Freshwater Biology (2009) 54, 621634 doi:10.1111/j.1365-2427.2008.02136.

APPLIED ISSUES

Availability of and access to critical habitats in regulated

rivers: effects of low-head barriers on threatened
lampreys
M A R T Y N C . L U C A S * , D A M I A N H . B U B B * , M I N - H O J A N G * , , K Y O N G H A * , A N D J E R O M E E . G .
MASTERS*
*School of Biological and Biomedical Sciences, University of Durham, Science Laboratories, Durham, U.K.

Department of Biology Education, Kongju National University, Kongju, Chungnam, South Korea

SUMMARY
1. Conservation of freshwater animal populations requires their access to, as well as
sufficient availability of, critical habitats, such as those for reproduction. Abundant small-
scale barriers may cause extensive fragmentation of freshwater habitat but, by comparison
to larger structures their effects are rarely considered by catchment managers. The
relationship between the distribution of, and access to, spawning habitat in a regulated
river, characterized by abundant small barriers, was examined for river lamprey Lampetra
fluviatilis, a threatened migratory fish.
2. Telemetry of adult lamprey in the River Derwent, North East England was used to
quantify upriver migration and access to spawning habitat, together with surveys of
spawning habitat availability and spawning activity between 2002 and 2007.
3. Access in to the Derwent appeared severely restricted by a tidal barrage, beyond which
lamprey migrated rapidly in unobstructed reaches. Of all lamprey tagged in the lower
4 km of river, or ascending the barrage, 64% and 17% passed the first and second weirs
respectively, with high flows crucial for this. Although over 98% of lamprey spawning
habitat occurred more than 51 km upstream, on average just 1.8% of river lamprey
spawners were recorded there.
4. In order to protect or rehabilitate species or species assemblages, greater attention needs
to be paid to the relative spatial distribution of low-head barriers and the resultant
availability of key habitats within individual catchments. This is particularly important
given the renewed emphasis internationally on low-head hydropower solutions as a
source of renewable energy, and the rapid growth in numbers of low-head barriers in
many catchments.

Keywords: dam, habitat fragmentation, Lampetra fluviatilis, migration, telemetry

habitats is fundamental to life cycle completion and

Introduction
for their effective conservation or rehabilitation (Wer-
For animals displaying ontogenetic shifts in habitat ner & Gilliam, 1984; Taylor et al., 1993; Lucas & Baras,
use, the availability of, and access between these 2001). Lack of availability of one or more habitat or

Correspondence: Martyn C. Lucas, School of Biological and Biomedical Sciences, University of Durham, Science Laboratories, South
Road, Durham DH1 3LE, U.K. E-mail: m.c.lucas@durham.ac.uk
Present address: Jerome E. G. Masters, Environment Agency, Phoenix House, Global Avenue, Leeds LS11 8PG, U.K.

 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd 621
622 M. C. Lucas et al.
poor connectivity between these habitats is likely to
act as a bottleneck and lead to population decline
(Wilcox & Murphy, 1985; Law & Dickman, 1998).
Species that exhibit different habitat requirements for
life cycle completion are particularly susceptible to
disruption of connectivity between habitats (Taylor
et al., 1993; Amoros & Bornette, 2002; Mumby, 2006)
and this is true of most diadromous fish species
(McDowall, 1992; Baras & Lucas, 2001).
Anthropogenic barriers such as barrages, dams and
weirs dramatically reduce the longitudinal connectiv-
ity of rivers and also strongly alter the river environ-
ment and neighbouring habitats (Baxter, 1977; Pringle,
Freeman & Freeman, 2000; Malmqvist & Rundle, 2002;
Nilsson et al., 2005). They often exclude, fragment and
isolate fish populations (Baras & Lucas, 2001; Morita &
Yamamoto, 2002; McLaughlin et al., 2006), preventing
or disrupting natural patterns of migration and
dispersal between key habitats. This results in reduced
fitness of organisms relying on those conditions, in
some cases leading to population extinction (Pringle
et al., 2000; Baras & Lucas, 2001; Zabel & Williams,
2002). Due to the importance of the longitudinal river
channel axis for migration in their life cycles, high
proportions of diadromous fish assemblages across
most regions have declined in distribution and abun-
dance and are increasingly threatened by extinction
(McDowall, 1992; Pringle et al., 2000; Baras & Lucas,
2001; Duncan & Lockwood, 2001). The cumulative
construction of dams worldwide has continued to
increase steadily, within developed and developing
regions (World Commission on Dams, 2000; Jackson
et al., 2001) and river system fragmentation is wide-
spread (Dynesius & Nilsson, 1994; Nilsson et al., 2005).
Although smaller low-head barriers may have less
dramatic local effects than large dams, their numbers
are probably two to four orders of magnitude greater
than for large dams and their cumulative effects may
be significant (Lucas & Baras, 2001; Poff & Hart, 2002;
Cooke et al., 2005; McLaughlin et al., 2006). They may
not form absolute barriers but have limited perme-
ability to passage of fish and other megafauna,
restricting access to parts of the river system, often
under a limited range of environmental conditions
(Lucas & Frear, 1997; OConnor et al., 2006). They also
locally modify habitat, again influencing suitability for
particular species and contributing to changes in fish
species assemblage (Tiemann et al., 2004; Gillette et al.,
2005; Poulet, 2007).
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
Lampreys are one taxonomic group that are sensi-
tive to freshwater habitat alteration; most species have
declined in distribution and abundance and many are
regarded as threatened (Renaud, 1997; Close, Fitzpa-
trick & Li, 2002). They are often of economic, cultural
and ecological significance (Tuunainen, Ikonen &
Auvinen, 1980; Close et al., 2002). Although, within
their native ranges, population declines have been
attributed to several factors including channelization
(Ojutkangas, Aronen & Laukkanen, 1995) and poor
water quality (Myllynen, Ojutkangas & Nikinmaa,
1997), the most pervasive contributory factor appears
to be river regulation and obstruction (Tuunainen
et al., 1980; Beamish & Northcote, 1989; Renaud, 1997;
Almeida, Quintella & Dias, 2002; Close et al., 2002).
The river lamprey Lampetra fluviatilis (Linnaeus) is a
parasitic, principally anadromous, species which typi-
cally migrates upriver in winter and spawns in spring
(Hardisty, 1986). Formerly widespread in Western
Europe (Hardisty, 1986; Kelly & King, 2001), it is
regarded as endangered (Lelek, 1987; Renaud, 1997)
and receives protection through the EC Habitats
Directive. The aim of this study was to determine
the relationship between distribution of, and access to,
fragmented spawning habitat for river lamprey in
relation to the occurrence of low-head potential
barriers. While study of this threatened species pro-
vides necessary information on mechanisms affecting
lamprey populations, it is also an appropriate model
species for investigating interactions between gross
abundance and net availability of key habitats at the
catchment level.
Methods
Study area
The study was carried out in the River Derwent,
a tributary of the River Ouse that joins with the River
Trent to form the Humber Estuary and flows south-
eastwards to the North Sea in Northeast England
(Fig. 1). The Humber system, with a mean discharge
of 250 m3 s)1, provides the largest contribution of
freshwater to the North Sea of all British rivers (Law,
Wass & Grimshaw, 1997). It has an extensive, shallow,
productive estuary and coastal zone and this, com-
bined with freshwater spawning and larval lamprey
habitats in Ouse tributaries, including the Derwent,
provides suitable conditions for a substantial river
 Availability of and access to critical habitats in regulated rivers
Fig. 1 Location of the Derwent study area
in relation to the Humber catchment,
showing channel obstructions in the lower
Derwent catchment as bars across the
rivers and locations of acoustic loggers (o)
and radio loggers (x) in the Derwent. At
Elvington and Stamford Bridge, several
loggers were present and the symbols are
overlaid.
lamprey population (Lucas et al., 1998; Jang & Lucas,
2005; Masters et al., 2006). Under the EC Habitats
Directive, the Derwent is a Special Area for Conser-
vation (SAC) and the Humber is a proposed SAC, for
which river lamprey and sea lamprey Petromyzon
marinus Linnaeus are listed features.
The lower 60 km of the Derwent is distinctly
lowland in character (overall gradient of about
0.3 m km)1), but about half this drop occurs locally
at the five low-head weirs (23 m height) and one
tidal barrage along the main channel over this
distance (Fig. 1). Barmby Barrage and lock is a tidal
barrage at the mouth of the Derwent, with two
undershot sluices, and was commissioned in 1975 in
order to maintain freshwater levels three kilometres
upstream (river kilometre 3, rkm3) for water abstrac-
tion. Elvington Sluices and lock (rkm24.3) maintains
the water level for water abstraction upstream and
consists of two undershot, pivoting sluices situated on
a 1 : 5 sloping weir sill. The weir has a head loss of
approximately 3.0 m at Q75 flow, and the sluices open
wider automatically with increasing discharge. A pool
and weir fishway, enters the river below the right
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
623
hand sluice and has its upstream exit in the bypass
canal. Stamford Bridge (rkm35.6) has a mill weir of
2.1 m head loss at Q75, and a bypass canal and lock,
together with a 1 : 5 slope Denil fishway up the right-
hand edge of the weir. Buttercrambe Weir (rkm40.2) is
a Flat-V flow-gauging weir with a head loss of 2.0 m
at Q75 and with no fishway. Howsham Weir
(rkm45.7) is an old mill weir, with a head loss of
about 2.0 m at Q75 and with no fishway. The
obstruction at Kirkham (rkm50.3) consists of a mill
weir, together with two overshot sluices feeding a
short diversion canal, both with a head loss of
approximately 2.5 m at Q75, and with a Larinier
baffle fishway at the left bank of the weir.
The lower Derwent is typically 1520 m wide and
has a 10-year mean daily flow of 15 m3 s)1, but is
strongly responsive to precipitation. The river is
relatively deep and slow, typically with mid-channel
depths of 26 m. Silt and sand (habitat for lamprey
larvae) is abundant throughout the lower river, but
likely spawning habitat is most common in the middle
and upper reaches of the river and in localized areas
below the weirs. The Derwents main tributary, the
624 M. C. Lucas et al.
Rye, joins at rkm68. Swift, shallow water with
gravelcobble beds become progressively more abun-
dant in the Derwent, Rye and minor tributaries
upstream of the Rye-Derwent confluence.
Telemetry of lamprey
In order to determine the success of passage past
potential barriers to spawning areas adult river
lamprey were radio tracked (River Derwent, 2002
03 to 200304) or acoustic tracked (River Ouse
estuary and River Derwent, 200203 to 200506)
(Fig. 1, Table 1). Lamprey for tagging were trapped
in the lower Derwent and Ouse (Jang & Lucas, 2005;
Masters et al., 2006). Where possible, lamprey for
release in the lower Derwent, were captured in the
lower 4 km of the river, immediately above the tidal
barrage. Similarly, where possible, lamprey for
release in the Ouse at the mouth of the Derwent,
were captured from traps set immediately adjacent in
the tidal Ouse. However, because of difficulties in
capturing sufficient fish, these were supplemented by
use of lamprey captured further upstream in the tidal
Ouse, using the same methods. A total of 57 fish
were tagged and released immediately below the
barrage and 56 fish were tagged and released less
than 4 km above the barrage, between November
and March (Table 1), representing the middle to end
period of adult migration in the lower river (Masters
et al., 2006). In 200203 a further 16 lamprey were
captured (by trap and from tree roots) between
rkm22.9 and rkm50.3 km in March and April and
radio tagged to study spatial behaviour through the
spawning period (Table 1). In total 129 adult river
lamprey were telemetry tagged between 200203 and
200506.
Table 1 Details of telemetry tagged river lamprey released in the River Ouse at the mouth of the River Derwent and released in the
Derwent upstream of the tidal barrage
Year Release site
200203 Ouse
Derwent, 14 km upstream
Derwent, 2350 km upstream
200304 Ouse
Derwent, 14 km upstream
200405 Ouse
200506 Ouse
Derwent, 14 km upstream
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
Lamprey were anaesthetized in a 0.1 g L)1 buffered
solution of tricaine methansulphonate (MS-222) for tag
attachment which was carried out under U.K. Home
Office licence. In seasons 200203 to 200405 radio and
acoustic tags were attached externally, adjacent to the
anterior dorsal fin, based on the method of Almeida
et al. (2002). But in 200506, with the advent of
narrower acoustic tags than previously available, 32
of 46 lamprey were tagged by implantation into the
body cavity, with the remainder externally tagged
with larger tags. The range of temperatures over which
lamprey were tagged was 2.08.5 C (mean 4.9 C).
Under some circumstances temperature at tagging
may influence subsequent survival and performance
of tagged fish. For lamprey released in the lowest 4 km
of the Derwent (n = 56) there was no significant
difference (t-test, P > 0.05) in tagging temperature
between lamprey that subsequently reached the first
river section upstream containing spawning habitat, or
beyond, and those that did not.
Radio tags (PIP, AgO 393 cell, Biotrack, Wareham,
U.K.) measured 17.0 8.2 6.0 mm, with a whip
antenna length of 10 cm and were potted in synthetic
rubber compound (Plastidip, Petersfield, U.K.). Radio
tags operated at 173 MHz, with a nominal spacing of
10 kHz to identify individual lamprey and an inter-
pulse period of 2.5 seconds, a period appropriate for
active tracking and scanning of relatively small
numbers of tags on different frequencies. Radio tag
life was 1016 weeks. Ideal tag life would have been
2024 weeks, enabling tracking through the migration
period until spawning (e.g. NovemberApril), but
was not feasible without the use of radio tags that
were excessively large, or lower power output or
longer interpulse interval, both of which would have
compromised tracking.
Mean release date Length, mean
n (min, max) and SD (cm)
6 121202 40.3, 0.5
18 201202 (221102, 5203) 39.6, 2.8
16 19303 (08203, 16403) 35.6, 2.2
20 10122003 (131103, 2104) 41.2, 2.5
15 4104 (41203, 24204) 39.0, 2.1
8 281104 (171104, 91204) 39.9, 2.8
23 241205 (291105, 23306) 39.2, 1.9
23 61205 (31105, 28106) 39.4, 1.9
 Availability of and access to critical habitats in regulated rivers
Acoustic tags used operated at 69 kHz and each
transmitted a unique coded signal. Vemco V9-6L tags
[9.0 mm diameter 20.0 mm long, life of 913 weeks
(Vemco, Shad Bay, NS, Canada)] were attached
externally as described above. Vemco V7-2L (7.0 mm
diameter 18.5 mm long, life of 610 weeks) and
Thelma Smolt [7.3 mm diameter 18.0 mm long, life
of 610 weeks (Thelma AS, Trondheim, Norway)] tags
were implanted surgically into the body cavity and
the incision closed with three non-absorbable 40 silk
sutures. Half of the tags were set to repeat their
identity codes every 1020 s and half were set to
repeat them every 1530 s, each repeat being gener-
ated randomly, to minimize the likelihood of code
collisions with nearby tags. While these fast repeat
rates reduced tag life, they ensured high tag detection
efficiency.
Radio tracking was by a combination of active
tracking (modified Yaesu FT290R; Yaesu Musen Co.,
Ltd, Tokyo, Japan or Sika receiver; Biotrack, Ware-
ham, U.K., 3 or 5-element Yagi antenna), on foot,
usually every 23 days (200203, 52 dates; 200304, 29
dates), and the use of two automatic frequency
scanning and logging receivers (SRX 400, Lotek
Wireless, Newmarket, ON, Canada). The radio log-
gers were positioned 10.9 km and 35.4 km upstream
from the mouth of the Derwent to automatically
record the passage of radio-tagged lamprey (Fig. 1).
Acoustic tracking was carried out principally by the
use of automated acoustic loggers (Vemco VR2) with
omnidirectional hydrophones positioned in the river
at strategic locations. Effective range was approxi-
mately 100 m and under field conditions, with loggers
in acoustically suitable locations, efficiency of logging
tags was high (>97% based on tests and sequential
records of tagged fish by multiple loggers). A total of
up to 19 acoustic loggers were employed, with up to
12 in the lower Derwent, one at the mouth of the
Derwent, immediately below Barmby Barrage (Fig. 1)
and up to a further six deployed within the Ouse
system and other Ouse tributaries (locations not
shown in Fig. 1). Acoustic loggers were deployed
immediately below and above Barmby Barrage,
Elvington Sluices and Stamford Bridge, the first three
barriers on the Derwent. At the latter two sites,
loggers were placed in the river channel above the
weir and near the upstream exit of the navigation
canal to enable the route of fish passage to be
determined. Active tracking using a boat and a
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
625
directional hydrophone and receiver (Vemco VR100)
was carried out periodically to locate fish indepen-
dently of automatic logging (200203, 3 dates; 2003
04, 4 dates; 200405 1 date; 200506, 1 date).
Spawning habitat and distribution of spawning surveys
A survey of the Derwent catchment for suitable
spawning habitat was carried out using a combination
of topographic information and in-stream habitat
surveys in 2003 and re-checked annually. River
lamprey use gravel, sometimes mixed with sand
andor cobble, as a spawning substratum (Hagelin &
Steffner, 1958; Hardisty, 1986; Jang & Lucas, 2005).
This material is normally non-embedded, with water
able to percolate through the gravel, and is typically
found at the tail of a pool or glide, usually before the
water breaks into a riffle (Hardisty, 1986; Maitland,
2003). Water depths at spawning sites are usually 0.2
1.5 m and water velocities are usually 0.51.5 m s)1
(Hardisty, 1986; Jang & Lucas, 2005; M. Lucas,
unpubl. data). Lampetra ammocoete larvae normally
occur in areas with river gradients of 0.26 m km)1,
but under extreme circumstances have been recorded
at gradients of 1020 m km)1, reflecting the ability of
adults to migrate into such zones (Kainua & Valtonen,
1980; Kelly & King, 2001).
River gradient data for the Derwent catchment
were obtained from the U.K. Environment Agencys
digital elevation geographical information system at
1 km intervals from tributary sources and were
mapped. All areas including and upstream of a
1 km section having a gradient greater than
10 m km)1 were classed as unsuitable for adult river
lamprey spawning, since these were typically head-
water streams with cobble and boulder substrata
inappropriate for spawning and for larval habitat.
Field surveys were carried out on the remainder of
the Derwent system to assess the availability of
potentially suitable river lamprey spawning habitat.
These surveys were made in March to June by boat,
by walking the river bank, by wading and by
snorkelling; habitat being characterized by the pres-
ence of gravel substratum without settlement of silt,
in a water depth of 0.21.5 m. This approximates the
conditions described above as suitable for spawning
river lamprey. Additionally areas of river deeper
than 1.5 m were searched for deepwater spawning
habitat in 2003 and 2006 using a boat and an
626 M. C. Lucas et al.
underwater camera attached to a battery-powered
television monitor. Although river lamprey have not
been recorded spawning in deep water, our aim was
to ensure that such sites were not missed in our
surveying as a result of their limited accessibility to
human observation. Accordingly, underwater sur-
veying in deep areas was carried out during the
spawning period including dates when river lamprey
were known to be spawning elsewhere in the
Derwent. This surveying concentrated on areas below
river obstructions, especially Elvington Sluices, and
parts of the lower river downstream of Kirkham
(Fig. 1) at sites where hydraulic conditions suggested
the bed might include habitat suitable for spawning
by river lamprey.
Between 2003 and 2007, up to 66 sites per year
(identified as 1-km river segments), were selected and
searched for lamprey and evidence of lamprey
spawning during April and early May (the predom-
inant period of river lamprey spawning in the River
Ouse system; Jang & Lucas, 2005; M. Lucas, unpubl.
data), mostly at locations identified as potentially
suitable for spawning. In 2004 and 2005 poor water
clarity associated with elevated flows during the main
spawning period precluded detailed inspection for
lamprey spawning. In 2003, 2006 and 2007 each site
was visited between one and 30 times. During
spawning, river lamprey make characteristic nests in
gravel that allow spawning sites to be recognized
even in the absence of lamprey (Hagelin & Steffner,
1958; Jang & Lucas, 2005), but these do not give an
indication of abundance, because highly variable
numbers of lamprey can use a single nest area (Jang
& Lucas, 2005). Any lamprey observed was identified
to species and the maximum count of numbers
recorded. Numbers of lamprey spawning, nests in
use (definite nests) and empty nests (possible nests)
were also recorded. Most surveying was carried out
by day, although some night-time surveys were also
done. Although lamprey are nocturnal during migra-
Ascended No. returned
Number upstream of at least once
Years released tidal barrage after >1 h absence
200203 6 1 2 1
200304 20 0 16
200405 8 0 2 2
200506 23 9 2
Combined (%) 57 10 (17.5) 22 (38.6)
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
tion, at spawning time they lose their negative
phototaxic behaviour and can easily be observed
engaged in spawning during daylight (Hardisty, 1986;
Jang & Lucas, 2005).
Environmental factors and analysis
Data on Barmby Barrage sluice gate operation (gate
positions) and tidal water heights below Barmby
Barrage were obtained from the U.K. Environment
Agency records. Mean daily river discharge records for
the River Derwent were obtained from the Environ-
ment Agencys Buttercrambe gauging station at
rkm40.2. Modelled daily flows, taking into account
water abstractions and minor inflows downstream of
Buttercrambe, showed a very close correlation (R2 >
0.99) between gauged mean daily flows at Butter-
crambe and estimated mean daily flows at Barmby
Barrage and Elvington. Therefore, gauged mean daily
flows at Buttercrambe were used to determine the
relationship between river discharge and lamprey
migration, especially at Elvington and Stamford
Bridge. Water temperatures were measured at 0.5 h
intervals using an automatic logger (Tinytag, Gemini
Data Loggers, Chichester, U.K.) at Stamford Bridge or
Elvington. Statistical analyses were carried out using
S P S S (Release 11.0; SPSS Inc., Chicago, IL, U.S.A.).
Results
Entry to the river and upstream migration
Of the 57 acoustic tagged lamprey released at the
mouth of the Derwent over the study period 10 (18%)
ascended the barrage (Table 2). In 200304 and 2004
05 none of the lamprey passed the barrage, even
though 39% (combined across years) left the river
mouth area for at least 1 h, but then returned some
time later, often after several days or weeks. Several
tagged lamprey repeatedly visited the barrage vicin-
Table 2 Numbers of acoustic tagged
No. returned lamprey released at the mouth of the
repeatedly with Derwent in the tidal Ouse, those which
>1 h absence ascended the barrage, those that returned
to the mouth of the Derwent once only
after an absence of at least 1 h and those
2
that returned repeatedly at intervals of
more than 1 h
1
6 (10.5)
 Availability of and access to critical habitats in regulated rivers 627
8
(a) Lamprey 33 present downstream
Ouse level
6 Gate position

Height (m)
2

-2

17/12/04 19/12/04 21/12/04 23/12/04 25/12/04 27/12/04

(b) 10

Release
Fig. 2 (a) Segment of a track record for a Gate position
Ouse level
river lamprey repeatedly detected (grey 8 Lamprey 115
bars) below Barmby Barrage in relation to Lamprey 56
Lamprey 55
tidal cycle and barrage gate operation and 6
Lamprey 52
Height (m)

(b) detections above and below Barmby Lamprey 51

Barrage of six tagged lamprey released at 4 Lamprey 47

the mouth of the Derwent, together with

2
tidal height and barrage gate operation.
When fish were detected repeatedly
0
within a 10-min period only a single
detection is shown on graph for clarity. -2
All fish ascended the barrage during
an atypical period when the gates were -4
continuously open. 29/11/05 0 : 00 29/11/05 8 : 00 29/11/05 16 : 00 30/11/05 0 : 00 30/11/05 8 : 00

ity, suggesting a strong attraction to the area and 100

possible attempts to pass the barrage (Fig. 2a). In
200506 nine (39%) tagged lamprey passsed the 80
% of taggged lamprey

barrage, of which five were tagged externally. Six of

eight lamprey released on 29 November successfully 60

Stamford bridge weir

passed the barrage at a time when flows were
Elvington sluices

relatively high in the Derwent (41.5 m3 s)1) but not 40

in the much larger Ouse (44.2 m3 s)1). This Derwent
high-flow event coincided with ebb tides and the
20
barrage gates remained open through several tidal
cycles, which is atypical of barrage operation (Fig. 2b).
0
Once above the barrage, upstream migration of 5 10 15 20 25 30 35 40
river lamprey was characterized by rapid initial Distance from mouth of derwent (km)
movement over a period of several days to weeks as
Fig. 3 Percentage of all river lamprey tracked from immediately
they progressed upstream, largely but not exclusively upstream of Barmby Barrage on the River Derwent (n = 56)
at night. Combining data from all tagged lamprey that that migrated past particular sites. The shaded, vertical bars
successfully negotiated the barrage (n = 10) or were identify the positions of the two lowest weirs in the
released above the barrage in the lower 4 km of the freshwater section of the river.

Derwent (n = 56), the two obstructions upstream of

Barmby Barrage acted as partial obstructions with Bridge. There was no significant difference in median
substantial declines in the proportions of fish recorded upstream migration distance for tagged
recorded immediately upstream of these sites lamprey in the Derwent in 200203 (34.4 km), 2003
(Fig. 3). Sixty-four per cent of lamprey were recorded 04 (26.8 km) and 200506 (35.4 km) (MannWhitney:
passing Elvington Sluices but only 17% of the initial all P > 0.05). Maximum distances migrated for radio-
cohort were recorded passing upstream of Stamford located lamprey in 200203 and 200304 were 44.9 and
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
628 M. C. Lucas et al.
45.4 km, respectively (below Howsham Weir), while mean daily flow (Fig. 4a). Lamprey passed Elvington
in 200506 five fish passed the loggers above Stamford and Stamford Bridge weirs at significantly higher
Bridge (rkm35.8). By contrast lamprey tagged 22.9 flows than they approached them (Mann Whitney:
50.3 km upstream of the Derwent mouth immediately Elvington, W = 514, P = 0.004; Stamford Bridge
prior to and during the spawning season (2003) W = 173, P = 0.002). Although fish moved through
moved short distances only (median, 0.2 km; range the lower, unobstructed part of the river over a range
09.7 km). of flows, all passage past Elvington Sluices occurred
Using the tracking data for 200506 (n = 29), for during periods of elevated discharge at mean daily
which precise arrival and departure times at loggers flows greater than 27 m3 s)1 (within upper quartile of
was obtained, lamprey movements were related to all flows over the tracking period). At least 90% of
tagged lamprey passing the weir were recorded
80 moving through the sluices rather than by the fishway
or canal. Lamprey recorded at the logger downstream
of the next obstacle, Stamford Bridge, were also
60 recorded at elevated discharges, probably due to
continued migration during high flows immediately
Discharge (m3s1)

after passing Elvington (Fig. 4a). By contrast all

40
lamprey recorded passing Stamford Bridge weir in
200506 did so at mean daily flows greater than
20
44 m3 s)1, outside the 90th percentile of all flows (i.e. a
flow exceedance value of under 10%) during the
tracking period. Although data from seasons 200203
0 and 200304 could not be ascribed flows at passage
by

Br.

.
s

ton

ton

Br

with the same precision, passage at obstructions,

ow

rm

ing

ing

ord

ord
fl

Ba

2.0
ter

Elv

Elv

including those with fishways, was almost always

mf

mf
US
win

S ta

S ta
DS

US
All

associated with strongly elevated river discharge.

DS

US

X Data
1.5
Further evidence of the influence of Elvington and
Speed (km h-1)

1.0
Stamford Bridge in restricting passage is shown from
the rate of progress upstream by tagged lamprey
0.5
through unobstructed zones downstream and
upstream of Elvington and in the obstructed zones
0.0 at Elvington and Stamford Bridge (Fig. 4b). The
median speed for lamprey passing through Elvington
Sluices was similar to upstream and downstream
.4 r .

0.3 eir
0.8 ices
d

te d

D = mfo d

d
6k n

10 rd B
km
km

km
S E r ic te

Sta tricte

ord tricte
= 1 to
m

reaches, but the range of speeds was much wider,

D = Br. W
ing estric
D lving

D = S lu
o D rest

DS nres

s
Sta Re
to n

reflecting the flow experienced on arrival at Elvington

R
Un

mf
E lv

those lamprey arriving at low flows were delayed

t

to
by
rm

until a spate occurred. By contrast all lamprey were

gto
Ba

in
US

Elv

delayed for long periods at Stamford Bridge.

US

Fig. 4 (a) Distributions of mean daily discharge for the period

November 2005 to March 2006 and discharges occurring when Distribution of spawning habitat and river lamprey
river lamprey were first recorded passing upstream at a location. spawning
Boxes show the median and quartiles, whiskers show the 10th
and 90th percentiles, outliers shown as circles and (b) rates of The main areas of gravel spawning habitat at low to
upstream movement by tagged river lamprey recorded for the intermediate gradients occur in the Rye and upper
unrestricted sections downstream of and upstream of Elvington Derwent subcatchments. Within the lower 68 km of
Sluices, and for the restricted sections past Elvington Sluices and
Stamford Bridge Weir in 200506. Boxes show the median and
the Derwent main channel, following intensive
quartiles, whiskers show the 10th and 90th percentiles, outliers searches, a total of 0.4% river length was ascribed as
shown as circles. being potentially suitable for river lamprey spawning
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
 Availability of and access to critical habitats in regulated rivers
R.
Fig. 5 Locations of river lamprey spawn-
ing sites and the maximum counts of
spawning river lamprey on the Derwent
system in 2003, 2006 and 2007 in relation
to the distribution of suitable and unsuit-
able habitat for spawning of river lam-
prey. Sites are given to a scale of 1 km and R.
Ou
reflect the occurrence of any spawning
habitat within each 1-km river channel
section. Large and small bars represent
obstructions (mostly weirs) greater than 0 10 km
and less than 1 m in height, respectively.
(Fig. 5). Spawning habitat was much more abundant
upstream of that area, associated with elevated gra-
dient and related changes in stream geomorphology,
especially in the main tributary, the Rye (Fig. 5).
Despite underwater camera and snorkelling observa-
tions, no gravel spawning habitat was found in deep
water areas, including downstream of Elvington
Sluices. The downstream-most spawning habitat
recorded in the main river channel was below Stam-
ford Bridge weir (Fig. 5).
Evidence of river lamprey spawning was found at
six (9%) of all sites examined in 2003, four sites in 2006
and eight sites in 2007 including immediately below all
of the main channel weirs in the lower Derwent, except
at Elvington (Fig. 5). Although over 98% of potential
spawning habitat occurred more than 51 km upstream
(upstream of Kirkham Weir), an annual average of just
1.8% (range: 0.05.1%) of daily maximum counts of
lamprey spawners were made upstream of this.
Abundant habitat in the middle Rye, upper Derwent
and their tributaries was not used by river lamprey in
2003, 2006 or 2007. In 2004 and 2005 elevated flows and
low clarity during the spawning period precluded
effective surveying. River lamprey spawning, in 2003,
2006 and 2007, occurred mostly below weirs and
around bridges in areas of locally elevated gradient
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
629
North
Sea
R ye
No lamprey recorded in any year
2003 2006 2007
1-10 lamprey
11-50 lamprey
51-250 lamprey
R. Derwent
251-1000 lamprey
>1000 lamprey
Unsuitable for spawning
se Suitable for spawning
Barriers 1 m height
Barriers <1 m height
andor water velocity over gravel. The maximum
counts of spawning lamprey decreased at sites further
upstream in 2003, 2006 and 2007 but the breadth of
distribution and abundance of spawners was lower in
2006 compared with 2003 and 2007 (Fig. 5).
Discussion
This study demonstrates how small-scale potential
barriers can affect the distribution of key habitats, but
especially can impact on access to those habitats by
relatively mobile megafauna. These barriers are of
types that are abundant across catchments but rarely
considered in terms of their catchment fragmentation
impacts. Although, in absolute terms, large amounts of
spawning habitat are available for river lamprey in the
Derwent, its distribution is very poorly matched by
accessibility. Despite over 98% of suitable spawning
habitat for river lamprey being more than 51 km
upstream in the Derwent, an annual average of just
1.8% of the combined site-specific maximum daily
counts of spawning river lamprey occurred there.
Although it is possible that river lamprey deliber-
ately halted migration below barriers because poten-
tial spawning areas were present, several sources of
evidence indicate that lamprey were restricted in their
630 M. C. Lucas et al.
ability to utilize upriver areas. First, the decline in
numbers of lamprey found at successive spawning
sites upstream and the cumulative decline in propor-
tion of tagged lamprey negotiating successive barriers
strongly suggest a cumulative effect of multiple
partial barriers on upstream migration, similar to that
observed for salmonids (e.g. Gowans et al., 2003) and
for Pacific lamprey Lampetra tridentata (Richardson)
(Moser et al., 2002) at large hydroelectric dams.
Secondly, in 2003, 2006 and 2007 adult river lamprey
penetrated upstream into the River Swale, another
tributary of the Ouse, in large numbers at least 102 km
from the tidal limit of the Ouse (M. Lucas, unpubl.
data), even though abundant spawning habitat is
available within 30 km of the tidal limit. This dem-
onstrates that river lamprey from the same catchment,
in the same year will use upriver spawning habitat
even where substantial habitat is available down-
stream and it is notable that the Ouse-Swale system
retains fewer in-channel barriers, though they are of
similar size to those on the Derwent. Lastly, 36% of
river lamprey were not recorded passing Elvington
Sluices, including during the immediate pre-spawn-
ing and spawning period, even though no spawning
habitat was available below it.
Following construction of impassable hydroelectric
dams in Finland, populations of river lamprey have
declined (Tuunainen et al., 1980; Ojutkangas et al.,
1995). One way of enabling upstream passage of
lampreys is to employ fishways, yet for many species,
most of these are not very efficient (Haro & Kynard,
1997; Laine, Kamula & Hoolt, 1998; Lucas & Baras,
2001; Moser et al., 2002) due to the low swimming
performance of lamprey (Beamish, 1978; Mesa, Bayer
& Seelye, 2003), their behaviour (Haro & Kynard,
1997; Moser et al., 2002) and, for river lamprey, the
low temperatures at which they migrate (Hardisty,
1986; Masters et al., 2006). Studies of river lamprey
Median discharge % days >
Year m3 s)1 (2575&) 27 m3 s)1
200203 24.7 (13.741.0) 47.6
200304 16.3 (5.726.6) 24.1
200405 15.4 (13.220.2) 18.5
200506 17.3 (13.423.9) 20.5
20062007 19.1 (12.429.6) 30.2
10 year 199778 to 200607 18.5 (11.828.8) 28.2 (2.852.8)*
*Minimum and maximum annual values.
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
swimming performance are ongoing (P. Kemp, pers.
comm.) but it is likely that the temperature for
maximum swimming performance is near to the
upper limit over which migration normally occurs
(Beamish, 1978), probably about 812 C. In the
current study, all lamprey passed barriers during
periods of high flow and use of fishways was
unimportant, at least at Elvington for which good
data are available. The behaviour of lampreys seeking
to pass low-head barriers usually involves the use of
the sucker for attachment to the substratum or barrier,
interspersed with burst swimming, usually in lower-
flow areas and often around the edges of fully or
partially inundated barriers (Hardisty, 1986; Quintella
et al., 2004; M. Lucas, pers. obs.).
Over 10 years the proportions of time for which
critical discharge values for lamprey passage were
exceeded on the Derwent during the migration period
(SeptemberMarch) varied 13-fold for flows above
27 m3 s)1 and varied 36-fold for flows greater than
44 m3 s)1 (Table 3). The availability of high flows for
enabling passage is therefore likely to have a strong
effect on the access of lamprey to spawning habitat
fragments and hence on the extent of nursery area
downstream to which the weakly-swimming larvae
are recruited. Furthermore, of the lamprey tagged in
the Ouse, only 18% entered the Derwent past Barmby
Barrage. It would appear that large freshwater attrac-
tion flows, as well as physical access are needed to
enable lamprey to enter. During normal, automatic
operation the barrage gates are shut for a large
proportion of the tidal cycle and when open, water
passes through the gates at high velocity, far from
ideal conditions for encouraging immigration. Nunn
et al. (2008) found evidence of variable year class
strength of Lampetra larvae in the Ouse catchment,
including the Derwent, especially upstream of barri-
ers and suggested that this was a reflection of limited
Table 3 Characteristics of the average
% days > daily discharge and the percentage of
44 m3 s)1 days exceeding critical flows for the River
Derwent at Buttercrambe gauging weir
20.3
during the season of river lamprey
6.1
migration (SeptemberMarch) for years
4.0
200203 to 200607, and for this season
5.3
over the 10-year period 199798 to 2006
3.8
07
8.6 (0.924.5)*
 Availability of and access to critical habitats in regulated rivers
access by adult lampreys, linked to low flows in some
years. Our study provides evidence that access to
spawning localities by adults, rather than differences
in larval mortality, may be the causal factor for the
observed variability in larval recruitment.
If access to the main areas of spawning habitat in
the Derwent has been deprived for many decades,
how has a significant population of river lamprey
remained? It is possible that the tiny fragments of
remaining spawning habitat are sufficient for effective
spawning and do not represent a significant bottle-
neck. River lamprey are communal, promiscuous
spawners (Jang & Lucas, 2005), the young of which
disperse to silt habitats downstream (abundant in the
lower Derwent) soon after hatching (Hardisty, 1986)
and it is possible that tiny fragments of spawning
habitat are adequate for population maintenance in
species such as these. It is currently unknown whether
lamprey in the different Ouse tributaries, including
the Derwent, are metapopulations (sensu Hanski,
1999). There is some evidence that, like sea lamprey
(Bergstedt & Seelye, 1995), river lamprey do not
exhibit natal homing (Tuunainen et al., 1980), but
probably return to tributaries containing ammocoete
larvae, for which several odourants attractive to
returning adults have been isolated (Vrieze & Soren-
sen, 2001; Gaudron & Lucas, 2006). Thus dispersal
and mixing is likely to be high. Although discrete
metapopulations may be unlikely, the spawning
population in the Derwent may be supplemented by
straying adults which were spawned in nearby
tributaries. Thus, abundant river lamprey population
components in other parts of the Ouse catchment
could help maintain the Derwent subpopulation.
The small number of sites on the Derwent used
intensively for spawning by river lamprey (c. 90% of
spawners occurring at two sites in 2003 and 2006 and
at three sites in 2007) makes the likelihood of severe
population damage by catastrophic events such as
pollution or by ill-conceived river engineering or local
exploitation (Masters et al., 2006), much greater, par-
ticularly if it is currently a self-sustaining population.
Therefore, a major conservation priority for river
lamprey and similar species in river catchments like
the Derwent must be to increase connectivity to
multiple, larger areas of spawning habitat. For
lampreys this might be assisted by designing and
installing more efficient fishways around the obstruc-
tions or, perhaps more effectively for low-head
 2008 The Authors, Journal compilation  2008 Blackwell Publishing Ltd, Freshwater Biology, 54, 621634
631
barriers, by removing parts of weirs or the whole
obstructions. It is likely that nature-like fish passes
such as rock-ramp weirs (Harris, Thorncraft & Wem,
1998) would be effective for lamprey passage, but
lamprey-specific bypasses are also being trialled
(M. Moser, pers. comm.). Wherever possible, the
operating regimes of gated structures, such as
barrages, should be modified to act in concert with
environmental conditions, promoting free and effec-
tive movement of biota through such structures. This
could include pet door type modifications of the
main gates to allow access during low flows, when the
main gates might otherwise be closed (Giannico &
Souder, 2005).
For rivers with numerous low-head barriers, frag-
mentation of the river habitats may limit dispersal
and migration for a wide range of aquatic megafauna,
especially fishes, including freshwater species (Lucas
& Batley, 1996; Cooke et al., 2005; McLaughlin et al.,
2006). When the distribution of key habitats in the
catchment is overlaid relative to the distribution and
nature of such barriers, a system with selective
permeability to and from habitat fragments is created.
River channel barriers or levees within temperate
regions are often decades or centuries old and
progressive catchment management needs to consider
the spatial distribution of small, but key habitat
fragments in order to assess how best to reconnect
these in longitudinal or lateral dimensions, as has
been examined in several cases (Bednarek, 2001;
Schmetterling, 2003). In many subtropical and tropical
countries, ecological knowledge of the spatial avail-
ability of key aquatic habitats relative to low-head
barriers and its use in conservation plans to limit loss
of connectivity is even poorer, but arguably is more
important (Pringle et al., 2000; Baras & Lucas, 2001).
Given the renewed international emphasis on encour-
aging use of small-scale (low-head) hydroelectricity
structures to contribute to increased renewable power
production (e.g. European Renewable Electricity
Directive 200177EC), the issue of chronic fragmenta-
tion of running water habitats needs to be addressed
simultaneously. Otherwise opportunities for river
habitat rehabilitation are likely to be replaced by
increased ecological deterioration. Greater emphasis
on integrated catchment mapping and network anal-
ysis of additive freshwater habitat fragmentation
effects, especially in relation to multiple minor
breakages in connectivity, is needed in order to
632 M. C. Lucas et al.
address these issues (Peterson, Theobald & Ver Hoef, Threats, conservation strategies, and prognosis for
2007; Moilanen, Leathwick & Elith, 2008). suckers (Catostomidae) in North America: insights
from regional case studies of a diverse family of non-
game fishes. Biological Conservation, 121, 317331.
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(Manuscript accepted 25 September 2008)