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New Zealand Journal of Marine and

Freshwater Research
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A survey of filamentous algal

proliferations in New Zealand rivers
a

Barry J. F. Biggs & Geoff M. Price

Hydrology Centre , Ministry of Works and Development , P. O.

Box 1479, Christchurch, New Zealand
Published online: 29 Mar 2010.

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

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New Zealand Journal of Marine and Freshwater Research, 1987, Vol. 2 1 : 175-191
0028-8330/87/2102-0175$2.50/0
Crown copyright 1987

175

A survey of filamentous algal proliferations

in New Zealand rivers

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BARRY J. F. BIGGS
GEOFF M. PRICE
Hydrology Centre
Ministry of Works and Development
P. O. Box 1479, Christchurch
New Zealand
Abstract
New Zealand rivers were surveyed for
filamentous algal proliferations following prolonged periods of low flows to define their extent,
standing crop, and composition. Significant growths
were found at 167 of the 423 sites surveyed. The
geometric mean river bed cover of filamentous algae
was 26% in summer and 22% in winter. Proliferations 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 in
winter. Proliferations of >50 g/m2 (AFDW)
occurred at 8% of the sites. A taxonomic classification analysis of the summer data identified nine
main clusters of sites which appeared to correspond to different catchment land uses. One, or a
combination, of the following taxa dominated these
clusters: Ulothrix zonata, Phormidium spp., Stigeoclonium sp., Spirogyra spp., Compsopogon coeruleus, Oedogonium spp., Cladophora sp.,
Cladophora glomerata, Rhizoclonium sp., and
Melosira varians. Ulothrix zonata and Diatoma
hiemale var. mesodon dominated the sites sampled
in winter. There was a close positive relationship
between the average standing crop and the average
water conductivity of the nine site clusters.
Keywords
freshwater ecology; ecological associations; low flows; periphyton; algae; enrichment;
eutrophication; trophic; biomass; water quality

INTRODUCTION
Proliferations of filamentous algae are often
observed in New Zealand rivers during summer.
Problems caused by these growths include clogging
of abstraction structures, degradation of water
quality (through diel fluctuations in dissolved oxygen and pH), and degradation of aesthetic values
(Biggs 1985a).
Little is known of the composition or ecology of
these proliferations in New Zealand. Only limited
observations have been reported as part of invertebrate studies (e.g., Hirsch 1958; Winterbourn et
al. 1971), or as part of studies on the effects of the
algae on river water quality (Biggs 1982; Freeman
& McFarlane 1982). Recently, more detailed studies have been carried out on nutrient limitation of
Cladophora glomerata in the Manawatu River
(Freeman 1986).
Most European and North American studies on
river enrichment have focused on the ecology of
Cladophora glomerata, a major problem taxon in
these areas (the early studies have been reviewed
by Whitton (1970)), or on enrichment processes in
experimental channels and a few rivers (e.g.,
Eichenberger & Wuhrmann 1975; Horner & Welch
1981; Horner et al. 1983; Bothwell 1985). A limited
survey in England was carried out by Pitcairn &
Hawkes (1973).
The present study was aimed at determining the
extent and composition of algal proliferations in
New Zealand rivers, to measure their standing
crops, and to carry out a preliminary characterisation of their habitats. A complementary survey of
management problems caused by such proliferations has already been reported (Biggs 1985a).

METHODS

Received 18 August 1986; accepted 14 November 1986

During an extensive summer drought in February

1983, 378 sites were surveyed in the North and
South Island of New Zealand for tbe present of
filamentous algal growths. Flow records, from the
rivers with water level recorders (65 rivers), indicated that most of the catchments had not received
major fresh events for between 40 and 60 days
before sampling. The sampling programme was thus

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

176
I
174'E

174'E

1
178*

170'E

South
North

1
170'E

170'E

Island

Island

16S'E

36S-

-36S

\ - .

Auckland-'

.<,w

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-38

"7

38

-44

'

/
/

-40

h
/

J
Wellington

;> f

''

40

176'
I

46

i*, Dunedin

,k

J
174-E
1

;
40

i44

/
174'E

^
170 \

172\

Fig. 1A North Island map showing the distribution of Fig. IB South Island map showing the distribution of
sites visited in thefilamentousalgal survey ( sites visited sites visited in thefilamentousalgal survey ( sites visited
in summer; + sites sampled in summer).
in summer; + sampled in summer; sampled in winter).
designed to largely remove the influence of flow
variability on the communities which, if high, can
have a major retarding effect on community
development (Tett et al. 1978). A further 45 sites
were surveyed in the South Island during a period
of winter low flows in July and August 1983 (Fig.
1). Most sites were dominated by "run" habitats
and were unshaded by overhanging vegetation.
For sites where filamentous algae occurred, the
following environmental descriptors were compiled: channel form, approximate width and depth
of the flow, water conductivity (standardised to
25 C), substratum type, surrounding land use, a
visual description of the turbidity of the water, and
a classification of the origin of the river, based on
NZMS 18 topographical maps (following Biggs
1985b), as either alpine-fed, foothills-fed, lowlandfed, or lake-fed. A visual description of the filamentous algae was also compiled for each site (texture, colour, locality of the growths on the river
bed), together with a visual estimate of the percentage of the river bed covered. The accuracy of

the cover estimate was tested against a point intercept analysis and a good agreement between the
two approaches was found {r = 0.900; P < 0.001).
The algae were sampled by randomly retrieving
rocks with filamentous growths and placing a
62 mm diameter cylinder on the top surface. This
was then scribed around, which cut all filaments
crossing the perimeter of the cylinder. Algae within
the circumscribed area were then removed using a
scalpel. Where very thick mats of long filaments
occurred, sections of the mat were lifted from the
river bottom until just clear of the water and
10 X 10 cm2 cut from it using surgical scissors. Care
was taken not to disturb the lie of the filaments.
The sampling cylinder was then placed on the
square mat and cut around to define a set area.
This second method was generally necessary for
mats of Cladophora glomerata and Rhizoclonium
sp. The number of samples collected at each site
varied from one to five, depending on the heterogeneity of the community (i.e., stratified proportional variance sampling). At least three samples

Downloaded by [114.124.25.25] at 21:29 22 May 2015

Biggs & PriceFilamentous algae in New Zealand

were usually collected. These were preserved in a
10% formalin solution and stored at 4 C.
In the laboratory, each sample was thoroughly
mixed and three subsamples were removed and
pooled. Abundance of taxa was assessed using an
inverted microscope and an 8-point scale (1, rare;
8, dominant). Samples in which the associated unicellular diatom frustules were difficult to identify,
because of visual interference by filamentous taxa,
were rechecked following ashing. A detailed taxonomic analysis of the samples was not attempted.
Non-diatom taxa were identified using Whitford &
Schumacher (1973) and Prescott (1972); for the
diatom identifications Patrick & Reimer (1966,
1975) and Foged (1979) were used. In the majority
of samples some of the common filamentous taxa
could not be defined to species level because many
of the essential distinguishing characters were not
present (e.g., reproductive structures in Spirogyra
spp. and Oedogonium spp.). Due to their high variability diatoms could usually not be identified to
the "variety" level. New distribution records for
the taxa were determined using Cassie (1984) and
Cassie (1986).
The mean standing crop of the filamentous dominated community at each site was estimated as
mean replicate ash-free dry weight (AFDW) (dry
weight at 105 C; ash weight at 550 C) multiplied
by the proportion of the stream bed covered by the
algae.
Statistical analyses were carried out using the
Minitab computer programme (Minitab 1984).
Where the data were not normally distributed, geometric means were calculated these are identified in the text; all other means are arithmetic.
A normal (sites by taxa) classification analysis
was carried out on the summer survey data to identify 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 during

the summer North and South Island field surveys.
Sites visited
Filamentous periphyton present
sampled
not sampled1
No filamentous periphyton
favourable habitat
unfavourable habitat
Dominated by macrophytes
Dry
Total number of sites visited
Mainly owing to difficulty of access.

N.I.

S.I.

79
9

38
5

40
61
50
1
240

30
9
7
49
138

177
with the 52 most common taxa (i.e., taxa recorded
at >10% of the sites) included. Squared euclidean
distance (PROC CLUSTER) was used as a measure of
similarity which, because of its tendency to weight
higher values, was particularly appropriate for use
with the ordered multistate data-set of the survey.
The similarity coefficients were converted into a
dendrogram using the agglomerative hierarchical
unweighted pair-group method with arithmetic
averages (METHOD AVERAGE). After clustering, a
complementary nodal analysis (Boesch 1977) was
carried out to show the pattern of taxonomic dominance between the site clusters. Nodal constancy
(a quantitative measure of the consistency with
which a taxon occurs at a high abundance in a particular cluster of sites) was determined as:
Cfl = [ajn^ ]100
where a{i is the sum of the abundance scores for
taxon i in the site cluster j , n{is the maximum possible abundance score for taxon i, n} is the total
number of sites in site cluster j , and Q i s expressed
as the percentage dominance of a node. Nodal
fidelity (the abundance of a taxon in a particular
cluster in relation to its abundance across all the
site clusters) was determined as:
Fn = (dry + al2 +
ain)100
where au to a^are the sums of the abundance scores
for clusters 1 to n and the other terms are the same
as in the previous equation. Fi} is expressed as a
percentage of the taxon's overall dominance.

RESULTS
General
Mats of filamentous algae were recorded at 131
(35%) of the sites visited in summer, 117 of which
were sampled (Table 1). The majority were in
moderately swift, shallow (<0.5 m deep), stony
bedded streams in Hawkes Bay, Canterbury, Taranaki, Manawatu, and Wellington. The 247 sites
where filamentous algae were not present were
either silty and dominated by aquatic macrophytes
(15%, Table 1) such as Elodea canadensis and Potamogeton spp. (lowland rivers of Northland, Waikato, and the Hauraki Plains), or had physical
conditions unfavourable for algal growth, such as
deep and turbid water. Although 70 (19%) of the
sites had favourable habitat conditions (e.g., rivers
with clear rapid flowing waters and a coarse bed
sediment), they did not support algal growth. Most
of the 50 (13%) dry sites were in braided rivers in
the South Island.
Of the 45 South Island sites surveyed during
winter, 36 (80%) had filamentous algal growths. The
majority of these were in Canterbury rivers. A

Downloaded by [114.124.25.25] at 21:29 22 May 2015

P
5.
Q

I
70
a

F 8
f ;^ 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 clumps
of Cladophora sp. and Oedogonium spp. in a foothills-fed river near Christchurch (right).
"*"!

Biggs & PriceFilamentous algae in New Zealand

Fig. 3 Site clusters for the summer survey data based on 6084
data points. A nodal constancy
analysis in a two-way table of
community clusters and the dominant filamentous taxa are also
shown. The sites in each cluster
are given in Table 2.

179
Site clusters

Nodal constancy

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51-75%

, "

summary of the main data collected at each site is

presented in Appendix 1. Appendix 2 lists the common taxa identified in the survey and the sites at
which they were found.
Cover of the river bed by filamentous algae

The geometric mean cover of the sites sampled in

summer was 26% ( = 114). In winter, the geometric mean cover was 22% (n = 34). The cover
of the river bed by filamentous algae became very
conspicuous from the banks of most rivers when
it exceeded about 40%. This occurred at 67 sites,
44% of the 148 sampled, or 16% of the total of 423
sites surveyed in the study. Algal cover was greater
than 90% at 17 sites (4% of the total number
visited).
River bed standing crop

The geometric mean summer standing crop was

15.3 g/m2, and the geometric mean winter standing
crop was 1.87 g/m2. This difference between summer and winter crops is significant (P < 0.05) when
those rivers within similar conductivity ranges are
compared. However, it should be noted that winter
sampling was only carried out in the South Island.
Standing crops higher than 50 g/m2 occurred at
32 sites. These growths were usually composed of
thick mats of streaming filaments (up to 1.0 m long
and 0.2 m thick, Fig. 2) which appeared to smother
the bed sediments and were also conspicuous from
the bank. Eleven sites, mainly small rivers around
Hawke's Bay, had standing crops higher than

2-25%

100 g/m2. The highest standing crop (291 g/m2) was

recorded in the Maraekakaho River, Hawke's Bay.
Community composition

The taxa most commonly encountered on the survey are listed in Table 3 and a summary of the sites
at which all taxa were found is given in Appendix
2. Cluster analysis of the summer abundance data
identified nine main groups of sites with different
communities in each group (Fig. 3). These groups
included 104 of the 114 sites, the remaining sites
having poor taxonomic similarities. The sites and
dominant taxa in each site group are listed in Table
2. Five primary groups were identified (1, 3, 6, 7,
and 9) which were dominated by taxa with high
levels of constancy. The remaining groups (2, 4, 5,
8) were usually dominated by several of the major
filamentous taxa and thus appeared to be intergrades between the primary groups.
A qualitative association of catchment land use
and the river community groups appeared to occur.
The Ulothrix zonata-dominated group of sites typically included rivers with poorly developed tussock and rocky catchments (low-conductivity
waters); groups of sites dominated by Stigeoclonium sp., Phormidium spp., and Spirogyra spp. were
typically in rivers with mixed tussock and forest
catchments (low-conductivity waters); dominance
of Cladophora glomerata and Rhizoclonium sp.
typically occurred in rivers with pastoral catchments (high-conductivity waters); and dominance
of Cladophora sp. typically occurred in rivers with

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

180
Community clusters
1

7
Taxa

Nodal fidelity

Z.coarulsuS

1 76 -100%

Stigeoclonium

sp.

Phormidium

spp.

Oedogonium

spp.

C.giomerata
Rhizoclonium

sp.

Fig. 4 Nodal fidelity analysis in

a two-way table of site clusters and
dominant taxa. The sites in each
cluster are given in Table 2.

^_
^

26-50%

2-25%

M.varians
U. zonata
Cladophora

sp.
S1%

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Spirogyra

spp.

mixed scrub and pastoral snow grass catchments

(moderate-conductivity waters).
Of the 10 community dominants, only Ulothrix
zonata and Cladophora glomerata displayed very
high levels of nodal fidelity (i.e., they were infrequently found in other site groups and thus had
high habitat specificity); and Stigeoclonium sp. and
Rhizoclonium sp. displayed moderately high levels
(Fig. 4). By contrast, Compsopogon coeruleus,
Phormidium spp., Oedogonium spp., Melosira varians, Cladophora sp., and Spirogyra spp. had poorer
specificity in their habitat requirements (Fig. 4), and
were commonly found in several of the site groups
and thus over wider ranges of habitat conditions.

Fig. 5 shows the location of the site groups along

the two-dimensional continua of water conductivity and standing crop. A change in both the dominance and the standing crop of algae was evident
between the rivers with different levels of conductivity/enrichment. The Cladophora glomerata and
Rhizoclonium sp. communities formed distinctly
higher standing crops than the other communities
(Fig. 5).
The South Island sites surveyed in winter were
usually dominated by Ulothrix zonata. The chainforming diatom Diatoma hiemale var. mesodon was
also abundant although it was rarely recorded in
summer.

Table 2 Site groups, and their dominant taxa, shown in Fig. 3, 4 and 5.
Site Community
groups dominants
1

Ulothrix zonata

Stigeoclonium sp.
Phormidium spp.
Spirogyra spp.

3
4
5

Spirogyra spp.
Oedogonium sp.
Oedogonium spp.
Cladophora sp.

Cladophora glomerata

Rhizoclonium sp.

8
9

Melosira varians
Cladophora sp.

Sites
1, 6, 16, 23, 24, 51, 58, 65,
66, 67, 68, 70, 74, 81, 87
17, 18, 21, 59, 72, 75, 91,
93, 100, 101, 109, 115
4, 5, 13, 15, 39, 40, 55, 73,
94,98, 111, 112
3, 20, 28, 36, 38, 48, 53, 56,
62,64, 77,82, 99, 102, 117
30, 37, 47, 57, 80, 96
10, 11, 12, 14,27, 52, 69, 84,
85, 92, 104, 108
32, 41, 42, 43, 44, 45, 46, 49,
50, 54, 95, 114
9, 29, 35, 60, 76, 97, 107
2, 7, 8, 19, 22, 31, 61, 71, 79,
86, 88, 89, 90, 103, 110, 116

Biggs & PriceFilamentous algae in New Zealand

181

70

Fig. 5 Relationship between the

mean conductivity of the water
and geometric mean standing crop
of algae at the site cluster nodes.
The error bars are 1 standard
deviation of each mean and are
included to show the overlap in
cluster nodes along the continua.
The cluster numbers and their
dominant filamentous taxa are
given on the right.

Communities

60""

7-Rhizoclonium sp.
6-C.glomerata

50 "

40"

8 -M.varians
9 - Cladophora sp.
5 - Oedogonium spp, Cladophora sp.

30"

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(ft

20 "
4-Spirogyra

spp., Oedogonlum spp.

10 "
1-U.zonata
3-Splrogyra
spp.
2- Sligeoclonium sp., Phormidium spp.
20

30

40

50

Conductivity26 (mS/m)

regulated Lower Clutha River, Otago, and as an

infrequent component of communities in several
small rivers around New Zealand in summer
(Appendix 2). Although this species was abundant
in the Lower Clutha River it has not been previously recorded in New Zealand.
Taxonomy A variety of synonyms are in use, the
most common being Rhodochorton violaceum
(Kiitz) Drew, Audouinella violacea (Ktttz) Hamel,
and Chantransia hermanii (Roth) Desuaux.
Description Specimens from the Lower Clutha
River are consistent with the description of material
from south-eastern Australia given by Entwistle &
Kraft (1984).
Habitat Audouinella hermanii formed extensive,
fine, burgundy-red coloured mats (like velvet) over
bedrock substrata; often found growing in conjunction with Musci or coating rootlets of crack
willows (Salix fragilus) where these protruded into
flowing waters.
The community formed a cover of 50% and a
standing crop of 15 g/m2 at site 140. It was usually
securely anchored to the substratum and survived
twofold daily changes in flow associated with upA new record for a mat-forming filamentous
stream power generation, as well as regular daily
Rhodophyta species in New Zealand
dewatering in shallow areas during the night and
Audouinella hermanii (Roth) Duby was recorded in the early morning. Extensive communities were
in winter as a filamentous dominant at a site in the found on bedrock in normally deep ( > 3 m), and

Overall, the filamentous taxa most frequently

dominating the communities were Ulothrix zonata
(36 sites), followed by Spirogyra spp. (23 sites)
(Appendix 1). Other filamentous taxa which occasionally dominated the communities included Zygenma sp., Binucleria sp., Vaucheria sp., Mougeotia
sp., and Audouinella hermanii.
A total of 27 unicellular taxa were commonly
found 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 were
ubiquitous. Taxa which were more abundant with
particular filamentous dominants included: Epithemia sorex and Gomphonema olivaceum (Table
3). These two taxa, together with Chamaesiphon
sp., Cocconeis placentula, Gomphonema parvulum,
Rhoicosphenia curvata, and Synedra ulna often
occurred as epiphytes on the larger cells of Cladophora sp., Cladophora glomerata, and Rhizoclonium sp.

182

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

turbid water (16.6 g/m3 suspended solids, 8.4 FTU;

Davies-Colley 1985).
New distribution records for the common algal
Appendix 2 shows new distribution records for all
the common algae collected in the survey, a total
of 2148 new species records and 980 new generic
records. Many of these taxa are cosmopolitan and,
with further collecting, would probably be found
extensively throughout New Zealand. The largest
number of new distribution records were found for

the following species: Synedra ulna, 128; Cymbella

kappii, 110; Cocconeis placentula, 99; Gomphoneis
herculeana, 99; Gomphonema parvulum, 98; Navicula rhyncocephala, 88; and Navicula cryptocephala, 86.

DISCUSSION
A large number of algal proliferations were recorded
in New Zealand streams and rivers in the survey,
many of which had the potential to cause degra-

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Table 3 The sum of of the abundance scores of the common taxa (lines) separated according to whichfilamentoustaxa dominated the sample (columns). Taxa
in brackets are tentative identifications.
Filamentous dominants
A B C D E F G H
Filamentous
A Cladophom sp.
B C. glomerata
C Compsopogon coeruleus
D Melosira varians
B Oedogonium spp.
F Phormidium spp.
G Rhizoclonium sp.
H Spirogyra spp.
I Stigeoclonium sp.
J Ulothrix zonata
Unicellular
Achnanthes lanceolata
A. linearis/minutissima
Chamaesiphon sp.
Closterium sp.
Cymbella kappii
C. minuta
Cocconeis placentula
Diatoma vulgare
Epithemia sorex
Fragilaria vaucheriae
Gomphoneis herculeana
Gomphonema (angustatum)
G. olivaceum
G. parvulum
G. (subclavatum)
G. truncatum
G. truncatum var. capitatum
(Lyngbya sp.)
Navicula cryptocephala
N. rhyncocephala
N. viridula var. avenacea
Nitzschia sp. (a)
N. palea
Rhoicosphenia curvata
Scenedesmus sp.
Synedra ulna
S. ulna var. contracta

_ 13 0 0 13 0 9 12 0
0 - 0 4 0 0 12 0 0
0 5 - 0 0 0 6 4 0
31 12 0 - 28 3 14 0 0

0
4
0

r\

35 10
0 5
0 10
19 10
0 0
0 4

0 20 _ 4 6 u 9 0
0 6 3 - 4 33
10 7 12
6 4 0 0 _ 5 0 0
5 3 29 5 0 - 11 6
0 0 10 5 0 10 - 0
0 5 0 3 0 5 5 _

16
0
0
6
37
5
34
0
21
4
29
5
4
41
0
4
0
4
12
33
10
0
8
9
15
36
6

0
0
0
0
0
0
5
0
3
0
3
0
0
0
0
0
0
0
0
0
0
0
0
3
0
6
0

5
0
6
0
32
4
36
0
52
0
31
5
6
12
12
0
0
6
25
23
4
13
5
3
3
36
12

12
0
0
4
19
13
23
0
5
4
13
3
3
21
4
10
9
0
20
24
9
3
9
3
11
53
10

7
11
0
8
17
10
25
0
11
8
17
0
0
19
4
0
0
0
10
22
0
0
0
0
14
43
10

11
6
0
0
18
11
12
0
0
4
12
0
0
15
0
0
0
0
6
7
3
4
3
3
7
24
7

9
0
12
3
13
5
43
0
5
0
12
0
0
8
0
4
11
0
5
7
0
3
0
12
13
21
0

22
6
0
0
43
18
21
0
12
11
54
0
10
30
0
0
0
7
24
14
0
12
0
10
13
49
6

10
0
0
0
0
15
7
0
0
0
22
0
0
29
0
0
0
11
3
0
0
4
0
0
3
14
0

22
5
3
0
56
24
13
12
5
28
53
11
3
26
6
0
0
3
11
20
17
21
6
6
0
57
39

Biggs & PriceFilamentous algae in New Zealand

dation of water quality and aesthetic values. The
growths smothered river lied sediments and severely
impeded water movement in some rivers (Fig. 2).
The standing crops exceeded most natural maximum values reported in the literature (Table 4).
Only under artificial conditions do the crops
reported elsewhere approach the higher levels
recorded in this survey (Appendix 1). The proliferations occurred even though large communities
of grazing insects were often observed, which would
be expected to have checked their accrual (Eichen-

183
berger & Schlatter 1978). The presence of coarse
cobble substrates, high light intensities, and diffuse
source enrichment is likely to have aided this proliferation of algae at the sites. For example, zones
of seepage from farmlands into braided rivers were
often evident as conspicuous peripheral accumulations of algae or as plumes of filaments from the
banks (Fig. 2). Temperature gradients of up to 10 C
were recorded from the back of some of these zones
to where the cooler seepage water became fully
mixed with the warmer river waters. In contrast,

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

Crop (g/m2) Locality/Treatment

Natural streams

Bahls 1971
Cushing 1967
Cushing et al. 1983

Dunn 1976
Elwood & Nelson 1972
Ertl et al. 1972
Fisher et al. 1982
Grimm et al. 1981
Kobayashi 1961
Liaw & MacCrimmon 1978
Lyford & Gregory 1975
Mulholland et al. 1985
Muller 1978
Murphy et al. 1981
Rose (unpubl. date)
Rounick & Gregoiy 1981
Stockner & Shortreed 1976
Stockner & Shortreed 1978

18.9
4.2
6.5
6.7
18.7
7.5
13.2
9.5
22.6
67.7
0.7
65
160
290
7.0
82.6
8.8
1.8
5.2
2.1
15.3
3.5
2.7
7.5
6.6
4.4

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., Austria
Desert Stm, Arizona, U.S.A.
Desert Stm, Arizona, U.S.A.
Arakawa R., Japan
Grand R., Ontario, Canada
Mountain Stm, Oregon, U.S.A.
Walker Branch, Tennesee, U.S.A.
Trydean R., Sweden
Tolangaan R., Sweden
Verkean R., Sweden
McKenzie R., Oregon, U.S.A.
Shite Oak Ck, Tennesee, U.S.A.
Mountain Stm, Oregon, U.S.A.
Carnation Ck, Vancouver I., Canada
Carnation Ck, Vancouver I., Canada

7.06
40.0
70.5
50.0
250
10.4
13.9
11.1
12.0
14.9
65.7
141.3
107.3
37.5

Outdoor troughs, no enrichment

Outdoor troughs, sewage enrichment
Outdoor troughs, inorganic enrichment
Outdoor troughs, enriched groundwater
Outdoor troughs, herbivores removed
Lab. streams, inorganic enrichment
Lab. streams, organic enrichment
Outdoor troughs
Lab. streams
Outdoor troughs, enriched groundwater
Lab. streams, high light, mod. current
Lab. streams, snails removed
Lab. streams, high light intensity
Outdoor troughs, inorganic enrichment

Experimental streams

Clark et. al. 1979

Eichenberger 1975
Eichenberger 1979
Eichenberger & Wuhrmann 1975
Eichenberger & Schlatter 1978
Erhlich & Slack 1969
Kaufman 1980
Kevern et al. 1966
Marker & Casey 1982
Mclntire 1968a
Mclnitre 1968b
Mclntire & Phinney 1965
Stockner & Shortreed 1978

Downloaded by [114.124.25.25] at 21:29 22 May 2015

184

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

rivers draining native bush catchments had poorly

developed filamentous algal mats.
Cladophora glomerata appears to be the main
component of algal proliferations in the rivers of
North America and Europe (Whitton 1970; Pitcairn & Hawkes 1973). Only occasionally are other
taxa recorded as forming major growths. Prescott
(1981) reports that Rhizoclonium sp. forms "long,
stringy, sometimes ropelike strands in flowing
water" in North America, and Ulothrix zonata has
been recorded forming proliferations below organic
pollution discharges in Europe (Fjerdingstad 1964)
and after forest removal in North America (Likens
et al. 1970). Although Cladophora glomerata dominated a significant number of proliferations in New
Zealand (20%), Rhizoclonium sp. had the highest
average covers and standing crops, and most frequently formed a crop > 50 g/m2. Ulothrix zonata
most frequently formed proliferations of >40%
cover. In contrast to the findings of Fjerdingstad
(1964), the Ulothrix zonata proliferations in New
Zealand were associated with high-quality (i.e., lowconductivity) waters.
Little information appears to be available in the
literature on the tolerance of periphytic taxa to siltation. Moderate siltation of communities was
noted at several sites (particularly in alpine-fed
rivers). Spirogyra spp., Ulothrix zonata, Gomphoneis herculeana, and Melosira varians dominated
these sites, and thus appeared to tolerate limited
silt deposition. These taxa are all characterised by
their high growth stature which would enable their
photosynthetic portions to remain above the zone
of basal silt accumulation. Species which often
occurred among the filaments of these communities included Synedra ulna, Cymbella kappii, Cymbella minuta, Fragilaria vaucheriae, and
Gomphonema parvulum.
Many of the taxa commonly encountered in the
survey displayed habitat preferences consistent with
their reported ranges in North America and Europe.
For example, Diatoma hiemale var. mesodon and
Ulothrix zonata are also found dominating cold
water mountain rivers in Europe (Blum 1956). The
only major difference occurred for Ulothrix zonata
which competes most successfully with other periphyton in unenriched streams in New Zealand, but
in Europe has both a clean-water form and a pollution-tolerant form (Fjerdingstad 1964; BensonEvans et al. 1975).
The relationship between water conductivity and
standing crop of the community types was close
(Fig. 5). Future work will need to define the
nutrients that are actually limiting algal development in New Zealand rivers and how long a period
of base flow is required (with different nutrient
levels) for proliferations to develop. This type of
information should enable the likely occurrence and

extent of proliferations to be predicted, thus aiding

the management of our river systems.

ACKNOWLEDGMENTS
We are grateful to our colleague Mr M. Close for valuable
discussion of the survey data, and to Dr P. A. Broady
(University of Canterbury), Dr J. M. Quinn (Water Quality
Centre, Ministry of Works and Development (MWD),
Hamilton), and Ms B. Vaile (Hydrology Centre, MWD,
Christchurch) for their constructive reviews. The assistance of Dr A. A. Davoren (Hydrology Centre, MWD,
Christchurch) with the SAS clustering procedures and the
verification of Audouinella hermanii by Dr T. J. Entwistle
(University of Melbourne) are also gratefully
acknowledged.

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Biggs & PriceFilamentous algae in New Zealand

187

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

Cover
(%)

AFDW
(g/m!)

7.17
8.37
15.23
10.64
11.51
17.79
6.48
22.34

10
5
5
20
5
90
80
80

30
85
50
95
5
30
20
10
30
10
15
50
5
95
15
5
10
5

30
108
24
185
1
13
2

23
14
19
17
17
16

25.45
31.04
28.52
34.80
8.90
25.34
17.53
10.59
4.43
16.56
15.75
19.52
4.30
15.36
6.89
6.48
10.19
9.00

9
2
24
10
99
0.3
94
15
8
8
1

2
3
1
1
2

F
F

16
22

13.76
14.84

40
40

13
19

2
3

N21:39:38
N32:14:62
N31:17:73
N32:91:79
N32:44:29
N32:27:17
N21:29:27
N23:80:47
N13:99:56
N18:85:02
N18:02:59

F
F
F
F
F
F
F
F
F

18
_
15
16
15
15
18
16
21

90
90
20
80
35
25
55
15
10

31
59
50
82
8
3
70
3
5

2
3
2
3
1
1
2

19
21

24.44
20.60
23.25
28.50
13.29
10.81
23.84
8.10
13.69
23.53
13.71

Manganui
Mangaone
Mangaone
Mangaone
Mangaonuku

N13:86:84
N32:09:38
N43:13:41
N43:13:41
N32:96:92

F
F
F
F
F

13
18
19
17
19

11.53
46.64
35.40
32.25
29.31

30
95
90
95
100

10
264
101
168
84

3
3
3
3
3

45
46
47
48
49
50
51

Mangaonuku
Mangaoraka
Mangapiko
Mangatangi
Mangawhero
Maraekakaho
Matakitaki

N32:97:89
Nl 3:76:92
N25:47:78
N26:69:17
N23:78:04
N43:05:20
S57:75:66

F
F
F
F
F
F
F/A

18
18
17
23
19
21
15

18.43
19.70
20.30
9.90
14.13
44.70
8.64

15
65
50
90
65
84
5

5
125
57
22
26
291
0.1

2
3
3

52
53
53
55
56

Ngaruroro
Ohariu
Okawa
Okuku
Opihi

N43:19:27
N21:29:29
N43:17:28
S55:86:91
S44:41:88

F
F
F
F
F

17
19
17
17
14

13.79
28.06
58.25
9.76
6.44

55
20
80
20
50

47
20
63

3
4

6
10

57
58
59
60
61
62
63
64

Oroua
Oturere
Pakuratahi
Pauatahanui
Percival
Piakonui
Poutu
Puketotara

N32:11:52
N33:25:78
N21:72:47
N21:48:42
S66:18:75
N35:12:52
N33:30:93
N18:43:55

F
F/A

24
10
16
17
14
19
12
-

11.76
12.21
8.81
16.77
10.52
11.00
10.77
-

5
35
90
75
40
80
20
60

1
31
41
12
75
58
18
70

3
3
3
1
3

65
66
67
68
69
70

Rakaia
Rangitata
Rangitata
Rangitikei
Ruakokoputuna
Ruamahanga

S55:14:58
S45:82:04
S45:92:93
N33:32:08
N21:98:32
N31:06:45

A
A
A
F/A

13
19
16
18
18
18

6.55
5.54
6.94
13.20
39.48
13.23

5
5
20
20
60
35

1
3

82
5
60
10

1
1
1
3
4
2

71
72
73
74
75

School Ck
Selwyn
Speargrass
Station
Tauherenikau

S66:21:48
S55:37:03
S67:15:69
S57:65:19
N21:88:39

F
F
F
F
F

17
13
17
13

90
5
5
25
15

30
4
0.4
16
3

3
1
1
1

18

21.58
8.85
4.13
12.75
7.68

S77:24:99
N35:49:44

F
F

14
18

13.84
5.84

5
25

1
7

2
3

NZMS-18 map
reference

Class

Cl

Summer survey
Ashburton (Nth)
Ashley
Awatere
Camping Gully
Clarence
Cla rence
Cobbolds
Conway

S55:03:48
S65:06:87
S77:34:81
S55:16:57
S77:18:20
S57:86:81
S66:42:28
S66:72:67

A
F
A
F
A
F
F
F

15
16
24

Doughboy
Esk
Flaxboume
Gorge
Hae Hae Te Moana
Hangaroa
Hapuku
Hope
Hope
Horokiwi
Horomanga
Hoteo
Howard
Huddlestones
Hurunui
Hurunui
Hutt
Hutt

S57:71:68
N43:25:52
S77:40:62
N43:01:25
S44:75:86
N44:98:31
S67:99:02
S56:85:71
S67:02:79
N21:46:46
N44:21:75
N27:03:17
S67:08:75
S67:87:94
S66:12:36
S66:42:28
N21:51:39
N21:63:44

L
F
L
F
F
F
F
A
F

14
20
22
25
11
23
15
16
17
19
19

F
A
A
F
F

27 Irrig. Canal
28 Kaeo

N43:09:21
N 18:29:71

29
30
31
32
33
34
35
36
37
38
39

Kenepuru
Kiwitea
Kopuaranga
Maharakeke
Makairo
Makakahi
Makara
Makotuku
Makuri
Mangakino
Mangamuka

40
41
42
43
44

Site
no. River

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1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26

76 Taylor
77 TeAhura

Temp

F
F
F
F

F
F

F
F
F
L
F

F/A

13
18
20
17
22

20

5
2
17

3
108
110
118

n Dominant taxa
1
2
3
4
3
2
2
2
1
3
2
3
2
3
1
1
3
1

1
1
2

3
3
2
2

3
3
1
1

3
2
3

Ulothrix zonata
Cladophora sp.
Spirogyra spp.
Phormidium spp.
Spirogyra spp.
Ulothrix zonata
Zygnema sp.
Zygnema sp.,
Cladophora sp.
Vaucheria sp.
Cladophora glomerata
Cladophora glomerata
Cladophora glomerata
Phormidium spp..
Cladophora glomerata
Spirogyra spp.
Ulothrix zonata
Spirogyra spp.
Stigeodonium sp.
Cladophora sp.
Cladophora sp.
(Lyngbya sp.)
Cladophora sp.
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Spirogyra spp.,
Stigeodonium sp.
Cladophora glomerata
Binucleria sp.,
Spirogyra spp.
Melosira varians
Cladophora sp.
Cladophora sp.
Rhizodonium sp,
Cladophora glomerata
Cladophora glomerata
Spirogyra spp.
Spirogyra spp.
Oedogonium spp.
Compsopogon coeruleus
Cladophora sp.,
Oedogonium spp.
Spirogyra spp.
Rhizodonium sp.
Cladophora sp.
Compsopogon coeruleus
Melosira varians,
Rhizodonium sp.
Rhizodonium sp.
Rhizodonium sp,.
Oedogonium spp.
Spirogyra spp.
Rhizodonium sp.
Rhizodonium sp.
Melosira varians,
Ulothrix zonata
Cladophora glomerata
Spirogyra spp.
Rhizodonium sp.
Spirogyra spp.
(Lyngbya sp.),
Spirogyra spp.
Melosira varians
Ulothrix zonata
Ulothrix zonata
Melosira varians
Cladophora sp.
Oedogonium spp.
Binudearia sp.
Spirogyra spp.,
Zygnema sp.
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Cladophora glomerata
Cladophora glomerata,
Cladophorag lomerata,
Vaucheria sp.
Phormidium spp.
Spirogyra spp.
Ulothrix zonata
Spirogyra spp.,
Stigeodonium sp.
Melosira varians
Spirogyra spp.

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

188
Appendix 1 (Continued).

NZMS-18 map
reference

Cl

Cond.
mS/m

Cover

Class

Dominant taxa

78 Temuka

S44:81:69

16

8.89

80

48

79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96

Tengawai
Toko
Tongariro
Topehahae
Tukipo
Turakina
Tutaekuri
Victoria
Waiau
Waihi
Waihopai
Waihuka
Waikanae
Waikari
Waikawa
Waimana
Waingongoro
Wainui

S44:66:70
N13:98:57
N34:30:01
N35:04:55
N32:92:81
N22:72:74
N43:21:37
N08:92:64
S66:58:52
S44:78:89
S77:03:98
N54:15:63
N21:59:71
N43:43:83
N21:75:94
N45:52:02
N13:85:47
N18:12:76

F
F
F/A

18.59
16.49
10.77
12.59
14.74
47.80
30.12
13.93
8.34
8.52
6.85

60

42
72
17
27
2
93
98
31
2
17

F
F
F
F
F
F

16
20
13
20
17
19
19
20
20
20
17
_
17
19
18
19
18

22

10.11
41.85
8.34
8.43
12.20
17.25

40

20
1
42
3
6
17
23

1
3
3
3
1
4
4
3
2
2
1
3
3
3
3
3
3
3

97 Wainuiomata

N21:46:16

17

15.29

70

17

98 Waioeka
99 Waiohine
100 Waiopoua

N45:71:ll
N21:99:49
N31:15:64

F
F
F

_
17
14

6.38
7.98

25
-

7
_
1

3
1
2

101
102
103
104

Waiotahi
Waipa
Waipara
Waipaoa

N45:62:05
N24:72:98
S66:ll:12
N54-.29-.56

F
F
F
F

19
19
19
21

7.26
7.26
34.40
50.21

20
90
30

1
6
27
88

2
3
2
3

Binuclearia sp.,
Phormidium spp.
Melosira varians
Melosira varians
Ulothrix zonata
Oedogonium spp.
Cladophora glomerata
Rhizoclonium sp.
Cladophora glomerata
Cladophora sp.
Ulothrix zonata
Oedogonium spp.
Cladophora sp.
Cladophora sp.
Stigeodonium sp.
Cladophora glomerata
Stigeodonium sp.
Spirogyra spp.
Rhizoclonium sp.
Cladophora sp.,
Oedogonium spp.
Melosira varians,
Oedogonium spp.
Spirogyra spp.
Ulothrix (variabilis)
Microspora sp.,
Stigeodonium sp.

105 Waipawa
106 Waipawa

N32:88:95
N42:02:83

F
F

18
20

9.77
15.30

5
5

3
11

1
1

107
108
109
110
111
112
113
114
115
116
117

S5 5:36:02
N44:85:ll
N35:53:48
Nl 7:61:96
Nl 3:83:93
N13:71:82
N24:85:13
N23:56:41
S57:64:05
N 18:23:49
N45:43:02

F
F
F
F
F
F
F
F
F
F
F

16
20
16
21
20
15
20
20
13
22
19

14.70
27.81
5.58
15.67
10.74
15.43
9.52
12.03
5.14
18.51
10.59

100
80
80
65
40

117
51
27
15
22
7
8
6
55
13

1
3
1
2
3
3
3
3
2
3
3

S65:O6:87
S65:O6:87
S32:32:77
S56:12:17

F
F
F
F

8
6
2
7

8.08
5.77
3.50
7.45

122 Cass

S56:22:18

4.26

123
124
125
126

Clarence
Clarence
Countess
Craigiebum

S67:12:16
S66-.20-.07
S66:21:55
S56:23:05

LK
LK
F
F

_
7
10
6

3.21
10.31
4.42

127
128
129
130
131

Fraser
Glentui
Grays
Hanmer
Hororata

S33:13:49
S55:70:94
S44:05:71
S66:16:72
S55:40:57

LK
F
F
F
F

3
4

13.12

10
9

9.06
6.46

132
133
134
135
136
137
138
139
140
141
142

Hurunui
Jacks
Jollies
Kowai
Kowai
Kowhai
Lower Clutha
Lower Clutha
Lower Clutha
Lower Clutha
Maruia

S66:12:36
S66:20:06
S66:20:06
S55:33:81
S65:ll:97
S66:85:99
S32:28:59
S32:29:85
S32:21:96
S32:21:96
S56:81:98

A
F
F
F
F
F
LK
LK
LK
LK
A

7
6
7
7
9
_
7
6
5
5
-

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Site
no. River

118
119
120
121

Wairiri
Wairoa
Wairoa
Wairua
Waitara
Waiwhakaiho
Wanganui
Wanganui
Warwick
Whakanekeneke
Whakatane
Winter survey
Ashley
Bealey
Beaumont
Bruces

Temp

F
F
F
F
A
F

(%)

75
75

55
10

85

50
20

20

75
10
75
15
10

25

65
5
45
20
75
45

AFDW
(g/m!)

10

0.2
1
0.4
0.3

0.4

30

0.5
_

3
3
5
3

0.3

3
3
3

65
5
80
5

_
1
68
0.7
14

2
6
3
3
3

6.34
2.24
3.69
6.18
16.44

5
35
30
5
80

1
8
3
0.3

7.24
7.37
7.87
7.42

10
20

3
3
3
3
3
3
5
5
6
3
2

95
20

95

12.0

45
20

_
1
8
15
30
-

50

143 Maruia-side ck

S57:66:01

70

144 Pahua
145 Percival

S66:19:45
S66:18:75

F
F

8
10

9.54
6.09

50
5

3
0.1

3
3

146
147
148
149
150
151
151
152

S66-.21-.48
S55:37:03
S32:33:53
S66:38:62
S56:22:21
S66:ll:12
S66:ll:12
S66:11:34

F
F
F
A
A
F
F
F

12
8
3
9
8
7
8
9

20.38
8.18
10.94
7.41
5.08
21.29
16.67
40.40

100
70

3
3
5
3
3
3
3
2

School Ck
Selwyn
Tuapeka
Waiau
Waimakariri
Waipara
Waipara
Wait

5
5

40
25
85
100

35

2
1
5
2
12
-

Stigeodoniumsp.Stigeodonium s p .

Spirogyra spp.
Cladophora sp.
Cladophora glomerata,
Rhizoclonium sp.,
Compsopogon coeruleus
Melosira varians
Cladophora glomerata,
Rhizoclonium sp.
Vaucheria sp.
Cladophora glomerata
Phormidium spp.
Cladophora sp.
Spirogyra spp.
Spirogyra spp.
Phormidium spp.
Rhizoclonium sp.
Stigeodonium sp.
Oedogonium spp.
Spirogyra spp.
Ulothrix zonata
Ulothrix zonata
Phormidium spp.
Diatoma hiemale
var. mesodon
Diatoma hiemale
var. mesodon
Spirogyra spp.
Ulothrix zonata
Ulothrix zonata
Diatoma hiemale
var. mesodon
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Diatoma hiemale
var. mesodon
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Melosira varians
Ulothrix zonata
Ulothrix zonata
Vaucheria sp.
Audouinella hermanii
Oedogonium spp.
Diatoma hiemale
var. mesodon
Diatoma hiemale
var. mesodon
Ulothrix zonata
Diatoma hiemale
var. mesodon
Ulothrix zonata
Ulothrix zonata
Phormidium spp.
Ulothrix zonata
Ulothrix zonata
Ulothrix zonata
Stigeodonium sp.
Ulothrix zonata

Biggs & PriceFilamentous algae in New Zealand

Appendix 2 Taxa of algae and the sites at which they were found, during the survey of New Zealand
rivers. Taxa preceded by ? are tentative designations. River names for each site are given in Appendix
1. * Denotes that the occurrence is not a new distribution record.
Taxon

Sites

Chlorophyta

Ankistrodesmus falcatus (Corda) Ralfs

Binuclearia sp.
Bulbochaete sp.
Characium sp.
Cladophora sp.
Cladophora glomerata Kiitz
Cladophora ?oligoclona Kutz
Closterium sp.

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Coelastrum sp.
Cosmarium sp.
Cylindrocapsa sp.
Dictyosphaerium sp.
Geminella crenulatocollis Prescott
TGloeocystis sp.
Microspora sp.
Mougeotia sp.
Oedogonium spp.
Pediastrum sp.
Rhizoclonium sp.
Scenedesmus sp.
Schroederia sp.
Selenastrum sp.
Spirogyra spp.

Staurastrum sp.
Stigeoclonium Tftagelliferum Ktitz
Stigeoclonium sp.
Stigeoclonium ?pachydermum Prescott
Ulothrix aequalis Kiltz
Vlothrix tenuissima Kiitz
Ulothrix variabilis Kutz
Vlothrix zonata (Weber & Mohr) Ktltz

Zygnema sp.
Chrysophyta

Dinobryon sp.
Tribonema sp.
Vaucheria sp.
Bacillariophyta

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, 128
7, 17, 28, 30, 37, 58, 63, 67, 77, 78, 97, 102, 103, 105
140
3, 11, 12, 17,23, 32, 115
2, 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, 117
5, 8, 10, 11, 12, 14, 27, 33, 34, 44, 52, 53, 57, 69, 70, 83, 84, 85, 92, 104, 105,
106, 108
42
4, 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, 110
13, 55, 71, 79, 88
2, 4, 5, 8, 11, 17, 25, 26, 41, 55, 56, 59, 61, 73, 75, 79, 87, 88, 94, 97, 100, 115
7, 55, 71, 89, 97
17, 55, 76, 79
44, 106
2, 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, 152
13, 17, 22, 55, 56, 73, 76, 94, 100, 115, 141, 143
3, 17,25, 55,65, 93, 99, 115, 128
2, 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, 141
11, 35, 55,61, 79,88
10, 12, 31, 32, 35, 40, 41, 42, 43, 44, 45, 46, 49, 50, 53, 54, 69, 84, 85, 95, 104,
105, 106, 114
2, 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, 116
3, 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, 128
36
2, 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, 117
2, 4, 13, 17, 55, 78, 79, 105, 128
56
2, 17, 18, 20, 25, 26, 29, 35, 45, 56, 58, 59, 68, 69, 70, 75, 91, 93, 96, 100, 101,
115, 128, 136, 151, 152
56, 61, 93, 97
58, 128
25, 44, 59, 75, 81
17, 59,99, 115
1, 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, 153
2, 7, 8, 16, 19, 24, 25, 45, 55, 59, 64, 94, 103, 105, 115, 123, 137
64
55, 115
9, 22, 71, 107, 138, 139, 140, 148

4, 5, 7, 17, 19, 23, 51, 76, 87, 106, 128

6, 17,24, 51, 71, 115
1, 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, 137
Achnanthes linearis (W. Smith) Grun
4, 13, 25, 26, 29, 35, 53, 59, 61, 88, 91, 97, 128
Achnanthes minutissima Kutz
12, 44, 59, 106
51, 68, 95, 120
Amphora sp.
23, 109
Asterionella formosa Hassall
43, 52, 54, 63, 98
Cocconeis sp.
1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 17, 19, 20, 21, 22, 24, 25, 26, 27,
Cocconeis placentula Ehr
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, 143
19, 29, 30, 35, 57, 60, 65, 90, 95
Cyciotella sp.
4, 13, 31, 44, 61, 71, 76, 79, 92, 106, 107, 108
Cyclotella menhigiana Ktitz
6, 23, 24, 88, 89, 123
Cyciotella stelligera Cleve & Grun.
2, 16, 22, 26, 33, 47, 58, 61, 63, 69, 91, 96, 102, 106, 111, 115, 123, 128, 143
Cymbella sp.
13, 15, 55, 61, 79, 88, 98, 100, 103, 105, 128
Cymbella aspera (Ehr.) Cleve
67, 88
Cymbella cesatii (Robh.) Grun
8, 23, 45, 59, 103, 107
Cymbella ?cistula (Hempr.) Grun
Cymbella cymbiformis (Ktttz) Van Heurok 44, 68, 98, 103, 128
Achnanthes sp.
Achnanthes Tdelicatula (Kutz) Gnin
Achnanthes lanceolata (Breb.) Grun

189

190

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

Appendix 2 (Continued).
Taxon

Sites

Cymbella kappii Cholnoky

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, 153
1, 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, 147
1, 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, 143
1, 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, 67
19, 112
2, 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, 137
5, 14, 17, 24, 61, 63, 98, 108, 128, 140

Cymbella minuta Hilse ex Rabh.

Cymbella sinuata Greg
Diatoma hiemale var. mesodon (Ehr)
Grun

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Diatoma vulgare Bory

Epithemia sp.
Epithemia sorex Kiitz
Epithemia zebra var. saxonica
(Kutz) Grun
Eunotia sp.
Fragilaria sp.
Fragilaria capucina Desmaz
Fragilaria crotonensis (Ehr) Grun
Fragilaria vaucheriae (KUtz) Petersen
Fragilaria virescens var. capitata Ostr
Frustulia rhomboides (Ehr) De Toni
Frustulia vulgaris (Thwaites) De Toni
Gomphoneis elegans (Grun) Cleve
Gomphoneis eriense (Grun) Skv & Meyer
Gomphoneis herculeana (Ehr) Cleve

Gomphoneis herculeana var. clavata

Cleve
Gomphoneis herculeana var. robusta
(Grun) Cleve
Gomphonema sp.
Gomphonema accuminatum Ehr
Gomphonema ?anguslatum (Kiitz) Rabh
Gomphonema dichotomum Kiitz
Gomphonema intricatum var. v/Z?ro
(Ehr) Cleve
Gomphonema olivaceum A-V.H.
Gomphonema parvulum (Kiitz) Grun

Gomphonema ?subclavatum (Grun) Grun

Gomphonema truncatum Ehr
Gomphonema truncatum var. capitatum
(Ehr) Patr
Gyrosigma accuminatum (Kutz) Rabh
Melosira granulata (Ehr) Ralfs
Melosira granulata var. angustissima
O. Muller
Melosira varians Ag

Meridian circulare (Grev) Ag

Navicula sp.
Navicula capitata Ehr
Navicula cryptocephala Kutz

Navicula cuspidata Kutz

19, 36, 37, 47, 80, 111, 114, 143

4, 8, 12, 27, 28, 30, 37, 41, 45, 47, 53, 57, 58, 68, 71, 77, 86, 88, 94, 95, 102,
110, 113, 115. 128
4, 83
12, 13, 128, 131
1, 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, 151
1, 13, 66, 78, 79
4, 23, 35, 37, 51, 58, 115, 120, 138, 139, 140, 141
9, 61, 115, 128
81
128
1, 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, 153
91
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,
112,113,114,115,128,144
64, 71, 79, 96, 107, 116
1, 3, 6, 21, 29, 30, 31, 33, 41, 51, 55, 64, 68, 74, 76. 83, 84, 87, 88, 93, 115, 128
3, 4, 5, 6, 8, 12, 13, 17, 21, 24, 52, 55, 61, 66, 67, 69, 71, 77, 78, 79, 89, 98,
105, 115, 129
1, 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, 151
1, 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, 129
1, 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, 137
2, 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, 137
12, 14, 32, 44, 45, 50, 52, 54, 71, 75, 92, 106, 107, 108
1, 31, 34, 44, 83, 87
87, 120, 129, 139
63
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, 148
21, 100, 115, 120, 129
7, 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,
111,112,128
35, 43, 61, 69, 90, 92
1, 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, 89
90, 91, 92, 93, 95, 97, 98, 102, 103, 104, 105, 107, 111, 113, 115, 116, 128, 129,
136, 137, 142
7, 8, 29, 57, 104

Biggs & PriceFilamentous algae in New Zealand

Appendix 2 (Continued).
Taxon

Sites

Navicula decussis Ostrup

Navicula exigua (Greg) Grun
Navicula pupula Kfitz

2, 3, 4, 5, 10, 11, 15, 16, 24, 27, 72, 74, 89

8, 25, 28, 31, 57, 83, 84, 86, 87, 89, 97, 98, 116
6, 7, 10, 13, 17, 25, 26, 33, 41, 43, 44, 55, 58, 63, 71, 74, 76, 86, 88, 89, 93, 97,
102, 115, 129, 137
4, 13,21,29, 55, 56, 60, 65, 88, 128
2, 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, 143
1, 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, 137
7, 22, 23, 27, 31, 35, 39, 46, 58, 81, 84, 90, 95, 96, 102, 110, 113, 114, 116, 128,
129, 131, 140, 146, 147, 152
2, 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, 148
44, 106, 128
22,35, 74, 115
1, 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, 140
2, 5, 6, 8, 17, 31, 51, 53, 59, 68, 70, 87, 99, 100
2, 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, 128
2, 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, 128
37, 39, 52, 61, 67, 80, 96, 107, 115, 120, 128, 129, 139
30, 35, 81, 88
1, 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, 137
58, 61, 87
5, 7, 8, 11, 61, 63, 81, 87, 98, 108, 123, 129, 137, 140
41, 92, 107, 108, 120, 128
17, 22, 35, 70, 72, 73, 76, 89, 100, 107, 115
79, 128
9, 12, 15, 22, 26, 29, 31, 33, 35, 49, 51, 69, 71, 76, 95, 97, 100, 128, 129
1, 2, 13, 16, 24, 34, 61, 65, 66, 69, 71, 72, 88, 113, 114, 127, 128
3, 5, 6, 16, 23, 24, 33, 46, 61, 63, 67, 72, 73, 74, 75, 76, 98, 115, 123, 127, 128
4, 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, 137
5,8, 16,24,30, 57,63,67,79,88, 111
13, 23, 42, 55, 61, 66, 67, 71, 85, 87, 105, 128
1, 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, 153
2, 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, 129
7, 10, 13, 14, 17, 22, 27, 42, 43, 55, 56, 69, 78, 79, 85, 88, 90, 92, 98, 104, 128,
129
37, 44
17, 63, 115, 123, 127, 129, 143

Naviada pusio Cleve

Navicula radiosa KUtz
Navicula rhyncocephala Kutz

Navicula viridula Kiitz

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

Downloaded by [114.124.25.25] at 21:29 22 May 2015

Nedium sp.
Nedium affine (Ehr) Cleve
Nitzschia spp. Hassall
Nitzschia (disspiata) (Kutz) Grun
Nitzschia ?linearis W. Smith
Nilzschia palea (Kutz) W. Smith
Pinnularia sp.
Rhoicosphenia adolfi M. Schmidt
Rhoicosphenia curvata (Kiitz) Grun

Rhopalodia sp.
Rhopalodia ?novae zelandiae Hust
Surirella sp.
Surirella angusta Kiitz
Surirella Hnearis W. Smith
Surirella ovata Kutz
Synedra sp.
Synedra acus Kiitz
SynedraacusKiitzSynedra ?minuscula G r u n
Synedra radians Kutz
Synedra ?rumpens Kiitz
Synedra ulna (Nitzsch) Ehr

Synedra ulna ?var. contracta Ostrup

Synedra ulna ?var. ramesi (Herib)

Hust
Tabellaria sp.
Tabellariaflocculosa(Rabhi) Kutz
Rhodophyta

Audouinella hermanii (Roth) Duby

Compsopogon coeruleus (Balb) Montagne

34, 56, 78, 88, 129, 138, 139, 140, 141

20, 28, 32, 38, 41, 43, 78, 102, 104, 140

Cyanobacteria

Anabaena sp.
Calothrix sp.
Chamaesiphon sp.
Chroococcus sp.
Gloeotrichia sp.
Merismopedia sp.
Nostoc sp.
Phormidium spp.

Schizothrix sp.
Spirulina sp.

1, 2, 3, 5, 6, 7, 8, 11, 13, 16, 17, 23, 28, 37, 39, 40, 45, 53, 55, 58, 61, 64, 68, 69
78, 79, 81, 83, 86, 88, 89, 90, 95, 96, 97, 98, 100, 103, 105, 115, 116, 117, 128
17, 61, 103
2, 17, 32, 34, 45, 49, 51, 55, 56, 58, 70, 81, 84, 88, 91, 95, 100, 114, 123, 128
13, 55, 61, 79, 88
21, 36, 57, 58, 84, 93
2, 11, 15, 17, 21, 35, 55, 56, 61, 74, 76, 83, 91, 93, 100, 115, 128
2, 5, 13, 55, 67, 128
3,61
1, 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, 152
17, 140
3, 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

191