, 

Family, Genus, original genus and species name, current genus and species name, species 

authority], common name, taxonomic hierarchy, with fields for notes and for comments 

on the generation of the host affected); 

• location data (including world region [based on FAO fisheries regions - Attachment 3], 

latitude, longitude, map references, country, location, depth, exposure, temperature, 

salinity, water clarity, habitat type, agent stability, timing of occurrence, as well as fields 

to record data on epidemiology, seasonality, culture information, disease control, host 

impact). 

In many cases data were not available, particularly with respect to location data and 

epidemiological information, as very little detail was provided in the original literature. 

The majority of the required fields for the database were determined at an initial stage and 

there was an on-going review of the database effectiveness with additional fields identified 

and included after the initial phase of the study. 


2.4. MAPPING 

The tender document specified a requirement to “map the known distribution of the diseases, 

pathogens and parasites which have been recorded”. Fields for longitude and latitude data 

were included in the database to enable this information to be extracted quickly from the 

database. 

3. Results 

3.1. GENERAL COMMENTS 

3.1.1. Data Sources 

Literature Review: 

The database includes a total of 927 references of which 549 pertinent papers are included in 

the reference list in this report. The Reference list also contains other literature cited in this 

report. The breakdown of papers by category is as follows: direct relevance (Laminariales), 

91; direct relevance (other algae), 363; generic/review, 70; source of additional references, 25; 

irrelevant, 292; unsourced, 86. The 292 entries considered irrelevant include, for example, 

papers dealing with epiphytes, saprobic organisms, freshwater, terrestrial and/or microalgae 

etc. Only relevant references are cited in the text of this report: the reference list provided lists 

all publications scored as ‘relevant’, ‘generic/reviews’ and ‘references only’, as well as papers 

cited in the text but not included in the database. The database contains a complete listing of 

all papers considered in relation to algal diseases. Some additional papers that were identified 

through electronic search engines were discarded based on abstract, keywords or titles. 

Electronic search engines cover mainstream journals and publications, generally from the 

1970s onwards. A number of the papers relevant to this project fell outside these parameters 

i.e. published in the early 20 th century and/or in specialist or limited edition 

publications/journals. Although we sought “grey literature” and anticipated there would be 

guides and manuals available from marine farming centres or aquaculture institutions, almost 

none of this type of literature was forthcoming. Obtaining material from some overseas 

sources took much longer than anticipated and was sometimes extremely costly. The database 

includes bibliographic information for 87 references which are categorised as “unsourced”. 

These include post-graduate theses from outside New Zealand, informal publication of 

abstracts from congresses and conferences, and grey literature which we have been unable to 

source, particularly from Asian research institutes (Japan, China, Korea). 

E-mail and personal contacts: 

There was only minor interest generated by our posting in the ALGAE-L list, and of the 14 

responses to our email, most did not provide information, but instead were interested in the 

outcome of this study and its public availability. Three of the responses provided references, 

and one directed us to another potential contact. Additional personal and targeted contacts 

yielded only a small amount of additional information, although there was interest in the 

results of this study. Initially we had intended to include data from email searches and through 

personal contacts (via phone or email) but as these were very few in number and did not 

contain new information they were not included. 

3.1.2. Mapping 

Mapping distribution information obtained through the data sources was determined to be of 

limited value. Only 72 of the 927 references (7.7%), or 192 of the more than 2300 agent 

entries in the database (~8%) included GIS compatible data (i.e. longitude and latitude) that 


could be mapped for sites where diseases were observed in seaweeds. More than 500 agents 

and 600 hosts were referred to in the relevant references reviewed, the majority of which were 

cited on a single occasion. It was concluded that mapping would not assist with visualization 

of these data. The data are summarised in Table 1 and 2 below. 

Two maps are provided illustrating the data obtained for the distribution of diseases/ 

pathogens/ parasites reported for Undaria pinnatifida, using the FAO regions map (Appendix 

3) and a separate map showing the pathogens present in the native range of U. pinnatifida 

(Appendix 4). 

Table 1. Summary of the number of records for each pest group in each algal host group. The 

numbers for the Ochrophyta exclude the Laminariales. NB. The numbers in the table do not 

relate directly to the number of references in the database, as references may contain multiple 

records. 

Viruses Bacteria Fungi Animals Other algae Total 

Rhodophyta 4 51 183 12 632 882 

Ochrophyta 95 3 120 14 84 316 

Chlorophyta 0 0 53 3 6 62 

Total 99 54 356 29 722 1260 

Table 2. Summary of the number records from each FAO region. Numbers for the Ochrophyta 

exclude the Laminariales. NB. Numbers in the table do not relate directly to the number of 

references in the database, as references may contain multiple records. Total numbers differ 

from those in Table 1 as not all references contained location information. 

Rhodophyta Ochrophyta Chlorophyta Total 

21 - Atlantic, Northwest 69 42 12 123 

27 - Atlantic, Northeast 114 99 10 223 

31 - Atlantic, Western Central 11 10 1 22 

34 - Atlantic, Eastern Central 15 6 2 23 

37 - Mediterranean and Black Sea 30 8 2 40 

41 - Atlantic, Southwest 25 8 0 33 

47 - Atlantic, Southeast 50 1 0 51 

48 - Atlantic, Antarctic 10 7 0 17 

51 - Indian Ocean, Western 10 4 11 25 

57 - Indian Ocean, Eastern 38 12 9 59 

58 - Indian Ocean, Antarctic 0 1 0 1 

61 - Pacific, Northwest 105 14 1 120 

67 - Pacific, Northeast 117 7 3 127 

71 - Pacific, Western Central 42 3 0 45 

77 - Pacific, Eastern Central 114 12 3 129 

81 - Pacific, Southwest 19 30 2 51 

87 - Pacific, Southeast 36 19 0 55 

Total 805 283 56 1144 

3.2. DESCRIPTION OF KNOWN PATHOGEN-HOST RELATIONSHIPS 

3.2.1. Undaria 

A summary of the records for each pest group in each algal host group, and number records 

from each FAO region are presented in Tables 3 & 4 respectively for members of the 

Laminariales, including Undaria. (Note – all papers refer to the sporophyte phase and not to 

the gametophyte phase of Undaria). 




3.2.2. Known pathogens in native range 

Viruses 

There are no virus diseases known from Undaria pinnatifida or other Undaria species. 

Bacteria 

Gram-negative bacteria such as Aeromonas, Flavobacterium, Moraxella, Pseudomonas, and 

Vibrio are associated with the "spot-rotting" disease ("Anaaki sho"; Kimura et al. 1976) and 

the so-called "shot hole disease" (Tsukidate 1991) in Japanese Undaria. Severe outbreaks of 

infections with Vibrio especially affect young sporophytes ("sporelings") of U. pinnatifida 

(Anon. 1991). The "shot hole disease" is characterised by brown spots appearing on the 

thallus blade near the midrib which subsequently fuse together and spread onto the pinnate 

part of the blade (Tsukidate 1991). 

The "green spot disease/rot" caused by unspecified bacteria in Japan (Ishikawa & Saga 1989; 

Vairappan et al. 2001) and South Korea (Kang 1982) manifests with similar symptoms, first 

as green spots of rotting host tissue that result in small holes with green margins, and in the 

distal parts of the frond these enlarge and finally coalesce, accelerating the decay of the frond 

(Kang 1982). Japanese Undaria is furthermore infected by an unspecified bacterium causing 

the "yellow hole disease" (Ishikawa & Saga 1989; Vairappan et al. 2001) and “spot-rotting” 

disease (Kito et al. 1976). 

Bacteria enter the thallus of U. pinnatifida through openings like dead mucilage channels, and 

digest cells and cell walls in the medulla. Cells of the cortex and meristoderm show ultrastructural 

damage (e.g. vacuolation of the dictyosome). When the host cells die, the disease 

symptoms become macroscopically visible (Kito et al. 1976). 

In China, the bacterium Halomonas venusta has been identified as a causative agent in “spot 

decay” (Ma et al. 1997a, b, 1998), and Vibrio logei in “green decay diseases” (Jiang et al. 

1997) of U. pinnatifida. 

Animals 

Some small crustacean species are associated with diseases in Undaria: The “pin hole 

disease” is caused by frond-mining nauplii of harpacticoid copepoda in Undaria from Japan 

(Anon. 1991) and South Korea (Tsukidate 1991), e.g. by species such as Amenophia 

orientalis, Parathalestris infestus, Scutellidium sp. (Ho & Hong 1988; Park et al. 1990; Anon. 

1991; Rho et al. 1993; Shimono et al. 2004) and Thalestris sp. (Kang 1982). 

Ceinina japonica, a gammeride amphipod from South Korea, invades the midrib of U. 

pinnatifida through the holdfast and bores a tunnel which may cause the longitudinal 

separation of the entire frond through the midrib. In heavily damaged thalli the holdfast may 

depart from the substrate (Kang 1982). 

Fungi 

A fungal infection occurs in Undaria from Japan, the so-called “chytrid blight” (Tsukidate 

1991). The name implies that this disease is caused by a true fungus of the class 

Chytridiomycetes, however, the culprit is an oomycete of the genus Olpidiopsis (Akiyama 

1977a). The fungus affects sporophytes, where it grows inside host cells, killing them slowly. 

Infected thalli gradually loose colour and disintegrate, juvenile thalli suffer severe damage or 

eventually die. 


Other algae 

Laminariocolax aecidioides is an endophytic brown alga infecting farmed U. pinnatifida in 

Japan (Akiyama 1977b; Yoshida & Akiyama 1978; Veiga et al. 1997). Infections result in 

host thalli becoming thicker and stiffer, lowering their market value (Yoshida & Akiyama 

1978). 

3.2.3. Known pathogens in the introduced range other than New Zealand (Australia, UK, 

France, Spain, USA (west coast), Argentina, Mexico, Taiwan) 

The endophyte Laminariocolax aecidioides (as Gononema aecidioides) has been found in 

farmed Undaria pinnatifida thalli from Spain (Veiga et al. 1997) and has also been found in 

Undaria in Argentina (Gauna et al. pers. comm.). 

3.2.4. Occurrence of known pathogens in New Zealand 

Even though members of the genus Laminariocolax occur in New Zealand kelps, none have 

so far been observed in Undaria pinnatifida. Instead, in New Zealand U. pinnatifida hosts 

another endophyte, Microspongium tenuissimum, which is also found in Ecklonia radiata and 

various red algae. The infection of U. pinnatifida with M. tenuissimum was not associated 

with obvious macroscopic disease symptoms (Heesch 2005). 

3.3. LAMINARIALES 

3.3.1. Known pathogens worldwide 

Viruses 

There are no viral diseases reported from members of the Laminariales outside New Zealand 

(see 3.2.2). 

Bacteria 

Most bacteria affecting kelps belong to the phylum Proteobacteria. Pathogenic species of 

Alteromonas, Pseudoalteromonas, Pseudomonas and Vibrio have been recorded from 

Saccharina japonica in China (e.g. Tang et al. 2001; Liu et al. 2002; Wang et al. 2006) and 

Japan (e.g. Ezura et al. 1990; Yamada et al. 1990; Sawabe et al. 1998; Sawabe et al. 2000a, b; 

Narita et al. 2001; Vairappan et al. 2001) resulting in holes and lesions on thalli and 

eventually “rot disease”. Some proteobacteria indirectly affect gametophytes and young 

sporophytes in culture when red spot disease of the culture bed (i.e. the culture ropes) causes 

the young Saccharina japonica to detach from infected ropes (e.g. Ezura et al. 1988; Yumoto 

et al. 1989a, b). Alteromonas sp. and Vibrio sp. are also associated with lesions and thallus 

bleaching of Saccharina ochotensis and S. religiosa in Japan (Vairappan et al. 2001). A 

species of Acinetobacter causes “white rot” in Nereocystis luetkeana resulting in rot of stipes 

and pneumatocysts, which collapse and become covered in white slime within 7-10 days 

(Andrews 1977). 

In China, both the gametophytes and sporophytes of Saccharina japonica are prone to 

“malformation disease” caused by the firmicute Macrococcus sp. (Anon. 1989). 

Unspecified bacteria have been reported as pathogens in Macrocystis pyrifera, Pelagophycus 

porra and Egregia laevigata in America (Brandt 1923), Saccharina japonica in China (Wu et 

al. 1983; Ding 1992; Yang et al. 2001; Huang et al. 2002a, b). The “black rot” of Macrocystis 

pyrifera in California is assumed to be caused by a unidentified parasitic microorganism 

invading already damaged host thalli (Rheinheimer 1992). 


A mycoplasma-like organism (MLO) causes the "twisted frond disease" or “coiling-stunt 

disease” in Saccharina japonica from China (e.g. Wang et al. 1983; Wu et al. 1983; 

Tsukidate 1991). 

Animals 

A number of amphipods are known to bore in kelp stipes and hollow them, causing 

considerable damage which may eventually lead to the death of the host. In Alaska and 

California, Peramphithoe stypotrupetes infests stipes of Laminaria setchellii damaged by 

gastropod grazing (Chess 1993). Also in California and Alaska, it occurs in Saccharina 

dentigera, and in southern California it is found in Eisenia arborea and Pterygophora 

californica (Conlan & Chess 1992), while Californian Macrocystis pyrifera populations are 

infested by the related amphipod P. humeralis (Chess 1993). In Ireland, Alaria esculenta is 

inhabited by Amphitholina cuniculus (Myers 1974; Chess 1993). In Japan, Saccharina 

japonica is similarly affected by Ceinina japonica (Akaike et al. 2002). 

Fungi 

The ascomycete Phycomelaina laminariae causes the “stipe blotch disease” in laminarian 

species from the north-western and north-eastern Atlantic. Its hyphae penetrate the surface of 

Alaria esculenta, Saccharina latissima, S. longicruris and Laminaria digitata, leading to 

necrotic tissue and reduced overall performance of the host thalli (Sutherland 1915b, c; 

Kohlmeyer 1968; Kohlmeyer 1979; Schatz et al. 1979; Schatz 1980, 1983, 1984a, c; Goff & 

Glasgow 1980; Porter & Farnham 1986a). 

Several other ascomycete fungi attack members of the Laminariales: Pontogeneia erikae is a 

parasite in Egregia menziesii from California (Kohlmeyer & Demoulin 1981), Sigmoidea 

marina causes lesions in the surface of Saccharina latissima from Britain (Haythorn et al. 

1980), Ophiobolus laminariae causes blackened patches on the stipes of Laminaria digitata in 

Scotland (Sutherland 1915c), and in California Asteromyces cruciatus has been reported from 

Egregia menziesii, however their relationship is uncertain (Nolan 1972). 

Oomycetes have also been reported from members of the Laminariales in the north-western 

and north-eastern Atlantic: Petersenia sp. causes damage to the stipes of Laminaria digitata, 

Laminaria sp. and Saccharina longicrucis (Kohlmeyer 1968) and Pleotrachelus minutus 

infects the apical hairs of Chorda filum in Sweden (Aleem 1952a). 

In France Labyrinthomyxa sauvageaui infects Laminaria ochroleuca (Duboscq 1921). An 

unknown hyphomycete causes contortion of the blade and blackening of the stipe in 

Laminaria digitata in Maine, USA (Kohlmeyer 1968) and in Russia an undetermined fungus 

has been isolated from farmed populations of Saccharina japonica (Zvereva 1998). 

Other algae 

Green algae are occasionally observed growing in kelps, however very little information is 

available on their impact on the host species. Acrochaete repens, for example, grows in 

Chorda filum from the North American east coast (O’Kelly et al. 2004), Canada (South 1968) 

and from Denmark, Ireland and the Isle of Man in the north eastern Atlantic (South 1968; 

Nielsen 1979). The related species A. geniculata infects kelps along the North American 

Pacific coast, such as Egregia menziesii, Cymathere triplicata, Laminaria sinclairii, 

Saccharina dentigera and Dictyoneurum californicum (O’Kelly 1983). Egregia menziesii 

from British Columbia also hosts another Acrochaete species, A. apiculata (C. O’Kelly, pers. 

com.). 

The green endophyte Bolbocoleon piliferum is found on the east and west coast of the USA, 

and eastern Canada, growing in the kelps Alaria marginata, Chorda filum, Cymathere 


triplicata and Pleurophycus gardneri (South 1968; O’Kelly et al. 2004). It is also recorded in 

Chorda filum from Denmark, Wales, Ireland and the Isle of Man (South 1968; Nielsen 1979) 

and in Laminaria hyperborea from Denmark (Nielsen 1979). Another green endophyte 

Entocladia viridis is also known from several countries in the north-eastern and north-western 

Atlantic, growing in Laminaria digitata and Saccharina latissima (Nielsen 1979). In Chile, 

another green endophyte, reported as Sporocladopsis novae-zelandiae grows in Lessonia 

nigrescens (Correa & Martinez 1996). 

Pigmented endophytic brown algae are very common in kelps (Lein et al. 1991; Ellertsdottir 

& Peters 1995). Their presence is often associated with brown spots (“dark-spot disease”, 

Lein et al. 1991), hyperplasia leading to warts or galls, and, in severe cases, thallus 

deformations (Andrews 1977; Apt 1988b). Traditionally, kelp endophytes have been 

classified as Streblonema species (Goff & Glasgow 1980), for example, the endophytes that 

affect Saccharina sessilis, Alaria tenuifolia, Laminaria setchellii and Nereocystis luetkeana 

along the North American west coast (Setchell & Gardner 1922). However, genetically, most 

kelp endophytes belong to the genera Laminariocolax and Microspongium. 

North Atlantic kelp populations are infected by two species of Laminariocolax: L. 

tomentosoides and L. aecidioides. The former is mainly found in Laminaria digitata, but 

occasionally also in L. hyperborea, Saccharina latissima and Alaria sp. (Lund 1959; Pedersen 

1976; Ellertsdottir & Peters 1997; Burkhardt & Peters 1998; Küpper et al. 2002). 

Laminariocolax tomentosoides ssp. deformans is associated with galls and stipe coiling in 

Laminaria digitata from France (Dangeard 1931b; Peters 2003). 

Laminariocolax aecidioides is found throughout the Northern Hemisphere. In the North 

Atlantic, it has been observed in Laminaria hyperborea and Saccharina latissima from 

Germany, France and Denmark (e.g. Peters & Ellertsdottir 1996; Burkhardt & Peters 1998; 

Heesch & Peters 1999; Peters 2003), in S. groenlandica, Laminaria sp. and S. longicruris 

from Greenland (Pedersen 1981), and on the North American east coast in Laminaria digitata 

(Peters 2003). In the North Pacific, it infects not only U. pinnatifida, but also Costaria sp. 

from Japan, and is furthermore known from Californian Hedophyllum sp. populations 

(Yoshida & Akiyama 1978). 

Southern hemisphere kelp populations are infected by two other members of the genus 

Laminariocolax, L. macrocystis and L. eckloniae. The former endophyte grows in 

Macrocystis pyrifera from Chile, the latter in Ecklonia maxima from South Africa (Peters 

1991; Burkhardt & Peters 1998). Heesch (2005) considers L. macrocystis and L. eckloniae to 

be synonymous. 

Laminariocolax sp. is recorded from the North Atlantic in Laminaria hyperborea (Lein et al. 

1991; Peters & Schaffelke 1996; Ellertsdottir & Peters 1997) and the Pacific in Macrocystis 

integrifolia, Saccharina latissima and Nereocystis luetkeana (Andrews 1977; Apt 1988a). 

Another endophytic brown alga, Laminarionema elsbetiae, occurs in Japanese Saccharina 

japonica as well as in the German kelps S. latissima and Laminaria digitata (Kawai & 

Tokuyama 1995; Peters & Ellertsdottir & 1996; Ellertsdottir & Peters 1997; Peters & 

Burkhardt 1998; Heesch & Peters 1999; Peters 2003). 

The genus Microspongium is occasionally found as endophyte in kelps. On the east coast of 

North America, Alaria esculenta and Saccharina longicruris are infected by Microspongium 

alariae, with symptoms ranging from dark spots to twisted stipes (Peters 2003). 


Gametophytes of kelps themselves colonise other algae as endophytes. A genetic study has 

revealed that endophytic brown algae growing in Lessoniopsis littoralis from British 

Columbia, Canada, are gametophytes of other kelps growing near the host, i.e. of Alaria sp., 

Macrocystis integrifolia and Nereocystis luetkeana (Lane & Saunders 2005). 

The ectocarpalean endophytes Phaeostroma parasiticum and Dermatocelis laminariae occur 

in Saccharina latissima and Laminaria sp. respectively in Greenland (Pedersen 1976). In 

Germany, unspecified ectocarpalean endophytes are reported to infect up to 85% of their 

hosts, Laminaria saccharina, L. digitata and L. hyperborea (Ellertsdottir & Peters 1995). 

An obligate epiphyte, Porphyra moriensis, infests Chorda filum in Japan (Notoya & 

Miyashita 1999). 


In northern New Zealand, mass diebacks of Ecklonia radiata where reported in the mid 1990s 

(e.g. Cole & Babcock 1996). Subsequent research indicates that the diebacks are caused by 

primary and secondary agents: E. radiata is affected by the amphipod Orchomenella aahu, 

which burrows into the stipes of the host and hollows them out, thus accelerating death of the 

fronds. The simultaneously occurring bleaching of the fronds is probably due to a secondary 

infection with a virus (Haggitt & Babcock 2003) and the diebacks have been associated with 

both virus-like particles (VLPs) and a poty virus (Easton 1995, Easton et al. 1997). 

Three species of pigmented endophytic brown algae infect kelps from New Zealand (Heesch 

2005): Laminariocolax macrocystis (which in this treatment includes L. eckloniae) is 

associated with galls and thallus deformations in Macrocystis pyrifera (North and South 

Islands) and Ecklonia radiata from the North, South and Chatham Islands. Additionally, E. 

radiata hosts Microspongium tenuissimum (which includes M. radians), an endophyte mostly 

observed in red algae. The third endophyte, an undescribed ectocarpalean species so far only 

known from New Zealand, was found in a gall on Lessonia tholiformis from the Chatham 

Islands (Heesch 2005). 

An unidentified green endophyte (probably a species belonging to the genus Acrochaete, 

O’Kelly pers. com.) was frequently observed in stipes of Macrocystis pyrifera along the 

Otago coast (Heesch, unpublished data). 

3.4. BROWN ALGAE OTHER THAN LAMINARIALES 


Viruses 

Virus-like particles (VLPs) in several members of the order Ectocarpales (Phaeophyceae), e.g. 

Ectocarpus and Pylaiella species (Markey 1974; Dodds 1979) have subsequently been 

identified as DNA viruses. Viruses have been found in Ectocarpus siliculosus (Ectocarpus 

siliculosus virus EsV), E. fasciatus (EfasV), Feldmannia irregularis (FirrV), F. simplex 

(FlexV), an unidentified Feldmannia species (FsV), Myriotricha clavaeformis (MclaV), 

Pylaiella littoralis (PlitV), Hincksia hincksiae (HincV), and also in Kuckuckia sp. and 

Leptonematella fasciata. The viruses infect naked spores, leading to a latent infection in 

vegetative thalli. Upon maturation, reproductive organs develop abnormally producing new 

virus particles instead of spores (Clitheroe & Evans 1974; Müller et al. 1990, 1996 (a, b, c), 

1998, 2000; Müller 1991a, b; Henry & Meints 1992, 1994; Müller & Stache 1992; Lanka et 

al. 1993; Müller & Frenzer 1993; Friess-Klebl et al. 1994; Kuhlenkamp & Müller 1994; 


Parodi & Müller 1994; Robledo et al. 1994; Bräutigam et al. 1995; Krueger et al. 1996; 

Müller & Schmid 1996; Sengco et al. 1996; Del Campo et al. 1997; Kapp et al. 1997; Maier 

et al. 1997, 1998, 2002; Kapp 1998; Lee et al. 1995, 1998; Maier & Müller 1998; Wolf et al. 

1998, 2000; Van Etten & Meints 1999; Delaroque et al. 2000a, b, 2003; Dixon et al. 2000; 

Van Etten et al. 2002; Chen et al. 2005; Dunigan et al. 2006). EsV and EfasV are known from 

host populations world-wide (Müller & Stache 1992). 

In Botrytella micromora, virus-like particles (VLPs) are associated with tissue necroses and 

zoospores that fail to germinate and lyse instead (Oliveira & Bisalputra 1978; Henry & 

Meints 1994). Likewise, VLPs affect zoospore germination in Halosiphon tomentosus, while 

thalli of Streblonema sp. containing VLPs in their vegetative cells do not appear to be 

negatively affected (Toth & Wilce 1972; LaClaire & West 1977; Dodds 1979; Henry & 

Meints 1992, 1994; Müller et al. 1998). 

Bacteria 

Bacteria associated with galls and thallus deformations occur in Fucus vesiculosus, F. spiralis 

and Saccorhiza polyschides (Cantacuzene 1930; Apt 1988b; Rheinheimer 1992). In France a 

proteobacterium infects Cystoseira nodicaulis causing damage to the thallus (Pellegrini & 

Pellegrini 1982), and in Russia’s Kurile Islands, Pseudoalteromonas issachenkonii degrades 

the thallus of its host Fucus evanescens (Ivanova et al. 2002). 

Animals 

Protozoan pathogens are reported from members of the Ectocarpales and the Fucales. The 

infection of Ectocarpus siliculosus from Chile with the plasmodiophorid Maulinia ectocarpii 

results in the sterility of the host sporangia (Maier et al. 2000). Also in Chile, another 

plasmodiophorid infects Durvillaea antarctica causing galls and internal hypertrophy of cells 

(Aguilera et al. 1988). An unspecified brown alga is also reported to be infected by the 

plasmodiophorid Phagomyxa algarum (Porter & Farnham 1986a). Amoeba are found in 

Sargassum muticum and in British Fucus serratus, the latter affected by the species 

Trichosphaerium sieboldi. The amoeba digest the walls and invade the cytoplasm of the host 

cells leading to a dissociation of the host tissue (Polne-Fuller & Gibor 1987; Rogerson et al. 

1998). 

In Japan, the harpacticoid copepods Dactylopusioides fodiens and D. macrolabris feed on the 

internal tissue of Dictyota dichotoma and live in the resulting galleries; Dactylopusioides 

fodiens also parasitises Pachydictyon coriaceum (Shimono et al. 2003, 2004). Copepoda are 

furthermore associated with galls in Desmarestia aculeata from Scotland (Barton 1892). 

Nematodes of the genus Halenchus are found in galls on members of the Fucales: H. fucicola 

affects Ascophyllum nodosum while H. dumnonicus inhabits Fucus vesiculosus and F. 

serratus (Barton 1892; Coles 1958; Tokida 1958; Apt 1988b). 

Fungi 

There is a large body of literature relating to fungi and seaweeds, however many contain little 

information about the fungal parasite’s impact on the algal host. 

Ascomycete fungi frequently form galls in members of the Fucales, e.g. Massarina 

cystophorae in Cystophora retroflexa and C. subfarcinata. Members of the ascomycete genus 

Haloguignardia are widespread and occur in a range of hosts, e.g. Haloguignardia irritans in 

Cystoseira osmundea, Cystoseira sp., Halidrys dioica and Halidrys sp.; Haloguignardia sp. in 

Cystoseira balearica, Cystoseira sp., Halydris dioica and various Sargassum species (S. 

decipiens, S. fallax, S. fluitans, S. natans and S. sinclairii) (Estee 1913; Cribb & Herbert 1954; 

Tokida 1958; Kohlmeyer 1979; Apt 1988c; Alongi et al. 1999). Haloguignardia cystoseirae 

infects Cystoseria spp. in the Mediterranean (Kohlmeyer & Demoulin 1981; Alongi et al. 

1999), whereas Haloguignardia tumefaciens, H. oceanica, H. decidua and H. longispora 


infect Sargassum spp. in Australia, Japan, America and the Sargasso Sea (e.g. Ferdinandsen 

& Winge 1920; Tokida 1958; Cribb & Cribb 1960; Kohlmeyer 1971, 1972; Alongi et al. 

1999). A secondary agent, the hyperparasite Sphaceloma cecidii has also been reported from 

Cystoseira sp., Sargassum sp. and Halidrys sp., where its infection is restricted to areas of the 

host already affected by Haloguignardia (Kohlmeyer 1979). 

Further members of the ascomycetes that affect marine algae include Thalassoascus 

treboubovii, recorded from Cutleria chilosa, C. multifida, Cystoseira sp. and Zanardinia typus 

(Ollivier 1929; Kohlmeyer 1979), Lindra thalassiae from Sargassum spp. (Meyers 1969; 

Kohlmeyer 1979; Raghukumar et al. 1992), Chadefaudia gymnogongri from Xiphophora 

chondrophylla (Kohlmeyer 1973a), Orcadia ascophylli and Trailia ascophylli from 

Ascophyllum nodosum (Sutherland 1915c), and Asteromyces cruciatus from Cystoseira 

osmundacea (Nolan 1972). Ascomycetes reported from Fucus spp. include Cephalosporium 

sp., Sigmoidea marina, Didymella fucicola, Orcadia ascophylli and Trailia ascophylli 

(Sutherland 1915c; Kohlmeyer 1968; Andrews 1977; Haythorn et al. 1980; Miller & Whitney 

1981; Schatz 1984a). Pelvetia canaliculata is also infected by a range of ascomycete fungi, 

including Didymella fucicola, Orcadia ascophylli, Dothidella pelvetiae, Pharcidia pelvetiae, 

Pleospora pelvetiae and Stigmatea pelvetiae (Sutherland 1915a). Additionally, 

Scolecobasidium salinum degrades alginates of brown algae (Moen et al. 1995). 

In the Archemycota, Chytridium polysiphoniae, C. megastomum and Olpidium sphacellarum 

have been reported from Sphacelaria cirrosa, Sphacelaria sp., Striaria attenuata and 

Pylaiella littoralis, disintegrating cell contents (Sparrow 1934, 1936; Raghukumar 1987b; 

Hyde et al. 1998; Küpper & Müller 1999; Müller et al. 1999). 

The term mycophycobiosis was created for obligate symbioses between algae and fungi 

which are without a detrimental effect for both symbionts, and in which, unlike in lichens, the 

alga is the partner that provides the structure. An example is Mycophycias ascophylli growing 

in Ascophyllum nodosum and in Pelvetia canaliculata in the Northwest Atlantic (e.g. 

Kohlmeyer & Kohlmeyer 1972; Miller & Whitney 1981; Porter & Farnham 1986a; Kingham 

& Evans 1986; Stanley 1992; Deckert & Garbary 2005a). The symbiosis between A. 

nodosum, M. ascophylli and Polysiphonia lanosa has been intensely studied (e.g. Garbary & 

London 1995; Garbary & MacDonald 1995; Deckert & Garbary 2005b; Garbary et al. 2005). 

In the pseudofungi, oomycetes are also common pathogens of seaweeds. In particular they are 

found in members of the Ectocarpales: Pylaiella littoralis, Ectocarpus siliculosus, Striaria 

attenuata and Hincksia spp. are infected by species of Eurychasma, Anisolpidium, 

Pleotrachelus, Petersenia and Olpidiopsis (Sparrow 1934, 1936; Karling 1943; Aleem 

1950a,d, 1952a; Küpper & Müller 1999; Müller et al. 1999; West et al. 2006). Some members 

of the Sphacelariales are also affected by oomycetes (Aleem 1952a). 

In the Sagenista, the Labyrinthulomycetes include two groups that are frequently associated 

with seaweeds; the thraustochytrids and the labyrinthulids. From the former, Aplanochytrium 

spp. occur as endophytes in Sargassum spp. and Padina atillarum (Raghukumar et al. 1992; 

Sathe-Pathak et al. 1993; Ulken et al. 1985; Raghukumar 2002); in the latter Labyrinthula sp. 

is reported from Lobophora variegata (Raghukumar 1987b). 

Other algae 

A few members of the Rhodophyta occur as epi- or endophytes of brown algal hosts. 

Polysiphonia lanosa is an obligate epiphyte on Ascophyllum nodosum (Turner & Evans 1977; 

Garbary et al. 1991; Cardinal & Lesage 1992; Lining & Garbary 1992; Garbary & London 

1995), which has occasionally also been observed on Fucus vesiculosus (Pearson & Evans 

1990; Rindi & Guiry 2004). It is often found on damaged host fronds (Lobban & Baxter 

1983; Rindi & Guiry 2004), deeply penetrating the host with its rhizoids (Rawlence 1972; 


Rawlence & Taylor 1972; Garbary et al. 2005). There is some evidence suggesting a transfer 

of substances occurs between the two symbionts (Citharel 1972; Penot 1974), while other 

studies doubt the translocation of synthetates from basiphyte to epiphyte (Turner & Evans 

1977; Harlin & Craigie 1975). Colacodictyon reticulatum is a small endophytic red alga 

growing in Desmarestia ligulata, and Haplodasya urceolata endophytises Cystophora 

retroflexa (Kylin 1956). 

Brown algae occur as endophytes of other brown algae. The endophytic brown algae 

Herponema valiantei and Streblonemopsis irritans are associated with galls in Cystoseira 

tamariscifolia and C. zosteroides, respectively (Apt 1988a, 1988b). Pedersen (1976) reports 

Herponema desmarestiae and Streblonema fasciculatum from Desmarestia viridis and 

Eudesme virescens respectively. The endophytic brown algae Microspongium alariae and 

Myriactula clandestina occur in Fucus vesiculosus from Finland and Greenland (Pedersen 

1976; Peters 2003). In California, Desmarestia ligulata is affected by Streblonema 

transfixum, and S. penetrale penetrates the stipe of Hesperophycus californicus (Setchell & 

Gardner 1922). Laminariocolax sp. occurs in Chordaria flagelliformis in Greenland (Pedersen 

1976) and Laminariocolax aecidioides in Sphacelaria arctica from multiple sites in the North 

Atlantic (Yoshida & Akiyama 1978). Notheia anomala is a hemi-parasitic brown alga 

occurring in Australia and New Zealand (Adams 1994). In Australia it infects Hormosira 

banksii and occasionally also Xiphophora chondrophylla (Gibson & Clayton 1987; Raven et 

al. 1995). 

Some small members of the Ectocarpales are on the border between epi- and endophytism. 

For example, Elachista fucicola is an obligate epiphyte of Fucus vesiculosus (Rindi & Guiry 

2004), but it also grows on Ascophyllum nodosum penetrating the host surface with its 

rhizoids and leading the host to form a tissue callus around the penetrating filaments (Deckert 

& Garbary 2005a, b). Filaments of Trachynema groenlandicum grow in the loosely organised 

cortex of Chordaria linearis in southern South America, but do not penetrate into the compact 

subcortex or medulla of the host (Peters 1992). Gononema pectinatum was isolated from a 

culture of Dictyosiphon hirsutum from Chile, however, the origin of the contaminant (epi- or 

endophytic) was not determined (Burkhardt & Peters 1998). 

Three endophytic brown algae have been reported from the Antarctic Peninsula: 

Laminariocolax eckloniae in Himanthothallus grandifolius, Geminocarpus austro-georgiae in 

Desmarestia menziesii, and Ascoseirophila violodora in Ascoseira mirabilis (Peters 2003). In 

addition, Antarctic Adenocystis utricularis specimens are epiphytised by Austrofilum 

incommodum, a small phaeophyte that is anchored in its host with endophytic filaments 

(Müller et al. 1992; Peters 2003). 

Antarctic Ascoseira mirabilis furthermore hosts the unicellular endophytic green alga 

Chlorochytrium sp. (Peters 2003) which is, like Codiolum sp., considered to be the 

endophytic sporophyte of an Acrosiphonia species. Codiolum petrocelidis, for example, 

grows in the crustose brown alga Ralfsia pacifica from the Pacific coast of Canada (Sussmann 

& DeWreede 2002). Chlorochytrium dermatocolax has been recorded from the North Atlantic 

and the Pacific coast of North America in Sargassum muticum and Sphacelaria spp. (Lund 

1959; Pedersen 1976; Polne-Fuller 1987). 

A filamentous green alga, Acrochaete repens, grows endophytically in Fucus serratus from 

the German Bight, Denmark and the Channel Islands (Kremer 1975; Nielsen 1979) and F. 

vesiculosus from Denmark (Nielsen 1979). The closely related Entocladia species E. viridis 

and E. wittrockii grow endophytically in Desmarestia aculeata, Dictyota dichotoma and 

Elasticha fucicola from locations in the Pacific, Mediterranean and North Atlantic (Nielsen 

1979; O’Kelly 1981). 


Navicula endophytica is a diatom living in the intercellular mucilage of receptacles of Fucales 

from the northern hemisphere. It has been reported from species such as Ascophyllum 

nodosum, Fucus vesiculosus, F. serratus, F. spiralis, F. ceranoides, F. evanescens, 

Furcellaria lumbricalis and Pelvetia canaliculata from Great Britain and from Norway 

(Wardlaw & Boney 1984, Armstrong et al. 2000). 


The DNA virus EsV was first isolated from specimens of Ectocarpus siliculosus from New 

Zealand infecting gametangia and sporangia (Müller et al. 1990) and has subsequently been 

found in host populations around the world (Müller & Stache 1992) giving rise to a large 

body of literature on aspects of this virus and its hosts (e.g. Sengco et al. 1996; Kapp et al. 

1997). 

The ascomycete fungus Haloguignardia tumefaciens has been reported parasitizing 

Sargassum sinclairii from Wellington and the west coast of the South Island (Cribb & Cribb 

1960; Kohlmeyer & Demoulin 1981). Also from Wellington, thraustochytrids of either the 

genus Thraustochytium or Schizochytrium have been isolated from drift Zonaria 

aureomarginata, Durvillaea antarctica and Marginariella boryana (Serena Cox, pers 

comm.). Karling (1968) isolated Schizochytrium aggregatum from algal debris, which 

potentially included brown algae. 

Pleurostichidium falkenbergii is an obligate epiphytic red alga on Xiphophora chondrophylla 

from northern New Zealand (Bay of Islands, Three Kings and North Cape) (Heydrich 1893; 

Kylin 1956; Phillips 2000). 

In addition to Notheia anomala partially-parasitising Hormosira banksii (Adams 1994) 

another parasitic brown alga occurs in New Zealand: Herpodiscus durvillaeae, which is 

restricted to New Zealand populations of Durvillaea antarctica. It grows epi-endophytically 

in its host and, in its emergent phase, leads to an erosion of the host surface, which may result 

in the eventual loss of the host phylloid (South 1974; Hay 1978; Peters 1990; Heesch 2005). 

An as yet undescribed pigmented endophytic ectocarpalean brown alga is associated with 

galls or pale spots on Durvillaea antarctica, D. willana, Marginariella urvilleana, and 

Xiphophora gladiata (Heesch 2005). 

3.5. RED ALGAE 


Viruses 

Virus-like particles have been observed in the single-celled Porphyridium purpureum 

(Chapman & Lang 1973), in Gracilaria conferta and in G. epihippisora from the 

Mediterranean Sea (Weinberger et al. 1994), as well as in Audouinella saviana from the east 

coast of USA (Pueschel 1995). 

Bacteria 

Bacteria are associated with tumour-like growth occurring on the fronds of Chondracanthus 

teedii (Tsekos 1982). Galls and proliferating tissue associated with bacteria are furthermore 

found in Acrochaetium species, Ahnfeltia plicata, Bonnemaisonia asparagoides, Ceramium 

virgatum, Chondracanthus teedii, Chondrus crispus, Curdiea angustata, Cystoclonium 

purpureum, Delesseria sanguinea, Dumontia contorta, Grateloupia filicina, Palmaria 

palmata, Plocamium cartilagineum, Polyneuropsis stolonifera, Prionitis decipiens, P. 


filiformis, P. lanceolata, Pterocladiella capillacea and Schizymenia dubyi (Cantacuzene 1930; 

Apt 1988b; Apt & Gibor 1989; Rheinheimer 1992; Ashen & Goff 1996, 1998, 2000). 

A number of bacterial diseases of Porphyra have been reported, particularly in relation to 

farmed Porphyra, including “green spot rotting-like deterioration” (Ryokuhan-byo) of 

Porphyra yezoensis (Nakao et al. 1972), "filament bacterial felt" disease (agent not specified) 

(Song et al. 1993), and "white wasting disease"/ white spot"/ "Gijishirogusare-sho" (Tsukidate 

1971, 1977). Tsukidate (1983) examined the symbiotic relationship between Porphyra species 

and attached bacteria that occurred in conjunction with white rot, the disease which has 

caused the most serious damage to the Porphyra cultivation industry in Japan. Anaaki-disease 

causes severe damage to the red alga Porphyra yezoensis; Hayashi et al. (1984) identified the 

agent as Vibrio fischeri and reported on how it attaches to host thalli (Porphyra sp.), digests 

host cells and makes holes in the thalli. Sunairi et al. (1995) reported Flavobacterium sp. to 

be the causative agent of Anaaki-disease, as a result of several repeated single-colony 

isolations and infection experiments. In order to ascertain the role of bacteria in the process of 

rotting or decaying of cultured laver, Fujita et al. (1972) examined 24 strains of bacteria 

isolated from diseased fronds of Porphyra yezoensis, including species of Pseudomonas, 

Vibrio, Beneckea. 

Weinberger et al. (1994) quantified the bacterial epiphytes of Gracilaria conferta and found 

that saprophytic bacteria reached 350 times and agar degraders 25,000 times higher numbers 

per gram of wet weight on tissues infected with the “white tips disease”, as compared to 

healthy tissues. Jaffray & Coyne (1996) developed an in situ assay to detect bacterial 

pathogens of the red alga Gracilaria gracilis responsible for causing lesions, thallus 

bleaching, and Jaffray et al. (1997) examined bacterial epiphytes on Gracilaria gracilis. The 

cause for the “white canopy disease” or “colourless disease” described from Gracilaria 

tenuistipitata cultivated in Vietnam is not known (Phap & Thuan 2002) although it is 

probably similar to “ice-ice disease” in farmed Eucheuma/Kappaphycus species. 

Uyenco et al. (1977) isolated strains of Pseudomonas, Flavobacterium, and Actinobacterium 

associated with "ice-ice disease" in diseased Eucheuma striatum. The symptoms of this 

disease include the presence of a white powdery growth on the thallus which causes loss of 

pigments, and the gradual consumption and subsequent fragmentation of the host. Largo et al. 

(1995a), found that pathogenic bacteria identified as Vibrio sp. and Cytophaga sp. promoted 

ice-ice disease in stressed host branches in the carrageenan-producing red algae Kappaphycus 

alvarezii and Eucheuma denticulatum. Largo et al. (1999) examined the time-dependent 

attachment mechanism of bacterial pathogens during ice-ice infection in Kappaphycus 

alvarezii. 

Ghirardelli (1998) reported on small sheathed Cyanophyta that occur in the cell walls of live 

and dead crustose rhodophytes, collected in the lower intertidal zone in the Gulf of Trieste 

(Northern Adriatic Sea, Italy). Pectonema terebrans is a cyanobacterium that grows in the 

calcified cell walls of coralline algae in Italy, such as Hydrolithon sp., Lithophyllum sp., 

Sporolithon sp. and Titanoderma sp., and it leaves characteristic holes behind and thus can be 

identified even in ancient host material (Ghirardelli 1998). The endophytic cyanobacterium 

Pleurocapsa sp. is associated with galls and the “deformative disease” in Chilean Mazzaella 

laminarioides (Correa et al. 1993, 1997, 2000; Sanchez et al. 1996; Buschmann et al. 1997; 

Faugeron et al. 2000). Pleurocapsa triggers the development of tumours that can result in 

major changes in frond morphology and texture and negatively affect the number of spores, 

settlement rates, germination success and offspring survival (Correa et al. 2000). 

An unspecified bacterium is the cause of “Coralline Lethal Orange Disease” (CLOD) in the 

crustose coralline alga Hydrolithon onkodes from central west Pacific. CLOD is characterised 


y conspicuous bright orange dots associated with tissue necroses that develop into rings 

moving over the host thallus as a front and leaving completely dead white carbonate skeletons 

behind (Littler & Littler 1995; Morcom & Woelkerling 2000). 

Animals 

Amoeba perforate the cell walls of farmed Gracilaria sp., e.g. G. chilensis from Chile, and 

digest the protoplast. Macroscopically the disease manifests as whitish spots spreading rapidly 

throughout the host thallus, similar to “ice-ice disease” (Correa & Flores 1995; Largo et al. 

1995a; Buschmann et al. 2001). 

Larvae and adults of the harpacticoid copepoda Diathrodes cystoceus and D. feldmanni live in 

burrows inside the tissue of red algae and feed on their hosts. Another species, Thalestris 

rhodymeniae, burrows in thalli of Palmaria palmata. The presence of copepoda in red algae is 

associated with galls or pinholes (Barton 1892; Apt 1988b; Park et al. 1990; Shimono et al. 

2004). 

Nematodes are associated with gall formation in Chondrus crispus and Furcellaria 

lumbricalis, however causality has not been demonstrated for these symbioses (Barton 1901; 

Apt 1988b). 

Fungi 

The most intensively studied fungi in red algae are oomycetes of the genus Pythium, 

particularly P. marinum and P. porphyrae, the latter a pathogen causing “red rot” in Porphyra 

species, one of the serious epidemics in laver cultures (e.g. Arasaki 1947; Fuller et al. 1966; 

Sasaki & Sato 1969; Kazama & Fuller 1970; Sasaki & Sakurai 1972; Sakurai et al. 1974; 

Fujita & Zenitani 1976, 1977; Takahashi et al. 1977; Aleem 1980; Tsukidate 1983; Kerwin et 

al. 1992; Amano et al. 1995, 1996; Uppalapati & Fujita 2000a, b, 2001; Uppalapati et al. 

2001; Park et al. 2001, 2007; Shin 2003a, b; Ding & Ma 2005). The majority of this research 

has been carried out in Japan and Korea although a number of studies have also been 

conducted in the eastern Pacific in Washington, USA. 

Diseases of the economically important Porphyra include “chytrid blight” disease (Migita 

1973; Song et al. 1993), the ascomycete Verrucaria consequens causing Kamenoko disease in 

conchocelis cultivation (Migita 1971), Olpidiopsis (Arasaki 1960; Arasaki et al. 1960) and 

also simultaneous infection by red rot and chytrid disease reported by Ding & Ma (2005). 

One basidiomycete pathogen has been reported for red algae (Porter & Farnham 1986b; 

Stanley 1992; Binder et al. 2006). Mycaureola dilseae is a pathogen of the subtidal 

rhodophyte Dilsea carnosa in the Atlantic north-east. This agent causes necrotic lesions, 

which degrade and leave holes in the host frond while the fruiting bodies of the agent develop 

on the margins of the holes. 

In the Ascomycetes species in the genera Chaudefaudia, Hispidicarpomyces, Spathulospora 

have been described from a range of hosts (e.g. Cribb & Herbert 1954; Feldmann 1957; Cribb 

& Cribb 1960; Kohlmeyer 1963b, 1973a, b, c; Sanson et al. 1990; Nakagiri 1993; Nakagiri & 

Ito 1997). Lautitia danica, a pathogen of Chondrus crispus, is found in the reproductive tissue 

of the host, infecting both cystocarpic and tetrasporangial regions (Wilson & Knoyle 1961; 

Schatz 1984b; Stanley 1992). 

Two ascomycetes have been reported to affect commercially important carrageenophytes. 

Dewey et al. (1983) reported on Microascus brevicaulis affecting Eucheuma in the 

Philippines, and Dewey et al. (1984) recorded Penicillium waksmanii isolated from 

Kappaphycus in Micronesia. 


Taxa belonging to the Bigyra, including species of Olpidiopsis, Eurychasmidium, Petersenia, 

Pontisma have been described from a number of hosts (e.g. Sparrow 1934, 1936; Aleem 

1950b, c, 1952b; Feldmann & Feldmann 1967; Whittick & South 1972; van der Meer & 

Pueschel 1985; Molina 1986; West et al. 2006). 

The geographic coverage of studies on marine fungi in red algae is very incomplete and 

currently reflects the regions where the key workers have been based. For example, the 

studies reporting on Chytridium and Rhizophidium species are Sparrow (1936) in the northwest 

Atlantic, Aleem (1952b) in Sweden and Raghukumar (1987a, b) in India. Similarly, the 

only papers focusing on thraustochytrids are Quick (1974) in Florida, USA and Raghukumar 

(1986b, 1987a, b) and Raghukumar et al. (1992) in India. 

Other algae 

Parasitic red algae 

More research has been published on red algal parasites than on any other area of algal 

diseases, pathogens and parasites considered in this study. Red algal parasites are quite 

common, making up to 15% of all named red algal genera, although this figure needs revising 

as the last comprehensive review of red algal parasites was by Goff (1982). Red algal 

parasitism is the most specific symbiosis between two red algae: parasitic red algae are 

symbionts that have a reduced size and are either completely colourless or have reduced 

pigmentation and must rely on their nutrition from their host. In the past two decades a lot has 

been learned about the origin of red algal parasites (Goff et al. 1996, 1997, Zuccarello et al. 

2004), their development (Goff & Coleman 1984, 1985; Goff & Zuccarello 1994; Zuccarello 

& West 1994a), and host specificity (Nonomura & West 1981b; Goff & Zuccarello 1994; 

Goff et al. 1997, Zuccarello & West 1994b, c). However only a very small percentage of 

these parasitic red algae have been studied in any detail beyond their first description. 

The host specificity and evolutionary studies are especially revealing in the context of algal 

pathogenicity and the effects of new interactions. Evolutionary studies have revealed that 

many red algal parasites are derived directly from their hosts (Goff et al. 1996, 1997). This 

was able to be understood once the development of parasites on their hosts was elucidated 

(Goff & Coleman 1984, 1985; Goff & Zuccarello 1994), showing that their unusual 

development was very similar to post-fertilisation processes in red algae. Early in their 

development, either upon spore germination or soon after, red algal parasites transfer into a 

host cell the cytoplasm of an entire cell, or the complete contents of a spore. This 

“transforms” the host cell as it now develops as a parasite, presumably under the control of 

the transferred parasite nucleus, producing more parasite nuclei and dividing to form new 

parasite cells. Finally reproductive structures are formed which eventually will lead to new 

infections. These developmental processes are similar to the nuclear transfer and subsequent 

events that occur during post-fertilisation development in most red algae, and thus red algal 

parasitism has been hypothesised to have been derived from these post-fertilisation processes 

(Goff et al. 1996, 1997). 

Nuclear transfer places constraints on the host range of red algal parasites. The majority of red 

algal parasites appear to be host specific, although this could be an artifact of naming new 

parasite species when parasites are found on new hosts. However, when host range has been 

tested in culture it has been shown to be quite limited (e.g. Goff & Zuccarello 1994). 

Occasionally parasites can grow on closely related host species (Nonomura & West 1981b; 

Zuccarello & West 1994b, c), but often this alternate host development is reduced and 

reproductive structures are not produced. Thus evidence to date supports a high level of host 

specificity. However, on an evolutionary time scale this is shown not to be so, as “host 

jumps” have been discovered using molecular markers. Parasites of the Gigartinales family 


Choreocolacaceae have been shown to infect several species or genera of hosts, although their 

original host was not determined. For example, the parasite Holmsella pachyderma infects 

species of two genera, Gracilaria and Gracilariopsis. The parasite Harveyella mirabilis 

infects several species of the family Rhodomelaceae plus one member of the family 

Delesseriaceae. Other studies have shown that while “host jumps” have been accomplished, 

the parasite is still found on its original host (Goff et al. 1996, 1997). So although not 

confirmed to date in culture studies, parasites appear to be able to jump hosts. This means that 

if introduced to new locations, and given there are appropriate host taxa in the new location, 

parasites may be able to infect new hosts in these environments. 

The question of the detrimental effects and nutritional requirements of red algal parasites on 

their hosts has been barely studied. The few studies that have been conducted show that 

although fixed carbon is translocated into the photosynthesis-lacking parasite, this is often a 

small fraction of the total fixed carbon (Goff 1979; Kremer 1983). No studies have looked at 

the effect of parasitism on host reproductive success, host recruitment, or the host ability to 

withstand perturbations. 

Endophytic red algae 

Endophytic red algae other than parasites are pigmented and do not form cellular connections 

to host cells. Usually they can be cultivated outside their hosts. Photosynthetic red algae 

found within the tissues of other algae are common. Species of Auduoinella sp. (also under 

the name of Acrochaetium sp., Colaconema sp., Rhodochorton sp.) are often found 

intercellularly within thallose red algae (e.g. West 1979). There have been few experimental 

studies of the specificity of these endophytes, although host range is considered to be fairly 

broad. It is possible that these endophytes could infect the tissue of new organisms given the 

opportunity. Acrochaetium yamadae grows in the tissue of Izziella orientalis from Taiwan 

(Kylin 1956) and of Liagora canariensis from the Canary Islands (Afonso-Carrillo et al. 

2003). The former Acrochaetium species, Colaconema asparagopsis and C. bonnemaisoniae, 

are found in British Bonnemaisonia hamifera and Asparagopsis sp., while the related species 

C. endophyticum grows in Heterosiphonia sp., (Kylin 1956; White & Boney 1969). 

Colaconema ophioglossum is an endophyte of Dudresnaya crassa from both sides of the 

central Atlantic (Afonso-Carrillo et al. 2003). 

Some semi-endophytic rhodophytes are found among non-geniculate coralline algae from the 

Central Pacific. The thallus of Lithophyllum cuneatum from Fiji is wedged into the thalli of its 

hosts, Neogoniolithon sp. and Hydrolithon onkodes. Endophyte and hosts do not form cellular 

connections; however the growth of the host may be disturbed by the presence of the 

endophyte (Keats 1995; Chamberlain 1999; Morcom & Woelkerling 2000). Similarly, 

Amphiroa species (such as A. kuetzingiana) are embedded into their hosts Hydrolithon 

onkodes, Neogoniolithon brassica-florida and Mesophyllum expansum, but apparently do not 

parasitise them (Chamberlain 1999). 

In contrast, the epiphyte Titanoderma corallinae has a detrimental effect on its basiphytes 

Corallina elongata and C. officinalis from France; contact with its spores leads to bleaching 

of the host tissue, from which the host may not recover (Cabioch 1979; Chamberlain 1999). 

Red algal epiphytes 

Most fouling red algae will grow on any surfaces (e.g. Stylonema, Erythrotrichia). These are 

often small algae, with asexual means of reproduction that can quickly colonise new surfaces. 

Most of these algae grow on the surface of the host without causing any structural damage to 

the host, though shading of the host could lead to slowed host growth. Some other red algae 

are generalist epiphytes, or at least much more common on algal surfaces (e.g. Microcladia 

coulteri - Gonzalez & Goff 1989). These algae have different ways of interacting with the 


host (Leonardi et al. 2006) with some of these interactions leading to damage and other 

detrimental effects to the host (e.g. tissue loss and secondary infections), as these epiphytes 

can penetrate host tissue to varying degrees. These algal-epiphyte interactions can especially 

be detrimental to cultivated algae (e.g. Gracilaria chilensisi) where hosts are in high 

concentration and economic loss is possible (Leonardi et al. 2006; Vairappan 2006). A 

number of filamentous red algae grow as epiphytes on Kappaphycus alvarezii, a 

carrageenophyte commercially cultivated throughout Asia. The predominant epiphyte species 

is Neosiphonia savatieri, but other species also occur such as Acanthophora sp., Ceramium 

sp., Centroceras sp. and N. apiculata. The epiphytes are anchored on their host by penetrating 

rhizoids. Their presence weakens the host and increases its susceptibility to bacteria 

(Vairappan 2006). 

Ostiophyllum sonderopeltae is an obligate epiphyte of Sonderopelta coriacea in Australia 

(Kraft 2003). Lembergia allanii is known only on Vidalia colensoi from New Zealand, 

whereas Dasyptilon pellucidum is predominantly found on Euptilota formossissima but may 

also be found growing on Hymenocladia and Cenacrum (Adams 1994). As the specificity of 

the epiphyte habit has not been determined for a number of species, and where these epiphyte 

taxa appear to cause no disease symptoms, epiphyte taxa have not been included in the 

database. 

Endophytic green algae in red algae 

Red algae are the hosts for a number endophytic green algae. The economically important 

carrageenophyte Chondrus crispus is infected by Acrochaete heteroclada and A. operculata 

on both sides of the northern Atlantic. Acrochaete heteroclada disrupts the tissue of the host 

cortex and has an overall negative effect on the host performance, slowing the growth and 

decreasing the capacity for regeneration, leading to lower yields of carrageenan. A. operculata 

likewise penetrates the cortex of its host. However, while gametophytes are not invaded 

beyond the outer cortex, sporophytes become completely endophytised, resulting in severe 

damage of the host tissue, secondary bacterial infections, and eventually disintegration and 

death of the host thallus (Correa & McLachlan 1991, 1992, 1994, Bouarab et al. 1999, 2001b; 

Potin et al. 1999, 2002; Bown et al. 2003; Weinberger et al. 2005). 

Achrochaete heteroclada is also found in Ahnfeltiopsis furcellata, A. linearis, Chondrus 

canaliculatus, Gracilaria chilensis and G. mammilaris (Correa & McLachlan 1991), while 

another Acrochaete species, A. leptochaete, infects Polysiphonia sp. and Champia sp. 

(O’Kelly et al. 2004). In Britain, Chondrus crispus hosts another green endophyte, Entocladia 

viridis (Bown et al. 2003), a species also found in Phycodrys rubens along the North Atlantic 

coasts of the USA (O’Kelly et al. 2004). 

The endophyte Endophyton ramosum causes “green patch disease” in Chilean Mazzaella 

laminarioides. This disease is characterised by fronds which lose their red pigmentation and 

turn green. The host tissue starts decaying, opening the way for secondary bacterial invasions. 

Lesions on the stipes lead to their breaking in heavy wave action (Correa et al. 1994, 1997; 

Sanchez et al. 1996; Buschmann et al. 1997; Faugeron et al. 2000). Eucheuma ramosum also 

inhabits the related host species Mazzaella oregona in the Northeast Pacific (O’Kelly et al. 

2004). 

Endophytic unicellular sporophytes of Acrosiphonia species, originally described as 

Chlorochytrium inclusum and Codiolum petrocelidis, were observed in a number of foliose 

and crustose red algae, respectively, from British Columbia: Callophyllis sp., Chondrus 

crispus, Constantinea subulifera, Dilsea californica, D. integra, Farlowia sp., Haemescharia 

hennedyi, Halymenia sp., Hildenbrandia occidentalis, Kallymenia sp., Mastocarpus 

papillatus, Mazzaella sanguinea, M. splendens, Palmaria mollis, Porphyra sp., Schizymenia 


pacifica, Sparlingia pertusa, and Weeksia sp. (Sussmann et al. 1999, 2005, Sussmann & 

DeWreede 2001, 2002, 2005). Acrosiphonia sporophytes also occur in Palmaria mollis and 

Polyides rotundus from the Northeast Atlantic (Sussmann & DeWreede 2002). 

Spongomorpha aeruginosa occurs in Haemescharia hennedyi from Germany, and the related 

species S. mertensii in Mastocarpus papillatus from Canada (Sussmann & DeWreede 2001). 

Two endophytic green algae observed in Curdiea racovitzae and Iridea cordata from the 

Antarctic Peninsula were not further identified (Peters 2003). 

Endophytic brown algae in red algae 

Brown algae living as endophytes in red algae are from three orders of the Phaeophyceae: 

Ectocarpales, Laminariales and Desmarestiales. Setchell & Gardner (1922) described a 

number of new species of Streblonema, both epiphytic and endophytic taxa, including the 

endophytes S. corymbiferum (in Cumagloia andersonii) and S. investiens (in Helminthocladia 

calvadosii). Microspongium tenuissimum occurs in Aeodes orbitosa from South Africa, 

Grateloupia doryphora from Canary Islands, and Grateloupia intestinalis from Chile (Peters 

2003). A second Microspongium species, M. radians, which has been described from Chilean 

Grateloupia doryphora and also grows in Mazzaella laminarioides from South Africa 

(Burkhardt & Peters 1998; Peters 2003) is considered synonymous to M. tenuissimum 

(Heesch 2005). Another endophyte, genetically identified as Microspongium sp., was isolated 

from Polysiphonia elongata growing in the Western Baltic Sea (Burkhardt & Peters 1998). 

This species may be synonymous with Mikrosyphar polysiphoniae described from Baltic 

Polysiphonia stricta. Pedersen (1976) reported Mikrosyphar polysiphoniae in Polysiphonia 

arctica in collections from Greenland. Other Mikrosyphar species, such as M. porphyrae, an 

endophyte of Porphyra sp. in the Baltic Sea, may likewise belong to the genus 

Microspongium (Heesch 2005). 

Kelp gametophytes have recently been discovered living endophytically in filamentous and 

foliose red algae. Most of the hosts belong to the order Ceramiales, such as Antithamnion 

densum, Callithamnion acutum, C. biseriatum, Ceramium gardneri, Delesseria decipiens, 

Griffithsia pacifica, Herposiphonia plumula, Irtugovia pacifica, Membranoptera platyphylla, 

Pleonosporium vancouverianum, Polyneura latissima, Polysiphonia paniculata, 

Pterosiphonia dendroidea, Pterosiphonia sp., Pterothamnion pectinatum and Scagelia 

pylaisei. Kelp gametophytes are furthermore hosted by Fryeella gardneri (Rhodymeniales) 

and Euthora cristata, Orculifilum denticulatum (Gigartinales). In earlier studies, the species 

of Laminariales involved were not identified further (Garbary et al. 1999a, b, Garbary & Kim 

2000), although more recently Sasaki et al. (2003) were able to identify Agarum clathratum in 

Orculifilum denticulatum and Hubbard et al. (2004) identified gametophytes of Alaria 

esculenta and Nereocystis luetkeana growing in a number of hosts. Gametophytes of 

Desmarestia antarctica grow in Antarctic Curdiea racovitzae (Moe & Silva 1989; Peters 

2003). 

Although some taxa are predominantly epiphytic, they may also affect the host through some 

endophytic development, as found in the epiphyte Elachista antarctica which is anchored 

within its Antarctic host Palmaria decipiens by endophytic filaments (Peters 2003). 

Endophytic diatoms 

Diatoms may either live as endo- or epiphytes in/on red algae. Diatoms such as Achnanthes 

longipes, Melosira nummoloides, Synedra gracilis and Ligmophora sp. heavily epiphytise 

Porphyra species, e.g. P. yezoense, in Japan and South Korea. The epiphyte load inhibits 

normal growth of the basiphyte, leading to a condition called “diatom felt disease” (Tsukidate 

1983, 1991; Fujita 1990; Song et al. 1993). Examples of endophytic diatoms are Gyrosigma 

coelophilum, which has been observed in Coelarthrum opuntia in Japan (Okamoto et al. 


2003), and Pseudogomphonema sp., an endophyte of Pachymenia sp. from the Antarctic 

Peninsula (Peters 2003). 


Red algal parasites are poorly documented in New Zealand although species belonging to the 

following genera are known, either reported in publications (e.g. Adams 1994) or recorded in 

herbaria: Callocolax, ?Ceratocolax sp., Champiocolax, Choreonema, Colacodasya, 

Colacopsis, Gloiocolax, Janczewskia, Levringiella, Microcolax, Plocamiocolax, 

Pterocladiophila, Rhodymeniocolax, Sporoglossum, Tylocolax. A great deal more work is 

required on the red algal parasites in the New Zealand region. 

Three species of the ascomycete Spathulospora have been described from New Zealand 

collections - Spathulospora lanata in Camontagnea oxyclada, S. adelpha and S. calva on 

Ballia callitricha (Kohlmeyer 1973a). Kohlmeyer & Demoulin (1981) described two 

ascomycete fungi that are found in association with the New Zealand endemic genus 

Apophlaea - Mycophycias apophlaeae and Polystigma apophlaeae Kohlm. 

The endophytic brown alga Microspongium tenuissimum (incl. M. radians) occurs in three red 

algae from New Zealand: Pachymenia lusoria, Grateloupia intestinalis and in a so far 

undescribed species of the family Kallymeniaceae (Heesch 2005). Another species, 

Mikrosyphar pachymeniae was described from northern populations of P. lusoria, but may be 

synonymous with Microspongium tenuissimum (Heesch 2005). 

3.6. GREEN ALGAE 


Viruses 

No virus infections have been reported for marine green algae. 

Bacteria 

No bacterial diseases have been reported for marine green algae. 

Animals 

Two unidentified protozoa, a ciliate and a flagellate, live endophytically in Codium bursa 

(Armstrong et al. 2000), while an amoeba has been reported from Blidingia chadefaudii 

(Feldmann & Feldmann 1967). In the Florida Keys an amphipod (Erichthonius brasiliensis) 

affects the growth of the green alga Halimeda tuna by rolling its terminal segments (Sotka et 

al. 1999). 

Fungi 

Species of the genus Cladophora host a number of pathogenic fungi, such as Labyrinthula 

spp. (e.g. L. coenocystis), Coenomyces sp., Achlyogeton salinus, Entophlyctis maxima, 

Olpidium rostiferum and Sirolpidium bryopsidis (Dangeard 1931a; Sparrow 1936; 

Raghukumar 1986a, 1987b; Rheinheimer 1992; Hyde et al. 1998; Raghukumar 2002). In 

India, a thraustochytrid fungus infects Cladophora liebetruthii (Raghukumar 1986a). 

Blodgettiomyces bornetii is a fungus occuring in Cladophora catenata and other Cladophora 

species, as well as in Siphonocladus rigidus (Kohlmeyer & Kohlmeyer 1972; Porter & 

Farham 1986a). Blodgettia sp. occurs in Cladophora dalmatica (Saccardo 1882a). 

Labyrinthula sp. also occurs in Chaetomorpha and Rhizoclonium species. The latter moreover 


hosts Coenomyces sp. and Olpidium rostiferum (Raghukumar 1986b, 1987a, 2002; Hyde et al. 

1998). 

Pontisma lagenidioides causes the “browning disease” in Chaetomorpha antennina 

(Raghukumar 1987a; Raghukumar & Chandramohan 1988). Thraustochytrium proliferum, 

Rhizophydium littoreum, R. globosum and Phlyctochytrium sp. are pathogens of Bryopsis 

plumosa (Sparrow 1936; Kazama 1972; Amon 1984; Hyde et al. 1998) and the former also 

infects Codium sp. (Amon 1984). Olpidiopsis andreii infects filamentous green algae, e.g. 

Acrosiphonia sp. and Spongomorpha sp. (Aleem 1952a; Porter & Farnham 1986a; West et al. 

2006). 

The ascomycetes Guignardia alaskana and G. prasiolae have been reported from Prasiola 

borealis and Prasiola tesselata, respectively. The former is also parasitised by Laestadia 

alaskana. Both algae furthermore host Turgidosculum complicatulum, while the related 

species T. ulvae occurs in Blidingia minima and B. minima var. vexata (Saccardo 1882b; Reed 

1902; Kohlmeyer & Kohlmeyer 1972, 1973; Kohlmeyer 1979) and Ulva californica (Reed 

1902). In France, the ascomycete Chadefaudia corallinarum infests Flabellia petiolata and 

Halimeda tuna (Kohlmeyer 1963b). In Russia’s Sea of Japan, Ulva fenestrata is endophytised 

by Ulocladium littoreum (Pivkin & Zvereva 2000). 

Ostreobium queketti, an endolithic alga growing in corals from French-Polynesia, is 

parasitised by an aspergillus-like fungus, causing a black banding pattern on the coral host 

(Priess et al. 2000). In Sweden, an unspecified fungus has been reported to parasitise 

Elasticha fucicola (Aleem 1952a). 

A unidentified heterokont biflagellate parasite lives inside Codium fragile from the North 

American Atlantic coast, consuming the plastids of its host (Lee & Kugrens 2003). 

Other algae 

Members of the genus Achrochaete occur as endophytes in Ulva rigida and Codium fragile. 

Another endophyte, Entocladia viridis has been found in Bryopsis duplex and Chaetomorpha 

linum from Italy and Denmark, respectively (O’Kelly 1981; Nielsen 1979; del Campo et al. 

1998). Another green seaweed, Chlorochytrium dermatocolax, has been reported as a parasite 

of a green host, Bryopsis plumosa (Sparrow 1936). 

There is a single record of a red seaweed (Schmitziella endophloea) as an endophyte in 

Cladophora pellucida (Kylin 1956). 


The labyrinthulid Thraustochytrium proliferum has been isolated from Bryopsis plumosa and 

Cladophora sp. from Dunedin (Karling 1968). 

3.7. XANTHOPHYCEAE 


Viruses 

No virus infections have been reported for the Xanthophyceae. 

Bacteria 

No bacterial infections have been reported for the Xanthophyceae. 


Animals 

A rotifer was reported to cause galls in Vaucheria sp. (Apt 1988b). 

Fungi 

No fungal infections have been reported for the Xanthophyceae. 

Other algae 

No algal infections have been reported for the Xanthophyceae. 


No infections have been reported for the Xanthophyceae in New Zealand. 


4. Discussion 

I: ASSESSMENT OF INFORMATION AVAILABLE ON SEAWEED DISEASES 

WORLDWIDE AND IN NEW ZEALAND 

A number of general reviews have dealt with pathogens of marine algae, e.g. Evans et al. 

(1978) and Goff (1982) focusing on parasitic red algae; Andrews (1979a, b) on the pathology 

of seaweeds; Apt (1988b) on galls and tumour-like growths; Correa (1997) examining the 

current knowledge and approaches to infectious diseases of marine algae; Bouarab et al. 

(2001a) examining the ecological and biochemical aspects of algal infectious diseases. Fujita 

(1990) authored a review specifically on the diseases of cultivated Porphyra in Japan. 

Biosecurity NZ requested that data in this project should be compiled for each pathogen 

including: 

• agent stability and inactivation data; 

• epidemiological features: 

− geographical range and features of distribution (international spread); 

− host range (including prevalence and incidence, resistant strains/species, life stage 

susceptibility and course of infection, habitat and seasonality); 

− morbidity/mortality rates; 

− transmission (including route and infectious dose). 

• host impact: 

− tissue tropism (site of infection); 

− brief description of major pathological and biological effects. 

• diagnostics and disease control: 

− key diagnostic features; 

− overview of diagnostic methods, including sensitivity and specificity; 

− disease management activities worldwide; 

− able to be eradicated? 

The majority of papers did not include data in these areas. Generally, the information on 

diseases of seaweeds is very patchy and the emphasis of published work lies in two main 

areas: 

• diseases occurring in monocultures of farmed species, mainly in East and Southeast Asia 

(particularly affecting the key economic genera Porphyra. Laminaria, Undaria, 

Gracilaria, Eucheuma and Kappaphycus); 

• observations of certain groups of pathogens in particular geographic regions as a 

consequence of the research interests of a particular team or research group, leading to 

“pockets of information”. 

The amount of information contained in the references we investigated varied greatly between 

articles, ranging from reports of the occurrence of pathogens to multi-paper treatments of 

certain diseases. The latter are especially numerous for farmed macroalgae e.g. Pythium 

porphyrae, the agent causing the red rot disease in Porphyra species (Porphyra cultivation is 

a billion dollar industry in Asian countries). Other agents, in contrast, have only been 

observed once and often only incidentally in the course of other research. 

Problems with the correct identifications and classification of pathogens may lead to different 

names for the same agent or the same name for different agents. For example, small algae 

such as endophytes may have a reduced morphology, leaving only few characters for 


identification. Often, they are not fertile when observed, making the correct identification 

difficult. 

Host symptoms are not always a good character for identifying pathogens/ parasites either, as 

these may depend on the susceptibility of a host species to a specific agent. Susceptibility can 

vary between generations of the same host species e.g. the sporophyte of the red alga 

Chondrus crispus is susceptible to infections of the green endophyte Acrochaete operculata, 

while the host gametophyte shows some resistance (Correa & McLachlan 1991). Life stages 

of pathogens may also display different morphologies e.g. nauplii of harpacticoid copepoda 

burrowing in kelp stipes may have to be reared to adult stages in order to correctly identify 

them by morphology. Most references describe a disease by the symptoms expressed in the 

host, but fall short of demonstrating causality, meaning for example, more obvious secondary 

invaders could be mistakenly attributed as the primary cause of a disease. There is almost no 

work that has examined more complex pathogen/host systems. 

Information on diseases, pathogens and parasites occurring in New Zealand is scant. 

Generally the publications available reflect the activities of a few overseas workers who have 

visited or received specimens and have published on particular agents (e.g. Kohlmeyer). 

There have been no focused studies incorporating field and laboratory investigations other 

than the work of Heesch (2005) on endophytes of brown algae in New Zealand, completed as 

research for a Ph.D. at the University of Otago. 

II: ASSESSMENT OF THREATS BY PATHOGENS OF UNDARIA TO NEW ZEALAND 

NATIVE MARINE FLORA 

The only disease reported in Undaria from its introduced range is the infection of thalli with 

the pigmented endophytic brown alga Laminariocolax aecidioides, both in Spain (Veiga et al. 

1997) and in Argentina (Gauna et al. personal communication). It is not clear whether this 

endophyte originates from Japanese populations introduced with the host or from European or 

Argentinian populations respectively. Laminariocolax aecidioides is known from other, 

native European kelps such as Laminaria hyperborea in the German Bight and Norway, and 

Saccharina latissima in the Western Baltic Sea (Lein et al. 1991; Ellerstdottir & Peters 1995, 

1997; Peters & Schaffelke 1996), but it has not been reported from southern Europe. It also 

occurs in the native range of U. pinnatifida, in Japan (Yoshida & Akiyama 1978). Genetic 

studies may determine the origin of the Spanish and Argentinean populations and thus shed 

some light on whether endophytes were or can be transmitted with host sporophytes (or other 

disease agents). 

In the Western Baltic, thalli of Saccharina latissima infected with Laminariocolax aecidioides 

show more severe symptoms in shallow water, due to the endophyte growth being accelerated 

in better light conditions. Increased severity of infection symptoms prevent host thalli 

surviving in water depths of 2 m, in contrast to deeper water where growth of the endophyte is 

light limited (Schaffelke et al. 1996). In New Zealand, Laminariocolax macrocystis, a closely 

related species in this genus, infects native kelps, such as Macrocystis pyrifera and Ecklonia 

radiata, and in severe cases this leads to crippled thalli (M. pyrifera) and/or stunted growth 

(E. radiata) (Heesch 2005). It is not known if this endophyte species has an influence on the 

depth distribution of its hosts. Further, it is not known if L. aecidioides would be able to infect 

New Zealand native kelps, and if so, what the consequences would be for native kelp 

populations. 

Reports in the international literature have highlighted the occurrence of kelp gametophytes as 

endophytes in a range of hosts (e.g. Garbary et al. 1999a, b; Garbary & Kim 2000; Sasaki et 


al. 2003; Hubbard et al. 2004). One of the potential threats of Undaria pinnatifida to New 

Zealand native kelps may consist of competition with other kelp gametophytes as endophytes. 

III: FUTURE STRATEGY FOR SCREENING POPULATIONS AND INCREASING 

KNOWLEDGE OF RISK POSED BY DISEASES/PARASITES/PATHOGENS TO NEW 

ZEALAND MACROALGAE AND COASTAL COMMUNITIES 

None of the known pathogens of Undaria have so far been observed in/on U. pinnatifida in 

New Zealand, however, populations of U. pinnatifida around New Zealand have not been 

screened for the presence of diseases, pathogens and parasites. Given that there is evidence 

that New Zealand has received at least 10 separate introduction events of Undaria pinnatifida 

(Uwai et al. 2006), it would be important to construct a sampling regime that reflected this 

known genetic diversity within New Zealand populations of Undaria. 

Correa (1997) recommends an operational approach to the study of infectious diseases in 

seaweeds: 

1. field and laboratory observations aiming to individualize a potential pathogen and to 

describe the lesions associated with the presence of that organism, 

2. laboratory experiments and observations to establish causality i.e. applying Koch’s 

postulates (Andrews & Goff 1984), as well as manipulative experiments to understand 

aspects of the host-pathogen relationship and thus develop methods to manage the 

disease, e.g. in marine cultures 

3. epidemiology to “evaluate... the population segment ...affected..., the severity of the 

disease and the occurrence of seasonal and spatial patterns of disease expression”, which 

includes the study of the reproduction, mortality and physiological performance of the 

host population and individuals. 

From the research conducted by Heesch (2005) it is clear that it is necessary to identify host 

populations and look for disease symptoms both intra- and inter-annually, with seasonal 

sampling occurring ca. quarterly. Given the range of environments, water temperatures, and 

photoperiods experienced through the New Zealand region, the sampling would need to be 

stratified and targeted on priority taxa. Depending on the biology of the target taxa the 

sampling regime would need to incorporate considerations of the species life history (i.e. 

whether the species has isomorphic or heteromorphic alternation of generations or direct 

development; if life history phases have differing cell wall chemistry as found for example in 

isomorphic phases of members of the Gigartinales), ecology and distribution (light, depth, 

exposure/shelter, substrate). Causality between disease and symptoms requires both field and 

detailed laboratory investigations. 

A number of authors point to the importance of considering diseases, pathogens and parasites 

in the wider context, testing hypotheses about the roles they may play in shaping population 

and community structure (Correa 1997; Prenter et al. 2004; Tompkins & Poulin 2006). 


5. Conclusions 

Seaweeds that are diseased are under-collected in New Zealand and, as a consequence, the 

status of knowledge about biotic diseases, pathogens and parasites is deficient: it is not 

possible to evaluate risk posed by introduced diseases, pathogens and parasites on the basis of 

current understanding of the native biota. 

Whilst experts in the field of algal diseases such as Correa (1997) stress the need for studies 

on the mechanisms of infection and the spread of the pathogens within and among host 

individuals, as well as on the genetics of the host-pathogen interaction, the basic underpinning 

surveys and research are required in New Zealand to document the biodiversity and 

distribution of diseases, pathogens and parasites within macroalgae. 

6. Acknowledgements 

The following people are thanked for their contributions to this study: Megan Gee for 

literature searching and acquisition, Helen Sui for developing the database structure, Joe 

Zuccarello for assistance with reviewing literature on red algal parasites, Joe Buchanan and 

Peter Martin for assistance with literature reviews, Tracy Farr for assistance with maps, Hoe 

Chang, Janet Grieve and Roberta D’Archino for assistance with translations. 


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Appendix 4. Map of known pathogens affecting Undaria pinnatifida in its native range of Japan, China and Korea. Locations are approximate 

and inferred from placenames available in the literature. 

120˚E 

150˚E 

30˚N 

0 500 

km 

☼ 

Amenophia (copepod) 

Parathalestris (copepod) 

Thalestris (copepod) 

Scutellidium (copepod) 

Ceinina (amphipod) 

SOUTH 

KOREA 

Honshu 

JAPAN 

☼ Olpidiopsis (fungi) 

Unspecified bacteria 

Vibrio (bacteria) 

40˚N 

Dalian 

CHINA 

Moraxella (bacteria) 

Flavobacterium (bacteria) 

Pseudomonas (bacteria) 

Hokkaido 

Halomonas (bacteria) 

Laminariocolax (algae) 

Microspongium (algae)

Appendix 3. Map showing FAO geographic regions, distribution of Undaria in these regions and pathogens ( Laminariocolax (algae); 

Microspongium (algae)) affecting Undaria in its introduced range. Native range (dark grey shading); introduced range (mid grey shading).

APPENDIX 1: 

Hierarchical Classification (based on Cavalier –Smith 1998) and Species 2000 

(D.Gordon, pers. comm. NIWA) 

EMPIRE OR SUPERKINGDOM 1. PROKARYOTA 

Kingdom 1. Bacteria 

Subkingdom 1. Negibacteria 

Infrakingdom 1. Eobacteria 

Phylum 1. Eobacteria 

Class 1. Chlorobacteria [e.g. Chloroflexus, Heliothrix, Thermomicrobium] 

Class 2. Hadobacteria [e.g. Deinococcus, Thermus] 

Infrakingdom 2. Glycobacteria 

Phylum 1. Cyanobacteria 

Subphylum 1. Gloeobacteria 

Class 1. Gloeobacteria 

Order 1. Gloeobacterales [e.g. Gloeobacter] 

Subphylum 2. Phycobacteria 

Class 1. Chroobacteria 

Order 1. Chroococcales [e.g. Anabaena, Prochloron] 

Order 2. Pleurocapsales [e.g. Pleurocapsa] 

Order 3. Oscillatoriales [e.g. Oscillatoria] 

Class 2. Hormogoneae 

Order 1. Nostocales [e.g. Nostoc] 

Order 2. Stigonemates [e.g. Stigonema] 

Phylum 2. Spirochaetae 

Class Spirochaetes [e.g. Leptospira, Spirochaeta, Treponema] 

Phylum 3. Sphingobacteria 

Class 1. Flavobacteria [Fibrobacter, Flavobacterium] 

Class 2. Chlorobia [e.g. Cytophaga, Flavobacteria] 

SUPERPHYLUM EXOFLAGELLATA 

Phylum 1. Planctobacteria 

Class 1. Planctomycea [e.g. Pirellula, Planctomyces] 

Class 2. Verrucomicrobeae [e.g. Verrucomicrobium] 

Class 3. Chlamydiae [e.g. Chlamydia] 

Phylum 2. Proteobacteria 

Subphylum 1. Rhodobacteria 

Class 1. Chromatibacteria [e.g. Chromatium, Escherichia, Haemophilus, Methylococcus, Pseudomonas, 

Spirillum, Vibrio] 

Class 2. Alphabacteria [e.g. Agrobacterium, Caulobacter, Hyphomicrobium, Rhizobium, Rhodospirillum, 

Rickettsia] 

Subphylum 2. Thiobacteria 

Class 1. Deltabacteria[e.g. Bdellovibrio, Desulfovibrio, Myxococcus] 

Class 2. Epsilobacteria [e.g. Aquifex, Helicobacter, Hydrogenobacter, Thermotoga] 

Subphylum 3. Geobacteria 

Class 1. Ferrobacteria [e.g. Geobacter, Leptospirillum, Magnetobacterium] 

Class 2. Acidobacteria [e.g. Acidobacterium, Holophaga, Geothrix] 

Subkingdom 2. Unibacteria 

Phylum 1. Posibacteria 

Subphylum 1. Endobacteria 

Class 1. Togobacteria [e.g. Heliobacterium, Selenomonas, Thermotoga] 

Class 2. Teichobacteria [e.g. Bacillus, Clostridium, Staphylococcus, Streptococcus] 

Class 3. Mollicutes [e.g. Mycoplasma] 

Subphylum 2. Actinobacteria 

Class 1. Arthrobacteria [e.g. Arthrobacter, Actinomyces] 

Class 2. Arabobacteria 

Order 1. Actinoplanales [e.g. Actinoplanes] 

Order 2. Mycobacteriales [Mycobacterium] 

Class 3. Streptomycetes [e.g. streptomyces] 

Phylum 2. Archaebacteria 


Subphylum 1. Euryarchaeota 

Superclass 1. Neobacteria 

Class 1. Methanothermea [e.g. Methanococcus] 

Class 2. Archaeoglobea 

Class 3. Halomebacteria [e.g. Halobacterium, Methanospirillum] 

Superclass 2. Eurythermea 

Class 1. Protoarchaea [e.g. Palaeococcus, Protococcus] 

Class 2. Picrophilea [e.g. Ferroplasma, Thermoplasma] 

Subphylum 2. Crenarchaeota 

Class 1. Crenarchaeota [e.g. Sulfolobus, Pyrobaculum] 

EMPIRE OR SUPERKINGDOM 2. EUKARYOTA 

Kingdom 1. Protozoa 

Subkingdom 1. Sarcomastigota 

Phylum 1. Amoebozoa [Rhizopoda] 

Subphylum 1. Protamoebae 

Class 1. Breviatea 

Class 2. Lobosea 

Order 1. Euamoebida [e.g. Amoeba, Rhizamoeba] 

Order 2. Copromyxida [e.g. Copromyxa] 

Order 3. Arcellinida [e.g. Arcella, Difflugia] 

Class 3. Discosea 

Order 1. Glycostylida [e.g. Paramoeba, Vannella] 

Order 2. Himatismenida [e.g. Cochliopodium] 

Order 3. Dermamoebida [e.g. Thecamoeba] 

Class 4. Variosea 

Order 1. Phalansteriida [e.g. Phalansterium] 

Order 2. Centramoebida [e.g. Acanthamoeba] 

Order 3. Varipodida [e.g. Filamoeba, Gephyramoeba] 

Subphylum 2. Conosa 

Infraphylum 1. Archamoebae 

Class 1. Archamoebea 

Order 1. Pelobiontida [e.g. Entamoeba, Pelomyxa] 

Order 2. Mastigamoebida [e.g. Endolimax, Mastigamoeba] 

Infraphylum 2. Mycetozoa 

Class 1. Stelamoebea 

Order 1. Protostelida [e.g. Protostelium, Schizoplasmodium] 

Order 2. Dictyosteliida [e.g. Dictyostelium] 

Class 2. Myxogastrea 

Order 1. Parastelida [e.g. Ceratiomyxa] 

Order 2. Echinosteliida [e.g. Echinostelium] 

Order 3. Liceida [e.g. Listerella] 

Order 4. Trichiida [e.g. Dianema] 

Order 5. Stemonitida [e.g. Stemonitis] 

Order 6. Physarida [e.g. Didymium, Elaeomyxa, Physarum] 

Phylum 2. Choanozoa 

Class 1. Choanoflagellatea 

Order 1. Craspedida [e.g. Codosiga, Monosiga, Salpingoeca] 

Order 2. Acanthoecida [e.g. Acanthoeca, Diaphanoeca] 

Class 2. Corallochytrea 

Order 1. Corallochytrida [e.g. Corallochytrium] 

Class 3. Ichthyosporea 

Order 1. Ichthyosporida [e.g. Dermocystidium, Ichthyophonus] 

Class 4. Cristidiscoidea 

Order 1. Ministeriida [e.g. Ministeria] 

Order 2. Nucleariida [e.g. Fonticula, Nuclearia] 

Subkingdom 2. Biciliata 

Infrakingdom 1. Rhizaria 

Phylum 1. Cercozoa [Zooflagellata] 

Subphylum 1. Filosa 

Superclass 1. Reticulofilosa 

Class 1. Chlorarachnea [e.g. Chlorarachnion] 

Class 2. Proteomyxidea [e.g. Dimorpha, Gymnophrys, Reticulamoeba] 


Superclass 2. Monadofilosa 

Class 1. Sarcomonadea [e.g. Cercomonas, Heteromita, Metopion] 

Class 2. Thecofilosea [e.g. Cryothecomonas, Cryptodifflugia] 

Class 3. Spongomonadea [e.g. Spongomonas] 

Class 4. Imbricatea [e.g. Euglypha, Thaumatomonas] 

Class 5. Phaeodaria [e.g. Collosphaera] 

Subphylum 2. Endomyxa 

Class 1. Phytomyxea 

Order 1. Phagomyxida [e.g. Phagomyxa] 

Order 2. Plasmodiophorida [e.g. Plasmodiophora] 

Class 2. Ascetosporea 

Order 1. Haplosporida [e.g. Bonamia, Haplosporidium, Urosporidium] 

Order 2. Paramyxida [e.g. Paramyxa] 

Order 3. Claustrosporida [e.g. Claustrosporidium] 

Class 3. Gromiidea 

Order 1. Gromiida [e.g. Gromia] 

Phylum 2. Foraminifera 

Class 1. Athalamea [e.g. Reticulomyxa] 

Class 2. Polythalamea [e.g. Allogromia, Globigerina, Textularia] 

Class 3. Xenophyophorea [e.g. Psammina] 

Phylum 3. Radiozoa 

Class 1. Acantharea [e.g. Acanthometra] 

Class 2. Sticholonchea [e.g. Sticholonche] 

Class 3. Polycystinea [e.g. Collozoum] 

Infrakingdom 1. Excavata 

SUPERPHYLUM 1. APUSOZOA 

Phylum Apusozoa 

Class 1. Diphyllatea 

Order 1. Diphylleida [e.g. Collodictyon, Diphylleia] 

Class 2. Thecomonadea 

Order 1. Apusomonadida [e.g. Amastigomonas, Apusomonas] 

Order 2. Ancyromonadida [e.g. Ancyromonas] 

Order 3. Hemimastigida [e.g. Spironema] 

Class 3. Teonemea [e.g. Nephromyces, Telonema] 

SUPERPHYLUM 2. EOZOA 

Phylum 1. Loukozoa 

Class 1. Jakobea 

Order 1. Jakobida [e.g. Histiona, Jakoba, Reclinomonas] 

Class 2. Malawimonadea 

Order 1. Malawimonadida [e.g. Malawimonas] 

Phylum 2. Metamonada 

Subphylum 1. Anaeromonada 

Class 1. Anaeromonadea [e.g. Dinenympha, Personympha, Trimastix] 

Order 1. Trimastigida [e.g. Trimastix] 

Order 2. Oxymonadida [e.g. Dinenympha, Pyrsonympha] 

Subphylum 2. Trichozoa 

Superclass 1. Parabasalia 

Class 1. Trichomonadea 

Order 1. Trichomonadida [e.g. Calonympha, Trichomonas] 

Order 2. Lophomonadida [e.g. Microjoenia, Lophomonas] 

Order 3. Spirotrichonymphida [e.g. Holomastigotoides] 

Class 2. Trichonymphea 

Order 1. Trichonymphida [e.g. Trichonympha] 


Superclass 2. Carpediemonadia 

Class 1. Carpediemonadea 

Order 1. Carpediemonadida [e.g. Carpediemonas] 

Superclass 3. Eopharyngia 

Class 1. Trepomonadea 

Subclass 1. Diplozoa 

Order 1. Distomatida [e.g. Hexamita, Spironucleus, Trepomonas] 

Order 2. Giardiida [e.g. Giardia, Octomitus] 

Subclass 2. Enteromonadia 

Order 1. Enteromonadida [e.g. Enteromonas] 

Class 2. Retortamonadea 

Order 1. Retortamonadida [e.g. Chilomastix, Retortamonas] 

SUPERPHYLUM 3. DISCICRISTATA 

Phylum 1. Percolozoa 

Class 1. Heterolobosea 

Order 1. Schizopyrenida [ e.g. Naegleria, Tetramitus, Vahlkampfia] 

Order 2. Acrasida [ e.g. Acrasis] 

Order 3. Lyromonadida [ e.g. Lyromonas, Psalteriomonas] 

Class 2. Percolatea 

Order 1. Percolomonadida [e.g. Percolomonas] 

Order 2. Pseudociliatida [e.g. Stephanopogon] 

Phylum 2. Euglenozoa 

Subphylum 1. Plicostoma 

Class 1. Euglenoidea 

Order 1. Petalomonadida [e.g. Calycimonas, Petalomonas] 

Order 2. Peranemida [e.g. Entosiphon, Peranema] 

Order 3. Rhabdomonadida [e.g. Distigma, Menoidium] 

Order 4. Euglenida [e.g. Astasia, Euglena, Eutreptia, Phacus] 

Class 2. Diplonemea 

Order 1. Diplonemida [e.g. Diplonema, Rhynchopus] 

Subphylum 2. Saccostoma 

Class 1. Kinetoplastea 

Order 1. Bodonida [e.g. Bodo, Cryptobia, Dimastigella, Ichthyobodo] 

Order 2. Trypanosomatida [e.g. Crithidia, Leishmannia, Trypanosoma] 

Class 2. Postgaardea 

Order 1. Postgaadida [e.g. Calkinsia, Postgaardi] 

Infrakingdom 2. Alveolata 

Phylum 1. Myzozoa 

Subphylum 1. Dinozoa 

Infraphylum 1. Protalveolata 

Class 1. Colponemea [e.g. Algovora, Colponema] 

Class 2. Myzomonadea [e.g. Alphamonas, Chilovora, Voromonas] 

Class 3. Perkinsea [e.g. Parvilucifera, Perkinsus, Phagodinium, Rastromonas] 

Class 4. Ellobiopsea [e.g. Elliobiopsis, Thalassomyces] 

Infraphylum 2. Dinoflagellata 

Superclass 1. Syndina 

Class 1. Syndinea [e.g. Amoebophrya] 

Superclass 2. Dinokaryota 

Class 1. Noctilucea [e.g. Noctiluca] 

Class 2. Peridinea 

Subclass 1. Peridinoidia [e.g. Amylodinium, Heterocapsa, Prorocentrum] 

Subclass 2. Dinophysoidia [e.g. Dinophysis] 

Subclass 3. Gonyaulacoidia 

Order 1. Gonyaulacida [e.g. Ceratium, Cryptothecodinium] 

Subclass 4. Suessioidia 

Order 1. Suessiida [e.g. Polarella, Symbiodinium] 

Subclass 5. Oxyrrhia 

Order 1. Oxyrrhida [e.g. Oxyrrhis] 

Subphylum 2. Apicomplexa 

Infraphylum 1. Apicomonada 

Class 1. Apicomonadea [ e.g. Acrocoelus, Colpodella] 

Infraphylum 2. Sporozoa 

Class 1. Coccidea [e.g. Cryptosporidium, Hepatozoon, Toxoplasma] 


Class 2. Gregarinea [e.g. Monocystis, Ophriocystis] 

Class 3. Haematozoa [e.g. Babesia, Plasmodium, Theileria] 

Phylum 2. Ciliophora 

Subphylum 1. Postciliodesmatophora 

Class 1. Karyorelictea [e.g. Kentrophoros, Loxodes, Tracheloraphis] 

Class 2. Heterotrichea [e.g. Blepharisma, Folliculina, Stentor] 

Subphylum 2. Intramacronucleata 

Infraphylum 1. Spirotrichia 

Class 1. Spirotrichea [e.g. Euplotes, Metopus, Oxytricha, Tintinnus] 

Infraphylum 2. Rhabdophora 

Class 1. Litostomatea [e.g. Didinium, Entodinium, Lacrymaria] 

Infraphylum 3. Ventrata 

Class 1. Phyllopharyngea [e.g. Dysteria, Podophrya] 

Class 2. Colpodea [e.g. Colpoda] 

Class 3. Nassophorea [e.g. Nassula] 

Class 4. Prostomatea [e.g. Coleps] 

Class 5. Plagiopylea 

Class 6. Oligohymenophorea [e.g. Paramecium, Tetrahymena, Vorticella] 

Kingdom 2. Animalia 

Subkingdom 1. Radiata 

Infrakingdom 1. Spongiaria 

Phylum 1. Porifera 

Subphylum 1. Hyalospongiae 

Subphylum 2. Calcispongiae 

Subphylum 3. Archaeocyatha 

Infrakingdom 2. Coelenterata 

Phylum 1. Cnidaria 

Subphylum 1. Anthozoa 

Subphylum 2. Medusozoa 

Phylum 2. Ctenophora 

Infrakingdom 3. Placozoa 

Phylum 1. Placozoa 

Subkingdom 2. Myxozoa 

Phylum 1. Myxosporidia 

Subkingdom 3. Bilateria 

Branch 1. PROTOSTOMIA 

Infrakingdom 1. Lophozoa 

SUPERPHYLUM POLYZOA 

Phylum 1. Bryozoa 

Subphylum 1. Stelmatopoda 

Subphylum 2. Lophopoda 

Phylum 2. Kamptozoa 

Subphylum 1. Entoprocta 

Subphylum 2. Cycliophora 

SUPERPHYLUM CONCHOZOA 

Phylum 1. Mollusca 

Subphylum 1. Bivalvia 

Subphylum 2. Glossophora 

Infraphylum 1. Univalvia 

Infraphylum 2. Spiculata 

Infraphylum 3. Cephalopoda 

Phylum 2. Brachiozoa 

Subphylum 1. Brachiopoda 

Subphylum 2. Phoronida 

SUPERPHYLUM 3. SIPUNCULA 

Phylum 1. Sipuncula 

SUPERPHYLUM 4. VERMIZOA 

Phylum 1. Annelida 

Subphylum 1. Polychaeta 

Subphylum 2. Clitellata 

Subphylum 3. Echiura 

Subphylum 4. Pogonophora 


Phylum 2. Nemertina 

Infrakingdom 2. Chaetognathi 

Phylum 1. Chaetognatha 

Infrakingdom 3. Ecdysozoa 

SUPERPHYLUM 1. HAEMOPODA 

Phylum 1. Arthropoda 

Subphylum 1. Cheliceromorpha 

Infraphylum 1. Pycnogonida 

Infraphylum 2. Chelicerata 

Subphylum 2. Trilobitomorpha 

Subphylum 3. Mandibulata 

Infraphylum 1. Crustacea 

Infraphylum 2. Myriapoda 

Infraphylum 3. Insecta 

Phylum 2. Lobopoda 

Subphylum 1. Onychophora 

Subphylum 2. Tardigrada 

SUPERPHYLUM NEMATHELMINTHES 

Phylum Nemathelminthes 

Subphylum 1. Scalidorhyncha 

Infraphylum 1. Priapozoa 

Infraphylum 2. Kinorhyncha 

Subphylum 2. Nematoida 

Infraphylum 1. Nematoda 

Infraphylum 2. Nematomorpha 

Infrakingdom 4. Platyzoa 

Phylum 1. Acanthognatha 

Subphylum 1. Trochata (Gnathifera) 

Infraphylum 1. Rotifera 

Infraphylum 2. Acanthocephala 

Subphylum 2. Monokonta 

Phylum 2. Platyhelminthes 

Subphylum 1. Turbellaria 

Infraphylum 1. Mucorhabda 

Infraphylum 2. Rhabditophora 

Subphylum 2. Neodermata 

Infraphylum 1. Trematoda 

Infraphylum 2. Cercomeromorpha 

BRANCH 2. DEUTEROSTOMIA 

Infrakingdom 1. Coelomopora 

Phylum 1. Hemichordata 

Subphylum 1. Pterobranchia 

Subphylum 2. Enteropneusta 

Phylum 2. Echinodermata 

Subphylum 1. Homalozoa 

Subphylum 2. Pelmatozoa 

Infraphylum 1. Blastozoa 

Infraphylum 2. Crinozoa 

Subphylum 3. Eleutherozoa 

Infraphylum 1. Asterozoa 

Infraphylum 4. Echinozoa 

Infrakingdom 2. Chordonia 

Phylum 1. Urochorda 

Subphylum 1. Tunicata 

Infraphylum 1. Ascidiae 

Infraphylum 2. Thaliae 

Subphylum 2. Appendicularia 

Phylum 2. Chordata 

Subphylum 1. Acraniata 

Infraphylum 1. Cephalochordata 

Infraphylum 2. Conodonta 

Subphylum 2. Vertebrata 

Infraphylum 1. Agnatha 


Infraphylum 2. Gnathostomata 

Subkingdom 4. Mesozoa 

Phylum 1. Mesozoa 

Kingdom 3. Fungi 

Subkingdom 1. Eomycota 

Phylum 1. Archemycota 

Subphylum 1. Dictyomycotina 

Class 1. Chytridiomycetes 

Class 2. Enteromycetes 

Subphylum 2. Melanomycotina 

Infraphylum 1. Allomycotina 

Class 1. Allomycetes 

Infraphylum 2. Zygomycotina 

Superclass 1. Eozygomycetia 

Class 1. Bolomycetes 

Class 2. Glomomycetes 

Superclass 2. Neozygomycetia 

Class 1. Zygomycetes 

Class 2. Zoomycetes 

Phylum Microsporidia 

Class 1. Minisporea 

Class 2. Microsporea 

Subkingdom 2. Neomycota 

Phylum 1. Ascomycota 

Subphylum 1. Hemiascomycotina 

Class 1. Taphrinomycetes 

Class 2. Geomycetes 

Class 3. Endomycetes 

Subphylum 2. Euascomycotina 

Class 1. Discomycetes 

Class 2. Pyrenomycetes 

Class 3. Loculomycetes 

Class 4. Plectomycetes 

Phylum 2. Basidiomycota 

Subphylum 1. Septomycotina 

Class 1. Septomycetes 

Subphylum 2. Orthomycotina 

Superclass 1. Hemibasidiomycetia 

Class 1. Ustomycetes 

Superclass 2. Hymenomycetia 

Class 1. Gelimycetes 

Class 2. Homobasidiomycetes 

Kingdom 4. Plantae 

Subkingdom 1. Biliphyta 

Infrakingdom 1. Glaucophyta 

Phylum 1.Glaucophyta [e.g. Cyanophora] 

Infrakingdom 2. Rhodophyta 

Phylum 1. Rhodophyta 

Subphylum 1. Rhodellophytina 

Class 1. Rhodellophyceae [e.g. Porphyridium] 

Subphylum 2. Macrorhodophytina 

Class 1. Bangiophyceae [e.g. Bangia, Porphyra] 

Class 2. Florideophyceae [e.g. Batrachospermum, Corallina] 

Subkingdom 2. Viridiplantae 

Infrakingdom 1. Chlorophyta 

Phylum 1. Chlorophyta 

Subphylum 1. Chlorophytina 

Infraphylum 1. Prasinophytae 

Class 1. Micromonadophyceae [e.g. Mesostigma, Micromonas] 

Class 2. Nephrophyceae [e.g. Nephroselmis, Pseudoscourfieldia] 


Infraphylum 2. Tetraphytae 

Class 1. Chlorophyceae [e.g. Chlamydomonas, Tetraselmis] 

Class 2. Trebouxiophyceae [e.g. Chlorella] 

Class 3. Ulvophyceae [e.g. Acetabularia, Bryopsis, Codium, Ulva] 

Subphylum 2. Phragmophytina 

Infraphylum 1. Charophytae 

Class 1. Charophyceae [e.g. Chara, Nitella] 

Infraphylum 2. Rudophytae 

Class 1. Eophyceae [e.g. Coleochaete, Klebsormidium] 

Class 2. Conjugophyceae [e.g. Spirogyra] 

Infrakingdom 2. Cormophyta 

Phylum 1. Bryophyta 

Subphylum 1. Hepaticae 

Subphylum 2. Anthocerotae 

Subphylum 3. Musci 

Phylum 2. Tracheophyta 

Subphylum 1. Pteridophytina 

Infraphylum 1. Psilophytae 

Infraphylum 2. Lycophytae 

Infraphylum 3. Sphenophytae 

Infraphylum 4. Filices 

Subphylum 2. Spermatophytina 

Infraphylum 1. Gymnospermae 

Infraphylum 2. Angiospermae 

Kingdom 5. Chromista 

Subkingdom 1. Cryptista 

Phylum 1. Cryptista 

Subphylum 1. Cryptomonada 

Class 1. Cryptophyceae [e.g. Chilomonas, Cryptomonas, Guillardia] 

Class 2. Goniomonadea [e.g. Goniomonas] 

Subphylum 2. Leucocrypta 

Class 1. Leucocryptea [e.g. Kathablepharis, Leucocryptos] 

Subkingdom 2. Chromobiota 

Infrakingdom 1. Heterokonta 

Phylum 1. Ochrophyta 

Subphylum 1. Phaeista 

Infraphylum 1. Hypogyrista 

Class 1. Pelagophyceae [e.g. Pelagomonas, Sarcinochrysis] 

Class 2. Actinochrysea (Dictyochophyceae) [e.g. Dictyocha, Pedinella] 

Class 3. Pinguiophyceae [e.g. Glossomastix, Pinguiochrysis] 

Infraphylum 2. Chrysista 

Class 1. Raphidophyceae [e.g. Heterosigma] 

Class 2. Eustigmatophyceae [e.g. Vischeria] 

Class 3. Chrysophyceae [e.g. Ochromonas, Oikomonas, Spumella, Synura] 

Class 4. Chrysomerophyceae [e.g. Chrysomeris, Giraudyopsis] 

Class 5. Phaeothamniophyceae [e.g. Phaeothamnion, Pleurochloridella] 

Class 6. Xanthophyceae [e.g. Chloromeson, Vaucheria] 

Class 7. Phaeophyceae [e.g. Fucus, Laminaria] 

Subphylum 2. Khakista 

Class 1. Bolidophyceae [e.g. Bolidomonas] 

Class 2. Diatomeae [e.g. Coscinodiscus, Bacillaria, Nitzschia] 

Phylum 2. Bigyra 

Subphylum 1. Bigyromonada 

Class 1. Bigyromonadea [e.g. Developopayella] 

Subphylum 2. Pseudofungi 

Class 1. Oomycetes [e.g. Achlya, Phytophthora] 

Class 2. Hyphochytrea [e.g. Rhizidiomyces] 

Subphylum 3. Opalinata 

Class 1. Proteromonadea [e.g. Proteromonas] 

Class 2. Blastocystea [e.g. Blastocystis] 

Class 3. Opalinea [e.g. Cepedea, Opalina] 

Phylum 3. Sagenista 


Class 1. Labyrinthulea [e.g. Labyrinthula, Thraustochytrium] 

Class 2. Bisoecea [e.g. Bicosoeca, Caecitellus, Cafeteria] 

Class 3. Placididea [e.g. Pendulomonas, Placidia, Wobblia] 

Infrakingdom 2. Haptophyta 

Phylum 1. Haptophyta 

Class 1. Pavlovophyceae [e.g. Pavlova] 

Class 2. Prymnesiophyceae [e.g. Emiliania, Isochrysis, Prymnesium] 

Phylum 2. Heliozoa [e.g. Acanthocystis, Acanthophrys] 


Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100640 (VWL9231): Waitata, 

Russell, Bay of Islands 9 Feb 1948 - on 

C. maschalocarpum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Syncoryne reinkei R.Nielsen & 

P.M.Pedersen 

CHR401337: (slide CHR1203-1209) 

Kaikoura, 5 Sept 1979, O.Moestrup 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100639 (VWL9135): Pawa Bay, 

Russell, Bay of Islands, 18 Jan 1948 - 

on C. maschalocarpum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100636 (VWL9048): Long Beach, 

Russell, Bay of Islands, 1 Jan 1948 - on 

Carpophyllum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU 100641 (VWL9324): Long Beach, 

Russell, Bay of Islands, 11 Feb 1948 - 

on C.plumosum 



on C.plumosum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100643 (VWL9379): Temple Bar, 


on C.plumosum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. WELT A001108: Russell, Long 

Beach, 12 Jan 1948, VWLindauer 


Russell, Bay of Islands, 14 Mar 1948 - 

on Carpophyllum plumosum 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. WELT A017693: North east end, 

Heaphy Shoal, Chatham Island, 04 

Nov 1986, CH Hay 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. WELT A003919: Island Bay, 26 Aug 

1970, N.Adams - on Lessonia 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100647 (VWL9823): Temple Bar, 


on Pachymenia 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100644 (VWL9652): Long Beach 

Russell, Bay of Islands, 9 Mar 1948 - on 

Pachymenia 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100645 (VWL9671): Long Beach, 


on Pachymenia 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100648 (VWL10159): Long Beach, 

Russell, Bay of Islands, 27 Apr 1948 - 

on Pachymenia 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100650 (VWL13298): Stormy Bay, 

Russell, Bay of Islands, 20 Jan 1953 - 

on Pachymenia 

Division Class Order Family Genus Species Authority AK CHR Te Papa 

Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zelandiae V.J.Chapm. AKU100651 (VWL13330): Kaikoura, 

U.V.Dellow, Dec 1949 - on Lessonia 

APPENDIX 2:

Chlorophyta Bryopsidophyceae Bryopsidales Ostreobiaceae Ostreobium quekettii Bornet & Flahault CHR401340: (slide) Kakanui, 18 

May 1980, O.Moestrup 

Chlorophyta Bryopsidophyceae Bryopsidales Ostreobiaceae Ostreobium quekettii Bornet & Flahault CHR401339: (slide) Portobello, 17 


Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Gomontia polyrhiza (Lagerh.) Bornet & 

Flahault 


Flahault 


Flahault 


Flahault 

Chlorophyta Chlorophyceae Phaeophilales Phaeophilaceae Phaeophila dendroides (P.Crouan & H.Crouan) 

Batters 


Batters 


Batters 


Batters 

Chlorophyta Ulvophyceae Ulvales Ulvellaceae Entocladia viridis Reinke VWL13256: Stewart Is., May 1950, - on 

Epymenia 

Chlorophyta ? Endoderma - (?) VWL no number: Harriet Kings, 

Coromandel, 5 Apr 1931 - on 

Pachymenia lusoria 

CHR401340: (slide) Kakanui, 18 


CHR401339: (slide) Portobello, 17 


CHR401338: (slide) Piha, 16 Apr 

1980, O.Moestrup 

CHR401337: (slide) Kaikoura, 5 

Sept 1979, O.Moestrup 

CHR401340: (slide) Kakanui, 18 


CHR401339: (slide) Portobello, 17 


CHR401338: (slide) Piha, 16 Apr 

1980, O.Moestrup 

CHR401337: (slide) Kaikoura, 5 

Sept 1979, O.Moestrup 


Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Eugomontia stelligera R.Nielsen Syntype - CHR219311: Kakanui, 

South I., 18 May 1980, O. Moestrup - 

green empty shell from 

intertidalzone, grown in culture

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A13520: Port William, Stewart 

Is, 29 Jan 1983, W.A.Nelson - on 

Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A7450: RingaRinga, Stewart 

Is, 30 Nov 1959, E.A.Willa - on 

Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A1498: Gore Bay, South Is, 

Nov 1925, R.M.Laing - on 

Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A12982: Tautuku Peninsula, 

SE Otago, 7 Dec 1973, C.H.Hay - on 

Xiphophora gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A7449: Lonneker's Nugget, 

Stewart Is, 29 Jan 1960, E.A.Willa - 

on X. gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A6674: Lonneker's Nugget, 

Stewart Is., 2 Dec 1971, E.Conway & 

N.M.Adams - on Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh Manurewa Reef, Te Awhaite, 

Wairarapa, 4 Nov 1973, N.M.Adams - 

on Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A7945: Pukerua Bay, 

Wellington, 18 Nov 1972, 

N.M.Adams - on Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR315689: Cape Palliser, North I., 

11 Nov 1962, M.J.Parsons - on 

Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A4052a+b: Karaka Bay, 

Wellington, Nov 1970, J.McCredie - 

on Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A6584: Cape Palliser, 7 Nov 

1971, N.M.Adams - on Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR38194: Whareponga, East 

Cape, North I., 12 Dec 1942, 

L.B.Moore - on C. torulosa 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR385155: Titahi Bay, Wellington, 

21 Nov 1942, L.B.Moore - on C. 

retroflexa 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR219394: Lonneker's Bay, 

Stewart I., 2 Dec 1971, M.J.Parsons 

- on C. retroflexa 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR354184: Big Solander I., 17 

Nov 1973, P.N.Johnson - on 

Cystophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR230929: Akitio, North I., 2 Jan 

1972, M.J.Parsons - on C. scalaris 


Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh AK22498 (=ANZE185): Pihama, CHR385154 (=ANZE185): Pihama, WELT A985 (=ANZE185): Pihama, 

Taranaki, North I., 2 Dec 1944, Taranaki, North I., 2 Dec 1944, Taranaki, North I., 2 Dec 1944, 

V.W.Lindauer - on C. retroflexa V.W.Lindauer - on C. retroflexa V.W.Lindauer - on C. retroflexa

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing AK22516: (= ANZE203): Pegasus Bay, 

Stewart I., 6 Oct 1945, V.W.Lindauer - 

on Xiphophora gladiata 

CHR62508 (= ANZE203): Pegasus 

Bay, Stewart I., 6 Oct 1945, 

V.W.Lindauer - on Xiphophora 

gladiata 

WELT A1003 (= ANZE203): Pegasus 

Bay, Stewart I., 6 Oct 1945, 

V.W.Lindauer - on Xiphophora 

gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR62503 (=VWL6698): Stewart I., 

22 Oct 1945, E.Willa - on 

Xiphophora gladiata 

Herponema maculaeforme (J.Agardh) Laing AK146242: Dunedin, S.Berggren - on 

Xiphophora ISOTYPE 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. CHR230702: Shag Pt, Otago, South 

I., 9 Sept 1971, M.J.Parsons - on 

Hormosira 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. CHR219443: Lonneker's Nugget, 


- on Hormosira 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. WELT A6669: Lonneler's Nugget, 

Stewart Is, 3 Dec 1971, E.Conway & 

N.M.Adams 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. WELT A7445: Lonneker's Nugget, 

Stewart Is, 6 Feb 1963, E.Awilla 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR219445: Lonneker's Nugget, 


- on Xiphophora gladiata 

J.Ag.) 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae (as 

pulvinatum 

(Harv.MS) non 

AK30304: ANZE334: Waitangi, Bay of 

Isdlands, North I., 19 Aug 1950, 

VWLindauer - on Hormosira SYNTYPE 

CHR227375 (=ANZE334): Waitangi, WELT A1134 (=ANZE334): Waitangi, 

Bay of Islands, North I., 19 Aug Bay of Islands, North I., 19 Aug 1950, 

1950, V.W.Lindauer - on Hormosira V.W.Lindauer - on Hormosira 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Hecatonema stewartensis V.J.Chapm. AK295761: Chris's Bay, Pegasus, 

Stewart Is, 10 Apr 1948 ex VWL10256 

TYPE 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A15913: Landing Bay, 

Burgess I, Mokohinau Is, 31 Dec 

1984, M.Francis - on X.chondrophylla 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A13850: Cable Bay, 

Doubtless Bay, 27 Oct. 1982, 

W.A.Nelson - on X.chondrophylla 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A13446: Tapeka Point, Bay of 

Islands, 30 Oct 1982, W.A.Nelson - 

on X.chondrophylla 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A18903: Katherine Bay, Great 

Barrier I, 7 Dec 1989, F.I.Dromgoole - 

on X.chondrophylla 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A2509: The Pinnacles, Little 

Barrier I., no date, U.V.Dellow - on 

X.chondrophylla 


Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A6569: Cape Palliser, 


on X.gladiata

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A26060: Doubtful Sound, 

Fiordland, 21 Jan 2000, C.Duffy 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. CHR63461: Kaikoura, South I., 14 

Nov 1948, L.B.Moore 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A25591" Bradshaw Sound, 

Fiordland, 3 Oct 2000, K.Neill 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema sp. WELT A13952: Northeast I, Three 

Kings Is, 25 Nov 1983, M.Francis - 

on Sargassum johnsonii 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Mikrosyphar pachymeniae Lindauer Isotype - CHR68937: Russell, Bay 

of Islands, North I., 1 Apr 1944, 

V.W.Lindauer (4267) 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. AK22467: ANZE184 On Ulva lactuca , 

Kaikoura, 31 Dec 1944 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. CHR219406: Lonneker's Bay, 


Xiphophora 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae as "Hecatonema?" in 

ANZE 

AK22532: (=ANZE229) Stewart Is, 14 

Jun 1945, V.W.Lindauer - on 

WELT A1029: (=ANZE229) Stewart 

Is, 14 Jun 1945, V.W.Lindauer 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A13525: Port William, Stewart 

Is, 29 Jan 1983, W.A.Nelson - on 

X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A7446: Ringaringa, Stewart 

Is, 27 Mar 1963, E.A.Willa - on 

X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A6576: Cape Palliser, 


on X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A7784: Ranui Cove, Auckland 

Is, 30 Nov 1972, A.N.Baker - on 

X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A18813a+b: Cape Young, 

Chatham I, 6 Mar 1987, W.A.Nelson - 

on X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A18261: Long I, Dusky 

Sound, Fiordland, 14 May 1986, 

L.A.Bolton - on X.gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR46581: Auckland Is., 26 Dec 

1943, W.Dawbin - on Xiphophora 

gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR67782: Pencarrow Head, Cook 

Strait, North I., 14 Jan 1950, 

L.B.Moore - on Xiphophora gladiata 

(drift) 


Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR52070: Waitangi, Chatham Is., 

12 July 1945, A.M.Rapson - on 

Xiiphophora gladiata 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR230701: Shag Pt, Otago, South 

I,, 8 Oct 1971, M.J.Parsons - on 

Xiphophora gladiata

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Pilinia rimosa Kuetz. WELT A022656: Piha, west 

Auckland, 03 Jul 1994, E.Henry 

Heterokontophyta Phaeophyceae Sphacelariales Sphacelariaceae Sphacelaria pulvinata Hook.f. & Harv. WELT A4369: Wharariki Beach, NW 

Nelson, 19 Mar 1971, F.M.Climo - on 

Carpophyllum maschalocarpum 

Heterokontophyta Phaeophyceae Sphacelariales Sphacelariaceae Sphacelaria pulvinata Hook.f. & Harv. AK146468: ANZE131, on Carpophyllum 

maschalocarpum, Mangonui, 24 Oct 

1942 

WELT A931: (=ANZE131) Mangonui, 

Northland, 24 Oct 1942, 

V.W.Lindauer - on Carpophyllum 

maschalocarpum 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A12907: Katiki, North Otago, 

4 Jun 1973, C.H.Hay 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A6331: Lonneker's Nugget, 

Stewart Is, 26 May 1971, E.Conway 

& N.M.Adams 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A3644a+b: Makara, 

Wellington, 2 Jun 1970, N.M.Adams - 

on drift 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A7447: RiongaRinga, Stewart 

Is, 18 Mar 1960, E.A.Willa 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema sp. WELT A8619: Tasman Bay, Three 

Kings Is, Feb 1974, A.N.Baker - on 

Landsburgia quercifolia 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Nemacystus novae-zelandiae Kylin WELT A18016: Parnell Reef, 

Waitemata Harbour, Auckland, 13 

Oct 1987, K.W.Glombitza - on 

Sargassum scabridum 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK295758: (VWL6253) Stewart Is, 12 

Jun 1945, E.Willa - ISOTYPE 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK22533: (ANZE230) (VWL6253), 

Stewart Is, 12 Jun 1945, E.Willa 

ISOTYPE 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK295759: (VWL6253) Stewart Is, 12 

Jun 1945, E.Willa - TYPE 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South CHR49999: Princess Bay Bay, 

Wellington, North I., 30 May 1943, 

L.B.Moore (drift) 

Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South CHR248256: Oaro, Kaikoura, South 

I., 4 July 1973, M.J.Parsons 


Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A16379: Perserverance 

Harbour, Campbell I, 12 Feb 1985, 

J.C.Yaldwyn 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A984: ANZE 184: Kaikoura, 

31 Dec 1944, V.W.Lindauer 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A18688: Monau, Chatham I, 3 

Mar 1987, W.A.Nelson 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A1592a+b: York Bay, 

Wellington, 31 May 1953, R.K.Dell 

Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A7073: Rosa I, Port Pegasus, 

Stewart Is, 29 Feb 1972, N.M.Adams

galls on Durvillaea CHR243660: Kaitangata, Otago, 

South I., 26 Feb 1973, R.Mason & 

E.M.Chapman - on Durvillaea 

antarctica drift -"galls not caused by 

fungus - possibly bacteria" det. 

J.Kohlmeyer 


galls on Macrocystis CHR47805: Native Island, Paterson 

Inlet, Stewart Island, 2 Dec 1944, 

L.B.Moore - on Macrocystis pyrifera - 

"not fungal but possibly caused by 

filamentous brown algae (see 

Andrews 1976 Biol. Rev. 51: 211- 

253, Can J. Bot 55:1019-1027)" det 

J.Kohlmeyer

Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra woolhousiae Harv. CHR 209060: Lyall Bay, Wellington, 

Sept 1931, Scarfe 

Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra woolhousiae Harv. CHR55566: Hokio Beach, Levin, 

Nov 1946, Moore. 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. AK147200: Bluff, 1874, Berggren 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. AK147201: Ringaringa, Stewart Is, 15 

Jan 1940, L.M.Cranwell 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. AK223832: Preservation Inlet, Fiordland, 

20 Jul 1995, M.S.Morley 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. AK22605: (=ANZE214) Stewart Is, 15 

Jan 1946, V.W.Lindauer 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR24001: Bluff, South I., 4 Jan 

1940, L.B.Moore 

Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra adamsiae W.A.Nelson WELT A10322: Crater Bay, 

Antipodes Is, 23 Nov 1978, C.H.Hay 

Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra adamsiae W.A.Nelson WELT A8038: Port Ross, Auckland 

Is, 19 Feb 1973, K.Johnson, 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon subtumens (J.Agardh ex Laing) 

W.A.Nelson 

WELT A15999: Brighton, Otago, 2 

Feb 1983, W.A.Nelson - on 

D.antarctica 

D.chathamensis 


W.A.Nelson 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon cameronii (W.A.Nelson) 

W.A.Nelson 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon cameronii (W.A.Nelson) 

W.A.Nelson 


W.A.Nelson 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Chlidophyllon kaspar (W.A.Nelson et N.M. 

Adams) W.A.Nelson 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Chlidophyllon kaspar (W.A.Nelson et N.M. 

Adams) W.A.Nelson 

WELT A26849: Three Kings Is, Jan 

1994, V.Staines 

WELT A16714: Princes Rocks, Three 

Kings Is, 18 Jan 1985, M.Francis & 

M.A.Williams 

WELT A26851: Wharekauri, 

Chatham I, Feb 2001, R.Russell 

WELT A17785: Heaphy Shoal, 

Chatham I, 4 Nov 1986, C.H.Hay 

WELT A3669: Makara, Wellington, 2 

Jun 1970, N.M.Adams - on drift 

D.antarctica 

WELT A7466a+b: Point Webb, 

Chatham I, 4 Nov 1986, C.H.Hay - on 

Phylum Class Order Family Genus Species Authority AK CHR Te Papa 

Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Chroodactylon ornatum (C.Agardh) Basson AK30298: (=ANZE 341) Glendowie, 

WELT A1141: (=ANZE 341) 

Auckland, 20 Dec 1949, V.W.Lindauer 

Glendowie, Auckland, 20 Dec 1949, 

V.W.Lindauer 

Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Chroodactylon ornatum (C.Agardh) Basson WELT A6707: Oban, Stewart I, 3 Dec 

1971, E.Conway & N.M.Adams - 

epiphyte 

Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Stylonema alsidii (Zanardini) K.M.Drew WELT A4403: Days Bay, Wellington, 

13 Jun 1971, N.M.Adams - on 

Chaetomorpha 

Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Erythrocladia sp. WELT A18578: Durham & Gap Pts, 

Chatham I, 4 Mar 1987, W.A.Nelson - 

on Cladophora sp. 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Erythrotrichia foliiformis South et N.M.Adams WELT A17692: Petre Bay, Chatham 

I, 4 Nov 1986, C.H.Hay - on Lessonia 

tholiformis 

Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Erythrotrichia foliiformis South et N.M.Adams WELT A6570: Cape Palliser, 


on Marginariella (oval)

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. WELT A3988: Owhiro Bay, 

Wellington, 19 Sep 1970, N.M.Adams 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. CHR357193: Rimu Bay, Pelorus 

Sound, South I., 5 Oct 1958, 

L.B.Moore 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. CHR49433: Matarangi Beach, 

Kuaotunu, Coromandel, North I., 29 

Mar 1945, N.M.Adams 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. CHR248233: Leigh Marine Station, 

North I., 22 May 1974, M.J.Parsons 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. AK290043: Waikawau Bay, Coromandel, 

7 Oct 2004, M.N.Lee 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. AK239454: Henderson Pt, Northland, 1 

Jul 1999, E.K.Cameron 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. CHR191680: Whangamumu 

Harbour, North I., 26 May 1969, 

E.Godley 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. AK147207: (=ANZE46) Bay of Islands, 

14 Aug 1938, V.W.Lindauer 

WELT A846: (=ANZE46) Bay of 

Islands, 14 Aug 1938, V.W.Lindauer 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. WELT A16131: Senecio Pool, Snares 

I, 18 Dec 1984, G.S.Hardy 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. WELT A25595, Deas Cove, 3 Oct 

2000, A.Loughnan 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. WELT A21795: Cascade I, South 

Westland, 21 Feb 1996, D.Neale 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. WELT A4010: Brighton, Otago, 5 Dec 

1970, N.M.Adams 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. WELT A26159: Port Hutt, Chatham 

Is, 12 Mar 2001, W.Nelson, J.Broom, 

W.Jones, T. Farr 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR354186: Big Solander I., 17 Nov 

1973, P.N.Johnson 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR324992: Mangere I., Chatham 

Is, 21 Aug 1968, I. & M.Ritchie 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR509075: Ackers Pt, Stewart I., 4 

Jan 1987, D.R.Given 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR316773: Western Chain, 

Snares Is, 26 Nov 1974, 

D.S.Horning 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR57281: Tautuku, Otago, South 

I., Dec 1947, I.Coulter 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR368067: Secretary I., Doubtful 

Sound, Fiordland, South I., 18 May 

1981, D.J.Brasch 

Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR205876: Blackhead, Dunedin, 

Otago, Dec 1919, W.A.Scarfe 


Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea lyallii Hook.f. et Harv. CHR379617: Bruce Rocks, Brighton, 

Otago, South I., Feb 1948, 

K.W.Allison

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin CHR319934: Atia Point, Kaikoura, 

South I., 14 Nov 1973, M.J.Parsons - 

on L. hookeriana 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR367973: South Bay, Kaikoura, 

South I., 3 Dec 1980, G.D.Fenwick - 

on Pterosiphonia 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR368028: Shap Point, Otago, 

South I., 10 Feb 1981, M.J.Parsons 

& M.Stolp - on Pterosiphonia 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. WELT A17700: McClatchie Reef, 

Chtaham I, 4 Nov 1986, C.H.Hay - on 

Chondria 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR368051: Macrocarpa Point, 

Katiki Beach, Otago, South I., 9 Feb 

1981, M.J.Parsons - on 

Pterosiphonia, drift 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. WELT A17498a+b: Port Webb, 

Chatham I, 6 Nov 1986, C.H.Hay - on 

Chondria 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. CHR230816: Oaro, Kaikoura, South 

I., 6 Nov 1971, M.J.Parsons - on 

Chondria 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. CHR360270: Pier Wharf, Kaikoura, 

South I., 10 Sept 1974, M.J.Parsons - 

on Chondria 

Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248307: Oamaru, South I., 31 

Oct 1972, M.J.Parsons - on 

Heterosiphonia concinna 

Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248099: Penguin Bay, 

Campbell I., 18 Feb 1971, 

C.D.Meurk - on Heterosiphonia 

concinna 

Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR316964: Cod Cavern Gutway, 

Snares Is, 24 Jan 1975, D.S.Horning 

- on Heterosiphonia concinna 

Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248303: Shag Point, Otago, 

South I., 1 Nov 1972, M.J.Parsons - 

on Heterosiphonia concinna 

Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya inconspicua (Reinsch) Schmitz CHR66045: French I., Auckland Is, 

16 Aug 1976, C.A.Fleming - on 

Heterosiphonia berkeleyi (slide only 

No.66) 


Rhodophyta Florideophyceae Hildenbrandiales Hildenbrandiaceae Apophlaea sinclairii Hook.f. et Harv. WELT A13938: West I, Three Kings 

Is, 25 Nov 1983, M.Francis 

Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT A027067: Wairarapa east 

coast, Mataikona reef, Feb 1969, 

N.M.Adams 

Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT AA027066: Cape Palliser, Nov 

1971, N.M.Adams 

Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT A027038: Kaikoura, barbeque 

area just south of Rakautara, Sept 

2004, Nelson, Farr & Neill

Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zelandiae Sparling CHR64545: Eastbourne, Wellington, 

20 Mar 1949, L.B.Moore, 

N.M.Adams & G.F.Papenfuss 

(NB:"Type collection by CHR64545 

not seen by author of species - 

Sparling 1979) - on Gloioderma 

saccatum 

Rhodophyta Florideophyceae Rhodymeniales Champiaceae Champiocolax sp. WELT A18631a+b: Inner Chetwode 

Is, Marlborough, 11 Aug 1987, 

C.H.Hay - on C. chathamensis 

Rhodophyta Florideophyceae Gracilariales Pterocladiophyllaceae Pterocladiophila hemisphaerica K.C.Fan et Papenf. CHR117794: locality unknown - from 

commercial collection, identity 

confirmed by K.C.Fan (UC 

Berkeley); on Pterocladiella 

capillacea 

Rhodophyta Florideophyceae Gigartinales Kallymeniaceae Callocolax sp. CHR367972: South Bay, Kaikoura, 


on Echinothamnion sp. 

Rhodophyta Florideophyceae Gigartinales Kallymeniaceae Callocolax neglectus Schmitz ex Batters CHR248213: Oaro, Kaikpoura, 

South I., 25 Oct 1972, M.J.Parsons - 

on Callophyllis calliblepharoides 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Tylocolax 

microcarpus ? 

CHR219462: Lonneker's Nugget, 

Stewart I., 3 Dec 1971, M.J.Parsons - 

on Adamsiella chauvinii 


microcarpus ? 

CHR368033: Shag Point, Otago, 

South I., 10 Feb 1981, M.J.Parsons 

& M.Stolp - on Adamsiella chauvinii 

chauvinii 


microcarpus ? 

CHR364690: Baxters Reef, 

Kaikoura, South I., 5 Feb 1980, 

G.D.Fenwick - on Adamsiella 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR319388: Curio Bay, SE Otago, 

South I., 16 Feb 1977, M.J.Parsons - 

on Echinothamnion (liq coll) 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR367972: South Bay, Kaikoura, 


on Echinothamnion sp. 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin WELT A18232: George Sound, 

Fiordland, 14 Feb 1987, M.Francis - 

on L. hookeriana 

Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR399500: Katiki Beach, Otago, 

South I., 9 Feb 1981, M.J.Parsons - 

on Polysiphonia rhododactyla (liq 

coll) 


Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin CHR319947: Lighthouse Reef, 

Kaikoura, South I., 13 Nov 1973, 

M.J.Parsons - on L. hookeriana

parasite on 

Rhodophyllis 

CHR364774:Old Wharf, Kaikoura, 

South I., 4 Feb 1980, G.D.Fenwick - 

on Rhodophyllis (liq coll.) 

parasite on 

Rhodophyllis 

CHR364762: South Bay, Kaikoura, 


on Rhodophyllis (liq coll.) 

sanguinea 

G.D.Fenwick (liq coll.) 

parasite on 

Hymenocladia 

CHR364678: Baxters Reef, 

Kaikoura, South I., 5 Feb 1980, 

sanguinea 

parasite on 

Hymenocladia 

CHR219369: Ringaringa, Stewart I., 

30 Nov 1971, M.J.Parsons 

parasite on 

Dasyclonium 

CHR319896: Seal Reef, Kaikoura, 

South I., 6 Oct 1971, M.J.Parsons - 

on Dasyclonium incisum 

oblongifolia 

parasite on 

Cladhymenia 

oblongifolia 

CHR319472: Old Wharf, Kaikoura, 

South I., 13 Nov 1973, V.Hoggard & 

G.D.Fenwick - on Cladhymenia 

on A. lyallii 

parasite on 

Apophlaea lyallii 

CHR219466: Lonneker's Nugget, 

Stewart I., 2 Dec 1971, M.J.Parsons - 

Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax WELT A026739: Te Werahi Beach, 

Northland, North I., 25 Oct 2003, 

W.Nelson 

Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR364760: South Bay, Kaikoura, 


on Plocamium 2x2 fine 

Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR367983: South Bay, Kaikoura, 


on Plocamium 2 x 2 fine 

Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR360413: Katiki Beach, Otago, 

South I., 27 Apr 1975, M.J.Parsons - 

on Plocamium 2x2 


Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zelandiae Sparling WELT A26638: Wharariki Beach, 19 

Mar 2003, W.Nelson & J.Dalen - on 

Gloioderma saccata 

Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zelandiae Sparling WELT A17670: Okawa Beach, 

Chatham I, 6 Jan 1987, A.N.Baker - 

on Gloioderma saccata 

Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zelandiae Sparling WELT A6599: Ringaringa, Stewart I, 

26 Apr 1963, E.A.Willa 

Rhodophyta Florideophyceae Rhodymeniales Rhodymeniaceae Rhodymeniocolax sp. WELT A14130: Antipodes I, 4 Dec 

1978, C.H.Hay - on Rhodymenia 

epimenioides 

Rhodophyta Florideophyceae Rhodymeniales Rhodymeniaceae Rhodymeniocolax sp. WELT A7568: Golden Bay, Paterson 

Inlet, Stewart I, 29 Feb 1960, 

E.A.Willa - on Rhodymenia linearis

parasite on 

Rhodophyllis 

WELT A4167: West Lyall Bay, 

Wellington, 12 Jan 1971, N.M.Adams - 

"cf Ceratocolax" 


parasite on 

WELT A6798: Harrold's Bay, 

Rhodophyllis 

Halfmoon Bay, Stewart Is, 1 Dec 

1971, N.M.Adams - "cf Ceratocolax"

galls on Chaetangium "pycnidia - unfortunately cannot CHR248185a: Monument Harbour, Campbell I., 14 Feb 

be further identified as long as 1971, C.D.Meurk - on Chaetangium fastigiatum 

perfect (ascigerous) state is 

unknown. Many marine 

algicolous Ascomyctes have 

similar pycnidia." det J.Kohlmeyer 

Chaudefaudia corallinarum (Crouan et Crouan) Muller et 

v.Arx 

Chaudefaudia corallinarum (Crouan et Crouan) Muller et 

v.Arx 

det Kohlmeyer CHR248265: Mollymawk Bay, Snares Is, 6 Dec 1974, 

D.S.Horning - on Euptilota formosissma 

det Kohlmeyer CHR248266: Cod Cavern Gutway, Snares Is, 24 Jan 

1975, D.S.Horning - on Euptilota formosissima 

Eurychasma dicksonii (Wright) Magnus Saprolegniales - "forms peculiar 

"netsporangia" with encysted 

zoospores" det Kohlmeyer 

CHR248343: Shag Point, Otago, South I., 8 Sept 1971, 

M.J.Parsons (liq coll + photomicrograph) - on 

Ectocarpus on Scytosiphon (CHR219500) 

Spathulospora lanata Kohlmeyer CHR:64534: Runaround, Wellington, North I., 18 Mar 

1949, N.M.Adams - on Ballia scoparia - det Kohlmeyer 

CHR357143: Open Bay Islands, Westland, South I., 4 

Feb 1976, G.D.Fenwick - on Sargassum undulatum (det 

Kohlmeyer) 

CHR315947c: Houghton Bay, 16 Oct 1962, M.J.Parsons 

- on Sargassum sinclairii (det Kohlmeyer) 

Haloguignardia tumefaciens (Cribb et Herbert) Cribb et 

Cribb 

Haloguignardia tumefaciens (Cribb et Herbert) Cribb et 

Cribb 

Genus species Authority comments CHR 

Mycosphaerella apophlaeae Kohlm. Bot Mar 24: 13- Kohlmeyer & CHR391939: South Promontory, Snares Is, 14 Dec 

Demoulin 1981 

1974, C.E.Holmes 

Polystigma apophlaeae Kohlm. Bot Mar 24: 13- Kohlmeyer & Herb - Holotype NY 

Demoulin 1981

APPENDIX 5: 

Data storage 

Dataset supplied to the Ministry in the form of an Access database and an electronic copy of 

the report. 

Utility of the Access database 

The database was operated at NIWA through a Delphi web application, enabling multiple 

users. Below are examples of the web interface pages we used, configured for data entry. 

Search functions will need to be developed as part of the front end of this database.

Diseases, pathogens and parasites of Undaria pinnatifida

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