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Is the breeding distribution of

Dippers influenced by stream
acidity?
a b c
S. J. Ormerod , Stephanie J. Tyler & J. M. S. Lewis
a
Department of Applied Biology , University of Wales
Institute of Science and Technology , King Edward VII Avenue,
Cardiff, CH1 3NU
b
RSPB Wales Office , Frolic Street, Newtown, Powys, SY16
1AP
c
High Leas, Coldbrook, Abergavenny, Gwent
Published online: 24 Jun 2009.

To cite this article: S. J. Ormerod , Stephanie J. Tyler & J. M. S. Lewis (1985) Is the
breeding distribution of Dippers influenced by stream acidity?, Bird Study, 32:1, 32-39, DOI:
10.1080/00063658509476852

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 Bird Study (1985) 32, 32-39

Is the breeding distribution of Dippers

influenced by stream acidity?
S.J. ORMEROD, STEPHANIE J. TYLER and J.M.S. LEWIS
Downloaded by [North Dakota State University] at 16:12 29 October 2014

There is growing concern amongst conservationists about the consequences

of 'acid rain', though this is not the only way that watercourses can
increase in acidity. This study shows that fewer Dippers are present on
acidic reaches of rivers, probably because changed water quality affects
the birds' food supply.

oncern has been expressed recently over
C the acidity of streams and rivers in
some upland regions of the United Kingdom
(e.g. Harriman & Morrison 1982; Stoner et al.
1984; Fry & Cooke 1984). From these and
many other studies (e.g. Sutcliffe & Carrick
1973; Townsend et al. 1983) it is known that
acidic water courses support impoverished
macroinvertebrate faunas by comparison
with physiographically similar water courses
which are less acidic. Dippers Cinclus cinclus
are dependent on aquatic macroinvertebrates
as a food supply (Creutz 1966; Ormerod,
in press); therefore, it is conceivable that
surface-water acidity might indirectly
influence their distribution. An examination
of this possibility seems desirable in view of
the current interest in this topic (e.g. Haines
1981).
Whilst there has been little temporal
change in the numbers of Dippers breeding
on British census plots since at least 1974
(Taylor & Marchant 1982), there are striking
spatial differences in breeding abundance
both between regions (Sharrock 1976;
Marchant & Hyde 1980) and within them
(Mawby 1961; Round & Moss 1984). Rela-
tionships have been proposed between
breeding abundance and altitude (Sharrock
1976; Shaw 1978; Marchant & Hyde 1980),
river gradient (Marchant & Hyde 1980;
Round & Moss 1984) and the availability of
shallow riffles (Robson 1956; Shooter 1970).
With the exception of a qualitative study by
Da Prato & Langslow (1976), there are few
published data on relationships between
breeding distribution and prey availability.
Recent indications from the River Wye were
that breeding Dippers were absent from the
upper reaches which were known to have
low densities of aquatic macroinvertebrates
(Brooker & Morris 1980; Round & Moss
1984).
This paper describes the breeding abun-
dance of Dippers (as the number of pairs per
10 km of river) in 1982 in the catchment of
the Welsh River Wye and on the Grwyne
Fawr (an adjacent tributary of the River Usk)
in relation to dietary requirements, prey-
abundance and some environmental factors.
Data on breeding abundance are also em-
ployed from earlier studies on the same river
system (RSPB 1977; Round & Moss 1984).
The study area has been described else-
where (Edwards & Brooker 1982; Ormerod et
al., in press).
METHODS
Abundance of breeding Dippers
Breeding abundances on most tributaries
were assessed using methods similar to the
BTO Waterways Bird Survey (Marchant &
Hyde 1979). Over 180 km of tributaries were
visited on at least seven occasions between
9 March 1982 and 7 July 1982, and three
registrations was the minimum criterion to

32

confirm Dipper territories. Breeding abun-
dances on the Rivers Monnow, Honddu and
Grwyne Fawr were assessed from fewer
visits to territories which had been occupied
by actively breeding birds in previous years
(Tyler, unpublished data). Territory locations
were plotted on 1: 50,000 OS maps. Breeding
abundances were calculated firstly over each
10 km stretch from the source to the mouth
of each tributary and, secondly, over reaches
5 km upstream and downstream from 23
sites at which macroinvertebrate abundances
were also assessed. Similar data on breeding
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distribution over a further 50 km were
drawn from Round & Moss (1984) for the
upper Wye and from the RSPB (1977) for the
upper Wye, the River Tarenig and the River
Elan. The latter figures were used in con-
junction with macroinvertebrate data from
five further sites sampled in 1982.
Environmental variables
Altitudes and gradients were assessed for
each surveyed reach using 1: 50,000 OS
maps. Mean pH values, based in each case
on at least 15 determinations in different
seasons, were provided by the Welsh Water
Authority, together with historical data on
the pH of some rivers of contrasting water
chemistry.
Macroinvertebrate abundances
Macroinvertebrates were sampled in March—
April 1982, at each of 28 sites, by one oper-
ator using a standardized kick-sampling
technique (2 x 3 minutes per site; net mesh
aperture 440 p.m). Abundances for each
macroinvertebrate order (numbers of indivi-
duals per sampling interval) were deter-
mined in the laboratory. This method gave
abundance estimates which correlated
strongly with values from more quantitative
techniques (e.g. Armitage et al. 1974).
Dietary analysis
The diet of adult and nestling Dippers was
assessed at 16 sites throughout the study
area by the analysis of 72 and 75 faecal
pellets respectively. These were collected in
Dippers and stream acidity 33
1983, either during ringing operations or
from stones in the vicinity of nests. After the
deflocculation and examination of faecal
pellets at x40, macroinvertebrate prey-items
could be identified to family and quantified
by counting mouthparts. Fish remains were
more difficult to quantify, and it was as-
sumed that the presence of fish bones in any
faecal pellet indicated that one fish had been
ingested. Probably this resulted in an over-
estimate, since no pellet contained more than
a few fish vertebrae.
Prey-weight contributions were estimated
for each type of prey item by multiplying the
recorded number of faecal occurrences by a
mean dry weight. For macroinvertebrates,
the latter were derived for each family from
field collections (n = 10-70 individuals). For
fish, a dry weight (220 mg) equivalent to
that of a brown trout Salm° trutta of length 7
cm was derived from data given by Milner et
al. (1978). Total dietary dry weights were
then converted to calorific values using
tables provided by Cummins (Si Wuycheck
(1971).
RESULTS
Abundance of breeding Dippers and
environmental variables
On the Rivers Monnow, Honddu, Grwyne
Fawr (Black Mountains rivers) and Ithon, the
abundance of breeding Dippers was greatest
on upper reaches which had steep gradients
(15-30 m/km), and the first occupied terri-
tory occurred within 1-2 km of the source
(Figure 1). However, the upper reaches of the
Rivers Irfon and Wye held few or no breed-
ing Dippers, despite the presence of suitable
gradients, and the first occupied territories
were 8 and 15 km from their respective
sources (Figure 1). These latter rivers had, in
their upper reaches, low pH (means: 6.16,
6.38; long-term minima: 4.5) and soft waters
(mean total hardnesses 10-12 mg CaCO 3 /1)
by contrast with the upper reaches of the
River Ithon and the rivers draining the Black
Mountains (pH means: 7.51-8.20; minima:
6.8-7.3. Mean total hardnesses 60-150 mg
CaCO 3 /1). There was also some evidence of a
historical fall in the pH of the upper Irfon,
 34 S.J. Ormerod et al.
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Figure 1. The distribution of breeding Dippers on five rivers in the Wye catchment and on
the adjacent Grwyne Fawr.
The upper figure shows the altitudinal profile and the symbols indicate the position of the
nearest occupied territory to the source. The histogram represents the abundance of breeding
Dippers in each 10 km stretch.
and of some tributaries of the upper Wye,
which was not apparent for the upper Ithon
(Figure 2).
Overall, the abundance of breeding
Dippers, on the upper reaches of the 17
tributaries for which data were available,
showed a strong correlation with mean pH
(Figure 3). For the complete data-set, breed-
ing abundance showed no significant corre-
lation with either altitude (r = —0.22, ns) or
gradient (r =- 0.09, ns), although there was
a significant correlation with the latter
(r = 0.70, n = 21; P < 0.001) after the removal
from the analysis of data from the acidic
5
1
k
rwy ne Fawr
2
Monnow
10 20 30 40 50 60
reaches of the upper River Irfon and the
upper Wye.
Abundances of Dippers and
macroinvertebrates
Some macroinvertebrate groups, notably
trichopteran larvae (caddis-flies) and
ephemeropteran nymphs (may-flies), were
also scarce in the acidic tributaries and the
abundance of breeding Dippers showed a
particularly strong correlation with the
abundance of the former (Figure 4c). There
was also a significant positive correlation
 Dippers and stream acidity 35

a
as

. • • • • So
• •
• • •
75.

• •• • • se • •
• . • • S • • IS—.•
• •
• •
65
Downloaded by [North Dakota State University] at 16:12 29 October 2014

5.5

4-5
pH
85

•• •
6-5
•• •
• • • •
•
• 00 0
5-5

4.5

1963 6.5 67 69 71 73 75 77 79 81 83

Year

Figure 2. Winter spot-metred pH readings on (a) the upper River Ithon and (b) the upper
River Irfon (open symbols) and two tributaries of the upper Wye (closed symbols). The lines
were fitted by least squares regression to combined data.

between the abundances of breeding ephemeropteran nymphs important to the

Dippers and nymphal ephemeropterans, former and trichopteran larvae important to
although this was due mostly to a cluster of the latter (Table 1). Mean dry weights for
outlying points (Figure 4a). trichopteran families (4.5-52.9 mg) were
greater than for all the other macroinver-
tebrate families recorded (1.0-3.7 mg) and
Dietary analysis
the former provided nearly 70% of the
In numerical terms, there was a significant caloric content of nestling diet. For adults,
difference between the diets of adults and fish were the single most important item in
nestlings (xi = 200.61, P < 0.001), with caloric terms, with trichopteran larvae
 36 S.J. Ormerod et al.
9
-90 (-76)
6
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6-0 8-0
pH
Figure 3. The abundance of breeding Dippers on
the upper reaches (nominally 10 km) of 16 tribu-
taries of the Wye and on the adjacent Grwyne
Fawr, in relation to mean long-term pH. Open
symbols = Round & Moss (1984); closed symbols
= this study. The correlation coefficients were
calculated excluding (and including) the encircled
values, which were for streams of lower gradient
(< 8 m/km).
second. Overall, macroinvertebrate prey,
particularly Trichoptera and Ephemeroptera,
provided over 65% of the caloric value of the
diet (assessed through faecal analysis) of
Dippers in the study area in the 1983 breed-
ing season (Table 1).
DISCUSSION
The study by faecal analysis of the diets of
insectivorous birds has been discussed by
Davies (e.g. 1977), who recorded a high
correlation between ingested prey and faecal
remains in wagtails (Motacillidae) and fly-
catchers (Muscicapidae). Recent experi-
mental evidence (Ormerod, unpublished)
indicates that the same is true for Dipper
nestlings, although for adults the production
of regurgitated pellets and the difficulty in
quantifying the caloric importance of fish are
possible sources of error when analysing
faeces.
Few authors (Shaw 1979) have attempted
to quantify the diets of breeding Dippers or
their nestlings. In the present study, the diet
of nestlings was composed mostly of macro-
invertebrates whilst, for adults, fish and
macroinvertebrates provided roughly similar
proportions of the dietary caloric content.
The importance of trichopteran larvae to
Dipper nestlings has been discussed else-
where (Ormerod, in press) and probably
reflects their size and immobility, relative to
other macroinvertebrate orders, which
permit an efficient capture and transfer of
energy to the nest by adult Dippers when
feeding young (cf. Davies 1977; Carlson
1983).
Relationships between food supply and
the breeding densities of other bird species
are well documented (e.g. Newton et al. 1977;
Enokson & Nilsson 1983) and, from the data
provided here, the abundance of breeding
Dippers correlated strongly with the abun-
dance of those macroinvertebrate groups
which were shown to be important prey.
Moreover, the scarcity of Dippers along
those rivers which had suitable gradients,
but which had low abundances of Trichop-
tera and Ephemeroptera, indicated that prey
availability was a dominant influence on the
distribution of Dippers on the upper reaches
of the rivers studied. The absence of breed-
ing Dippers from the upper River Wye,
noted by previous authors (Round & Moss
1984), was not surprising in view of the low
abundance of macroinvertebrates along this
stretch (Brooker & Morris 1980; this study).
A further feature of those rivers along
which Dippers were scarce, in spite of suit-
able gradients, was a low pH, with minima
of 4.5 recorded on the upper Irfon and on
some tributaries of the upper Wye. However,
the relationship between pH and the abun-
dance of breeding Dippers (Figure 3) was
probably an indirect consequence of the
abundances of macroinvertebrates and fish,
both of which are also relatively scarce at low
pH (Townsend et al. 1983; Stoner et al. 1984;
this study). Nevertheless, the relationship
provides a pathway by which human in-
fluences on surface water acidity might
influence indirectly the breeding distri-
bution of Dippers: Harriman & Morrison
(1982) and Stoner et al. (1984) have described
an acidifying effect of conifer plantations on
the chemistry of soft-water streams in Wales
and Scotland, in both cases with effects on
the quantity and quality of fish and macro-
invertebrates. Additionally, some ornitho-
 Dippers and stream acidity 37

10 b

r ..54

6 6

•
MO •
4 4 M • •
SIX

•
2 2

0
Downloaded by [North Dakota State University] at 16:12 29 October 2014

0
10 100 1000 15 100 1000
Ephemeroptera N 1j°g103 Plecoptera N Dog io ]

10

-78
8.

6
•

,0 •

1 10 1000 125 250 500 1000 2000 400C)

TricItoptera N Elog io ] Combined N gog io l

Figure 4. Relationships between the abundances of breeding Dippers (pairs per 10 km) and
selected macroinvertebrate groups (N = numbers per 2 x 3 minute kick sample; log. scale).
Breeding abundances assessed in 1977 (RSPB 1977) = open symbols; assessed in 1982 =
closed symbols. The lines were fitted by least squares regression and the encircled values, for
the impounded River Elan, excluded.

Table 1. The diets of breeding Dippers and their nestlings in the study area in 1983 assessed by faecal
analysis

Prey item Total number of items Caloric value (kcal)

Adults (n=72) Nestlings (n=75) Adults (%) Nestlings (%) Combined (%)

Trichopteran larvae 176 430 8.17 (31.8) 32.29 (68.9) 40.46 (55.8)
Ephemeropteran nymphs 484 269 4.74 (18.4) 2.76 ( 5.7) 7.50 (10.3)
Plecopteran nymphs 90 30 1.15 ( 4.5) 0.41 ( 0.9) 1.56 ( 2.1)
Others 54 36 0.37 ( 1.4) 0.29 ( 0.6) 0.66 ( 0.9)
Fish 10 10 11.22 (43.7) 11.22 (23.9) 22.44 (30.9)

Totals 814 775 25.65 46.87 72.52

Note: The caloric values for each macroinvertebrate order have been summed from calculations made at
family level. Method in text.
 38 S.J. Ormerod et al.
logical consequences of surface-water acidity
have been described in North America and
Sweden. Haines (1982) noted a reduction in
the breeding distribution of Black Ducks
Anas rubripes in areas sensitive to acidifi-
cation, and alluded to an impoverished food
supply. Nyholm & Myhrberg (1977) and
Nyholm (1981) described shell-thinning and
a reduction in breeding success in passerines
feeding on insects emerging from moder-
ately acid lakes. High concentrations of
aluminium may be implicated.
Downloaded by [North Dakota State University] at 16:12 29 October 2014
In the present study, all the streams which
showed evidence of an historical fall in pH
drained from catchments which were
25-40% covered by mature forests; none
had breeding Dippers within 8 km of its
source in 1982. No scientific data are avail-
able on the historical distribution of Dippers
over these reaches. However, Colonel
Morrey Salmon (pers. comm.) provided
information on occupied nest-sites which
indicated that Dippers bred on the upper
River Irfon in the mid-1950s at abundances
in excess of nine pairs per 10 km, and other
anecdotal accounts (Smith 1967) corroborate
this observation. Few other changes, for
example in the availability of nest-sites, have
occurred along this reach (Ormerod, un-
published). Such an apparent decrease in
population, which accompanied an increase
in the acidity of this river, is not consistent
with data from other studies of Dipper
populations which have either remained
constant (Taylor & Marchant 1982) or have
shown considerable increase (Perry 1983)
over recent years. Hence a more widespread
regional survey of the distribution and
breeding success of Dippers in soft-water
areas of upland Scotland or Wales might be
desirable in the light of the results presented
here.
ACKNOWLEDGEMENTS
We are grateful to Colonel H. Morrey Salmon
and Welsh Water Authority for allowing
access to unpublished data, and to Dr A. S.
Gee and R. A. Jenkins for comments on the
manuscript. Our thanks are also due to
Roger Lovegrove, Mark Boilstone and
Dr Lennox Campbell of the RSPB. S.J.O. was
in receipt of a NERC studentship for work
on macroinvertebrates.
SUMMARY
Studies of the distribution of breeding
Dippers in relation to prey availability and
environmental factors were undertaken in
the River Wye catchment in 1982. Dietary
studies were undertaken in 1983. Breeding
abundance correlated strongly with the
abundance of those groups of macroinver-
tebrates, notably larval Trichoptera, which
were also prey of importance to Dippers
when feeding nestlings. Breeding abundance
also correlated with stream acidity, although
the latter probably had an indirect effect
through the food supply. Trends in historical
data indicated that several rivers in the
catchment had become more acidic over
recent decades and at least one of these
showed a concomitant reduction in the
number of breeding Dippers.
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