Diseases, pathogens and parasites of Undaria pinnatifida
Diseases, pathogens and parasites of Undaria pinnatifida
Diseases, pathogens and parasites of Undaria pinnatifida
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<strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong><br />
<strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong><br />
MAF Biosecurity New Zeal<strong>and</strong> Technical Paper No: 2009/44<br />
Authors:<br />
Neill, K., Heesch, S., Nelson, W.<br />
National Institute <strong>of</strong> Water <strong>and</strong> Atmospheric Research, Private<br />
Bag 14-901, Wellington<br />
Prepared for BNZ Post-clearance Directorate<br />
By National Institute <strong>of</strong> Water <strong>and</strong> Atmospheric Research<br />
As contract No: ZBS2005-01<br />
ISSN 1176-838X (Print)<br />
ISSN 1177-6412 (Online)<br />
ISBN 978-0-478-35752-3(Print)<br />
ISBN 978-0-478-35753-0(Online<br />
April 2008
Disclaimer<br />
While every effort has been made to ensure the information in this publication is accurate, the<br />
Ministry <strong>of</strong> Agriculture <strong>and</strong> Forestry does not accept any responsibility or liability for error or<br />
fact omission, interpretation or opinion which may be present, nor for the consequences <strong>of</strong><br />
any decisions based on this information.<br />
Any view or opinions expressed do not necessarily represent the <strong>of</strong>ficial view <strong>of</strong> the Ministry<br />
<strong>of</strong> Agriculture <strong>and</strong> Forestry.<br />
The information in this report <strong>and</strong> any accompanying documentation is accurate to the best <strong>of</strong><br />
the knowledge <strong>and</strong> belief <strong>of</strong> the National Institute <strong>of</strong> Water & Atmospheric Research Ltd<br />
(NIWA) acting on behalf <strong>of</strong> the Ministry <strong>of</strong> Agriculture <strong>and</strong> Forestry. While NIWA has<br />
exercised all reasonable skill <strong>and</strong> care in preparation <strong>of</strong> information in this report, neither<br />
NIWA nor the Ministry <strong>of</strong> Agriculture <strong>and</strong> Forestry accept any liability in contract, tort or<br />
otherwise for any loss, damage, injury, or expense, whether direct, indirect or consequential,<br />
arising out <strong>of</strong> the provision <strong>of</strong> information in this report.<br />
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P O Box 2526<br />
WELLINGTON<br />
Telephone: (04) 474 4100<br />
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This publication is also available on the MAF website at www.maf.govt.nz/publications<br />
© Crown Copyright - Ministry <strong>of</strong> Agriculture <strong>and</strong> Forestry
Contents Page<br />
Executive Summary 1<br />
Overall objective 2<br />
Specific objectives 2<br />
1. Introduction 3<br />
2. Methods <strong>and</strong> Materials 5<br />
2.1. Definitions 5<br />
2.2. Data Sources 6<br />
2.3. Database 7<br />
2.4. Mapping 8<br />
3. Results 8<br />
3.1. General Comments 8<br />
3.2. description <strong>of</strong> known pathogen-host relationships 9<br />
3.3. Laminariales 13<br />
3.4. Brown algae other than Laminariales 16<br />
3.5. Red algae 20<br />
3.6. Green algae 27<br />
3.7. Xanthophyceae 28<br />
4. Discussion 30<br />
I: Assessment <strong>of</strong> information available on seaweed diseases worldwide <strong>and</strong> in New Zeal<strong>and</strong> 30<br />
II: Assessment <strong>of</strong> threats by <strong>pathogens</strong> <strong>of</strong> <strong>Undaria</strong> to New Zeal<strong>and</strong> native marine flora 31<br />
III: Future strategy for screening populations <strong>and</strong> increasing knowledge <strong>of</strong> risk posed by<br />
diseases/<strong>parasites</strong>/<strong>pathogens</strong> to New Zeal<strong>and</strong> macroalgae <strong>and</strong> coastal communities 32<br />
5. Conclusions 33<br />
6. Acknowledgements 33<br />
7. References 34<br />
i
Executive Summary<br />
A detailed desk study was carried out on diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>and</strong><br />
other macroalgae. In additional to published literature, data sources included specimens<br />
housed in New Zeal<strong>and</strong> herbaria, <strong>and</strong> information obtained through email <strong>and</strong> personal<br />
contacts. An Access database was established to enter data from the relevant papers <strong>and</strong> to<br />
record details <strong>of</strong> the diseases, <strong>parasites</strong> <strong>and</strong> <strong>pathogens</strong>. The database contains a complete<br />
listing <strong>of</strong> all papers considered (927 references) <strong>of</strong> which 549 pertinent papers are included in<br />
the reference list in this report.<br />
The information on diseases <strong>of</strong> seaweeds is very patchy <strong>and</strong> the emphasis <strong>of</strong> published work<br />
lies in two main areas: diseases occurring in monocultures <strong>of</strong> farmed species, mainly in East<br />
<strong>and</strong> Southeast Asia (particularly affecting the key economic genera Porphyra, Laminaria,<br />
<strong>Undaria</strong>, Gracilaria, Eucheuma <strong>and</strong> Kappaphycus), <strong>and</strong> observations <strong>of</strong> certain groups <strong>of</strong><br />
<strong>pathogens</strong> in particular geographic regions as a consequence <strong>of</strong> the research interests <strong>of</strong> a<br />
particular team or research group, leading to “pockets <strong>of</strong> information”. The amount <strong>of</strong><br />
information contained in the references we investigated varied greatly between articles,<br />
ranging from reports <strong>of</strong> the occurrence <strong>of</strong> <strong>pathogens</strong> to multi-paper treatments <strong>of</strong> certain<br />
diseases. The latter are especially numerous for farmed macroalgae e.g. Pythium porphyrae,<br />
the agent causing the red rot disease in Porphyra species (Porphyra cultivation is a billion<br />
dollar industry in Asian countries). Other agents, in contrast, have only been observed once<br />
<strong>and</strong> <strong>of</strong>ten only incidentally in the course <strong>of</strong> other research.<br />
The only disease reported in <strong>Undaria</strong> from its introduced range is the infection <strong>of</strong> thalli with<br />
the pigmented endophytic brown alga Laminariocolax aecidioides, both in Spain (Veiga et al.<br />
1997) <strong>and</strong> in Argentina (Gauna et al. personal communication). It is not clear whether this<br />
endophyte originates from Japanese populations introduced with the host or from European or<br />
Argentinian populations respectively. Laminariocolax aecidioides is known from other,<br />
native European kelps such as Laminaria hyperborea in the German Bight <strong>and</strong> Norway, <strong>and</strong><br />
Saccharina latissima in the Western Baltic Sea (Lein et al. 1991; Ellerstdottir & Peters 1995,<br />
1997; Peters & Schaffelke 1996), but it has not been reported from southern Europe. It also<br />
occurs in the native range <strong>of</strong> U. <strong>pinnatifida</strong>, in Japan (Yoshida & Akiyama 1978). Genetic<br />
studies may determine the origin <strong>of</strong> the Spanish <strong>and</strong> Argentinean populations <strong>and</strong> thus shed<br />
some light on whether endophytes were or can be transmitted with host sporophytes (or other<br />
disease agents).<br />
None <strong>of</strong> the known <strong>pathogens</strong> <strong>of</strong> <strong>Undaria</strong> have so far been observed in/on U. <strong>pinnatifida</strong> in<br />
New Zeal<strong>and</strong>, however, populations <strong>of</strong> U. <strong>pinnatifida</strong> around New Zeal<strong>and</strong> have not been<br />
screened for the presence <strong>of</strong> diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong>. Given that there is evidence<br />
that New Zeal<strong>and</strong> has received at least 10 separate introduction events <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong><br />
(Uwai et al. 2006), it would be important to construct a sampling regime that reflected this<br />
known genetic diversity within New Zeal<strong>and</strong> populations <strong>of</strong> <strong>Undaria</strong>.<br />
Seaweeds that are diseased are under-collected in New Zeal<strong>and</strong> <strong>and</strong>, as a consequence, the<br />
status <strong>of</strong> knowledge about biotic diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> is deficient: it is not<br />
possible to evaluate risk posed by introduced diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> on the basis <strong>of</strong><br />
current underst<strong>and</strong>ing <strong>of</strong> the native biota. Whilst experts in the field <strong>of</strong> algal diseases such as<br />
Correa (1997) stress the need for studies on the mechanisms <strong>of</strong> infection <strong>and</strong> the spread <strong>of</strong> the<br />
<strong>pathogens</strong> within <strong>and</strong> among host individuals, as well as on the genetics <strong>of</strong> the host-pathogen<br />
interaction, the basic underpinning surveys <strong>and</strong> research are required in New Zeal<strong>and</strong> to<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 1
document the biodiversity <strong>and</strong> distribution <strong>of</strong> diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> within<br />
macroalgae.<br />
OVERALL OBJECTIVE:<br />
To determine <strong>and</strong> assess the threats that known diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong><br />
<strong>pinnatifida</strong> pose to native New Zeal<strong>and</strong> macroalgae<br />
SPECIFIC OBJECTIVES:<br />
1. To undertake a review (literature, email, telephone) <strong>and</strong> map the known distribution <strong>of</strong> the<br />
diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> which have been recorded to affect <strong>Undaria</strong> <strong>pinnatifida</strong><br />
(<strong>and</strong>/or closely related members <strong>of</strong> the Laminariales)<br />
a. in its native range (Japan, Korea <strong>and</strong> the Kamchatka Peninsula <strong>of</strong> Russia), <strong>and</strong>,<br />
b. in its introduced range (Australia, United Kingdom, France, USA, Argentina, New<br />
Zeal<strong>and</strong>).<br />
2. To undertake a review (literature, email, telephone) <strong>and</strong> map the geographic distribution <strong>of</strong><br />
the status <strong>of</strong> known diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> in macroalgae.<br />
3. To determine if any <strong>of</strong> the diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> identified in specific objective<br />
1 are present in New Zeal<strong>and</strong>.<br />
4. To determine if any <strong>of</strong> the diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> identified in specific objective<br />
2 are present in native New Zeal<strong>and</strong> macroalgae.<br />
2 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
1. Introduction<br />
<strong>Undaria</strong> <strong>pinnatifida</strong> is a large kelp (Laminariales, Phaeophyceae) native to the north western<br />
Pacific (Japan, Korea, China <strong>and</strong> the Kamchatka Peninsula <strong>of</strong> Russia) (Akiyama & Kurogi<br />
1982; Silva et al. 2002; Guiry & Guiry 2007). It was introduced to Europe in the 1970s<br />
associated with the transport <strong>of</strong> oysters from Asia (Perez et al. 1981; Bourdouresque et al.<br />
1985; Castric-Fay et al. 1993; Fletcher & Farrell 1998). In the 1980s <strong>Undaria</strong> was recorded in<br />
New Zeal<strong>and</strong> (Hay & Luckens 1987; Hay 1990), Tasmania, Australia (S<strong>and</strong>erson 1990), in<br />
the 1990s in Argentina (Casas & Piriz 1996), Victoria, Australia (Campbell & Burridge<br />
1998), <strong>and</strong> in the 2000s from California, USA (Silva et al. 2002) <strong>and</strong> Baja California, Mexico<br />
(Aguilar-Rosas et al. 2004).<br />
Since its detection in New Zeal<strong>and</strong>, <strong>Undaria</strong> has spread primarily by human-mediated vectors<br />
such as vessel hulls <strong>and</strong> marine farming equipment. This species has the potential to displace<br />
native macroalgae (environmental impact), alter habitat for commercial species<br />
(environmental <strong>and</strong> economic impact), disrupt aquaculture activities (economic impact) <strong>and</strong><br />
may affect the cultural values <strong>of</strong> particular sites.<br />
At present, <strong>Undaria</strong> in New Zeal<strong>and</strong> has been reported from Great Barrier Isl<strong>and</strong>, Auckl<strong>and</strong><br />
(Waitemata Harbour), Corom<strong>and</strong>el, Tauranga, Gisborne, Napier, Port Taranaki, Wellington<br />
<strong>and</strong> the Wellington region <strong>of</strong> Cook Strait in the North Isl<strong>and</strong>, in the Marlborough Sounds,<br />
Nelson, Golden Bay, Kaikoura, Lyttelton, Akaroa, Timaru, Oamaru, Dunedin Harbour, Bluff<br />
in the South Isl<strong>and</strong> <strong>and</strong> also from Stewart Isl<strong>and</strong> <strong>and</strong> the Snares Isl<strong>and</strong>s. The potential exists<br />
for this species to be spread further to regions regarded as having high conservation values<br />
such as the other sub-Antarctic isl<strong>and</strong>s, Stewart Isl<strong>and</strong> (apart from Paterson Inlet <strong>and</strong> Oban),<br />
the Chatham Isl<strong>and</strong>s, Fiordl<strong>and</strong>, Abel Tasman National Park, as well as the isl<strong>and</strong>s <strong>of</strong> the<br />
Hauraki Gulf, <strong>and</strong> <strong>of</strong>fshore isl<strong>and</strong>s <strong>of</strong> north eastern New Zeal<strong>and</strong>. A genetic study <strong>of</strong> <strong>Undaria</strong><br />
populations in New Zeal<strong>and</strong> has revealed multiple introductions have occurred with 10<br />
distinct haplotypes present in New Zeal<strong>and</strong>, although only a single haplotype was found in<br />
current North Isl<strong>and</strong> populations (Uwai et al. 2006).<br />
As well as the direct impact this species has on indigenous biota, it also poses a potential risk<br />
to native New Zeal<strong>and</strong> macroalgae through diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong>. Infectious<br />
diseases in macroalgae are caused by a wide variety <strong>of</strong> organisms ranging from viruses,<br />
bacteria, cyanobacteria, fungi (phycomycetes, ascomycetes, fungi imperfecti),<br />
heterokontophytes (oomycetes, labyrinthulids), nematodes, protozoans, through to endophytic<br />
<strong>and</strong> parasitic macroalgae (Chlorophyta, Phaeophyceae, Rhodophyta) (Andrews 1976; G<strong>of</strong>f<br />
1982; Apt 1988b; Correa 1994; Bouarab et al. 2001a; Van Etten et al. 2002). In addition, the<br />
grazing <strong>of</strong> macroalgae by herbivores results in vulnerability to disease/access for pathogenic<br />
organisms. Within its native distribution <strong>Undaria</strong> has a number <strong>of</strong> known diseases, <strong>pathogens</strong><br />
<strong>and</strong> <strong>parasites</strong> (e.g. Yoshida & Akiyama 1978; Rho et al. 1993; Jiang et al. 1997). <strong>Undaria</strong>, as<br />
a member <strong>of</strong> the Laminariales, has a heteromorphic life history with the 2 life stages having<br />
entirely different morphologies – the sporophyte grows up to several metres in length whereas<br />
the gametophyte is microscopic growing to only several hundred microns in size. Any<br />
consideration <strong>of</strong> diseases in macroalgae needs to consider the vulnerability <strong>of</strong> different life<br />
history stages to diseases/<strong>pathogens</strong>/<strong>parasites</strong>.<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 3
Biological invasions are understood to pose a significant threat to biodiversity, <strong>and</strong> <strong>parasites</strong><br />
are considered to play a role in determining the outcomes <strong>of</strong> invasions. Transmission <strong>of</strong><br />
<strong>parasites</strong> to native species from the invading species can influence the fitness <strong>of</strong> native taxa,<br />
mediating competitive interactions. Introduced diseases may have catastrophic impacts or<br />
may result in persistent <strong>and</strong> sub-lethal effects on natives <strong>and</strong> consequent impacts on<br />
community structure (Prenter et al. 2004). Introduced hosts may also play a role as reservoirs<br />
for native diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> from which potentially deleterious “spillback” <strong>of</strong><br />
infection to native hosts may occur (Prenter et al. 2004; Tompkins & Poulin 2006). Tompkins<br />
& Poulin (2006) observe that although many <strong>parasites</strong> are apparently lost from hosts when<br />
they are introduced to a new environment, the introduced hosts tend to acquire generalist<br />
<strong>parasites</strong> from the native biota. Impacts <strong>of</strong> disease are <strong>of</strong>ten dependent on the context, with<br />
multiple abiotic <strong>and</strong> biotic factors implicated in the emergence <strong>of</strong> <strong>parasites</strong>, invasion<br />
processes, <strong>and</strong> the impacts experienced by native biota (Blaustein & Kiesecker 2002). Factors<br />
which increase host susceptibility to infection, including a range <strong>of</strong> stressors such as habitat<br />
alteration <strong>and</strong> degradation, may make them more prone to introduced <strong>parasites</strong>. Artificial<br />
rearing <strong>and</strong> aquaculture increase the potential for disease transmission as well as increasing<br />
potential host susceptibility in crowded or sub-optimal growth conditions. Correa (1997)<br />
considers that long term strategies for disease control in macroalgal farms will only succeed if<br />
the genetics <strong>of</strong> disease resistance in the host <strong>and</strong> virulence in the pathogen are understood.<br />
The risks that diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> pose to the New Zeal<strong>and</strong> marine<br />
environment have yet to be quantified. To fully underst<strong>and</strong> <strong>Undaria</strong>’s impacts <strong>and</strong> to<br />
effectively implement control or management options, diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong><br />
associated with this species as well as with other macroalgae need to be documented, both<br />
internationally <strong>and</strong> nationally.<br />
In this study the status <strong>of</strong> known diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> was<br />
determined (Objective 1) <strong>and</strong> literature was reviewed for reports <strong>of</strong> these diseases, <strong>pathogens</strong><br />
<strong>and</strong> <strong>parasites</strong> in <strong>Undaria</strong> populations in New Zeal<strong>and</strong> (Objective 3). <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong><br />
<strong>parasites</strong> <strong>of</strong> other macroalgae known internationally were summarised (Objective 2), as was<br />
information relating to those present in macroalgal populations in New Zeal<strong>and</strong> (Objective 4).<br />
Within a wider consideration <strong>of</strong> diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> macroalgae there are<br />
difficulties in confirming a causal role <strong>of</strong> specific organisms that have been implicated in<br />
disease or infection. The confirmation <strong>of</strong> Koch’s postulates is the exception rather than the rule.<br />
Thus, the database developed in this proposal has considered all organisms that have been<br />
associated with infection/disease/pathology with a clear indication <strong>of</strong> the evidence linking<br />
specific organisms to disease states/symptomology.<br />
4 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
2. Methods <strong>and</strong> Materials<br />
2.1. DEFINITIONS<br />
A disease is defined by various authors as:<br />
• either “… a continuing disturbance to the plant’s normal structure or function such that it<br />
is altered in growth rate, appearance or economic importance” (Andrews 1976),<br />
• “... the abnormal, injurious <strong>and</strong> continuous interference with physiological activities <strong>of</strong> the<br />
host" (Andrews 1979a, page 429; 1979b, page 448),<br />
• or a "... disturbance <strong>of</strong> the normal appearance <strong>and</strong> function <strong>of</strong> a plant” (Correa 1994)<br />
<strong>Diseases</strong> can have a variety <strong>of</strong> causes. This study only deals with infectious diseases, i.e.<br />
diseases caused by another organism (i.e. viruses, bacteria, protozoa, animals, fungi, other<br />
algae). This excludes diseases due to adverse abiotic conditions, i.e. physiological diseases<br />
caused by factors such as UV light, high or low temperature, or by dehydration (Gäumann<br />
1951; Andrews 1976). Exceptions are diseases caused by organisms affecting hosts weakened<br />
by adverse abiotic conditions.<br />
A pathogen is an organism that causes a disease.<br />
In the literature, an agent is <strong>of</strong>ten called a pathogen when it is found to be associated with a<br />
disease or aberrant appearance. However, for true pathogenicity, causality has to be<br />
demonstrated according to Koch’s postulates (Andrews & G<strong>of</strong>f 1984):<br />
1. the agent must be associated in every case with the disease under natural conditions, <strong>and</strong><br />
the disease must not appear in the absence <strong>of</strong> the agent<br />
2. the agent must be isolated in pure culture <strong>and</strong> characterised.<br />
3. typical symptoms must develop when the host is inoculated with the agent under suitable<br />
conditions, <strong>and</strong> the appropriate control inoculations must be made concurrently.<br />
4. the causal agent must be re-isolated <strong>and</strong> demonstrated to be identical to the agent isolated<br />
originally.<br />
An endophyte is an "organism living within a host plant" (Greek: éndon = inside;<br />
phytón = plant; Womersley 1987)<br />
An epiphyte is an organism living on the surface <strong>of</strong> a plant (its basiphyte). Epiphyte species<br />
can be opportunists (i.e. grow on all available surfaces), generalist epiphytes (i.e. grow on a<br />
variety <strong>of</strong> algal substrates), or specialists (i.e. grow on one or a few algal surfaces). Obligate<br />
specialist epiphytes are restricted to the epiphyte habit <strong>and</strong> to particular hosts. In the database<br />
we primarily focused on obligate or specialist epiphytes. In some cases <strong>of</strong> obligate<br />
epiphytism, such as in Polysiphonia lanosa growing on Ascophyllum nodosum, a directional<br />
exchange <strong>of</strong> nutrients from basi- to epiphyte has been experimentally demonstrated (Citharel<br />
1972), however, such information is lacking in the majority <strong>of</strong> cases. Organisms that were<br />
isolated from the surface <strong>of</strong> macroalgae but can also grow on other substrates are not<br />
considered in this study.<br />
A parasite is an organism which benefits to the detriment <strong>of</strong> its host organism. Usually, this<br />
means a physiological dependence, e.g. parasitic algae are unpigmented <strong>and</strong> thus, as<br />
heterotrophic organisms, rely at least to some extent on their host for nutrition, especially<br />
carbohydrates (G<strong>of</strong>f 1983; Correa 1994, 1997).<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 5
Especially in older literature the terms pathogen <strong>and</strong> parasite, or parasite <strong>and</strong> endophyte, tend<br />
to be used interchangeably. This should be avoided. For example, endophytic algae may but<br />
need not be parasitic, while not all <strong>parasites</strong> live inside the tissue <strong>of</strong> their host. Likewise, the<br />
term symbiosis is <strong>of</strong>ten used as opposite to parasitism. However, in its original definition, i.e.<br />
sensu de Bary (1879), a symbiosis is "...a phenomenon in which dissimilar organisms live<br />
together..." (de Bary 1879, cited in Paracer & Ahmadjian 2000; G<strong>of</strong>f 1983; Correa 1994), thus<br />
including parasitism.<br />
Classification system: We have based the hierarchical classification used in this study on the<br />
work <strong>of</strong> Cavalier-Smith (1998) with modifications adopted for the New Zeal<strong>and</strong> Species 2000<br />
project (pers. comm. D. Gordon, NIWA). The hierarchy is provided as Appendix 1. In this<br />
study the term “fungus” comprises true fungi, such as Ascomycetes, but also taxa that are<br />
traditionally treated as fungi, but really belong to the Ochrophyta/Chromista, i.e. oomycetes,<br />
Labyrinthula sp., etc.<br />
The organisms are treated as follows:<br />
Section in report Kingdoms/phyla included<br />
Viruses Viruses, Virus-like particles (VLPs)<br />
Bacteria Bacteria including phyla Eubacteria, Cyanobacteria, Proteobacteria, & Mycoplasmalike<br />
Organisms<br />
Fungi Fungi, Chromista (phyla Bigyra, Sagenista)<br />
Animals Animalia, Protozoa<br />
Other algae Plantae, Chromista (phylum Ochrophyta)<br />
2.2. DATA SOURCES<br />
2.2.1. Literature Review:<br />
A detailed literature search was carried out by NIWA information management staff to locate<br />
literature on diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>and</strong> other macroalgae.<br />
The literature searching strategy <strong>and</strong> terms were 1+3 <strong>and</strong> 2+3, where 1= seaweeds,<br />
macroalgae, <strong>Undaria</strong>, Laminaria, Macrocystis, Laminariales, Phaeophyceae, Phaeophyta; 2=<br />
Rhodophyta, Chlorophyta, Phaeophyceae, Phaeophyta; 3= disease, pathogen, parasite,<br />
endophyte, symbiosis, ascomycetes, bacteria, fungi, cyanobacteria, bluegreen/blue-green/blue<br />
green alga*e, chytrid*iomycetes, labrinthulids, virus*es, nematodes, copepod*s. The<br />
databases searched included st<strong>and</strong>ard marine bibliographic sources (e.g. SCOPUS, Web <strong>of</strong><br />
Science, ASFA, Google Scholar) <strong>and</strong> also web sites <strong>of</strong> marine research organisations were<br />
explored. The references obtained were entered into an EndNote database.<br />
Titles <strong>and</strong> abstracts <strong>of</strong> literature were scrutinised to determine relevance to the review <strong>and</strong><br />
papers were scored (immediate acquisition, later acquisition, possible inclusion, no<br />
relevance). The scoring <strong>of</strong> literature was carried out by 2 people <strong>and</strong> cross–checked by a third<br />
to check for consistent treatment. Relevant literature was obtained, <strong>and</strong> there was an iterative<br />
review <strong>of</strong> key words. Additional papers to be scored <strong>and</strong> entered into the database were<br />
located through scrutiny <strong>of</strong> reference lists <strong>and</strong> earlier review articles.<br />
Translations were made <strong>of</strong> key papers in Chinese, Spanish, French <strong>and</strong> German. Generally<br />
papers in Japanese (<strong>and</strong> some in Chinese) included English abstracts/ summaries as well as<br />
captions in English for tables, <strong>and</strong> graphs.<br />
6 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
2.2.2. E-mail <strong>and</strong> personal contacts:<br />
A message about the project requesting literature <strong>and</strong> general information was sent to Algae-<br />
L, a bulletin-board-type forum for people interested in any aspect <strong>of</strong> algae (terrestrial,<br />
freshwater <strong>and</strong> marine). In addition the archives <strong>of</strong> Algae-L were searched (May 1995 -<br />
October 2007) for any messages containing the words “disease” (37 hits) “pathogen” (19<br />
hits), “parasite” (14 hits). The majority <strong>of</strong> these were found to be references to books,<br />
microalgae, or to be otherwise irrelevant.<br />
Personal contacts <strong>and</strong>/or email messages were sent to key researchers in this field including<br />
Pr<strong>of</strong>essor Juan Correa (Universidad Catolica de Chile, Santiago, Chile), Mr Smith (Australian<br />
Centre for International Agricultural Research, Australia), Dr M. Polne-Fuller (University <strong>of</strong><br />
California, Santa Barbara, USA), Dr Bruce Harger (Sunshine Marine Farms, USA), Pr<strong>of</strong>essor<br />
Ma & Dr Bin Sun (Shanghai Fisheries University, China), Pr<strong>of</strong>essor Sung Min Boo<br />
(Chungnam National University, Korea), Dr M. Gauna (Universidad Nacional del Sur, Bahia<br />
Blanca, Argentina), Dr Danilo Largo (University <strong>of</strong> San Carlos, Philippines).<br />
2.2.3. Herbaria:<br />
The collections <strong>of</strong> the herbaria holding the majority <strong>of</strong> macroalgal specimens in New Zeal<strong>and</strong><br />
were examined (Auckl<strong>and</strong> Museum – including the Lindauer <strong>and</strong> ex-Auckl<strong>and</strong> University<br />
collections [AK/AKU], Museum <strong>of</strong> New Zeal<strong>and</strong> Te Papa Tongarewa [WELT], L<strong>and</strong>care<br />
Manaaki Whenua [CHR]). In addition the NZFungi database <strong>of</strong> L<strong>and</strong>care was searched for<br />
specimens <strong>of</strong> algal parasitic taxa known to be reported from New Zeal<strong>and</strong>. The data are<br />
presented in Appendix 2.<br />
2.3. DATABASE<br />
An Access database was established to enter data from the relevant papers <strong>and</strong> to record<br />
details <strong>of</strong> the diseases, <strong>parasites</strong> <strong>and</strong> <strong>pathogens</strong>. The following fields were included:<br />
• bibliographic data (including author, year, title, book/journal/publication details, abstract,<br />
keywords, comments, language);<br />
• characteristics <strong>of</strong> the agent (including classification [Kingdom, Phylum, Class, Order,<br />
Family, Genus, original genus <strong>and</strong> species name, current genus <strong>and</strong> species name, species<br />
authority], common name, agent type, associated species/community, secondary agent);<br />
• characteristics <strong>of</strong> the host (including classification [Kingdom, Phylum, Class, Order,<br />
Family, Genus, original genus <strong>and</strong> species name, current genus <strong>and</strong> species name, species<br />
authority], common name, taxonomic hierarchy, with fields for notes <strong>and</strong> for comments<br />
on the generation <strong>of</strong> the host affected);<br />
• location data (including world region [based on FAO fisheries regions - Attachment 3],<br />
latitude, longitude, map references, country, location, depth, exposure, temperature,<br />
salinity, water clarity, habitat type, agent stability, timing <strong>of</strong> occurrence, as well as fields<br />
to record data on epidemiology, seasonality, culture information, disease control, host<br />
impact).<br />
In many cases data were not available, particularly with respect to location data <strong>and</strong><br />
epidemiological information, as very little detail was provided in the original literature.<br />
The majority <strong>of</strong> the required fields for the database were determined at an initial stage <strong>and</strong><br />
there was an on-going review <strong>of</strong> the database effectiveness with additional fields identified<br />
<strong>and</strong> included after the initial phase <strong>of</strong> the study.<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 7
2.4. MAPPING<br />
The tender document specified a requirement to “map the known distribution <strong>of</strong> the diseases,<br />
<strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> which have been recorded”. Fields for longitude <strong>and</strong> latitude data<br />
were included in the database to enable this information to be extracted quickly from the<br />
database.<br />
3. Results<br />
3.1. GENERAL COMMENTS<br />
3.1.1. Data Sources<br />
Literature Review:<br />
The database includes a total <strong>of</strong> 927 references <strong>of</strong> which 549 pertinent papers are included in<br />
the reference list in this report. The Reference list also contains other literature cited in this<br />
report. The breakdown <strong>of</strong> papers by category is as follows: direct relevance (Laminariales),<br />
91; direct relevance (other algae), 363; generic/review, 70; source <strong>of</strong> additional references, 25;<br />
irrelevant, 292; unsourced, 86. The 292 entries considered irrelevant include, for example,<br />
papers dealing with epiphytes, saprobic organisms, freshwater, terrestrial <strong>and</strong>/or microalgae<br />
etc. Only relevant references are cited in the text <strong>of</strong> this report: the reference list provided lists<br />
all publications scored as ‘relevant’, ‘generic/reviews’ <strong>and</strong> ‘references only’, as well as papers<br />
cited in the text but not included in the database. The database contains a complete listing <strong>of</strong><br />
all papers considered in relation to algal diseases. Some additional papers that were identified<br />
through electronic search engines were discarded based on abstract, keywords or titles.<br />
Electronic search engines cover mainstream journals <strong>and</strong> publications, generally from the<br />
1970s onwards. A number <strong>of</strong> the papers relevant to this project fell outside these parameters<br />
i.e. published in the early 20 th century <strong>and</strong>/or in specialist or limited edition<br />
publications/journals. Although we sought “grey literature” <strong>and</strong> anticipated there would be<br />
guides <strong>and</strong> manuals available from marine farming centres or aquaculture institutions, almost<br />
none <strong>of</strong> this type <strong>of</strong> literature was forthcoming. Obtaining material from some overseas<br />
sources took much longer than anticipated <strong>and</strong> was sometimes extremely costly. The database<br />
includes bibliographic information for 87 references which are categorised as “unsourced”.<br />
These include post-graduate theses from outside New Zeal<strong>and</strong>, informal publication <strong>of</strong><br />
abstracts from congresses <strong>and</strong> conferences, <strong>and</strong> grey literature which we have been unable to<br />
source, particularly from Asian research institutes (Japan, China, Korea).<br />
E-mail <strong>and</strong> personal contacts:<br />
There was only minor interest generated by our posting in the ALGAE-L list, <strong>and</strong> <strong>of</strong> the 14<br />
responses to our email, most did not provide information, but instead were interested in the<br />
outcome <strong>of</strong> this study <strong>and</strong> its public availability. Three <strong>of</strong> the responses provided references,<br />
<strong>and</strong> one directed us to another potential contact. Additional personal <strong>and</strong> targeted contacts<br />
yielded only a small amount <strong>of</strong> additional information, although there was interest in the<br />
results <strong>of</strong> this study. Initially we had intended to include data from email searches <strong>and</strong> through<br />
personal contacts (via phone or email) but as these were very few in number <strong>and</strong> did not<br />
contain new information they were not included.<br />
3.1.2. Mapping<br />
Mapping distribution information obtained through the data sources was determined to be <strong>of</strong><br />
limited value. Only 72 <strong>of</strong> the 927 references (7.7%), or 192 <strong>of</strong> the more than 2300 agent<br />
entries in the database (~8%) included GIS compatible data (i.e. longitude <strong>and</strong> latitude) that<br />
8 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
could be mapped for sites where diseases were observed in seaweeds. More than 500 agents<br />
<strong>and</strong> 600 hosts were referred to in the relevant references reviewed, the majority <strong>of</strong> which were<br />
cited on a single occasion. It was concluded that mapping would not assist with visualization<br />
<strong>of</strong> these data. The data are summarised in Table 1 <strong>and</strong> 2 below.<br />
Two maps are provided illustrating the data obtained for the distribution <strong>of</strong> diseases/<br />
<strong>pathogens</strong>/ <strong>parasites</strong> reported for <strong>Undaria</strong> <strong>pinnatifida</strong>, using the FAO regions map (Appendix<br />
3) <strong>and</strong> a separate map showing the <strong>pathogens</strong> present in the native range <strong>of</strong> U. <strong>pinnatifida</strong><br />
(Appendix 4).<br />
Table 1. Summary <strong>of</strong> the number <strong>of</strong> records for each pest group in each algal host group. The<br />
numbers for the Ochrophyta exclude the Laminariales. NB. The numbers in the table do not<br />
relate directly to the number <strong>of</strong> references in the database, as references may contain multiple<br />
records.<br />
Viruses Bacteria Fungi Animals Other algae Total<br />
Rhodophyta 4 51 183 12 632 882<br />
Ochrophyta 95 3 120 14 84 316<br />
Chlorophyta 0 0 53 3 6 62<br />
Total 99 54 356 29 722 1260<br />
Table 2. Summary <strong>of</strong> the number records from each FAO region. Numbers for the Ochrophyta<br />
exclude the Laminariales. NB. Numbers in the table do not relate directly to the number <strong>of</strong><br />
references in the database, as references may contain multiple records. Total numbers differ<br />
from those in Table 1 as not all references contained location information.<br />
Rhodophyta Ochrophyta Chlorophyta Total<br />
21 - Atlantic, Northwest 69 42 12 123<br />
27 - Atlantic, Northeast 114 99 10 223<br />
31 - Atlantic, Western Central 11 10 1 22<br />
34 - Atlantic, Eastern Central 15 6 2 23<br />
37 - Mediterranean <strong>and</strong> Black Sea 30 8 2 40<br />
41 - Atlantic, Southwest 25 8 0 33<br />
47 - Atlantic, Southeast 50 1 0 51<br />
48 - Atlantic, Antarctic 10 7 0 17<br />
51 - Indian Ocean, Western 10 4 11 25<br />
57 - Indian Ocean, Eastern 38 12 9 59<br />
58 - Indian Ocean, Antarctic 0 1 0 1<br />
61 - Pacific, Northwest 105 14 1 120<br />
67 - Pacific, Northeast 117 7 3 127<br />
71 - Pacific, Western Central 42 3 0 45<br />
77 - Pacific, Eastern Central 114 12 3 129<br />
81 - Pacific, Southwest 19 30 2 51<br />
87 - Pacific, Southeast 36 19 0 55<br />
Total 805 283 56 1144<br />
3.2. DESCRIPTION OF KNOWN PATHOGEN-HOST RELATIONSHIPS<br />
3.2.1. <strong>Undaria</strong><br />
A summary <strong>of</strong> the records for each pest group in each algal host group, <strong>and</strong> number records<br />
from each FAO region are presented in Tables 3 & 4 respectively for members <strong>of</strong> the<br />
Laminariales, including <strong>Undaria</strong>. (Note – all papers refer to the sporophyte phase <strong>and</strong> not to<br />
the gametophyte phase <strong>of</strong> <strong>Undaria</strong>).<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 9
10 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 11
3.2.2. Known <strong>pathogens</strong> in native range<br />
Viruses<br />
There are no virus diseases known from <strong>Undaria</strong> <strong>pinnatifida</strong> or other <strong>Undaria</strong> species.<br />
Bacteria<br />
Gram-negative bacteria such as Aeromonas, Flavobacterium, Moraxella, Pseudomonas, <strong>and</strong><br />
Vibrio are associated with the "spot-rotting" disease ("Anaaki sho"; Kimura et al. 1976) <strong>and</strong><br />
the so-called "shot hole disease" (Tsukidate 1991) in Japanese <strong>Undaria</strong>. Severe outbreaks <strong>of</strong><br />
infections with Vibrio especially affect young sporophytes ("sporelings") <strong>of</strong> U. <strong>pinnatifida</strong><br />
(Anon. 1991). The "shot hole disease" is characterised by brown spots appearing on the<br />
thallus blade near the midrib which subsequently fuse together <strong>and</strong> spread onto the pinnate<br />
part <strong>of</strong> the blade (Tsukidate 1991).<br />
The "green spot disease/rot" caused by unspecified bacteria in Japan (Ishikawa & Saga 1989;<br />
Vairappan et al. 2001) <strong>and</strong> South Korea (Kang 1982) manifests with similar symptoms, first<br />
as green spots <strong>of</strong> rotting host tissue that result in small holes with green margins, <strong>and</strong> in the<br />
distal parts <strong>of</strong> the frond these enlarge <strong>and</strong> finally coalesce, accelerating the decay <strong>of</strong> the frond<br />
(Kang 1982). Japanese <strong>Undaria</strong> is furthermore infected by an unspecified bacterium causing<br />
the "yellow hole disease" (Ishikawa & Saga 1989; Vairappan et al. 2001) <strong>and</strong> “spot-rotting”<br />
disease (Kito et al. 1976).<br />
Bacteria enter the thallus <strong>of</strong> U. <strong>pinnatifida</strong> through openings like dead mucilage channels, <strong>and</strong><br />
digest cells <strong>and</strong> cell walls in the medulla. Cells <strong>of</strong> the cortex <strong>and</strong> meristoderm show ultrastructural<br />
damage (e.g. vacuolation <strong>of</strong> the dictyosome). When the host cells die, the disease<br />
symptoms become macroscopically visible (Kito et al. 1976).<br />
In China, the bacterium Halomonas venusta has been identified as a causative agent in “spot<br />
decay” (Ma et al. 1997a, b, 1998), <strong>and</strong> Vibrio logei in “green decay diseases” (Jiang et al.<br />
1997) <strong>of</strong> U. <strong>pinnatifida</strong>.<br />
Animals<br />
Some small crustacean species are associated with diseases in <strong>Undaria</strong>: The “pin hole<br />
disease” is caused by frond-mining nauplii <strong>of</strong> harpacticoid copepoda in <strong>Undaria</strong> from Japan<br />
(Anon. 1991) <strong>and</strong> South Korea (Tsukidate 1991), e.g. by species such as Amenophia<br />
orientalis, Parathalestris infestus, Scutellidium sp. (Ho & Hong 1988; Park et al. 1990; Anon.<br />
1991; Rho et al. 1993; Shimono et al. 2004) <strong>and</strong> Thalestris sp. (Kang 1982).<br />
Ceinina japonica, a gammeride amphipod from South Korea, invades the midrib <strong>of</strong> U.<br />
<strong>pinnatifida</strong> through the holdfast <strong>and</strong> bores a tunnel which may cause the longitudinal<br />
separation <strong>of</strong> the entire frond through the midrib. In heavily damaged thalli the holdfast may<br />
depart from the substrate (Kang 1982).<br />
Fungi<br />
A fungal infection occurs in <strong>Undaria</strong> from Japan, the so-called “chytrid blight” (Tsukidate<br />
1991). The name implies that this disease is caused by a true fungus <strong>of</strong> the class<br />
Chytridiomycetes, however, the culprit is an oomycete <strong>of</strong> the genus Olpidiopsis (Akiyama<br />
1977a). The fungus affects sporophytes, where it grows inside host cells, killing them slowly.<br />
Infected thalli gradually loose colour <strong>and</strong> disintegrate, juvenile thalli suffer severe damage or<br />
eventually die.<br />
12 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Other algae<br />
Laminariocolax aecidioides is an endophytic brown alga infecting farmed U. <strong>pinnatifida</strong> in<br />
Japan (Akiyama 1977b; Yoshida & Akiyama 1978; Veiga et al. 1997). Infections result in<br />
host thalli becoming thicker <strong>and</strong> stiffer, lowering their market value (Yoshida & Akiyama<br />
1978).<br />
3.2.3. Known <strong>pathogens</strong> in the introduced range other than New Zeal<strong>and</strong> (Australia, UK,<br />
France, Spain, USA (west coast), Argentina, Mexico, Taiwan)<br />
The endophyte Laminariocolax aecidioides (as Gononema aecidioides) has been found in<br />
farmed <strong>Undaria</strong> <strong>pinnatifida</strong> thalli from Spain (Veiga et al. 1997) <strong>and</strong> has also been found in<br />
<strong>Undaria</strong> in Argentina (Gauna et al. pers. comm.).<br />
3.2.4. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
Even though members <strong>of</strong> the genus Laminariocolax occur in New Zeal<strong>and</strong> kelps, none have<br />
so far been observed in <strong>Undaria</strong> <strong>pinnatifida</strong>. Instead, in New Zeal<strong>and</strong> U. <strong>pinnatifida</strong> hosts<br />
another endophyte, Microspongium tenuissimum, which is also found in Ecklonia radiata <strong>and</strong><br />
various red algae. The infection <strong>of</strong> U. <strong>pinnatifida</strong> with M. tenuissimum was not associated<br />
with obvious macroscopic disease symptoms (Heesch 2005).<br />
3.3. LAMINARIALES<br />
3.3.1. Known <strong>pathogens</strong> worldwide<br />
Viruses<br />
There are no viral diseases reported from members <strong>of</strong> the Laminariales outside New Zeal<strong>and</strong><br />
(see 3.2.2).<br />
Bacteria<br />
Most bacteria affecting kelps belong to the phylum Proteobacteria. Pathogenic species <strong>of</strong><br />
Alteromonas, Pseudoalteromonas, Pseudomonas <strong>and</strong> Vibrio have been recorded from<br />
Saccharina japonica in China (e.g. Tang et al. 2001; Liu et al. 2002; Wang et al. 2006) <strong>and</strong><br />
Japan (e.g. Ezura et al. 1990; Yamada et al. 1990; Sawabe et al. 1998; Sawabe et al. 2000a, b;<br />
Narita et al. 2001; Vairappan et al. 2001) resulting in holes <strong>and</strong> lesions on thalli <strong>and</strong><br />
eventually “rot disease”. Some proteobacteria indirectly affect gametophytes <strong>and</strong> young<br />
sporophytes in culture when red spot disease <strong>of</strong> the culture bed (i.e. the culture ropes) causes<br />
the young Saccharina japonica to detach from infected ropes (e.g. Ezura et al. 1988; Yumoto<br />
et al. 1989a, b). Alteromonas sp. <strong>and</strong> Vibrio sp. are also associated with lesions <strong>and</strong> thallus<br />
bleaching <strong>of</strong> Saccharina ochotensis <strong>and</strong> S. religiosa in Japan (Vairappan et al. 2001). A<br />
species <strong>of</strong> Acinetobacter causes “white rot” in Nereocystis luetkeana resulting in rot <strong>of</strong> stipes<br />
<strong>and</strong> pneumatocysts, which collapse <strong>and</strong> become covered in white slime within 7-10 days<br />
(Andrews 1977).<br />
In China, both the gametophytes <strong>and</strong> sporophytes <strong>of</strong> Saccharina japonica are prone to<br />
“malformation disease” caused by the firmicute Macrococcus sp. (Anon. 1989).<br />
Unspecified bacteria have been reported as <strong>pathogens</strong> in Macrocystis pyrifera, Pelagophycus<br />
porra <strong>and</strong> Egregia laevigata in America (Br<strong>and</strong>t 1923), Saccharina japonica in China (Wu et<br />
al. 1983; Ding 1992; Yang et al. 2001; Huang et al. 2002a, b). The “black rot” <strong>of</strong> Macrocystis<br />
pyrifera in California is assumed to be caused by a unidentified parasitic microorganism<br />
invading already damaged host thalli (Rheinheimer 1992).<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 13
A mycoplasma-like organism (MLO) causes the "twisted frond disease" or “coiling-stunt<br />
disease” in Saccharina japonica from China (e.g. Wang et al. 1983; Wu et al. 1983;<br />
Tsukidate 1991).<br />
Animals<br />
A number <strong>of</strong> amphipods are known to bore in kelp stipes <strong>and</strong> hollow them, causing<br />
considerable damage which may eventually lead to the death <strong>of</strong> the host. In Alaska <strong>and</strong><br />
California, Peramphithoe stypotrupetes infests stipes <strong>of</strong> Laminaria setchellii damaged by<br />
gastropod grazing (Chess 1993). Also in California <strong>and</strong> Alaska, it occurs in Saccharina<br />
dentigera, <strong>and</strong> in southern California it is found in Eisenia arborea <strong>and</strong> Pterygophora<br />
californica (Conlan & Chess 1992), while Californian Macrocystis pyrifera populations are<br />
infested by the related amphipod P. humeralis (Chess 1993). In Irel<strong>and</strong>, Alaria esculenta is<br />
inhabited by Amphitholina cuniculus (Myers 1974; Chess 1993). In Japan, Saccharina<br />
japonica is similarly affected by Ceinina japonica (Akaike et al. 2002).<br />
Fungi<br />
The ascomycete Phycomelaina laminariae causes the “stipe blotch disease” in laminarian<br />
species from the north-western <strong>and</strong> north-eastern Atlantic. Its hyphae penetrate the surface <strong>of</strong><br />
Alaria esculenta, Saccharina latissima, S. longicruris <strong>and</strong> Laminaria digitata, leading to<br />
necrotic tissue <strong>and</strong> reduced overall performance <strong>of</strong> the host thalli (Sutherl<strong>and</strong> 1915b, c;<br />
Kohlmeyer 1968; Kohlmeyer 1979; Schatz et al. 1979; Schatz 1980, 1983, 1984a, c; G<strong>of</strong>f &<br />
Glasgow 1980; Porter & Farnham 1986a).<br />
Several other ascomycete fungi attack members <strong>of</strong> the Laminariales: Pontogeneia erikae is a<br />
parasite in Egregia menziesii from California (Kohlmeyer & Demoulin 1981), Sigmoidea<br />
marina causes lesions in the surface <strong>of</strong> Saccharina latissima from Britain (Haythorn et al.<br />
1980), Ophiobolus laminariae causes blackened patches on the stipes <strong>of</strong> Laminaria digitata in<br />
Scotl<strong>and</strong> (Sutherl<strong>and</strong> 1915c), <strong>and</strong> in California Asteromyces cruciatus has been reported from<br />
Egregia menziesii, however their relationship is uncertain (Nolan 1972).<br />
Oomycetes have also been reported from members <strong>of</strong> the Laminariales in the north-western<br />
<strong>and</strong> north-eastern Atlantic: Petersenia sp. causes damage to the stipes <strong>of</strong> Laminaria digitata,<br />
Laminaria sp. <strong>and</strong> Saccharina longicrucis (Kohlmeyer 1968) <strong>and</strong> Pleotrachelus minutus<br />
infects the apical hairs <strong>of</strong> Chorda filum in Sweden (Aleem 1952a).<br />
In France Labyrinthomyxa sauvageaui infects Laminaria ochroleuca (Duboscq 1921). An<br />
unknown hyphomycete causes contortion <strong>of</strong> the blade <strong>and</strong> blackening <strong>of</strong> the stipe in<br />
Laminaria digitata in Maine, USA (Kohlmeyer 1968) <strong>and</strong> in Russia an undetermined fungus<br />
has been isolated from farmed populations <strong>of</strong> Saccharina japonica (Zvereva 1998).<br />
Other algae<br />
Green algae are occasionally observed growing in kelps, however very little information is<br />
available on their impact on the host species. Acrochaete repens, for example, grows in<br />
Chorda filum from the North American east coast (O’Kelly et al. 2004), Canada (South 1968)<br />
<strong>and</strong> from Denmark, Irel<strong>and</strong> <strong>and</strong> the Isle <strong>of</strong> Man in the north eastern Atlantic (South 1968;<br />
Nielsen 1979). The related species A. geniculata infects kelps along the North American<br />
Pacific coast, such as Egregia menziesii, Cymathere triplicata, Laminaria sinclairii,<br />
Saccharina dentigera <strong>and</strong> Dictyoneurum californicum (O’Kelly 1983). Egregia menziesii<br />
from British Columbia also hosts another Acrochaete species, A. apiculata (C. O’Kelly, pers.<br />
com.).<br />
The green endophyte Bolbocoleon piliferum is found on the east <strong>and</strong> west coast <strong>of</strong> the USA,<br />
<strong>and</strong> eastern Canada, growing in the kelps Alaria marginata, Chorda filum, Cymathere<br />
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triplicata <strong>and</strong> Pleurophycus gardneri (South 1968; O’Kelly et al. 2004). It is also recorded in<br />
Chorda filum from Denmark, Wales, Irel<strong>and</strong> <strong>and</strong> the Isle <strong>of</strong> Man (South 1968; Nielsen 1979)<br />
<strong>and</strong> in Laminaria hyperborea from Denmark (Nielsen 1979). Another green endophyte<br />
Entocladia viridis is also known from several countries in the north-eastern <strong>and</strong> north-western<br />
Atlantic, growing in Laminaria digitata <strong>and</strong> Saccharina latissima (Nielsen 1979). In Chile,<br />
another green endophyte, reported as Sporocladopsis novae-zel<strong>and</strong>iae grows in Lessonia<br />
nigrescens (Correa & Martinez 1996).<br />
Pigmented endophytic brown algae are very common in kelps (Lein et al. 1991; Ellertsdottir<br />
& Peters 1995). Their presence is <strong>of</strong>ten associated with brown spots (“dark-spot disease”,<br />
Lein et al. 1991), hyperplasia leading to warts or galls, <strong>and</strong>, in severe cases, thallus<br />
deformations (Andrews 1977; Apt 1988b). Traditionally, kelp endophytes have been<br />
classified as Streblonema species (G<strong>of</strong>f & Glasgow 1980), for example, the endophytes that<br />
affect Saccharina sessilis, Alaria tenuifolia, Laminaria setchellii <strong>and</strong> Nereocystis luetkeana<br />
along the North American west coast (Setchell & Gardner 1922). However, genetically, most<br />
kelp endophytes belong to the genera Laminariocolax <strong>and</strong> Microspongium.<br />
North Atlantic kelp populations are infected by two species <strong>of</strong> Laminariocolax: L.<br />
tomentosoides <strong>and</strong> L. aecidioides. The former is mainly found in Laminaria digitata, but<br />
occasionally also in L. hyperborea, Saccharina latissima <strong>and</strong> Alaria sp. (Lund 1959; Pedersen<br />
1976; Ellertsdottir & Peters 1997; Burkhardt & Peters 1998; Küpper et al. 2002).<br />
Laminariocolax tomentosoides ssp. deformans is associated with galls <strong>and</strong> stipe coiling in<br />
Laminaria digitata from France (Dangeard 1931b; Peters 2003).<br />
Laminariocolax aecidioides is found throughout the Northern Hemisphere. In the North<br />
Atlantic, it has been observed in Laminaria hyperborea <strong>and</strong> Saccharina latissima from<br />
Germany, France <strong>and</strong> Denmark (e.g. Peters & Ellertsdottir 1996; Burkhardt & Peters 1998;<br />
Heesch & Peters 1999; Peters 2003), in S. groenl<strong>and</strong>ica, Laminaria sp. <strong>and</strong> S. longicruris<br />
from Greenl<strong>and</strong> (Pedersen 1981), <strong>and</strong> on the North American east coast in Laminaria digitata<br />
(Peters 2003). In the North Pacific, it infects not only U. <strong>pinnatifida</strong>, but also Costaria sp.<br />
from Japan, <strong>and</strong> is furthermore known from Californian Hedophyllum sp. populations<br />
(Yoshida & Akiyama 1978).<br />
Southern hemisphere kelp populations are infected by two other members <strong>of</strong> the genus<br />
Laminariocolax, L. macrocystis <strong>and</strong> L. eckloniae. The former endophyte grows in<br />
Macrocystis pyrifera from Chile, the latter in Ecklonia maxima from South Africa (Peters<br />
1991; Burkhardt & Peters 1998). Heesch (2005) considers L. macrocystis <strong>and</strong> L. eckloniae to<br />
be synonymous.<br />
Laminariocolax sp. is recorded from the North Atlantic in Laminaria hyperborea (Lein et al.<br />
1991; Peters & Schaffelke 1996; Ellertsdottir & Peters 1997) <strong>and</strong> the Pacific in Macrocystis<br />
integrifolia, Saccharina latissima <strong>and</strong> Nereocystis luetkeana (Andrews 1977; Apt 1988a).<br />
Another endophytic brown alga, Laminarionema elsbetiae, occurs in Japanese Saccharina<br />
japonica as well as in the German kelps S. latissima <strong>and</strong> Laminaria digitata (Kawai &<br />
Tokuyama 1995; Peters & Ellertsdottir & 1996; Ellertsdottir & Peters 1997; Peters &<br />
Burkhardt 1998; Heesch & Peters 1999; Peters 2003).<br />
The genus Microspongium is occasionally found as endophyte in kelps. On the east coast <strong>of</strong><br />
North America, Alaria esculenta <strong>and</strong> Saccharina longicruris are infected by Microspongium<br />
alariae, with symptoms ranging from dark spots to twisted stipes (Peters 2003).<br />
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Gametophytes <strong>of</strong> kelps themselves colonise other algae as endophytes. A genetic study has<br />
revealed that endophytic brown algae growing in Lessoniopsis littoralis from British<br />
Columbia, Canada, are gametophytes <strong>of</strong> other kelps growing near the host, i.e. <strong>of</strong> Alaria sp.,<br />
Macrocystis integrifolia <strong>and</strong> Nereocystis luetkeana (Lane & Saunders 2005).<br />
The ectocarpalean endophytes Phaeostroma parasiticum <strong>and</strong> Dermatocelis laminariae occur<br />
in Saccharina latissima <strong>and</strong> Laminaria sp. respectively in Greenl<strong>and</strong> (Pedersen 1976). In<br />
Germany, unspecified ectocarpalean endophytes are reported to infect up to 85% <strong>of</strong> their<br />
hosts, Laminaria saccharina, L. digitata <strong>and</strong> L. hyperborea (Ellertsdottir & Peters 1995).<br />
An obligate epiphyte, Porphyra moriensis, infests Chorda filum in Japan (Notoya &<br />
Miyashita 1999).<br />
3.3.2. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
In northern New Zeal<strong>and</strong>, mass diebacks <strong>of</strong> Ecklonia radiata where reported in the mid 1990s<br />
(e.g. Cole & Babcock 1996). Subsequent research indicates that the diebacks are caused by<br />
primary <strong>and</strong> secondary agents: E. radiata is affected by the amphipod Orchomenella aahu,<br />
which burrows into the stipes <strong>of</strong> the host <strong>and</strong> hollows them out, thus accelerating death <strong>of</strong> the<br />
fronds. The simultaneously occurring bleaching <strong>of</strong> the fronds is probably due to a secondary<br />
infection with a virus (Haggitt & Babcock 2003) <strong>and</strong> the diebacks have been associated with<br />
both virus-like particles (VLPs) <strong>and</strong> a poty virus (Easton 1995, Easton et al. 1997).<br />
Three species <strong>of</strong> pigmented endophytic brown algae infect kelps from New Zeal<strong>and</strong> (Heesch<br />
2005): Laminariocolax macrocystis (which in this treatment includes L. eckloniae) is<br />
associated with galls <strong>and</strong> thallus deformations in Macrocystis pyrifera (North <strong>and</strong> South<br />
Isl<strong>and</strong>s) <strong>and</strong> Ecklonia radiata from the North, South <strong>and</strong> Chatham Isl<strong>and</strong>s. Additionally, E.<br />
radiata hosts Microspongium tenuissimum (which includes M. radians), an endophyte mostly<br />
observed in red algae. The third endophyte, an undescribed ectocarpalean species so far only<br />
known from New Zeal<strong>and</strong>, was found in a gall on Lessonia tholiformis from the Chatham<br />
Isl<strong>and</strong>s (Heesch 2005).<br />
An unidentified green endophyte (probably a species belonging to the genus Acrochaete,<br />
O’Kelly pers. com.) was frequently observed in stipes <strong>of</strong> Macrocystis pyrifera along the<br />
Otago coast (Heesch, unpublished data).<br />
3.4. BROWN ALGAE OTHER THAN LAMINARIALES<br />
3.4.1. Known <strong>pathogens</strong> worldwide<br />
Viruses<br />
Virus-like particles (VLPs) in several members <strong>of</strong> the order Ectocarpales (Phaeophyceae), e.g.<br />
Ectocarpus <strong>and</strong> Pylaiella species (Markey 1974; Dodds 1979) have subsequently been<br />
identified as DNA viruses. Viruses have been found in Ectocarpus siliculosus (Ectocarpus<br />
siliculosus virus EsV), E. fasciatus (EfasV), Feldmannia irregularis (FirrV), F. simplex<br />
(FlexV), an unidentified Feldmannia species (FsV), Myriotricha clavaeformis (MclaV),<br />
Pylaiella littoralis (PlitV), Hincksia hincksiae (HincV), <strong>and</strong> also in Kuckuckia sp. <strong>and</strong><br />
Leptonematella fasciata. The viruses infect naked spores, leading to a latent infection in<br />
vegetative thalli. Upon maturation, reproductive organs develop abnormally producing new<br />
virus particles instead <strong>of</strong> spores (Clitheroe & Evans 1974; Müller et al. 1990, 1996 (a, b, c),<br />
1998, 2000; Müller 1991a, b; Henry & Meints 1992, 1994; Müller & Stache 1992; Lanka et<br />
al. 1993; Müller & Frenzer 1993; Friess-Klebl et al. 1994; Kuhlenkamp & Müller 1994;<br />
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Parodi & Müller 1994; Robledo et al. 1994; Bräutigam et al. 1995; Krueger et al. 1996;<br />
Müller & Schmid 1996; Sengco et al. 1996; Del Campo et al. 1997; Kapp et al. 1997; Maier<br />
et al. 1997, 1998, 2002; Kapp 1998; Lee et al. 1995, 1998; Maier & Müller 1998; Wolf et al.<br />
1998, 2000; Van Etten & Meints 1999; Delaroque et al. 2000a, b, 2003; Dixon et al. 2000;<br />
Van Etten et al. 2002; Chen et al. 2005; Dunigan et al. 2006). EsV <strong>and</strong> EfasV are known from<br />
host populations world-wide (Müller & Stache 1992).<br />
In Botrytella micromora, virus-like particles (VLPs) are associated with tissue necroses <strong>and</strong><br />
zoospores that fail to germinate <strong>and</strong> lyse instead (Oliveira & Bisalputra 1978; Henry &<br />
Meints 1994). Likewise, VLPs affect zoospore germination in Halosiphon tomentosus, while<br />
thalli <strong>of</strong> Streblonema sp. containing VLPs in their vegetative cells do not appear to be<br />
negatively affected (Toth & Wilce 1972; LaClaire & West 1977; Dodds 1979; Henry &<br />
Meints 1992, 1994; Müller et al. 1998).<br />
Bacteria<br />
Bacteria associated with galls <strong>and</strong> thallus deformations occur in Fucus vesiculosus, F. spiralis<br />
<strong>and</strong> Saccorhiza polyschides (Cantacuzene 1930; Apt 1988b; Rheinheimer 1992). In France a<br />
proteobacterium infects Cystoseira nodicaulis causing damage to the thallus (Pellegrini &<br />
Pellegrini 1982), <strong>and</strong> in Russia’s Kurile Isl<strong>and</strong>s, Pseudoalteromonas issachenkonii degrades<br />
the thallus <strong>of</strong> its host Fucus evanescens (Ivanova et al. 2002).<br />
Animals<br />
Protozoan <strong>pathogens</strong> are reported from members <strong>of</strong> the Ectocarpales <strong>and</strong> the Fucales. The<br />
infection <strong>of</strong> Ectocarpus siliculosus from Chile with the plasmodiophorid Maulinia ectocarpii<br />
results in the sterility <strong>of</strong> the host sporangia (Maier et al. 2000). Also in Chile, another<br />
plasmodiophorid infects Durvillaea antarctica causing galls <strong>and</strong> internal hypertrophy <strong>of</strong> cells<br />
(Aguilera et al. 1988). An unspecified brown alga is also reported to be infected by the<br />
plasmodiophorid Phagomyxa algarum (Porter & Farnham 1986a). Amoeba are found in<br />
Sargassum muticum <strong>and</strong> in British Fucus serratus, the latter affected by the species<br />
Trichosphaerium sieboldi. The amoeba digest the walls <strong>and</strong> invade the cytoplasm <strong>of</strong> the host<br />
cells leading to a dissociation <strong>of</strong> the host tissue (Polne-Fuller & Gibor 1987; Rogerson et al.<br />
1998).<br />
In Japan, the harpacticoid copepods Dactylopusioides fodiens <strong>and</strong> D. macrolabris feed on the<br />
internal tissue <strong>of</strong> Dictyota dichotoma <strong>and</strong> live in the resulting galleries; Dactylopusioides<br />
fodiens also parasitises Pachydictyon coriaceum (Shimono et al. 2003, 2004). Copepoda are<br />
furthermore associated with galls in Desmarestia aculeata from Scotl<strong>and</strong> (Barton 1892).<br />
Nematodes <strong>of</strong> the genus Halenchus are found in galls on members <strong>of</strong> the Fucales: H. fucicola<br />
affects Ascophyllum nodosum while H. dumnonicus inhabits Fucus vesiculosus <strong>and</strong> F.<br />
serratus (Barton 1892; Coles 1958; Tokida 1958; Apt 1988b).<br />
Fungi<br />
There is a large body <strong>of</strong> literature relating to fungi <strong>and</strong> seaweeds, however many contain little<br />
information about the fungal parasite’s impact on the algal host.<br />
Ascomycete fungi frequently form galls in members <strong>of</strong> the Fucales, e.g. Massarina<br />
cystophorae in Cystophora retr<strong>of</strong>lexa <strong>and</strong> C. subfarcinata. Members <strong>of</strong> the ascomycete genus<br />
Haloguignardia are widespread <strong>and</strong> occur in a range <strong>of</strong> hosts, e.g. Haloguignardia irritans in<br />
Cystoseira osmundea, Cystoseira sp., Halidrys dioica <strong>and</strong> Halidrys sp.; Haloguignardia sp. in<br />
Cystoseira balearica, Cystoseira sp., Halydris dioica <strong>and</strong> various Sargassum species (S.<br />
decipiens, S. fallax, S. fluitans, S. natans <strong>and</strong> S. sinclairii) (Estee 1913; Cribb & Herbert 1954;<br />
Tokida 1958; Kohlmeyer 1979; Apt 1988c; Alongi et al. 1999). Haloguignardia cystoseirae<br />
infects Cystoseria spp. in the Mediterranean (Kohlmeyer & Demoulin 1981; Alongi et al.<br />
1999), whereas Haloguignardia tumefaciens, H. oceanica, H. decidua <strong>and</strong> H. longispora<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 17
infect Sargassum spp. in Australia, Japan, America <strong>and</strong> the Sargasso Sea (e.g. Ferdin<strong>and</strong>sen<br />
& Winge 1920; Tokida 1958; Cribb & Cribb 1960; Kohlmeyer 1971, 1972; Alongi et al.<br />
1999). A secondary agent, the hyperparasite Sphaceloma cecidii has also been reported from<br />
Cystoseira sp., Sargassum sp. <strong>and</strong> Halidrys sp., where its infection is restricted to areas <strong>of</strong> the<br />
host already affected by Haloguignardia (Kohlmeyer 1979).<br />
Further members <strong>of</strong> the ascomycetes that affect marine algae include Thalassoascus<br />
treboubovii, recorded from Cutleria chilosa, C. multifida, Cystoseira sp. <strong>and</strong> Zanardinia typus<br />
(Ollivier 1929; Kohlmeyer 1979), Lindra thalassiae from Sargassum spp. (Meyers 1969;<br />
Kohlmeyer 1979; Raghukumar et al. 1992), Chadefaudia gymnogongri from Xiphophora<br />
chondrophylla (Kohlmeyer 1973a), Orcadia ascophylli <strong>and</strong> Trailia ascophylli from<br />
Ascophyllum nodosum (Sutherl<strong>and</strong> 1915c), <strong>and</strong> Asteromyces cruciatus from Cystoseira<br />
osmundacea (Nolan 1972). Ascomycetes reported from Fucus spp. include Cephalosporium<br />
sp., Sigmoidea marina, Didymella fucicola, Orcadia ascophylli <strong>and</strong> Trailia ascophylli<br />
(Sutherl<strong>and</strong> 1915c; Kohlmeyer 1968; Andrews 1977; Haythorn et al. 1980; Miller & Whitney<br />
1981; Schatz 1984a). Pelvetia canaliculata is also infected by a range <strong>of</strong> ascomycete fungi,<br />
including Didymella fucicola, Orcadia ascophylli, Dothidella pelvetiae, Pharcidia pelvetiae,<br />
Pleospora pelvetiae <strong>and</strong> Stigmatea pelvetiae (Sutherl<strong>and</strong> 1915a). Additionally,<br />
Scolecobasidium salinum degrades alginates <strong>of</strong> brown algae (Moen et al. 1995).<br />
In the Archemycota, Chytridium polysiphoniae, C. megastomum <strong>and</strong> Olpidium sphacellarum<br />
have been reported from Sphacelaria cirrosa, Sphacelaria sp., Striaria attenuata <strong>and</strong><br />
Pylaiella littoralis, disintegrating cell contents (Sparrow 1934, 1936; Raghukumar 1987b;<br />
Hyde et al. 1998; Küpper & Müller 1999; Müller et al. 1999).<br />
The term mycophycobiosis was created for obligate symbioses between algae <strong>and</strong> fungi<br />
which are without a detrimental effect for both symbionts, <strong>and</strong> in which, unlike in lichens, the<br />
alga is the partner that provides the structure. An example is Mycophycias ascophylli growing<br />
in Ascophyllum nodosum <strong>and</strong> in Pelvetia canaliculata in the Northwest Atlantic (e.g.<br />
Kohlmeyer & Kohlmeyer 1972; Miller & Whitney 1981; Porter & Farnham 1986a; Kingham<br />
& Evans 1986; Stanley 1992; Deckert & Garbary 2005a). The symbiosis between A.<br />
nodosum, M. ascophylli <strong>and</strong> Polysiphonia lanosa has been intensely studied (e.g. Garbary &<br />
London 1995; Garbary & MacDonald 1995; Deckert & Garbary 2005b; Garbary et al. 2005).<br />
In the pseud<strong>of</strong>ungi, oomycetes are also common <strong>pathogens</strong> <strong>of</strong> seaweeds. In particular they are<br />
found in members <strong>of</strong> the Ectocarpales: Pylaiella littoralis, Ectocarpus siliculosus, Striaria<br />
attenuata <strong>and</strong> Hincksia spp. are infected by species <strong>of</strong> Eurychasma, Anisolpidium,<br />
Pleotrachelus, Petersenia <strong>and</strong> Olpidiopsis (Sparrow 1934, 1936; Karling 1943; Aleem<br />
1950a,d, 1952a; Küpper & Müller 1999; Müller et al. 1999; West et al. 2006). Some members<br />
<strong>of</strong> the Sphacelariales are also affected by oomycetes (Aleem 1952a).<br />
In the Sagenista, the Labyrinthulomycetes include two groups that are frequently associated<br />
with seaweeds; the thraustochytrids <strong>and</strong> the labyrinthulids. From the former, Aplanochytrium<br />
spp. occur as endophytes in Sargassum spp. <strong>and</strong> Padina atillarum (Raghukumar et al. 1992;<br />
Sathe-Pathak et al. 1993; Ulken et al. 1985; Raghukumar 2002); in the latter Labyrinthula sp.<br />
is reported from Lobophora variegata (Raghukumar 1987b).<br />
Other algae<br />
A few members <strong>of</strong> the Rhodophyta occur as epi- or endophytes <strong>of</strong> brown algal hosts.<br />
Polysiphonia lanosa is an obligate epiphyte on Ascophyllum nodosum (Turner & Evans 1977;<br />
Garbary et al. 1991; Cardinal & Lesage 1992; Lining & Garbary 1992; Garbary & London<br />
1995), which has occasionally also been observed on Fucus vesiculosus (Pearson & Evans<br />
1990; Rindi & Guiry 2004). It is <strong>of</strong>ten found on damaged host fronds (Lobban & Baxter<br />
1983; Rindi & Guiry 2004), deeply penetrating the host with its rhizoids (Rawlence 1972;<br />
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Rawlence & Taylor 1972; Garbary et al. 2005). There is some evidence suggesting a transfer<br />
<strong>of</strong> substances occurs between the two symbionts (Citharel 1972; Penot 1974), while other<br />
studies doubt the translocation <strong>of</strong> synthetates from basiphyte to epiphyte (Turner & Evans<br />
1977; Harlin & Craigie 1975). Colacodictyon reticulatum is a small endophytic red alga<br />
growing in Desmarestia ligulata, <strong>and</strong> Haplodasya urceolata endophytises Cystophora<br />
retr<strong>of</strong>lexa (Kylin 1956).<br />
Brown algae occur as endophytes <strong>of</strong> other brown algae. The endophytic brown algae<br />
Herponema valiantei <strong>and</strong> Streblonemopsis irritans are associated with galls in Cystoseira<br />
tamariscifolia <strong>and</strong> C. zosteroides, respectively (Apt 1988a, 1988b). Pedersen (1976) reports<br />
Herponema desmarestiae <strong>and</strong> Streblonema fasciculatum from Desmarestia viridis <strong>and</strong><br />
Eudesme virescens respectively. The endophytic brown algae Microspongium alariae <strong>and</strong><br />
Myriactula cl<strong>and</strong>estina occur in Fucus vesiculosus from Finl<strong>and</strong> <strong>and</strong> Greenl<strong>and</strong> (Pedersen<br />
1976; Peters 2003). In California, Desmarestia ligulata is affected by Streblonema<br />
transfixum, <strong>and</strong> S. penetrale penetrates the stipe <strong>of</strong> Hesperophycus californicus (Setchell &<br />
Gardner 1922). Laminariocolax sp. occurs in Chordaria flagelliformis in Greenl<strong>and</strong> (Pedersen<br />
1976) <strong>and</strong> Laminariocolax aecidioides in Sphacelaria arctica from multiple sites in the North<br />
Atlantic (Yoshida & Akiyama 1978). Notheia anomala is a hemi-parasitic brown alga<br />
occurring in Australia <strong>and</strong> New Zeal<strong>and</strong> (Adams 1994). In Australia it infects Hormosira<br />
banksii <strong>and</strong> occasionally also Xiphophora chondrophylla (Gibson & Clayton 1987; Raven et<br />
al. 1995).<br />
Some small members <strong>of</strong> the Ectocarpales are on the border between epi- <strong>and</strong> endophytism.<br />
For example, Elachista fucicola is an obligate epiphyte <strong>of</strong> Fucus vesiculosus (Rindi & Guiry<br />
2004), but it also grows on Ascophyllum nodosum penetrating the host surface with its<br />
rhizoids <strong>and</strong> leading the host to form a tissue callus around the penetrating filaments (Deckert<br />
& Garbary 2005a, b). Filaments <strong>of</strong> Trachynema groenl<strong>and</strong>icum grow in the loosely organised<br />
cortex <strong>of</strong> Chordaria linearis in southern South America, but do not penetrate into the compact<br />
subcortex or medulla <strong>of</strong> the host (Peters 1992). Gononema pectinatum was isolated from a<br />
culture <strong>of</strong> Dictyosiphon hirsutum from Chile, however, the origin <strong>of</strong> the contaminant (epi- or<br />
endophytic) was not determined (Burkhardt & Peters 1998).<br />
Three endophytic brown algae have been reported from the Antarctic Peninsula:<br />
Laminariocolax eckloniae in Himanthothallus gr<strong>and</strong>ifolius, Geminocarpus austro-georgiae in<br />
Desmarestia menziesii, <strong>and</strong> Ascoseirophila violodora in Ascoseira mirabilis (Peters 2003). In<br />
addition, Antarctic Adenocystis utricularis specimens are epiphytised by Austr<strong>of</strong>ilum<br />
incommodum, a small phaeophyte that is anchored in its host with endophytic filaments<br />
(Müller et al. 1992; Peters 2003).<br />
Antarctic Ascoseira mirabilis furthermore hosts the unicellular endophytic green alga<br />
Chlorochytrium sp. (Peters 2003) which is, like Codiolum sp., considered to be the<br />
endophytic sporophyte <strong>of</strong> an Acrosiphonia species. Codiolum petrocelidis, for example,<br />
grows in the crustose brown alga Ralfsia pacifica from the Pacific coast <strong>of</strong> Canada (Sussmann<br />
& DeWreede 2002). Chlorochytrium dermatocolax has been recorded from the North Atlantic<br />
<strong>and</strong> the Pacific coast <strong>of</strong> North America in Sargassum muticum <strong>and</strong> Sphacelaria spp. (Lund<br />
1959; Pedersen 1976; Polne-Fuller 1987).<br />
A filamentous green alga, Acrochaete repens, grows endophytically in Fucus serratus from<br />
the German Bight, Denmark <strong>and</strong> the Channel Isl<strong>and</strong>s (Kremer 1975; Nielsen 1979) <strong>and</strong> F.<br />
vesiculosus from Denmark (Nielsen 1979). The closely related Entocladia species E. viridis<br />
<strong>and</strong> E. wittrockii grow endophytically in Desmarestia aculeata, Dictyota dichotoma <strong>and</strong><br />
Elasticha fucicola from locations in the Pacific, Mediterranean <strong>and</strong> North Atlantic (Nielsen<br />
1979; O’Kelly 1981).<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 19
Navicula endophytica is a diatom living in the intercellular mucilage <strong>of</strong> receptacles <strong>of</strong> Fucales<br />
from the northern hemisphere. It has been reported from species such as Ascophyllum<br />
nodosum, Fucus vesiculosus, F. serratus, F. spiralis, F. ceranoides, F. evanescens,<br />
Furcellaria lumbricalis <strong>and</strong> Pelvetia canaliculata from Great Britain <strong>and</strong> from Norway<br />
(Wardlaw & Boney 1984, Armstrong et al. 2000).<br />
3.4.2. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
The DNA virus EsV was first isolated from specimens <strong>of</strong> Ectocarpus siliculosus from New<br />
Zeal<strong>and</strong> infecting gametangia <strong>and</strong> sporangia (Müller et al. 1990) <strong>and</strong> has subsequently been<br />
found in host populations around the world (Müller & Stache 1992) giving rise to a large<br />
body <strong>of</strong> literature on aspects <strong>of</strong> this virus <strong>and</strong> its hosts (e.g. Sengco et al. 1996; Kapp et al.<br />
1997).<br />
The ascomycete fungus Haloguignardia tumefaciens has been reported parasitizing<br />
Sargassum sinclairii from Wellington <strong>and</strong> the west coast <strong>of</strong> the South Isl<strong>and</strong> (Cribb & Cribb<br />
1960; Kohlmeyer & Demoulin 1981). Also from Wellington, thraustochytrids <strong>of</strong> either the<br />
genus Thraustochytium or Schizochytrium have been isolated from drift Zonaria<br />
aureomarginata, Durvillaea antarctica <strong>and</strong> Marginariella boryana (Serena Cox, pers<br />
comm.). Karling (1968) isolated Schizochytrium aggregatum from algal debris, which<br />
potentially included brown algae.<br />
Pleurostichidium falkenbergii is an obligate epiphytic red alga on Xiphophora chondrophylla<br />
from northern New Zeal<strong>and</strong> (Bay <strong>of</strong> Isl<strong>and</strong>s, Three Kings <strong>and</strong> North Cape) (Heydrich 1893;<br />
Kylin 1956; Phillips 2000).<br />
In addition to Notheia anomala partially-parasitising Hormosira banksii (Adams 1994)<br />
another parasitic brown alga occurs in New Zeal<strong>and</strong>: Herpodiscus durvillaeae, which is<br />
restricted to New Zeal<strong>and</strong> populations <strong>of</strong> Durvillaea antarctica. It grows epi-endophytically<br />
in its host <strong>and</strong>, in its emergent phase, leads to an erosion <strong>of</strong> the host surface, which may result<br />
in the eventual loss <strong>of</strong> the host phylloid (South 1974; Hay 1978; Peters 1990; Heesch 2005).<br />
An as yet undescribed pigmented endophytic ectocarpalean brown alga is associated with<br />
galls or pale spots on Durvillaea antarctica, D. willana, Marginariella urvilleana, <strong>and</strong><br />
Xiphophora gladiata (Heesch 2005).<br />
3.5. RED ALGAE<br />
3.5.1. Known <strong>pathogens</strong> worldwide<br />
Viruses<br />
Virus-like particles have been observed in the single-celled Porphyridium purpureum<br />
(Chapman & Lang 1973), in Gracilaria conferta <strong>and</strong> in G. epihippisora from the<br />
Mediterranean Sea (Weinberger et al. 1994), as well as in Audouinella saviana from the east<br />
coast <strong>of</strong> USA (Pueschel 1995).<br />
Bacteria<br />
Bacteria are associated with tumour-like growth occurring on the fronds <strong>of</strong> Chondracanthus<br />
teedii (Tsekos 1982). Galls <strong>and</strong> proliferating tissue associated with bacteria are furthermore<br />
found in Acrochaetium species, Ahnfeltia plicata, Bonnemaisonia asparagoides, Ceramium<br />
virgatum, Chondracanthus teedii, Chondrus crispus, Curdiea angustata, Cystoclonium<br />
purpureum, Delesseria sanguinea, Dumontia contorta, Grateloupia filicina, Palmaria<br />
palmata, Plocamium cartilagineum, Polyneuropsis stolonifera, Prionitis decipiens, P.<br />
20 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
filiformis, P. lanceolata, Pterocladiella capillacea <strong>and</strong> Schizymenia dubyi (Cantacuzene 1930;<br />
Apt 1988b; Apt & Gibor 1989; Rheinheimer 1992; Ashen & G<strong>of</strong>f 1996, 1998, 2000).<br />
A number <strong>of</strong> bacterial diseases <strong>of</strong> Porphyra have been reported, particularly in relation to<br />
farmed Porphyra, including “green spot rotting-like deterioration” (Ryokuhan-byo) <strong>of</strong><br />
Porphyra yezoensis (Nakao et al. 1972), "filament bacterial felt" disease (agent not specified)<br />
(Song et al. 1993), <strong>and</strong> "white wasting disease"/ white spot"/ "Gijishirogusare-sho" (Tsukidate<br />
1971, 1977). Tsukidate (1983) examined the symbiotic relationship between Porphyra species<br />
<strong>and</strong> attached bacteria that occurred in conjunction with white rot, the disease which has<br />
caused the most serious damage to the Porphyra cultivation industry in Japan. Anaaki-disease<br />
causes severe damage to the red alga Porphyra yezoensis; Hayashi et al. (1984) identified the<br />
agent as Vibrio fischeri <strong>and</strong> reported on how it attaches to host thalli (Porphyra sp.), digests<br />
host cells <strong>and</strong> makes holes in the thalli. Sunairi et al. (1995) reported Flavobacterium sp. to<br />
be the causative agent <strong>of</strong> Anaaki-disease, as a result <strong>of</strong> several repeated single-colony<br />
isolations <strong>and</strong> infection experiments. In order to ascertain the role <strong>of</strong> bacteria in the process <strong>of</strong><br />
rotting or decaying <strong>of</strong> cultured laver, Fujita et al. (1972) examined 24 strains <strong>of</strong> bacteria<br />
isolated from diseased fronds <strong>of</strong> Porphyra yezoensis, including species <strong>of</strong> Pseudomonas,<br />
Vibrio, Beneckea.<br />
Weinberger et al. (1994) quantified the bacterial epiphytes <strong>of</strong> Gracilaria conferta <strong>and</strong> found<br />
that saprophytic bacteria reached 350 times <strong>and</strong> agar degraders 25,000 times higher numbers<br />
per gram <strong>of</strong> wet weight on tissues infected with the “white tips disease”, as compared to<br />
healthy tissues. Jaffray & Coyne (1996) developed an in situ assay to detect bacterial<br />
<strong>pathogens</strong> <strong>of</strong> the red alga Gracilaria gracilis responsible for causing lesions, thallus<br />
bleaching, <strong>and</strong> Jaffray et al. (1997) examined bacterial epiphytes on Gracilaria gracilis. The<br />
cause for the “white canopy disease” or “colourless disease” described from Gracilaria<br />
tenuistipitata cultivated in Vietnam is not known (Phap & Thuan 2002) although it is<br />
probably similar to “ice-ice disease” in farmed Eucheuma/Kappaphycus species.<br />
Uyenco et al. (1977) isolated strains <strong>of</strong> Pseudomonas, Flavobacterium, <strong>and</strong> Actinobacterium<br />
associated with "ice-ice disease" in diseased Eucheuma striatum. The symptoms <strong>of</strong> this<br />
disease include the presence <strong>of</strong> a white powdery growth on the thallus which causes loss <strong>of</strong><br />
pigments, <strong>and</strong> the gradual consumption <strong>and</strong> subsequent fragmentation <strong>of</strong> the host. Largo et al.<br />
(1995a), found that pathogenic bacteria identified as Vibrio sp. <strong>and</strong> Cytophaga sp. promoted<br />
ice-ice disease in stressed host branches in the carrageenan-producing red algae Kappaphycus<br />
alvarezii <strong>and</strong> Eucheuma denticulatum. Largo et al. (1999) examined the time-dependent<br />
attachment mechanism <strong>of</strong> bacterial <strong>pathogens</strong> during ice-ice infection in Kappaphycus<br />
alvarezii.<br />
Ghirardelli (1998) reported on small sheathed Cyanophyta that occur in the cell walls <strong>of</strong> live<br />
<strong>and</strong> dead crustose rhodophytes, collected in the lower intertidal zone in the Gulf <strong>of</strong> Trieste<br />
(Northern Adriatic Sea, Italy). Pectonema terebrans is a cyanobacterium that grows in the<br />
calcified cell walls <strong>of</strong> coralline algae in Italy, such as Hydrolithon sp., Lithophyllum sp.,<br />
Sporolithon sp. <strong>and</strong> Titanoderma sp., <strong>and</strong> it leaves characteristic holes behind <strong>and</strong> thus can be<br />
identified even in ancient host material (Ghirardelli 1998). The endophytic cyanobacterium<br />
Pleurocapsa sp. is associated with galls <strong>and</strong> the “deformative disease” in Chilean Mazzaella<br />
laminarioides (Correa et al. 1993, 1997, 2000; Sanchez et al. 1996; Buschmann et al. 1997;<br />
Faugeron et al. 2000). Pleurocapsa triggers the development <strong>of</strong> tumours that can result in<br />
major changes in frond morphology <strong>and</strong> texture <strong>and</strong> negatively affect the number <strong>of</strong> spores,<br />
settlement rates, germination success <strong>and</strong> <strong>of</strong>fspring survival (Correa et al. 2000).<br />
An unspecified bacterium is the cause <strong>of</strong> “Coralline Lethal Orange Disease” (CLOD) in the<br />
crustose coralline alga Hydrolithon onkodes from central west Pacific. CLOD is characterised<br />
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y conspicuous bright orange dots associated with tissue necroses that develop into rings<br />
moving over the host thallus as a front <strong>and</strong> leaving completely dead white carbonate skeletons<br />
behind (Littler & Littler 1995; Morcom & Woelkerling 2000).<br />
Animals<br />
Amoeba perforate the cell walls <strong>of</strong> farmed Gracilaria sp., e.g. G. chilensis from Chile, <strong>and</strong><br />
digest the protoplast. Macroscopically the disease manifests as whitish spots spreading rapidly<br />
throughout the host thallus, similar to “ice-ice disease” (Correa & Flores 1995; Largo et al.<br />
1995a; Buschmann et al. 2001).<br />
Larvae <strong>and</strong> adults <strong>of</strong> the harpacticoid copepoda Diathrodes cystoceus <strong>and</strong> D. feldmanni live in<br />
burrows inside the tissue <strong>of</strong> red algae <strong>and</strong> feed on their hosts. Another species, Thalestris<br />
rhodymeniae, burrows in thalli <strong>of</strong> Palmaria palmata. The presence <strong>of</strong> copepoda in red algae is<br />
associated with galls or pinholes (Barton 1892; Apt 1988b; Park et al. 1990; Shimono et al.<br />
2004).<br />
Nematodes are associated with gall formation in Chondrus crispus <strong>and</strong> Furcellaria<br />
lumbricalis, however causality has not been demonstrated for these symbioses (Barton 1901;<br />
Apt 1988b).<br />
Fungi<br />
The most intensively studied fungi in red algae are oomycetes <strong>of</strong> the genus Pythium,<br />
particularly P. marinum <strong>and</strong> P. porphyrae, the latter a pathogen causing “red rot” in Porphyra<br />
species, one <strong>of</strong> the serious epidemics in laver cultures (e.g. Arasaki 1947; Fuller et al. 1966;<br />
Sasaki & Sato 1969; Kazama & Fuller 1970; Sasaki & Sakurai 1972; Sakurai et al. 1974;<br />
Fujita & Zenitani 1976, 1977; Takahashi et al. 1977; Aleem 1980; Tsukidate 1983; Kerwin et<br />
al. 1992; Amano et al. 1995, 1996; Uppalapati & Fujita 2000a, b, 2001; Uppalapati et al.<br />
2001; Park et al. 2001, 2007; Shin 2003a, b; Ding & Ma 2005). The majority <strong>of</strong> this research<br />
has been carried out in Japan <strong>and</strong> Korea although a number <strong>of</strong> studies have also been<br />
conducted in the eastern Pacific in Washington, USA.<br />
<strong>Diseases</strong> <strong>of</strong> the economically important Porphyra include “chytrid blight” disease (Migita<br />
1973; Song et al. 1993), the ascomycete Verrucaria consequens causing Kamenoko disease in<br />
conchocelis cultivation (Migita 1971), Olpidiopsis (Arasaki 1960; Arasaki et al. 1960) <strong>and</strong><br />
also simultaneous infection by red rot <strong>and</strong> chytrid disease reported by Ding & Ma (2005).<br />
One basidiomycete pathogen has been reported for red algae (Porter & Farnham 1986b;<br />
Stanley 1992; Binder et al. 2006). Mycaureola dilseae is a pathogen <strong>of</strong> the subtidal<br />
rhodophyte Dilsea carnosa in the Atlantic north-east. This agent causes necrotic lesions,<br />
which degrade <strong>and</strong> leave holes in the host frond while the fruiting bodies <strong>of</strong> the agent develop<br />
on the margins <strong>of</strong> the holes.<br />
In the Ascomycetes species in the genera Chaudefaudia, Hispidicarpomyces, Spathulospora<br />
have been described from a range <strong>of</strong> hosts (e.g. Cribb & Herbert 1954; Feldmann 1957; Cribb<br />
& Cribb 1960; Kohlmeyer 1963b, 1973a, b, c; Sanson et al. 1990; Nakagiri 1993; Nakagiri &<br />
Ito 1997). Lautitia danica, a pathogen <strong>of</strong> Chondrus crispus, is found in the reproductive tissue<br />
<strong>of</strong> the host, infecting both cystocarpic <strong>and</strong> tetrasporangial regions (Wilson & Knoyle 1961;<br />
Schatz 1984b; Stanley 1992).<br />
Two ascomycetes have been reported to affect commercially important carrageenophytes.<br />
Dewey et al. (1983) reported on Microascus brevicaulis affecting Eucheuma in the<br />
Philippines, <strong>and</strong> Dewey et al. (1984) recorded Penicillium waksmanii isolated from<br />
Kappaphycus in Micronesia.<br />
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Taxa belonging to the Bigyra, including species <strong>of</strong> Olpidiopsis, Eurychasmidium, Petersenia,<br />
Pontisma have been described from a number <strong>of</strong> hosts (e.g. Sparrow 1934, 1936; Aleem<br />
1950b, c, 1952b; Feldmann & Feldmann 1967; Whittick & South 1972; van der Meer &<br />
Pueschel 1985; Molina 1986; West et al. 2006).<br />
The geographic coverage <strong>of</strong> studies on marine fungi in red algae is very incomplete <strong>and</strong><br />
currently reflects the regions where the key workers have been based. For example, the<br />
studies reporting on Chytridium <strong>and</strong> Rhizophidium species are Sparrow (1936) in the northwest<br />
Atlantic, Aleem (1952b) in Sweden <strong>and</strong> Raghukumar (1987a, b) in India. Similarly, the<br />
only papers focusing on thraustochytrids are Quick (1974) in Florida, USA <strong>and</strong> Raghukumar<br />
(1986b, 1987a, b) <strong>and</strong> Raghukumar et al. (1992) in India.<br />
Other algae<br />
Parasitic red algae<br />
More research has been published on red algal <strong>parasites</strong> than on any other area <strong>of</strong> algal<br />
diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> considered in this study. Red algal <strong>parasites</strong> are quite<br />
common, making up to 15% <strong>of</strong> all named red algal genera, although this figure needs revising<br />
as the last comprehensive review <strong>of</strong> red algal <strong>parasites</strong> was by G<strong>of</strong>f (1982). Red algal<br />
parasitism is the most specific symbiosis between two red algae: parasitic red algae are<br />
symbionts that have a reduced size <strong>and</strong> are either completely colourless or have reduced<br />
pigmentation <strong>and</strong> must rely on their nutrition from their host. In the past two decades a lot has<br />
been learned about the origin <strong>of</strong> red algal <strong>parasites</strong> (G<strong>of</strong>f et al. 1996, 1997, Zuccarello et al.<br />
2004), their development (G<strong>of</strong>f & Coleman 1984, 1985; G<strong>of</strong>f & Zuccarello 1994; Zuccarello<br />
& West 1994a), <strong>and</strong> host specificity (Nonomura & West 1981b; G<strong>of</strong>f & Zuccarello 1994;<br />
G<strong>of</strong>f et al. 1997, Zuccarello & West 1994b, c). However only a very small percentage <strong>of</strong><br />
these parasitic red algae have been studied in any detail beyond their first description.<br />
The host specificity <strong>and</strong> evolutionary studies are especially revealing in the context <strong>of</strong> algal<br />
pathogenicity <strong>and</strong> the effects <strong>of</strong> new interactions. Evolutionary studies have revealed that<br />
many red algal <strong>parasites</strong> are derived directly from their hosts (G<strong>of</strong>f et al. 1996, 1997). This<br />
was able to be understood once the development <strong>of</strong> <strong>parasites</strong> on their hosts was elucidated<br />
(G<strong>of</strong>f & Coleman 1984, 1985; G<strong>of</strong>f & Zuccarello 1994), showing that their unusual<br />
development was very similar to post-fertilisation processes in red algae. Early in their<br />
development, either upon spore germination or soon after, red algal <strong>parasites</strong> transfer into a<br />
host cell the cytoplasm <strong>of</strong> an entire cell, or the complete contents <strong>of</strong> a spore. This<br />
“transforms” the host cell as it now develops as a parasite, presumably under the control <strong>of</strong><br />
the transferred parasite nucleus, producing more parasite nuclei <strong>and</strong> dividing to form new<br />
parasite cells. Finally reproductive structures are formed which eventually will lead to new<br />
infections. These developmental processes are similar to the nuclear transfer <strong>and</strong> subsequent<br />
events that occur during post-fertilisation development in most red algae, <strong>and</strong> thus red algal<br />
parasitism has been hypothesised to have been derived from these post-fertilisation processes<br />
(G<strong>of</strong>f et al. 1996, 1997).<br />
Nuclear transfer places constraints on the host range <strong>of</strong> red algal <strong>parasites</strong>. The majority <strong>of</strong> red<br />
algal <strong>parasites</strong> appear to be host specific, although this could be an artifact <strong>of</strong> naming new<br />
parasite species when <strong>parasites</strong> are found on new hosts. However, when host range has been<br />
tested in culture it has been shown to be quite limited (e.g. G<strong>of</strong>f & Zuccarello 1994).<br />
Occasionally <strong>parasites</strong> can grow on closely related host species (Nonomura & West 1981b;<br />
Zuccarello & West 1994b, c), but <strong>of</strong>ten this alternate host development is reduced <strong>and</strong><br />
reproductive structures are not produced. Thus evidence to date supports a high level <strong>of</strong> host<br />
specificity. However, on an evolutionary time scale this is shown not to be so, as “host<br />
jumps” have been discovered using molecular markers. Parasites <strong>of</strong> the Gigartinales family<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 23
Choreocolacaceae have been shown to infect several species or genera <strong>of</strong> hosts, although their<br />
original host was not determined. For example, the parasite Holmsella pachyderma infects<br />
species <strong>of</strong> two genera, Gracilaria <strong>and</strong> Gracilariopsis. The parasite Harveyella mirabilis<br />
infects several species <strong>of</strong> the family Rhodomelaceae plus one member <strong>of</strong> the family<br />
Delesseriaceae. Other studies have shown that while “host jumps” have been accomplished,<br />
the parasite is still found on its original host (G<strong>of</strong>f et al. 1996, 1997). So although not<br />
confirmed to date in culture studies, <strong>parasites</strong> appear to be able to jump hosts. This means that<br />
if introduced to new locations, <strong>and</strong> given there are appropriate host taxa in the new location,<br />
<strong>parasites</strong> may be able to infect new hosts in these environments.<br />
The question <strong>of</strong> the detrimental effects <strong>and</strong> nutritional requirements <strong>of</strong> red algal <strong>parasites</strong> on<br />
their hosts has been barely studied. The few studies that have been conducted show that<br />
although fixed carbon is translocated into the photosynthesis-lacking parasite, this is <strong>of</strong>ten a<br />
small fraction <strong>of</strong> the total fixed carbon (G<strong>of</strong>f 1979; Kremer 1983). No studies have looked at<br />
the effect <strong>of</strong> parasitism on host reproductive success, host recruitment, or the host ability to<br />
withst<strong>and</strong> perturbations.<br />
Endophytic red algae<br />
Endophytic red algae other than <strong>parasites</strong> are pigmented <strong>and</strong> do not form cellular connections<br />
to host cells. Usually they can be cultivated outside their hosts. Photosynthetic red algae<br />
found within the tissues <strong>of</strong> other algae are common. Species <strong>of</strong> Auduoinella sp. (also under<br />
the name <strong>of</strong> Acrochaetium sp., Colaconema sp., Rhodochorton sp.) are <strong>of</strong>ten found<br />
intercellularly within thallose red algae (e.g. West 1979). There have been few experimental<br />
studies <strong>of</strong> the specificity <strong>of</strong> these endophytes, although host range is considered to be fairly<br />
broad. It is possible that these endophytes could infect the tissue <strong>of</strong> new organisms given the<br />
opportunity. Acrochaetium yamadae grows in the tissue <strong>of</strong> Izziella orientalis from Taiwan<br />
(Kylin 1956) <strong>and</strong> <strong>of</strong> Liagora canariensis from the Canary Isl<strong>and</strong>s (Afonso-Carrillo et al.<br />
2003). The former Acrochaetium species, Colaconema asparagopsis <strong>and</strong> C. bonnemaisoniae,<br />
are found in British Bonnemaisonia hamifera <strong>and</strong> Asparagopsis sp., while the related species<br />
C. endophyticum grows in Heterosiphonia sp., (Kylin 1956; White & Boney 1969).<br />
Colaconema ophioglossum is an endophyte <strong>of</strong> Dudresnaya crassa from both sides <strong>of</strong> the<br />
central Atlantic (Afonso-Carrillo et al. 2003).<br />
Some semi-endophytic rhodophytes are found among non-geniculate coralline algae from the<br />
Central Pacific. The thallus <strong>of</strong> Lithophyllum cuneatum from Fiji is wedged into the thalli <strong>of</strong> its<br />
hosts, Neogoniolithon sp. <strong>and</strong> Hydrolithon onkodes. Endophyte <strong>and</strong> hosts do not form cellular<br />
connections; however the growth <strong>of</strong> the host may be disturbed by the presence <strong>of</strong> the<br />
endophyte (Keats 1995; Chamberlain 1999; Morcom & Woelkerling 2000). Similarly,<br />
Amphiroa species (such as A. kuetzingiana) are embedded into their hosts Hydrolithon<br />
onkodes, Neogoniolithon brassica-florida <strong>and</strong> Mesophyllum expansum, but apparently do not<br />
parasitise them (Chamberlain 1999).<br />
In contrast, the epiphyte Titanoderma corallinae has a detrimental effect on its basiphytes<br />
Corallina elongata <strong>and</strong> C. <strong>of</strong>ficinalis from France; contact with its spores leads to bleaching<br />
<strong>of</strong> the host tissue, from which the host may not recover (Cabioch 1979; Chamberlain 1999).<br />
Red algal epiphytes<br />
Most fouling red algae will grow on any surfaces (e.g. Stylonema, Erythrotrichia). These are<br />
<strong>of</strong>ten small algae, with asexual means <strong>of</strong> reproduction that can quickly colonise new surfaces.<br />
Most <strong>of</strong> these algae grow on the surface <strong>of</strong> the host without causing any structural damage to<br />
the host, though shading <strong>of</strong> the host could lead to slowed host growth. Some other red algae<br />
are generalist epiphytes, or at least much more common on algal surfaces (e.g. Microcladia<br />
coulteri - Gonzalez & G<strong>of</strong>f 1989). These algae have different ways <strong>of</strong> interacting with the<br />
24 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
host (Leonardi et al. 2006) with some <strong>of</strong> these interactions leading to damage <strong>and</strong> other<br />
detrimental effects to the host (e.g. tissue loss <strong>and</strong> secondary infections), as these epiphytes<br />
can penetrate host tissue to varying degrees. These algal-epiphyte interactions can especially<br />
be detrimental to cultivated algae (e.g. Gracilaria chilensisi) where hosts are in high<br />
concentration <strong>and</strong> economic loss is possible (Leonardi et al. 2006; Vairappan 2006). A<br />
number <strong>of</strong> filamentous red algae grow as epiphytes on Kappaphycus alvarezii, a<br />
carrageenophyte commercially cultivated throughout Asia. The predominant epiphyte species<br />
is Neosiphonia savatieri, but other species also occur such as Acanthophora sp., Ceramium<br />
sp., Centroceras sp. <strong>and</strong> N. apiculata. The epiphytes are anchored on their host by penetrating<br />
rhizoids. Their presence weakens the host <strong>and</strong> increases its susceptibility to bacteria<br />
(Vairappan 2006).<br />
Ostiophyllum sonderopeltae is an obligate epiphyte <strong>of</strong> Sonderopelta coriacea in Australia<br />
(Kraft 2003). Lembergia allanii is known only on Vidalia colensoi from New Zeal<strong>and</strong>,<br />
whereas Dasyptilon pellucidum is predominantly found on Euptilota formossissima but may<br />
also be found growing on Hymenocladia <strong>and</strong> Cenacrum (Adams 1994). As the specificity <strong>of</strong><br />
the epiphyte habit has not been determined for a number <strong>of</strong> species, <strong>and</strong> where these epiphyte<br />
taxa appear to cause no disease symptoms, epiphyte taxa have not been included in the<br />
database.<br />
Endophytic green algae in red algae<br />
Red algae are the hosts for a number endophytic green algae. The economically important<br />
carrageenophyte Chondrus crispus is infected by Acrochaete heteroclada <strong>and</strong> A. operculata<br />
on both sides <strong>of</strong> the northern Atlantic. Acrochaete heteroclada disrupts the tissue <strong>of</strong> the host<br />
cortex <strong>and</strong> has an overall negative effect on the host performance, slowing the growth <strong>and</strong><br />
decreasing the capacity for regeneration, leading to lower yields <strong>of</strong> carrageenan. A. operculata<br />
likewise penetrates the cortex <strong>of</strong> its host. However, while gametophytes are not invaded<br />
beyond the outer cortex, sporophytes become completely endophytised, resulting in severe<br />
damage <strong>of</strong> the host tissue, secondary bacterial infections, <strong>and</strong> eventually disintegration <strong>and</strong><br />
death <strong>of</strong> the host thallus (Correa & McLachlan 1991, 1992, 1994, Bouarab et al. 1999, 2001b;<br />
Potin et al. 1999, 2002; Bown et al. 2003; Weinberger et al. 2005).<br />
Achrochaete heteroclada is also found in Ahnfeltiopsis furcellata, A. linearis, Chondrus<br />
canaliculatus, Gracilaria chilensis <strong>and</strong> G. mammilaris (Correa & McLachlan 1991), while<br />
another Acrochaete species, A. leptochaete, infects Polysiphonia sp. <strong>and</strong> Champia sp.<br />
(O’Kelly et al. 2004). In Britain, Chondrus crispus hosts another green endophyte, Entocladia<br />
viridis (Bown et al. 2003), a species also found in Phycodrys rubens along the North Atlantic<br />
coasts <strong>of</strong> the USA (O’Kelly et al. 2004).<br />
The endophyte Endophyton ramosum causes “green patch disease” in Chilean Mazzaella<br />
laminarioides. This disease is characterised by fronds which lose their red pigmentation <strong>and</strong><br />
turn green. The host tissue starts decaying, opening the way for secondary bacterial invasions.<br />
Lesions on the stipes lead to their breaking in heavy wave action (Correa et al. 1994, 1997;<br />
Sanchez et al. 1996; Buschmann et al. 1997; Faugeron et al. 2000). Eucheuma ramosum also<br />
inhabits the related host species Mazzaella oregona in the Northeast Pacific (O’Kelly et al.<br />
2004).<br />
Endophytic unicellular sporophytes <strong>of</strong> Acrosiphonia species, originally described as<br />
Chlorochytrium inclusum <strong>and</strong> Codiolum petrocelidis, were observed in a number <strong>of</strong> foliose<br />
<strong>and</strong> crustose red algae, respectively, from British Columbia: Callophyllis sp., Chondrus<br />
crispus, Constantinea subulifera, Dilsea californica, D. integra, Farlowia sp., Haemescharia<br />
hennedyi, Halymenia sp., Hildenbr<strong>and</strong>ia occidentalis, Kallymenia sp., Mastocarpus<br />
papillatus, Mazzaella sanguinea, M. splendens, Palmaria mollis, Porphyra sp., Schizymenia<br />
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pacifica, Sparlingia pertusa, <strong>and</strong> Weeksia sp. (Sussmann et al. 1999, 2005, Sussmann &<br />
DeWreede 2001, 2002, 2005). Acrosiphonia sporophytes also occur in Palmaria mollis <strong>and</strong><br />
Polyides rotundus from the Northeast Atlantic (Sussmann & DeWreede 2002).<br />
Spongomorpha aeruginosa occurs in Haemescharia hennedyi from Germany, <strong>and</strong> the related<br />
species S. mertensii in Mastocarpus papillatus from Canada (Sussmann & DeWreede 2001).<br />
Two endophytic green algae observed in Curdiea racovitzae <strong>and</strong> Iridea cordata from the<br />
Antarctic Peninsula were not further identified (Peters 2003).<br />
Endophytic brown algae in red algae<br />
Brown algae living as endophytes in red algae are from three orders <strong>of</strong> the Phaeophyceae:<br />
Ectocarpales, Laminariales <strong>and</strong> Desmarestiales. Setchell & Gardner (1922) described a<br />
number <strong>of</strong> new species <strong>of</strong> Streblonema, both epiphytic <strong>and</strong> endophytic taxa, including the<br />
endophytes S. corymbiferum (in Cumagloia <strong>and</strong>ersonii) <strong>and</strong> S. investiens (in Helminthocladia<br />
calvadosii). Microspongium tenuissimum occurs in Aeodes orbitosa from South Africa,<br />
Grateloupia doryphora from Canary Isl<strong>and</strong>s, <strong>and</strong> Grateloupia intestinalis from Chile (Peters<br />
2003). A second Microspongium species, M. radians, which has been described from Chilean<br />
Grateloupia doryphora <strong>and</strong> also grows in Mazzaella laminarioides from South Africa<br />
(Burkhardt & Peters 1998; Peters 2003) is considered synonymous to M. tenuissimum<br />
(Heesch 2005). Another endophyte, genetically identified as Microspongium sp., was isolated<br />
from Polysiphonia elongata growing in the Western Baltic Sea (Burkhardt & Peters 1998).<br />
This species may be synonymous with Mikrosyphar polysiphoniae described from Baltic<br />
Polysiphonia stricta. Pedersen (1976) reported Mikrosyphar polysiphoniae in Polysiphonia<br />
arctica in collections from Greenl<strong>and</strong>. Other Mikrosyphar species, such as M. porphyrae, an<br />
endophyte <strong>of</strong> Porphyra sp. in the Baltic Sea, may likewise belong to the genus<br />
Microspongium (Heesch 2005).<br />
Kelp gametophytes have recently been discovered living endophytically in filamentous <strong>and</strong><br />
foliose red algae. Most <strong>of</strong> the hosts belong to the order Ceramiales, such as Antithamnion<br />
densum, Callithamnion acutum, C. biseriatum, Ceramium gardneri, Delesseria decipiens,<br />
Griffithsia pacifica, Herposiphonia plumula, Irtugovia pacifica, Membranoptera platyphylla,<br />
Pleonosporium vancouverianum, Polyneura latissima, Polysiphonia paniculata,<br />
Pterosiphonia dendroidea, Pterosiphonia sp., Pterothamnion pectinatum <strong>and</strong> Scagelia<br />
pylaisei. Kelp gametophytes are furthermore hosted by Fryeella gardneri (Rhodymeniales)<br />
<strong>and</strong> Euthora cristata, Orculifilum denticulatum (Gigartinales). In earlier studies, the species<br />
<strong>of</strong> Laminariales involved were not identified further (Garbary et al. 1999a, b, Garbary & Kim<br />
2000), although more recently Sasaki et al. (2003) were able to identify Agarum clathratum in<br />
Orculifilum denticulatum <strong>and</strong> Hubbard et al. (2004) identified gametophytes <strong>of</strong> Alaria<br />
esculenta <strong>and</strong> Nereocystis luetkeana growing in a number <strong>of</strong> hosts. Gametophytes <strong>of</strong><br />
Desmarestia antarctica grow in Antarctic Curdiea racovitzae (Moe & Silva 1989; Peters<br />
2003).<br />
Although some taxa are predominantly epiphytic, they may also affect the host through some<br />
endophytic development, as found in the epiphyte Elachista antarctica which is anchored<br />
within its Antarctic host Palmaria decipiens by endophytic filaments (Peters 2003).<br />
Endophytic diatoms<br />
Diatoms may either live as endo- or epiphytes in/on red algae. Diatoms such as Achnanthes<br />
longipes, Melosira nummoloides, Synedra gracilis <strong>and</strong> Ligmophora sp. heavily epiphytise<br />
Porphyra species, e.g. P. yezoense, in Japan <strong>and</strong> South Korea. The epiphyte load inhibits<br />
normal growth <strong>of</strong> the basiphyte, leading to a condition called “diatom felt disease” (Tsukidate<br />
1983, 1991; Fujita 1990; Song et al. 1993). Examples <strong>of</strong> endophytic diatoms are Gyrosigma<br />
coelophilum, which has been observed in Coelarthrum opuntia in Japan (Okamoto et al.<br />
26 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
2003), <strong>and</strong> Pseudogomphonema sp., an endophyte <strong>of</strong> Pachymenia sp. from the Antarctic<br />
Peninsula (Peters 2003).<br />
3.5.2. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
Red algal <strong>parasites</strong> are poorly documented in New Zeal<strong>and</strong> although species belonging to the<br />
following genera are known, either reported in publications (e.g. Adams 1994) or recorded in<br />
herbaria: Callocolax, ?Ceratocolax sp., Champiocolax, Choreonema, Colacodasya,<br />
Colacopsis, Gloiocolax, Janczewskia, Levringiella, Microcolax, Plocamiocolax,<br />
Pterocladiophila, Rhodymeniocolax, Sporoglossum, Tylocolax. A great deal more work is<br />
required on the red algal <strong>parasites</strong> in the New Zeal<strong>and</strong> region.<br />
Three species <strong>of</strong> the ascomycete Spathulospora have been described from New Zeal<strong>and</strong><br />
collections - Spathulospora lanata in Camontagnea oxyclada, S. adelpha <strong>and</strong> S. calva on<br />
Ballia callitricha (Kohlmeyer 1973a). Kohlmeyer & Demoulin (1981) described two<br />
ascomycete fungi that are found in association with the New Zeal<strong>and</strong> endemic genus<br />
Apophlaea - Mycophycias apophlaeae <strong>and</strong> Polystigma apophlaeae Kohlm.<br />
The endophytic brown alga Microspongium tenuissimum (incl. M. radians) occurs in three red<br />
algae from New Zeal<strong>and</strong>: Pachymenia lusoria, Grateloupia intestinalis <strong>and</strong> in a so far<br />
undescribed species <strong>of</strong> the family Kallymeniaceae (Heesch 2005). Another species,<br />
Mikrosyphar pachymeniae was described from northern populations <strong>of</strong> P. lusoria, but may be<br />
synonymous with Microspongium tenuissimum (Heesch 2005).<br />
3.6. GREEN ALGAE<br />
3.6.1. Known <strong>pathogens</strong> worldwide<br />
Viruses<br />
No virus infections have been reported for marine green algae.<br />
Bacteria<br />
No bacterial diseases have been reported for marine green algae.<br />
Animals<br />
Two unidentified protozoa, a ciliate <strong>and</strong> a flagellate, live endophytically in Codium bursa<br />
(Armstrong et al. 2000), while an amoeba has been reported from Blidingia chadefaudii<br />
(Feldmann & Feldmann 1967). In the Florida Keys an amphipod (Erichthonius brasiliensis)<br />
affects the growth <strong>of</strong> the green alga Halimeda tuna by rolling its terminal segments (Sotka et<br />
al. 1999).<br />
Fungi<br />
Species <strong>of</strong> the genus Cladophora host a number <strong>of</strong> pathogenic fungi, such as Labyrinthula<br />
spp. (e.g. L. coenocystis), Coenomyces sp., Achlyogeton salinus, Entophlyctis maxima,<br />
Olpidium rostiferum <strong>and</strong> Sirolpidium bryopsidis (Dangeard 1931a; Sparrow 1936;<br />
Raghukumar 1986a, 1987b; Rheinheimer 1992; Hyde et al. 1998; Raghukumar 2002). In<br />
India, a thraustochytrid fungus infects Cladophora liebetruthii (Raghukumar 1986a).<br />
Blodgettiomyces bornetii is a fungus occuring in Cladophora catenata <strong>and</strong> other Cladophora<br />
species, as well as in Siphonocladus rigidus (Kohlmeyer & Kohlmeyer 1972; Porter &<br />
Farham 1986a). Blodgettia sp. occurs in Cladophora dalmatica (Saccardo 1882a).<br />
Labyrinthula sp. also occurs in Chaetomorpha <strong>and</strong> Rhizoclonium species. The latter moreover<br />
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hosts Coenomyces sp. <strong>and</strong> Olpidium rostiferum (Raghukumar 1986b, 1987a, 2002; Hyde et al.<br />
1998).<br />
Pontisma lagenidioides causes the “browning disease” in Chaetomorpha antennina<br />
(Raghukumar 1987a; Raghukumar & Ch<strong>and</strong>ramohan 1988). Thraustochytrium proliferum,<br />
Rhizophydium littoreum, R. globosum <strong>and</strong> Phlyctochytrium sp. are <strong>pathogens</strong> <strong>of</strong> Bryopsis<br />
plumosa (Sparrow 1936; Kazama 1972; Amon 1984; Hyde et al. 1998) <strong>and</strong> the former also<br />
infects Codium sp. (Amon 1984). Olpidiopsis <strong>and</strong>reii infects filamentous green algae, e.g.<br />
Acrosiphonia sp. <strong>and</strong> Spongomorpha sp. (Aleem 1952a; Porter & Farnham 1986a; West et al.<br />
2006).<br />
The ascomycetes Guignardia alaskana <strong>and</strong> G. prasiolae have been reported from Prasiola<br />
borealis <strong>and</strong> Prasiola tesselata, respectively. The former is also parasitised by Laestadia<br />
alaskana. Both algae furthermore host Turgidosculum complicatulum, while the related<br />
species T. ulvae occurs in Blidingia minima <strong>and</strong> B. minima var. vexata (Saccardo 1882b; Reed<br />
1902; Kohlmeyer & Kohlmeyer 1972, 1973; Kohlmeyer 1979) <strong>and</strong> Ulva californica (Reed<br />
1902). In France, the ascomycete Chadefaudia corallinarum infests Flabellia petiolata <strong>and</strong><br />
Halimeda tuna (Kohlmeyer 1963b). In Russia’s Sea <strong>of</strong> Japan, Ulva fenestrata is endophytised<br />
by Ulocladium littoreum (Pivkin & Zvereva 2000).<br />
Ostreobium queketti, an endolithic alga growing in corals from French-Polynesia, is<br />
parasitised by an aspergillus-like fungus, causing a black b<strong>and</strong>ing pattern on the coral host<br />
(Priess et al. 2000). In Sweden, an unspecified fungus has been reported to parasitise<br />
Elasticha fucicola (Aleem 1952a).<br />
A unidentified heterokont biflagellate parasite lives inside Codium fragile from the North<br />
American Atlantic coast, consuming the plastids <strong>of</strong> its host (Lee & Kugrens 2003).<br />
Other algae<br />
Members <strong>of</strong> the genus Achrochaete occur as endophytes in Ulva rigida <strong>and</strong> Codium fragile.<br />
Another endophyte, Entocladia viridis has been found in Bryopsis duplex <strong>and</strong> Chaetomorpha<br />
linum from Italy <strong>and</strong> Denmark, respectively (O’Kelly 1981; Nielsen 1979; del Campo et al.<br />
1998). Another green seaweed, Chlorochytrium dermatocolax, has been reported as a parasite<br />
<strong>of</strong> a green host, Bryopsis plumosa (Sparrow 1936).<br />
There is a single record <strong>of</strong> a red seaweed (Schmitziella endophloea) as an endophyte in<br />
Cladophora pellucida (Kylin 1956).<br />
3.6.2. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
The labyrinthulid Thraustochytrium proliferum has been isolated from Bryopsis plumosa <strong>and</strong><br />
Cladophora sp. from Dunedin (Karling 1968).<br />
3.7. XANTHOPHYCEAE<br />
3.7.1. Known <strong>pathogens</strong> worldwide<br />
Viruses<br />
No virus infections have been reported for the Xanthophyceae.<br />
Bacteria<br />
No bacterial infections have been reported for the Xanthophyceae.<br />
28 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Animals<br />
A rotifer was reported to cause galls in Vaucheria sp. (Apt 1988b).<br />
Fungi<br />
No fungal infections have been reported for the Xanthophyceae.<br />
Other algae<br />
No algal infections have been reported for the Xanthophyceae.<br />
3.7.2. Occurrence <strong>of</strong> known <strong>pathogens</strong> in New Zeal<strong>and</strong><br />
No infections have been reported for the Xanthophyceae in New Zeal<strong>and</strong>.<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 29
4. Discussion<br />
I: ASSESSMENT OF INFORMATION AVAILABLE ON SEAWEED DISEASES<br />
WORLDWIDE AND IN NEW ZEALAND<br />
A number <strong>of</strong> general reviews have dealt with <strong>pathogens</strong> <strong>of</strong> marine algae, e.g. Evans et al.<br />
(1978) <strong>and</strong> G<strong>of</strong>f (1982) focusing on parasitic red algae; Andrews (1979a, b) on the pathology<br />
<strong>of</strong> seaweeds; Apt (1988b) on galls <strong>and</strong> tumour-like growths; Correa (1997) examining the<br />
current knowledge <strong>and</strong> approaches to infectious diseases <strong>of</strong> marine algae; Bouarab et al.<br />
(2001a) examining the ecological <strong>and</strong> biochemical aspects <strong>of</strong> algal infectious diseases. Fujita<br />
(1990) authored a review specifically on the diseases <strong>of</strong> cultivated Porphyra in Japan.<br />
Biosecurity NZ requested that data in this project should be compiled for each pathogen<br />
including:<br />
• agent stability <strong>and</strong> inactivation data;<br />
• epidemiological features:<br />
− geographical range <strong>and</strong> features <strong>of</strong> distribution (international spread);<br />
− host range (including prevalence <strong>and</strong> incidence, resistant strains/species, life stage<br />
susceptibility <strong>and</strong> course <strong>of</strong> infection, habitat <strong>and</strong> seasonality);<br />
− morbidity/mortality rates;<br />
− transmission (including route <strong>and</strong> infectious dose).<br />
• host impact:<br />
− tissue tropism (site <strong>of</strong> infection);<br />
− brief description <strong>of</strong> major pathological <strong>and</strong> biological effects.<br />
• diagnostics <strong>and</strong> disease control:<br />
− key diagnostic features;<br />
− overview <strong>of</strong> diagnostic methods, including sensitivity <strong>and</strong> specificity;<br />
− disease management activities worldwide;<br />
− able to be eradicated?<br />
The majority <strong>of</strong> papers did not include data in these areas. Generally, the information on<br />
diseases <strong>of</strong> seaweeds is very patchy <strong>and</strong> the emphasis <strong>of</strong> published work lies in two main<br />
areas:<br />
• diseases occurring in monocultures <strong>of</strong> farmed species, mainly in East <strong>and</strong> Southeast Asia<br />
(particularly affecting the key economic genera Porphyra. Laminaria, <strong>Undaria</strong>,<br />
Gracilaria, Eucheuma <strong>and</strong> Kappaphycus);<br />
• observations <strong>of</strong> certain groups <strong>of</strong> <strong>pathogens</strong> in particular geographic regions as a<br />
consequence <strong>of</strong> the research interests <strong>of</strong> a particular team or research group, leading to<br />
“pockets <strong>of</strong> information”.<br />
The amount <strong>of</strong> information contained in the references we investigated varied greatly between<br />
articles, ranging from reports <strong>of</strong> the occurrence <strong>of</strong> <strong>pathogens</strong> to multi-paper treatments <strong>of</strong><br />
certain diseases. The latter are especially numerous for farmed macroalgae e.g. Pythium<br />
porphyrae, the agent causing the red rot disease in Porphyra species (Porphyra cultivation is<br />
a billion dollar industry in Asian countries). Other agents, in contrast, have only been<br />
observed once <strong>and</strong> <strong>of</strong>ten only incidentally in the course <strong>of</strong> other research.<br />
Problems with the correct identifications <strong>and</strong> classification <strong>of</strong> <strong>pathogens</strong> may lead to different<br />
names for the same agent or the same name for different agents. For example, small algae<br />
such as endophytes may have a reduced morphology, leaving only few characters for<br />
30 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
identification. Often, they are not fertile when observed, making the correct identification<br />
difficult.<br />
Host symptoms are not always a good character for identifying <strong>pathogens</strong>/ <strong>parasites</strong> either, as<br />
these may depend on the susceptibility <strong>of</strong> a host species to a specific agent. Susceptibility can<br />
vary between generations <strong>of</strong> the same host species e.g. the sporophyte <strong>of</strong> the red alga<br />
Chondrus crispus is susceptible to infections <strong>of</strong> the green endophyte Acrochaete operculata,<br />
while the host gametophyte shows some resistance (Correa & McLachlan 1991). Life stages<br />
<strong>of</strong> <strong>pathogens</strong> may also display different morphologies e.g. nauplii <strong>of</strong> harpacticoid copepoda<br />
burrowing in kelp stipes may have to be reared to adult stages in order to correctly identify<br />
them by morphology. Most references describe a disease by the symptoms expressed in the<br />
host, but fall short <strong>of</strong> demonstrating causality, meaning for example, more obvious secondary<br />
invaders could be mistakenly attributed as the primary cause <strong>of</strong> a disease. There is almost no<br />
work that has examined more complex pathogen/host systems.<br />
Information on diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> occurring in New Zeal<strong>and</strong> is scant.<br />
Generally the publications available reflect the activities <strong>of</strong> a few overseas workers who have<br />
visited or received specimens <strong>and</strong> have published on particular agents (e.g. Kohlmeyer).<br />
There have been no focused studies incorporating field <strong>and</strong> laboratory investigations other<br />
than the work <strong>of</strong> Heesch (2005) on endophytes <strong>of</strong> brown algae in New Zeal<strong>and</strong>, completed as<br />
research for a Ph.D. at the University <strong>of</strong> Otago.<br />
II: ASSESSMENT OF THREATS BY PATHOGENS OF UNDARIA TO NEW ZEALAND<br />
NATIVE MARINE FLORA<br />
The only disease reported in <strong>Undaria</strong> from its introduced range is the infection <strong>of</strong> thalli with<br />
the pigmented endophytic brown alga Laminariocolax aecidioides, both in Spain (Veiga et al.<br />
1997) <strong>and</strong> in Argentina (Gauna et al. personal communication). It is not clear whether this<br />
endophyte originates from Japanese populations introduced with the host or from European or<br />
Argentinian populations respectively. Laminariocolax aecidioides is known from other,<br />
native European kelps such as Laminaria hyperborea in the German Bight <strong>and</strong> Norway, <strong>and</strong><br />
Saccharina latissima in the Western Baltic Sea (Lein et al. 1991; Ellerstdottir & Peters 1995,<br />
1997; Peters & Schaffelke 1996), but it has not been reported from southern Europe. It also<br />
occurs in the native range <strong>of</strong> U. <strong>pinnatifida</strong>, in Japan (Yoshida & Akiyama 1978). Genetic<br />
studies may determine the origin <strong>of</strong> the Spanish <strong>and</strong> Argentinean populations <strong>and</strong> thus shed<br />
some light on whether endophytes were or can be transmitted with host sporophytes (or other<br />
disease agents).<br />
In the Western Baltic, thalli <strong>of</strong> Saccharina latissima infected with Laminariocolax aecidioides<br />
show more severe symptoms in shallow water, due to the endophyte growth being accelerated<br />
in better light conditions. Increased severity <strong>of</strong> infection symptoms prevent host thalli<br />
surviving in water depths <strong>of</strong> 2 m, in contrast to deeper water where growth <strong>of</strong> the endophyte is<br />
light limited (Schaffelke et al. 1996). In New Zeal<strong>and</strong>, Laminariocolax macrocystis, a closely<br />
related species in this genus, infects native kelps, such as Macrocystis pyrifera <strong>and</strong> Ecklonia<br />
radiata, <strong>and</strong> in severe cases this leads to crippled thalli (M. pyrifera) <strong>and</strong>/or stunted growth<br />
(E. radiata) (Heesch 2005). It is not known if this endophyte species has an influence on the<br />
depth distribution <strong>of</strong> its hosts. Further, it is not known if L. aecidioides would be able to infect<br />
New Zeal<strong>and</strong> native kelps, <strong>and</strong> if so, what the consequences would be for native kelp<br />
populations.<br />
Reports in the international literature have highlighted the occurrence <strong>of</strong> kelp gametophytes as<br />
endophytes in a range <strong>of</strong> hosts (e.g. Garbary et al. 1999a, b; Garbary & Kim 2000; Sasaki et<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 31
al. 2003; Hubbard et al. 2004). One <strong>of</strong> the potential threats <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> to New<br />
Zeal<strong>and</strong> native kelps may consist <strong>of</strong> competition with other kelp gametophytes as endophytes.<br />
III: FUTURE STRATEGY FOR SCREENING POPULATIONS AND INCREASING<br />
KNOWLEDGE OF RISK POSED BY DISEASES/PARASITES/PATHOGENS TO NEW<br />
ZEALAND MACROALGAE AND COASTAL COMMUNITIES<br />
None <strong>of</strong> the known <strong>pathogens</strong> <strong>of</strong> <strong>Undaria</strong> have so far been observed in/on U. <strong>pinnatifida</strong> in<br />
New Zeal<strong>and</strong>, however, populations <strong>of</strong> U. <strong>pinnatifida</strong> around New Zeal<strong>and</strong> have not been<br />
screened for the presence <strong>of</strong> diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong>. Given that there is evidence<br />
that New Zeal<strong>and</strong> has received at least 10 separate introduction events <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong><br />
(Uwai et al. 2006), it would be important to construct a sampling regime that reflected this<br />
known genetic diversity within New Zeal<strong>and</strong> populations <strong>of</strong> <strong>Undaria</strong>.<br />
Correa (1997) recommends an operational approach to the study <strong>of</strong> infectious diseases in<br />
seaweeds:<br />
1. field <strong>and</strong> laboratory observations aiming to individualize a potential pathogen <strong>and</strong> to<br />
describe the lesions associated with the presence <strong>of</strong> that organism,<br />
2. laboratory experiments <strong>and</strong> observations to establish causality i.e. applying Koch’s<br />
postulates (Andrews & G<strong>of</strong>f 1984), as well as manipulative experiments to underst<strong>and</strong><br />
aspects <strong>of</strong> the host-pathogen relationship <strong>and</strong> thus develop methods to manage the<br />
disease, e.g. in marine cultures<br />
3. epidemiology to “evaluate... the population segment ...affected..., the severity <strong>of</strong> the<br />
disease <strong>and</strong> the occurrence <strong>of</strong> seasonal <strong>and</strong> spatial patterns <strong>of</strong> disease expression”, which<br />
includes the study <strong>of</strong> the reproduction, mortality <strong>and</strong> physiological performance <strong>of</strong> the<br />
host population <strong>and</strong> individuals.<br />
From the research conducted by Heesch (2005) it is clear that it is necessary to identify host<br />
populations <strong>and</strong> look for disease symptoms both intra- <strong>and</strong> inter-annually, with seasonal<br />
sampling occurring ca. quarterly. Given the range <strong>of</strong> environments, water temperatures, <strong>and</strong><br />
photoperiods experienced through the New Zeal<strong>and</strong> region, the sampling would need to be<br />
stratified <strong>and</strong> targeted on priority taxa. Depending on the biology <strong>of</strong> the target taxa the<br />
sampling regime would need to incorporate considerations <strong>of</strong> the species life history (i.e.<br />
whether the species has isomorphic or heteromorphic alternation <strong>of</strong> generations or direct<br />
development; if life history phases have differing cell wall chemistry as found for example in<br />
isomorphic phases <strong>of</strong> members <strong>of</strong> the Gigartinales), ecology <strong>and</strong> distribution (light, depth,<br />
exposure/shelter, substrate). Causality between disease <strong>and</strong> symptoms requires both field <strong>and</strong><br />
detailed laboratory investigations.<br />
A number <strong>of</strong> authors point to the importance <strong>of</strong> considering diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong><br />
in the wider context, testing hypotheses about the roles they may play in shaping population<br />
<strong>and</strong> community structure (Correa 1997; Prenter et al. 2004; Tompkins & Poulin 2006).<br />
32 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
5. Conclusions<br />
Seaweeds that are diseased are under-collected in New Zeal<strong>and</strong> <strong>and</strong>, as a consequence, the<br />
status <strong>of</strong> knowledge about biotic diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> is deficient: it is not<br />
possible to evaluate risk posed by introduced diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> on the basis <strong>of</strong><br />
current underst<strong>and</strong>ing <strong>of</strong> the native biota.<br />
Whilst experts in the field <strong>of</strong> algal diseases such as Correa (1997) stress the need for studies<br />
on the mechanisms <strong>of</strong> infection <strong>and</strong> the spread <strong>of</strong> the <strong>pathogens</strong> within <strong>and</strong> among host<br />
individuals, as well as on the genetics <strong>of</strong> the host-pathogen interaction, the basic underpinning<br />
surveys <strong>and</strong> research are required in New Zeal<strong>and</strong> to document the biodiversity <strong>and</strong><br />
distribution <strong>of</strong> diseases, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> within macroalgae.<br />
6. Acknowledgements<br />
The following people are thanked for their contributions to this study: Megan Gee for<br />
literature searching <strong>and</strong> acquisition, Helen Sui for developing the database structure, Joe<br />
Zuccarello for assistance with reviewing literature on red algal <strong>parasites</strong>, Joe Buchanan <strong>and</strong><br />
Peter Martin for assistance with literature reviews, Tracy Farr for assistance with maps, Hoe<br />
Chang, Janet Grieve <strong>and</strong> Roberta D’Archino for assistance with translations.<br />
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Appendix 4. Map <strong>of</strong> known <strong>pathogens</strong> affecting <strong>Undaria</strong> <strong>pinnatifida</strong> in its native range <strong>of</strong> Japan, China <strong>and</strong> Korea. Locations are approximate<br />
<strong>and</strong> inferred from placenames available in the literature.<br />
120˚E<br />
150˚E<br />
30˚N<br />
0 500<br />
km<br />
☼<br />
Amenophia (copepod)<br />
Parathalestris (copepod)<br />
Thalestris (copepod)<br />
Scutellidium (copepod)<br />
Ceinina (amphipod)<br />
SOUTH<br />
KOREA<br />
Honshu<br />
JAPAN<br />
☼ Olpidiopsis (fungi)<br />
Unspecified bacteria<br />
Vibrio (bacteria)<br />
40˚N<br />
Dalian<br />
CHINA<br />
Moraxella (bacteria)<br />
Flavobacterium (bacteria)<br />
Pseudomonas (bacteria)<br />
Hokkaido<br />
Halomonas (bacteria)<br />
Laminariocolax (algae)<br />
Microspongium (algae)
Appendix 3. Map showing FAO geographic regions, distribution <strong>of</strong> <strong>Undaria</strong> in these regions <strong>and</strong> <strong>pathogens</strong> ( Laminariocolax (algae);<br />
Microspongium (algae)) affecting <strong>Undaria</strong> in its introduced range. Native range (dark grey shading); introduced range (mid grey shading).
APPENDIX 1:<br />
Hierarchical Classification (based on Cavalier –Smith 1998) <strong>and</strong> Species 2000<br />
(D.Gordon, pers. comm. NIWA)<br />
EMPIRE OR SUPERKINGDOM 1. PROKARYOTA<br />
Kingdom 1. Bacteria<br />
Subkingdom 1. Negibacteria<br />
Infrakingdom 1. Eobacteria<br />
Phylum 1. Eobacteria<br />
Class 1. Chlorobacteria [e.g. Chlor<strong>of</strong>lexus, Heliothrix, Thermomicrobium]<br />
Class 2. Hadobacteria [e.g. Deinococcus, Thermus]<br />
Infrakingdom 2. Glycobacteria<br />
Phylum 1. Cyanobacteria<br />
Subphylum 1. Gloeobacteria<br />
Class 1. Gloeobacteria<br />
Order 1. Gloeobacterales [e.g. Gloeobacter]<br />
Subphylum 2. Phycobacteria<br />
Class 1. Chroobacteria<br />
Order 1. Chroococcales [e.g. Anabaena, Prochloron]<br />
Order 2. Pleurocapsales [e.g. Pleurocapsa]<br />
Order 3. Oscillatoriales [e.g. Oscillatoria]<br />
Class 2. Hormogoneae<br />
Order 1. Nostocales [e.g. Nostoc]<br />
Order 2. Stigonemates [e.g. Stigonema]<br />
Phylum 2. Spirochaetae<br />
Class Spirochaetes [e.g. Leptospira, Spirochaeta, Treponema]<br />
Phylum 3. Sphingobacteria<br />
Class 1. Flavobacteria [Fibrobacter, Flavobacterium]<br />
Class 2. Chlorobia [e.g. Cytophaga, Flavobacteria]<br />
SUPERPHYLUM EXOFLAGELLATA<br />
Phylum 1. Planctobacteria<br />
Class 1. Planctomycea [e.g. Pirellula, Planctomyces]<br />
Class 2. Verrucomicrobeae [e.g. Verrucomicrobium]<br />
Class 3. Chlamydiae [e.g. Chlamydia]<br />
Phylum 2. Proteobacteria<br />
Subphylum 1. Rhodobacteria<br />
Class 1. Chromatibacteria [e.g. Chromatium, Escherichia, Haemophilus, Methylococcus, Pseudomonas,<br />
Spirillum, Vibrio]<br />
Class 2. Alphabacteria [e.g. Agrobacterium, Caulobacter, Hyphomicrobium, Rhizobium, Rhodospirillum,<br />
Rickettsia]<br />
Subphylum 2. Thiobacteria<br />
Class 1. Deltabacteria[e.g. Bdellovibrio, Desulfovibrio, Myxococcus]<br />
Class 2. Epsilobacteria [e.g. Aquifex, Helicobacter, Hydrogenobacter, Thermotoga]<br />
Subphylum 3. Geobacteria<br />
Class 1. Ferrobacteria [e.g. Geobacter, Leptospirillum, Magnetobacterium]<br />
Class 2. Acidobacteria [e.g. Acidobacterium, Holophaga, Geothrix]<br />
Subkingdom 2. Unibacteria<br />
Phylum 1. Posibacteria<br />
Subphylum 1. Endobacteria<br />
Class 1. Togobacteria [e.g. Heliobacterium, Selenomonas, Thermotoga]<br />
Class 2. Teichobacteria [e.g. Bacillus, Clostridium, Staphylococcus, Streptococcus]<br />
Class 3. Mollicutes [e.g. Mycoplasma]<br />
Subphylum 2. Actinobacteria<br />
Class 1. Arthrobacteria [e.g. Arthrobacter, Actinomyces]<br />
Class 2. Arabobacteria<br />
Order 1. Actinoplanales [e.g. Actinoplanes]<br />
Order 2. Mycobacteriales [Mycobacterium]<br />
Class 3. Streptomycetes [e.g. streptomyces]<br />
Phylum 2. Archaebacteria<br />
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Subphylum 1. Euryarchaeota<br />
Superclass 1. Neobacteria<br />
Class 1. Methanothermea [e.g. Methanococcus]<br />
Class 2. Archaeoglobea<br />
Class 3. Halomebacteria [e.g. Halobacterium, Methanospirillum]<br />
Superclass 2. Eurythermea<br />
Class 1. Protoarchaea [e.g. Palaeococcus, Protococcus]<br />
Class 2. Picrophilea [e.g. Ferroplasma, Thermoplasma]<br />
Subphylum 2. Crenarchaeota<br />
Class 1. Crenarchaeota [e.g. Sulfolobus, Pyrobaculum]<br />
EMPIRE OR SUPERKINGDOM 2. EUKARYOTA<br />
Kingdom 1. Protozoa<br />
Subkingdom 1. Sarcomastigota<br />
Phylum 1. Amoebozoa [Rhizopoda]<br />
Subphylum 1. Protamoebae<br />
Class 1. Breviatea<br />
Class 2. Lobosea<br />
Order 1. Euamoebida [e.g. Amoeba, Rhizamoeba]<br />
Order 2. Copromyxida [e.g. Copromyxa]<br />
Order 3. Arcellinida [e.g. Arcella, Difflugia]<br />
Class 3. Discosea<br />
Order 1. Glycostylida [e.g. Paramoeba, Vannella]<br />
Order 2. Himatismenida [e.g. Cochliopodium]<br />
Order 3. Dermamoebida [e.g. Thecamoeba]<br />
Class 4. Variosea<br />
Order 1. Phalansteriida [e.g. Phalansterium]<br />
Order 2. Centramoebida [e.g. Acanthamoeba]<br />
Order 3. Varipodida [e.g. Filamoeba, Gephyramoeba]<br />
Subphylum 2. Conosa<br />
Infraphylum 1. Archamoebae<br />
Class 1. Archamoebea<br />
Order 1. Pelobiontida [e.g. Entamoeba, Pelomyxa]<br />
Order 2. Mastigamoebida [e.g. Endolimax, Mastigamoeba]<br />
Infraphylum 2. Mycetozoa<br />
Class 1. Stelamoebea<br />
Order 1. Protostelida [e.g. Protostelium, Schizoplasmodium]<br />
Order 2. Dictyosteliida [e.g. Dictyostelium]<br />
Class 2. Myxogastrea<br />
Order 1. Parastelida [e.g. Ceratiomyxa]<br />
Order 2. Echinosteliida [e.g. Echinostelium]<br />
Order 3. Liceida [e.g. Listerella]<br />
Order 4. Trichiida [e.g. Dianema]<br />
Order 5. Stemonitida [e.g. Stemonitis]<br />
Order 6. Physarida [e.g. Didymium, Elaeomyxa, Physarum]<br />
Phylum 2. Choanozoa<br />
Class 1. Choan<strong>of</strong>lagellatea<br />
Order 1. Craspedida [e.g. Codosiga, Monosiga, Salpingoeca]<br />
Order 2. Acanthoecida [e.g. Acanthoeca, Diaphanoeca]<br />
Class 2. Corallochytrea<br />
Order 1. Corallochytrida [e.g. Corallochytrium]<br />
Class 3. Ichthyosporea<br />
Order 1. Ichthyosporida [e.g. Dermocystidium, Ichthyophonus]<br />
Class 4. Cristidiscoidea<br />
Order 1. Ministeriida [e.g. Ministeria]<br />
Order 2. Nucleariida [e.g. Fonticula, Nuclearia]<br />
Subkingdom 2. Biciliata<br />
Infrakingdom 1. Rhizaria<br />
Phylum 1. Cercozoa [Zo<strong>of</strong>lagellata]<br />
Subphylum 1. Filosa<br />
Superclass 1. Reticul<strong>of</strong>ilosa<br />
Class 1. Chlorarachnea [e.g. Chlorarachnion]<br />
Class 2. Proteomyxidea [e.g. Dimorpha, Gymnophrys, Reticulamoeba]<br />
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Superclass 2. Monad<strong>of</strong>ilosa<br />
Class 1. Sarcomonadea [e.g. Cercomonas, Heteromita, Metopion]<br />
Class 2. Thec<strong>of</strong>ilosea [e.g. Cryothecomonas, Cryptodifflugia]<br />
Class 3. Spongomonadea [e.g. Spongomonas]<br />
Class 4. Imbricatea [e.g. Euglypha, Thaumatomonas]<br />
Class 5. Phaeodaria [e.g. Collosphaera]<br />
Subphylum 2. Endomyxa<br />
Class 1. Phytomyxea<br />
Order 1. Phagomyxida [e.g. Phagomyxa]<br />
Order 2. Plasmodiophorida [e.g. Plasmodiophora]<br />
Class 2. Ascetosporea<br />
Order 1. Haplosporida [e.g. Bonamia, Haplosporidium, Urosporidium]<br />
Order 2. Paramyxida [e.g. Paramyxa]<br />
Order 3. Claustrosporida [e.g. Claustrosporidium]<br />
Class 3. Gromiidea<br />
Order 1. Gromiida [e.g. Gromia]<br />
Phylum 2. Foraminifera<br />
Class 1. Athalamea [e.g. Reticulomyxa]<br />
Class 2. Polythalamea [e.g. Allogromia, Globigerina, Textularia]<br />
Class 3. Xenophyophorea [e.g. Psammina]<br />
Phylum 3. Radiozoa<br />
Class 1. Acantharea [e.g. Acanthometra]<br />
Class 2. Sticholonchea [e.g. Sticholonche]<br />
Class 3. Polycystinea [e.g. Collozoum]<br />
Infrakingdom 1. Excavata<br />
SUPERPHYLUM 1. APUSOZOA<br />
Phylum Apusozoa<br />
Class 1. Diphyllatea<br />
Order 1. Diphylleida [e.g. Collodictyon, Diphylleia]<br />
Class 2. Thecomonadea<br />
Order 1. Apusomonadida [e.g. Amastigomonas, Apusomonas]<br />
Order 2. Ancyromonadida [e.g. Ancyromonas]<br />
Order 3. Hemimastigida [e.g. Spironema]<br />
Class 3. Teonemea [e.g. Nephromyces, Telonema]<br />
SUPERPHYLUM 2. EOZOA<br />
Phylum 1. Loukozoa<br />
Class 1. Jakobea<br />
Order 1. Jakobida [e.g. Histiona, Jakoba, Reclinomonas]<br />
Class 2. Malawimonadea<br />
Order 1. Malawimonadida [e.g. Malawimonas]<br />
Phylum 2. Metamonada<br />
Subphylum 1. Anaeromonada<br />
Class 1. Anaeromonadea [e.g. Dinenympha, Personympha, Trimastix]<br />
Order 1. Trimastigida [e.g. Trimastix]<br />
Order 2. Oxymonadida [e.g. Dinenympha, Pyrsonympha]<br />
Subphylum 2. Trichozoa<br />
Superclass 1. Parabasalia<br />
Class 1. Trichomonadea<br />
Order 1. Trichomonadida [e.g. Calonympha, Trichomonas]<br />
Order 2. Lophomonadida [e.g. Microjoenia, Lophomonas]<br />
Order 3. Spirotrichonymphida [e.g. Holomastigotoides]<br />
Class 2. Trichonymphea<br />
Order 1. Trichonymphida [e.g. Trichonympha]<br />
76 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Superclass 2. Carpediemonadia<br />
Class 1. Carpediemonadea<br />
Order 1. Carpediemonadida [e.g. Carpediemonas]<br />
Superclass 3. Eopharyngia<br />
Class 1. Trepomonadea<br />
Subclass 1. Diplozoa<br />
Order 1. Distomatida [e.g. Hexamita, Spironucleus, Trepomonas]<br />
Order 2. Giardiida [e.g. Giardia, Octomitus]<br />
Subclass 2. Enteromonadia<br />
Order 1. Enteromonadida [e.g. Enteromonas]<br />
Class 2. Retortamonadea<br />
Order 1. Retortamonadida [e.g. Chilomastix, Retortamonas]<br />
SUPERPHYLUM 3. DISCICRISTATA<br />
Phylum 1. Percolozoa<br />
Class 1. Heterolobosea<br />
Order 1. Schizopyrenida [ e.g. Naegleria, Tetramitus, Vahlkampfia]<br />
Order 2. Acrasida [ e.g. Acrasis]<br />
Order 3. Lyromonadida [ e.g. Lyromonas, Psalteriomonas]<br />
Class 2. Percolatea<br />
Order 1. Percolomonadida [e.g. Percolomonas]<br />
Order 2. Pseudociliatida [e.g. Stephanopogon]<br />
Phylum 2. Euglenozoa<br />
Subphylum 1. Plicostoma<br />
Class 1. Euglenoidea<br />
Order 1. Petalomonadida [e.g. Calycimonas, Petalomonas]<br />
Order 2. Peranemida [e.g. Entosiphon, Peranema]<br />
Order 3. Rhabdomonadida [e.g. Distigma, Menoidium]<br />
Order 4. Euglenida [e.g. Astasia, Euglena, Eutreptia, Phacus]<br />
Class 2. Diplonemea<br />
Order 1. Diplonemida [e.g. Diplonema, Rhynchopus]<br />
Subphylum 2. Saccostoma<br />
Class 1. Kinetoplastea<br />
Order 1. Bodonida [e.g. Bodo, Cryptobia, Dimastigella, Ichthyobodo]<br />
Order 2. Trypanosomatida [e.g. Crithidia, Leishmannia, Trypanosoma]<br />
Class 2. Postgaardea<br />
Order 1. Postgaadida [e.g. Calkinsia, Postgaardi]<br />
Infrakingdom 2. Alveolata<br />
Phylum 1. Myzozoa<br />
Subphylum 1. Dinozoa<br />
Infraphylum 1. Protalveolata<br />
Class 1. Colponemea [e.g. Algovora, Colponema]<br />
Class 2. Myzomonadea [e.g. Alphamonas, Chilovora, Voromonas]<br />
Class 3. Perkinsea [e.g. Parvilucifera, Perkinsus, Phagodinium, Rastromonas]<br />
Class 4. Ellobiopsea [e.g. Elliobiopsis, Thalassomyces]<br />
Infraphylum 2. Din<strong>of</strong>lagellata<br />
Superclass 1. Syndina<br />
Class 1. Syndinea [e.g. Amoebophrya]<br />
Superclass 2. Dinokaryota<br />
Class 1. Noctilucea [e.g. Noctiluca]<br />
Class 2. Peridinea<br />
Subclass 1. Peridinoidia [e.g. Amylodinium, Heterocapsa, Prorocentrum]<br />
Subclass 2. Dinophysoidia [e.g. Dinophysis]<br />
Subclass 3. Gonyaulacoidia<br />
Order 1. Gonyaulacida [e.g. Ceratium, Cryptothecodinium]<br />
Subclass 4. Suessioidia<br />
Order 1. Suessiida [e.g. Polarella, Symbiodinium]<br />
Subclass 5. Oxyrrhia<br />
Order 1. Oxyrrhida [e.g. Oxyrrhis]<br />
Subphylum 2. Apicomplexa<br />
Infraphylum 1. Apicomonada<br />
Class 1. Apicomonadea [ e.g. Acrocoelus, Colpodella]<br />
Infraphylum 2. Sporozoa<br />
Class 1. Coccidea [e.g. Cryptosporidium, Hepatozoon, Toxoplasma]<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 77
Class 2. Gregarinea [e.g. Monocystis, Ophriocystis]<br />
Class 3. Haematozoa [e.g. Babesia, Plasmodium, Theileria]<br />
Phylum 2. Ciliophora<br />
Subphylum 1. Postciliodesmatophora<br />
Class 1. Karyorelictea [e.g. Kentrophoros, Loxodes, Tracheloraphis]<br />
Class 2. Heterotrichea [e.g. Blepharisma, Folliculina, Stentor]<br />
Subphylum 2. Intramacronucleata<br />
Infraphylum 1. Spirotrichia<br />
Class 1. Spirotrichea [e.g. Euplotes, Metopus, Oxytricha, Tintinnus]<br />
Infraphylum 2. Rhabdophora<br />
Class 1. Litostomatea [e.g. Didinium, Entodinium, Lacrymaria]<br />
Infraphylum 3. Ventrata<br />
Class 1. Phyllopharyngea [e.g. Dysteria, Podophrya]<br />
Class 2. Colpodea [e.g. Colpoda]<br />
Class 3. Nassophorea [e.g. Nassula]<br />
Class 4. Prostomatea [e.g. Coleps]<br />
Class 5. Plagiopylea<br />
Class 6. Oligohymenophorea [e.g. Paramecium, Tetrahymena, Vorticella]<br />
Kingdom 2. Animalia<br />
Subkingdom 1. Radiata<br />
Infrakingdom 1. Spongiaria<br />
Phylum 1. Porifera<br />
Subphylum 1. Hyalospongiae<br />
Subphylum 2. Calcispongiae<br />
Subphylum 3. Archaeocyatha<br />
Infrakingdom 2. Coelenterata<br />
Phylum 1. Cnidaria<br />
Subphylum 1. Anthozoa<br />
Subphylum 2. Medusozoa<br />
Phylum 2. Ctenophora<br />
Infrakingdom 3. Placozoa<br />
Phylum 1. Placozoa<br />
Subkingdom 2. Myxozoa<br />
Phylum 1. Myxosporidia<br />
Subkingdom 3. Bilateria<br />
Branch 1. PROTOSTOMIA<br />
Infrakingdom 1. Lophozoa<br />
SUPERPHYLUM POLYZOA<br />
Phylum 1. Bryozoa<br />
Subphylum 1. Stelmatopoda<br />
Subphylum 2. Lophopoda<br />
Phylum 2. Kamptozoa<br />
Subphylum 1. Entoprocta<br />
Subphylum 2. Cycliophora<br />
SUPERPHYLUM CONCHOZOA<br />
Phylum 1. Mollusca<br />
Subphylum 1. Bivalvia<br />
Subphylum 2. Glossophora<br />
Infraphylum 1. Univalvia<br />
Infraphylum 2. Spiculata<br />
Infraphylum 3. Cephalopoda<br />
Phylum 2. Brachiozoa<br />
Subphylum 1. Brachiopoda<br />
Subphylum 2. Phoronida<br />
SUPERPHYLUM 3. SIPUNCULA<br />
Phylum 1. Sipuncula<br />
SUPERPHYLUM 4. VERMIZOA<br />
Phylum 1. Annelida<br />
Subphylum 1. Polychaeta<br />
Subphylum 2. Clitellata<br />
Subphylum 3. Echiura<br />
Subphylum 4. Pogonophora<br />
78 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Phylum 2. Nemertina<br />
Infrakingdom 2. Chaetognathi<br />
Phylum 1. Chaetognatha<br />
Infrakingdom 3. Ecdysozoa<br />
SUPERPHYLUM 1. HAEMOPODA<br />
Phylum 1. Arthropoda<br />
Subphylum 1. Cheliceromorpha<br />
Infraphylum 1. Pycnogonida<br />
Infraphylum 2. Chelicerata<br />
Subphylum 2. Trilobitomorpha<br />
Subphylum 3. M<strong>and</strong>ibulata<br />
Infraphylum 1. Crustacea<br />
Infraphylum 2. Myriapoda<br />
Infraphylum 3. Insecta<br />
Phylum 2. Lobopoda<br />
Subphylum 1. Onychophora<br />
Subphylum 2. Tardigrada<br />
SUPERPHYLUM NEMATHELMINTHES<br />
Phylum Nemathelminthes<br />
Subphylum 1. Scalidorhyncha<br />
Infraphylum 1. Priapozoa<br />
Infraphylum 2. Kinorhyncha<br />
Subphylum 2. Nematoida<br />
Infraphylum 1. Nematoda<br />
Infraphylum 2. Nematomorpha<br />
Infrakingdom 4. Platyzoa<br />
Phylum 1. Acanthognatha<br />
Subphylum 1. Trochata (Gnathifera)<br />
Infraphylum 1. Rotifera<br />
Infraphylum 2. Acanthocephala<br />
Subphylum 2. Monokonta<br />
Phylum 2. Platyhelminthes<br />
Subphylum 1. Turbellaria<br />
Infraphylum 1. Mucorhabda<br />
Infraphylum 2. Rhabditophora<br />
Subphylum 2. Neodermata<br />
Infraphylum 1. Trematoda<br />
Infraphylum 2. Cercomeromorpha<br />
BRANCH 2. DEUTEROSTOMIA<br />
Infrakingdom 1. Coelomopora<br />
Phylum 1. Hemichordata<br />
Subphylum 1. Pterobranchia<br />
Subphylum 2. Enteropneusta<br />
Phylum 2. Echinodermata<br />
Subphylum 1. Homalozoa<br />
Subphylum 2. Pelmatozoa<br />
Infraphylum 1. Blastozoa<br />
Infraphylum 2. Crinozoa<br />
Subphylum 3. Eleutherozoa<br />
Infraphylum 1. Asterozoa<br />
Infraphylum 4. Echinozoa<br />
Infrakingdom 2. Chordonia<br />
Phylum 1. Urochorda<br />
Subphylum 1. Tunicata<br />
Infraphylum 1. Ascidiae<br />
Infraphylum 2. Thaliae<br />
Subphylum 2. Appendicularia<br />
Phylum 2. Chordata<br />
Subphylum 1. Acraniata<br />
Infraphylum 1. Cephalochordata<br />
Infraphylum 2. Conodonta<br />
Subphylum 2. Vertebrata<br />
Infraphylum 1. Agnatha<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 79
Infraphylum 2. Gnathostomata<br />
Subkingdom 4. Mesozoa<br />
Phylum 1. Mesozoa<br />
Kingdom 3. Fungi<br />
Subkingdom 1. Eomycota<br />
Phylum 1. Archemycota<br />
Subphylum 1. Dictyomycotina<br />
Class 1. Chytridiomycetes<br />
Class 2. Enteromycetes<br />
Subphylum 2. Melanomycotina<br />
Infraphylum 1. Allomycotina<br />
Class 1. Allomycetes<br />
Infraphylum 2. Zygomycotina<br />
Superclass 1. Eozygomycetia<br />
Class 1. Bolomycetes<br />
Class 2. Glomomycetes<br />
Superclass 2. Neozygomycetia<br />
Class 1. Zygomycetes<br />
Class 2. Zoomycetes<br />
Phylum Microsporidia<br />
Class 1. Minisporea<br />
Class 2. Microsporea<br />
Subkingdom 2. Neomycota<br />
Phylum 1. Ascomycota<br />
Subphylum 1. Hemiascomycotina<br />
Class 1. Taphrinomycetes<br />
Class 2. Geomycetes<br />
Class 3. Endomycetes<br />
Subphylum 2. Euascomycotina<br />
Class 1. Discomycetes<br />
Class 2. Pyrenomycetes<br />
Class 3. Loculomycetes<br />
Class 4. Plectomycetes<br />
Phylum 2. Basidiomycota<br />
Subphylum 1. Septomycotina<br />
Class 1. Septomycetes<br />
Subphylum 2. Orthomycotina<br />
Superclass 1. Hemibasidiomycetia<br />
Class 1. Ustomycetes<br />
Superclass 2. Hymenomycetia<br />
Class 1. Gelimycetes<br />
Class 2. Homobasidiomycetes<br />
Kingdom 4. Plantae<br />
Subkingdom 1. Biliphyta<br />
Infrakingdom 1. Glaucophyta<br />
Phylum 1.Glaucophyta [e.g. Cyanophora]<br />
Infrakingdom 2. Rhodophyta<br />
Phylum 1. Rhodophyta<br />
Subphylum 1. Rhodellophytina<br />
Class 1. Rhodellophyceae [e.g. Porphyridium]<br />
Subphylum 2. Macrorhodophytina<br />
Class 1. Bangiophyceae [e.g. Bangia, Porphyra]<br />
Class 2. Florideophyceae [e.g. Batrachospermum, Corallina]<br />
Subkingdom 2. Viridiplantae<br />
Infrakingdom 1. Chlorophyta<br />
Phylum 1. Chlorophyta<br />
Subphylum 1. Chlorophytina<br />
Infraphylum 1. Prasinophytae<br />
Class 1. Micromonadophyceae [e.g. Mesostigma, Micromonas]<br />
Class 2. Nephrophyceae [e.g. Nephroselmis, Pseudoscourfieldia]<br />
80 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Infraphylum 2. Tetraphytae<br />
Class 1. Chlorophyceae [e.g. Chlamydomonas, Tetraselmis]<br />
Class 2. Trebouxiophyceae [e.g. Chlorella]<br />
Class 3. Ulvophyceae [e.g. Acetabularia, Bryopsis, Codium, Ulva]<br />
Subphylum 2. Phragmophytina<br />
Infraphylum 1. Charophytae<br />
Class 1. Charophyceae [e.g. Chara, Nitella]<br />
Infraphylum 2. Rudophytae<br />
Class 1. Eophyceae [e.g. Coleochaete, Klebsormidium]<br />
Class 2. Conjugophyceae [e.g. Spirogyra]<br />
Infrakingdom 2. Cormophyta<br />
Phylum 1. Bryophyta<br />
Subphylum 1. Hepaticae<br />
Subphylum 2. Anthocerotae<br />
Subphylum 3. Musci<br />
Phylum 2. Tracheophyta<br />
Subphylum 1. Pteridophytina<br />
Infraphylum 1. Psilophytae<br />
Infraphylum 2. Lycophytae<br />
Infraphylum 3. Sphenophytae<br />
Infraphylum 4. Filices<br />
Subphylum 2. Spermatophytina<br />
Infraphylum 1. Gymnospermae<br />
Infraphylum 2. Angiospermae<br />
Kingdom 5. Chromista<br />
Subkingdom 1. Cryptista<br />
Phylum 1. Cryptista<br />
Subphylum 1. Cryptomonada<br />
Class 1. Cryptophyceae [e.g. Chilomonas, Cryptomonas, Guillardia]<br />
Class 2. Goniomonadea [e.g. Goniomonas]<br />
Subphylum 2. Leucocrypta<br />
Class 1. Leucocryptea [e.g. Kathablepharis, Leucocryptos]<br />
Subkingdom 2. Chromobiota<br />
Infrakingdom 1. Heterokonta<br />
Phylum 1. Ochrophyta<br />
Subphylum 1. Phaeista<br />
Infraphylum 1. Hypogyrista<br />
Class 1. Pelagophyceae [e.g. Pelagomonas, Sarcinochrysis]<br />
Class 2. Actinochrysea (Dictyochophyceae) [e.g. Dictyocha, Pedinella]<br />
Class 3. Pinguiophyceae [e.g. Glossomastix, Pinguiochrysis]<br />
Infraphylum 2. Chrysista<br />
Class 1. Raphidophyceae [e.g. Heterosigma]<br />
Class 2. Eustigmatophyceae [e.g. Vischeria]<br />
Class 3. Chrysophyceae [e.g. Ochromonas, Oikomonas, Spumella, Synura]<br />
Class 4. Chrysomerophyceae [e.g. Chrysomeris, Giraudyopsis]<br />
Class 5. Phaeothamniophyceae [e.g. Phaeothamnion, Pleurochloridella]<br />
Class 6. Xanthophyceae [e.g. Chloromeson, Vaucheria]<br />
Class 7. Phaeophyceae [e.g. Fucus, Laminaria]<br />
Subphylum 2. Khakista<br />
Class 1. Bolidophyceae [e.g. Bolidomonas]<br />
Class 2. Diatomeae [e.g. Coscinodiscus, Bacillaria, Nitzschia]<br />
Phylum 2. Bigyra<br />
Subphylum 1. Bigyromonada<br />
Class 1. Bigyromonadea [e.g. Developopayella]<br />
Subphylum 2. Pseud<strong>of</strong>ungi<br />
Class 1. Oomycetes [e.g. Achlya, Phytophthora]<br />
Class 2. Hyphochytrea [e.g. Rhizidiomyces]<br />
Subphylum 3. Opalinata<br />
Class 1. Proteromonadea [e.g. Proteromonas]<br />
Class 2. Blastocystea [e.g. Blastocystis]<br />
Class 3. Opalinea [e.g. Cepedea, Opalina]<br />
Phylum 3. Sagenista<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 81
Class 1. Labyrinthulea [e.g. Labyrinthula, Thraustochytrium]<br />
Class 2. Bisoecea [e.g. Bicosoeca, Caecitellus, Cafeteria]<br />
Class 3. Placididea [e.g. Pendulomonas, Placidia, Wobblia]<br />
Infrakingdom 2. Haptophyta<br />
Phylum 1. Haptophyta<br />
Class 1. Pavlovophyceae [e.g. Pavlova]<br />
Class 2. Prymnesiophyceae [e.g. Emiliania, Isochrysis, Prymnesium]<br />
Phylum 2. Heliozoa [e.g. Acanthocystis, Acanthophrys]<br />
82 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100640 (VWL9231): Waitata,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s 9 Feb 1948 - on<br />
C. maschalocarpum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Syncoryne reinkei R.Nielsen &<br />
P.M.Pedersen<br />
CHR401337: (slide CHR1203-1209)<br />
Kaikoura, 5 Sept 1979, O.Moestrup<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100639 (VWL9135): Pawa Bay,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 18 Jan 1948 -<br />
on C. maschalocarpum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100636 (VWL9048): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 1 Jan 1948 - on<br />
Carpophyllum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU 100641 (VWL9324): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 11 Feb 1948 -<br />
on C.plumosum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100642 (VWL9367): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 25 Feb 1948 -<br />
on C.plumosum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100643 (VWL9379): Temple Bar,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 26 Feb 1948 -<br />
on C.plumosum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. WELT A001108: Russell, Long<br />
Beach, 12 Jan 1948, VWLindauer<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis sp. AKU100646 (VWL9687): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 14 Mar 1948 -<br />
on Carpophyllum plumosum<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. WELT A017693: North east end,<br />
Heaphy Shoal, Chatham Isl<strong>and</strong>, 04<br />
Nov 1986, CH Hay<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. WELT A003919: Isl<strong>and</strong> Bay, 26 Aug<br />
1970, N.Adams - on Lessonia<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100647 (VWL9823): Temple Bar,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 24 Mar 1948 -<br />
on Pachymenia<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100644 (VWL9652): Long Beach<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 9 Mar 1948 - on<br />
Pachymenia<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100645 (VWL9671): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 13 Mar 1948 -<br />
on Pachymenia<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100648 (VWL10159): Long Beach,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 27 Apr 1948 -<br />
on Pachymenia<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100650 (VWL13298): Stormy Bay,<br />
Russell, Bay <strong>of</strong> Isl<strong>and</strong>s, 20 Jan 1953 -<br />
on Pachymenia<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Chlorophyta Chlorophyceae Chaetophorales Chaetophoraceae Sporocladopsis novae-zel<strong>and</strong>iae V.J.Chapm. AKU100651 (VWL13330): Kaikoura,<br />
U.V.Dellow, Dec 1949 - on Lessonia<br />
APPENDIX 2:
Chlorophyta Bryopsidophyceae Bryopsidales Ostreobiaceae Ostreobium quekettii Bornet & Flahault CHR401340: (slide) Kakanui, 18<br />
May 1980, O.Moestrup<br />
Chlorophyta Bryopsidophyceae Bryopsidales Ostreobiaceae Ostreobium quekettii Bornet & Flahault CHR401339: (slide) Portobello, 17<br />
May 1980, O.Moestrup<br />
Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Gomontia polyrhiza (Lagerh.) Bornet &<br />
Flahault<br />
Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Gomontia polyrhiza (Lagerh.) Bornet &<br />
Flahault<br />
Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Gomontia polyrhiza (Lagerh.) Bornet &<br />
Flahault<br />
Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Gomontia polyrhiza (Lagerh.) Bornet &<br />
Flahault<br />
Chlorophyta Chlorophyceae Phaeophilales Phaeophilaceae Phaeophila dendroides (P.Crouan & H.Crouan)<br />
Batters<br />
Chlorophyta Chlorophyceae Phaeophilales Phaeophilaceae Phaeophila dendroides (P.Crouan & H.Crouan)<br />
Batters<br />
Chlorophyta Chlorophyceae Phaeophilales Phaeophilaceae Phaeophila dendroides (P.Crouan & H.Crouan)<br />
Batters<br />
Chlorophyta Chlorophyceae Phaeophilales Phaeophilaceae Phaeophila dendroides (P.Crouan & H.Crouan)<br />
Batters<br />
Chlorophyta Ulvophyceae Ulvales Ulvellaceae Entocladia viridis Reinke VWL13256: Stewart Is., May 1950, - on<br />
Epymenia<br />
Chlorophyta ? Endoderma - (?) VWL no number: Harriet Kings,<br />
Corom<strong>and</strong>el, 5 Apr 1931 - on<br />
Pachymenia lusoria<br />
CHR401340: (slide) Kakanui, 18<br />
May 1980, O.Moestrup<br />
CHR401339: (slide) Portobello, 17<br />
May 1980, O.Moestrup<br />
CHR401338: (slide) Piha, 16 Apr<br />
1980, O.Moestrup<br />
CHR401337: (slide) Kaikoura, 5<br />
Sept 1979, O.Moestrup<br />
CHR401340: (slide) Kakanui, 18<br />
May 1980, O.Moestrup<br />
CHR401339: (slide) Portobello, 17<br />
May 1980, O.Moestrup<br />
CHR401338: (slide) Piha, 16 Apr<br />
1980, O.Moestrup<br />
CHR401337: (slide) Kaikoura, 5<br />
Sept 1979, O.Moestrup<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Chlorophyta Chlorophyceae Chlorococcales Endosphaeraceae Eugomontia stelligera R.Nielsen Syntype - CHR219311: Kakanui,<br />
South I., 18 May 1980, O. Moestrup -<br />
green empty shell from<br />
intertidalzone, grown in culture
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A13520: Port William, Stewart<br />
Is, 29 Jan 1983, W.A.Nelson - on<br />
Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A7450: RingaRinga, Stewart<br />
Is, 30 Nov 1959, E.A.Willa - on<br />
Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A1498: Gore Bay, South Is,<br />
Nov 1925, R.M.Laing - on<br />
Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A12982: Tautuku Peninsula,<br />
SE Otago, 7 Dec 1973, C.H.Hay - on<br />
Xiphophora gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A7449: Lonneker's Nugget,<br />
Stewart Is, 29 Jan 1960, E.A.Willa -<br />
on X. gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A6674: Lonneker's Nugget,<br />
Stewart Is., 2 Dec 1971, E.Conway &<br />
N.M.Adams - on Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh Manurewa Reef, Te Awhaite,<br />
Wairarapa, 4 Nov 1973, N.M.Adams -<br />
on Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A7945: Pukerua Bay,<br />
Wellington, 18 Nov 1972,<br />
N.M.Adams - on Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR315689: Cape Palliser, North I.,<br />
11 Nov 1962, M.J.Parsons - on<br />
Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A4052a+b: Karaka Bay,<br />
Wellington, Nov 1970, J.McCredie -<br />
on Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh WELT A6584: Cape Palliser, 7 Nov<br />
1971, N.M.Adams - on Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR38194: Whareponga, East<br />
Cape, North I., 12 Dec 1942,<br />
L.B.Moore - on C. torulosa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR385155: Titahi Bay, Wellington,<br />
21 Nov 1942, L.B.Moore - on C.<br />
retr<strong>of</strong>lexa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR219394: Lonneker's Bay,<br />
Stewart I., 2 Dec 1971, M.J.Parsons<br />
- on C. retr<strong>of</strong>lexa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR354184: Big Sol<strong>and</strong>er I., 17<br />
Nov 1973, P.N.Johnson - on<br />
Cystophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh CHR230929: Akitio, North I., 2 Jan<br />
1972, M.J.Parsons - on C. scalaris<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Corynophlaea cystophorae J.Agardh AK22498 (=ANZE185): Pihama, CHR385154 (=ANZE185): Pihama, WELT A985 (=ANZE185): Pihama,<br />
Taranaki, North I., 2 Dec 1944, Taranaki, North I., 2 Dec 1944, Taranaki, North I., 2 Dec 1944,<br />
V.W.Lindauer - on C. retr<strong>of</strong>lexa V.W.Lindauer - on C. retr<strong>of</strong>lexa V.W.Lindauer - on C. retr<strong>of</strong>lexa
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing AK22516: (= ANZE203): Pegasus Bay,<br />
Stewart I., 6 Oct 1945, V.W.Lindauer -<br />
on Xiphophora gladiata<br />
CHR62508 (= ANZE203): Pegasus<br />
Bay, Stewart I., 6 Oct 1945,<br />
V.W.Lindauer - on Xiphophora<br />
gladiata<br />
WELT A1003 (= ANZE203): Pegasus<br />
Bay, Stewart I., 6 Oct 1945,<br />
V.W.Lindauer - on Xiphophora<br />
gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR62503 (=VWL6698): Stewart I.,<br />
22 Oct 1945, E.Willa - on<br />
Xiphophora gladiata<br />
Herponema maculaeforme (J.Agardh) Laing AK146242: Dunedin, S.Berggren - on<br />
Xiphophora ISOTYPE<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. CHR230702: Shag Pt, Otago, South<br />
I., 9 Sept 1971, M.J.Parsons - on<br />
Hormosira<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. CHR219443: Lonneker's Nugget,<br />
Stewart I., 3 Dec 1971, M.J.Parsons<br />
- on Hormosira<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. WELT A6669: Lonneler's Nugget,<br />
Stewart Is, 3 Dec 1971, E.Conway &<br />
N.M.Adams<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae Lindauer & V.J.Chapm. WELT A7445: Lonneker's Nugget,<br />
Stewart Is, 6 Feb 1963, E.Awilla<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR219445: Lonneker's Nugget,<br />
Stewart I., 5 Dec 1971, M.J.Parsons<br />
- on Xiphophora gladiata<br />
J.Ag.)<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema hormosirae (as<br />
pulvinatum<br />
(Harv.MS) non<br />
AK30304: ANZE334: Waitangi, Bay <strong>of</strong><br />
Isdl<strong>and</strong>s, North I., 19 Aug 1950,<br />
VWLindauer - on Hormosira SYNTYPE<br />
CHR227375 (=ANZE334): Waitangi, WELT A1134 (=ANZE334): Waitangi,<br />
Bay <strong>of</strong> Isl<strong>and</strong>s, North I., 19 Aug Bay <strong>of</strong> Isl<strong>and</strong>s, North I., 19 Aug 1950,<br />
1950, V.W.Lindauer - on Hormosira V.W.Lindauer - on Hormosira<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Hecatonema stewartensis V.J.Chapm. AK295761: Chris's Bay, Pegasus,<br />
Stewart Is, 10 Apr 1948 ex VWL10256<br />
TYPE<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A15913: L<strong>and</strong>ing Bay,<br />
Burgess I, Mokohinau Is, 31 Dec<br />
1984, M.Francis - on X.chondrophylla<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A13850: Cable Bay,<br />
Doubtless Bay, 27 Oct. 1982,<br />
W.A.Nelson - on X.chondrophylla<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A13446: Tapeka Point, Bay <strong>of</strong><br />
Isl<strong>and</strong>s, 30 Oct 1982, W.A.Nelson -<br />
on X.chondrophylla<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A18903: Katherine Bay, Great<br />
Barrier I, 7 Dec 1989, F.I.Dromgoole -<br />
on X.chondrophylla<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A2509: The Pinnacles, Little<br />
Barrier I., no date, U.V.Dellow - on<br />
X.chondrophylla<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Elachista australis J.Agardh WELT A6569: Cape Palliser,<br />
Wairarapa, 7 Nov 1971, N.M.Adams -<br />
on X.gladiata
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A26060: Doubtful Sound,<br />
Fiordl<strong>and</strong>, 21 Jan 2000, C.Duffy<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. CHR63461: Kaikoura, South I., 14<br />
Nov 1948, L.B.Moore<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A25591" Bradshaw Sound,<br />
Fiordl<strong>and</strong>, 3 Oct 2000, K.Neill<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema sp. WELT A13952: Northeast I, Three<br />
Kings Is, 25 Nov 1983, M.Francis -<br />
on Sargassum johnsonii<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Mikrosyphar pachymeniae Lindauer Isotype - CHR68937: Russell, Bay<br />
<strong>of</strong> Isl<strong>and</strong>s, North I., 1 Apr 1944,<br />
V.W.Lindauer (4267)<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. AK22467: ANZE184 On Ulva lactuca ,<br />
Kaikoura, 31 Dec 1944<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. CHR219406: Lonneker's Bay,<br />
Stewart I., 2 Dec 1971, M.J.Parsons<br />
Xiphophora<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae as "Hecatonema?" in<br />
ANZE<br />
AK22532: (=ANZE229) Stewart Is, 14<br />
Jun 1945, V.W.Lindauer - on<br />
WELT A1029: (=ANZE229) Stewart<br />
Is, 14 Jun 1945, V.W.Lindauer<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A13525: Port William, Stewart<br />
Is, 29 Jan 1983, W.A.Nelson - on<br />
X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A7446: Ringaringa, Stewart<br />
Is, 27 Mar 1963, E.A.Willa - on<br />
X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A6576: Cape Palliser,<br />
Wairarapa, 7 Nov 1971, N.M.Adams -<br />
on X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A7784: Ranui Cove, Auckl<strong>and</strong><br />
Is, 30 Nov 1972, A.N.Baker - on<br />
X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A18813a+b: Cape Young,<br />
Chatham I, 6 Mar 1987, W.A.Nelson -<br />
on X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing WELT A18261: Long I, Dusky<br />
Sound, Fiordl<strong>and</strong>, 14 May 1986,<br />
L.A.Bolton - on X.gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR46581: Auckl<strong>and</strong> Is., 26 Dec<br />
1943, W.Dawbin - on Xiphophora<br />
gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR67782: Pencarrow Head, Cook<br />
Strait, North I., 14 Jan 1950,<br />
L.B.Moore - on Xiphophora gladiata<br />
(drift)<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR52070: Waitangi, Chatham Is.,<br />
12 July 1945, A.M.Rapson - on<br />
Xiiphophora gladiata<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Herponema maculaeforme (J.Agardh) Laing CHR230701: Shag Pt, Otago, South<br />
I,, 8 Oct 1971, M.J.Parsons - on<br />
Xiphophora gladiata
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Pilinia rimosa Kuetz. WELT A022656: Piha, west<br />
Auckl<strong>and</strong>, 03 Jul 1994, E.Henry<br />
Heterokontophyta Phaeophyceae Sphacelariales Sphacelariaceae Sphacelaria pulvinata Hook.f. & Harv. WELT A4369: Wharariki Beach, NW<br />
Nelson, 19 Mar 1971, F.M.Climo - on<br />
Carpophyllum maschalocarpum<br />
Heterokontophyta Phaeophyceae Sphacelariales Sphacelariaceae Sphacelaria pulvinata Hook.f. & Harv. AK146468: ANZE131, on Carpophyllum<br />
maschalocarpum, Mangonui, 24 Oct<br />
1942<br />
WELT A931: (=ANZE131) Mangonui,<br />
Northl<strong>and</strong>, 24 Oct 1942,<br />
V.W.Lindauer - on Carpophyllum<br />
maschalocarpum<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A12907: Katiki, North Otago,<br />
4 Jun 1973, C.H.Hay<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A6331: Lonneker's Nugget,<br />
Stewart Is, 26 May 1971, E.Conway<br />
& N.M.Adams<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A3644a+b: Makara,<br />
Wellington, 2 Jun 1970, N.M.Adams -<br />
on drift<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South WELT A7447: RiongaRinga, Stewart<br />
Is, 18 Mar 1960, E.A.Willa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema sp. WELT A8619: Tasman Bay, Three<br />
Kings Is, Feb 1974, A.N.Baker - on<br />
L<strong>and</strong>sburgia quercifolia<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Nemacystus novae-zel<strong>and</strong>iae Kylin WELT A18016: Parnell Reef,<br />
Waitemata Harbour, Auckl<strong>and</strong>, 13<br />
Oct 1987, K.W.Glombitza - on<br />
Sargassum scabridum<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK295758: (VWL6253) Stewart Is, 12<br />
Jun 1945, E.Willa - ISOTYPE<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK22533: (ANZE230) (VWL6253),<br />
Stewart Is, 12 Jun 1945, E.Willa<br />
ISOTYPE<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South AK295759: (VWL6253) Stewart Is, 12<br />
Jun 1945, E.Willa - TYPE<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South CHR49999: Princess Bay Bay,<br />
Wellington, North I., 30 May 1943,<br />
L.B.Moore (drift)<br />
Heterokontophyta Phaeophyceae Ectocarpales incertae sedis Herpodiscus durvillaeae (Lindauer) South CHR248256: Oaro, Kaikoura, South<br />
I., 4 July 1973, M.J.Parsons<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A16379: Perserverance<br />
Harbour, Campbell I, 12 Feb 1985,<br />
J.C.Yaldwyn<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A984: ANZE 184: Kaikoura,<br />
31 Dec 1944, V.W.Lindauer<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A18688: Monau, Chatham I, 3<br />
Mar 1987, W.A.Nelson<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A1592a+b: York Bay,<br />
Wellington, 31 May 1953, R.K.Dell<br />
Heterokontophyta Phaeophyceae Ectocarpales Chordariaceae Myrionema strangulans Grev. WELT A7073: Rosa I, Port Pegasus,<br />
Stewart Is, 29 Feb 1972, N.M.Adams
galls on Durvillaea CHR243660: Kaitangata, Otago,<br />
South I., 26 Feb 1973, R.Mason &<br />
E.M.Chapman - on Durvillaea<br />
antarctica drift -"galls not caused by<br />
fungus - possibly bacteria" det.<br />
J.Kohlmeyer<br />
Division Class Order Family Genus Species Authority AK CHR Te Papa<br />
galls on Macrocystis CHR47805: Native Isl<strong>and</strong>, Paterson<br />
Inlet, Stewart Isl<strong>and</strong>, 2 Dec 1944,<br />
L.B.Moore - on Macrocystis pyrifera -<br />
"not fungal but possibly caused by<br />
filamentous brown algae (see<br />
Andrews 1976 Biol. Rev. 51: 211-<br />
253, Can J. Bot 55:1019-1027)" det<br />
J.Kohlmeyer
Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra woolhousiae Harv. CHR 209060: Lyall Bay, Wellington,<br />
Sept 1931, Scarfe<br />
Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra woolhousiae Harv. CHR55566: Hokio Beach, Levin,<br />
Nov 1946, Moore.<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. AK147200: Bluff, 1874, Berggren<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. AK147201: Ringaringa, Stewart Is, 15<br />
Jan 1940, L.M.Cranwell<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. AK223832: Preservation Inlet, Fiordl<strong>and</strong>,<br />
20 Jul 1995, M.S.Morley<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. AK22605: (=ANZE214) Stewart Is, 15<br />
Jan 1946, V.W.Lindauer<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR24001: Bluff, South I., 4 Jan<br />
1940, L.B.Moore<br />
Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra adamsiae W.A.Nelson WELT A10322: Crater Bay,<br />
Antipodes Is, 23 Nov 1978, C.H.Hay<br />
Rhodophyta Bangiophyceae Bangiales Bangiaceae Porphyra adamsiae W.A.Nelson WELT A8038: Port Ross, Auckl<strong>and</strong><br />
Is, 19 Feb 1973, K.Johnson,<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon subtumens (J.Agardh ex Laing)<br />
W.A.Nelson<br />
WELT A15999: Brighton, Otago, 2<br />
Feb 1983, W.A.Nelson - on<br />
D.antarctica<br />
D.chathamensis<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon subtumens (J.Agardh ex Laing)<br />
W.A.Nelson<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon cameronii (W.A.Nelson)<br />
W.A.Nelson<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon cameronii (W.A.Nelson)<br />
W.A.Nelson<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Pyrophyllon subtumens (J.Agardh ex Laing)<br />
W.A.Nelson<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Chlidophyllon kaspar (W.A.Nelson et N.M.<br />
Adams) W.A.Nelson<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Chlidophyllon kaspar (W.A.Nelson et N.M.<br />
Adams) W.A.Nelson<br />
WELT A26849: Three Kings Is, Jan<br />
1994, V.Staines<br />
WELT A16714: Princes Rocks, Three<br />
Kings Is, 18 Jan 1985, M.Francis &<br />
M.A.Williams<br />
WELT A26851: Wharekauri,<br />
Chatham I, Feb 2001, R.Russell<br />
WELT A17785: Heaphy Shoal,<br />
Chatham I, 4 Nov 1986, C.H.Hay<br />
WELT A3669: Makara, Wellington, 2<br />
Jun 1970, N.M.Adams - on drift<br />
D.antarctica<br />
WELT A7466a+b: Point Webb,<br />
Chatham I, 4 Nov 1986, C.H.Hay - on<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Chroodactylon ornatum (C.Agardh) Basson AK30298: (=ANZE 341) Glendowie,<br />
WELT A1141: (=ANZE 341)<br />
Auckl<strong>and</strong>, 20 Dec 1949, V.W.Lindauer<br />
Glendowie, Auckl<strong>and</strong>, 20 Dec 1949,<br />
V.W.Lindauer<br />
Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Chroodactylon ornatum (C.Agardh) Basson WELT A6707: Oban, Stewart I, 3 Dec<br />
1971, E.Conway & N.M.Adams -<br />
epiphyte<br />
Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Stylonema alsidii (Zanardini) K.M.Drew WELT A4403: Days Bay, Wellington,<br />
13 Jun 1971, N.M.Adams - on<br />
Chaetomorpha<br />
Rhodophyta Rhodellophyceae Stylonematales Stylonemataceae Erythrocladia sp. WELT A18578: Durham & Gap Pts,<br />
Chatham I, 4 Mar 1987, W.A.Nelson -<br />
on Cladophora sp.<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Erythrotrichia foliiformis South et N.M.Adams WELT A17692: Petre Bay, Chatham<br />
I, 4 Nov 1986, C.H.Hay - on Lessonia<br />
tholiformis<br />
Rhodophyta Compsopogonophyceae Erythropeltidiales Erythrotrichiaceae Erythrotrichia foliiformis South et N.M.Adams WELT A6570: Cape Palliser,<br />
Wairarapa, 7 Nov 1971, N.M.Adams -<br />
on Marginariella (oval)
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. WELT A3988: Owhiro Bay,<br />
Wellington, 19 Sep 1970, N.M.Adams<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. CHR357193: Rimu Bay, Pelorus<br />
Sound, South I., 5 Oct 1958,<br />
L.B.Moore<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. CHR49433: Matarangi Beach,<br />
Kuaotunu, Corom<strong>and</strong>el, North I., 29<br />
Mar 1945, N.M.Adams<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. CHR248233: Leigh Marine Station,<br />
North I., 22 May 1974, M.J.Parsons<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. AK290043: Waikawau Bay, Corom<strong>and</strong>el,<br />
7 Oct 2004, M.N.Lee<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. AK239454: Henderson Pt, Northl<strong>and</strong>, 1<br />
Jul 1999, E.K.Cameron<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. CHR191680: Whangamumu<br />
Harbour, North I., 26 May 1969,<br />
E.Godley<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. AK147207: (=ANZE46) Bay <strong>of</strong> Isl<strong>and</strong>s,<br />
14 Aug 1938, V.W.Lindauer<br />
WELT A846: (=ANZE46) Bay <strong>of</strong><br />
Isl<strong>and</strong>s, 14 Aug 1938, V.W.Lindauer<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. WELT A16131: Senecio Pool, Snares<br />
I, 18 Dec 1984, G.S.Hardy<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. WELT A25595, Deas Cove, 3 Oct<br />
2000, A.Loughnan<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. WELT A21795: Cascade I, South<br />
Westl<strong>and</strong>, 21 Feb 1996, D.Neale<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. WELT A4010: Brighton, Otago, 5 Dec<br />
1970, N.M.Adams<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. WELT A26159: Port Hutt, Chatham<br />
Is, 12 Mar 2001, W.Nelson, J.Broom,<br />
W.Jones, T. Farr<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR354186: Big Sol<strong>and</strong>er I., 17 Nov<br />
1973, P.N.Johnson<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR324992: Mangere I., Chatham<br />
Is, 21 Aug 1968, I. & M.Ritchie<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR509075: Ackers Pt, Stewart I., 4<br />
Jan 1987, D.R.Given<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR316773: Western Chain,<br />
Snares Is, 26 Nov 1974,<br />
D.S.Horning<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR57281: Tautuku, Otago, South<br />
I., Dec 1947, I.Coulter<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR368067: Secretary I., Doubtful<br />
Sound, Fiordl<strong>and</strong>, South I., 18 May<br />
1981, D.J.Brasch<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR205876: Blackhead, Dunedin,<br />
Otago, Dec 1919, W.A.Scarfe<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea lyallii Hook.f. et Harv. CHR379617: Bruce Rocks, Brighton,<br />
Otago, South I., Feb 1948,<br />
K.W.Allison
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin CHR319934: Atia Point, Kaikoura,<br />
South I., 14 Nov 1973, M.J.Parsons -<br />
on L. hookeriana<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR367973: South Bay, Kaikoura,<br />
South I., 3 Dec 1980, G.D.Fenwick -<br />
on Pterosiphonia<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR368028: Shap Point, Otago,<br />
South I., 10 Feb 1981, M.J.Parsons<br />
& M.Stolp - on Pterosiphonia<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. WELT A17700: McClatchie Reef,<br />
Chtaham I, 4 Nov 1986, C.H.Hay - on<br />
Chondria<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Levringiella CHR368051: Macrocarpa Point,<br />
Katiki Beach, Otago, South I., 9 Feb<br />
1981, M.J.Parsons - on<br />
Pterosiphonia, drift<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. WELT A17498a+b: Port Webb,<br />
Chatham I, 6 Nov 1986, C.H.Hay - on<br />
Chondria<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. CHR230816: Oaro, Kaikoura, South<br />
I., 6 Nov 1971, M.J.Parsons - on<br />
Chondria<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Janczewskia sp. CHR360270: Pier Wharf, Kaikoura,<br />
South I., 10 Sept 1974, M.J.Parsons -<br />
on Chondria<br />
Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248307: Oamaru, South I., 31<br />
Oct 1972, M.J.Parsons - on<br />
Heterosiphonia concinna<br />
Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248099: Penguin Bay,<br />
Campbell I., 18 Feb 1971,<br />
C.D.Meurk - on Heterosiphonia<br />
concinna<br />
Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR316964: Cod Cavern Gutway,<br />
Snares Is, 24 Jan 1975, D.S.Horning<br />
- on Heterosiphonia concinna<br />
Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya sp. CHR248303: Shag Point, Otago,<br />
South I., 1 Nov 1972, M.J.Parsons -<br />
on Heterosiphonia concinna<br />
Rhodophyta Florideophyceae Ceramiales Dasyaceae Colacodasya inconspicua (Reinsch) Schmitz CHR66045: French I., Auckl<strong>and</strong> Is,<br />
16 Aug 1976, C.A.Fleming - on<br />
Heterosiphonia berkeleyi (slide only<br />
No.66)<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
Rhodophyta Florideophyceae Hildenbr<strong>and</strong>iales Hildenbr<strong>and</strong>iaceae Apophlaea sinclairii Hook.f. et Harv. WELT A13938: West I, Three Kings<br />
Is, 25 Nov 1983, M.Francis<br />
Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT A027067: Wairarapa east<br />
coast, Mataikona reef, Feb 1969,<br />
N.M.Adams<br />
Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT AA027066: Cape Palliser, Nov<br />
1971, N.M.Adams<br />
Rhodophyta Florideophyceae Corallinales Corallinaceae Choreonema thuretii (Bornet) F.Schmitz WELT A027038: Kaikoura, barbeque<br />
area just south <strong>of</strong> Rakautara, Sept<br />
2004, Nelson, Farr & Neill
Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zel<strong>and</strong>iae Sparling CHR64545: Eastbourne, Wellington,<br />
20 Mar 1949, L.B.Moore,<br />
N.M.Adams & G.F.Papenfuss<br />
(NB:"Type collection by CHR64545<br />
not seen by author <strong>of</strong> species -<br />
Sparling 1979) - on Gloioderma<br />
saccatum<br />
Rhodophyta Florideophyceae Rhodymeniales Champiaceae Champiocolax sp. WELT A18631a+b: Inner Chetwode<br />
Is, Marlborough, 11 Aug 1987,<br />
C.H.Hay - on C. chathamensis<br />
Rhodophyta Florideophyceae Gracilariales Pterocladiophyllaceae Pterocladiophila hemisphaerica K.C.Fan et Papenf. CHR117794: locality unknown - from<br />
commercial collection, identity<br />
confirmed by K.C.Fan (UC<br />
Berkeley); on Pterocladiella<br />
capillacea<br />
Rhodophyta Florideophyceae Gigartinales Kallymeniaceae Callocolax sp. CHR367972: South Bay, Kaikoura,<br />
South I., 3 Dec 1980, G.D.Fenwick -<br />
on Echinothamnion sp.<br />
Rhodophyta Florideophyceae Gigartinales Kallymeniaceae Callocolax neglectus Schmitz ex Batters CHR248213: Oaro, Kaikpoura,<br />
South I., 25 Oct 1972, M.J.Parsons -<br />
on Callophyllis calliblepharoides<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Tylocolax<br />
microcarpus ?<br />
CHR219462: Lonneker's Nugget,<br />
Stewart I., 3 Dec 1971, M.J.Parsons -<br />
on Adamsiella chauvinii<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Tylocolax<br />
microcarpus ?<br />
CHR368033: Shag Point, Otago,<br />
South I., 10 Feb 1981, M.J.Parsons<br />
& M.Stolp - on Adamsiella chauvinii<br />
chauvinii<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Tylocolax<br />
microcarpus ?<br />
CHR364690: Baxters Reef,<br />
Kaikoura, South I., 5 Feb 1980,<br />
G.D.Fenwick - on Adamsiella<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR319388: Curio Bay, SE Otago,<br />
South I., 16 Feb 1977, M.J.Parsons -<br />
on Echinothamnion (liq coll)<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR367972: South Bay, Kaikoura,<br />
South I., 3 Dec 1980, G.D.Fenwick -<br />
on Echinothamnion sp.<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin WELT A18232: George Sound,<br />
Fiordl<strong>and</strong>, 14 Feb 1987, M.Francis -<br />
on L. hookeriana<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum sp. CHR399500: Katiki Beach, Otago,<br />
South I., 9 Feb 1981, M.J.Parsons -<br />
on Polysiphonia rhododactyla (liq<br />
coll)<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
Rhodophyta Florideophyceae Ceramiales Rhodomelaceae Sporoglossum lophurellae Kylin CHR319947: Lighthouse Reef,<br />
Kaikoura, South I., 13 Nov 1973,<br />
M.J.Parsons - on L. hookeriana
parasite on<br />
Rhodophyllis<br />
CHR364774:Old Wharf, Kaikoura,<br />
South I., 4 Feb 1980, G.D.Fenwick -<br />
on Rhodophyllis (liq coll.)<br />
parasite on<br />
Rhodophyllis<br />
CHR364762: South Bay, Kaikoura,<br />
South I., 4 Feb 1980, G.D.Fenwick -<br />
on Rhodophyllis (liq coll.)<br />
sanguinea<br />
G.D.Fenwick (liq coll.)<br />
parasite on<br />
Hymenocladia<br />
CHR364678: Baxters Reef,<br />
Kaikoura, South I., 5 Feb 1980,<br />
sanguinea<br />
parasite on<br />
Hymenocladia<br />
CHR219369: Ringaringa, Stewart I.,<br />
30 Nov 1971, M.J.Parsons<br />
parasite on<br />
Dasyclonium<br />
CHR319896: Seal Reef, Kaikoura,<br />
South I., 6 Oct 1971, M.J.Parsons -<br />
on Dasyclonium incisum<br />
oblongifolia<br />
parasite on<br />
Cladhymenia<br />
oblongifolia<br />
CHR319472: Old Wharf, Kaikoura,<br />
South I., 13 Nov 1973, V.Hoggard &<br />
G.D.Fenwick - on Cladhymenia<br />
on A. lyallii<br />
parasite on<br />
Apophlaea lyallii<br />
CHR219466: Lonneker's Nugget,<br />
Stewart I., 2 Dec 1971, M.J.Parsons -<br />
Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax WELT A026739: Te Werahi Beach,<br />
Northl<strong>and</strong>, North I., 25 Oct 2003,<br />
W.Nelson<br />
Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR364760: South Bay, Kaikoura,<br />
South I., 4 Feb 1980, G.D.Fenwick -<br />
on Plocamium 2x2 fine<br />
Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR367983: South Bay, Kaikoura,<br />
South I., 3 Dec 1980, G.D.Fenwick -<br />
on Plocamium 2 x 2 fine<br />
Rhodophyta Florideophyceae Plocamiales Plocamiaceae Plocamiocolax CHR360413: Katiki Beach, Otago,<br />
South I., 27 Apr 1975, M.J.Parsons -<br />
on Plocamium 2x2<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zel<strong>and</strong>iae Sparling WELT A26638: Wharariki Beach, 19<br />
Mar 2003, W.Nelson & J.Dalen - on<br />
Gloioderma saccata<br />
Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zel<strong>and</strong>iae Sparling WELT A17670: Okawa Beach,<br />
Chatham I, 6 Jan 1987, A.N.Baker -<br />
on Gloioderma saccata<br />
Rhodophyta Florideophyceae Rhodymeniales Faucheaeceae Gloiocolax novae-zel<strong>and</strong>iae Sparling WELT A6599: Ringaringa, Stewart I,<br />
26 Apr 1963, E.A.Willa<br />
Rhodophyta Florideophyceae Rhodymeniales Rhodymeniaceae Rhodymeniocolax sp. WELT A14130: Antipodes I, 4 Dec<br />
1978, C.H.Hay - on Rhodymenia<br />
epimenioides<br />
Rhodophyta Florideophyceae Rhodymeniales Rhodymeniaceae Rhodymeniocolax sp. WELT A7568: Golden Bay, Paterson<br />
Inlet, Stewart I, 29 Feb 1960,<br />
E.A.Willa - on Rhodymenia linearis
parasite on<br />
Rhodophyllis<br />
WELT A4167: West Lyall Bay,<br />
Wellington, 12 Jan 1971, N.M.Adams -<br />
"cf Ceratocolax"<br />
Phylum Class Order Family Genus Species Authority AK CHR Te Papa<br />
parasite on<br />
WELT A6798: Harrold's Bay,<br />
Rhodophyllis<br />
Halfmoon Bay, Stewart Is, 1 Dec<br />
1971, N.M.Adams - "cf Ceratocolax"
galls on Chaetangium "pycnidia - unfortunately cannot CHR248185a: Monument Harbour, Campbell I., 14 Feb<br />
be further identified as long as 1971, C.D.Meurk - on Chaetangium fastigiatum<br />
perfect (ascigerous) state is<br />
unknown. Many marine<br />
algicolous Ascomyctes have<br />
similar pycnidia." det J.Kohlmeyer<br />
Chaudefaudia corallinarum (Crouan et Crouan) Muller et<br />
v.Arx<br />
Chaudefaudia corallinarum (Crouan et Crouan) Muller et<br />
v.Arx<br />
det Kohlmeyer CHR248265: Mollymawk Bay, Snares Is, 6 Dec 1974,<br />
D.S.Horning - on Euptilota formosissma<br />
det Kohlmeyer CHR248266: Cod Cavern Gutway, Snares Is, 24 Jan<br />
1975, D.S.Horning - on Euptilota formosissima<br />
Eurychasma dicksonii (Wright) Magnus Saprolegniales - "forms peculiar<br />
"netsporangia" with encysted<br />
zoospores" det Kohlmeyer<br />
CHR248343: Shag Point, Otago, South I., 8 Sept 1971,<br />
M.J.Parsons (liq coll + photomicrograph) - on<br />
Ectocarpus on Scytosiphon (CHR219500)<br />
Spathulospora lanata Kohlmeyer CHR:64534: Runaround, Wellington, North I., 18 Mar<br />
1949, N.M.Adams - on Ballia scoparia - det Kohlmeyer<br />
CHR357143: Open Bay Isl<strong>and</strong>s, Westl<strong>and</strong>, South I., 4<br />
Feb 1976, G.D.Fenwick - on Sargassum undulatum (det<br />
Kohlmeyer)<br />
CHR315947c: Houghton Bay, 16 Oct 1962, M.J.Parsons<br />
- on Sargassum sinclairii (det Kohlmeyer)<br />
Haloguignardia tumefaciens (Cribb et Herbert) Cribb et<br />
Cribb<br />
Haloguignardia tumefaciens (Cribb et Herbert) Cribb et<br />
Cribb<br />
Genus species Authority comments CHR<br />
Mycosphaerella apophlaeae Kohlm. Bot Mar 24: 13- Kohlmeyer & CHR391939: South Promontory, Snares Is, 14 Dec<br />
Demoulin 1981<br />
1974, C.E.Holmes<br />
Polystigma apophlaeae Kohlm. Bot Mar 24: 13- Kohlmeyer & Herb - Holotype NY<br />
Demoulin 1981
APPENDIX 5:<br />
Data storage<br />
Dataset supplied to the Ministry in the form <strong>of</strong> an Access database <strong>and</strong> an electronic copy <strong>of</strong><br />
the report.<br />
Utility <strong>of</strong> the Access database<br />
The database was operated at NIWA through a Delphi web application, enabling multiple<br />
users. Below are examples <strong>of</strong> the web interface pages we used, configured for data entry.<br />
Search functions will need to be developed as part <strong>of</strong> the front end <strong>of</strong> this database.<br />
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 99
100 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 101
102 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>
MAF Biosecurity New Zeal<strong>and</strong> <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> • 103
104 • <strong>Diseases</strong>, <strong>pathogens</strong> <strong>and</strong> <strong>parasites</strong> <strong>of</strong> <strong>Undaria</strong> <strong>pinnatifida</strong> MAF Biosecurity New Zeal<strong>and</strong>