!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( !

The importance of size-fecundity relationships in the management of the European lobster, Homarus gammarus.

Daryl Gunning (Queens University Belfast)

Supervisor: Prof. Paulo Prodhl Co-supervisor: Dr. Dai Roberts

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A project supported by the European Union's INTERREG IVA Programme managed by the Special EU Programmes Body!

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( !

The!importance!of!size1fecundity! relationships!in!the!management!of! the!European!lobster,!Homarus( gammarus( (

MSc!Sustainable!Aquaculture!and!Inshore! Fisheries!
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Daryl&Gunning& Supervisor:&Prof.&Paulo&Prodhl& Co:Supervisor:&Dr.&Dai&Roberts& October&2012& & & &

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

Non-technical Summary
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THE IMPORTANCE OF SIZE-FECUNDITY RELATIONSHIPS IN THE MANAGEMENT OF THE EUROPEAN LOBSTER, HOMARUS GAMMARUS

The lobster fisheries will be managed in a more sustainable way by protecting the most reproductively active and fecund individuals

BACKGROUND
The European lobster (Homarus gammarus) is distributed throughout the coastal areas of the northeast Atlantic from mid-Norway to the North Sea, south to the North African coast and into the eastern Mediterranean. Over the past 70 years, the total annual European landings of H. gammarus have varied from 1,600 to 4,800 tonnes per annum. During the early 1960s it was not uncommon for landings to regularly reach 3,000 to 3,500 tonnes per annum. Historically, the European lobster fishery has been overfished and completely unregulated or only mildly regulated by a national minimum landing size (MLS). As management plans, such as restrictions on berried females and Vnotching, now focus on improving recruitment by increasing egg production, an understanding of lobster fecundity is essential if they are to be successful. MAIN FINDINGS Egg number, egg size, ovary mass, and the capacity to hold eggs increased with female size. As larger females produce more eggs and only the largest and most dominant males mate with females, the inclusion of large males in V-notching programmes and the introduction of a maximum landing size for both sexes could provide benefits to reproductive success by protecting the most reproductively active and fecund individuals within a fishery. A large proportion of the catch is below the minimum landing size.! As fecundity appears to vary geographically, it is important that such data is available for specific managed stocks. This study focused on Northern Irish stocks in the Irish Sea and showed egg numbers increased significantly with female size.

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

ABSTRACT
Historically, the European lobster fishery has been overfished and completely unregulated or only mildly regulated by a national minimum landing size (MLS). As management plans, such as restrictions on berried females and V-notching, now focus on improving recruitment by increasing egg production, an understanding of lobster fecundity is essential if they are to be successful. Egg production is dependent on a number of biological parameters such as; size at maturity, frequency of spawning, number of lobsters per size class, and fecundity. As important population parameters vary with latitude, it is important to quantify them for specific managed stocks. Fecundity at length of H. gammarus stocks managed under a V-notch scheme off the northeast coast of Northern Ireland was estimated from commercial catch samples. It was found that egg number increased significantly with carapace length (CL) (mm) and individual fecundity ranged from 4,080 to 15,522 eggs (F=176.76*CL 7313.6). These findings correlate with previous similar studies carried out in North-east Ireland, the Republic of Ireland and Norway. Size-fecundity investigations indicated that a number of morphological parameters may be responsible for the higher reproductive capacity of larger females. Egg size, ovary mass, and the capacity of pleopods to attach egg masses increased with female size (P = >0.05).! Protecting egg-bearing females by marking them with a V-notch in the tail capitalises on these characteristics and allows them to contribute more to future generations, ensuring a more sustainable fishery. As larger females produce more eggs and only the largest and most dominant males mate with females, the inclusion of large males in V-notching programmes and the introduction of a maximum landing size for both sexes could provide benefits to reproductive success by protecting the most reproductively active and fecund individuals within a fishery.

Keywords: Female size; fecundity; V-notch; egg mass; pleopod

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

ACKNOWLEDGEMENTS
First of all, I would like to thank both my supervisors for their continued support and assistance throughout the difficult planning stage of this project. To Colin Nelson, for helping source berried females for sampling and assisting in the organisation of commercial surveys. Special thanks must be given to Francis Young and Paul Donnan, who kindly allowed me aboard their fishing vessels for the commercial surveys. Finally, I would like to thank my family and my partner for their continued support throughout my academic endeavours.

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

TABLE OF CONTENTS 1! INTRODUCTION&.......................................................................................................&1! 2! METHODS AND MATERIALS&.................................................................................&5! 2.1! Collection&of&Berried&Females&............................................................................................&5! 2.2! Morphometric&measurements&...........................................................................................&5! 2.3! Pleopod&and&egg&mass&preparation&....................................................................................&6! 2.4! Potential&fecundity&............................................................................................................&6! 2.5! Reproductive&capacity&.......................................................................................................&7! 2.6! Commercial&catch&surveys&.................................................................................................&8! 3! RESULTS&..................................................................................................................&8! 3.1! Potential&Fecundity&...........................................................................................................&8! 3.1.1! Size&Fecundity!Relationships!.............................................................................................!8! 3.1.2! Egg!mass!distribution!.......................................................................................................!11! 3.2! Reproductive&capacity&.....................................................................................................&15! 3.3! Commercial&catch&surveys&...............................................................................................&17! 4! DISCUSSION&..........................................................................................................&18! 5! CONCLUSION& .........................................................................................................&23! 6! REFERENCES&........................................................................................................&25!
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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

LIST OF TABLES AND FIGURES

Table 1: Size range and location of lobsters collected.5 Table 2: Models fitted to the relationship between fecundity and CL, and fecundity and AW..9 Figure 1: a) Tail fan being V-notched; b) V-notched tail fans..2

Figure 2: Relationship between total egg number and (a) carapace length (mm) and (b) abdomen width (mm) for all samples of females (n=10). The solid line represents a linear fit model, and the dashed line represents a power fit model..10 Figure 3: Relationship between average egg size (mm) and carapace length (mm) for all samples of females (n=10)...11 Figure 4: The distribution of eggs across the 5 pair of pleopods, averaged for all samples of females (n=10). The dark lines in the boxes represent the median; the top of the box above the median shows the 75th percentile; the bottom of the box shows the 25th percentile; the whole box represents the middle 50% of the data; the whiskers show the maximum and minimum values12 Figure 5: The proportion of eggs across the 5 pairs of pleopods for the small size class of female lobster (<87mm CL, n=4)..13 Figure 6: The proportion of eggs across the 5 pairs of pleopods for the medium size class of female lobster (88-100mm CL, n=3).14 Figure 7: The proportion of eggs across the 5 pairs of pleopods for the large size class of female lobster (101mm CL, n=3)14 Figure 8: Relationship between pleopod capacity (g) and carapace length (mm) for all samples of female lobster (n=10).15 Figure 9: The average mass (g) of each pleopod pair for all samples of female lobsters (n=10). The dark line in the boxes represents the median; the top of the box above the median shows the 75th percentile; the bottom of the box shows the 25th percentile; the whole box represents the middle 50% of the data; the whiskers show the maximum and minimum values.16 Figure 10: Relationship between ovary mass (g), ovary colour and carapace length (mm) for all samples of female lobster (n=10)...16 Figure 11: Summary of female catch not landed during the a) Portavogie (9/08/12) and b) Newcastle harbour (5/09/12) commercial survey. *These females had not previously been V-notched but were V-notched on-board...17 Figure 12: Size-class frequency of lobsters hauled during the a) Portavogie (9/08/12) and b) Newcastle harbour (5/09/12) commercial survey..18

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

INTRODUCTION
The European lobster (Homarus gammarus Linnaeus, 1758) is distributed throughout the

coastal areas of northeast Atlantic from mid-Norway to the North Sea, south to the North African coast and into the eastern Mediterranean (Tully, et al. 2006). Over the past 70 years, the total annual European landings of H. gammarus have varied from 1,600 to 4,800 tonnes per annum. During the early 1960s it was not uncommon for landings to regularly reach 3,000 to 3,500 tonnes per annum. However, catches began to drop during the mid to late 1960s and by the early 1980s, landings were below 2,000 tonnes! (Prodhl, et al. 2006). Prior to the 1950s, Norway was the main supplier of H. gammarus to the European market, accounting for 24% of landings. These landings, however, decreased dramatically from 1960-1980 at less than 10% of pre-1960 levels (Agnalt, et al. 2007). Currently, the main commercial range of H. gammarus is limited to Northern France, Britain and Ireland, with stocks in Norway and the Shetlands significantly declined or collapsed (Tully, et al. 2006). H. gammarus is of particular economic importance to Britain and Ireland with combined landings during the 2000s at an estimated 3,500 tonnes per annum and an approximate value of 35 million (FAO, 2003 and Seafish, 2011). A number of discreet areas around Northern Ireland are actively fished and in the late 2000s, total landings were recorded at about 100 tonnes per annum, with an estimated value of 500,000 to 600,000 (Marine Management, 2010). Historically, the European lobster fishery has been overfished and completely unregulated or only mildly regulated by a national minimum landing size (MLS) (Gibson, 1967). Currently, European lobster fisheries are regulated by an EU MLS of 87mm carapace length (CL) and are supported in some countries by national or regional prohibitions on landing berried (egg-bearing) females (e.g. Cumbria and Devon in the UK), closed seasons (e.g. Norway) and/or licencing (e.g. UK, France, Ireland) (Tully, 2004 and Tully et al. 2006). In addition to these management strategies, V-notching schemes have been implemented in many countries as a method to conserve stock.

V-notching is a conservation measure, which involves marking the uropods of oviparous females by removing a small triangular piece of exoskeleton and somatic tissue (Figure 1). V-notched females are then returned to the water (Deangelis, et al. 2010). The main aim of this scheme is to remove breeding females from the fishery, thereby increasing the potential fecundity of the population. It has been reported that, for small scale fisheries at least, substantial increases in the reproductive potential of the population can be achieved through V-notching schemes (Tully, 2001). A study of lobster movements conducted by Smith, et al. (2001) showed that lobsters (50-80mm CL) do not undertake extensive along-shore or offshore movements, with most recaptures being made within 3.8km of the release position. This study did not assess the dispersal of larvae or larger adults, however, it is believed that

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( movements would probably be restricted to a regional scale of <100km (Tully, 2004). Consequently, the outcomes of V-notching schemes will most likely impact fisheries regionally.

Figure 1 a) Tail fan being V-notched;

b) V-notched tail fans

The practice of V-notching was first implemented in Maine, USA, in 1917 to protect breeding females of the American lobster (Homarus americanus), however, it wasnt until the 1970s that fisherman began to voluntary V-notch on a large scale (Acheson and Gardner, 2011). V-notching has been in operation both on a voluntary basis, or supported through several grant aid schemes, in Northern Ireland for the past 10 years. Currently, the scheme is run by both the North Coast Lobster Fishermans Association (NCLFA) and the North East Lobster Cooperative (NELCO) through financial aid coming from the European Fisheries Fund. Under the NELCO scheme, the V-notch and approximately 20-30 eggs are collected, preserved, and sent to Queens University Belfast (QUB) for genetic analysis. A database which records the progeny of previously notched females is being created to provide a measure of the effectiveness of the Vnotching scheme (AFBI, 2012). This scheme is being run in conjunction with the GEL project (Genetics of the European Lobster), whose objectives include: determining the paternity of egg masses from individual females and thus elucidating the breeding structure in various lobster populations, and developing and testing microsatellite and mitochondrial DNA markers in order to optimize screening conditions, enabling high-resolution studies of European lobster genetics (Ferguson, et al. 2002). Although the V-notching scheme is voluntary, once notched, the female lobsters are protected by national UK legislation (statutory instruments No. 874, lobster and crawfish order 2000), which prohibits the landing of lobsters with v-notched or mutilated tails (Seafish, 2011). As returning V-notched lobsters to the water provides a lower profit in most cases than landing unmarked lobsters, fishermen who participate must recognise the long-term benefits of the scheme in ensuring sustainability of stocks. By participating, they are also consciously contributing to the 2! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( conservation of stocks (Acheson and Gardner, 2011). On the other hand, breeding males are not warranted the same protection and may suffer a higher level of exploitation. Therefore, the Vnotching scheme has the potential to skew the operational sex ratio (OSR) towards females (Atema and Voight, 1995, Ferguson, et al. 2002, Gosselin et al. 2003, and Leslie et al. 2006). As management plans such as MLS, restrictions on berried females and V-notching focus on improving recruitment by increasing egg production, an understanding of the complex social and reproductive behaviour (e.g. fecundity) of lobsters is essential if they are to be successful (Gosselin, et al. 2003). The time between larval settlement and recruitment to the fishery is 4-8 years, therefore, such management plans will take 4-8 years to have an impact on the fishery stock (Tully, et al. 2006). However, it is important to note that the expected return will be affected by local environmental conditions, and consequently, will not be the same for all areas (Tully, 2004). Although research on the size-fecundity relationships of H. gammarus and the biologically similar H. americanus has demonstrated that potential fecundity increases significantly with the size of the female (Ennis, 1981, Estrella and Cadrin, 1995, Tully, et al. 2001 and Lizarragna-Cubedo, et al. 2003), it must be noted that this important population biological parameter can vary with latitude. Therefore, it is important to accurately determine the geographical scales over which these biological characteristics vary in order to apply appropriate and effective stock management measures to individual regions (Tully, et al. 2001). Additionally, research has indicated that egg size increases with an increase in maternal size for several lobster species. Ouellet and Plante, (2004), found that larger females of H. americanus tended to produce larger eggs than smaller females (<79mm CL). These larger eggs tend to have a larger reserve of lipids, subsequently, the hatched larvae tend to be larger and have a better chance of survival than those that have hatched from smaller eggs (Ouellet and Plante, 2004). This improvement in survival characteristics with increasing egg size was also demonstrated for the European lobster in a study conducted by Moland, et al. (2010). Nevertheless, the relationship between egg size and female size is positive but weak in H. gammarus (Tully, et al. 2001, Lizarraga-Cubedo, et al. 2003, Agnalt, 2008, and Moland, et al. 2010). It has been suggested by Ouellet and Plante, (2004) that egg quality may be influenced by the reproductive experience of the female rather than by her size. Egg production is a function of size at onset of maturity (SOM), frequency of spawning, the size structure of the stock and the fecundity at length of lobsters (Tully, et al. 2001 and Green, et al. 2009). Therefore, an understanding of such life-history parameters is of paramount importance in the creation of biological reference points that can be used to effectively manage exploited stocks. Unlike most crustacean species, which mate when the female is hard shelled, the biologically similar H. americanus usually mate shortly after the female has moulted. However, when necessary, mating 3! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( will take place during the intermolt period. This is referred to as a dual mating strategy (Waddy and Aiken, 1991). Intermolt mating occurs in smaller lobsters that for some reason did not mate at molt. Large lobsters (120mm CL) frequently spawn twice without molting and often fail to store sufficient sperm to fertilise consecutive spawnings. Consequently, intermolt mating will ensure these females produce fertile second broods (Waddy and Aiken, 1990). The female of H. americanus usually begins to show an interest in mating when she is ready to moult and this becomes apparent by her repeated approaches to a male den over several days. The male and female then cohabit for approximately 12 days (less if the female is at the intermoult stage) (Atema and Voight, 1995). Mating is initiated by the male within moments of the females moult, when she is at peak receptivity (Talbot and Helluy, 1995). The two lobsters continue to cohabit for several days post-mating. It is believed that the male cohabits with the female to protect her from predation until she hardens and to limit the opportunity for other males to inseminate her, thus, ensuring parentage of the brood (Waddy and Aiken, 1991). Eggs are held on the pleopods (under the abdomen) following extrusion, where they are incubated for nine months until they are ready to hatch. In the majority of cases, reproduction takes place in the summer and the eggs hatch in April or May (Atema, 1986). Females tend to spawn biannually; however, large American lobsters (120mm CL) have been shown to display successive year spawning (i.e. two consecutive egg extrusions without an intervening moult), which suggests they have the capacity for sperm storage. This was found to alternate with a cycle of typical biennial spawning, thus, spawning would occur within 3 out of every 5 years (Atema, 1986 and Waddy and Aiken, 1986). Due to the biological similarity of the European lobster, this pattern may also be possible in this species. Moulting will not take place while the female is carrying eggs as they may be lost. Therefore, the growth and reproductive cycles are antagonistic and the completion of both in one year is dependent upon timing and having a sufficient energy budget (Tully, et al. 2006). The age at which the European lobster reaches maturity is dependent on water temperature and usually occurs before 5-8 years (Prodhl, et al. 2006). Genetic studies have indicated that females tend to mate with a single male in the wild (Ferguson, et al. 2002). However, tank experiments have demonstrated that individual males can fertilise several females in the one season. Therefore, further studies of the breeding system of H. gammarus in the wild are required before a definitive conclusion about polygamous behaviour is made (Ferguson, et al. 2002, and Prodhl, et al. 2006). Presently, there is a lack of published literature available on the population biology of the European lobster in the waters of Northern Ireland. The main aim of this study was to determine key population characteristics of European lobsters that have been have been protected by V-notching off the northeast coast of Northern Ireland. It is hoped that knowledge gained from this work can be used to better inform future stock management. The objectives were to: 1) examine the fecundity of H. 4! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( gammarus in North-east Ireland and; 2) determine whether the fecundity estimate obtained from this study differ from estimates found within the literature.! This study aimed to test the hypothesis that: the number of eggs per brood increases with female size (carapace length and abdomen width {mm}).

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2.1

METHODS AND MATERIALS

Collection of Berried Females Berried Homarus gammarus females were purchased from local lobster fishermen at the

beginning of the spawning cycle (August-September). Data for size-fecundity analysis was collected from lobsters that were purchased 0-2 days after they were captured to minimize egg loss due to handling or storage. Once collected, lobsters were transported live in individual plastic bags to prevent egg loss or transfer. A total of ten berried females of varying sizes were collected from Portavogie (54 24.777 N, 5 23.875 W) and Newcastle harbours (54 12 N, 54 8 W) (Table 1). The sample size was limited to ten due to time and financial constraints as well as a desire to minimize the impact on the stock surveyed. Table 1: Size range and location of lobsters collected Carapace Length (mm) 76.6 77 82.8 83.4 93.9 94.6 98.6 103.3 104.9 115.6 Location 54 12 N, 54 8 W 54 12 N, 54 8 W 54 12 N, 54 8 W 54 12 N, 54 8 W 54 24.777 N, 5 23.875 W 54 24.777 N, 5 23.875 W 54 24.777 N, 5 23.875 W 54 24.777 N, 5 23.875 W 54 24.777 N, 5 23.875 W 54 24.777 N, 5 23.875 W

2.2

Morphometric measurements Each lobster was weighed to the nearest 1g using a METTLER analytical balance. A vernier

calliper was used to record the following measurements to the nearest 0.1mm; Carapace Length (CL) measured from the posterior edge of either eye-socket to the dorsal midpoint of the posterior edge of 5! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( the carapace. Abdomen width (AW) measured at the widest point across the pleura of the second abdominal segment.

2.3

Pleopod and egg mass preparation Females were sacrificed for analysis by placing them in a freezer at -20C overnight.

Individuals were thawed fully prior to sampling eggs. Each lobster was laid flat on her back and scissors were used to cut the pleopods from the abdominal flap at the basal segment, with the egg mass still attached. Each corresponding pleopod pair and attached egg mass was stored in a separately labelled specimen-jar containing 100% ethanol to ensure independent sampling.

2.4

Potential fecundity Each pleopod pair and attached egg mass was analysed independently as follows: Each

pleopod pair and attached egg mass was removed from the ethanol and laid on a paper-towel for approximately 10 minutes to absorb excess moisture. The wet weight of the pleopod pair and attached eggs (total wet weight) was recorded to the nearest 0.001g on a METTLER analytical balance. The egg mass was then separated from the pleopod pair. When the eggs could not be easily removed from the pleopods, they were immersed in a solution of 2% sodium hypochlorite for a maximum of 5 minutes (Tully et al. 2001). Following the removal of the eggs, the pleopod pair was weighed to the nearest 0.001g. The pleopod pair weight was then subtracted from the total wet weight in order to obtain the weight of the total egg mass minus the pleopods (TEW). Three separate subsamples, each representing 1/15 of the pleopod pair egg mass were taken and laid on a paper-towel prior to being weighed to the nearest 0.001g. Each subsample was weighed to the exact same mass (weight of 1/15 of the total egg mass [EM1/15]). The number of eggs in each subsample was counted manually and averaged (average egg number in 1/15 of the egg mass [EN1/15]). EN1/15 was divided into EM1/15 to estimate the average individual egg weight in the whole sample (IEW). IEW was then divided into TEW to obtain an estimate for the total number of eggs per pleopod pair. This process was repeated across the five pairs of pleopods to give separate fecundity estimates for each pleopod pair. The sum of these five values (and the number of loose eggs remaining in the bag) provided an estimate of the potential fecundity for an individual female (total number of eggs).

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( It must be noted that the eggs were removed from the pleopods with setae still attached. This was done in order to minimize egg damage, however, their effects on wet weight and subsequent fecundity estimates is likely to be negligible. Due to the unknown egg loss during capture and handling (especially in the case of the Portavogie lobsters, which were stored in a keep-cage for 2 days prior to collection), these values should be considered as a minimum value. Linear regression analysis was applied to the untransformed data of both CL and AW (mm) and total number of eggs and to CL and ovary mass (g). A power-fit model was also applied to the regression of CL and AW and total number of eggs. Correlation analysis was performed on all these relationships (Note: all statistical analysis was performed using the R 2.14.1 statistical software). The distribution of eggs across the five pairs of pleopods was investigated for all lobsters. A one-way analysis of variance (ANOVA) was performed to test for an overall difference in the average number of eggs held on the different pleopods of all individuals. Pair-wise comparison tests (i.e. Tukey HSD) were then performed to directly compare pleopod pairs (e.g. the difference between pairs 1 and 2 or 1 and 3 etc.). These tests were also carried out on each size class of lobsters (i.e. small {<87mm CL}, medium {88-100mm CL} and large {101mm CL}). The diameter of the eggs was measured using a dissecting microscope (x10) that had a graticule with an accuracy of 0.1mm. Ten randomly selected eggs per pleopod pair (i.e. a total of 50 eggs per lobster) were measured along their longest and widest axes and an average was obtained. An average egg size was obtained for individual lobsters by taking an average of the 50 eggs measured. As the samples were not close to hatching, most eggs were spherical in shape, with less than 10% of eggs measured being ovoid in shape. A note was made of those eggs that had developing embryos (i.e. eyed). Linear regression was applied to the untransformed data of CL (mm) and egg size (mm) and correlation analysis was performed on this relationship.

2.5

Reproductive capacity The reproductive capacity of each berried female was assessed under the following 3 criteria

as follows: i. Pleopod capacity Pleopod weight is considered to be a justifiable measure of pleopod capacity as it will be affected by both pleopod size and setal hair amount and size (Tallack, 2007). Corresponding pairs of pleopods were blotted on a piece of paper to remove excess ethanol and measured to the nearest 0.001g.

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( ii. Ovary mass Following the removal of the egg mass (section 2.3), the lobster was dissected and the ovary was located and removed. It was then weighed to the nearest 0.001g. iii. Vitellogenesis stage Each ovary was classified as either cream or pale green (primary vitellogenesis) or dark green (secondary vitellogenesis). Linear regression was applied to the untransformed data of CL (mm) and both pleopod capacity (g) and ovary mass (g) and correlation analysis was performed on both these relationships. A one-way analysis of variance (ANOVA) was performed to test for an overall difference in the average mass of each pleopod pair for all individuals. Pair-wise comparison tests (i.e. Tukey HSD) were then performed to make a direct comparison between pleopod pairs.

2.6

Commercial catch surveys Field surveys of commercial lobster fishing operations took place on-board single-manned

fishing vessels at Portavogie (54 24.777 N, 5 23.875 W) and Newcastle (54 12 N 5, 48W) harbours on the 9th of August and 5th of September 2012 respectively. Each lobster hauled on-board was sexed, sized and recorded as: berried; previously V-notched prior to being hauled on-board; Vnotched on-board and/or returned to sea or kept for sale. Due to time constraints and to avoid interference with fishing operations, the size of each lobsters was not measured with a vernier calliper, but classed as either small (<87mm CL), medium (87-100mm CL) or large (101mm CL) using a square-grid (laminated A4 page with boxes 1mm in diameter). This information was collected per individual string and pot. On the Portavogie survey, a total of 14 strings with an average of 10 pots per string were sampled. Unfortunately, due to equipment malfunction aboard the vessel, only 3 strings out of 10 were sampled on the Newcastle survey. There was an average of 14 pots per string.

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3.1 3.1.1

RESULTS
Potential Fecundity Size-Fecundity Relationships The relationship of size and fecundity in Homarus gammarus was investigated in a total of

ten berried females. The total number eggs per individual ranged from 4,080 for the smallest female (76.6mm CL) to 15,522 for the third largest (103.3mm CL). The number of eggs per individual ranged from 4,080-8,422 for females below the minimum landing size of 87mm; 7,163-10,816 for medium-sized females (88-100mm CL) and 10,788-15,522 for large females (101mm CL). A positive trend (r = 0.735) existed between CL (mm) and the estimated number of eggs per brood 8! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( (fecundity) (Figure 2a). This trend was found to be significant (r2 = 0.540, d.f. = 8, P < 0.05). As the eggs are stored ventrally on the abdomen, the relationship of AW and fecundity was also determined (Figure 2b). This relationship also showed a positive trend (r = 0.757) that was significant (r2=0.573, d.f. = 8, P < 0.05). A power-fitted model of data was provided in addition to the linear model. The power function produced exponent values less than 3.0 for the relationship between fecundity and CL (2.0) and fecundity and AW (1.5). This implies the presence of a simple volumetric relationship and suggests that the number of eggs produced is fully explained by body size (Table 2). Table 2: Models fitted to the relationship between fecundity and CL, and fecundity and AW Model a b r2 Fecundity vs. CL Linear fit (y=aCL+b) Power fit (y=aCLb) Fecundity vs. AW Linear fit (y=aAW+b) Power fit (y=aAW )
b

176.76 0.992

-7313.6 2.005

0.540 0.596

226.95 19.92

-3278.9 1.524

0.573 0.657

CL = Carapace length, AW = Abdomen width

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

18,000! 16,000! 14,000! Total&no.&of&eggs&per&brood& 12,000! 10,000! 8,000! 6,000! 4,000! 2,000! 0!

a)!

70!

75!

80!

85!

90!

95!

100!

105!

110!

115!

120!

Carapace&Length&(mm)&

18,000! 16,000! 14,000! Total&No.&of&eggs&per&brood& 12,000! 10,000! 8,000! 6,000! 4,000! 2,000! 0! 35! 40! 45! 50! 55! 60! 65! 70! 75! Abdomen&Width&(mm)&

Figure 2: Relationship between total egg number and (a) carapace length (mm) and (b) abdomen width (mm) for all samples of females (n=10). The solid line represents a linear fit model, and the dashed line represents a power fit model

10! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( Linear regression was also applied to average egg size (mm) and CL (Figure 3). The relationship was found to be positive (r=0.665) and also significant (r2=0.437, d.f.=8, P < 0.05). This relationship was in contrast to previous results obtained in the Republic of Ireland and Scotland, which found that the relationship was not statistically significant (Tully et al. 2001, Lizarraga-Cubedo et al. 2003, and McBride, 2011). The egg diameter ranged from 1.7 2.4mm, with a mean of 2.05mm. No eggs were found to be eyed.

2.5! 2.4! 2.3! Av.&Egg&Size&(mm)& 2.2! 2.1! 2.0! 1.9! 1.8! 1.7! 1.6! 70! 75! 80! 85! 90! 95! 100! 105! 110! 115! 120! Carapace&Length&(mm)&

Figure 3: Relationship between average egg size (mm) and carapace length (mm) for all samples of females (n=10)

3.1.2 !

Egg mass distribution

The distribution of eggs across the five pairs of pleopods was investigated. Pleopod pair 1 refers to the pleopods closest to the thorax and pair 5 refers to those closest to the tail fans. The distribution of eggs across the pleopod pairs are not even, with most being found on the third pleopod pair, located towards the middle of the abdomen. The least amount of eggs was found on the 1st pleopod pair (Figure 4). The average number of eggs differed significantly across the five pairs of pleopods (F4,45=15.99, P < 0.001). It was found that there was a significant difference between pleopod pair 1

11! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( and 2 (P < 0.05), 1 and 3 (P < 0.001), 1 and 4 (P < 0.001), 2 and 3 (P < 0.01), 3 and 5 (P < 0.001), and 4 and 5 (P < 0.01). Differences between all other pairs of pleopods were not significant.!

Av. number of eggs

1000

2000

3000

4000

Pair.1

Pair.2

Pair.3 Pleopod

Pair.4

Pair.5

Figure 4: The distribution of eggs across the 5 pair of pleopods, averaged for all samples of females (n=10). The dark lines in the boxes represent the median; the top of the box above the median shows the 75th percentile; the bottom of the box shows the 25th percentile; the whole box represents the middle 50% of the data; the whiskers show the maximum and minimum values

The distribution of egg mass across the pleopod pairs was also considered for these lobsters under 3 different size classes; small (<87mm CL, n=4), medium (88-100mm CL, n=3) and large (101mm CL, n=3). The average egg number per pleopod pair was converted to a percentage to allow for a direct comparison amongst the three size classes. For all size class, the distribution of eggs across the pleopod pairs was not even, with most being found on the third pleopod pair (Figure 5-7).

12! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( There was a significant difference in the number of eggs per pleopod pair for the small size class (F4,15=5.895, P < 0.01). The significant difference occurred between pleopod pair 1 and 3 (P < 0.01) and between 3 and 5 (P < 0.05), all other differences were not significant. The average number of eggs differed significantly across the pleopod pairs of the medium size class (F4,15=4.74, P < 0.05), however, a significant difference only occurred between pleopod pair 1 and 3 (P < 0.05), with all other differences being non-significant. The average number of eggs also differed significantly across the pleopod pairs of the large size class (F4,15=13, P < 0.001). There was a significant difference between pleopod pair 1 and 2 (P < 0.05), 1 and 3 (P < 0.01), 1 and 4 (P < 0.01), 3 and 5 (P < 0.01), and 4 and 5 (P < 0.05)

45! 40! 35! ProporVon&of&eggs&(%)& 30! 25! 20! 15! 10! 5! 0! 1! 2! 3! Pleopod&Pair& 4! 5!

Figure 5: The proportion of eggs across the 5 pairs of pleopods for the small size class of female lobster (<87mm CL, n=4)

13! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

35! 30! 25! 20! 15! 10! 5! 0! 1! 2! 3! Pleopod&Pair& 4! 5!

Figure 6: The proportion of eggs across the 5 pairs of pleopods for the medium size class of female lobster (88-100mm CL, n=3)
40! 35! 30! ProporVon&of&eggs&(%)& 25! 20! 15! 10! 5! 0! 1! 2! 3! Pleopod&Pair& 4! 5!

Figure 7: The proportion of eggs across the 5 pairs of pleopods for the large size class of female lobster (101mm CL, n=3)

ProporVon&of&eggs&(%)&

14! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( 3.2 Reproductive capacity Total pleopod capacity, defined as total pleopod weight in this study, had a very strong positive linear relationship (r = 0.987) with CL (mm) that was highly significant (r2=0.9744, d.f. = 8, P < 0.001) (Figure 8). A summary of the average weight of each pleopod pair for all individuals is provided in Figure 9. The average mass differed significantly across the five pairs of pleopods (F4,45=12.59, P < 0.001). It was found that there was a significant difference between pleopod pair 1 and 2 (P < 0.001), 1 and 3 (P < 0.001), 1 and 4 (P < 0.001), and 1 and 5 (P < 0.01). Differences between all other pairs of pleopods were not significant. The relationship of ovary weight and CL was also evaluated to assess reproductive capacity. It was found that ovary weight increased with females size (CL) and followed a very strong positive linear relationship (r = 0.952) that was highly significant (r2 = 0.905, d.f. = 8, P < 0.001). The reproductive capacity of the females ovaries was further assessed by noting ovary colouration. Five of the females studied had dark green ovaries, indicating a later stage of development (i.e. secondary vitellogenesis). The remaining five females had pale green or cream ovaries, indicating an early stage of development (i.e. primary vitellogenesis) (Figure 10). Although five females appeared to be at the secondary vitellogenesis stage, no females appeared to be carrying eyed eggs.

6! 5! Pleopod&Capacity&(g)& 4! 3! 2! 1! 0! 70! 75! 80! 85! 90! 95! 100! 105! 110! 115! 120! Carapace&Length&(mm)&

Figure 8: Relationship between pleopod capacity (g) and carapace length (mm) for all samples of female lobster (n=10)

15! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

Average Mass (g)

0.0

0.5

1.0

1.5

Pair.1

Pair.2

Pair.3 Pleopod

Pair.4

Pair.5

Figure 9: The average mass (g) of each pleopod pair for all samples of female lobsters (n=10). The dark line in the boxes represents the median; the top of the box above the median shows the 75th percentile; the bottom of the box shows the 25th percentile; the whole box represents the middle 50% of the data; the whiskers show the maximum and minimum values

16! 14! 12! Ovary&Mass&(g)& 10! 8! 6! 4! 2! 0! 70! 75! 80! 85! 90! 95! 100! 105! 110! 115! 120! Carapace&Length&(mm)& Dark!Green! Pale!Green! Cream!

Figure 10: Relationship between ovary mass (g), ovary colour and carapace length (mm) for all samples of female lobster (n=10)

16! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( 3.3 Commercial catch surveys

A total of 104 lobsters were hauled during the Portavogie commercial catch survey, with an average of seven individuals per string and one individual per pot. Of this total, only 8.65% were landed (4 males and 5 females of size class M).!Of the 91.35% of the haul that was returned to the sea, 45 males and 49 females were below the minimum landing size (87mm CL) and one size-class L (101mm CL) non-berried female was V-notched on-board. A total of 65 lobsters were hauled during the Newcastle commercial catch survey, with an average of 22 individuals per string and two individuals per pot. Similar to the Portavogie survey, only 10.77% of the total haul was landed (4 males and 1 female of size class M and 2 males of size class L). Of the 89.23% of the haul that was returned to the sea, a large proportion was also under the legal landing size of 87mm CL (38 males and 15 females). Figure 11 provides a summary of the female catch that was returned to the sea. The bulk of the Portavogie catch were of size-class S (<87mm), with females showing a higher level of frequency across all sizeclasses. No males in size-class L were hauled on-board. Similiarly, the bulk of the Newcastle catch were also of size-class S (<87mm), however, males showed a higher level of frequency across all size-classes (figure 12).

100! Frequency&of&females&(%)& 80! 60! 40! 20! 0! Below!MLS,!not! berried!

a)!
Frequency&of&females&(%)&

70! 60! 50! 40! 30! 20! 10! 0!

b)!

Very!Large!

Below!MLS,! Below!MLS,! V&notched,! not!berried! berried*! not!berried! Reason&for&not&landing&

V&notched,! berried!

Reason&for&not&landing&

Figure 11: Summary of female catch not landed during the a) Portavogie (9/08/12) and b) Newcastle harbour (5/09/12) commercial survey. *These females had not previously been Vnotched but were V-notched on-board

17! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

50! 45! 40! 35! Frequency&(%)& 30! 25! 20! 15! 10! 5! 0! S! M! Size&Class*& L! Frequency&(%)&

70! 60! 50! 40! 30! 20! 10! 0! S! M! Size&Class*&

Male! Female!

*S=<87mm;!M=88&100mm;! L=101mm!CL!

L!

Figure 12: Size-class frequency of lobsters hauled during the a) Portavogie (9/08/12) and b) Newcastle harbour (5/09/12) commercial survey

DISCUSSION
The size-fecundity analysis conducted during this study revealed a significant increase in the

estimated number of eggs per brood as lobster size increased (both carapace length {CL} and abdomen width {AW}). This finding is consistent with previous studies on the biologically similar Homarus americanus (Ennis, 1981 and Estrella and Cadrin, 1995) and on H. gammarus (Tully, et al. 2001, Lizarragna-Cubedo, et al. 2003 and McBride, 2011). The study conducted by McBride (2011) also took place off the coast of northeastern Northern Ireland and the results were very similar to those found in this study. The estimated number of eggs per brood ranged from 4,080 for the smallest female at 76.6mm CL to 15,522 for a female of 103.3mm CL. The female with the most eggs was the third largest, while the largest female produced the second largest number of eggs, at 11,050. It must be noted that all lobsters in the large size-class (101mm CL) were collected from Newcastle harbour. Due to collection delays, these specimens were kept in a keep-cage for 2 days prior to collection. Therefore, some individuals may have lost eggs during their transfer to the keep-cage. Alternatively, egg loss may have occurred due to natural means prior to collection or the largest female may simply have only produced this number of eggs. For the McBride (2011) study the total number of eggs per 18! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( lobster varied from 3,575 for the smallest female of 80mm CL to 18, 685 for the largest female of 116mm CL. Both a linear and power-fit model was fitted to the relationship of fecundity with CL and AW. In previous studies of H. gammarus, a linear function was most commonly applied (Hepper and Gough, 1978, Bennet and Howard, 1987 and Free et al. 1992), however, recently the power function has been preferred (Tully, et al. 2001 and Lizarrangna-Cubedo, et al. 2003) and was the function with the strongest correlation in this study. It has also been chosen in previous studies due to its volumetric nature (Agnalt, 2008). As female body size has an influence on egg weight (Tully, et al. 2001) and egg weight was used in the fecundity estimations, the shape of the size-fecundity relationship will be partially dependent on the size range of the sample. This is not recognised by the linear model and may result in the underestimation of fecundity, particularly in large females. As the relationship between length and fecundity follows a cuboidal trajectory (i.e. a coefficient of approximately 3) (Green et al. 2009), power-fitted models were applied to the relationship of brood size versus CL and AW. This provided exponent values <3.0 in both cases, implying the presence of a simple volumetric relationship. This may be an indication that the number of eggs produced can be fully explained by body size. Previous studies on berried females from Northern Irish (Irish Sea) stocks found these exponent values for both relationships to be above 3.0. This implies the absence of a simple volumetric relationship and indicates that the number of eggs produced is only partly explained by body size (McBride, 2011) and may also be influenced by other factors such as the females reproductive experience and the quality and quantity of sperm received. Both studies had a small sample size of ten berried females over a limited study period. Further studies with a larger sample size spread over each season are required. Ovary weight also increased significantly with increasing CL (r = 0.951, P < 0.01). This relationship was also seen in McBrides (2011) study of lobsters off the Antrim coast, however, in this study the relationship was highly variable and only weakly significant. Egg size was found to increase with female size (CL), however, this trend was only weakly significant (r=0.665, P < 0.05). It has been demonstrated that larger eggs contain a higher quantity of yolk, which has been shown to correlate with a higher rate of initial survival after spawning and an improved growth rate. They also tend to produce larger larvae that have a greater chance of avoiding predation and surviving to the recruitment stage (Ouellet and Plante, 2004, and Moland et al. 2010). Despite this, previous egg size and body size relationship analysis of H. gammarus has found the relationship to be not statistically significant (Tully, et al. 2001, Lizarraga-Cubedo, et al. 2003, Agnalt, 2008, Moland, et al. 2010, and McBride, 2011).

19! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( In the wild, there is a limit to the resources a female can allocate to reproduction; therefore, a trade-off is likely between the size of an egg and the number of eggs produced. The weak correlation of egg and body size would suggest that the European lobster increases its reproductive fitness by concentrating more effort on the number of eggs produced rather than the size and quality of the eggs. During the incubation phase of both H. americanus and Nephrops norvegicus, egg loss is quite high with a loss of up to 36% and 20-60% respectively (MacDiarmid and Sainte-Maire, 2006 and McQuaid, et al. 2009). The rate of natural egg loss at the incubation stage in H. gammarus is unclear (Agnalt, 2008). However, if losses are comparable to the biologically similar H. americanus, it would be in H. gammarus best interest to focus efforts on producing a large number of eggs rather than producing eggs of a larger size or better quality. The absence of eyespots on the eggs indicates that fertilization has not yet taken place. Females lay eggs regardless of whether they have mated or not (Ennis, 1975). It would appear that large females with heavier ovaries produce more eggs. However, this reproductive effort would be wasted if efforts, such as increased abdomen width and pleopod capacity, were not made to accommodate the extra eggs (Alunno-Bruscia and Sainte-Maire, 1998). The findings from this and McBrides (2011) study show an increase in egg number as abdomen width increases. Tallack (2007) reported that the pleopod capacity of Cancer pagurus increased significantly upon reaching maturity. McBride (2011) fitted a linear model of pleopod capacity and CL and found the relationship to be strong. This same relationship was investigated in this study and it was discovered that pleopod capacity increased significantly with increasing female size (r=0.987, P < 0.001). It would appear that a combination of morphological features is responsible for the females reproductive capacity as she increases in size and age. The spawning cycle of H. gammarus is generally biennial. Spawning occurs in early autumn, and the females remain berried for a nine month period. The eggs hatch the following summer and the female then moults in late summer/autumn. It is possible that mating also occurs at this time. Following the moult, females remain unberried until spawning occurs the following year (MacDiarmid and SainteMaire, 2006). With an alternating cycle of brooding and moulting, one might expect that approximately half the population of reproductive females would be carrying eggs. For both commercial surveys, this was not the case. During the Portavogie survey, no berried females were hauled on-board and during the Newcastle survey only five of the 19 females hauled were berried. A possible explanation for this low frequency of berried females is the fact that sampling took place in August and September. A large proportion of these sexually mature females may have recently extruded their larvae or were going to spawn over the coming months.

20! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( Using data obtained from the commercial surveys, the number of eggs that can be saved on a typical fishing trip due to a ban on landing V-notched females can be estimated. Two non-berried, legalsized, V-notched females in the medium size-class (88-100mm CL) were hauled on-board during the Newcastle survey. From the size-fecundity analysis conducted, potential egg production for a female in the medium size-class ranges from 7,163-10,816. If both females produced a minimum of 7,163 eggs, an approximate total egg production of 14,326 would be obtained. Assuming the V-notch remains for four years, the females would be protected for two reproductive cycles under a biennial spawning cycle (Leslie et al. 2006). This would result in a total egg production of 28,652 between the two females. Recruitment is highly variable, but assuming a low of 0.1% (Waddy and Aiken, 1991), these two females would contribute 29 individuals to the recruitment population over a four year period. However, it is important to note that this is a rough calculation based on only a half-days fishing effort. It also ignores the fact that the females will grow and produce a larger number of eggs over the four year period. Therefore, this figure must be taken as a minimum estimation. As the V-notching scheme only offers protection to female lobsters, one can assume that mature males are exploited more intensely than females. This has the potential to skew the operational sex ratio (OSR) towards females. The commercial survey off the Portavogie coast found a higher frequency of females for all size classes. Although a higher frequency of males for all size classes was found during the Newcastle survey, it must be noted that a full survey could not be conducted due to equipment malfunction. Studies by McBride (2011), found that the population in Dundrum Bay was skewed towards females in individuals above the minimum landing size (MLS). Further investigations of OSR are required as commercial surveys can be limited by trap-bias. For several species of lobster, males tend to be more catchable than females (Trembley and Smith, 2001). This may explain the higher proportion of males encountered on the partial survey in Newcastle and for individuals below the MLS for the McBride (2011) survey. A greater catchability of males could increase the potential of fisheries to alter the OSR in areas that undertake the V-notching scheme. However, there are seasonal, size, and sex-specific changes in catchability. This makes it difficult to determine if variations in size distribution and sex ratio are due to natural cycles or fishing mortality (Morgan, 1974). Tagging studies and visual census through diver hand-collection could be used to avoid trapbias (MacDiarmid, 1991). Such an alteration to the OSR could have a number of negative impacts, such as; altering the supply and quality of sperm (MacDiarmid and Sainte-Maire, 2006), and lowering the encounter rates of potential mates (Gosselin, et al. 2003). Reproduction is a costly process and therefore it is essential that individuals find and recognise a potential mate quickly. Previous studies indicate that the biologically similar H. americanus exhibits synchronous mating and limited female receptivity. It appears that the males accept a female mate that is mature and has not yet mated. The female, 21! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( however, demonstrates a greater level of mate choice, competing for a mate that will maximize personal reproductive success (MacDiarmid and Sainte-Maire, 2006). Studies have shown that the females of H. americanus actively search out the dominant (usually the largest) male to mate with (Atema and Voight, 1995 and Leslie et al. 2006). Females may be choosing the largest, dominant males due their greater offering of sperm, greater protection value, and larger dens that can accommodate their size (Atema and Voight, 1995 and Gosselain et al. 2003). If females are mate-limited, they may be forced to mate with a smaller male of lower value. Gosselain, et al. (2003) found that females cohabit dens five times longer with males >114mm CL than those of 8184mm CL. In some cases, the den of smaller males cannot accommodate the size of large females (Cowan and Atema, 1990 and Goesselain, et al. 2003). According to Cowan and Atema (1990), it may be the case that H. americanus staggers its moult to mate with a preferred male. The female depends upon a male for protection during the moult and they may suffer injury or mortality when they cannot find a suitable mate (Cowan and Atema, 1990). It is not yet known whether this behaviour occurs in H. gammarus. Further investigation is required to better understand the population characteristics of H. gammarus and how they influence stock management. Although the majority of female H. gammarus display a biennial pattern of spawning, studies have shown moulting frequency to decrease with increasing size (Gibson, 1967 and Tully, 2001). Wady and Aiken, (1986) found that H. americanus displays successive-year spawning, a cycle consisting of two consecutive egg extrusions without an intervening moult. This cycle usually alternates with a biennial cycle, resulting in spawning in three out of every five years. Consequently, a larger proportion of large females would be carrying eggs. Females also exhibit alternate-year spawning (spawning occurs the year following moult and again two years later without an intervening moult), which means spawning would only occur in two of every five years however, sperm can remain viable in the receptacle of a female for at least two years, allowing a female to fertilize two consecutive egg broods with the sperm of a single mate (Waddy and Aiken, 1991). The inclusion of large males in V-notching programmes and the introduction of a maximum landing size for both sexes could provide benefits to reproductive success by protecting the most reproductively active and fecund individuals within a fishery. It has been shown that the larger females produce more eggs and only the largest and most dominant males mate with females (Leslie, et al. 2006). The distribution of the egg mass across the five pairs of pleopods was investigated to assess any potential influence it may have on the development of larvae and the manner in which they are released. The distribution was found to be non-uniform, with the third pleopod pair carrying most 22! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( eggs for all individuals. The fourth pleopod also carried a similarly large quantity of eggs and the difference in quantity between the 3rd and 4th pair was not significantly different. The lowest number of eggs was found on the first pair of pleopods for all females. This distribution pattern can be explained by the carrying capacity of each pleopod. The third and fourth pairs of pleopods were the largest in weight, whereas, the first pair weighed the least. Since the weight of the pleopods relates to their size and the number and length of the setae, the third and fourth pleopod pairs have a greater capacity for carrying eggs. A similar pattern of egg mass distribution was seen in a study conducted by McBride (2011). During the incubation period, the female cleans her eggs with her last pair of walking legs and aerates them by waving her tail fan, providing them with oxygen. The third and fourth pleopod pair contains a thicker mass of eggs than the other pleopod pairs and subsequently, the exchange of oxygen and wastes with the surrounding water may be reduced. The literature provides evidence of oxygen limitation within large egg masses for many crustacean and marine invertebrate species. Oxygen concentrations have been shown to decline from the surface of an egg mass to the centre and eggs contained in thicker egg masses develop slower than those in thinner egg masses. Eggs contained centrally within thick egg masses may be at the threshold oxygen concentration, beyond which, development is retarded (Chaffee and Strathmann, 1984, Strathmann and Strathmann, 1995, and Cohen and Strathmann, 1996). Larval release occurs nocturnally and in rhythmical batches over a two to six week period (Templeman, 1937). As the hatching of eggs does not appear to be under the females direct control (Davis, 1964 and Ennis, 1975), an uneven distribution of eggs on the pleopods may be an evolutionary strategy to incur different developmental times of the larvae. Pandian (1970) observed that larvae released earlier in the hatching process had softer integuments but more energy and those released later had harder integuments but less energy. By varying the development condition of released larvae, the types of predators that can be evaded increases. Also, releasing them at night, over a prolonged period, reduces the occurrence of predation events substantially. If they were released in one batch, it is quite likely that a large proportion would fall prey to predators before having the opportunity to disperse (Templeman, 1937, Davis, 1964, and Ennis, 1975).

CONCLUSION
Given that fecundity appears to vary geographically (Tully, et al. 2001), it is important that

such data is available for specific managed stocks. This study, similar to previous work carried out by McBride, (2011) focused on Northern Irish stocks in the Irish Sea. The results of both these studies 23! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( were very similar, with egg numbers increasing significantly with female size. However, it is important to note that the seasons assessed in these studies were limited (July-September). For a more accurate assessment of size-fecundity relationships in this, and any other region, future studies should take place throughout the year. The V-notching practice has the benefit of protecting breeding females, even when they are not in a berried state. The notch will remain on a female for a number of reproductive cycles, over which time she will increase in size and subsequently increase her reproductive output. However, there is a potential flaw to the V-notching scheme. Numerous studies (Cowan and Atema, 1990, Atema and Voight, 1995, Gosselain et al. 2003 and Leslie, et al. 2006) have shown that females select the most dominant and large males to mate with. Despite this, there is currently no legislation in Ireland that protects males above the minimum landing size and, as the V-notching scheme only protects females, males suffer greater levels of exploitation. If the number and size of males in a population diminish, the supply of sperm would undoubtedly be reduced. Although the Portavogie commercial survey in this study and the Dumdrum Bay survey by McBride, (2011) demonstrated a skew in the population towards females, it must be noted that both studies only involved one day of fishing and did not account for seasonality differences in catchability of either sex. Further studies that involve a more detailed census and/or tagging component, conducted throughout the year, may be required to fully understand the impacts that a V-notching scheme may have on local lobster population dynamics. There is growing agreement amongst fishery scientists that all components of a mating system are essential for the conservation of marine resources (Conover and Munch, 2002). It would appear that this understanding also exists amongst lobster fishermen. An interview conducted with the fisherman involved in the Newcastle survey revealed that some lobster fishermen who operate in waters off the coast of northeastern Northern Ireland are, on a volunteer basis, V-notching random male lobsters in order to balance the ratio of males and females (Donnan, personal communication, 2012). The fisherman involved in the Portavogie survey V-notched a previously un-notched, unberried, large female. Following an interview with this fisherman, it was clear he understood that larger females provide a larger number of eggs, potentially adding a greater number of offspring to his stock. In addition to V-notching, a maximum landing size for both sexes now exists in the US. This additional management scheme was introduced to increase the contribution of large males and females to future generations (Fogarty and Gendron, 2004 and Tully, 2004). Considering the mating preference of female H. gammarus, it may benefit the European lobster fishery to adopt a more holistic approach to fisheries management.

24! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus(

REFERENCES

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( Deangelis, B. M., Cooper, R., Clancy, M., Cooper, C., Angell, T., Olszewski, S., Colburn, W., and Catena, J. (2010). Impacts of V-notching the American lobster. Journal of Shellfish Research, 29, pp. 489-496. Ennis, G.P. (1981). Fecundity of the American lobster, Homarus gammarus, in Newfoundland waters. Fishery Bulletin, 82, pp. 796-800. Ennis, G.P. (1975). Observations on hatching and larval release in lobster Homarus americanus. Journal of the Fisheries Research Board of Canada, 32, pp. 2210-2213. Estrella, B.T., and Cadrin, S.X. (1995). Fecundity of the American lobster (Homarus gammarus) in Massachusetts coastal waters. ICES Marine Science Symposia, 199, pp. 61-72. Ferguson, A. et al. (2002). Genetic diversity in the European lobster (Homarus gammarus): population structure and impacts of stock enhancement. Accessed through: http://www.qub.ac.uk/bb-old/prodohl/GEL/gel.html on 10/07/2012. Fogarty, M.J., and Gendron, L. (2004). Biological reference points for American lobster (Homarus americanus). Canadian Technical report of Fisheries and Aquatic Sciences, 2369, pp. 1-31. Food and Agriculture Organisation. (2003). Fishery and aquaculture country profile Ireland. Accessed through: http://www.fao.org/fishery/countrysector/FI-CP_IE/en on 09/07/2012. Free, E.K., Tyler, P.A., and Addison, J.T. (1992). Lobster (Homarus gammarus) fecundity and maturity in England and Wales. ICES document, CM, 43. Gibson, A. (1967). Irish investigations of the lobster. Irish Fisheries Investigations Series B, 5, pp. 1345. Gosselin, T., Sainte-Marie, B., and Bernatchez, L. (2003). Patterns of sexual cohabitation and female ejaculate storage in the American lobster (Homarus americanus). Behaviour Ecology Sociobiology, 55, pp. 151-160. Green, B.S., Gardner, C., and Kennedy, R.B. (2009). Generalised linear modelling of fecundity at length in southern rock lobsters, Jasus edwardsii. Marine Biology, 156, pp. 1941-1947. Hepper, B.T. and Gough, C.J. (1978). Fecundity and rate of embryonic development of the lobster, Homarus gammarus (L), off the coast of North Wales. Journal du Conseil International pour IExploration de le Mer, 38, pp. 54-57. Leslie, B., Henderson, S., and Riley, D. (2006). Lobster Stock Conservation V-notching. NAFC Marine Centre, Fisheries Development Note, 22, pp. 1-4. Lizrraga-Cubedo, H.A., Tuck, I., Bailey, N., Pierce, G.J., and Kinnear, J.A.M. (2003). Comparisons of size at maturity and fecundity of two Scottish populations of the European lobster, Homarus gammarus. Fisheries Research, 65, pp. 137-152. MacDiarmid, A.B., and Sainte-Maire, B. (2006). Reproduction. In Lobster: Biology, Management, Aquaculture, and Fisheries (ed. B.F. Philips).Blackwell Publishing, Oxford, pp. 45-68. Marine Management (2010). The UK fishing industry in 2010 landings. Accessed through: http://marinemanagement.org.uk/fisheries/statistics/annual.htm on 09/07/2012. 26! ! !

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( McBride, S. (2011). The importance of V-notching for fisheries management of the European lobster, Homarus gammarus. MSc Project, School of Biological Sciences, Queens University Belfast, pp. 1-36. McQuaid, N., Briggs, R.P., and Roberts, D. (2009). Fecundity of Nephrops norvegicus from the Irish Sea. Journal of the Marine Biological Association of the United Kingdom, 89, pp. 1181-1188. Moland, E., Olsen, E.M., and Stenseth, N.C. (2010). Maternal influences on offspring size variation and viability in wild European lobster Homarus gammarus. Marine Ecology Progress Series, 400, pp. 165-173. Ouellet, P., and Plante, F. (2004). An investigation of the source of variability in American lobster eggs and larvae size: Maternal effects, and inter-annual and inter-regional comparisons. Journal of Crustacean Biology, 24, pp. 481-495. Pandian, T.J. (1970). Ecophysiological studies studies on the developing eggs and embryos of the European lobster, Homarus gammarus. Marine Biology, 5, pp. 154-167. Prodhl P.A., Jrstad K.E., Triantafyllidis A., Katsares V. and Triantaphyllidis C. (2006). European lobster Homarus gammarus. In: Genetic effects of domestication, culture and breeding of fish and shellfish, and their impacts on wild populations. (ed. D. Crosetti, S. Lapgue, I. Olesen, T. Svaasand). GENIMPACT project: Evaluation of genetic impact of aquaculture activities on native populations. A European network. WP1 workshop Genetics of domestication, breeding and enhancement of performance of fish and shellfish, Viterbo, Italy, 12-17th June, 2006, 6 pp. Seafish (2011). Responsible sourcing guide: crabs and lobsters. Version 3, pp. 1-9. Smith, I.P., Jensen, A.C., Collins, K.J., and Mattey, E.L. (2001). Movement of wild European Lobster Homarus gammarus in natural habitat. Marine Ecology Progress Series, 222, pp. 177-186. Strathmann, R.R., and Srathmann, M.F. (1995). Oxygen supply and limits on aggregation of embryos. Journal of Marine Biological Society of the United Kingdom, 75, 413-428. Talbot, P., and Helluy, S. (1995). Reproduction and embryonic development. In Biology of the lobster Homarus americanus. Academic Press, New York, pp. 177-216. Tallack, S.M.L. (2007). Size-fecundity relationships for Cancer pagurus and Necora puber in the Shetland Islands, Scotland: how is reproductive capacity facilitated? Journal of the Marine Biological Association of the United Kingdom, 87, pp. 507-515. Templeman, W. (1937). Habits and distribution of larval lobsters (Homarus americanus). Journal of the Biological Board of Canada, 3, pp. 343-347. Tully, O., Bell, M., OLeary, A., McCarthy, A., ODonovan, V., and Nee, D. (2006). The lobster (Homarus gammarus L.) fishery: Analysis of the resource in 2004/2005. Fisheries Resource Series, Bord Iascaigh Mhara (Irish Sea Fisheries Board), Dun Laoghaire, Ireland, 6, pp. 1-62.

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!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Size&fecundity!relationships!in!Homarus(gammarus( Tully, O. (2004). The Biology and Management of Clawed Lobster (Homarus gammarus L.) in Europe. Fisheries Resource Series, Bord Iascaigh Mhara (Irish Sea Fisheries Board), Dun Laoghaire, Ireland, 2, pp. 1-31. Tully, O. (2001). Impact of v-notch technical conservation measure on reproductive potential in a lobster (Homarus gammarus L.) fishing in Ireland. Marine Freshwater Research, 52, pp. 1551-1557. Tully, O., Roantree, V., and Robinson, M. (2001). Maturity, fecundity and reproductive potential of the European lobster (Homarus gammarus) in Ireland. Journal of the marine Biological Association of the United Kingdom, 81, pp. 61-68. Waddy, S.L., and Aiken, D.E. (1991). Mating and insemination in the American lobster Homarus americanus. In Crustacean Sexual Biology (ed. R.T. Bauer and J.W. Marin). Columbia University Press,!New York and Oxford, pp. 126-144. Waddy, S.L., and Aiken, D.E. (1990). Intermolt insemination, and alternative mating strategy for the American Lobster (Homarus americanus). Canadian Journal of Fisheries and Aquatic Sciences, 47, pp. 2402-2406. Waddy, S.L., and Aiken, D.E. (1986). Multiple fertilizations and consecutive spawning in large American lobsters, Homarus americanus. Canadian Journal of Fisheries and Aquatic Science, 43, pp. 2291-2294.

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