ABSTRACT

Dipturus batis, the largest of the European Rajiforms, was once common around the coasts
of Britain (Brander, 1981). In recent years due to fishing pressure, its numbers have
receded dramatically from the Irish Sea (Brander, 1981) and the North Sea (Walker and
Hislop, 1998). Little is known about the life history characteristics of this species,
therefore, to better understand its biological characteristics a study was done, mainly using
the data collected by the Glasgow Museum Tagging study, but also with data collected by
the author. The data was split into three groups, A, B and C after it became clear that bias
due to fishing effort and local changes in bathymetry influenced the results as a whole to
greatly to be analysed. Results showed a significant association between the weight and
depth for female fish in all areas (A, B and C), with larger fish being caught in shallower
depths, this correlation was strongest in area A (Spearmans Rank Correlation 2-tailed test,
p=<0.01). The same correlation was found for all males in the study area as well as
(p=<0.05) and there was a strong association in area A (p=<0.01). Using age at maturity as
11 years old (Brander, 1981, Ellis and Walker, 2005 and Little 1995), weight at maturity
was estimated to be between 33-42kg for males and 69-96kg for females, these estimates
are substantially higher than those quoted in the literature and suggest, especially for
males, that either D. batis matures earlier, or is heavier than previously thought at
maturation. It was found that the economic worth of D. batis in area C is higher than its
commercial value and that charter boat trips annually bring £292,500 to the local economy.

Keywords: D. batis, Rajiformes, seasonal migration, maturation, depth, weight, economic

worth.

i
 ACKNOWLEDGEMENTS

My thanks go to:
Dr Alastair Lyndon from Heriot Watt University who supervised this project, his continual
enthusiasm, assistance and guidance throughout this study was invaluable and greatly
appreciated.
Richard Sutcliffe from Glasgow Museums for initially allowing me the use of the Glasgow
Museum Tagging Study data, which he himself has collected since its initiation, but also
for the continued help and assistance he gave during the project.
SNH, SSMEI Sound of Mull Project and Argyll and Bute Council Local Biodiversity
Action Plan for funding the project.
Dr Jane Dodd for accepting me to undertake the research and for her continued enthusiasm,
help and support throughout this study, it made the collection of data in Oban run much
more smoothly. Thanks also to all staff at the Oban SNH office who’s patience and support
was much appreciated.
Dr Sarah Cunningham for all of her help and support from beginning to end, especially her
guidance with GIS software
Ronnie Campbell for all the boat time he provided for research trips. His continued help
and support throughout the write up of this report was greatly appreciated.
I would also like to thank the following people for their contribution to this report: Davy
Holt, Professor Jon Side, Adrian Lauder, SSACN, all the anglers who have contributed to
the Glasgow museum tagging project throughout its course

ii
 CONTENTS
Table of Contents Page number

ABSTARCT i

ACKNOWLEDGEMENTS ii

TABLE OF CONTENTS iii

TABLE OF FIGURES AND TABLES v

ABREVIAITONS vii

CHAPTER .1 INTRODUCTION 1
1.1 Background on D. batis 1
1.2 Distribution 1
1.3 Feeding 3
1.4 Migratory Behaviour 4
1.4.1 Seasonal depth migrations 4
1.4.2 Population movements and aggregations 5
1.5 Life History 6
1.6 Pressures 8
1.7 Potential for Re-populating 11

CHAPTER 2. MATERIALS AND METHODS 12

2.1 The Study Area 12
2.2 Data Collection 13
2.2.1 Collection of biological data 13
2.2.2 Extra Field Work undertaken towards this project 16
2.2.3 Measurements from landed fish 18
2.2.4 Collection of economic data 19
2.3 Data Analysis 20
2.3.1 Population segregation 20
2.3.2 Life history 21
2.3.3 Movement 22
2.3.4 Population estimates 23
2.3.5 Economic importance of D. batis 24
2.3.6 Value of skate compared to commercial market 25
2.3.7 Statistics 26

CHAPTER 3. RESULTS 27
3.1 Behavioural Ecology of D. batis 27

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3.1.1General Observation of the population throughout the study area 27
3.1.2 Seasonal observations of the population throughout the study area 33
3.1.3 Areas A, B and C 37
3.1.4 Annual migration 48
3.1.5 Population estimates 52
3.1.6 Fishing mortality 52
3.2 Economic importance of D. batis 53

CHAPTER 4 DISCUSSION 58
4.1 Maturation 58
4.2.1 Population segregation 61
4.2.2 Areas A, B and C 63
4.3 Migratory Behaviour 64
4.4 Population Estimates 66
4.5 Economic Importance 68
4.6 Life History 69

CHAPTER 5. CONCLUSIONS 70
Conclusions from this study 70
Further areas for Study 71

REFERENCES 73

APPENDICES 78
Appendix 1 Individual Growth rates 78
Appendix 2 Growth rate Scatter plots 79
Appendix 3 Spearmans Rank Results 81
Appendix 4 Distance travelled by each fish 82
Appendix 5 total number of fish caught each year 83
Appendix 6 Tables used for population estimates 84
Appendix 7 Logsheets used for data collection 88
Appendix 8 Questionnaires 89
Appendix 9 Weight Charts 91

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 Table of figures

Figure 1.1 Map of D. batis Distribution 2

Figure 1.2 Relative numbers of eggs laid per species on an annual basis 7
Figure 1.3 Increase of smaller species in commercial landings 10
Figure 2.1 Map of the Study Area 12
Figure 2.2 Photograph of the Jumbo Rototags 13
Figure 2.3 Diagram of restrictive nature of Rototag 13
Figure 2.4 Diagram of D. batis showing tagging site for Rototag 13
Figure 2.5 Floy Dart Tag 14
Figure 2.6 Needle Applicator for Floy Dart tag 14
Figure 2.7 Diagram of D. batis showing tagging site for Dart Tag 15
Figure 2.8 Fishing gear used for catching D. batis 16
Figure 2.9 Diagram of D. batis showing Gaffing sites 16
Figure 2.10 Diagram of D. batis showing measurements taken 17
Figure 3.1 Map showing the distribution of both sexes throughout the study area 28
Figure 3.2 Histogram showing the number of Males caught in each weight class 29
Figure 3.3 Histogram showing the number of Females caught in each weight class 29
Figure 3.4 Scatter Plot showing weight related depth distribution for males 31
Figure 3.5 Scatter Plot showing weight related depth distribution for females 31
Figure 3.6 Sex ratio over time 32
Figure 3.7 Map showing the seasonal distribution of males 33
Figure 3.8 Map showing the seasonal distribution of females 34
Figure 3.9 Median depth of males and females 35
Figure 3.10 Growth rate of male D. batis 36
Figure 3.11 Growth rate of female D. batis 36
Figure 3.12 Sex ratio in areas A, B and C 38
Figure 3.13 Histogram showing numbers of males and females caught in area A 39
Figure 3.14 Histogram showing numbers of males and females caught in area B 39
Figure 3.15 Histogram showing numbers of males and females caught in area C 40
Figures 3.16 Histograms showing numbers of males in each weight class for all areas 41
Figures 3.16 Histograms showing numbers of females in each weight class for all areas 41
Figure 3.18 Map showing depths fish were caught at 42
Figure 3.19 Median seasonal depth values for males and females in Area A 43
Figure 3.20 Median seasonal depth values for males and females in Area B 44
Figure 3.21 Median seasonal depth values for males and females in Area C 44
Figure 3.22 Seasonal weight related depth distribution for females in area A 46
Figure 3.23 Seasonal weight related depth distribution for males in area A 46
Figure 3.24 Seasonal weight related depth distribution for females in area B 47
Figure 3.25 Seasonal weight related depth distribution for males in area B 47

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Figure 3.26 Map of longest migration routes 48
Figure 3.27 Map showing localised travel of males 49
Figure 3.28 Map showing localised travel of males 49
Figure 3.29 Map showing localised travel of females 50
Figure 3.30 Map showing localised travel of females 51
Figure 3.31 Mean distance travelled by males and females 51
Figure 3.32 Population estimates 52
Figure 3.33 Fisheries mortality for D. batis 53
Figure 3.34 Percentage of participants on D. batis angling trips compared to Highlands
and Islands 56
Figure 3.35 Percentage contribution of charter trips from Oban to Highlands and
Islands watersports Industry 56
Figure 3.36 Amount spent on the top four water sports in the Highlands and Islands 57
Figure 3.37 Comparative value of fish to t he sport and commercial fishing industries 57
Figure 4.1 Sigmoid growth curve 60
Figure 4.2 Diagram of hypothetical movement 65

Table 3.1 Estimated amount of money brought to the local economy by anglers 54
Table 3.2 Estimated value of an individual fish 54
Table 3.3 Estimated value of fish to area C 55

vi
 ABBREVIATIONS USED THROUGHOUT THIS REPORT:

CEFAS: Centre for Environment, Fisheries and Aquaculture Science

FRS: Fisheries Research Service

ICES: International Council for the Exploration of the Sea

IUCN: International Union for the Conservation of Nature

MarLIN: Marine Life Information Network: The website of the Marine Biological
Association of the UK and Ireland

SAH: Scottish Angling Homepage

SNH: Scottish natural Heritage

SSMEI: Scottish Sustainable Marine Environment Initiative

SSACN: Scottish Sea Anglers Conservation Network

vii
 1. INTRODUCTION

1.1 Background on D. batis

The Common skate Dipturus batis (formerly Raja batis or Raia batis Linnaeus 1758)
was once common around the coast of the British Isles (Brander, 1981 and Fowler &
Cavanagh, 2005) but due to increased fishing efforts, both commercial and sport
angling, their numbers have rapidly decreased (Brander, 1981). D. batis is a favoured
sport fish and has been pursued by anglers for many years, this has led to its depletion
around the coast of Orkney, where a productive sport fishery has reduced its numbers
(Anon, 2000). The number of D. batis caught in commercial trawls began to decline in
the 1920s and, after a brief recovery period during World War II, they had all but
disappeared from North Sea trawls by the early 1980’s (Walker & Hislop, 1998). One of
the main reasons that these industries have impacted D. batis so much is partly due to
our lack of knowledge of the species. The species is currently on the IUCN red list as an
endangered species throughout its range (IUCN, 2007) and the UK Biodiversity Action
Plan (Anon, 1999) although neither of these offers it any protection in terms prevention
of landing and killing.
The aim of this project was to gain more insight into the life history
characteristics and behaviour for the species as well as assessing the population around
the Isle of Mull, this was accomplished by using the data set provided by Richard
Sutcliffe at the Museum of Glasgow. This report also looks at the economic value of D.
batis to the local economy to the Oban area.

1.2 Distribution
D. batis is the largest of the European Rajiform (IUCN, 2007) with females reaching a
length of 285cm (approx 120kg, weight estimated from weight chart in appendix 9) and
males, 205cm (approx 75kg, weight estimated from weight chart in appendix 9) (Ellis
and Walker, 2005) (length is measured from snout tip to tail tip and can be seen in
Figure 2.10), although there is evidence from commercial catches suggesting that they
may grow up to 180kg in weight. In common with many other species of
elasmobranchs, D. batis is slow growing, late to mature and has relatively low fecundity
(Brander, 1981). They can be found in the northeast region of the Atlantic, ranging from
the Mediterranean near Madeira and northern Morocco up to the Barents Sea and across

1
the Atlantic to Iceland (MarLIN, 2008) (Figure 1.1). They are found at varying depths
throughout their range but are most commonly found in water around 200m deep on the
edge of continental shelves (IUCN, 2007), although in the Barents Sea in the north of
their range, they have been found in water deeper than 300m, and were most prevalent
at depths of 550-800m (Dolgov et al 2005b). A CEFAS trawl study around the British
coast classed D. batis as a deep-water species, distributed along the outer continental
shelf edge of the Celtic and northern North Sea (Ellis et al, 2005). However, this does
not represent the distribution of the species fully as it is often taken by shore anglers,
suggesting that the preferential depth for D. batis cannot be easily assigned, it has also
been hypothesised that depth distribution may be linked to the life history of the species.
They are tolerant to temperature variation as shown by Dolgov et al (2005a) in the
Barents Sea, where it the species occupied a wide temperature range even within a
small geographic region.

Figure 1.1: Map of the known distribution of D. batis

throughout Europe. Range marked in orange fill.
Adapted from ZeeInZicht.

2
1.3 Feeding
It is recognised that D. batis has a varied predatory diet ranging from benthic feeding on
crustacea and molluscs (Little 1995) to demersal species such as cod (Dolgov, 2005c).
In a study on the stomach contents of different species of skate in the Barents Sea by
Dolgov (2005c) it was shown that fish contributed approximately 70% of the diet for D.
batis with young cod, haddock, halibut and long rough dab being prevalent. Fisheries
waste was also a major constituent of their diet, contributing 25% by mass, Dolgov
(2005c) noted that this could not be seen as an indicator of poor abundance of prey
species, but was more indicative of the large amount of available fisheries waste in the
area due to high fishing intensity. There are some noted morphological differences in
tooth shape between the sexes, these have been noted and substantiated by numerous
anglers. Males appear to have more cuspidate teeth associated more with pisciverous
fish, while females have more molariform teeth, more in common with bottom feeding
fish, these differences in tooth shape suggest different feeding strategies, which might in
turn suggest different habitat selection due to different prey items. As mentioned, D.
batis is known for having a varied diet, but the possibility that this might vary with sex
has never been studied and the sex of the fish studied in the Barents Sea was either not
looked at or not disclosed. If it is the case that there is a difference in the feeding
regimes between the sexes, then we may expect to see a difference in behaviour,
especially migratory. It is possible that males, relying on migratory fish species as prey,
follow the migration routes of fish such as cod while females, having a more varied diet,
are able to stay within a smaller hydrographical region, opportunistically feeding on the
benthos and in the demersal water column. However, it is known that males do feed on
the bottom as anglers use benthic targeting equipment, and dead bait for skate fishing
and regularly catch males. Dolgov’s study looked at 39 specimens of D.batis over 6
years, not a large number, which is perhaps a reflection of decreasing population levels.
There was no breakdown of results by size for D. batis in his study so no change in diet
for size/age is noted; other species however which were caught in larger numbers did
have results broken down into stomach contents by length. For the Thronback Ray
(Raja Clavata) it can be seen that as the fish grows, its diet changes, smaller fish having
a more benthic based diet, 70% consisting of largely polychaetes and benthic
crustaceans these diet components not being present in larger fish, where the diet
consists mainly of fish (Dolgov et al, 2005c). If we assume a similar behavioural change
in D. batis we might expect a similar change in diet, although we may anticipate fish to

3
appear in stomach contents at an earlier age due to the larger size of D. batis allowing it
to handle larger prey items at earlier. If this is true, then we would expect only larger
fish to migrate to follow fish stocks, while smaller immature fish may remain in the
same place, feeding on benthic invertebrates.

1.4 Migratory Behaviour

1.4.1 Seasonal depth migrations
The annual movements of D. batis are unknown and previous tagging programmes have
been aimed at trying to understand any patterns in the movement of adult fish. However,
since it is mainly anglers undertaking the tagging, it tends to be the larger, adult fish that
are targeted. The initial thinking was that, due to their shape, they remain in the same
place throughout the year, although Stephen (1929) proposed the idea that skate of all
sizes and sex undertake annual migration to deeper water throughout the winter,
returning to the same shallower waters in the spring time, other species of Rajiform
have been shown to make such annual migrations. One of the best examples of this is
shown research carried out by Hunter et al (2005 a & b) focusing on the migratory
behaviour of R. clavata. Electronic data tags measuring depth and temperature every 10
minutes were used, which revealed that this species made an annual depth migration
(Hunter et al, 2005a). The animals moved from deeper water (20-35m) during the
winter months (November –February) into shallower water (<20m) in spring and early
summer months (March – June). This supported previous studies (Walker et al 1997)
and the timing was in direct relationship to the R, clavata reproduction cycle (Holden,
1975). If we put forward the hypothesis that D. batis performs similar migrations to R.
Clavata, then we would expect to see an annual movement of the species from deeper
water (600-200m) during winter months to shallower water (200-50m) in summer
months in direct relationship to its breeding cycle, which is thought to be between
March and June (MarLIN, 2008). Hunter et al (2005b) also focused on the movements
of a single mature female R. clavata. Using the information gathered by an electronic
data tag and a tidal location method (Hunter et al, 2005b), a migration route for this
individual could be plotted over 423 days. A clear pattern was found as the female
moved to the shallower water within the Thames Estuary in spring through to late
summer time, presumably to spawn, then to the deeper waters off the Dutch coast
during autumn and winter months. It repeated this migration 1.5 times, being released in
London and recaptured off the Dutch coast. Although this was only one animal and

4
therefore not representative of the whole species, it does highlight that annual migration
can occur within the Rajiformes and that some individuals are likely to remain in a
regional population. The migration work of Hunter et al, (2005a&b) on R. clavata
could explain the noted recapture of D. batis specimens within the same area (Little,
1995), Although it is known that some of the population remain in shallower coastal
waters during winter months as some specimens are caught (Little 1997&1998) and
there is anecdotal evidence from anglers that they catch D. batis during the winter
months, although few assumptions about the decline in caught and recaptured fish
during the winter months can be made as this may be down to decreased fishing effort
alone (Little, 1995).

1.4.2 Population movements and aggregations

It is unknown if D. batis populations are closed or recruit from other regional
populations; it could be that individuals of both sexes move between populations. If this
inter-population recruitment does take place it could explain why some fish were caught
further away from their original catch site, they may have been en-route to another
breeding area; there have been no genetic studies to prove or disprove this though.
Trawl surveys undertaken in the North Sea have shown that skates and rays can have
reasonably discrete species distributions, with each species occupying a slightly
different environment, often depth related (Walker and Hislop, 1998) which suggests
that species congregate, either all year round situation or only during spawning. The
Glasgow tagging study has previously shown some possible congregation areas for D.
batis on the west of Mull (Anon, 2000) where animals seem to gather and then disperse
in northerly and southerly movements.
As well as species segregation between Rajiformes, it is also thought that there
is a degree of segregation within the same species based on sex, size and maturity (Ebert
et al, 2008). It is unclear as to what extent this segregation takes place and whether
segregated groups all move within the same population area, or whether each group
occupy different habitats, based mainly on feeding and predator avoidance. In support
of different groups occupying different areas, anecdotal evidence from anglers report
that in the Crinan basin, off the west coast of Scotland, there seems to be a population of
larger juvenile males and few if any females are caught. However, whilst the anglers
claim these male fish to be larger specimens, the angling gear used would very rarely
catch smaller individuals so a full picture of the resident weight range is unclear. Some

5
trawl surveys have caught mostly immature fish (Walker and Hislop, 1998) suggesting
possible congregation of this age group in this region, although the data are not clear
enough to determine if this is a single sex congregation or not. It was also noted in a
Barents Sea study that some long line catches could compromise 100% of D. batis and
other times, no D. batis were caught (Dolgov, 2005b), further suggesting that the
species moves in groups, there was no breakdown of these catches with regard to sex or
size mentioned.
D. batis has also been shown to travel individually; a trawl survey along the
edge of the continental shelf off Norway caught all its specimens of D. batis singly
(Dolgov et al, 2005a). All of these differing reports suggest that the distribution and
segregation of D. batis is complicated and little is fully understood. However, it is likely
that in shallower coastal waters, mature populations come together for spawning as
suggested by Little (1995 and 1997). Using the data collected through the Glasgow
tagging programme, his study produced an interesting pattern regarding the movements
of males into and out of the area. The proportion of males caught increased in the
spring, suggesting an increase in numbers and declined again in the early summer, then
increased again in September/October (Little, 1997). There is evidence from other
Rajiform species that as their population levels decrease, populations have aggregated in
certain key areas, with fewer pressures allowing for increased survival, these are
becoming known as essential fish habitats (Payne et al (2008). If D. batis follows this
survival behaviour we would expect to see a population increase within an area, as well
as an increase in the diversity of sex, size and maturity as groups come together.
Currently there is not enough information to map out segregation within species
accurately.

1.5 Life History

Growth rates for D. batis have been shown to be 1-9kg per year for males and 4-13kg
per year for females (Little, 1998) and they are generally thought to live to 50yrs of age
(IUCN). Both sexes reach maturity at approximately 11 years old, with males being
125cm long (Fowler & Cavanagh, 2005) and females 180cm (Walker & Hislop, 1998).
The number of eggs laid per female is dependent on her size but is estimated to
be approximately 40 eggs (Brander 1981) which get deposited on the sea bed once
every two years (Little 1995), which is low compared to other European species of
Rajiform. In a study by Koop (2005) on five Raja species (Raja brachyura, R. clavata,

6
R. microocellata, R. montaggui and R. undulate) it was shown that the average
fecundity of females was 117 on an annual basis.

Figure 1.2: Relative numbers of eggs laid per

month of five species of rays in the ‘Ray Reef’ of
Dolphinarium Harderwijk, 1993-2001. Raja
brachyura, R. clavata, R. microocellata, R. montagui
and R. undulate. Adapted from Koop (2005)

However, the skate studied were all smaller species than D. batis (R. brachyura is the
largest of the group at 125cm long, less than half the size of D. batis) and lay smaller
eggs. If the fecundity of D. batis is compared to a Rajiform of similar size such as
Dipturus trachyderma (Roughskin Skate), from the southeastern Pacific Ocean, which
grows up to 258cm long (D. batis 285cm) there is similarity in their fecundity with D.
trachyderma laying between 28 – 68 eggs, depending on the size of the female. This is
similar to the estimate per female for D. batis, supporting it as a reasonable estimate.
The exact breeding time for D. batis is unknown and is hard to estimate because
Rajiformes are able to mate many weeks in advance of the females laying their eggs.
The females are able to store the sperm and then lay eggs throughout a specific season
Figure 1.2 (Koop, 2005). In the case of D. batis is in the spring and summer months
(Ellis and Walker, 2005 and MarLIN, 2008). If we assume that sex segregation occurs,
then breeding can only occur when the sexes mingle, according to previous analysis of
the Glasgow data, this would appear to be in spring time when the proportion of males
increases, this assumes that the sexes do not come together in another area (Little,
1995). The eggs are 14.4-24.5cm long (Ellis and Walker, 2005) and the length of time
before the eggs hatch varies from 2-5 months dependent on temperature (Little, 1995).

7
These life history characteristics suggest that instead of a ‘bloom’ of hatchling skate,
there is a more constant recruitment throughout the spring and summer. A more in depth
study of the spawning cycle of D. batis could confirm fecundity, mating, laying and
hatching season as the timing of the reproduction cycle of different species of Rajiform
can vary considerably. This is highlighted in a study by Koop (2005) in which the
reproduction of different species of captive Rajiform where looked at. All species
displayed a similar reproductive cycle, although the timing and duration of this cycle
varied considerably with some species laying large numbers during April to June season
while other species lay lower numbers but more consistently between November and
July Figure 1.2.
The young of D. batis hatch from the eggs at approximately 22cm long with a
13-15cm wingspan (Brander 1981). They have fully developed spinal columns and
sturdy spines (Walker and Hisplop, 1998) the movement of these hatchling fish is
unknown as they are too small to be targeted by sport anglers and capture by
commercial fishing is unrecorded. If D. batis does segregate by sex and size, then we
would expect to see a mass movement of young out of the area. It is thought that
breeding, laying and hatching occurs within the Firth of Lorne and Sound of Mull due to
the presence of mature males and females during the spring and summer months (Little
1998) and anecdotal evidence from commercial trawl skippers confirming D. batis egg
cases are brought up in trawl nets.

1.6 Pressures
Elasmobranchs are well known for their vulnerability to commercial fisheries, and of
these, skates and rays are arguably the most sensitive (Dulvy et al, 2000). For the same
level of mortality among Rajiform species, it is expected that the longer the maturation
period (Jennings et al, 1998) and the larger the body size (Dulvy and Reynolds, 2001)
the greater the population decline. This places D. batis as one of the most sensitive
species of Rajiform as it is a strong K-strategist (Stevens et al, 2000), has the longest
maturation period for any species of Rajifrom and is the largest species in Europe. Even
the hatchlings of D. batis are vulnerable to commercial fishing due to their size and
morphology (Brander 1981). It is also true that species in higher trophic levels are more
susceptible to decline (Jennings et al, 1999) and the size of D. batis puts it at a higher
level than other Rajifrom species due to the size of prey it can consume, including
smaller species of Rajifrom (Dolgov et al, 2005c). Current estimates suggest that the

8
population of D. batis could be declining by as much as 35% a year (IUCN, 2007). This
large depletion of the population probably leading to significant changes in distribution
(Oddone and Vooren, 2005) as is with the case of R. clavata which appears to be
aggregating in ‘Essential Fish Habitats’ (Payne et al 2008) these are areas were
pressures are fewer and the population is more likely to survive. The history of D. batis
rebuilding former populations is not promising; after the initial depletion of skate
around Orkney the population has shown little sign of restabilising itself after 30 years
(Anon, 2000). This may be down to populations of D. batis being isolated so there are
very few if any resident mature fish to enable repopulation. Elasmobranch species with
the most potential to rebuild populations are short lived, early maturing inshore coastal
species (Stevens et al, 2000) such as smaller species of Rajifrom, which after
exploitation could rapidly (compared to D. batis) rebuild population levels.
One of the main problems is that the UK commercial fishing fleet do not log
Raja catches as individual species, instead grouping all catches of Raja species under
‘Skate and Ray’ (Dulvy et al 2000). Not only does this make it impossible to set catch
quotas for specific species (Agnew et al, 2000) but it also makes determining the impact
a commercial fishery has on a particular species using landing data very inaccurate. D.
batis, however, is vulnerable to commercial fishing (Stevens et al, 2000), and is often
taken as by-catch from long line, benthic and demersal trawl fisheries (Agnew et al,
2000). All skates and rays have some commercial value and commercial vessels will
retain them especially as the exclusion of one species from a fishery is not an easy task.
There is currently only a Total Allowable Catch limit on skate and rays set in the North
Sea but this has historically always been above the total landings for this group (Payne
et al, 2008), rendering it a very ineffective management tool.
It has been shown that in the Irish sea, while larger species of skate such as D.
batis have reduced in number due to commercial pressure (Gallagher et al 2004),
smaller species such as R. montagui (spotted ray) and L. naevus (cuckoo ray) have
increased in number. This may be due to the reasons mentioned previously, with the
smaller species being able to rebuild their population more rapidly than larger species.
This change in community structure has also been noted in the North Sea (Gallagher et
al 2004). The proportion of species being landed is constantly changing and whilst the
landings of skate and rays as a whole may not be substantially declining, the proportion
of D. batis may be declining, but this cannot be detected due to mixed species landing
data (Figure 1.3).

9
 Figure 1.3: Hypothetical situation: Increase in
proportion of smaller species being landed masking
the decline of larger species being landed

Another reason for the sudden increase in proportion of smaller skate could be due to a
change in their life history. It has been shown that long term exploitation of
elasmobranches can lead them to change their life history (Tanaka et al, 1990, Walker,
1999), most noticeably, their growth rates (Stevens et al, 2000) and age of maturity
(Tanaka et al, 1990) which means that these individuals will have a greater chance of
surviving to breeding age (Licandeo et al, 2007). As these are selective attributes which
are partly inheritable, it is expected that the population will evolve over time (Stevens et
al 2000). Walker (1999) showed that this was true of certain Raja species (although not
as yet noted for D. batis), although even if larger species do change life history patterns,
the evolutionary nature of these changes on the population will take longer to occur in
more K-selective species such as D. batis putting them at a further competitive
disadvantage.
As mentioned previously, D. batis was once common around the coasts of the
UK. However, even at the beginning of the 1900s it was clear that the population levels
were rapidly declining. In a study by Rogers and Ellis (2000) survey trawl data from the
Irish and North Seas (in which specimens were identified by species) from 1901-1907
and 1989-1997 was compared. Even in the 1901-1907 D. batis was absent from the
North Sea and only had low numbers in the Irish Sea (maximum 1 per hour trawl time
(Roger and Ellis, 2000)) showing the reaction of the population to early commercial

10
pressure. In the trawls conducted between 1989 and 1997, D. batis was absent from all
trawls. This has led to D. batis being declared extinct from the Irish Sea (Brander, 1981)
and numbers disappearing completely from North Sea trawls (Walker and Hislop, 1998
and Philippart, 1998). Trawl data for the Celtic Sea up until 2003 showed that D. batis
was still prevalent in the area (Walker & Hislop, 1998). However, the trawl survey
around the coast of Britain showed that D. batis was found in the northern North Sea
and Celtic sea only, in water 84-271m deep (Ellis et al, 2005), a lot shallower then the
fish in the Barents sea. It is important to note that during these trawls, there was very
low occurrence of immature fish.

1.7 Potential for Re-populating

For a species with such a late maturation age and a low fecundity such as D. batis,
survival to maturation is the most important factor (Walker and Hislop, 1998). The
response to exploitation showed by some species of Rajiformes of increased fecundity
would do little to increase the population size of D. batis as it is so susceptible to
immature animals being caught. Far more important for species survival is the number
of individual females reaching maturity. However, in a species with such a slow
maturation period the initial survival of adults is paramount for population growth. This
was shown to be the case for D. laevis (Barndoor skate), another large Rajifrom
populating the Western Atlantic (Frisk et al, 2002). It has been documented that the
species is robust and can survive the trauma of commercial capture well (IUCN) so if
commercial vessels can be encouraged to return D. batis, there is a good chance the fish
will survive, dramatically reducing fisheries mortality for this species.

11
 2. MATERIALS AND METHODS

2.1 The Study Area

The data used in this thesis were collected around the Isle of Mull, which lies between
6º23’18” - 5º38’45” and 56º39’25” - 56º15’52” and is located km off the West coast of
Scotland (Figure 2.1)

57
42E3
42E4

B
Passage of Tiree
Loch Aline
56.5 Sound of Mull
A
Isle of Mull
Oban
C
Firth of Lorn

56 41E4

-7 -6.5 -6
Figure 2.1: Study Area around the Isle of Mull, western Scotland, with ICES rectangle numbers
marked for the three used for Fisheries mortality analysis. Also marked a the three Skate angling
areas, A, B and C used to split the Glasgow data into geographic areas.

This area was chosen due to the availability of historical angling records dating back to
1975 collected by various anglers in the area and compiled by Richard Sutcliffe,
(Sutcliffe, 1994) at Glasgow Museums who kindly made these data available through
SNH and they comprise the majority of those used for further analysis presented here.
There were two phases of skate tagging in the area (1974-1988 and 1988-present day),

12
originally initiated (in 1974) by Dr Deitrich Burkel as a result of a dramatic decline in
population numbers around the Shetland and Orkney Isles (Little, unknown date).

2.2 Data Collection

2.2.1 Collection of biological data

Project ID

Tag number

Figure 2.2: Photograph of the Jumbo Rototags used in phase 1 of the Glasgow Museum
Study. Picture courtesy of Richard Sutcliffe.

Figure 2.3: Diagram representing the Jumbo Rototags initially used during phase 1 of the
tagging study. Source: http://www.ketchum.ca/imgs/livestock/p7/versa.gif. viewed on
12/08/2008.

Figure 2.4: Showing the tagging location on the skate for the Jumbo Rototags.

13
In phase 1 (1974 – 1988) Dalton Jumbo Rototags (Little, unknown) were used (Figure
2.2) (normally used for ear tagging livestock). However these proved to be ineffective
because, firstly, the tags offered a large surface area which allowed fouling organisms to
attach, rendering it unreadable, while secondly, the constricting nature of the tag
restricted growth (Figure 2.3) in the area of the fish around the tag, causing sores. The
build up of fouling matter on the tag, which included hard shelled organisms such as
barnacles, along with the rotational movement of the tag also caused damages to the
skin of the animals. (Little, 1995). The Jumbo Rototag was placed on the trailing edge
of the wing of the animal on either side as shown in Figure 2.4.

Figure 2.5: Showing the design of the current Floy Dart Tag with
the barbed anchor at the left end of the tag. Picture courtesy of
Richard Sutcliffe

Figure 2.6: Showing the Needle Applicator and the insertion of the tag into the needle ready
for application. Picture courtesy of Richard Sutcliffe

14
Figure 2.7: Showing the tagging areas on the fish for the new Flow Dart Tag. The body core
surrounding the internal organs is outlined in Grey

Due to the unsuitably of the Rototags, they were replaced by Floy FT-1 Dart tags
(Figure 2.5)(Little, unknown) in 1988. These tags have a single anchoring point, do not
cause injury and offer very little surface area for fouling. A needle applicator (Figure
2.6) is used to insert the anchoring point beneath the skin but above the muscle. The
changeover of tags also represented the end of phase 1 and the beginning of phase 2. At
the time of writing, Floy dart tags are still in use, although they have undergone a design
change by the manufacturer, shortening the tags and changing the colour from orange to
yellow. Floy Dart Tags could not be secured in the area previously used, as the tissue
here was thin and the area subject to a lot of movement during swimming, which had
the potential to dislodge the tags. Consequently they were secured in the more muscular
areas towards the centre of the body (Figure 2.7), but avoiding areas above the internal
organs. These areas provide a more secure anchor point for the tags and also are subject
to less movement, reducing the risk of the tags being lost. (pers. Comm. Davy Holt).
When Fish were initially caught and if they were not already tagged a tag was anchored
to the fish, if a Rototag was present, this was replaced with a dart tag. Log Sheets
provided by Glasgow museum were filled in for each new tag used. The details
recorded were: tag Number, date caught/tagged, time, length (inches), longitude and
latitude, location in words, sex, wingspan (inches), weight (pounds), depth the fish was
caught at (depth of location opposed to length of line out), tidal state (open to taggers
interpretation), bait used, tagger/captor and comments (such as condition of fish, any
unusual features or circumstances). If the fish was already tagged, then the same details
were taken but instead of a new tag being fitted, the number form the existing tag was

15
recorded and the tag left in the fish. There are some circumstances where the tag may be
replaced, if the existing tag is a Jumbo Rototag then this is replaced with a Floy Dart
Tag or if the existing Dart tag is loose and likely to fall out then this should try to be re-
anchored or if the anchoring point is damaged then the tag is replaced.

2.2.2 Extra Field work undertaken towards this project

Swivel to main
line

Line between split Line between split

link and main Size 10 hook link and main
line. 40cm line. 240cm

Swivel with split

15mm plastic pipe link
approximately 30cm. Snap
swivel tied on with cable tie
for attachment of lead
weight (1.5-2kg)

Figure 2.8: Fishing gear used to catch D. batis and prevent deep hooking and cause
minimal damage in the situation where the main line snaps, the weight and
majority of line being detached from the hook. Picture Courtesy of Ronnie
Campbell.

Figure 2.9: Showing the gaffing sights on D. batis

16
 Figure 2.10: Showing the length and Wingspan measurements on D. batis

For the Field work undertaken in this project the Floy Dart Tags (Figure 2.6), provided
by Glasgow Museums, were used along with a needle application tool (Figure 2.7). Log
sheets were created (see appendix 7) which recorded name of skipper, date, weather,
start time of fishing, end time of fishing, time fish caught, longitude and latitude (in
WGS1984) existing tag number, new tag number, sex, depth (of location), weight
(pounds or kg), length (inches or cm), width (inches or cm), tidal state, bait and
comments. Also included were two images representing the dorsal and ventral side of
the fish to for the purpose of marking any areas of damage, this damage could be further
described in the comments box. The addition of start and end time is to allow for future
analysis of fishing time or effort as this is one thing that is currently missing from the
Glasgow data set. These logsheets were given to all skippers in the area running D. batis
targeted charters. Any completed log sheets were collected at the end of the sampling
period. Sampling took place form the angling boat “Laura Dawn” skippered by Ronnie
Campbell operating from the Oban Times Pier in Oban between the dates 06/06/2008 –
26/06/2008 (not fishing every day). Fishing trips started at 9am and lasted 8 hours until
5pm, the amount of time spent fishing for D. batis varied each day due to a number of
external influences. Firstly, the weather was the deciding factor as to the location for
fishing as some, although offering better fishing, were very exposed and were un-
fishable in rough weather so the travel distance between Oban times Pier and the fishing
location varied considerably which affected time spent fishing. Also, the first part of the

17
trip was spent fishing for bait fish, usually mackerel. The length of time spent fishing
for mackerel depended on the number of fish caught, needed a minimum of 20 (personal
observation) to use as bait for D. batis and also if the anglers bought any bait on board
with them such as frozen squid, salmon then not as many mackerel were needed. The
fishing gear used can be seen in Figure 2.8, this was attached to a nylon main line (50-
60lb approx 500 metres long) run from good quality reel with harness links attached to a
50lb class rods, there were commonly 6 rods over the side of the boat, but this was
condition dependent, in rougher weather were the boat was subject to greater
movement, fewer rods were put out to prevent entanglement with each other and the
anchor rope. Bait was routinely checked as other benthic and demersal organisms in the
area removed the bait from the hook. The bait was changed upon every inspection to try
and maximise the scent trail along the sea floor. When a Specimen of D. baits was
caught, the fish was brought to the surface and landed on the boat using gaff hooks, one
for smaller fish, two for larger specimens, through the muscular tissue on the wings of
the fish (Figure 2.9) avoiding the main body cavity. This procedure has been shown to
cause no lasting damage to the fish and is the currently accepted method of landing the
fish (Sharktrust, 2004) although some boat anglers chose not to land the fish at all,
preferring to take measurements over the side of the boat while the fish is still in the
water.

2.2.3. Measurements from landed fish

Once the fish was landed, the length of the fish from the snout tip to the tail tip and the
wingspan, the longest possible width remaining perpendicular to the length axis (Figure
2.10) were measured. The weight was often estimated using weight tables (appendix 9)
as the fish are often too big to be accurately weighted at sea and the vast majority of
anglers do not wish to kill the fish to take it back to shore for an official weight to be
taken. Weights, wingspans and lengths were collected in imperial units as the equipment
the skippers use is in these units as in the angling community, fish are still described in
pounds and the weight charts use length and wingspan units of inches to estimate weight
(in pounds) As information on the relationship between length, wingspan and weight
has gradually been compiled over the years weight tables have been created for each sex
to allow anglers to accurately estimate the weight of their fish from the length and
wingspan measurements, these tables can be viewed in appendix 9. Sex is determined
by the presence of claspers in male fish (Hamlett, 1999) although this can be harder to

18
spot in smaller specimens where immature males have very underdeveloped claspers.
The current practice for anglers is to try and limit the time the fish stays out the water to
three minutes maximum (pers. comm. Ronnie Campbell, Davy Holt and Adrian Lauder)
Readings of position and depth should be taken before the fish has been landed, during
the time the angler is playing the fish on the line, this is often in excess of 20 minutes
due to the length of line out, but these readings can be taken afterwards, before the boat
moves and should ideally be taken directly after the fish has been released. During the
course of the field work which compromised of 12 days at sea, a total of six fish were
caught and tagged, three aboard the Laura Dawn.

2.2.4 Collection of Economic data

As part of the project, it was important to find out how much the population of D. batis
was worth to the local economy as not only do 5 charter boats from the area run regular
trips to fish for the species, but the anglers paying the charter also contribute to the local
economy. To try and find out the worth of the species locally a socio-economic survey
was undertaken. Questionnaires were handed out to the charter boat skippers with
questions aimed at finding out how much they relied on the population of D. batis for
their income. The information they gave also gave insight into how many charter trips
they ran per year and which months they ran fishing trips for D. batis.
The skippers were also given a questionnaire for the anglers to fill in (examples
of both questionnaires can be found in appendix 8). These questionnaires were aimed at
trying to find out if anglers were staying in the area, and if so, which type of
accommodation they used and the group size of the party (including non-anglers). It was
also designed to establish the expenditure of the group in the area, in order to evaluate
out how much each charter boat contributed to the local economy. Aboard the Laura
Dawn, these questionnaires were often filled out by the author directly, while the
skippers from other boats gave the questionnaire out at the beginning of the trip and
collected them at the end. A modified version of the anglers questionnaire, aimed to
target all anglers of D. batis including shore based anglers and anglers with private
boats was given to the SSACN to pass on to their members and also to Davy Holt to
distribute on the skate tagging forum, directed through the SAH. All filled in
questionnaires were returned via email or fax to the author

19
2.3 Data analysis
Before any analysis of the data took place, the Glasgow data fully updated and filtered
for any errors, erroneous and missing data. They were then converted to metric units
(metres, centimetres and kilograms) using the following conversion factors:

1lb = 0.453 592 37 kilogram and 1 inch = 2.54 cm

The majority of the data collected has been measured in imperial units, in all instances,
the data has been converted to metric units (metres, centimetres and kilograms). All
entries for longitude and latitude were converted to decimal degrees, the method for this
varying depending upon the original form of entries. The majority of data were entered
as degrees and decimal minutes DD° MM.MM” although some entries were in degrees,
minutes and seconds DD°MM”SS’. To convert these into decimal degrees the following
formulae were used:
Decimal Degrees = Degrees + minutes/60 + seconds/3600 or
Decimal Degrees = Degrees +MM.MM”/60
The calculations for these were carried out in Microsoft© Excel© 2003 spreadsheet.

2.3.1 Population Segregation

In order to determine if there was any segregation within the population of D. batis in
the Mull area, the data were split into two groups by sex and weight class which were
then both compared against depth in Microsoft© Excel© 2003 and geographic location
(determined by longitude and latitude) in ESRI ARCview. Initially, a Spearmans-rank
correlation was used in the software SPSS to see if there was any realtionship between
weight of fish and capture depth the fish was caught for each sex. Further splitting of
the data into seasonal groups, winter, spring, summer and autumn, allowed for annual
patterns and movements to be highlighted. Each season covered three months with
winter grouping catches in November, December and January, Spring grouping
February, March and April, Summer grouping May, June and July and Autumn
grouping August, September and October. The months for each season were chosen
selected based on three variables, namely months spent fishing by different anglers and
the breeding cycle of other species of Rajifrom taken from the work carried out by
Koop (2005) and are also linked to the breeding cycle of R. clavata as shown in the

20
studies by Hunter et al 2005 a&b (the data were analysed again in Microsft Excel and
ARCview).
The data collected by the Glasgow Museum project were mostly collected by
anglers and as such cannot be considered random sampling, since each angler who
contributed data probably have preferred fishing areas decided by their own personal
fishing history in the area and limitations on where they moore and operate their boat
from. Also, different anglers fish during different times of the year with charter skippers
mainly fishing during summer and Autumn, whereas the pleasure anglers fish all year
round. In order to distinguish any movement patterns displayed by the skate from
patterns created due to angling effort, the data were split into three geographic locations
based on angling activity. These areas were the Sound of Mull (around the mouth of
Loch Aline), referred to as Area A, the Passage of Tiree, referred to as Area B and the
Firth of Lorne below the Junction of the Sound of Mull and Loch Linnhe, referred to as
Area C. These areas are shown on Figure 2.1. Although these areas do not contain all
fish caught, they do represent the majority of catches of D. batis in the study area and
contain the majority of the records with geographic location given. The variables sex
and weight class were then compared to depth within these geographic constraints.

2.3.2 Life History

It has been shown that other Species of Rajifrom have changed their life history
characteristics in response to mortality pressure (Tanaka et al, 1990 and Walker et al,
1999). We wanted to know if this might be the case for D. batis in the study area. As the
Glasgow Museum data has been collected over the last thirty years it was possible to
compare historical information to more recent data. The first thing looked at was the
proportion of males and females and how this altered over time. As anglers target both
sexes equally when they are fishing, it was assumed that fishing effort for both sexes
was equal as no evidence on geographic preference by either sex could be found and the
sexes showed no preference to bait, so proportions of each sex should be representative
of population. Recapture data within the same year was discounted as this might count
the same individual more than once and could lead to false values. Sex ratios for each
year were looked at within each area, as fishing within each area covered different time
periods.
Another variable which was looked at was growth rate of each sex of the
species. Individuals captured more than once with weight data for each catch incidence

21
were included in this analysis. An average value of kilograms change per day was
calculated by dividing weight change by days between captures. This was compared to
growth rates from a smoothed set of data (method outlined below) as most species of
elasmobranchs have different growth rates throughout the year (Kusher et al, 1992). To
smooth the data out and discount large changes caused by annual growth patterns,
changes caused by feeding regimes and the carrying of gametes by females, the total
weight change per individual between initial capture and final recapture was divided by
total day’s freedom to gain an average growth rate in kg/day. In both cases, the figure
for the growth rate was paired with the initial starting weight. The equation of this line
was marked using Excel and used in calculations to create Figures for an estimated
growth curve. Assuming a start weight of 0.5kg the line equation from the growth rate –
weight scatter plot was used to work out the growth rate in kg/day for this weight of
fish. 0.5 was used as an initial start weight by adapting the weight charts in appendix 9
as this figure was added onto the original starting weight (0.5) to give the weight after
one day. This process was repeated for 5475 days (15 years, assuming 365 days per
year) to create growth data for each sex over a period exceeding the cited maturation
period of 11 years (Brander, 1981). This process was repeated four times, twice for each
sex using the smoothed and unsmoothed results.

2.3.3 Movement
The second objective of this project was to try and ascertain if individuals of
either sex undertook any migrations as displayed by individual R. clavata in the
southern North Sea (Hunter et al, 2005b). The recapture data were used to test this. Any
fish that had been caught more than twice i.e. had two recaptures after its initial tagging
catch were spatially analysed using ESRI© ARCview, individuals again being analysed
by sex categories. The distance between each capture location for the same individual
was marked in km and for each fish a total distance between capture sites was
calculated, this was used to see if there were any annual migration patterns for either sex
and to see if there was a difference between the sexes for total distance between capture
points. The literature on maturity size for both sexes of D. batis is unclear and
unconfirmed, so there is no definite weight split to distinguish mature from immature
fish, to overcome this, the data was split into weight classes of 10kg intervals - 0-10kg,
11-20kg, 21-30kg, 31-40kg, 41-50kg, 51-60kg, 61-70kg, 71-80kg, 81-90kg, 91-100kg
and 101-110kg and compared to each other.

22
2.3.4 Population Estimates
As there is no unit of fishing time or effort the only model which could be used
to estimate the population was the Schnabel model (McCallum , 2000). This is similar
to the Peterson method but deals with ongoing sampling as opposed to just two discrete
sampling episodes. The present data were split up into sampling periods so that
recaptures and total catches could be compared over a given time. Three sampling
periods were selected as the basis of population estimates. For all sampling periods, the
same assumptions were applied, these being that the population of D. batis was closed,
the mortality rate was zero, the tagged animals were not more vulnerable to recapture,
death and had no competitive disadvantage compared to untagged fish, there was
complete mixing between tagged and untagged individuals and finally, that none of the
tags were lost. All of the above assumptions will lead to overestimates of the population
but will provide a starting point for further analyses and evaluation of restricting
parameters such as mortality.
Daily sampling period
The first population estimate used all the recapture data except for recaptures occurring
on the same day, but does include fish caught in consecutive days. This is perhaps too
short a time to allow mixing between the tagged and untagged individuals and is likely
to provide the lowest population estimate.
Annual sampling period
For the second population estimate, each year was treated as a discrete sampling
episode, discounting recaptures of fish within the same year, as this discounted many of
the recaptures, as many fish were caught within the same year, this was expected to give
the highest population estimate out of the three methods applied.
30 Days sampling period
The third method is aimed at giving an intermediate estimate by including many of the
recaptures but trying to leave enough time for mixing of the tagged fish with the rest of
the population. As we know so little about the movements and behaviour of D. batis it is
hard to give an exact time period to allow the fish to mix within the population, a time
period of thirty days was used to allow for mixing of the population. This time length
was chosen as most recaptures within a month occur at the same point or within the
immediate vicinity of the original catch point, while recaptures after one month show
greater dispersal which implies that they have mixed with the rest of the population.

23
Fishing Mortality
In order to gain some level of understanding into the mortality from commercial fishing
in the area data provided by FRS in Aberdeen were used as it shows the total skate and
ray catch for the ICES squares 42E3, 42E4 and 41E4 which cover the study area (Figure
2.1). The original population estimates from the above sections are for the whole area so
to work out fishing mortality for this population estimate, the ICES data was added
together for each year to get a total skate and ray landings for the whole study area.
Previous studies on Rajifrom species have run survey trawls in the Irish, Celtic, Barents
and North Sea (Dolgov et al, 2005b and Ellis and Walker, 2005) and have noted skate
and ray landings by species. These studies gave catch levels for D. batis in the form of
either percentage of total catch or average number of individuals caught per hour
fishing. With the information from these studies, a range of hypothetical catch levels as
percentages could be applied to the FRS data to see what the landings of D. batis from
commercial fishing maybe, in kg of gutted D. batis. To convert this number into
individuals proved difficult as the commercial fishing is indiscriminate regarding sex or
size as all life stages of D. batis are vulnerable to being caught (Brander, 1981). The
Glasgow Museum Study was unsuitable to provide an average weight for D. batis as the
data is collected by anglers who target larger specimens. An average weight for D. batis
was gained by calculating the mean value of 50% maximum male weight and 50%
maximum female of fish caught in the area, this gave a mean weight of 88.495kg. The
kg catch was divided by this number to get a hypothetical value for individuals caught
by commercial fleets. This figure was felt to be too vague to apply to the population
estimates so has been left on its own as estimated mortality.

2.3.5 Economic Importance of D. batis

Although the study area focuses on the Isle of Mull and surrounding area, the
only number of viable returns for the economic survey all came from Oban, so the
results are focusing on the economic importance of D. batis to Oban. With the data
collected from the economic survey, the following results were worked out using
Microsoft Excel formula with Minitab used for all statistical analysis. Firstly an average
based on mean values, as the data was parametric as shown in Minitab using the
Anderson Darling test (P value = <0.05), was gained for Money spent in area and an
average based on the median value, as the data was shown to be non-parametric (P
value = >0.05), for money spent on fishing tackle in the area were added together to get

24
a total average spend figure. Then an average, based on mean values (P value <0.05) for
group size was calculated and the total spend figure was divided by average group size
to get average spend per person excluding charter boat costs.
The combined total for chartered days was estimated to be three hundred
between the four charter boats operating from Oban, this was estimated from the returns
of the skipper questionnaires (appendix 8). This multiplied by the average spend per
person per day gave a total contribution of the customers partaking in fishing charters
targeting D. batis to the local economy per year. This was then compared to an
economic report on tourism in the Highlands and Islands (George Street), within this
report was a section on watersports with total money spent in the Highlands and Islands.
The figures gained from the study could then be used to assess how much money skate
fishing contributes this as a whole. The figure for average spent per person on charter
was compared to other watersports, such as fishing, scuba diving and sailing, to see how
much money people spend on each activity.
To assess the value of the Charter boat industry in Oban an average cost of
charter hire per day was estimated from the returns of the skipper questionnaires and
personal observation. This value was divided by the average group size to calculate the
average spend on charter hire per person per day. This figure could then be used to
calculate the total amount spent on chartered trips per year assuming 300 days at sea
total.

2.3.6 Value of Skate compared to commercial market

The SSMEI for Mull project was interested in the average value of individual D. batis in
area. A population estimate had been worked out earlier in the thesis with varying levels
of mortality. The Highest population estimate, i.e. without any mortality added, would
give the lowest value per individual so this was used as a benchmark to estimate an
individual worth for D. batis. The total value of the charter boat industry was divided by
the population estimate to give a value per individual figure. In order to compare this to
current commercial values (which are given in £ per kg), the average weights as used in
the fishing mortality analysis were used to work out the average worth per kg of D.
batis to the Charter boat industry in Oban. This value was then compared to values
taken from the internet.

25
2.3.7 Statistics
The Anderson-Darling test for normal distribution and Levenes test for equal variances
were used in Minitab to determine whether further testing should be parametric or non-
parametric. In all instances in this study, data proved to be non-parametric. Spearmans
Rank tests in SPSS were used to compare the association between variables. Variables
used were depth and weight of fish (appendix 3).
As all the variables looked at were non-parametric, median values have been
used for averages opposed to mean values as they more accurately represent the average
values of the variables.

26
 3. RESULTS

3.1 Behavioural Ecology of D. batis

3.1.1 General Observations of the Population throughout the Study Area
The observed distribution of D. batis within the study area (Figure 3.1) shows that
concentrations of specimens were concentrated in three main areas, the Firth of Passage
of Tiree, the Sound of Mull and the northern end of the Passage of Tiree. These areas
are ringed in black in Figure 3.1. The sexes were distributed equally throughout this
range with neither showing preference for a particular area. Male specimens appeared to
be more closely grouped in the Passage of Tiree and were absent from the west coast of
the Isle of Mull while females were more widely distributed, including a scattering
along the west coast of the island.
There is a noticeable peak in the number of males being caught between 40.01
and 60kg with the largest number of male fish caught belonged to the 40.01-50kg
weight class as shown in Figure 3.2. There were very few fish caught in the 60.01-70kg
class. The numbers of females caught from each weight class shows a bimodal
distribution (Figure 3.3) with numbers of individuals peaking in the 10.01 – 20 kg and
70.01 – 80 kg weight classes. Very few fish heavier than 90.01 kg were caught.

27
 Area B: Based on fishing
effort

Area A: Based on fishing

effort

Area C: Based on fishing

effort

Figure 3.1: Map showing the distribution of both sexes of the D. batis population
around the Isle of Mull. The main map shows all captures of D. batis and the lower,
smaller maps show captures of males and females.

28
Figure 3.2: Histogram showing the number of male specimens caught in the study area for each
weight class between 1975 and 2008. Each weight class has a 10kg interval and the maximum
weight for adult male specimens of D. batis in the area was 74kg.

Figure 3.3: Histogram showing the number of female specimens caught in the study area for
each weight class between 1975-2008. Each weight class has a 10kg interval and the
maximum weight for adult female specimens of D. batis in the area was 102.06kg

29
Figure 3.4 shows the weight related depth distribution of male fish. There was a
significant association between the weight of the fish and the depth it was caught (result
after performing Spearmans Rank correlation 2-tailed p = <0.001). There are two
distinct groupings at different depths, group I, between 25 and 75m and group II,
between 75 and 145m (Figure 3.4). The association between weight and depth can be
seen when looking at the weights of fish caught in each group, group I only had fish
greater than 10kg caught in it while group II had fish between 2-70kg, encompassing
nearly the full range of weights showing that while large fish (greater than 10kg) are
caught at in all depths of water, smaller animals (less than 10kg) are only caught the
deeper water range encompassed by group II. In group I, the depth distribution of fish
between 40 and 60kg is greater (between 25-75m) than fish outside this weight range,
which are all caught at a similar depth (50-60m), there is no similar pattern in group II
with all weights of fish being distributed throughout the depth range (75-145m).
The weight related depth distribution for female fish (Figure 3.5) showed no
association between these two variables (result after performing Spearmans Rank
correlation 2-tailed p = >0.001). There are once again two distinct s depth groups, again
labelled group I, 25-75m and group II, between75-145m. In group I only female fish
weighing between 7kg – 83kg were caught, while in group B, fish between 1.36-
102.6kg were caught, showing larger weight range in group II. It can also be seen that in
group I that there is an even distribution of weights, with no weight being substantially
more represented than another, while in group II there are two groups based on weight,
one between 1.36-30kg and the other between 60-90kg, these two groups are ringed in
blue on Figure 3.5.

30
 0.00

I
-50.00

-100.00
Depth (m)

II
-150.00

-200.00

-250.00
0.00 20.00 40.00 60.00 80.00
Weight (kgs)

Figure 3.4: Scatter plot showing the depth distribution of male fish in relation to weight. Depth
groups are circled in black

0.00

I
-50.00

-100.00
Depth (m)

-150.00 II

-200.00

-250.00
0.00 20.00 40.00 60.00 80.00 100.00 120.00
Weight (kgs)

Figure 3.5: Scatter plot showing the depth distribution of female in relation to weight. Black
circles are showing depth groups and blue circles are highlighting the bimodal distribution of
weight in depth group II.

31
The proportion of males within the study area appears to have declined throughout the
tagging programme as shown in Figure 3.6, with the large drop between 1988 and 1990
where the proportion fell from 70% down to just over 20%. Before this large decline,
the sex ratio was even, with fluctuations, around 50:50, sometimes with a higher
proportion of male fish, sometimes, more females. After 1990, the proportion of males
did not rise above 45% and is currently at its lowest recorded value of below 20%.

Figure 3.6: Showing the change in sex ratio over time. Dashed line showing the trend for the
proportion of males

32
3.1.2 Seasonal observations of the population throughout the study area
When the geographic location of males was plotted with seasonal variation (Figure 3.7),
a pattern could be seen. The numbers of males caught in summer and autumn were
significantly higher than the number of males caught in the spring and winter. It can
also be seen that there is an absence of males from the Passage of Tiree during the
winter and spring seasons, while a large number of males were caught in this area
during summer and autumn.

Figure 3.7: Map showing seasonal distribution of males throughout the study area with
Seasons defined as winter: November-January, spring: February-April, summer: May-July and
autumn August -October.

When the geographic location of females was plotted with seasonal variation (Figure
3.8), it could be seen that the numbers of females caught in summer and autumn was
significantly higher than the number of females caught in the spring and winter. Another

33
pattern that could be seen, although not as distinctly as with males, is the dramatic drop
in numbers of females caught in the Passage of Tiree during the winter and spring
seasons while a large number of females were caught in this area during summer and
autumn.

Winter

Autumn
Figure 3.8: Map showing seasonal distribution of females throughout the study with Seasons
defined as winter: November-January, spring: February-April, summer: May-July and autumn August
-October.

Figure 3.9 shows that both sexes displayed a change in median depth throughout the
seasons, with both sexes being caught at a similar, deeper median depth in the winter
and spring and a shallower median depth during the summer. The median depth for
males was 118.5m deep in the winter and spring and 92m deep in the summer. Females
showed a similar pattern with average winter and spring depths being around 122m

34
during the winter and spring months and 102m in the summer months. During winter,
spring and summer, the average depth for females was lower than for the males with the
largest difference in the summer season. The median depths for both sexes were the
same for autumn and lie approximately halfway between the median spring and median
summer depths.

Season
0.00
Winter Spring Summer Autumn
Winter Spring Summer Autumn
-20.00

-40.00
Depth (m)

-60.00
Females
Males
-80.00

-100.00

-120.00

-140.00

Figure 3.9: Showing the median depth of male and female fish throughout the study.

35
 Figure 3.10: Line graph showing the estimated Growth Curves for Males using Smoothed
and Unsmoothed data for the first 15 years of the animal’s life.

Figure 3.11: Line graph showing the estimated Growth Curves for Females using
Smoothed and Unsmoothed data for the first 15 years of the animal’s life

Comparing the two estimated growth curves for males (Figure 3.10), the unsmoothed
data curve shows a faster initial growth rate, which gives weight differences of up to
12.5kg between the ages of 4 and 9 between the two curves (smoothed and

36
unsmoothed). This shows signs of levelling after 40kg to give a difference between the
two curves of less than 5kgs. The lessening of the difference between the curves is
caused as the smoothed data curve continues to rise relatively steadily, displaying more
linear progression, showing that after the age of six, males experience a higher growth
rate. The smoothed curve shows only a very small decline in growth rate between the
ages of 7 and 15 years.
Looking at the two estimated growth curves for the females, smoothed and
unsmoothed, there is a large difference between them which continues to get larger as
the fish get older instead of converging like the male growth curves in Figure 3.10. The
initial growth rates for females are very rapid, although this is not clear in Figure 3.11
due to the scale of the axis, the smoothed curve, showing the upper estimate, shows
females reaching a maximum of 30kgs towards the end of their first year, while the
estimated weight predicted by the lower, smoothed curve shows female weight to be
approximately 18kgs after the end of 2 years. Assuming a maturation age of eleven
years old (Brander, 1981) the maturity weight for males is 33-42kg and for females,
maturity weigh would be 69-96kg.

3.1.3 Areas A, B and C

As Figure 3.1 shows, there are three distinct clusters of records; this next section looks
at these areas individually in an attempt to remove local variability from the analysis.
Figure 2.1 in materials and methods shows the geographic split based on angling
activity which are similar to the three groups distinguished in Figure 3.1, it is these three
areas, A, B and C that are used in the following figures.
When the sex ratio in each geographic area was compared, it was found
that there was a higher proportion of females in Area A, 70% than in Areas B or C
which both had similar proportions 55 and 52% respectively, this is clearly shown in
Figure 3.12. These proportions were taken from total numbers of males and females
caught in each area as a cumulative total from all years.

37
 Figure 3.12: Column Graph showing the sex ratio in Areas A, B and C

The distribution of males and females in each weight class varies throughout the three
study areas. Figure 3.13 for area A shows a large increase in the number of females
caught in the two weight classes between 60-80kgs, with up to 50 fish per weight class
caught. The weight classes show little variation between the numbers caught, ranging
between 20 (0-20kgs) and 12 (40.04-50kg). Area A had the largest female fish caught in
the study area, weighing 102.6kg. The distribution of males in area A shows a steady
increase in numbers of fish caught up to the 50.01 – 60kg weight class, with no
substantial increase in numbers between each weight class. There were no larger males
than 60kg caught in this area.
In area B (Figure 3.14) there is no substantial peak of numbers of females in any
weight class such as apparent in Area A, with the largest number of females caught
being 40 in the 20.01-30kg class. Males, however, show a large increase in numbers, up
to 70 fish, in the 40.01-50kg and 50.01-60kg weight classes, this a substantial increase
of almost 50 fish from the weight class below (30.01-40kg). Numbers of males caught
in other weight classes are relatively low outside the two peak weight classes, with
numbers of fish caught not rising above 22.
Area C (Figure 3.15) shows large peaks in the number of fish caught for both
sexes. With females, the largest numbers of fish, are caught in the 10.01-20kg weight
class (26), this declines in subsequent weight class to a low of 2 fish caught in the

38
40.01-50kg and 50.01-60kg weight classes. The number of fish then rise again to a high
of 12 individuals caught in the 70.01-80kg class. Males have a substantial rise in
numbers in the 40.01-50kg weight class with 29 individuals being caught. This quickly
drops over the next two weight classes.

60

50

40
Number Caught

Male
30
Female

20

10

0
0-10 10.01- 20.01- 30.01- 40.01- 50.01- 60.01- 70.01- 80.01- 90.01- 10.01-
20 30 40 50 60 70 80 90 100 110
Weight class (kgs)

Figure 3.13: Histogram showing numbers of males and females caught from each weight
class in area A within 10 kg weight classes

80

70

60
Number Caught

50

Male
40
Female

30

20

10

0
0-10 10.01- 20.01- 30.01- 40.01- 50.01- 60.01- 70.01- 80.01- 90.01- 10.01-
20 30 40 50 60 70 80 90 100 110
Weight Class (kgs)

Figure 3.14: Histogram showing numbers of males and females caught from each weight class
in area B within 10kg weight classes

39
 35

30

25
Number Caught

20
Male
Female
15

10

0
0-10 10.01- 20.01- 30.01- 40.01- 50.01- 60.01- 70.01- 80.01- 90.01- 10.01-
20 30 40 50 60 70 80 90 100 110

Weight Class (kgs)

Figure 3.15: Histogram showing numbers of males and females caught from each weight class
in Area C within 10kg weight classes

Figure 3.16 shows the number of males caught per weight class from each area
compared against each other. It can be seen that most fish were caught in area B. In all
three areas there is a similar pattern of gradually increasing numbers of male fish caught
in each weight class until the 40.01-50kg class, which contains the highest number of
fish caught for all three areas. This is most noticeable in area B where the numbers
rapidly increase. The numbers of males in this weight class in areas A and C do rise, but
not as sharply as area B. In all three areas, the peak of the numbers is within either the
40.01-50kg class (areas B and C) or the 50.01-60kg class (area A).
There is no clear pattern in weight class proportion displayed by females (Figure
3.17), all three areas do have some level of bimodal distribution but the highest
representation varies between weight classes. In area A, the largest numbers of females
were caught in the 70.01-80kg weight class, in area B 20.01-30kg and in area C 10.01-
20.01kg.

40
 80

70

60
Number Caught

50
Area A
40 Area B
Area C
30

20

10

0
0-10 10.01-20 20.01-30 30.01-40 40.01-50 50.01-60 60.01-70
Weight Class (kgs)

Figure 3.16: Histogram showing numbers of males per weight class in Areas A, B and C

60

50

40
Number Caught

Area A
30 Area B
Area C

20

10

0
0-10 10.01- 20.01- 30.01- 40.01- 50.01- 60.01- 70.01- 80.01- 90.01- 10.01-
20 30 40 50 60 70 80 90 100 110
Weight Class (kgs)

Figure 3.17: Histogram showing numbers of females per weight class in Areas A, B and C

41
The map in Figure 3.18 shows that most fish caught at less than 75m deep were caught
in area B near in the Passage of Tiree apart from one individual caught from the shore in
area A. Fish caught at more than 145m deep were only caught in Area C in the Firth of
Passage of Tiree. Fish caught between 75m-145m deep were caught in areas A and C.

Figure 3.18: Map showing distribution of depths fish were caught at throughout Areas A, B and C.

Figure 3.19 shows the median depth for fish in area A, which display similar seasonal
depth distribution to the fish caught throughout the whole study area, with both males
and females moving to shallower water in the summer. There is a difference of 11m
between the summer median depth for males and the summer median depth for females
in area A. The median depths for autumn are very similar for males and females, around
107m, being deeper than the median summer depth for males, approximately 114m, and
shallower than the median summer depth for females, approximately 103m. Winter and
spring depths are similar for each sex and only vary by a maximum of 2.5 metres, with
females being caught at a median depth of 116.5m and males, 119m. Both sexes were
caught at the deepest median depth in the spring.

42
 In area B, shown in Figure 3.20, there were no fish caught in the winter and only
1 female caught in spring. The median depth value for the female caught in the spring is
a lot shallower than the median depth for females in the summer. The median catch
depth for both males and females is the same in the summer, approximately 56.5m deep
and in both cases, this rises to a shallower median depth in the autumn, females being
caught at a median depth of approximately 54.75m and males 53.5m. This is the
opposite pattern that we see in area A.
In area C, the sexes display very different seasonal variation in the depth they
were caught in, Figure 3.21. In areas A and B, although the actual median value
differed, the seasonal pattern of depth caught in was the same for each sex between the
seasons. Females show progressively shallow median depths throughout the year, with
their shallowest median depth (111.25m) in autumn and the deepest in the winter
(115.8), while males were caught at their shallowest median depth in the spring
(112.8m), and their deepest during the autumn (116.28m). This is the exact opposite of
the seasonal depths in Figure 3.19.

-90.00
Winter Spring Summer Autumn

-95.00

-100.00

-105.00
Depth (m)

Male
Female
-110.00

-115.00

-120.00

-125.00

Figure 3.19: Column graph showing the median depth values for males and females in area A

43
 -44.00
Winter Spring Summer Autumn

-46.00

-48.00

-50.00
Depth (m)

Male
Female
-52.00

-54.00

-56.00

-58.00

Figure 3.20: Column graph showing the median depth values for males and females in area B

-108.00
Winter Spring Summer Autumn

-109.00

-110.00

-111.00

-112.00
Depth (m)

Male
Female
-113.00

-114.00

-115.00

-116.00

-117.00

Figure 3.21: Column graph showing the median depth values for males and females in area C

Figure 3.22 shows a scatter plot of depth against weight with seasonal data for females
in area A, the association between weight and depth was shown to be significant using
Spearmans Rank, 2-tail correlation test, p=<0.01. The general trend shows larger fish,
50kg+, being caught in shallower water. This is especially noticeable during the
summer, where larger animals are found in shallower water, this can be seen in Figure

44
3.22 ringed in black. The fish caught in the shallowest water is 70kg in weight, heavier
fish were found in deeper water again. In winter, there are three distinct groups, the first
being 7 animals up to 25kg in weight caught in approximately 120m of water, the
second group, two individuals between 20 and 40kg, being caught in 76m of water and
the third group, 5 individuals, between being caught in 115m of water. These three
groups are ringed in blue on Figure 3.22. Most females are caught at the same depth
(between 130-140m deep) in the spring with a few outlying results, mainly between 50-
80kg, in shallower water. Most females caught in the autumn were above 60 kg in
weight and were caught at shallower depths than most fish caught in the spring and
summer time. There was one small specimen caught in the autumn under 10kg at a
depth of 87m, shallower than any of the larger females fish. Larger males, (35kg+) in
area A, in common with females, were found in shallower water up until 55kg, after this
weight, they are caught in deeper water as within the circle region on Figure 3.23, once
again, the association between weight of male fish and the depth was found to be
significant (Spearmans Rank 2-tail correlation test, p=<0.01). This is most apparent in
the summer, where fish between 35-60kg were caught at shallower depths, no fish
lighter than 35kg were caught in less than 100m of water. Two groups are seen in the
spring, with a fish between 5 and 45kg all being caught at a depth of 120m while larger
fish 50-55kg are caught between 90-120m (ringed in blue on Figure 3.23).
Figures 3.24 and 3.25 show the seasonal weight related depth distribution for
males (Figure 3.24) and females (Figure 3.25) in area B. There were no winter results
for either sex and only 1 spring result for females. Looking at the female data for
summer and autumn, we can see that smaller individuals, less than 40kg, are caught
between 50-60m, heavier females, 40kg+, have a greater depth range between 28-72m.
In the summer, these larger animals, 40kg+, were more prevalent at shallower depths,
50-30m deep. Only one individual weighing less than 40kg was caught at a depth of less
than 50m. The autumn female data appears to follow a linear progression (the ranges of
this are marked with black lines on Figure 3.24) with larger fish being found at
progressively shallower depths. The association between weight of female fish in area B
and the depth they were caught at was found to be significant to a 0.05 level (Spearmans
rank correlation test, p=<0.05). This is not true for males, although it appears that some
larger fish, up to 50kg, are found at shallower depths, heavier fish are found at
increasingly deeper depths down to 60m. The depth range for summer increases with
weight, with males above 40kg being caught between 30 and 75m, below 40kg, the fish

45
are concentrated between 50-60m. There was a significant association between weight
and depth for females in area C also (p=<0.05) but the scatter plot for this area did not
show this and so scatter plots fort area C have not been included.

Figure 3.22: Scatter plot showing seasonal weight related depth distribution for females in area A

Figure 3.23: Scatter plot showing seasonal weight related depth distribution for males in area A

46
 0.00

-10.00

-20.00

-30.00
Depth (m)

-40.00

-50.00

-60.00

-70.00

-80.00
0.00 10.00 20.00 30.00 40.00 50.00 60.00 70.00 80.00 90.00 100.00
Weight (kg) Spring Summer Autumn

Figure 3.24: Scatter plot showing seasonal weight related depth distribution for females in area B

0.00

-10.00

-20.00

-30.00
Depth (m)

-40.00

-50.00

-60.00

-70.00

-80.00
0.00 10.00 20.00 30.00 40.00 50.00 60.00 70.00 80.00
Weight (kg) Summer Autumn

Figure 3.25: Scatter plot showing seasonal weight related depth distribution for males in area B

47
3.1.4 Annual Migration
Figure 3.26 shows the longest confirmed distance travelled by D. batis between catch
sites. There have been longer distances recorded, but these are by commercial boats and
their accuracy is dubious (Little, 1995) and so are not been presented here. Fish 1 and 3
where originally tagged in the Sound of Mull and then travelled south to the Firth of
Clyde. Fish 2 was originally tagged in Ballycastle Bay in Rathlin Sound then move
north to the Firth of Passage of Tiree where it was subsequently captured the following
year. These distances are all in excess of 200km. The median distance travelled by male
fish was 6613m and 746m for females (taken from total distance travelled by all fish
recaptured more than once). It can be seen from these median values that males, on
average, travelled greater distances than females.

Fish 1 Fish 2 Fish 3

Figure 3.26: Maps showing the longest recorded ‘anger to angler’ distances travelled by male and
female fish. Fish 1 and 3 were caught in the Sound of Mull and then the Firth of Clyde, fish 2 was
caught in Ballycastle bay, Ireland, then in the Firth of Lorn.

The two movement patterns shown in Figure 3.27 show that the fish were being
caught within the same area on a routine basis. The pattern between catch sites is
an approximate figure of eight shape, with fish moving away from the initial catch
site, and then moving back towards it after subsequent catches. The maximum
distance between recapture sites was 5km

48
Figure 3.27: map showing catch sites of two male D. batis in the Sound of Mull and the Firth
of Lorne. The lines and arrows shown on the map do not show the movement path of the fish,
the fish is likely to have moved beyond the pathway of the line. The lines show direction of
movement between the points and help to provide a time series between catch numbers of
each fish.

Figure 3.28: Map showing the catch sites of two male D. batis in the
Sound of Mull. One specimen is marked with blue circles and blue lines, the
other with blue squares and black lines. Again, the lines are not representing
the exact movement of the fish, but represent the direction travelled and time
series of recaptures.

49
Female fish show a similar pattern as male fish with individuals seeming to move away
from their initial capture site, then move back towards it after the first re-capture. The
fish in Figure 3.29 was shown moving south in the Firth of Passage of Tiree from its
original capture site, before moving north into the Sound of Mull, between the two
geographic areas C and A. This was a longer distance (20km) between re-capture sites
than displayed by males in Figures 3.27 and 3.28. The two females in Figure 3.30 show
similar catch site locations to the male fish caught within the same geographic location.
The fish move away from their initial site of tagging, only to move back towards it after
the first recapture. The fish marked by red circles shows a repeated movement pattern
along the Sound showing an almost circular movement.
Using median averages, the distance between catch sites was greater for males
than it was for females as shown Figure 3.30. On average, the distance between catch
sites or males was 6613m, while for females, this was 736m, this is a difference of
5877m (5.877km).

Figure 3.29: Map showing the catch sites of a female D. batis Between the Passage of
Tiree and the Sound of Mull. The lines are not representing the exact movement of the
fish, but represent the direction travelled and help represent the time progression of
recaptures.

50
Figure 3.30: Map showing the movements of two female D. batis within the Sound of Mull.
One is marked by red triangles, the other red squares. The lines are not representing the exact
movement of the fish, but represent the direction travelled and help represent the time
progression of recaptures.

7000

6000
Average distnce between catch sites (m)

5000

4000

3000

2000

1000

0
Male Sex Female

Figure 3.31: Median distance travelled overall between catch sites for male and females

51
3.1.5 Population Estimates
Figure 3.32 shows population estimates for the area using different assumed sampling
periods. All the estimates show an increasing population with highest and lowest
estimates for 2008 of 3318 and 1961 respectively; a difference of 1356. The middle
population estimate shown is 2341. All lines show a rapid increase in the population
after 1990 with the population continuing to rise until 2008. The annual sampling
estimate shows an initial population decrease after 1979, which is not apparent in the
other estimates. The annual sampling estimate also shows a slight decline in numbers in
2004, which is the same time period as a year of ‘no growth’ for the estimate gained
from the 30 day sampling period. All estimates show a decrease in population in 1985
with numbers dropping by a maximum 200 individuals.

Population Estimates

3500

3000

2500
Number of Fish

2000 All Recaptures

30 Day mixing
1500 Annual

1000

500

0
1978
1980
1982
1984
1986
1988
1990
1992
1994
1996
1998
2000
2002
2004
2006
2008

Year

Figure 3.32: Line graph showing population estimates for the whole study area. Each line
shows a population estimate assuming different study periods. The red line is an estimate
assuming an annual sampling period. The green line assumes a 30 day sampling period
which allows for mixing. The blue line is an estimate assuming a 1 day sampling period,
using all records except those caught on the same day. All estimates are with the
assumption of no mortality.

3.1.6 Fishing Mortality

The general trend is for the number of landed D. batis to decrease (Figure 3.33). This is
true of all assumed proportions of total landings. The highest number of D. batis landed

52
was in 1984 when 198 specimens were landed. This dropped by 62 the following year to
136. After 1984 the levels of D. batis landed did not rise above 85. There was another
dip (down to 49) in numbers being landed in 1994; this, however, recovered in 1996 to
73 fish landed. The lowest estimate shows 0.166 fish landed per year in 2007 while the
upper estimate is 16.65. This is substantially different from the lower estimate of 1.56
and 156 in 1980, almost by a factor of 10.

FIsheries Mortality for D. batis with different % of Total Catch

Scenarios

210

160
Number of Individual fish

0.10%
110
0.50%
1%
5%
60
10%

10
80

82

84

86

88

90

92

94

96

98

00

02

04

06
19

19

19

19

19

19

19

19

19

19

20

20

20

20

-40
Year

Figure 3.33: Line graph showing the estimated landings of D. batis by the commercial
fishing industry in the ICES rectangles 42E3, 42E4 and 41E4 assuming different
proportions of total landings. Assuming an average weight of 88.495kg for D. batis

3.2 Economic Importance of D. batis

From the results of the economic questionnaires it can be seen that charter trips for D.
batis contribute £292,500 to the local economy every year (based on results for 2008).
Of this, £135,000 goes to the charter boat industry and £157,500 goes to the other
businesses in Oban. Each member of an angling party spends £141.53 per day in Oban,
£65.2 of this goes towards charter hire, the rest is spent within Oban town on food and
drink, fuel, accommodation, other activities and goods.

53
 Table 3.1: Showing the estimated amount of money that the D. batis angling
fishery brought to Oban n 2008. The table shows the breakdown of this figure
and how it was achieved by showing average spent per person, number of
people a year the fishery brings to Oban and how much of this money goes to
the local economy.
Amount in
£'s
Total average money spent per trip per group excluding charter 1195.83
Total average spent per group per day excluding charter 525.00
Average spent per person per day excluding charter 76.21
Average spent per person on charter 65.32
Total spent in area per person per day 141.53
Estimated number of people per year 2066
Total spent per day fishing per charter boat 975.00
Total spent annually in area (300 days charter fishing) 292500.00
Contribution to local economy 157500.00
Charter boat industry Gross income 135000.00

Using the median number from the population estimates of 2342 from the entire study
area (this number excludes mortality and therefore is an overestimate; any value placed
on the worth of an individual fish is likely to be an underestimate) we can see that a
single specimen is worth £124.89 in total with a breakdown of worth to local economy
and the charter boat industry shown in Table 3.2
An estimate of the worth of fish in Area C is also useful, as this is the area most
frequented by the charter boats based in Oban (personal correspondence, Ronnie
Campbell) and so the population of D. batis in this area is the target of most charter boat
anglers. The population estimate for area C is 169 and, as the money spent in this area
and on the charter boat industry is all from Oban, we can assume that at least 80% of
this money is spent on targeting fish in Area C. The values for ‘total spent in the area’,
‘contribution to local economy’ and ‘charter boat income’ are used for the calculations
of fish value in Area C, as seen in Table 3.3.

Table 3.2: Showing the value of each fish throughout the study
area, broken down into value as a whole, value to local
economy and value to charter boats.

Value of
individual fish
Value to local economy £67.25
Value to Charter boat industry £57.64
Total Value of fish £124.89

54
 Table 3.3: Showing value for fish in area C, the area heavily targeted by Oban
based charter boats assuming an estimated worth of 80% of total money spent.

Minimum Worth of
0% of money spent in: Amount £'s individual fish
Total Spend on Area C 234000 £1384
Contribution to local economy of Area C 126000 £745
Contribution to charter boat of Area C 108000 £639

When the number of people who come to Oban to fish for D. batis (2066) is compared
to the number of people who participate in all water sports in the Highlands and Islands
(165, 000( George Street Research)), it can be seen that 1.25% of these participate in D.
batis charter trips from Oban.
Of the total revenue spent in the Highlands and Islands by water sports
participants’ 1.56% of that is spent in Oban by charter boat customers. On average,
charter boat customers spend £26.03 per person per day more than do other water sport
participants. The most money is spent on water activities. The money spent on each
activity is shown in Figure 3.36 where D. batis angling trips are the third most
expensive activity after Scuba diving and canoeing/kayaking.

55
 Oban angling trips for D.
batis

Total number of people

participating in water
sports in the Highlands
and Islands

99%

Figure 3.34: Percentage of watersports participants who fish for D. batis from Oban.
Actual figure is 1.25% of total watersprt participants from the highlands and Islands.

Figure 3.35: The percentage of total revenue to area brought by anglers fishing for
D. batis. Actual figures are 1.53%

56
Figure 3.36. Amount spent on each water based activity. Adapted from George Street Research

If the value of D. batis from area C and the entire Isle of Mull population are compared,
it can be seen that fish in area C are much more valuable to the local economy than fish
in the Isle of Mull population (including area C) (Figure 3.37). If this is then compared
to the current market value for skate (currently £2.40 per kg) which includes all species
because skate is sold as skate and is not species specific, it can be seen that although the
total value of the population is less per kg than the current market value, the value of
fish in area C (£13.84 per kg) is comparably much larger than the market value.

16

14

12

10
Price per kg

Fish Market
8 Isle of Mull
Frith of Lorn
6

0
Value of f ish in £ per kg

Figure 3.37: Comparative value of D. batis. Comparing current market price for skate (Grimsby
Fish market, 2008) (mixed box of flesh, so all species included) price per kg of Isle of mull stocks,
using lowest population estimate and price per kg for fish in Area C base on population estimate for
area C and assumed 80% spend of total revenue in area C.

57
 4. DISCUSSION

4.1 Maturation
Broadly speaking, a population of any species can be broken down into four main
groups, mature males, mature females, immature males and immature females, so it is
useful to first get growth parameters for these groups for D. batis so the population can
be studied with reference to them as we may reasonably expect D. batis to segregate its
population based on sex, size and maturity as it has been shown that other Rajifrom
species do so (Ebert et al, 2008). The current literature gives the age of maturation in D.
batis as 11years (Brander, 1981) with males being 125cm long (Fowler and Cavanagh,
2005) and females between 140cm (Walker and Heesen, 1996) and 180cm long (Walker
and Hislop, 1998). Using the weight charts in appendix 9, we can estimate the weight
range that these lengths give. For males, the estimated weight would be 12.7-16.33kg
and females, the lowest estimate would be between 17.23-21.77kg and the upper
estimate between 41.28-51.71kg. Using the estimated growth curves in Figures 3.10 and
3.11 which were taken from recapture data for animals caught more than twice, it can be
seen that the estimated weight range for 11year old fish in both sexes is substantially
higher than those estimated from lengths quoted in the literature for D. batis. The
growth curves given in Figures 3.10 and 3.11 are liable to be overestimates as they take
into account weight change caused by food in the gut in both sexes and the spawning
cycle in females. If fish are initially caught with an empty stomach and subsequently
caught just after feeding, there will be a large weight gain recorded which will
subsequently give a large weight gain from which the growth curves are estimated from.
This effect will be compensated to some extent by fish that were initially caught directly
after feeding with a full stomach then subsequently caught with an empty stomach,
causing the growth rate to be low if not negative, A similar situation occurs with
spawning females as female D. batis produce up to 40 eggs a year (Brander, 1981),
measuring up to 24.5cm long, assuming an average brood size of 20 eggs, and a weight
range of 10-20 g per egg, this could result in an apparent weight change (likely
negative) of 200-400g per fish (likely negative). The other variable which could affect
the estimated growth rate is the use of the weight charts to estimate weight. The
common practice is for anglers is to use the weight charts (personal correspondence
Davy Holt, Ronnie Campbell and Adrian Lauder) as opposed to weighing the fish with
scales at sea (which is in any case difficult on a moving boat). Apart from the 5% error

58
that the weight charts give, there is also error produced by inaccurate measuring of the
fish, as in rough sea conditions, or when measuring the fish in the water, the accuracy of
the measurements may be lessened, even an error of 2.54cm (1 inch) either way can
lead to a difference in weight estimate of up to 11kg (estimated from female weight
charts, appendix 9). This error can also be introduced by intermediate measurements
(i.e. 30.5 inches), the number will be rounded up to the nearest whole number (personal
observation), giving overestimates in weight although the anglers try to keep the
measurements as accurate as possible, allowance for errors should be taken into
account. These errors will overestimate the weight of the fish, in turn overestimating
growth rate.
These combined factors lead to an overestimate of weight gain per day, the
smoothed growth curves were produced to reduce this overestimate, they do however
reduce the growth rate of smaller fish as the growth rate is taken as an average over time
and any early periods of rapid growth are reduced by subsequent slower growth. The
growth curves for males (Figure 3.10) show these differences very clearly, with faster,
logarithmic growth being displayed by the unsmoothed data and lower, more linear
progression displayed by the smoothed. For the females however there is no
logarithmic progression for either estimated growth curves and the smoothed data curve
shows more rapid growth than the unsmoothed curve. The reasons that the opposite of
what we would expect occurred could be due to annual growth patterns displayed by
females, if we look at the growth lines for each fish shown in appendix 1, we can see
that females experience no growth between November and February, these periods of
no growth would show up on the unsmoothed data giving zero growth rate values which
would lower the average value causing the unsmoothed growth curve to be lower than
the smoothed growth curve as the smoothed growth rates would be averaged against
less zero values as the weight change is from a longer period and less likely to be zero
or negative.
Neither of the growth curves show the traditional sigmoid shape (Figure 4.1) but
this is to be expected as D. batis is strongly K selective (Stevens et al 2000) investing
resources into producing well developed hatchlings.

59
 Weight

Time

Figure 4.1: Sigmoid Growth curve.

This will remove the lower section of a typical sigmoid growth curve (ringed in black in
Figure 4.1) as this occurs within the egg case. This is shown to be the case with other
species of Rajifrom such as D. trachyderma (Licandeo et al, 2006) and Bathyraja
parmifera (Alaska skate) (Matta and Gunderson, 2007) and the curve that was produced
for the growth of these two species is similar in shape to the growth curves in Figure
3.10 (smoothed) and 3.11 (both) for D. batis suggesting that they are reasonable
estimates. As the estimated weight at maturity taken from the growth curves is between
33-42kg for males and 69-96kg for females we can assume, due to the overestimating
nature of the growth curves, that the lower value is closer to the true weight. This is
further supported by the scatter plots of weight against growth rate in appendix 2 where
after these estimated weights of maturity, animals started experiencing weight loss,
33kg for males and 64 kg for females, this suggests onset of maturity, as more energy
gets put into gamete production and weight loss is not uncommon in mature animals,
especially during breeding seasons (Ref?). These estimates are significantly different to
the current literature which, compared to this study, appear to be underestimates,
especially in the case of males suggesting that maturation for D .batis occurs at an
earlier age, or that D. batis are heavier upon maturity than previously thought.

60
4.2.1 Population Segregation
The number of animals caught in each weight class throughout the whole area shows
that there is a peak in the numbers of males (Figure 3.2) in the weight classes 40.01-50
and 50.01-60kg. When compared to the estimated weight at maturity it can be seen that
these two weight classes encompass putative mature fish, so it appears that a large
proportion of all males caught in the study area are probably mature. The numbers of
females in each weight class (Figure 3.3) shows bimodal distribution with a large
number of immature females being caught that were between 10.01-30kg and 60.01-90
kg. The largest number of females are in the 60.01-70 and the 70.01-80kg classes which
are mature fish when using the growth curves and allowing for the over estimate. The
high proportions of mature males and females in the region suggest behaviour in
common with other Rajifrom species whereby mature individuals move to shallower
coastal waters during the breeding season (Walker et al, 1997 and Hunter et al 2005b).
Using the growth curves it is possible to estimate the age of the fish in the 10.01-20kg
class to be between 6 months and 2 years old which suggest they are a generation below
the mature adults. If D. batis does move to shallower water to breed and lay eggs, then
the eggs will hatch in the shallower water and it is possible that they remain in this
depth and use it as a nursery until they are strong enough to move into deeper, offshore
waters to mature as we know that D. batis does occur in deep water up to 800m (Dolgov
et al 2005b) but there was no classification of the weight of the animals found at this
depth so it isn’t possible to estimate how old they are and this can’t be used to support
an offshore migration of smaller animals. A possible reason that there is no bimodal
distribution of males could be explained by 2 year old male fish being a maximum of
10kg (using the lower estimate from the growth curves, Figure 3.10) and are too small
to be caught by anglers in the area who target larger fish. The change in diet shown by
R. clavata shows smaller animals consuming largely benthic polychates and amphipos,
this diet would allow smaller fish to retreat to deeper waters and continue feeding while
also avoiding predation. As the fish grow and become less vulnerable to predation, they
also change their diet to more active prey, such as migratory fish, and they will move
into shallower water where these fish are present. (Dolgov, 2005c). It is known that D.
batis preys on Spurdog (Squalus acanthias) in the area (pers. comm. with local
fisherman), this is too large a prey item for smaller D. batis to take and the population

61
of S. acanthias and the Lesser Spotted Dogfish (Scyliorhinus canicula), of which there
are high numbers in the area, would compete with smaller skate to which they have
similar diets, both feeding opportunistically on macrobenthic fauna including hermit
crabs, cockles and whelks (Dolgov, 2005 and Lyle, 1983). These feeding patterns
support the theory that young D. batis move into deeper water to feed and avoid
competition and predation, and then move into shallower water in accordance with a
change of diet and fill a predatory niche with little competition from other species. This
is only a hypothesis and the results from this project cannot support this fully, but it is
an area that would benefit from further research with some targeting of fish smaller than
10kg in weight as well as sampling of D. batis in deeper, offshore water.
An initial look at the geographic distribution of both sexes of D. batis
throughout the study area shows three main areas of clustering at the Passage of Tiree,
the mouth of Loch Aline in the Sound of Mull and the Firth of LornLorn (Figure 3.1)
and it would be easy to assume, when linked to the large proportion of mature fish that
these are possible breeding sites for D. batis. However, with more investigation into
fishing effort it is revealed that the clustering pattern is purely down to fishing effort
throughout the area, with anglers focusing their fishing in these three regions. What is
clear from the geographic distribution throughout the study region is that both male and
female fish are prevalent throughout the area.
Fishing effort is also responsible for the grouping at depths, group I and II, seen
in Figures 3.4 and 3.5. By looking at the depth fish were caught at in each area, A, B
and C it can be seen that all catches in less than 75m were in area B, less than 145m in
area C and equal distribution of fish being caught between 75-145m between areas A
and C (Figure 3.18). Each angler tends to fish within the same area i.e. anglers fishing
area A will rarely fish in area B, they also fish at different times of the year which
appears to be the cause of the variance in seasonal geographic distribution. The main
angler who fished area B only fished the months of March to October with peak fishing
times around June/July (personal correspondence, Brian Swinbanks) while the main
angler fishing area A is often out all year (personal correspondence, Davy Holt) and
anglers fishing area C are generally out all year but with effort focused between March
and October (personal correspondence Adrian Lauder and Ronnie Campbell). This
explains the absence of fish in winter from area B and the large numbers in summer and
autumn; this also offers an explanation for the observed shallower median depth in these
seasons (Figure 3.9).

62
 The obvious biasing of the data by the methods used to collect it makes it
unsuitable to draw any conclusions on population segregation regarding depth and
geographic distribution as a whole. However, if we remove the variable of local
bathymetry and fishing effort by splitting the data into areas where the angling effort
has been focused so all fish caught are subject to the same effort and the same localised
bathymetry, where depth is more likely to be by choice rather than by location, then
actual depth changes displayed by the fish can be looked at. This was done for the fish
in nominal areas A, B and C shown in Figure 2.1 with each area being analysed
separately.

4.2.2 Areas A, B and C

The median depth for each sex between the areas show that both sexes in area A are in
deeper water during the winter and spring (November – April) and in shallow water
during the summer and autumn (June – October), in area B there were no records of
depth for winter and only one record for spring making it unrepresentative, the fish were
at their deepest during summer, although no depth data were available for winter and
spring so no comparison against these seasons can be made. In area C males and
females appear to follow different movements with females being at their deepest in the
winter, progressing to their shallowest depth in the autumn, males however are found in
deep water during the winter, their shallowest depth in the spring, and progressively
deeper water over the summer reaching their deepest depth in the autumn. These
patterns on their own might suggest that area A could be a breeding and/or hatchery
area as the fish are moving to shallower water later in the year in accordance with their
thought breeding season (Fowler and Cavanagh, 2005). Area C, with the rise of animals
to shallower depth in the spring might also be a breeding area from which the males
then descend to deeper water again, while the females remain to lay eggs. In an attempt
to support these hypotheses, the depth distribution was plotted against weight with each
season on the same area plot, the scatter plots for area C showed no patterns of depth
related to weight although, in general, the depth of fish appeared to be slightly shallower
in summer seasons, this was supported by a significant association between the weight
of female fish and the depth they were caught in (Spearmans rank, p=<0.05), there was
no association for the male fish in the area, more reslts from area C need ot be recorde
before any firm conclusions can be drawn. In area A there was a distinct cluster of
females larger than 60kg (borderline mature fish) found in shallower depths during the

63
summer, the records for autumn only included one fish smaller than 60 kg which
happened to be caught in shallow water, assuming this result to be unrepresentative then
a similar pattern during the autumn of larger fish at shallower depths can be seen. For
males, this pattern is even clearer, with no fish less than thirty five kilograms being
caught in less than 100m of water. During all four seasons, but most noticeably in
spring and summer, the larger male fish (mature animals of 35kg and above) were found
at shallower depths, this is similar behaviour displayed by other Rajifrom species
(Walker et al, 1997, hunter et al 2005a) during breeding seasons, supporting the
hypothesis that area A could be a potential breeding or hatchery ground with mainly the
larger, mature fish being found in the shallower depth, to strengthen this hypothesis,
there was a significant association between weight of fish (both male and female) and
the depth they were caught in (p=<0.01). Area B showed similar patterns to area A for
both sexes, although only females showed a significant association (p=<0.05), with
mature fish rising to shallower water during the summer and autumn, this was again
more distinct with males with no male fish less than 35kg being found in water
shallower than 50m. Although area B is locally shallower than area A, it might be
supposed that it is not the actual depth of water that is important, but the movement
from deeper to shallower water that matters, in order for mature adults to congregate
away from immature individuals or groups. This is assuming that both areas A and b are
used for breeding purposes and cannot be assumed to be true. There were no records for
winter and only one for spring so no comparisons against these seasons can be drawn
These are hypothesis based on the known behaviour of other species of Rajifrom
and although are only suggested by the data in this report are worth consideration and
further research. There is not enough information to draw any conclusions for area C.

4.3 Migratory Behaviour

Other species of Rajifrom do undertake annual migrations moving into offshore waters
during winter and summer months and moving back into coastal waters during the
summer months this movement appears to be linked to the breeding cycle, with the
population moving to coastal waters during the breeding season (Walker et al 1997 and
Hunter et al 2005b), it would be reasonable to assume from this that the fish breed and
lay eggs in shallower, coastal waters (Hunter et al, 2005a). The records of migratory
behaviour for D. batis are few, all using the Glasgow museum data, the conclusions
drawn from these papers were that individual D. batis are associated with a regional

64
population with a few individuals performing long migrations (Sutcliffe, 1994 and
Little, 1995). The results gained from this study generally support these hypotheses as
there are several examples of fish being caught repeatedly within a small geographic
area, displaying recapture patterns that suggest confinement to a small area,. Caution
must be used; however, when looking at the recapture sites as, although they show were
the fish has been caught and were the fish has moved to, they do not show movement in
between the catch sites, we may only be looking at a small section of a larger pattern.
Take the example in Figure 4.2, if the fish normally performs the migration shown by
the solid black line but we only sample it within the boxed section, we might assume
that the fish only moves within that area and the assumed route is shown by the dotted
line. So although it cannot be said from the recapture data that the fish remains within
one area throughout its life, it can be said that it returns to the same area on a regular
basis.

Figure 4.2: Diagram to of hypothetical migration route with

small sampling area in the centre of the route. Example of
misleading recaptures.

The long migrations shown in Figure 3.26 prove that not all fish remain within a small
geographic area. It cannot be assumed that the fish have moved from one population to
another as we might only be sampling two points on a circular route, this is shown in
Figure 4.2 where the two red circles represent sampling points, if we only catch a fish
once at point X and once at point Z afterwards, it would be easy to assume that the fish
had moved to point Z when in actual case the points are on a continual route. Of the fish
that have been confirmed to make long journeys two, a male and a female, are recorded
as having moved from the Isle of Mull to the Firth of Clyde in a southern movement, the
other is a female moving from Ballycastle Bay (Ireland) to the Isle of Mull in a
northerly movement. It is possible that these fish are performing similar routes between

65
the Isle of Mull and the Firth of Clyde but due the timing of catching, different
conclusions for each fish could be drawn. The frequency and number of these
migrations is unknown, partly due to the fact that fishing effort for D. batis is focused
around Mull so the low record of long movements may be due to the lack of targeting
outside of the study area rather than due to low numbers of fish moving. The other
records of long migrations are from commercial vessels and there is a certain amount
doubt about the location of the catch sites (Sutcliffe, 1994). A tagging programme in
other areas, such as the Firth of Clyde where some of the Isle of Mull fish have moved
to might increase our understanding of the movements of D. batis along the west coast
of Scotland and would give insight into the regularity of these movements and whether
or not they are one way migrations leading to recruitment into other populations or
circular movements such as displayed by R. clavata in the Southern North Sea (Hunter
et al 2005b).

4.4 Population Estimates

Figure 3.32 suggest that the population of D. batis in the area is increasing. After talking
to local fishermen, this does not seem unreasonable as some would agree that the
population has been increasing over the last decade. Yet it would be unwise to assume
from these figures that the population of D. batis in the area is increasing as there are
other factors which can affect the population estimates, one of these is tagging effort. In
the beginning of the project there were few anglers participating in the tagging
programme, the common practice was to return the fish to sure for confirmation of
weight landbased. This became less common and from the early 1990’s and more
anglers began to participate in tagging in 1993, a concentrated tagging effort began in
the Sound of Mull and in 1994, the Firth of LornLorn in; these years corresponds
directly to the beginning of a rapid increase in population levels shown on Figure3.32, it
can therefore be assumed that tagging effort is a major variable in the population
estimates and it cannot be assumed that the increase in population size shown in Figure
3.32 is due solely to an increase in population. If the graph for population estimate
shown in Figure 3.32 is compared to the graph in appendix 5 showing the number of
fish caught each year discounting recaptures within the same year, it can be seen that a
decrease in numbers caught directly relate to years with a decrease or zero increase in
population, this further demonstrates that population estimates are heavily related to
tagging effort. Little (1995) produce a population estimate of 60 -90 from the Glasgow

66
data, this was at the beginning of the increased tagging effort and so was likely to be an
underestimate due to the lack of data being returned. We now know that the population
is at least 110 fish as this was the maximum number of fish caught in one year and this
would assume that 100% of the D. batis in the area had been landed. We can, however,
use the population estimate in 2008 as a start point for future estimates and might prove
a useful baseline to compare to any future research, but it must be emphasised that this
is an overestimate and should not be used for any management plans, nor should it be
assumed that there is a resilient population of D. batis in the area.
If there is an increase in the population, this might not be solely attributable to
recruitment through breeding, since there have been examples of Rajifroms
congregating together in what is referred to as essential fish habitat (EFH) (Payne et al,
2005), it could be that the waters around the Isle of Mull are an EFH for D. batis. This
would certainly explain why, in most other areas around the UK, D. batis is reported as
being virtually extinct (Brander, 1981, Roger and Ellis, 2000). Although at initial glance
this could partially explain the large increase in population, the bias put on the data by
fishing effort does not permit this conclusion to be drawn directly, although it cannot be
discounted and needs further research with data collection more suitable for population
estimates.
On a cautionary note, all estimates are overestimates, as the Schnabel model
used does not add in any level of mortality. It also assumes that the population around
the Isle of Mull is closed, we know that this is not the case as there are examples of
individual animals both leaving and incoming to the area with up to 200km between
catch sites (Figure 3.26). This shows that there is some degree of mixing outside the
study area in this project although to an unknown degree. Another assumption for the
model is that fish caught and tagged recover 100% and are at no competitive
disadvantage to uncaught fish. This may not always be the case. One of the problems
reported by some skate anglers is ‘deep hooking’ This is where the hook is taken down
into the stomach and can either tear the inside of the stomach, resulting in death of the
fish, or can remain impaled in the lining, causing feeding problems and weight loss. The
main reason for deep hooking is the use of inappropriate fishing gear sometimes used by
private boat angler or shore anglers, especially those not directly targeting D. batis.
Deep hooking is sometimes a problem that cannot be avoided, due to the nature of skate
landing on the bait and consuming it without performing a ‘run’ alerting the angler to
‘strike’ the fish, which would tend to set the hook in the mouth. Taking this into

67
account, it is fair to assume that some of the fish returned may be at a disadvantage to
other fish and also may have a higher mortality rate(Committee on the Recreational
Fisheries et al). There is a fishing code of conduct for D. batis which should be
followed to minimise stress and damage to the animal, increasing its chances of survival
after capture (available from Richard Sutcliffe at Glasgow Museum Resource Centre)
The estimated fisheries mortalities all show the same pattern, a declining number
of Rajifromes being landed. Due to the nature of Rajifrom landings all being grouped
under ‘skates and rays’ the estimate for the proportion of this catch being D. batis works
around assumed proportions of total catch and assumed average weight for D. batis
which itself assumes that equal numbers from each weight class were caught. This
makes the estimates liable to be inaccurate and not reliably representative of the actual
D. batis take. It is likely that landings of D. batis have decreased, but to what extent is
unknown without species specific landing data. Assuming a level of 10% of total catch,
which is higher than any of the percentages in research trawls performed in other seas
(Dolgov et a, 2005, Wlaker and Heesen, 1996 and Ellis and Walker, 2005), the pattern
for most elasmobranch species in the southern North Sea is of declines in landings
(Philippart, 1998), which is also true for the skate and ray landings from around Mull.
Fishing effort in the area has not decreased substantially, and in fact the most days spent
out of harbour were between 1994 and 1999. Reported landings of skate and rays do
increase during this period, but not above levels pre-1986, suggesting that the numbers
of skate and rays in the area are decreasing, most likely in response to fishing pressure,
to which they are particularly vulnerable (Dulvy et al, 2000). Since D. batis is one of
the most sensitive species to fishing pressure (Stevens et al, 2000) we can assume that
there has been a proportionally greater drop in the numbers of D. batis being taken,
compared to other Rajifroms.

4.5 Economic Importance

The returns from the economic survey were not sufficient to conduct in-depth analysis
into the economic value of D. batis to the area. Nevertheless, there were enough to
make some general overviews as to how much money anglers brought to the area. For a
relatively small industry, the charter trips bring a lot of people to the area, most of them
travelling especially to the area for D. batis angling trips. Although the proportion of all
water sport participants is only 1.25%, it must be remembered that skate angling is a
specialist sport, with many anglers coming to Oban especially for it, staying for several

68
days and fishing for most of them. Proportionally, the amount of money spent by the
anglers chartering boats is higher than the average spend by other water sport
participants and contributes 1.53% of total income to the Highlands and Islands. It is
hard to work out what percentage of money coming into Oban from water sports skate
angling represents as there are no figures by town or even council area as to how much
money is contributed by the local economy. The figures for contribution from skate
anglers are estimates and would need more results to substantiate them; however, it is
clear that the money brought into Oban by skate anglers is relatively large per capita.
It appears from the results that fish in area C are far more valuable to the local
economy than they are to the commercial industry, and it must also be remembered that
skate caught by anglers and returned can be caught again, which doubles the value of
that fish as two charter boat trips have enjoyed the benefit of catching that animal,
which will in turn encourage anglers to return to the area to fish for D. batis again.
Although the average estimated value of each fish throughout the whole study area is
lower than the current commercial price per kg, it must be remembered that this value
was obtained from a probable overestimated population level, which in turn,
underestimates the value of the fish, so it is likely that the true value of D. batis is
higher than that quoted in this here (£1.14 per kg)

4.6 Life History

Although the Glasgow museum provides an excellent historical record of D. batis in the
area, there were not enough records for every year with all the necessary data recorded
to accurately assess any change in the life history of the population in response to
pressure. Other species of elasmobranchs have been shown to do this (Tanaka et al,
1990), changing growth rates and age of maturity (Tanaka et al, 1990 and Stevens et al,
2000). Although it is possible that D. batis could change its age at maturity by
increasing its growth rate and lessening its age of maturity, there is no evidence that this
is the case. The results from this study strongly suggest that the age of maturity for D.
batis around the Isle of Mull is 11 years old. This is the same age for maturity given 20
years ago (Brander, 1981) and cannot be seen to have altered. Due to the long life, late
maturity and strong K selectivity of D. batis any changes to life history characteristics
would be expected to occur over several generations so we would not expect any
changes to happen quickly.

69
 5. CONCLUSIONS

5.1 Conclusions from this study

In conclusion, this investigation was able to show that immature and mature individuals
of D. batis are present throughout the northern Passage of Tiree, the Sound of Mull and
the Firth of Lorn. There was no observed geographic preference of either sex throughout
the study area, although seasonal preference for one area was not attainable from the
results due to the seasonal change in fishing effort in different areas.

There were clear trends showing annual movement of mature individuals of D. batis, of
both sex, from deep water to shallow water in relation to the cited breeding cycle (Ellis
and Walker, 2005) in certain areas which strongly suggested the preference of D. batis
to breed and lay eggs in shallower water during the spring and summer in areas A and
B. There was no preference to an actual depth displayed by either mature or immature
individuals of either sex, this appeared to be related to local bathymetry. There is a
strong suggestion that larger animals are found in shallower water, this association was
shown to be significant for females in all areas, with the strongest association in area A
(Spearmans Rank 2-tailed correlation test, p=<0.01) and males in area A and the study
area as a whole (p=<0.01). Careful management of areas A and B might aid the
rebuilding of the population around the Isle of Mull, this in turn, due to proven
movements of D. batis, might help the population growth along the west coast of
Scotland.

It is also clear that the Isle of Mull population is not closed, this can be seen by the long
migrations of both sexes shown in figure 3.26. It is unknown to what extent these
migrations occur and whether or not they are one way or cyclic.

This study also calls into question previous estimates for length and associated weight at
maturity for D. batis. It is true that either D. batis matures at an earlier age than
previously thought, or, that, the previous estimates for length at maturity from other
papers are under estimates.

No clear conclusions can be drawn from the population estimates as the results are
dependent onto many assumptions and are heavily influenced by the number of tag

70
returns. What is clear is that the tagging data needs more information, such as a record
of time spent fishing for D. batis to enable a more in depth study into population levels
is needed to determine the current status of the Isle of Mull population. As with
population estimates, as we have no official record for the landings of D. batis it is
unclear what impact the commercial industry has on the Isle of Mull population as this
study assumes landing levels. Landing data recording individuals of D. batis is needed
to look into the fisheries mortality.

Charter trips for D. batis bring a substantial amount of money to Oban, but it is unclear
what proportion of total income this contributes to Oban itself, although to the
Highlands and Islands as a whole, it is a relatively large amount for such a small
industry. What is clear is that individual fish in the area are worth more to the angling
industry than they are to the commercial market and, due to their ability to survive the
trauma of netting, are worth returning to the sea upon capture.

5.2. Further Areas for Study

The main conclusion drawn from this study is the need for further research into D. batis,
not only in the waters around the Isle of Mull, but around the whole UK. If this species
is to be preserved effectively, more knowledge on its behaviour is desperately needed to
ensure its future, this study has opened the doors to many areas of potential research and
I would suggest the following areas as targets for further study:
• Genetic study. Looking into the mixing between regional populations of D.
baits. This would indicate to what level populations mix and whether or not
regional populations are closed or not. This is essential as closed populations are
more vulnerable to fisheries mortality and are less likely to survive in exploited
areas.

• Study into the annual migration of both sexes of D. batis. Using electronic data
tags or/and sonar tags to track the movement of individual skate more closely to
see how common long migrations are and if they are cyclic by nature or one
way. Also will show any annual migrations.

71
• Study into the weight classes found in various areas of sea including offshore.
This will further our understanding into the movements of young D. batis and
will substantiate if they do indeed move into deeper water offshore to mature.

• Sampling for D. batis throughout all waters around Mull. This will lead to better
understanding of any geographic congregation areas for D. batis and will help in
the allocation of breeding and hatchery areas for the species, allowing more
effective management.

• An in depth study into the breeding cycle of D. batis to confirm the months of
breeding and egg laying for the species, again, to allow more effective
management.

• A more in depth study into the economic worth of D. batis to the Oban area,
looking for contribution to local economy.

• Collection of egg case data via trawl surveys and commercial trawlers. This will
help us to understand the egg laying nature of D. batis and if females do
congregate in a common hatchery, where these areas are.

72
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the Barents Sea. Journal of Northwest Atlantic Fisheries Science. 35. 249-260

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Dolgov, A.V., (2005c) Feeding and Food Consumption by the Barents Sea Skates.
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Ellis, J. and Walker, P. Compiled by Fowler, S.L and Cavanagh, R.D. (2005) Grey,
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Chimeras. IUCN 333-335

Gallagher, M.J. Nolan, C.P and Jeal, F. (2004) Age, Growth and maturity of the
Commercial Ray Species from the Irish Sea. Journal of Northwest Atlantic Fisheries
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Grimsby Fish Market. Price List. Available from:

http://www.grimsbyfishmarket.co.uk/frames/Prices.html. Viewed on 15/08/2008

Hamlett, W.C (1999) Sharks, Skates and Rays: The Biology of Elasmobranch Fishes.
The Johns Hopkin University Press. Baltimore. 1

Hedges, Andrew. 2003 Convert Latitude/Longitude to Decimal. Direct link:

http://andrew.hedges.name/experiments/convert_lat_long/ viewed on 26/06/2008

Holden, M.J. (1975) The Fecundity of Raja clavata in British waters. Journal de
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Hunter, E., Buckley, A.A., Stewart, C. and Metcalfe, J.D. (2005a) Migratory behaviour
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Biological Association of the United Kingdom. 85. 1095-1105

Hunter, E., Buckley, A.A., Stewart, C. and Metcalfe, J.D. (2005b) Repeated seasonal
migration by a thornback ray in the Southern North Sea. Journal of the Marine
Biological Association of the United Kingdom. 85. 1199-1200

IUCN (2007) IUCN Red list of Threatened Species: Dipturus batis. [online]
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Jennings, S., Reynolds, J.D and Mills, S.C. (1998) Life history correlates of ersponses
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Jennings, S., Greenstreet, S.P.R. and Reynolds, J.D. (1999) Structural change in an
exploited fish community: a consequence of differential fishing effects on species
with contrasting life histories. Journal of Animal Ecology. 68. 617-627

Koop, J.S., (2005) Reproduction of captive Raja spp. in the Dolfinarium Harderwijk.
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Kusher, D.I. Smith, S.E and Cailliet, G, M.(1992) Validation age and growth of the
leopard shark, Triakis semifasciuta. Environmental Biology of Ifhses. 35. 187-203

Licandeo, R., Cerna, F. and Céspedes, R. (2007) Age, growth and reproduction of the
roughskin skate, Dipturus trachyderma, from the southeastern Pacific. Ices Journal
of Marine Science 64 141-148

Little, W. (1995) Common Skate and Tope: First Results of Glasgow Museum’s
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Little, W. (1997) Common Skate in the Sound of Mull. Glaucus. Vernal/Summer. 42-
43

Little, W. (1998) Tope and Skate Tagging off West Scotland: Part 2. Glaucus.
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Little, W. (date unknown) Tope & Common Skate tagging off the West of Scotland - an
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on 09/07/2008

Lyle, J. M. (1983) Food and feeding habits of the lesser spotted dogfish, Scyliorhinus
canicula (L.), in Isle of Man waters. Journal of Fish Biology. 23, issue 6. 725-737

MarLIN, The Marine Biological Association. Direct link to Common skate (Dipturus
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Blackwell Publishing. 87-95

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Oddone, M.C. and Vooren, C.M. (2005) Reproductive behaviour of Atlantoraja
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Philippart, C. J. M. (1998) Long-term impact of bottom fisheries on several by-catch

species of demersal fish and benthic invertebrates in the south-eastern North Sea.
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Rogers, S.I and Ellis, J.R. (2000) Changes in the demersal fish assemblages of British
coastal waters during the 20th century. ICES Journal of Marine Science. 57. 866-881

Sharktrust, (2004) Common Skate code of conduct. Available from Glasgow Museums.

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sharks, rays and chimeras (chondricthyans), and the implications for marine
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Stephen, A.C. (1929) Large skate from the Firth of Clyde. Scottish Naturalist. 175. 94

Sutcliffe, R. (1994) Twenty years of tagging common skate and tope off the west coast
of Scotland. Shark, Skate and Ray Workshop. London. 15-16 February. 14-16

Tanaka, S., Cailliet, G.M. and Yudin, K.G. (1990) Difference in growth of the Blue
Shark, Prionace Glauca: Technique or Population. Elasmobranchs as Living
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Fisheries. U.S Department of Commerce, Washington. 90. 177-188

Walker, P.A. and Heessen, H.J.L. (1996) Long-term changes in ray populations in the
North Sea. ICES Journal of Marine Science. 53. 1085-1093

Walker, P., Howlett, G. and Millner R. (1997) Distribution, movement and stock
structure of three ray species in the North Sea and eastern English Channel. ICES
Journal of Marine Science. 54. 797-808

Walker, P.A. and Hislop, J.R.G. (1998) Sensitive skates or resilient rays? Spatial and
temporal shifts in ray species composition in the central and north-western North Sea
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Walker, P.A. (1999) Fleeting Images Dynamics of North Sea Ray Populations. PhD
Thesis.

ZeeInZicht Digitale Encyclopedie. Direct Link:

http://www.zeeinzicht.nl/vleet/index.php?item=blue+skate. Viewed on 15/06/2008

77
 Appendix 1. Annual Growth patterns of individual fish

Figure A.1: Individual growth curves for females showing annual growth patterns.

Figure A.2: individual growth curves for males showing annual growth patterns

78
 Appendix 2: Growth rate versus Weight Scatter Plots

0.25

0.2

0.15

y = -0.0688Ln(x) + 0.306
0.1

0.05

0
0 10 20 30 40 50 60 70

-0.05

-0.1

Figure A.3: Scatter plot for weight against growth rate for males using smoothed data, showing
logarithmic trend line and equation.

0.05

0.04

0.03
Growth rate (kg per day)

0.02

0.01
y = -0.0127Ln(x) + 0.0644

0
0 10 20 30 40 50 60 70

-0.01

-0.02
Weight (kg)

Figure A.4: Scatter plot for weight against growth rate for males using unsmoothed data,
showing logarithmic trend line and equation.

79
 0.12

0.1

0.08
y = -0.0214Ln(x) + 0.1313
Growth rate (kg per day)

0.06

0.04

0.02

0
0 20 40 60 80 100
-0.02

-0.04

-0.06
Weight (kg)

Figure A.5: Scatter plot for weight against growth rate for females using smoothed data, showing
logarithmic trend line and equation.

0.35

0.3

0.25
Grwoth rate (kg per day)

0.2

0.15
y = -0.0492Ln(x) + 0.2888
0.1

0.05

0
0 20 40 60 80 100 120
-0.05

-0.1
Weight(kg)

Figure A.6: Scatter plot for weight against growth rate for females using unsmoothed data,
showing logarithmic trend line and equation.

80
 Appendix 3: Spearmans rank results for Weight against Depth

Table A.1: Spearmans rank results for all males and females in study area.
Significant association between weight and depth for males

Depth Weight
Spearman's rho Depth F Correlation Coefficient 1.000 -.065
Sig. (2-tailed) . .061
N 829 827
Weight Correlation Coefficient -.065 1.000
F Sig. (2-tailed) .061 .
N 827 830

Depth Weight
Spearman's rho Depth M Correlation Coefficient 1.000 .179(**)
Sig. (2-tailed) . .000
N 448 448
Weight Correlation Coefficient .179(**) 1.000
M Sig. (2-tailed) .000 .
N 448 468

Table A.2: Spearmans rank results for males and females within each are, A, B and C.

81
Appendix 4: Distance travelled by each fish, male and female

Table A.3: Total distance travelled by each fish re-caught

more than 3 times. Median values shown at the bottom.

Sex Female Male

134093 8530
0 5525
Distance travelled 0 4458
(metres) 5663 0
0 9383
7859 8543
100 0
150 8733
0 298
0 8561
0 7787
0 5768
16346 224990
509 10744
752 1555
1904 1295
250 895
2178 10243
8757 3079
11012 11046
4178 7458
4217 4765
5744
5527
9947
317
8062
150
200
150
243330
100
100
250
4887
1415
740
100
32104
800
8501
100
253020
0

Median 6613 746

82
Appendix 5: Total number of Fish caught each year within the study area.

120

100

80
Number caught

60

40

20

0
77

79

81

83

85

87

89

91

93

95

97

99

01

03

05

07
19

19

19

19

19

19

19

19

19

19

19

19

20

20

20

20

Year

Figure A.7: Number of fish caught discounting recaptures within the same year within
the study area

83
 Appendix 6: Tables used for population estimates.

Table A.4: Showing information needed for the Schnabel model to

estimate population assuming a sampling period of one day with no
same day recaptures included.

Numbers
using
YEAR Ct Rt Ut Mt CtMt (CtMt)/Rt
1973 0 0 0 0 0 0.00
1974 0 0 0 0 0 0.00
1975 1 0 1 0 0 0.00
1976 0 0 0 1 0 0.00
1977 28 1 27 1 28 0.00
1978 23 4 19 28 644 134.40
1979 28 3 25 47 1316 248.50
1980 12 2 10 72 864 285.20
1981 9 2 7 82 738 299.17
1982 15 3 12 89 1335 328.33
1983 15 4 11 101 1515 338.95
1984 16 3 13 112 1792 374.18
1985 46 6 40 125 5750 499.36
1986 8 7 1 165 1320 437.20
1987 24 8 16 166 3984 448.51
1988 26 5 21 182 4732 500.38
1989 48 5 43 203 9744 637.02
1990 38 4 34 246 9348 756.32
1991 16 6 10 280 4480 755.40
1992 30 3 27 290 8700 852.88
1993 65 15 50 317 20605 949.32
1994 45 19 26 367 16515 934.10
1995 70 24 46 393 27510 975.16
1996 54 29 25 439 23706 945.27
1997 38 7 31 464 17632 1014.11
1998 64 9 55 495 31680 1147.56
1999 62 12 50 550 34100 1259.88
2000 58 13 45 600 34800 1354.84
2001 55 15 40 645 35475 1427.33
2002 101 19 82 685 69185 1611.83
2003 100 26 74 767 76700 1748.81
2004 91 37 54 841 76531 1789.45
2005 109 39 70 895 97555 1873.59
2006 71 27 44 965 68515 1923.81
2007 41 14 27 1009 41369 1962.72
2008 7 4 3 1036 7252 1961.12

Population estimate using all recaptures except same day catches.

84
Table A.5: Showing information needed for the Schnabel model
to estimate population assuming a sampling period of one year

Numbers
using
YEAR Ct Rt Ut Mt CtMt (CtMt)/Rt
1973 0 0 0 0 0 0.00
1974 0 0 0 0 0 0.00
1975 1 0 1 0 0 0.00
1976 0 0 0 1 0 0.00
1977 27 0 27 1 27 0.00
1978 21 2 19 28 588 307.50
1979 25 0 25 47 1175 895.00
1980 12 2 10 72 864 663.50
1981 9 2 7 82 738 565.33
1982 14 2 12 89 1246 579.75
1983 14 3 11 101 1414 550.18
1984 16 3 13 112 1792 560.29
1985 44 4 40 125 5500 741.33
1986 8 7 1 165 1320 586.56
1987 24 8 16 166 3984 565.09
1988 25 4 21 182 4550 626.97
1989 46 3 43 203 9338 813.40
1990 38 4 34 246 9348 951.91
1991 16 6 10 280 4480 927.28
1992 30 3 27 290 8700 1038.94
1993 53 3 50 317 16801 1283.30
1994 32 6 26 367 11744 1348.53
1995 51 5 46 393 20043 1547.04
1996 33 8 25 439 14487 1575.19
1997 34 3 31 464 15776 1716.86
1998 59 4 55 495 29205 1989.27
1999 59 9 50 550 32450 2149.12
2000 48 3 45 600 28800 2386.91
2001 46 6 40 645 29670 2540.40
2002 92 10 82 685 63020 2882.36
2003 84 10 74 767 64428 3179.07
2004 80 26 54 841 67280 3073.75
2005 88 18 70 895 78760 3216.63
2006 59 15 44 965 56935 3265.16
2007 34 7 27 1009 34306 3326.72
2008 5 2 3 1036 5180 3318.88
Population estimate using an assumed annual sampling period

85
 Table A.6: Showing information needed for the Schnabel model
to estimate population assuming a sampling period of 30 days

Numbers
using
YEAR Ct Rt Ut Mt CtMt (CtMt)/Rt
1973 0 0 0 0 0 0.00
1974 0 0 0 0 0 0.00
1975 1 0 1 0 0 0.00
1976 0 0 0 1 0 0.00
1977 27 0 27 1 27 0.00
1978 23 4 19 28 644 167.75
1979 26 1 25 47 1222 378.60
1980 12 2 10 72 864 393.86
1981 9 2 7 82 738 388.33
1982 15 3 12 89 1335 402.50
1983 14 3 11 101 1414 416.27
1984 16 3 13 112 1792 446.44
1985 44 4 40 125 5500 615.27
1986 8 7 1 165 1320 512.28
1987 24 8 16 166 3984 509.19
1988 25 4 21 182 4550 570.49
1989 46 3 43 203 9338 743.82
1990 38 4 34 246 9348 876.58
1991 16 6 10 280 4480 862.15
1992 30 3 27 290 8700 969.40
1993 60 10 50 317 19020 1108.60
1994 40 14 26 367 14680 1098.22
1995 64 18 46 393 25152 1152.61
1996 40 15 25 439 17560 1154.98
1997 37 6 31 464 17168 1240.30
1998 61 6 55 495 30195 1420.88
1999 62 12 50 550 34100 1544.43
2000 52 7 45 600 31200 1685.04
2001 49 9 40 645 31605 1791.79
2002 97 15 82 685 66445 2025.92
2003 95 21 74 767 72865 2185.51
2004 87 33 54 841 73167 2190.19
2005 100 30 70 895 89500 2284.24
2006 69 25 44 965 66585 2318.34
2007 41 14 27 1009 41369 2348.86
2008 7 4 3 1036 7252 2341.62
Population estimate using assumed 30 day sampling period

86
Table A.7: Showing information needed for the Schnabel model
to estimate population for each area A, B and C assuming a 1 day
sampling period to get a lower overestimate. (no mortality so all
estimates are over estimates).

YEAR A B C Total
1973
1974
1975
1976
1977
1978 8.8 8.8
1979 19.441176 19.441176
1980 41.054054 41.054054
1981 47.775 47.775
1982 58.431818 58.431818
1983 70.3125 70.3125
1984 58.928571 58.928571
1985 73.597826 73.597826
1986 74.173469 74.173469
1987 83.436364 83.436364
1988 96.85 96.85
1989 127.21212 127.21212
1990 161.72993 161.72993
1991 173.51049 173.51049
1992 204.09459 204.09459
1993 220.68831 220.68831
1994 72.16 223.56129 295.72129
1995 97.626374 4 101.62637
1996 107.52137 9.5675676 117.08894
1997 110.85124 28.5 139.35124
1998 117.14173 44.857143 161.99888
1999 135.00758 69.925926 204.9335
2000 148.88387 84.835294 233.71917
2001 155.36686 106.58511 261.95197
2002 168.76796 118.52475 287.29271
2003 174.215 144.95556 319.17056
2004 191.14938 149.8169 340.96628
2005 201.51341 159.79874 361.31215
2006 212.53737 166.21687 378.75423
2007 216.50842 169.41667 385.92508

87
 Appendix 7: Log sheet used for data collection

Figure A.8: Log sheet used to collect data for the project. Data collected was similar to data
collected by the Glasgow Museum Tagging Programme, but with the addition of start and finish
time.

88
 Appendix 8: Questionnaires used to collect economic data

Figure A.11: Questionnaire aimed at anglers onboard charter vessels to collect economic data

89
Figure A.10: Questionnaire aimed at skippers of charter boat businesses to collect economic data

90
 Appendix 9: Weight charts for D. batis

Table A.8: Chart to convert Length and wingspan (inches) into Weight (lbs) for males,
accurate to 5%. Adapted from SAH, created by Bill Little.

91
Table A.9: Chart to convert Length and wingspan (inches) into Weight (lbs) for males,
accurate to 5%. Adapted from SAH, created by Bill Little.

92