00288330%2E1987%2E9516214-11

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A survey of filamentous algalproliferations in New Zealand riversBarry J. F. Biggs a & Geoff M. Price aa Hydrology Centre , Ministry of Works and Development , P. O.Box 1479, Christchurch, New ZealandPublished online: 29 Mar 2010.

To cite this article: Barry J. F. Biggs & Geoff M. Price (1987) A survey of filamentous algalproliferations in New Zealand rivers, New Zealand Journal of Marine and Freshwater Research, 21:2,175-191, DOI: 10.1080/00288330.1987.9516214

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New Zealand Journal of Marine and Freshwater Research, 1987, Vol. 2 1 : 175-1910028-8330/87/2102-0175$2.50/0 Crown copyright 1987

175

A survey of filamentous algal proliferationsin New Zealand rivers

BARRY J. F. BIGGSGEOFF M. PRICEHydrology CentreMinistry of Works and DevelopmentP. O. Box 1479, ChristchurchNew Zealand

Abstract New Zealand rivers were surveyed forfilamentous algal proliferations following pro-longed periods of low flows to define their extent,standing crop, and composition. Significant growthswere found at 167 of the 423 sites surveyed. Thegeometric mean river bed cover of filamentous algaewas 26% in summer and 22% in winter. Prolifer-ations of >40% cover occurred at 16% of the sites.The geometric mean standing crop was 15.3 g/m2(ash-free dry weight) in summer and 1.87 g/m2 inwinter. Proliferations of >50 g/m2 (AFDW)occurred at 8% of the sites. A taxonomic classifi-cation analysis of the summer data identified ninemain clusters of sites which appeared to corre-spond to different catchment land uses. One, or acombination, of the following taxa dominated theseclusters: Ulothrix zonata, Phormidium spp., Sti-geoclonium sp., Spirogyra spp., Compsopogon coe-ruleus, Oedogonium spp., Cladophora sp.,Cladophora glomerata, Rhizoclonium sp., andMelosira varians. Ulothrix zonata and Diatomahiemale var. mesodon dominated the sites sampledin winter. There was a close positive relationshipbetween the average standing crop and the averagewater conductivity of the nine site clusters.

Keywords freshwater ecology; ecological asso-ciations; low flows; periphyton; algae; enrichment;eutrophication; trophic; biomass; water quality

INTRODUCTIONProliferations of filamentous algae are oftenobserved in New Zealand rivers during summer.Problems caused by these growths include cloggingof abstraction structures, degradation of waterquality (through diel fluctuations in dissolved oxy-gen and pH), and degradation of aesthetic values(Biggs 1985a).

Little is known of the composition or ecology ofthese proliferations in New Zealand. Only limitedobservations have been reported as part of inver-tebrate studies (e.g., Hirsch 1958; Winterbourn etal. 1971), or as part of studies on the effects of thealgae on river water quality (Biggs 1982; Freeman& McFarlane 1982). Recently, more detailed stud-ies have been carried out on nutrient limitation ofCladophora glomerata in the Manawatu River(Freeman 1986).

Most European and North American studies onriver enrichment have focused on the ecology ofCladophora glomerata, a major problem taxon inthese areas (the early studies have been reviewedby Whitton (1970)), or on enrichment processes inexperimental channels and a few rivers (e.g.,Eichenberger & Wuhrmann 1975; Horner & Welch1981; Horner et al. 1983; Bothwell 1985). A limitedsurvey in England was carried out by Pitcairn &Hawkes (1973).

The present study was aimed at determining theextent and composition of algal proliferations inNew Zealand rivers, to measure their standingcrops, and to carry out a preliminary characteris-ation of their habitats. A complementary survey ofmanagement problems caused by such prolifera-tions has already been reported (Biggs 1985a).

Received 18 August 1986; accepted 14 November 1986

METHODSDuring an extensive summer drought in February1983, 378 sites were surveyed in the North andSouth Island of New Zealand for tbe present offilamentous algal growths. Flow records, from therivers with water level recorders (65 rivers), indi-cated that most of the catchments had not receivedmajor fresh events for between 40 and 60 daysbefore sampling. The sampling programme was thus

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176 New Zealand Journal of Marine and Freshwater Research, 1987, Vol. 21

-36S

-38

-40

I174'E

\ - .Auckland-'

WellingtonC

174-E1

North

; ;

J

174'E

Island

. f176'

I

1 178*

N

f1

"7

36S-

38 -

40 -

16S'E

-44

/h40

170'E

South

H

/

f

J ^

Island

/

' ' ,k

J

170'E

* ' *

if

i*, Dunedin

/

170-\

1170'E

i-172-

\

44 -

46 -

174'E

Fig. 1A North Island map showing the distribution ofsites visited in the filamentous algal survey ( sites visitedin summer; + sites sampled in summer).

Fig. IB South Island map showing the distribution ofsites visited in the filamentous algal survey ( sites visitedin summer; + sampled in summer; sampled in winter).

designed to largely remove the influence of flowvariability on the communities which, if high, canhave a major retarding effect on communitydevelopment (Tett et al. 1978). A further 45 siteswere surveyed in the South Island during a periodof winter low flows in July and August 1983 (Fig.1). Most sites were dominated by "run" habitatsand were unshaded by overhanging vegetation.

For sites where filamentous algae occurred, thefollowing environmental descriptors were com-piled: channel form, approximate width and depthof the flow, water conductivity (standardised to25 C), substratum type, surrounding land use, avisual description of the turbidity of the water, anda classification of the origin of the river, based onNZMS 18 topographical maps (following Biggs1985b), as either alpine-fed, foothills-fed, lowland-fed, or lake-fed. A visual description of the fila-mentous algae was also compiled for each site (tex-ture, colour, locality of the growths on the riverbed), together with a visual estimate of the per-centage of the river bed covered. The accuracy of

the cover estimate was tested against a point inter-cept analysis and a good agreement between thetwo approaches was found {r = 0.900; P < 0.001).

The algae were sampled by randomly retrievingrocks with filamentous growths and placing a62 mm diameter cylinder on the top surface. Thiswas then scribed around, which cut all filamentscrossing the perimeter of the cylinder. Algae withinthe circumscribed area were then removed using ascalpel. Where very thick mats of long filamentsoccurred, sections of the mat were lifted from theriver bottom until just clear of the water and10 X 10 cm2 cut from it using surgical scissors. Carewas taken not to disturb the lie of the filaments.The sampling cylinder was then placed on thesquare mat and cut around to define a set area.This second method was generally necessary formats of Cladophora glomerata and Rhizocloniumsp. The number of samples collected at each sitevaried from one to five, depending on the hetero-geneity of the community (i.e., stratified propor-tional variance sampling). At least three samples

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Biggs & PriceFilamentous algae in New Zealand 177

were usually collected. These were preserved in a10% formalin solution and stored at 4 C.

In the laboratory, each sample was thoroughlymixed and three subsamples were removed andpooled. Abundance of taxa was assessed using aninverted microscope and an 8-point scale (1, rare;8, dominant). Samples in which the associated uni-cellular diatom frustules were difficult to identify,because of visual interference by filamentous taxa,were rechecked following ashing. A detailed tax-onomic analysis of the samples was not attempted.Non-diatom taxa were identified using Whitford &Schumacher (1973) and Prescott (1972); for thediatom identifications Patrick & Reimer (1966,1975) and Foged (1979) were used. In the majorityof samples some of the common filamentous taxacould not be defined to species level because manyof the essential distinguishing characters were notpresent (e.g., reproductive structures in Spirogyraspp. and Oedogonium spp.). Due to their high var-iability diatoms could usually not be identified tothe "variety" level. New distribution records forthe taxa were determined using Cassie (1984) andCassie (1986).

The mean standing crop of the filamentous dom-inated community at each site was estimated asmean replicate ash-free dry weight (AFDW) (dryweight at 105 C; ash weight at 550 C) multipliedby the proportion of the stream bed covered by thealgae.

Statistical analyses were carried out using theMinitab computer programme (Minitab 1984).Where the data were not normally distributed, geo-metric means were calculated these are identi-fied in the text; all other means are arithmetic.

A normal (sites by taxa) classification analysiswas carried out on the summer survey data to iden-tify clusters of sites with similar communities.Computations were carried out using the S.A.S.computer programme (S.A.S. Institute Inc. 1985),

Table 1 Number and nature of river sites visited duringthe summer North and South Island field surveys.

Sites visited

Filamentous periphyton presentsamplednot sampled1

No filamentous periphytonfavourable habitatunfavourable habitat

Dominated by macrophytesDryTotal number of sites visited

N.I.

799

406150

1240

S.I.

385

3097

49138

with the 52 most common taxa (i.e., taxa recordedat >10% of the sites) included. Squared euclideandistance (PROC CLUSTER) was used as a measure ofsimilarity which, because of its tendency to weighthigher values, was particularly appropriate for usewith the ordered multistate data-set of the survey.The similarity coefficients were converted into adendrogram using the agglomerative hierarchicalunweighted pair-group method with arithmeticaverages (METHOD AVERAGE). After clustering, acomplementary nodal analysis (Boesch 1977) wascarried out to show the pattern of taxonomic dom-inance between the site clusters. Nodal constancy(a quantitative measure of the consistency withwhich a taxon occurs at a high abundance in a par-ticular cluster of sites) was determined as:

Mainly owing to difficulty of access.

Cfl = [ajn^ ]100where a{i is the sum of the abundance scores fortaxon i in the site cluster j , n{is the maximum pos-sible abundance score for taxon i, n} is the totalnumber of sites in site cluster j , and Qis expressedas the percentage dominance of a node. Nodalfidelity (the abundance of a taxon in a particularcluster in relation to its abundance across all thesite clusters) was determined as:

Fn = (dry + al2 + ain)100where au to a^are the sums of the abundance scoresfor clusters 1 to n and the other terms are the sameas in the previous equation. Fi} is expressed as apercentage of the taxon's overall dominance.

RESULTSGeneralMats of filamentous algae were recorded at 131(35%) of the sites visited in summer, 117 of whichwere sampled (Table 1). The majority were inmoderately swift, shallow (

OP5.Qg

I70a

Ff8;^ J M i " S f CladP^or,a glomerata and Rhizoclonium sp. in a periphery seepage zone of a foothills-fed river in the Wairarapa (left) and clumpsof Cladophora sp. and Oedogonium spp. in a foothills-fed river near Christchurch (right). "*"!

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Fig. 3 Site clusters for the sum-mer survey data based on 6084data points. A nodal constancyanalysis in a two-way table ofcommunity clusters and the dom-inant filamentous taxa are alsoshown. The sites in each clusterare given in Table 2.

Site clusters

Nodal constancy

51-75%

, " 2-25%

summary of the main data collected at each site ispresented in Appendix 1. Appendix 2 lists the com-mon taxa identified in the survey and the sites atwhich they were found.

Cover of the river bed by filamentous algaeThe geometric mean cover of the sites sampled insummer was 26% ( = 114). In winter, the geo-metric mean cover was 22% (n = 34). The coverof the river bed by filamentous algae became veryconspicuous from the banks of most rivers whenit exceeded about 40%. This occurred at 67 sites,44% of the 148 sampled, or 16% of the total of 423sites surveyed in the study. Algal cover was greaterthan 90% at 17 sites (4% of the total numbervisited).River bed standing cropThe geometric mean summer standing crop was15.3 g/m2, and the geometric mean winter standingcrop was 1.87 g/m2. This difference between sum-mer and winter crops is significant (P < 0.05) whenthose rivers within similar conductivity ranges arecompared. However, it should be noted that wintersampling was only carried out in the South Island.

Standing crops higher than 50 g/m2 occurred at32 sites. These growths were usually composed ofthick mats of streaming filaments (up to 1.0 m longand 0.2 m thick, Fig. 2) which appeared to smotherthe bed sediments and were also conspicuous fromthe bank. Eleven sites, mainly small rivers aroundHawke's Bay, had standing crops higher than

100 g/m2. The highest standing crop (291 g/m2) wasrecorded in the Maraekakaho River, Hawke's Bay.

Community compositionThe taxa most commonly encountered on the sur-vey are listed in Table 3 and a summary of the sitesat which all taxa were found is given in Appendix2. Cluster analysis of the summer abundance dataidentified nine main groups of sites with differentcommunities in each group (Fig. 3). These groupsincluded 104 of the 114 sites, the remaining siteshaving poor taxonomic similarities. The sites anddominant taxa in each site group are listed in Table2. Five primary groups were identified (1, 3, 6, 7,and 9) which were dominated by taxa with highlevels of constancy. The remaining groups (2, 4, 5,8) were usually dominated by several of the majorfilamentous taxa and thus appeared to be inter-grades between the primary groups.

A qualitative association of catchment land useand the river community groups appeared to occur.The Ulothrix zonata-dominated group of sites typ-ically included rivers with poorly developed tus-sock and rocky catchments (low-conductivitywaters); groups of sites dominated by Stigeoclon-ium sp., Phormidium spp., and Spirogyra spp. weretypically in rivers with mixed tussock and forestcatchments (low-conductivity waters); dominanceof Cladophora glomerata and Rhizoclonium sp.typically occurred in rivers with pastoral catch-ments (high-conductivity waters); and dominanceof Cladophora sp. typically occurred in rivers with

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Community clusters1 2 3 4 5 6 7

TaxaZ.coarulsuS

Stigeoclonium sp.

Phormidium spp.

Oedogonium spp.

C.giomerata

Rhizoclonium sp.

M.varians

U. zonata

Cladophora sp.

Spirogyra spp.

Nodal fidelity

1

_^

1 76 -100%

^ 26-50%

o 2-25%

S 1 %

Fig. 4 Nodal fidelity analysis ina two-way table of site clusters anddominant taxa. The sites in eachcluster are given in Table 2.

mixed scrub and pastoral snow grass catchments(moderate-conductivity waters).

Of the 10 community dominants, only Ulothrixzonata and Cladophora glomerata displayed veryhigh levels of nodal fidelity (i.e., they were infre-quently found in other site groups and thus hadhigh habitat specificity); and Stigeoclonium sp. andRhizoclonium sp. displayed moderately high levels(Fig. 4). By contrast, Compsopogon coeruleus,Phormidium spp., Oedogonium spp., Melosira var-ians, Cladophora sp., and Spirogyra spp. had poorerspecificity in their habitat requirements (Fig. 4), andwere commonly found in several of the site groupsand thus over wider ranges of habitat conditions.

Fig. 5 shows the location of the site groups alongthe two-dimensional continua of water conductiv-ity and standing crop. A change in both the dom-inance and the standing crop of algae was evidentbetween the rivers with different levels of conduc-tivity/enrichment. The Cladophora glomerata andRhizoclonium sp. communities formed distinctlyhigher standing crops than the other communities(Fig. 5).

The South Island sites surveyed in winter wereusually dominated by Ulothrix zonata. The chain-forming diatom Diatoma hiemale var. mesodon wasalso abundant although it was rarely recorded insummer.

Table 2 Site groups, and their dominant taxa, shown in Fig. 3, 4 and 5.

Site Communitygroups dominants Sites

1, 6, 16, 23, 24, 51, 58, 65,66, 67, 68, 70, 74, 81, 8717, 18, 21, 59, 72, 75, 91,93, 100, 101, 109, 1154, 5, 13, 15, 39, 40, 55, 73,94,98, 111, 1123, 20, 28, 36, 38, 48, 53, 56,62,64, 77,82, 99, 102, 11730, 37, 47, 57, 80, 96

10, 11, 12, 14,27, 52, 69, 84,85, 92, 104, 10832, 41, 42, 43, 44, 45, 46, 49,50, 54, 95, 1149, 29, 35, 60, 76, 97, 1072, 7, 8, 19, 22, 31, 61, 71, 79,86, 88, 89, 90, 103, 110, 116

1 Ulothrix zonata

2 Stigeoclonium sp.Phormidium spp.

3 Spirogyra spp.

4 Spirogyra spp.Oedogonium sp.

5 Oedogonium spp.Cladophora sp.

6 Cladophora glomerata

7 Rhizoclonium sp.

8 Melosira varians9 Cladophora sp.

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Fig. 5 Relationship between themean conductivity of the waterand geometric mean standing cropof algae at the site cluster nodes.The error bars are 1 standarddeviation of each mean and areincluded to show the overlap incluster nodes along the continua.The cluster numbers and theirdominant filamentous taxa aregiven on the right.

70

1

(ft

6 0 -

50 "

4 0 "

3 0 "

20 "

10 "

Communities

"" 7-Rhizoclonium sp.

6-C.glomerata

8 -M.varians9 - Cladophora sp.

5 - Oedogonium spp, Cladophora sp.

4-Spirogyra spp., Oedogonlum spp.

1-U.zonata3-Splrogyra spp.

2- Sligeoclonium sp., Phormidium spp.

20 30 40Conductivity26 (mS/m)

50

Overall, the filamentous taxa most frequentlydominating the communities were Ulothrix zonata(36 sites), followed by Spirogyra spp. (23 sites)(Appendix 1). Other filamentous taxa which occa-sionally dominated the communities included Zyg-enma sp., Binucleria sp., Vaucheria sp., Mougeotiasp., and Audouinella hermanii.

A total of 27 unicellular taxa were commonlyfound among the filamentous dominants (Table 3),of which Achnanthes lanceolata, Cymbella kappii,Cocconeis placentula, Gomphoneis herculeana,Gomphonema parvulum, Navicula cryptocephala,Navicula rhyncocephala, and Synedra ulna wereubiquitous. Taxa which were more abundant withparticular filamentous dominants included: Epi-themia sorex and Gomphonema olivaceum (Table3). These two taxa, together with Chamaesiphonsp., Cocconeis placentula, Gomphonema parvulum,Rhoicosphenia curvata, and Synedra ulna oftenoccurred as epiphytes on the larger cells of Cla-dophora sp., Cladophora glomerata, and Rhizo-clonium sp.

A new record for a mat-forming filamentousRhodophyta species in New ZealandAudouinella hermanii (Roth) Duby was recordedin winter as a filamentous dominant at a site in the

regulated Lower Clutha River, Otago, and as aninfrequent component of communities in severalsmall rivers around New Zealand in summer(Appendix 2). Although this species was abundantin the Lower Clutha River it has not been previ-ously recorded in New Zealand.Taxonomy A variety of synonyms are in use, themost common being Rhodochorton violaceum(Kiitz) Drew, Audouinella violacea (Ktttz) Hamel,and Chantransia hermanii (Roth) Desuaux.Description Specimens from the Lower CluthaRiver are consistent with the description of materialfrom south-eastern Australia given by Entwistle &Kraft (1984).Habitat Audouinella hermanii formed extensive,fine, burgundy-red coloured mats (like velvet) overbedrock substrata; often found growing in con-junction with Musci or coating rootlets of crackwillows (Salix fragilus) where these protruded intoflowing waters.

The community formed a cover of 50% and astanding crop of 15 g/m2 at site 140. It was usuallysecurely anchored to the substratum and survivedtwofold daily changes in flow associated with up-stream power generation, as well as regular dailydewatering in shallow areas during the night andin the early morning. Extensive communities werefound on bedrock in normally deep (> 3 m), and

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turbid water (16.6 g/m3 suspended solids, 8.4 FTU; the following species: Synedra ulna, 128; CymbellaDavies-Colley 1985). kappii, 110; Cocconeis placentula, 99; Gomphoneis

herculeana, 99; Gomphonema parvulum, 98; Navi-New distribution records for the common algal cula rhyncocephala, 88; and Navicula cryptoce-

phala, 86.Appendix 2 shows new distribution records for allthe common algae collected in the survey, a totalof 2148 new species records and 980 new generic DISCUSSIONrecords. Many of these taxa are cosmopolitan and,with further collecting, would probably be found A large number of algal proliferations were recordedextensively throughout New Zealand. The largest in New Zealand streams and rivers in the survey,number of new distribution records were found for many of which had the potential to cause degra-

Table 3 The sum of of the abundance scores of the common taxa (lines) sepa-rated according to which filamentous taxa dominated the sample (columns). Taxain brackets are tentative identifications.

FilamentousA Cladophom sp.B C. glomerataC Compsopogon coeruleusD Melosira varians

B Oedogonium spp.F Phormidium spp.G Rhizoclonium sp.H Spirogyra spp.I Stigeoclonium sp.J Ulothrix zonataUnicellularAchnanthes lanceolataA. linearis/minutissimaChamaesiphon sp.Closterium sp.Cymbella kappiiC. minutaCocconeis placentulaDiatoma vulgareEpithemia sorexFragilaria vaucheriaeGomphoneis herculeanaGomphonema (angustatum)G. olivaceumG. parvulumG. (subclavatum)G. truncatumG. truncatum var. capitatum(Lyngbya sp.)Navicula cryptocephalaN. rhyncocephalaN. viridula var. avenaceaNitzschia sp. (a)N. paleaRhoicosphenia curvataScenedesmus sp.Synedra ulnaS. ulna var. contracta

A

_

00

31

3500

1900

16006

375

340

214

2954

410404

123310089

15366

Filamentous

B

13-

512

105

101004

5060

324

360

520

3156

1212006

25234

13533

3612

C

00

-

0

006500

000000503030000000000003060

D

040

-

2064305

12004

191323054

1333

214

1090

20249393

115310

E

1300

28_

30

29100

71108

1710250

118

1700

194000

10220000

144310

dominants

F

0003

4-

0553

11600

181112004

1200

1500006734337

247

G

9126

14

64_

000

90

123

135

43050

1200804

11057030

1213210

H

12040r\u33105

-

105

22600

4318210

1211540

10300007

24140

120

101349

6

I

0000

970

11-

5

100000

157000

2200

29000

113004003

140

J

040

012060_

22530

562413125

2853113

266003

11201721660

5739

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dation of water quality and aesthetic values. Thegrowths smothered river lied sediments and severelyimpeded water movement in some rivers (Fig. 2).The standing crops exceeded most natural maxi-mum values reported in the literature (Table 4).Only under artificial conditions do the cropsreported elsewhere approach the higher levelsrecorded in this survey (Appendix 1). The prolif-erations occurred even though large communitiesof grazing insects were often observed, which wouldbe expected to have checked their accrual (Eichen-

berger & Schlatter 1978). The presence of coarsecobble substrates, high light intensities, and diffusesource enrichment is likely to have aided this pro-liferation of algae at the sites. For example, zonesof seepage from farmlands into braided rivers wereoften evident as conspicuous peripheral accumu-lations of algae or as plumes of filaments from thebanks (Fig. 2). Temperature gradients of up to 10 Cwere recorded from the back of some of these zonesto where the cooler seepage water became fullymixed with the warmer river waters. In contrast,

Table 4 Maximum periphyton standing crops (as ash-free dry weight) from various studies in natu-ral and experimental stream systems. Only values from rock substrata in the natural streams havebeen included.

Reference

Natural streamsBahls 1971Cushing 1967Cushing et al. 1983

Dunn 1976Elwood & Nelson 1972Ertl et al. 1972Fisher et al. 1982Grimm et al. 1981Kobayashi 1961Liaw & MacCrimmon 1978Lyford & Gregory 1975Mulholland et al. 1985Muller 1978

Murphy et al. 1981Rose (unpubl. date)Rounick & Gregoiy 1981Stockner & Shortreed 1976Stockner & Shortreed 1978Experimental streamsClark et. al. 1979Eichenberger 1975Eichenberger 1979Eichenberger & Wuhrmann 1975Eichenberger & Schlatter 1978Erhlich & Slack 1969

Kaufman 1980Kevern et al. 1966Marker & Casey 1982Mclntire 1968aMclnitre 1968bMclntire & Phinney 1965Stockner & Shortreed 1978

Crop (g/m2)

18.94.26.56.7

18.77.5

13.29.5

22.667.70.7

65160290

7.082.68.81.85.22.1

15.33.52.77.56.64.4

7.0640.070.550.0

25010.413.911.112.014.965.7

141.3107.337.5

Locality/Treatment

East Gallatin R., Montana, U.S.A.Columbia R., Washington, U.S.A.Mowitch Ck, Idaho, U.S.A.Indian Ck, Idaho, U.S.A.Middle Fork, Idaho, U.S.A.Loon Ck, Idaho, U.S.A.Big Ck, Idaho, U.S.A.Middle Fork, Salmon River, Idaho, U.S.A.Salmon R., Idaho, U.S.A.Desert Stm, Idaho, U.S.A.Watershed Stm, Tennesse, U.S.A.Danube R., AustriaDesert Stm, Arizona, U.S.A.Desert Stm, Arizona, U.S.A.Arakawa R., JapanGrand R., Ontario, CanadaMountain Stm, Oregon, U.S.A.Walker Branch, Tennesee, U.S.A.Trydean R., SwedenTolangaan R., SwedenVerkean R., SwedenMcKenzie R., Oregon, U.S.A.Shite Oak Ck, Tennesee, U.S.A.Mountain Stm, Oregon, U.S.A.Carnation Ck, Vancouver I., CanadaCarnation Ck, Vancouver I., Canada

Outdoor troughs, no enrichmentOutdoor troughs, sewage enrichmentOutdoor troughs, inorganic enrichmentOutdoor troughs, enriched groundwaterOutdoor troughs, herbivores removedLab. streams, inorganic enrichmentLab. streams, organic enrichmentOutdoor troughsLab. streamsOutdoor troughs, enriched groundwaterLab. streams, high light, mod. currentLab. streams, snails removedLab. streams, high light intensityOutdoor troughs, inorganic enrichment

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rivers draining native bush catchments had poorlydeveloped filamentous algal mats.

Cladophora glomerata appears to be the maincomponent of algal proliferations in the rivers ofNorth America and Europe (Whitton 1970; Pit-cairn & Hawkes 1973). Only occasionally are othertaxa recorded as forming major growths. Prescott(1981) reports that Rhizoclonium sp. forms "long,stringy, sometimes ropelike strands in flowingwater" in North America, and Ulothrix zonata hasbeen recorded forming proliferations below organicpollution discharges in Europe (Fjerdingstad 1964)and after forest removal in North America (Likenset al. 1970). Although Cladophora glomerata dom-inated a significant number of proliferations in NewZealand (20%), Rhizoclonium sp. had the highestaverage covers and standing crops, and most fre-quently formed a crop > 50 g/m2. Ulothrix zonatamost frequently formed proliferations of >40%cover. In contrast to the findings of Fjerdingstad(1964), the Ulothrix zonata proliferations in NewZealand were associated with high-quality (i.e., low-conductivity) waters.

Little information appears to be available in theliterature on the tolerance of periphytic taxa to sil-tation. Moderate siltation of communities wasnoted at several sites (particularly in alpine-fedrivers). Spirogyra spp., Ulothrix zonata, Gompho-neis herculeana, and Melosira varians dominatedthese sites, and thus appeared to tolerate limitedsilt deposition. These taxa are all characterised bytheir high growth stature which would enable theirphotosynthetic portions to remain above the zoneof basal silt accumulation. Species which oftenoccurred among the filaments of these communi-ties included Synedra ulna, Cymbella kappii, Cym-bella minuta, Fragilaria vaucheriae, andGomphonema parvulum.

Many of the taxa commonly encountered in thesurvey displayed habitat preferences consistent withtheir reported ranges in North America and Europe.For example, Diatoma hiemale var. mesodon andUlothrix zonata are also found dominating coldwater mountain rivers in Europe (Blum 1956). Theonly major difference occurred for Ulothrix zonatawhich competes most successfully with other per-iphyton in unenriched streams in New Zealand, butin Europe has both a clean-water form and a pol-lution-tolerant form (Fjerdingstad 1964; Benson-Evans et al. 1975).

The relationship between water conductivity andstanding crop of the community types was close(Fig. 5). Future work will need to define thenutrients that are actually limiting algal develop-ment in New Zealand rivers and how long a periodof base flow is required (with different nutrientlevels) for proliferations to develop. This type ofinformation should enable the likely occurrence and

extent of proliferations to be predicted, thus aidingthe management of our river systems.

ACKNOWLEDGMENTSWe are grateful to our colleague Mr M. Close for valuablediscussion of the survey data, and to Dr P. A. Broady(University of Canterbury), Dr J. M. Quinn (Water QualityCentre, Ministry of Works and Development (MWD),Hamilton), and Ms B. Vaile (Hydrology Centre, MWD,Christchurch) for their constructive reviews. The assist-ance of Dr A. A. Davoren (Hydrology Centre, MWD,Christchurch) with the SAS clustering procedures and theverification of Audouinella hermanii by Dr T. J. Entwistle(University of Melbourne) are also gratefullyacknowledged.

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Appendix 1 Summary of the main data collected at each site sampled during the survey of filamentous algal growthsin New Zealand rivers. (Class, classification of the river's source, where A, alpine-fed; F, foothills-fed; L, lowland-fed; LK, lake-fed; N, number of samples).

Siteno. River

Summer survey1 Ashburton (Nth)2 Ashley3 Awatere4 Camping Gully5 Clarence6 Cla rence7 Cobbolds8 Conway9 Doughboy

10 Esk11 Flaxboume12 Gorge13 Hae Hae Te Moana14 Hangaroa15 Hapuku16 Hope17 Hope18 Horokiwi19 Horomanga20 Hoteo21 Howard22 Huddlestones23 Hurunui24 Hurunui25 Hutt26 Hutt27 Irrig. Canal28 Kaeo29 Kenepuru30 Kiwitea31 Kopuaranga32 Maharakeke33 Makairo34 Makakahi35 Makara36 Makotuku37 Makuri38 Mangakino39 Mangamuka40 Manganui41 Mangaone42 Mangaone43 Mangaone44 Mangaonuku45 Mangaonuku46 Mangaoraka47 Mangapiko48 Mangatangi49 Mangawhero50 Maraekakaho51 Matakitaki52 Ngaruroro53 Ohariu53 Okawa55 Okuku56 Opihi57 Oroua58 Oturere59 Pakuratahi60 Pauatahanui61 Percival62 Piakonui63 Poutu64 Puketotara65 Rakaia66 Rangitata67 Rangitata68 Rangitikei69 Ruakokoputuna70 Ruamahanga71 School Ck72 Selwyn73 Speargrass74 Station75 Tauherenikau

76 Taylor77 TeAhura

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10.199.00

13.7614.8424.4420.6023.2528.5013.2910.8123.848.10

13.6923.5313.71

11.5346.6435.4032.2529.3118.4319.7020.309.90

14.1344.708.64

13.7928.0658.259.766.44

11.7612.218.81

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132323113123112112

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23231123322

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Dominant taxa

Ulothrix zonataCladophora sp.Spirogyra spp.Phormidium spp.Spirogyra spp.Ulothrix zonataZygnema sp.Zygnema sp.,Cladophora sp.Vaucheria sp.Cladophora glomerataCladophora glomerataCladophora glomerataPhormidium spp..Cladophora glomerataSpirogyra spp.Ulothrix zonataSpirogyra spp.Stigeodonium sp.Cladophora sp.Cladophora sp.(Lyngbya sp.)Cladophora sp.Ulothrix zonataUlothrix zonataUlothrix zonataSpirogyra spp.,Stigeodonium sp.Cladophora glomerataBinucleria sp.,Spirogyra spp.Melosira variansCladophora sp.Cladophora sp.Rhizodonium sp,Cladophora glomerataCladophora glomerataSpirogyra spp.Spirogyra spp.Oedogonium spp.Compsopogon coeruleusCladophora sp.,Oedogonium spp.Spirogyra spp.Rhizodonium sp.Cladophora sp.Compsopogon coeruleusMelosira varians,Rhizodonium sp.Rhizodonium sp.Rhizodonium sp,.Oedogonium spp.Spirogyra spp.Rhizodonium sp.Rhizodonium sp.Melosira varians,Ulothrix zonataCladophora glomerataSpirogyra spp.Rhizodonium sp.Spirogyra spp.(Lyngbya sp.),Spirogyra spp.Melosira variansUlothrix zonataUlothrix zonataMelosira variansCladophora sp.Oedogonium spp.Binudearia sp.Spirogyra spp.,Zygnema sp.Ulothrix zonataUlothrix zonataUlothrix zonataUlothrix zonataCladophora glomerataCladophora glomerata,Cladophorag lomerata,Vaucheria sp.Phormidium spp.Spirogyra spp.Ulothrix zonataSpirogyra spp.,Stigeodonium sp.Melosira variansSpirogyra spp.

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Appendix 1 (Continued).Siteno. River78 Temuka

79 Tengawai80 Toko81 Tongariro82 Topehahae83 Tukipo84 Turakina85 Tutaekuri86 Victoria87 Waiau88 Waihi89 Waihopai90 Waihuka91 Waikanae92 Waikari93 Waikawa94 Waimana95 Waingongoro96 Wainui97 Wainuiomata98 Waioeka99 Waiohine

100 Waiopoua101 Waiotahi102 Waipa103 Waipara104 Waipaoa

105 Waipawa106 Waipawa

107 Wairiri108 Wairoa109 Wairoa110 Wairua111 Waitara112 Waiwhakaiho113 Wanganui114 Wanganui115 Warwick116 Whakanekeneke117 Whakatane

Winter survey118 Ashley119 Bealey120 Beaumont121 Bruces

122 Cass123 Clarence124 Clarence125 Countess126 Craigiebum127 Fraser128 Glentui129 Grays130 Hanmer131 Hororata132 Hurunui133 Jacks134 Jollies135 Kowai136 Kowai137 Kowhai138 Lower Clutha139 Lower Clutha140 Lower Clutha141 Lower Clutha142 Maruia143 Maruia-side ck144 Pahua145 Percival146 School Ck147 Selwyn148 Tuapeka149 Waiau150 Waimakariri151 Waipara151 Waipara152 Wait

NZMS-18 mapreferenceS44:81:69

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N32:88:95N42:02:83

S5 5:36:02N44:85:llN35:53:48Nl 7:61:96Nl 3:83:93N13:71:82N24:85:13N23:56:41S57:64:05N 18:23:49N45:43:02

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3.2110.314.42

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13.129.066.466.342.243.696.18

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Dominant taxaBinuclearia sp.,Phormidium spp.Melosira variansMelosira variansUlothrix zonataOedogonium spp.Cladophora glomerataRhizoclonium sp.Cladophora glomerataCladophora sp.Ulothrix zonataOedogonium spp.Cladophora sp.Cladophora sp.Stigeodonium sp.Cladophora glomerataStigeodonium sp.Spirogyra spp.Rhizoclonium sp.Cladophora sp.,Oedogonium spp.Melosira varians,Oedogonium spp.Spirogyra spp.Ulothrix (variabilis)Microspora sp.,Stigeodonium sp.Stigeodoniumsp.Stigeodonium s p .Spirogyra spp.Cladophora sp.Cladophora glomerata,Rhizoclonium sp.,Compsopogon coeruleusMelosira variansCladophora glomerata,Rhizoclonium sp.Vaucheria sp.Cladophora glomerataPhormidium spp.Cladophora sp.Spirogyra spp.Spirogyra spp.Phormidium spp.Rhizoclonium sp.Stigeodonium sp.Oedogonium spp.Spirogyra spp.

Ulothrix zonataUlothrix zonataPhormidium spp.Diatoma hiemale

var. mesodonDiatoma hiemale

var. mesodonSpirogyra spp.Ulothrix zonataUlothrix zonataDiatoma hiemale

var. mesodonUlothrix zonataUlothrix zonataUlothrix zonataUlothrix zonataDiatoma hiemale

var. mesodonUlothrix zonataUlothrix zonataUlothrix zonataUlothrix zonataMelosira variansUlothrix zonataUlothrix zonataVaucheria sp.Audouinella hermaniiOedogonium spp.Diatoma hiemale

var. mesodonDiatoma hiemale

var. mesodonUlothrix zonataDiatoma hiemale

var. mesodonUlothrix zonataUlothrix zonataPhormidium spp.Ulothrix zonataUlothrix zonataUlothrix zonataStigeodonium sp.Ulothrix zonata

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Appendix 2 Taxa of algae and the sites at which they were found, during the survey of New Zealandrivers. Taxa preceded by ? are tentative designations. River names for each site are given in Appendix1. * Denotes that the occurrence is not a new distribution record.Taxon SitesChlorophytaAnkistrodesmus falcatus (Corda) RalfsBinuclearia sp.Bulbochaete sp.Characium sp.Cladophora sp.Cladophora glomerata KiitzCladophora ?oligoclona KutzClosterium sp.Coelastrum sp.Cosmarium sp.Cylindrocapsa sp.Dictyosphaerium sp.Geminella crenulatocollis PrescottTGloeocystis sp.

Microspora sp.Mougeotia sp.Oedogonium spp.

Pediastrum sp.Rhizoclonium sp.

Scenedesmus sp.

Schroederia sp.Selenastrum sp.Spirogyra spp.

Staurastrum sp.Stigeoclonium Tftagelliferum KtitzStigeoclonium sp.Stigeoclonium ?pachydermum PrescottUlothrix aequalis KiltzVlothrix tenuissima KiitzUlothrix variabilis KutzVlothrix zonata (Weber & Mohr) Ktltz

Zygnema sp.ChrysophytaDinobryon sp.Tribonema sp.Vaucheria sp.BacillariophytaAchnanthes sp.Achnanthes Tdelicatula (Kutz) GninAchnanthes lanceolata (Breb.) Grun

Achnanthes linearis (W. Smith) GrunAchnanthes minutissima KutzAmphora sp.Asterionella formosa HassallCocconeis sp.Cocconeis placentula Ehr

Cyciotella sp.Cyclotella menhigiana KtitzCyciotella stelligera Cleve & Grun.Cymbella sp.Cymbella aspera (Ehr.) CleveCymbella cesatii (Robh.) GrunCymbella ?cistula (Hempr.) GrunCymbella cymbiformis (Ktttz) Van Heurok

2, 4, 5, 11, 12, 17, 18, 25, 26, 29, 31, 34, 35, 40, 46, 55, 60, 61, 67, 71, 73, 75,76, 78, 79, 83, 87, 88, 89, 91, 93, 95, 97, 100, 111, 115, 1287, 17, 28, 30, 37, 58, 63, 67, 77, 78, 97, 102, 103, 1051403, 11, 12, 17,23, 32, 1152, 4, 7, 8, 11, 15, 19, 20, 22, 30, 31, 32, 37, 38, 39, 41, 44, 45, 47, 55, 57, 61,64, 69, 71, 80, 86, 88, 89, 90, 94, 96, 98, 103, 105, 106, 108, 110, 116, 1175, 8, 10, 11, 12, 14, 27, 33, 34, 44, 52, 53, 57, 69, 70, 83, 84, 85, 92, 104, 105,106, 108424, 7, 11, 14, 17, 18, 22, 26, 29, 31, 35, 37, 39, 41, 45, 53, 55, 79, 80, 86, 87, 88,89, 96, 97, 98, 102, 11013, 55, 71, 79, 882, 4, 5, 8, 11, 17, 25, 26, 41, 55, 56, 59, 61, 73, 75, 79, 87, 88, 94, 97, 100, 1157, 55, 71, 89, 9717, 55, 76, 7944, 1062, 4, 13, 17, 26, 45, 53, 55, 59, 69, 76, 78, 79, 87, 91, 93, 97, 115, 118, 119, 121,122, 124, 126, 128, 130, 135, 136, 144, 145, 149, 150, 151, 15213, 17, 22, 55, 56, 73, 76, 94, 100, 115, 141, 1433, 17,25, 55,65, 93, 99, 115, 1282, 4, 5, 7, 8, 11, 13, 17, 19, 20, 25, 26, 28, 30, 31, 34, 35, 36, 37, 39, 41, 47, 48,53, 55, 57, 60, 61, 62, 63, 64, 69, 71, 75, 76, 79, 80, 82, 84, 86, 88, 89, 95, 96,97, 100, 102, 103, 105, 110, 116, 123, 128, 130, 14111, 35, 55,61, 79,8810, 12, 31, 32, 35, 40, 41, 42, 43, 44, 45, 46, 49, 50, 53, 54, 69, 84, 85, 95, 104,105, 106, 1142, 3, 4, 5, 8, 11, 13, 14, 17, 19, 22, 23, 25, 26, 28, 29, 30, 31, 32, 34, 35, 36, 40,44, 45, 46, 51, 52, 53, 55, 60, 61, 67, 68, 69, 71, 72, 73, 76, 78, 79, 83, 85, 88,89, 91, 92, 93, 94, 95, 97, 98, 103, 105, 106, 111, 1163, 4, 6, 11, 13, 16, 17, 18, 21, 28, 29, 35, 45, 55, 57, 59, 60, 61, 67, 70, 71, 74,75, 76, 79, 87, 88, 89, 96, 97, 99, 102, 111, 116, 128362, 3, 4, 5, 6, 8, 11, 12, 13, 14, 15, 16, 17, 18, 24, 26, 28, 30, 35, 36, 37, 38, 39,40, 43, 45, 48, 53, 54, 55, 56, 57, 58, 59, 60, 63, 64, 70, 71, 73, 75, 77, 79, 80,81, 82, 85, 86, 88, 89, 90, 94, 96, 97, 98, 100, 101, 102, 103, 105, 108, 111, 112,115, 116, 1172, 4, 13, 17, 55, 78, 79, 105, 128562, 17, 18, 20, 25, 26, 29, 35, 45, 56, 58, 59, 68, 69, 70, 75, 91, 93, 96, 100, 101,115, 128, 136, 151, 15256, 61, 93, 9758, 12825, 44, 59, 75, 8117, 59,99, 1151, 4, 5, 6, 10, 13, 15, 16, 17, 21, 23, 24, 25, 29, 36, 51, 58, 65, 66, 67, 68, 70,74, 75, 76, 81*, 87, 91, 97*, 100, 115, 118, 120, 121, 124, 125, 127, 128, 129,130, 132, 133, 134, 135, 137, 138, 140, 141, 142, 143, 144, 146, 147, 148, 149,150, 151, 152, 1532, 7, 8, 16, 19, 24, 25, 45, 55, 59, 64, 94, 103, 105, 115, 123, 137

6455, 1159, 22, 71, 107, 138, 139, 140, 148

4, 5, 7, 17, 19, 23, 51, 76, 87, 106, 1286, 17,24, 51, 71, 1151, 2, 4, 6, 12, 13, 16, 17, 18, 19, 22, 24, 25, 27, 29, 30, 32, 34, 35, 36, 37, 40,41, 43, 44, 45, 46, 49, 51, 53, 55, 57, 58, 61, 63, 65, 66, 68, 69, 71, 73, 75, 76,78, 79, 80, 81, 83, 88, 91, 93, 94, 95, 97, 99, 100, 102, 105, 106, 109, 110, 111,113, 114, 115, 128, 1374, 13, 25, 26, 29, 35, 53, 59, 61, 88, 91, 97, 12812, 44, 59, 10651, 68, 95, 12023, 10943, 52, 54, 63, 981, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 17, 19, 20, 21, 22, 24, 25, 26, 27,28, 30, 31, 32, 33, 34, 35, 36, 37, 39, 40, 41, 42, 43, 44, 45, 46, 47, 49, 50, 51,52, 53, 54, 55, 56, 57, 58, 59, 61, 63, 66, 67, 68, 69, 71, 74, 76, 78, 79, 80, 83,84, 85, 86, 88, 89, 90, 91, 92, 93, 94, 95, 96, 98, 100, 102, 103, 104, 105, 106,107, 108, 110, 111, 113, 114, 115, 116, 117, 128, 129, 137, 139, 14319, 29, 30, 35, 57, 60, 65, 90, 954, 13, 31, 44, 61, 71, 76, 79, 92, 106, 107, 1086, 23, 24, 88, 89, 1232, 16, 22, 26, 33, 47, 58, 61, 63, 69, 91, 96, 102, 106, 111, 115, 123, 128, 14313, 15, 55, 61, 79, 88, 98, 100, 103, 105, 12867, 888, 23, 45, 59, 103, 10744, 68, 98, 103, 128

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190

Appendix 2 (Continued).

New Zealand Journal of Marine and Freshwater Research, 1987, Vol. 21

Taxon Sites

Cymbella kappii Cholnoky

Cymbella minuta Hilse ex Rabh.

Cymbella sinuata Greg

Diatoma hiemale var. mesodon (Ehr)GrunDiatoma vulgare BoryEpithemia sp.Epithemia sorex Kiitz

Epithemia zebra var. saxonica(Kutz) GrunEunotia sp.Fragilaria sp.Fragilaria capucina DesmazFragilaria crotonensis (Ehr) GrunFragilaria vaucheriae (KUtz) Petersen

Fragilaria virescens var. capitata OstrFrustulia rhomboides (Ehr) De ToniFrustulia vulgaris (Thwaites) De ToniGomphoneis elegans (Grun) CleveGomphoneis eriense (Grun) Skv & MeyerGomphoneis herculeana (Ehr) Cleve

Gomphoneis herculeana var. clavataCleveGomphoneis herculeana var. robusta(Grun) CleveGomphonema sp.

Gomphonema accuminatum EhrGomphonema ?anguslatum (Kiitz) RabhGomphonema dichotomum KiitzGomphonema intricatum var. v/Z?ro(Ehr) CleveGomphonema olivaceum A-V.H.

Gomphonema parvulum (Kiitz) Grun

Gomphonema ?subclavatum (Grun) Grun

Gomphonema truncatum EhrGomphonema truncatum var. capitatum(Ehr) PatrGyrosigma accuminatum (Kutz) RabhMelosira granulata (Ehr) RalfsMelosira granulata var. angustissimaO. MullerMelosira varians Ag

Meridian circulare (Grev) AgNavicula sp.

Navicula capitata EhrNavicula cryptocephala Kutz

Navicula cuspidata Kutz

1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 15, 16, 17, 18, 19, 21, 22, 23, 24, 25, 26,27, 30, 31, 35, 36, 37, 38, 39, 40, 42, 43, 44, 45, 46, 49, 50, 5), 52, 53, 54, 55,57, 58, 59, 61, 63, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76. 78, 79, 81, 83,84, 85, 87, 88, 89, 90, 91, 92, 93, 94, 96, 97, 98, 100, 101, 102, 103, 104, 105,106, 108, 110, 111, 112, 113, 114, 115, 116, 117, 123, 127, 128, 129, 131, 132,136, 137, 138, 139, 140, 143, 147, 151, 1531, 2, 4, 5, 6, 8. 13, 14, 15, 17, 23, 24, 25, 26, 30, 31, 33, 34, 35, 36, 40, 43, 45,46, 49, 51, 52, 53, 55, 56, 61, 63, 65, 66, 67, 68, 69, 71, 73, 74, 78, 79, 84, 85,87, 88, 90, 91, 92, 93, 95, 97*, 98, 100, 105, 106, 107, 111, 112, 113, 114, 115, 127,128, 129, 131, 137, 138, 142, 1471, 2, 5, 6, 8, 10, 11, 14, 16, 18, 21, 25, 26, 30, 33, 34, 35, 41, 51, 52, 55, 56, 57,58, 59, 60, 65, 66, 68, 69, 70, 74, 75, 78, 79, 83, 84, 88, 90, 91, 93, 98, 100, 105,114, 115, 128, 142, 1431, 4, 6, 13, 24, 51, 58, 61, 63, 73, 74, 79, 87, 88, 90. 92, 93, 95, 114, 115, 119,120, 121, 122. 123, 124, 126, 127, 129, 130, 131, 133, 134, 135, 137, 142, 143,145, 150*1, 4, 23, 24, 65, 66, 6719, 1122, 3, 5, 6, 7, 8, 10, 11, 12, 14, 17, 27, 30, 36, 39, 40, 42, 43, 46, 52, 54, 57, 58,61, 63, 64, 69, 71, 76, 79, 81, 83, 84, 85, 86. 89, 90, 92, 96, 98, 102, 103, 104,108, 111, 114. 123, 127, 128, 1375, 14, 17, 24, 61, 63, 98, 108, 128, 140

19, 36, 37, 47, 80, 111, 114, 1434, 8, 12, 27, 28, 30, 37, 41, 45, 47, 53, 57, 58, 68, 71, 77, 86, 88, 94, 95, 102,110, 113, 115. 1284, 8312, 13, 128, 1311, 2, 4, 5, 6, 7, 9, 10, 12, 13, 15, 17, 22, 23, 24, 25, 27, 32, 42, 43, 44, 45, 51,52, 55, 56, 57, 61, 63, 65, 66, 67, 73, 74, 78. 85, 89, 98, 104, 106, 108, 115, 118,120, 123, 127, 128, 129, 136, 137, 138, 140, 143, 1511, 13, 66, 78, 794, 23, 35, 37, 51, 58, 115, 120, 138, 139, 140, 1419, 61, 115, 128811281, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 21, 23, 24, 25, 26,27, 30, 31, 32, 33, 35, 36, 37, 39, 40, 41, 42, 43, 44, 45, 46, 49, 51, 52, 53, 55,56, 57, 58, 60, 61, 63, 65, 66, 67, 68, 69, 70; 71, 72, 74, 78, 79, 81, 84, 85, 86,87, 88, 89, 90, 91, 92, 93, 94, 95, 98, 101, 105, 106, 108, 111, 112, 114, 115,117, 118, 123, 127, 128, 129, 132, 134, 137, 144, 147, 148, 151, 152, 15391

5, 15, 69, 78, 128

2, 7, 10, 12, 22, 23, 26, 27, 28, 30, 31, 32, 34, 35, 37, 40, 41, 42, 43, 44, 45, 49,51, 52, 55, 57, 58, 59, 61, 63, 64, 65, 69, 71. 75, 83, 84, 90, 98, 99, 103, 106,112,113,114,115,128,14464, 71, 79, 96, 107, 1161, 3, 6, 21, 29, 30, 31, 33, 41, 51, 55, 64, 68, 74, 76. 83, 84, 87, 88, 93, 115, 1283, 4, 5, 6, 8, 12, 13, 17, 21, 24, 52, 55, 61, 66, 67, 69, 71, 77, 78, 79, 89, 98,105, 115, 1291, 13

1, 2, 4, 9, 19, 23, 24, 25, 29, 30, 31, 35, 36, 37, 41, 42, 43, 46, 47, 49, 53, 54,55, 56, 57, 59, 63, 66, 67, 68, 69, 71, 77, 84, 88, 89, 90, 91, 95, 97, 100, 102, 105,107, 111, 112, 1511, 2, 4, 5, 6, 7, 9, 10, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 25, 26, 27,28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 40, 41, 42, 43, 44, 45, 46, 47, 49, 51, 52,53, 54, 55, 56, 57, 58, 59, 60, 61, 63, 64, 67, 69, 70, 71, 72, 73, 75, 76, 77, 78,80, 83, 84, 85, 87, 88, 89, 90, 91, 92, 93, 9< 95, 97, 98, 99, 100, 101, 102, 104,105, 106, 107, 108, 109, 110, 111, 112, 113, 114, 115, 128, 1291, 2, 4, 5, 8, 9, 11, 12, 13, 14, 15, 24, 25, 27, 31, 32, 42, 43, 44, 45, 51, 52, 55,56, 57, 61, 67, 69, 79, 84, 85, 87, 88, 89, 90, 92, 97, 104, 105, 107, 108, 120,128, 1372, 4, 7, 11, 13, 27, 30, 32, 44, 45, 51, 55, 57, 61, 65. 67, 71, 79, 83, 88, 89, 92,94, 96, 97, 98, 105, 107, 128, 13712, 14, 32, 44, 45, 50, 52, 54, 71, 75, 92, 106, 107, 108

1, 31, 34, 44, 83, 8787, 120, 129, 13963

2, 4, 8, 9, 11, 13, 17, 18, 19, 22, 25, 29, 30, 31, 32, 33, 34, 35, 39, 40, 44, 45,46, 51, 52, 53, 55, 57, 58, 60, 61, 63, 69, 71. 72, 75, 76, 77, 78, 79, 80, 81, 83,86, 88, 89, 90, 91, 93, 94, 95, 96, 97, 98, 100, 102, 103, 105, 106, 107, 110, 111,112, 114, 115. 116, 128, 136, 138, 140, 141, 146, 14821, 100, 115, 120, 1297, 8, 10, 12, 13, 14, 27, 28, 36, 38, 40, 43, 54, 55, 56, 57, 58, 61, 63, 68, 69, 71,76, 77, 81, 83, 86, 88, 90, 91, 92, 93, 94, 97, 98, 100, 102, 104, 106, 107, 108,111,112,12835, 43, 61, 69, 90, 921, 2, 4, 5, 6, 7, 8, 9, 10, 11, 13, 14, 15, 17, 18, 19, 20, 22, 23, 24, 25, 26, 27,28, 29, 30, 31, 33, 34, 35, 37, 40, 42, 43, 44, 45, 46, 47, 50, 51, 52, 53, 54, 55,56, 57, 58, 60, 63, 67, 68, 69, 70, 71, 73, 74, 76, 79, 80, 83 84 85, 86, 88, 8990, 91, 92, 93, 95, 97, 98, 102, 103, 104, 105, 107, 111, 113, 115, 116, 128, 129,136, 137, 1427, 8, 29, 57, 104

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Appendix 2 (Continued).Taxon Sites

Navicula decussis OstrupNavicula exigua (Greg) GrunNavicula pupula Kfitz

Naviada pusio CleveNavicula radiosa KUtzNavicula rhyncocephala Kutz

Navicula viridula KiitzNavicula virdiula var. avenacea(Breb ex Grun) V.H.

Nedium sp.Nedium affine (Ehr) CleveNitzschia spp. Hassall

Nitzschia (disspiata) (Kutz) GrunNitzschia ?linearis W. SmithNilzschia palea (Kutz) W. Smith

Pinnularia sp.Rhoicosphenia adolfi M. SchmidtRhoicosphenia curvata (Kiitz) Grun

Rhopalodia sp.Rhopalodia ?novae zelandiae HustSurirella sp.Surirella angusta KiitzSurirella Hnearis W. SmithSurirella ovata KutzSynedra sp.Synedra acus KiitzSynedraacusKiitzSynedra ?minuscula G r u n

Synedra radians Ku tzSynedra ?rumpens Kii tzSynedra ulna (Nitzsch) Ehr

Synedra ulna ?var. contracta Ostrup

Synedra ulna ?var. ramesi (Herib)HustTabellaria sp.Tabellaria flocculosa (Rabhi) KutzRhodophytaAudouinella hermanii (Roth) DubyCompsopogon coeruleus (Balb) MontagneCyanobacteriaAnabaena sp.

Calothrix sp.Chamaesiphon sp.Chroococcus sp.Gloeotrichia sp.

Merismopedia sp.Nostoc sp.Phormidium spp.

Schizothrix sp.Spirulina sp.

2, 3, 4, 5, 10, 11, 15, 16, 24, 27, 72, 74, 898, 25, 28, 31, 57, 83, 84, 86, 87, 89, 97, 98, 1166, 7, 10, 13, 17, 25, 26, 33, 41, 43, 44, 55, 58, 63, 71, 74, 76, 86, 88, 89, 93, 97,102, 115, 129, 1374, 13,21,29, 55, 56, 60, 65, 88, 1282, 4, 5, 7, 8, 12, 13, 15, 17, 21, 24, 33, 36, 41, 43, 44, 55, 58, 61, 63, 64, 72, 73,79, 86, 87, 88, 92, 98, 103, 105, 111, 112, 116, 128, 137, 1431, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 19, 22, 24, 26, 28, 29, Ji,32, 33, 34, 35, 37, 40, 42, 43, 44, 45, 47, 49, 50, 51, 52, 53, 54, 55, 56, 57, 58,60, 61, 63, 67, 68, 69, 70, 72, 73, 76, 77, 78, 79, 80, 81, 84, 85, 86, 87, 88, 89,90, 92, 93, 94, 95, 96, 97, 98, 100, 101, 102, 104, 106, 107, 108, 111, 115, 116,117, 128, 129, 1377, 22, 23, 27, 31, 35, 39, 46, 58, 81, 84, 90, 95, 96, 102, 110, 113, 114, 116, 128,129, 131, 140, 146, 147, 1522, 6, 7, 9, 10, 12, 14, 18, 24, 26, 29, 31, 34, 35, 36, 37, 40, 42, 43, 47, 51, 53,57, 61, 63, 67, 68, 69, 70, 72, 73, 76, 77, 78, 79, 80, 81, 84, 85, 86, 87, 88, 89,90, 92, 93, 94, 95, 96, 97, 98, 100, 101, 102, 104, 106, 107, 108, 111, 115, 116,117, 120, 128, 138, 139, 140, 141, 14844, 106, 12822,35, 74, 1151, 2, 3, 4, 5, 7, 8, 10, 11, 13, 14, 17, 18, 30, 31, 39, 45, 52, 58, 61, 63, 64, 71,76, 79, 80, 81, 84, 88, 94, 95, 96, 100, 101, 102, 103, 104, 107, 108, 116, 128,129, 138, 1402, 5, 6, 8, 17, 31, 51, 53, 59, 68, 70, 87, 99, 1002, 4, 5, 7, 8, 9, 11, 15, 22, 33, 34, 35, 42, 43, 44, 45, 55, 61, 67, 71, 74, 76, 79,83, 85, 90, 92, 98, 104, 107, 108, 1282, 4, 5, 6, 8, 9, 11, 13, 15, 17, 18, 30, 35, 40, 41, 42, 44, 52, 61, 63, 64, 71, 80,81, 84, 88, 94, 95, 96, 100, 101, 102, 103, 104, 108, 12837, 39, 52, 61, 67, 80, 96, 107, 115, 120, 128, 129, 13930, 35, 81, 881, 4, 5, 6, 7, 8, 10, 11, 12, 13, 14, 16, 17, 22, 24, 26, 27, 29, 30, 31, 32, 33, 35.39, 40, 41, 42, 43, 44, 45, 46, 47, 49, 50, 51, 52, 53, 55, 57, 58, 61, 63, 68, 69,71, 72, 73, 74, 76, 79, 83, 84, 85, 86, 88, 90, 92, 93, 94, 95, 96, 97, 98, 100, 105,106, 107, 108, 111, 112, 114, 128, 129, 13758, 61, 875, 7, 8, 11, 61, 63, 81, 87, 98, 108, 123, 129, 137, 14041, 92, 107, 108, 120, 12817, 22, 35, 70, 72, 73, 76, 89, 100, 107, 11579, 1289, 12, 15, 22, 26, 29, 31, 33, 35, 49, 51, 69, 71, 76, 95, 97, 100, 128, 1291, 2, 13, 16, 24, 34, 61, 65, 66, 69, 71, 72, 88, 113, 114, 127, 1283, 5, 6, 16, 23, 24, 33, 46, 61, 63, 67, 72, 73, 74, 75, 76, 98, 115, 123, 127, 1284, 5, 6, 11, 13, 16, 17, 21, 23, 24, 25, 26, 32, 33, 34, 45, 51, 59, 61, 66, 72, 73,74, 75, 83, 87, 88, 89, 102, 111, 115, 123, 127, 128, 1375,8, 16,24,30, 57,63,67,79,88, 11113, 23, 42, 55, 61, 66, 67, 71, 85, 87, 105, 1281, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17, 19, 21, 22, 23, 24, 26, 27, 28,29, 30, 31, 32, 33, 34, 35, 36, 37, 39, 40, 41, 42, 43, 44, 45, 46, 47, 50, 51, 52,53, 55, 56, 57, 58, 59, 60, 61, 63, 64, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75,76, 77, 78, 79, 80, 83, 84, 85, 86, 87, 88, 90, 91, 92, 93, 94, 95, 96, 97, 98, 99,100, 101, 102, 103, 104, 105, 106, 107, 108, 109, 111, 113, 114, 115, 116, 117,118, 120, 121, 123, 125, 127, 128, 129, 131, 132, 133, 134, 136, 137, 139, 143,145, 146, 147, 148, 149, 150, 151, 152, 1532, 4, 6, 8, 9, 10, 12, 14, 17, 19, 23, 24, 25, 29, 31, 32, 33, 36, 42, 43, 44, 46, 47,49, 51, 52, 55, 57, 58, 63, 66, 68, 69, 70, 73, 74, 75, 76, 77, 79, 81, 85, 86, 87,89, 91, 92,94,97,98, 101, 102, 104, 108, 110, 111, 112, 113, 114, 115, 116, 117,128, 1297, 10, 13, 14, 17, 22, 27, 42, 43, 55, 56, 69, 78, 79, 85, 88, 90, 92, 98, 104, 128,12937, 4417, 63, 115, 123, 127, 129, 143

34, 56, 78, 88, 129, 138, 139, 140, 14120, 28, 32, 38, 41, 43, 78, 102, 104, 140

1, 2, 3, 5, 6, 7, 8, 11, 13, 16, 17, 23, 28, 37, 39, 40, 45, 53, 55, 58, 61, 64, 68, 6978, 79, 81, 83, 86, 88, 89, 90, 95, 96, 97, 98, 100, 103, 105, 115, 116, 117, 12817, 61, 1032, 17, 32, 34, 45, 49, 51, 55, 56, 58, 70, 81, 84, 88, 91, 95, 100, 114, 123, 12813, 55, 61, 79, 8821, 36, 57, 58, 84, 932, 11, 15, 17, 21, 35, 55, 56, 61, 74, 76, 83, 91, 93, 100, 115, 1282, 5, 13, 55, 67, 1283,611, 2, 3, 4, 5, 8, 9, 13, 15, 17, 18, 21, 23, 24, 25, 26, 32, 33, 34, 36, 45, 51, 53,55, 58, 59, 60, 61, 63, 64, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78, 80,83, 86, 87, 91, 94, 95, 97, 100, 101, 102, 103, 105, 106, 109, 111, 112, 113, 115,120, 129, 137, 141, 143, 148, 15217, 1403, 4, 6, 12, 18, 24, 26, 31, 35, 41, 49, 53, 55, 56, 58, 63, 70, 71, 74, 75, 78, 79,86, 87, 88, 93, 96, 97, 128

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