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platessa)
Introduction:
Plaice (Pleuronectes platessa) is the sole member of the Order Pleuronectiformes, and is one of
the several species within the Family Pleuronectidae occurring in the waters surrounding Ireland.
Pleuronectids are flatfishes characterised by the eyes being on the right side of the body in an
asymmetrical position. Living in the demersal environment, plaice are usually immediately
distinguished by their striking colour with the eyed side being warm brown with red or orange spots.
The blind, demersal side is a clear pearly white. Fig 1. External biology of plaice
The mouth is terminal and located slightly to
the right of the eyes. A series of four to seven
bony knobs run in a curved line from the eyes
back to the lateral line which, like in most
flatfish, is well-developed on both sides of
the body. Body scales are cyloid with the
body being smooth to touch. Plaice can
grow up to 91cm in length and weigh up to
7kg in exceptional cases. Most fish however
usually reach a maximum length of 50 – 60cm.
The plaice is the most familiar flatfish in
northern Atlantic waters largely due to its
excellent edible quality and its relatively high abundance. It is a typical shelf species found in shallow
water from the shoreline (juveniles & smaller specimens) to about 120m in depth. It is most
common in sandy benthic habitats but is also found on mud and gravel. “Occasionally in certain
areas, larger plaice may move up into intertidal areas and estuaries. However, plaice are not as
tolerant of fresh water as flounders and do not typically penetrate far up estuaries” (Wheeler, 1969).
Spawning takes place off the Irish coast in March – April. The spawning period differs in different
regions and at different latitudes. There are a number of particularly important spawning grounds
where mature fish congregate and to which they migrate over some distance. One of these grounds
is in the southern North Sea (ICES Division 4c) and being of particular importance to many plaice fish
stocks in North-West Europe, including Ireland. Other important spawning grounds exist in the Irish
Sea.
Plaice typically produce 10,000 to 600,000 eggs which drift as plankton for 2-3 weeks in the
upper water column. Upon hatching, the larvae and post larvae are surface living feeding on diatoms
and zooplanktonic copepods as they progressively grow and age. After around 6 weeks
metamorphosis occurs and the left eye migrates to the right side. At this stage, the juvenile plaice
become bottom living and spend their first year of life in shallow inshore water where feed on
crustaceans and amphipods. As they grow larger they gradually move into deeper waters, changing
their diets to principally feed on molluscs and other benthic organisms including polychaetes. The
food an individual eats depends on the availability in the particular bottom sediment. Plaice feed
mainly during daylight hours and feed less intensively in the winter months compared with other
times of the year. Females grow faster and live longer than males, with females reaching sexual
maturity between 3 – 7 years and males between 2 – 6 years. “The plaice is thus a long-lived flatfish,
growing rapidly early in life after which the growth rate slows” (Wheeler, 1969). The growth rate
varies from area to area due to availability of food, temperature and the population density.
The plaice is the most important flatfish to the fisheries of Europe. The total European catch
amounted to roughly 7kt (7,000 tonnes) tonnes of landed fish in 2015. Ireland holds rights for 47.6%
of the total allowable catch within its own waters within the E.U. Roughly 295 tonnes of plaice was
caught annually in mixed demersal fisheries by Irish vessels between 2013 and 2015, 49% of which
were caught in ICES Division 7.a (Irish Sea). The landings were worth approximately €708,454 for
Irish vessels in 2015 alone. Ground fishing gear is primarily used because plaice is a demersal
species. Several fishing types are used but mostly bean and otter trawls. ICES has set the minimum
landing size for plaice at 27cm so these fishing gears have to specifically tailored to avoid catching
smaller juvenile individuals which would damage the stock. Juveniles dwell in shallower water hence
destructive trawling is not carried out at shallower depths in order to preserve the recruitment stock
and juvenile habitat.
ICES sets TACs for five stocks around Ireland including the Celtic Sea, Irish Sea, South-west of Ireland,
West of Ireland and the West of Scotland & Rockhall. Over the years, the plaice has become over
exploited with the average size of landed fish dropping as the effort to land a given unit has
increased. Huge problems exist at this moment in time in relation to the plaice stock and fishery,
especially in the Celtic Sea (ICES Division 7f-g). Here, plaice are taken as a minor by-catch in mixed
fisheries and are landed with whiting and anglerfish. High discards of other non-target species occur
leading to a total of 73% of the total catch being discarded. Before 2004, the reporting of discards
was not mandatory so the magnitude of discards from the previous decades is not known.
Fig. 4(a) Showing Irish landings for gear type & (b) proportion of
discards vs landings.
Material and Methods:
A population of plaice (Pleuronectes platessa) was supplied by Galway Sea-foods and used for
the examination of the health of the stock and to analyse the various parameters of the fisheries
biology of plaice. The precise location or ICES division from which the population was taken was not
known nor whether or not the fish were taken from the same stock. For the purpose of the
investigation it was taken that the location from which the fish were taken was unknown but in
Ireland, and the data obtained used for comparison between known stocks.
A sample of 71 plaice were taken from the sample population and the various physical
properties for each were measured and recorded in the lab, i.e. length (cm), weight (g) and meristics
(dorsal fin ray count). An accuracy level of 0.1cm was established when measuring the length and
1.0cm when measuring the weight. The bodies of each fish were then cut open to expose the gonads
which were hence used in sex determination. An accurate cut was then performed diagonally
between the bony knobs on the lead in order to expose the two otholiths within the head. Both
otholiths were carefully removed, cleaned and placed under a stereoscopic microscope in order to
determine the age in years (which was then recorded). Once all of these vital properties (for fisheries
biology) were measured and recorded, the data was processed and interpreted using MS Excel into
appropriate graphs and tables from which the results were interpreted and compared to previous
fishery studies for plaice.
Results:
Fig 5. Morphology of outer and inner surface of otholith. The outer surface exhibits both opaque and
transparent growth rings. Opaque growth rings are much wider and represent summer growth, whilst
transparent growth rings represent winter growth and are much narrower. These two type of growth
rings combined are equal and deposited by one year of growth. By observing and counting the numerous
growth rings carefully, the age in years can be accurately estimated. The older a fish becomes the more
tightly packed the growth rings appear which makes it more difficult to estimate the age accurately.
Variation in width of growth rings signifies differing environmental conditions from year to year.
2).
Fig 6(a). Length – Weight Relationship in plaice sample Fig 6(b). Length – Age Relationship in plaice sample
Fig 6(a) shows the relationship between plaice length and weight in the sample population. There is
an evident positive correlation between the two parameters. Generally, as the length increases so
does the weight (as expected – growth). The graph also suggests an isometric growth pattern where
every part of the body is growing as an equal rate. The data series correlates quite closely with the
line of best fit (R2 = 0.78) with no significant outliers present.
Fig 6(b) shows the relationship between plaice age and length in the sample population. There is also
a positive correlation between the two parameters with plaice length gradually increasing with age.
The positive relationship is less prominent then that in Fig 6(a) however with the rate of change of
length decreasing as the fish gets older. The data series also do not fit the model very well (R2 = 0.2).
Some individuals from age 4 were actually measured to be shorter (at ~30cm) then individuals from
the age 3 group which were measured to be about 7-8cm longer. This indicates that different
individuals grow in length at different rates, possibly according to diet, habitat and population
density.
Parameter N Mean Range SE 95% CI Fig 7(b) Summary Statistics for total
Total length (cm) 71 37.5 14 0.31 0.6119 length (cm), weight (g), age (yrs)
Weight (g) 71 568.1 678.6 0.00 27.397 and meristics.
Age (years) 68 4.8 6 0.18 0.364
Mersitics 71 68.3 35 0.81 1.6159
Independent pooled t-test for significant difference in dorsal fin ray count between male and
female plaice (Pleuronectes platessa)
T-test for significant difference in mean dorsal fin ray count (meristics)
Data Assumptions:
Data is random, representative, replicated.
Populations independent.
Nature of data is continuous and discrete, not linked.
Data Assumptions:
Data is random, representative, replicated.
Populations independent.
Nature of data is continuous and discrete, not linked.
Conclusion:
Based on the data, there is not a significant difference in the mean length (cm) of male and
female plaice, on average (t-stat = 0.749, df = 69, p = 0.4564)
µFp - µMp = 0.5cm => ~0.00 (as per H0)
These differences are not significant.
Independent pooled t-test for significant difference in mean age between male and female
plaice (Pleuronectes platessa)
Data Assumptions:
Data is random, representative, replicated.
Populations independent.
Nature of data is continuous and discrete, not linked.
Conclusion:
Based on the data, there is not a significant difference in the mean weight (g) of male and female
plaice, on average (t-stat = 1.4115, df = 69, p = 0.1626)
µFp - µMp = 42.2g => ~0.00 (as per H0)
These differences are not significant.
4). Sex Ratio:
ii)
Observed Expected
No. of males 22 35.5 -13.5
No. of females 49 35.5 13.5
Total no. of fish 71
Fig 9. Chi2 Chi^2 10.26
calculations df 1
Chi^2 from table (df1, 0.05) 3.84
Chi x
2 2 Oi Ei 2
Chi2 formula: c
Ei
O = observed
E = expected
Larger the difference between the two, larger the value of Chi2
Df = n-1 = 2-1 = 1, where n is the number of classes => two in this case (females vs males)
Significance = 5% (p=0.05)
Decision rule:
If Chi2 calculation > Chi2 critical, there is a significance difference in the sex ratio in the
plaice population
Decision:
Chi2 (10.24) > Chi2 critical (3.84)
=> there is a significant difference in the sex ratio in the plaice population, heavily biased
towards females.
5).
Length Distribution:
Fig 10(b). Length distribution of Irish landings of plaice in Division 7(a) [1999]
Fig 10(a) shows the length distribution of plaice in the sample population. No fish where measured or were present that
were below the minimum landing size of 27cm. A relatively small proportion of smaller fish were found in the sample
(ranging from 30-34cm). The highest length frequency was measured to be 38cm with about 25% of the population
measuring within this length class. Large proportions also existed in the 37cm and 40cm length classes. These frequencies
within these length classes conformed to the average length (37.5cm). The frequency of larger individuals became less as
the length class increased which indicated that there were fewer larger, older individuals in the sample population (similar
to younger individuals). No individuals were present which measured greater than 44cm. When examining the plot as a
whole, it was concluded that the sample population had close to a normal pdf. When comparing the constructed length
distribution to the reference length frequency distribution (Fig 10(b)), it was concluded that it mirrored it quite closely.
However, Fig 10(b) showed numerous individuals which were below the minimum landing size and the graph contained a
higher frequency of smaller individuals. This is typically of a natural population. The highest frequency occurred at 28cm in
length. There were significantly fewer larger individuals. Fig 10(b) was hypothesised to contain or represent a population
with a higher sample size which is why it exhibits a more natural distribution.
6).
Age Distribution:
Fig 11(a) shows the age distribution of the sample plaice population. There was a complete absence of younger and juvenile
individuals from age groups 0 & 1 in the sample and this translated to the frequency chart. At ages 0 and 1, plaice are still
rapidly developing progressing towards stock recruitment size and age. It is very good that none of these individuals were in
the sample because this would signify illegal catching and would affect the recruitment and stock health of the wider plaice
population. Individuals from age group 4 occurred at the highest frequency and containing 20% of the sample. There was
also a large frequency of individuals from age group 5 (15% of the sample). Plaice begin to mature, both physically and
sexually, at around the age of 4. Fig. 11(b) shows a much more even spread of different age groups across the population,
with age groups 3, 4 and 5 occurring at similar and the highest frequencies. As in Fig 11(a), virtually no individuals existed
from ages 0 and 1. The frequency occurring in the age group gradually decreased as the age increased which indicated that
older individuals were much rarer or more allusive than younger individuals. The age range in Fig. 11(b) was also much
larger, ranging from 0 – 10 years.
7).
N 0 3 9 20 14 11 8 3
Stats for length data
Fig 13(a). Growth curve of plaice population plotting mean length (± C. L.) at age for the sample
Fig. 13(b). Growth curve of Pleuronectes platessa at age according to i) mean length & ii) % maturity in the North
Sea (ICES, 2015)
Fig 12 & 13(a) represent the length – age distribution in the sample plaice population
The total age observations/recordings were 68. This was in contrast to a total of 71 fish which were actually sampled at
the start. The missing age of the 3 fish was because of obscure or unfindable or otholiths which were damaged or nearly
impossible to age.
From examining the table of age - length distribution and the growth curve it was observed that
- the majority of the population were 4 - 5 age group, with numbers of individuals gradually decreasing with increased
age
-as the age increased, the length increased. Younger individuals tended to be shorter in length
-Individuals from juveniles and age group one were completely absent from the sample population. If they were there
would have been a large proportion and all being smaller than the existing population. This would have caused the graph
to be exponential in appearance (see fig 13(b) - rapidly rising at the start (with rapid growth rate and younger
individuals) but gradually levelling off as maximum lengths are reached.
The minimum landing size for plaice in the EU is 27cm. None of the sample population measured less than this - good
practise, sustainable, avoidance of younger individuals in order to let the stock recover and not become overly exploited
(if smaller, juveniles are caught in equal proportion to adults, then the stock would diminish with no patterns of
succession occuring and juvenile fish replacing the adult population)
Fig. 13(b) i) shows the growth curve for both male and female plaice (separately) at age. At ages 0-3 there is a clear rapid
growth rate. This is because juvenile plaice develop and grow at a rapid rate in their first five years. In contrast, in Fig
13(a) there is a complete absence of juvenile individuals younger than 2 years. This absence in turn does not result in a
clear growth curve where the growth rate is rapidly increasing at the start but levelling off. Females are also shown to
grow larger and faster than males.
Fig. 13(b) ii) shows a graphical representation of the % maturity of male and female plaice reached at age. As
represented in the graph, males develop far quicker than the females, with 50% of the population reaching maturity by
age two (0% for females). Hence, males develop and mature much quicker than females. However, the females catch up
by year 6 where the majority of both males and females have reached maturity. Developing and maturing quicker may
be a potential downfall for the males, in that they do not live as long as females.
8).
Condition Factor:
Fig 14(a) Non-linear regression of length vs weight Fig. 14(b) Linear regression of length vs weight
When the ln length is plotted against the ln weight, the resulting graph shows a linear regression (relationship) - fits the
model with moderate accuracy (R2 = 0.8216) – Fig 14(b)
W = aLb
W = 0.0154L2.897
a = condition factor / k
=> condition factor = 0.0154 (approach 1)
- autumn time (less condition factor because adults have been spawning, equivalent to juveniles)
Data are suggesting very poor conditions for the plaice, in comparison to k = 0.87 – possible reasons for this could be
that they were caught right after spawning or caught in winter.
Could have been after the spawning period, bad conditions of the gonads - more information could be gained by
examining the gonad condition
On average, autumn plaice are 16% heavier then spring plaice
9).
Age 2 3 4 5 6 7 8
Number 3 9 20 14 11 8 3
ln 1.099 2.197 2.996 2.639 2.398 2.079 1.099
Fig 15. Table of relative numbers by age (including natural log values)
Fig 16. Catch curve for sample plaice population – used to determine z (instantaneous coefficient of mortality
From examining the catch curve, it was observed that the age of full recruitment was in age group number 5 or 5 years old.
This age is the year after which the plaice reach full maturity and after they begin to spawn. A decreased number of fish from
older age groups which means that there are fewer older fish in the population, possibly because of mortality stresses or
possibly because the larger fish have been removed or prefer a different habitat. Recruitment into the adult stock occurs
between the ages of 5 & 7
Using slope of a line equation from catch curve to calculate z (instantaneous coefficient of mortality)
y2 - y1
x2 - x1
y1 - number at minimum recruitment (2.05)
y2 - number at max recruitment (2.6)
x1 - age at min recruitment (7)
x2 - age at max recruitment (5)
-z = -0.275
z = 0.275 yr-1
= total / instantaneous mortality coefficient (including natural and fishing mortality)
Z=F+M
z - total mort coefficient
f - fishing mort coefficient
m - natural mort coefficient (Europe)
M= 0.12
Z = 0.275
F = 0.155
Ratio of fishing mortality:natural mortality
= F/M
= 0.155/0.12
= 1.29
1.29:1
-> from the age of 5, plaice are 1.29 times more susceptible to fishing mortality then natural mortality when recruited into
the stock
For this investigation, Galway seafood’s supplied the fish/plaice for dissection, investigation and analysing, however it was
not conclusive about where (which sea/fishery/Ices area) the fish came from. For the purpose of analysis and comparison,
the plaice were taken to be from Irish waters with particular focus being put on the Irish sea plaice fishery.
There was a complete absence of younger and juvenile individuals from age groups 0 & 1 when
examining the age frequency distribution. At ages 0 and 1, plaice are still rapidly developing
progressing towards stock recruitment size and age. It is very good that none of these individuals
were in the sample because this would signify illegal catching and would affect the recruitment and
stock health of the wider plaice population. Individuals from age group 4 occurred at the highest
frequency and containing 20% of the sample. Plaice begin to mature, both physically and sexually, at
around the age of 4. The previously constructed graph shows a much more even spread of different
age groups across the population. Virtually no individuals existed from ages 0 and 1. The frequency
occurring in the age group gradually decreased as the age increased which indicated that older
individuals were much rarer or more allusive than younger individuals.
Individuals from juveniles and age group one were completely absent from the sample
population. If they were there would have been a large proportion and all being smaller than the
existing population. This would have caused the graph to be exponential in appearance, rapidly
rising at the start (with rapid growth rate and younger individuals) but gradually levelling off as
maximum lengths are reached. The minimum landing size for plaice in the EU is 27cm. None of the
sample population measured less than this which is good, sustainable practise. Younger individuals
are avoided in order to let the stock recover and not become overly exploited.
Fig 17. Landings and discards in the North Sea (ICES, 2016)
Upon graphing the linear regression between length and weight, the growth pattern of the
plaice was concluded to be allometric with different parts of the body growing at different rates.
This contrasted to the conclusions when plotting length against weight (in Results, section 2). The
condition factor for the plaice population was calculated to be 0.0154 using approach 1 calculations,
which compares very closely to the condition factor of juvenile plaice. This figure compared very
poorly with the constant or reference condition factor of 0.87. This much lower figure may be to do
with the fact that the plaice may have been caught in the autumn/winter season. Plaice feed much
less in winter which would possibly explain this. Another possible reason may have been that the
plaice were caught not long after spawning. Closer examination of the gonads when carrying out the
dissection may have shed light on this further. On average, autumn plaice are 16% heavier then
spring plaice.
From examining the catch curve, it was observed that the age of full recruitment was at 5 years
old. This age is the year after which the plaice reach full maturity and after they begin to spawn. A
decreased number of fish from older age groups which means that there are fewer older fish in the
population, possibly because of mortality stresses or possibly because the larger fish have been
removed or prefer a different habitat. Recruitment into the adult stock occurs between the ages of 5
& 7. From analysing the catch curve, the instantaneous coefficient of mortality was calculated to be
0.275yr-1. From this figure the fishing mortality was calculated to be 0.155yr-1. From the age of 5,
plaice are 1.29 times more susceptible to fishing mortality then natural mortality when recruited
into the stock. The mortality rate was calculated to be 24.04% and the survival rate calculated to be
75%.
The Marine Institute Stock Book 2015 presents some invaluable insights into the plaice fisheries
in the 5 ICES Divisions surround Ireland.
ICES advises that when the precautionary approach is applied, catches in 2017 should be no more
than 1,493 tonnes in the Irish Sea. Discard rates have been continuing to increase in recent years,
vastly outweighing the landings. Ireland has rights to 768 tonnes of the 1,098 tonnes of allowable
catch in the Irish Sea. The Irish plaice fishery is in decline, however sustainable fishing practises,
continued monitoring by ICES & M.I and the possibility of new, innovative and revolutionary fisheries
thinking could save the fishery and ensure that the popular plaice is around for years to come.
Bibliography:
ICES (2017) ‘Plaice (Pleuronectes platessa) in Division 7a (Irish Sea)’, ICES Advice of fishing
opportunities, catch & effort. Available at:
www.ices.dk/sites/pub/Publications%20Reports/Advice/2017/2017/ple.27.7a.pdf [Accessed 28th
Dec. 2017].
Marine Institute (2015) ‘The Stock Book 2015’, Marine Institute Stock Book. 2015 Edition.
[Downloaded 23rd Nov. 2017]
Wheeler, A. (1969) The Fishes of the British Isles and N.W. Europe. Michigan State University Press.
U.S.A.
Murphy, S. (2017) Plaice: Biology and the Fishery. [Lecture to BSc Applied Freshwater & Marine
Biology Stage 3] RMA49685: Resource Management & Assessment. G.M.I.T. 5th Dec.
Dutz, J., Støttrup, J.G., Stenberg, C. & Munk, P. (2016) Recent trends in the abundance of plaice
(Pleuronectes platessa) and cod (Gadus morhua) in shallow coastal waters of the North-eastern
Atlantic continental shelf – a review. vol. 1, pp 1.