ALMQ Data Analysis Final Report

P.O. Box 25125, Halifax, Nova Scotia B3M 4H4 Telephone: (902) 876-1160 Fax: (902) 876-1320 www.fsrs.ns.ca

Final Report Prepared for: Fishermen and Scientists Research Society Contract: FSRS2012-022

ATLANTIC LOBSTER MOULT & QUALITY PROJECT

Data analysis and descriptive statistics

September 22, 2013

www.lobstermoult.ca

ALMQ data analysis Final Report

Table of Content

Report Contribution 3 Executive Summary 4 Project Background and Introduction 6 Materials and Methods 8 Site Selection 8 Sampling Procedure 8 Data Analysis 8 Moult Timing Definitions 10 Results 12 Descriptive Statistics 12 Moult Timing Based on Brix Index Value 15 Moult Timing Based on Moult Stage Proportion 18 Moult Timing Based on Shell Hardness 19 Comparison of Moult Timing Classification Methods 19 Graphical Comparison of Moult Timing Classification Methods 20 Interpretation 23 Acknowledgement 27 References 28 Appendices 31

ALMQ data analysis Final Report

Report Contribution

Primary authour Jean Lavallée Principal Consultant Aquatic Science & Health Services 465 University Avenue, PO Box 21116 Charlottetown, PE C1A 9H6 902.628.7981 [email protected]

Other report resource Tim Burnley Melanie Burton Owner Field Biologist Eastern Epidemiological Services Aquatic Science & Health Services 23 Karen Drive 465 University Avenue, PO Box 21116 Cornwall, PE C0A 1H8 Charlottetown, PE C1A 9H6 902.626.8118 902.940.3788 [email protected] [email protected]

John Tremblay Shannon Scott-Tibbetts Research Scientist & Head, Lobster Unit Research Biologist Population Ecology Division Fishermen and Scientists Research Society Fisheries and Oceans PO Box 25125 Bedford Institute of Oceanography Halifax, NS B3M 4H4 1 Challenger Drive 902.461.8119 Dartmouth, NS B2Y 4A2 [email protected] 902.426.3926 [email protected]

Angelica Silva Douglas Pezzack Research Scientist Biologist Coastal Ecosystem Science Division Population Ecology Division Fisheries and Oceans Fisheries and Oceans Bedford Institute of Oceanography Bedford Institute of Oceanography 1 Challenger Drive 1 Challenger Drive Dartmouth, NS B2Y 4A2 Dartmouth, NS B2Y 4A2 902.426.6525 902.426.2099 [email protected] [email protected]

ALMQ data analysis Final Report

Executive Summary

Background Since the early 2000's, there has been a significant increase in the proportion of soft shelled lobsters caught in Southwest Nova Scotia, especially at the first few weeks of the commercial fishing season in late November-early December, which has resulted in severe economic hardship for both fishers and processors. The Atlantic Lobster Moult & Quality monitoring project began in 2004 at the request of harvesters and processors, as a collaborative effort to better understand the variation in their catch quality and all information collected for this project is available on the Internet (www.lobstermoult.ca). Lobster shell hardness (and implications for quality) is directly related to moult timing.

Understanding the extent of the occurrence of different lobster quality is vital to the sustainability and health of the Canadian lobster industry. Data from 2004 to 2012 generated from this project were analysed to generate descriptive statistics.

Methodology Whenever possible, lobsters from various sites were sampled every second week inside and outside the commercial fishing season. Lobsters were sexed, measured, checked for liveliness/vigour, damage, shell hardness, sampled for hemolymph protein levels and for premoult stage assessment. In total, 117,571 lobsters were sampled between 2004 and 2012 through 981 different sampling events in 15 different locations in LFAs 33, 34 and 35.

Descriptive statistics were generated for lobster gender and size by location and year. Only the sites with sufficient data to allow for proper data analysis were further investigated and these include: Jacquard’s Ridge, Lobster Bay, Port Latour Inside, Yarmouth Inside, and Yarmouth Outside. Additionally, data collected in 2004 were insufficient to allow for valid year-to-year comparisons and therefore, weren’t included in the in-depth analysis. Interest in the identification of moult timing among the five sampling locations over the eight year study period (2005 - 2012), resulted in the evaluation of potential dataset variables that could be used to identify moult timing. The lobster variables protein blood levels (Brix), moult stage, moult stage group, and shell hardness were evaluated for potential use in identifying moult timing.

Summary of results The extensive dataset on moult indicators collected from 2005-2012 was analyzed to assess differences in moult timing. Three different methods of assessing moult timing were used: hemolymph protein levels (Brix index), shell hardness and pleopods staging for premoult assessment. The agreement among those methods was only fair at the most, which may result from the semi-objective nature of some of these methods. Moulting appears to occur at different times in different areas. Areas further from shore appeared to have later moulting periods compared to areas closer to shore. There were differences in moult timing between males and females, and lobsters of different sizes; larger lobsters moulting first.

ALMQ data analysis Final Report

There was an overall tendency for the moult to become progressively earlier from 2005 to 2011; however, this was not uniform across all sites. Moulting appears to have been later in 2012, even though this was an exceptionally warm year. Additionally, 2012 appeared to have had more moulting activity as indicated by the presence of more soft-shelled lobsters.

Recommendations Sampling methodology for this project should be reviewed after consideration of the program goals. Possible expansion of the sampling program to extend throughout the entire study period to ensure that peak moulting is captured among all locations. Expansion could include greater frequency of sampling over entire study periods (e.g., weekly sampling), using the current sampling procedure. A cost benefit analysis of the current sampling strategy vs. alternatives should be evaluated with the program goals in mind. The choice of the proper variable for the determination of moult timing needs to be further evaluated with respect to choices in cut-points (protein levels), and the repeatability and reproducibility of the other more subjective moult scoring variables (shell hardness, premoult staging).

Further work with Gaussian curve fitting methodologies may provide alternatives to expanding or modifying the current sampling program and provide a greater accuracy in determination of moult timing.

Water temperature data and other environmental parameters should be incorporated in the sampling program and in further data analysis. This could allow for the establishment of correlations between these parameters and moult timing. The incorporation of ocean forecasting models should also be considered to generate long-term moult timing predictions. These novel prediction models could have fishery management applications with lower post- harvest product downgrading and greater economic return to the industry. Among other things, the following options could be considered: dynamic fishing season opening/closing dates, sub- area and smaller scale management, at-sea management procedures, and establishment of grading/quality standards with corresponding labeling on Canadian lobster harvests.

This document represents the Final Report on the data analysis and descriptive statistics for the ATLANTIC LOBSTER MOULT & QUALITY PROJECT. This was completed in fulfillment of Contract # FSRS2012-022 between the Fishermen & Scientist Research Society (Halifax, NS) and Aquatic Science & Health Services (Charlottetown, PE).

ALMQ data analysis Final Report

Project Background and Introduction

Since the early 2000's, there has been a significant increase in the proportion of soft shelled lobsters caught in Southwest Nova Scotia (SWNS), especially at the first few weeks of the commercial fishing season in late November-early December. Anecdotal reports have estimated the proportion of soft shell lobsters to have increased from traditionally 5-10% to over 30-40% in recent years. These soft shelled lobsters often have low meat yield and poor survivability during holding and transportation. This has resulted in severe economic hardship for both fishers and processors. It is crucial to understand the extent of the occurrence of lower quality lobster in terms of shell hardness to ensure the lobster fishery in SWNS remains sustainable and healthy. The Atlantic Lobster Moult & Quality (ALMQ) monitoring project began in 2004 at the request of harvesters and processors, as a collaborative effort to better understand the variation in their catch quality. The information collected for this project is available on the Internet and allows users to look at lobster sex, size, hemolymph protein levels, moult stage and shell hardness by sampling location or dates (www.lobstermoult.ca).

The ALMQ project provided a new focus and resource to the lobster fishery by building on knowledge and capacity developed during previous years of research. The accumulated data could be used to build predictive models for landed lobster quality. Funding was initially provided by Fisheries and Oceans Canada in 2004 to the Fishermen & Scientists Research Society (FSRS) with additional financial support from the Province of Nova Scotia. In 2005, the AVC Lobster Science Centre at the University of Prince Edward Island (AVCLSC) was able to transfer the oversight and management of the project under its umbrella through funding from the Atlantic Canada Opportunities Agency (ACOA) under Round 1 of the Atlantic Innovation Fund (AIF) with continued collaboration with the FSRS, DFO, the Provincial government, industry partners and local communities. In 2007, the AVCLSC received additional funding from ACOA under the 5th round of the AIF which allowed for continuation of the project until December 2011. In 2012, the project and associated data were transferred back to the FSRS with major financial support from the Province of Nova Scotia.

To grow, crustaceans must shed their exoskeleton through the natural physiological process known as moulting (Aiken, 1980). Throughout their life cycle, new shell is being formed underneath the old one. At the time of the extrusion of the shell (ecdysis), there is an active water intake allowing expansion of the new exoskeleton which is soft (Mykles, 1980; Waddy et al, 1995). Because of this excess water, recently moulted lobsters are usually considered of lower quality due to the decreased meat yield. Post-ecdysis or postmoult lobsters are in a recovery phase with re-mineralization of the newly formed exoskeleton. This is when the water is replaced with new tissue growth (Aiken, 1980; Parrish & Martinelli-Liedtke, 1999; Passano, 1960). Following the postmoult, lobsters will move into the intermoult stage. Brylawski & Miller (2006) showed that in blue crab, this stage is significantly related to temperature; the intermoult period decreases as water temperature increases, and the same should be expected with lobsters. Variation in the length of the intermoult could also be influenced by other parameters such as salinity, diet, and limb loss (Brylawski & Miller, 2006). Pleopod staging is a technique used to differentiate the premoult stages in lobster. The technique is based on morphological changes in cuticle formation in the pleopods (Aiken, 1973). When no signs of

ALMQ data analysis Final Report

cuticle formation are present, the pleopod would be scored as a stage zero, which encompasses all intermoult and postmoult lobsters.

There are many observations of lobster moulting in summer and it is generally accepted that the major moult period for lobsters occurs in the warmer months of the year. However, there has been little systematic evaluation of how the timing varies annually and how early and late moulting may occur. Normally, adult female lobsters will moult and mate in late summer, extrude their eggs the next summer and release them the following spring or summer, while smaller females may moult and extrude in the same summer and fall (Nelson et al, 1988). It is often assumed that lobsters will moult at a specific and constant time of the year. For example, Comeau & Savoie (2001) reported that the main moulting period in the southern Gulf of St. Lawrence was between early July to early September, while Campbell (1983) suggested that majority of lobsters moult during August to October each year in the Bay of Fundy. Additionally, adverse environmental conditions

Hemolymph protein levels have been used as an important health indicator in crustaceans (Stewart & Li, 1969; Mercaldo-Allen, 1991; Paterson et al, 1999; Huang & Chen, 2001; Perazzolo, 2002). Stewart & Li (1969) also suggested that serum protein concentration in the hemolymph of American lobsters could be used as physiological criterion of environmental conditions. Ozbay & Riley (2002) suggested that refractometers could be used by industry stakeholders to assess lobsters’ response to various holding and handling practices. Circulating proteins in the hemolymph consist primarily of hemocyanin, which function is as a copper based respiratory protein (Goodwin, 1960). Other circulating proteins can include moult and female specific proteins and fibrinogen (Ferrero et al, 1983). The moult cycle is likely the most important factor affecting hemolymph protein levels in crustaceans. Hemolymph protein concentrations have been shown to increase prior to the moult followed by a rapid decrease postmoult (Bursey & Lane, 1971; Mercaldo-Allen, 1991; Spindler-Barth, 1976; Travis, 1955). Therefore, using total hemolymph protein levels as a method of assessing moult in crustaceans should be useful in monitoring annual changes in the moult cycle.

Lobster shell hardness (and implications for quality) is directly related to moult timing. Understanding the extent of the occurrence of different lobster quality is vital to the sustainability and health of the Canadian lobster industry. Temperature, photoperiod, diet, and other ecosystem factors can affect moult-timing (Aiken & Waddy, 1992; Hammond et al, 2006; Mikami, 2005; Waddy & Aiken, 1992; 1999; Waddy et al, 1995). Timing of moult is important in its effects on lobster quality because a certain period after moulting is required before lobsters harden and are in premium market quality (Retzlaff et al., 2007).

Data generated from the ALMQ project were analysed to generate descriptive statistics and outline when lobsters were moulting during a 9 year span (2005-2012) for specific locations in SWNS.

ALMQ data analysis Final Report

Materials and Methods Site Selection The initial selection of sites that were included in the project was a combination of convenient sampling and targeted sampling. The targeted sites consisted of locations chosen for their representation of the fishing efforts in LFAs 33 and 34. The selection was done by a lobster science committee with members from industry, academia, research, Federal and Provincial representatives.

Sampling procedure Sampling was carried out during and outside of the commercial fishing seasons. Although the project was initiated in 2004, it wasn’t until 2005 that the sampling procedure was refined. Outside the fishing season, lobsters were sampled every 2-3 weeks, weather permitting, by chartering vessels. Traps would be set on one day, fished and retrieved the following day. During the season, the same 2-3 week sampling interval was generally the goal, but decreased fishing effort in the winter and inclement weather often resulted in longer intervals between sampling events.

For each sampling, approximately 200 lobsters were selected. Sample size was reduced to 125 in 2009 as the result of an analysis of sample size variability. Each lobster was sexed, measured (carapace length), checked for liveliness/vigour, damage, shell hardness, sampled for hemolymph protein levels and had the tip of one pleopod removed for pre-moult stage assessment. During the fishing season, the sampling was carried out within 48 hrs post-landing to allow lobsters to be re-immersed. When sampling on chartered vessels, lobsters were sampled as they were coming out of the water, and returned to the ocean once the sampling procedures were completed. To limit the potential negative impact of handling and sampling, ovigerous females were not targeted. Only legal-size lobsters were sampled during the commercial season while a certain proportion of sub-legal lobsters were assessed outside the season on board chartered vessels. In total, 117,571 lobsters were sampled between 2004 and 2012 through 981 different sampling events (Table 1).

Data Analysis The dataset was validated with errors replaced to missing values when appropriate. Descriptive statistics were generated for the dataset variables gender and size, by sampling location and sampling year (see Table 2 for a list and description of the data variables). Only the sites with sufficient data to allow for proper data analysis were further investigated and these include: Jacquard’s Ridge, Lobster Bay, Port Latour Inside, Yarmouth Inside, and Yarmouth Outside (Table 1). Additionally, data collected in 2004 were insufficient to allow for valid year-to-year comparisons and therefore, weren’t included in the in-depth analysis. ANOVA was used to investigate the effects of gender, sampling location, and sampling year on lobster size. Interest in the identification of moult timing among the five sampling locations (Jacquard’s Ridge, Lobster Bay, Port Latour Inside, Yarmouth Inside, and Yarmouth Outside) over the eight year study period (2005 - 2012), resulted in the evaluation of potential dataset variables that could be used to identify moult timing. The dataset variables, protein brix, moult stage, moult stage

ALMQ data analysis Final Report

group, and shell hardness were evaluated for potential use in identifying moult timing among the study populations.

Furthermore, the four dataset variables were dichotomized (0/1) to indicate moulting/not moulting status as follows. The continuous protein brix variable (brix 0 - 28.8) was dichotomized using a brix cut-point of 16. Lobsters with protein brix values of 16 and above, were deemed to be moulting. The ordinal moult stage variable (0 - 5.5) was dichotomized at a moult stage value of 4+, with lobsters having a moult stage value of 4 and above, deemed to be closest to moulting. The ordinal moult stage group variable (post/intermoult, passive premoult, active premoult) was dichotomized with those lobsters in active premoult deemed to be committed to moulting. The ordinal shell hardness variable (soft, medium, and hard shell) was dichotomized through combining soft and medium vs. hard.

Table 1. Sampling effort summary (years 2004-2012).

LFA Site # samplings # lobsters Date range

33 Sambro 30 4,837 Nov 04, 2004 – May 23, 2013 Lunenburg 207 5,301 Jul 13, 2004 – May 27, 2011 Moose Harbour 20 2,689 Nov 02, 2004 – Jan 16, 2013 Port Latour; Inside 111 14,503 Jun 01, 2004 – Jul 09, 2013 Outside 15 1,928 Nov 10, 2004 – Nov 02, 2012 34 Cape Sable, Inside 39 6,337 Nov 10, 2004 – Apr 16, 2013 Outside 19 3,175 Nov 09, 2004 – May 02, 2013 Lobster Bay 128 19,276 Nov 16, 2004 – Jul 04, 2013 Jacquard’s Ridge 102 15,714 Aug 09, 2004 – Jul 03, 2013 Yarmouth; Inside 122 18,361 Jul 23, 2004 – Jul 11, 2013 Outside 112 13,579 Oct 27, 2004 – Dec 02, 2012 German Bank 20 3,265 Dec 09, 2004 – Dec 12, 2012 St. Mary’s Bay 28 4,668 Dec 07, 2004 – Jul 11, 2013 Bay of Fundy 18 2,533 Nov 03, 2004 – Apr 05, 2011 35 Digby 3 375 Nov 16, 2009 – Nov 23, 2012 Total: 981 117,571 Jun 01, 2004 – Jul 11, 2013

ALMQ data analysis Final Report

Table 2. Dataset variables description.

Data field Description EDATE date of sample AREA area ID (1-27) LOCATION NAME sampling location SEX sex of lobster SIZE carapace length in mm SIZECODE size grouping (1 ≤ 82.5mm; 2 > 82.5mm) PROT_BRIX hemolymph Brix index PROT_PRE Brix as predictor of moult (0 < 16; 1 ≥ 16) SHELL shell hardness SHELLGRP shell hardness group (1:soft; 2:medium; 3:hard) SHELL_PRE Shell hardness as predictor of moult (0:soft/medium; 1:hard) MOLTSTAGE moult stage (0 - 5.5) MOLTGRP moult stage group (post/intermoult, passive premoult, active premoult) MOLT_PRE Moult stage as predictor of moult (0:post/inter/passive; 1:active) SAMPLEID unique Atlantic Veterinary College Lobster Science Centre sample ID LFA LFA area is in

Moult Timing Definitions For each of the dichotomized potential moult indicator variables (protein brix, moult stage, moult stage group and shell hardness), new variables were generated, which represented the proportion of lobsters having the specified condition within location and sampling date. The proportion variables were used to identify peak moulting using descriptive and graphical methods.

Hemolymph Protein Levels. For Brix value, peak moulting was identified when the highest proportion of lobster with protein brix values of 16 and higher were observed. It is assumed that lobsters would start moulting shortly after reaching a peak of protein levels as reflected by the Brix values. Other Brix values cut-offs were investigated; however the value of 16 was selected as being the one with the least variation.

Pleopod Staging Method. Two different moult timing definitions were used in regards of pleopod staging. For moult stage, peak moulting was identified when the highest proportion of lobster with a moult stage value of 4+ were observed. For moult stage group, peak moulting was identified when the highest proportion of active premoult lobster were observed. It is assumed that lobsters would start moulting shortly after reaching the premoult stage of 4+ or shortly after being in active premoult.

ALMQ data analysis Final Report

Shell Hardness. Moulting should occur shortly before the highest proportion of lobsters in soft-shelled condition is observed. However, the low number of lobsters in that category resulted in poor consistency and therefore, peak moulting was identified when the lowest proportion of hard shell lobster were observed

The methodology used to identify the timing of moulting had the potential to incorrectly identify peak moulting due to lack of regular and repeated sampling throughout the entire study time span (i.e. time of peak moulting may not have occurred within the sampling time span). The lack of regular and repeated sampling prohibited the generation of fitted curves to identify the time of peak (or lowest) moulting proportions, as the data were truncated with respect to time. With these limitations in mind, moult timing was solely based on either the highest or lowest proportion values for each of the moult indicator variables, as outlined above.

Statistical analysis of moult timing proved to be very challenging. Common statistical approaches, such as survival analysis (to investigate time to moult) and non-parametric methods to compare distributions, where not applicable in this study, due to the sampling methodology employed. Non-parametric methods could only be applied to investigate differences in moult distribution for factors (i.e. gender and size) within each specific location and year.

Differences across locations and years within a specific location could not be evaluated due to structural differences in moult distributions arising from the sampling methodology (i.e. differences would be due mostly to interval of sampling and sometimes to sampling timing). Graphical methods were used to compare moult timing within and across locations and years. Comparisons of moult timing for locations across years, was achieved by overlaying the distribution of the various proportion variables.

ALMQ data analysis Final Report

Results

Descriptive Statistics

Gender Table 3 shows a summary of the sampling effort for each location by year and gender. Ovigerous females, if sampled, have been grouped with non-ovigerous females. The overall proportion of male/female remained somewhat close to the 50|50 distribution, although both Port Latour Inside, Lobster Bay, St. Mary’s Bay, and Bay of Fundy had consistently more male sampled then female lobsters, while Yarmouth Outside had predominantly more female than male lobsters sampled (Table 3).

Table 3. Summary table of gender by sampling locations (years 2004 - 2012).

Sambro Lunenburg Moose Harbour Port Latour In. Port Latour Out. Year Male Female Male Female Male Female Male Female Male Female 2004 214 186 516 400 112 54 446 317 197 190 2005 271 328 339 279 214 186 1051 783 91 109 2006 531 416 37 25 86 76 1108 781 69 54 2007 335 264 317 253 34 43 883 583 - - 2008 313 212 855 615 97 103 1171 713 - - 2009 205 151 529 350 240 235 915 534 214 160 2010 251 209 166 178 237 233 1148 731 200 144 2011 254 197 236 206 190 173 804 671 214 161 2012 151 99 - - 132 118 788 712 62 63 Cape Sable Isl In. Cape Sable Isl Out. Jacquard’s Ridge Lobster Bay German Bank Year Male Female Male Female Male Female Male Female Male Female 2004 204 196 59 123 231 169 259 141 113 287 2005 605 569 250 349 955 1132 1522 874 71 129 2006 254 476 492 508 1101 1193 1855 1150 333 267 2007 316 397 121 79 1003 1097 1649 894 458 753 2008 489 436 82 118 766 879 1659 796 118 129 2009 127 123 120 124 743 757 1152 684 48 77 2010 331 294 56 69 771 729 1431 941 57 68 2011 386 239 110 140 759 741 1387 862 114 118 2012 188 187 130 120 761 739 885 740 63 62 Yarmouth In. Yarmouth Out. St. Mary’s Bay Bay of Fundy Digby Year Male Female Male Female Male Female Male Female Male Female 2004 413 325 183 241 110 49 90 87 - - 2005 1263 1251 823 1065 586 386 35 18 - - 2006 1481 1508 1128 1291 632 357 213 196 - - 2007 1607 1649 1310 1519 511 224 111 89 - - 2008 1034 1090 282 417 422 266 164 161 - - 2009 851 643 578 656 202 48 228 147 66 59 2010 956 899 510 495 138 112 298 196 92 33 2011 724 776 940 960 71 54 297 203 - - 2012 667 645 659 710 146 104 - - 53 72

ALMQ data analysis Final Report

Size Lobster size variability was evaluated through the use of ANOVA. The effects of location, year, and gender and their interactions were investigated (Table 4). Year, location, and gender were found to have significant effect (p < 0.001) on lobster size. Interaction between year and location, year and gender, location and gender, and year and location and gender, were found to have significant effect on mean lobster size (p < 0.001). All pairwise comparisons of the marginal linear predictions were made, adjusted for multiple comparisons (Scheffé’s method) (see Appendix 1). Yarmouth Outside had a significantly larger overall mean size than any other location (p < 0.05), while Yarmouth Inside had the smallest overall mean size for 2005 -2012 (Figure 1).

Table 4. ANOVA results for lobster size (mm) Source Partial SS df MS F Prob>F Model1 701511.74 79 8879.89545 78.02 < 0.005 Year 40524.3894 7 5789.19849 50.86 < 0.005 Location 92624.9999 4 23156.25 230.45 < 0.005 Year x Location 107209.189 28 3828.89961 33.64 < 0.005 Gender 339473.001 1 339473.001 2982.63 < 0.005 Year x Gender 17992.5885 7 2570.36979 22.58 < 0.005 Location x Gender 20774.9943 4 5193.74858 45.63 < 0.005 Year x Location x Gender 15247.7346 28 544.56195 4.78 < 0.005 1Adjusted R-squared = 0.0729

Mean Lobster Size for Locations by Year

Linear Prediction Linear

86 84

Jacquard’s Ridge Lobster Bay Port Latour Inside Yarmouth Inside Yarmouth Outside

2005 2006 2007 2008 2009 2010 2011 2012

Figure 1. Estimates of mean lobster size (mm) with 95% CI for all locations (2005 - 2012).

ALMQ data analysis Final Report

Significant differences in the mean lobster size were noted among the seven years. Lobster size was significantly highest overall in 2005 followed next in years 2006 and 2012 (p < 0.05). Significant differences in lobster size (p < 0.05) between sexes were noted within all locations and year combinations, where males were significantly larger than females (see Appendix 1).

The effect of gender on lobster size was also assessed (Table 5). Mean carapace length was consistently higher in male compared to female lobsters, for every year and location.

Table 5. Summary table of mean lobster size by sampling locations, year (2005 - 2012), and gender. Location Year Gender Jacquard’s Ridge Lobster Bay Port Latour Inside mean sd 95% CI mean sd 95% CI mean sd 95% CI 2005 Male 87.7 13.8 86.9 - 88.6 92.8 12.6 92.2 - 93.4 91.1 12.6 90.4 - 91.9 Female 82.6 7.91 82.1 - 83.1 86.6 8.27 86.0 - 87.1 86.4 8.47 85.8 - 86.9 2006 Male 88.1 12.5 87.4 - 88.8 91.0 11.9 90.5 - 91.6 92.2 13.2 91.4 - 93.0 Female 84.6 9.74 84.1 - 85.1 85.1 7.61 84.6 - 85.5 86.3 8.72 85.7 - 87.0 2007 Male 85.3 11.0 84.6 - 86.0 89.9 12.9 89.3 - 90.5 90.4 11.5 89.6 - 91.1 Female 83.6 9.47 83.0 - 84.2 83.3 8.19 82.7 - 88.0 86.0 8.16 85.3 - 86.7 2008 Male 87.1 11.7 86.3 - 88.0 89.0 13.1 88.4 - 89.7 89.9 10.5 89.3 - 90.6 Female 84.2 11.2 83.5 - 85.0 83.6 10.3 86.8 - 87.8 85.7 7.73 85.1 - 86.3 2009 Male 86.6 10.4 85.8 - 87.3 87.2 10.3 86.6 - 87.8 90.7 11.4 90.0 - 91.5 Female 83.6 8.89 83.0 - 84.2 83.1 7.33 85.2 - 86.1 84.9 7.46 84.3 - 85.6 2010 Male 88.8 11.9 87.9- 89.6 90.0 10.9 89.4 - 90.6 89.1 9.44 88.5 - 89.6 Female 84.6 9.69 83.9 - 85.3 83.1 7.03 82.7 - 83.6 84.7 7.25 84.2 - 85.3 2011 Male 90.6 14.2 89.5 - 91.6 90.0 11.8 89.4 - 90.6 89.2 10.6 88.5 - 89.9 Female 85.2 10.0 84.5 - 86.0 83.5 7.92 83.0 - 84.1 83.6 8.00 83.0 - 84.2 2012 Male 92.2 14.0 91.2 - 93.2 93.0 12.9 92.2 - 93.9 90.6 12.3 89.7 - 91.5 Female 86.3 11.7 85.5 - 87.1 84.9 10.5 84.2 - 85.7 84.9 9.18 84.2 - 85.5 Yarmouth Inside Yarmouth Outside Year Gender mean sd 95% CI mean sd 95% CI 2005 Male 87.4 13.3 86.7 - 88.2 97.3 18.8 96.0 - 98.5 Female 81.9 7.87 81.4 - 82.3 90.4 12.0 89.7 - 91.2 2006 Male 87.4 10.2 86.9 - 88.0 87.6 11.2 87.0 - 88.3 Female 84.6 84.3 84.2 - 85.0 86.8 9.64 86.2 - 87.3 2007 Male 87.1 9.80 86.7 - 87.6 87.5 11.5 86.9 - 88.1 Female 84.4 7.41 84.1 - 84.8 86.4 8.84 86.0 - 86.8 2008 Male 85.5 9.33 85.0 - 86.1 89.1 13.5 87.5 - 90.6 Female 83.5 7.64 83.0 - 84.0 88.4 10.9 87.4 - 89.5 2009 Male 87.4 11.1 86.7 - 88.2 90.2 10.6 87.9 - 89.6 Female 82.9 7.30 82.3 - 83.5 86.3 9.69 83.9 - 85.3 2010 Male 87.8 11.7 87.0 - 88.5 88.9 10.8 88.0 - 89.8 Female 83.5 7.13 83.0 - 83.9 86.3 6.77 85.7 - 86.9 2011 Male 87.1 11.9 86.2 - 87.9 92.6 15.2 91.6 - 93.6 Female 83.1 9.59 82.5 - 83.8 88.1 10.7 87.4 - 88.7 2012 Male 89.3 13.0 88.8 - 90.8 90.5 11.6 89.6 - 91.2 Female 83.8 7.62 83.2- 84.3 86.0 8.31 85.4 - 86.6

ALMQ data analysis Final Report

When looking at the Quantile-Quantile plot (Figure 2), we see that the majority of the points are to the left of the main diagonal, meaning that males are generally larger than females (with the exception of the smallest and largest lobster). The inequality in size is greatest in lobster between 95 - 160 mm. See Appendices 2-6 for box plots of lobster size by gender, according to locations, for the years 2005-2012.

Figure 2. Q-Q Plot for lobster size, all locations and years (2005 - 2012) combined.

Moult Timing Based on Brix Index Value (hemolymph protein levels)

Several options were considered in trying to define moult timing based on hemolymph protein levels assessed via the Brix index. Using a Brix index of 16 or higher yielded the most consistent results. When looking at moult timing based on Brix of 16 or higher, we see that the timing of the moult varied by location and year (Table 6). In general, sampled lobsters from Yarmouth Outside and Inside seemed to be moulting later in the year compared to sampled lobsters from Lobster Bay and Port Latour Inside (see appendices 7-11 for overlaid plots of protein Brix (16+) proportion for years 2005 – 2012).

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Table 6. Moult timing based on Brix value (protein levels) proportion (moult indicated by protein Brix value 16).

Port Latour Yarmouth Jacquard’s Ridge Lobster Bay Yarmouth Inside Year Inside Outside Date Prop. Date Prop. Date Prop. Date Prop. Date Prop. 2005 Aug 16 .219 May 06 .430 Jul 20 .062 Aug 25 .224 Sep 09 .189 2006 Jul 25 .206 May 09 .210 Aug 14 .051 Aug 31 .211 Aug 03 .197 2007 Aug 29 .300 May 10 .508 Jun 16 .135 Aug 08 .540 Aug 07 .526 2008 Jul 02 .231 Jun 19 .147 Jul 07 .033 Jul 10 .237 Aug 21 .412 2009 Jun 25 .168 May 13 .328 Jul 01 .080 Jul 14 .296 Jun 30 .267 2010 Jun 10 .176 May 25 .312 Apr 23 .352 Jun 29 .619 Jun 02 .256 2011 Jul 04 .184 May 25 .460 May 27 .232 May 04 .232 Jul 25 .255 2012 Jul 03 .320 Jul 04 .240 Aug 21 .024 Jun 29 .193 Jun 15 .261

Statistical Evaluation of Differences in Distribution of Protein Brix Moult Proportion by Gender Statistical evaluation of the differences in the distribution of moult proportion distributions for gender was evaluated using the Kruskal-Wallis test. Significantly different distributions were found among some comparisons within location and year, with no apparent common trends in ranking. Samples may not represent a truly random sample if there are differences in trapping rates between genders. Comparisons across locations and years cannot be made due to the effect of sampling procedures on the distribution of Brix index data. Differences would be artificial due to different data collection periods among the locations and years. (see Appendices 12-16 show overlaid plots of moult proportion by gender, for all years standardized by days since January 1st, and by location).

Statistical Evaluation of Differences in Distribution of Protein Brix Moult Proportion by Size Statistical evaluation of the differences in the distribution of moult proportion distributions for lobster size was evaluated using the Kruskal-Wallis test. A dichotomized lobster size variable was created by splitting lobsters in two categories: those < 82.5 mm and those ≥ 82.5 mm. Significantly different distributions were found among some comparisons within location and year, with a higher frequency of lobster < 82.5 mm ranking higher than expected. Samples may not represent a truly random sample if there were biases in trapping rates (see Appendices 17- 21 show overlaid plots of moult proportion by size, for all years standardized by days since January 1st, and by location).

Investigation in the moult timing by year and by size category was done for each location; Figure 3 shows the original estimation of moult timing for all sizes for Jacquard’s Ridge for 2005, and the estimates for the 2 different size categories (other years and locations not shown). The estimated difference in moult timing is approximately two weeks, with larger lobsters moulting first.

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| Summary of Brix value, all size date | Mean Std. Dev. Freq. ------+------18may2005 | .235 .01670051 200 21jun2005 | .10540633 .04439127 173 22jun2005 | .10144928 .02786156 69 05jul2005 | .065 .00011911 200 22jul2005 | .095 .04195613 200 02aug2005 | .18831169 .02107356 154 16aug2005 | .21978022 .11028984 91 30aug2005 | .18000001 .10803343 200 16sep2005 | .075 .0919551 200 27sep2005 | .01 .01336679 200 12oct2005 | .01 .01887663 200 05dec2005 | .08 .01839932 200 ------+------Total | .10744384 .09068697 2087

| Summary of Brix value, size ≤ 82.5mm date | Mean Std. Dev. Freq. ------+------18may2005 | 0 0 1 21jun2005 | .09032258 0 155 22jun2005 | .125 0 40 05jul2005 | .06493507 0 154 22jul2005 | .07954545 0 176 02aug2005 | .18115942 0 138 16aug2005 | .17105263 0 76 30aug2005 | .26890758 0 119 16sep2005 | .2 0 70 27sep2005 | .02777778 0 72 12oct2005 | .04545455 0 44 05dec2005 | 0 0 10 ------+------Total | .12417062 .07347649 1055

| Summary of Brix value, size > 82.5mm date | Mean Std. Dev. Freq. ------+------18may2005 | .2361809 0 199 21jun2005 | .23529412 0 18 22jun2005 | .06896552 0 29 05jul2005 | .06521739 0 46 22jul2005 | .20833333 0 24 02aug2005 | .25 0 16 16aug2005 | .46666667 0 15 30aug2005 | .04938272 0 81 16sep2005 | .00769231 0 130 27sep2005 | 0 0 128 12oct2005 | 0 0 156 05dec2005 | .08421053 0 190 ------+------Total | .09034428 .10267175 1032

Figure 3. Summary Brix values (mean proportion of lobsters > 16+) for Jacquard's Ridge (2005) for all size combined, and for the two different size categories. Highlighted lines represent the estimated time of moult.

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Moult Timing Based on Moult Stage Proportion

Several options were also considered in trying to define moult timing based on the pleopod method for premoult staging. Two different options were kept in the final analysis; moult was assumed to happen when lobsters reached premoult stage of 4+ (advance active premoult), and moult was assumed to happen when lobsters reached premoult stage grouping of 3 (commitment to moult). In both options, moult timing should occur shortly after a peak in these 2 premoult cut-off points.

When looking at moult timing based on premoult stage of 4+, we see that the timing of the moult varied by location and year (Table 7). The pattern of variation in moult timing is unclear and inconsistent among the locations while variation over the years showed a general tendency to have moult happening earlier in the year with the exception of 2012 (see appendices 22-26 for overlaid plots of moult stage (4+) proportion for years 2005 – 2012). When looking at the data based on the highest proportion of lobsters in active premoult (Table 8), we also observe a general shift towards earlier moult timing as the years progressed (see appendices 27-31 for overlaid plots of premoult stage group (4) proportion for years 2005 – 2012).

Table 7. Timing of moult based on premoult stage proportion (moult indicated by premoult stage of 4+).

Jacquard’s Ridge Lobster Bay Port Latour Inside Yarmouth Inside Yarmouth Outside Year Date Prop. Date Prop. Date Prop. Date Prop. Date Prop. 2005 Aug 02 .052 Sep 17 .015 Sep 02 .045 Sep 19 .045 Aug 09 .059 2006 Sep 06 .040 Aug 09 .020 Aug 29 .020 Sep 14 .060 Sep 15 .097 2007 Jul 14 .030 Aug 28 .026 Jul 10 .044 Sep 06 .040 Sep 05 .086 2008 Jul 02 .263 Jul 29 .196 Jun 24 .214 Aug 07 .410 Aug 21 .706 2009 Aug 05 .073 Jul 22 .040 Jul 01 .032 Jun 19 .056 Jul 27 .057 2010 Jul 20 .026 May 25 .024 Jun 30 .066 Jun 18 .072 Aug 30 .080 2011 Jul 20 .048 Aug 03 .024 May 27 .116 Sep 01 .024 Jul 25 .049 2012 Aug 01 .096 Sep 06 .020 Aug 07 .064 Oct 12 .085 Jul 11 .104

Table 8. Timing of moult based on premoult stage group proportion (moult indicated by premoult stage group of 3).

Jacquard’s Ridge Lobster Bay Port Latour Inside Yarmouth Inside Yarmouth Outside Year Date Prop. Date Prop. Date Prop. Date Prop. Date Prop. 2005 Sep 16 .135 Jul 06 .175 Jul 08 .303 Jul 12 .210 Sep 09 .170 2006 Jul 25 .175 Jun 29 .170 Jul 05 .128 Jul 04 .209 Aug 03 .171 2007 Aug 14 .173 May 22 .302 Jul 25 .150 Aug 08 .176 Sep 05 .298 2008 Jul 02 .409 Jun 19 .425 Jul 07 .386 Aug 07 .508 Aug 21 .706 2009 Aug 05 .234 Apr 20 .400 May 14 .192 Jun 29 .176 Jun 18 .202 2010 Jun 22 .232 Apr 08 .592 Apr 23 .576 Apr 21 .416 Jun 02 .302 2011 Jun 21 .336 May 10 .648 May 27 .642 May 04 .352 Aug 11 .540 2012 Jul 03 .176 Sep 06 .301 Jun 12 .136 Oct 12 .149 Jul 11 .248

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To account for the fact that lobsters will only moult after a certain period post premoult stage 4+ and/or moult stage group 3, an adjusted timing of the moult was calculated. This calculation was based on Campbell (1983) who reported that lobsters will moult an average of 3-6 weeks after reaching stage 3 and only 0.5-3 weeks after reaching stage 4. Twenty-one days were added to the date with the highest proportion of lobsters reaching premoult group stage 3 and 7 days to premoult stage 4+ (Table 9).

Table 9. Adjusted timing of moult based on premoult stage proportion (moult indicated by premoult stage of 4+ plus 7 days) and premoult stage group proportion (moult indicated by premoult stage group of 3 plus 21 days).

Yarmouth Jacquard’s Ridge Lobster Bay Port Latour Inside Yarmouth Inside Outside Year Stage Stage Stage Stage Stage Stage Stage Stage Stage Stage 4+ 3 4+ 3 4+ 3 4+ 3 4+ 3 2005 Aug 09 Oct 07 Sep 24 Jul 27 Sep 09 Jul 29 Sep 26 Aug 02 Aug 16 Sep 30 2006 Sep 13 Aug 15 Aug 16 Jul 20 Sep 05 Jul 26 Sep 21 Jul 25 Sep 22 Aug 24 2007 Jul 21 Sep 04 Sep 04 Jun 12 Jul 17 Aug 15 Sep 13 Aug 29 Sep 12 Sep 26 2008 Jul 09 Jul 23 Aug 05 Jul 10 Jul 01 Jul 28 Aug 14 Aug 28 Aug 28 Sep 11 2009 Aug 12 Aug 26 Jul 29 May 10 Jul 08 Jun 04 Jun 26 Jul 19 Aug 03 Jul 09 2010 Jul 27 Jul 13 Jun 01 Apr 29 Jul 07 May 14 Jun 25 May 12 Sep 06 Jun 23 2011 Jull 27 Jul 12 Aug 10 May 31 Jun 03 Jun 17 Sep 08 May 25 Aug 01 Sep 01 2012 Aug 08 Jul 24 Sep 13 Sep 27 Aug 14 Jul 02 Oct 19 Nov 02 Jul 18 Aug 01

Moult Timing Based on Shell Hardness

Table 10 shows the moult timing based on the lowest proportion of lobsters in hard-shell conditions. The span of moult timing seems higher over the years when looking at shell hardness. For example in Yarmouth Outside, lobsters appeared to have moulted as early as late May 30 in 2005 and as late as late October in 2008. Additionally, the general tendency for lobsters to moult earlier each year was not observed with shell hardness. See appendices 32-36 for overlaid plots of lowest hard-shell proportion for years 2005 – 2012.

Table 10. Timing of moult based on shell hardness (moult indicated by lowest proportion of hard-shell lobsters).

Jacquard’s Ridge Lobster Bay Port Latour Inside Yarmouth Inside Yarmouth Outside Year Date Prop. Date Prop. Date Prop. Date Prop. Date Prop. 2005 Sep 16 .465 Aug 17 .455 Sep 02 .300 Sep 08 .560 May 30 .408 2006 Aug 23 .535 Sep 07 .680 Jul 18 .515 Aug 17 .655 Sep 27 .630 2007 Sep 21 .590 Jul 13 .700 Aug 07 .375 Sep 20 .570 Oct 03 .557 2008 Aug 25 .793 Jul 29 .659 Aug 05 .294 Oct 02 .679 Oct 28 .321 2009 Sep 16 .584 Sep 03 .592 Aug 12 .336 Jul 28 .400 Aug 10 .432 2010 Oct 20 .848 Oct 28 .800 Jul 14 .728 Aug 10 .815 Aug 30 .800 2011 Aug 23 .504 Aug 03 .600 Jul 26 .312 Jul 27 .528 Sep 28 .648 2012 Jul 17 .568 Oct 01 .614 Aug 21 .815 Sep 28 .696 Sep 27 .650

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We see that moult indicator variables only show slight to fair agreement (Table 11), with the highest agreement between Brix value and moult stage group, and moult stage group and moult stage. These kappa values indicate that neither of the moult indicator variables is consistently capturing the same lobster moult status.

Table 11. Agreement between the four potential dichotomized moult indicator variables using the kappa statistic. Moult stage Moult stage group Shell hardness (> 4+) (active premoult) (lowest % of hard-shell) Brix value (16+) Kappa 0.16431 Kappa 0.35751 Kappa -0.12051 (slight agreement) (fair agreement) (disagreement) Moult stage (> 4+) - Kappa 0.33661 Kappa 0.02601 (fair agreement) (slight agreement) Moult stage group - - Kappa -0.11741 (active premoult) (disagreement) 1The level of agreement better than that expected due to chance alone, p < 0.001.

Graphical Comparison of Moult Timing Classification Methods

Graphical evaluation of moult timing based on the four moult classification variables illustrated the lack of strong agreement between the four moult timing indicator variables (see Figures 4 – 8). Agreement between the four methods of identifying moult timing varied and appears to be clustered within specific locations and years, with no clear trend.

Figure 4. Comparison of moult timing for Jacquard’s Ridge for each potential moult indicator predictor variable (2005 – 2012).

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Figure 5. Comparison of moult timing for Lobster Bay for each potential moult indicator predictor variable (2005 – 2012).

Figure 6. Comparison of moult timing for Port Latour Inside for each potential moult indicator predictor variable (2005 – 2012).

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Figure 7. Comparison of moult timing for Yarmouth Inside for each potential moult indicator predictor variable (2005 – 2012).

Figure 8. Comparison of moult timing for Yarmouth Outside for each potential moult indicator predictor variable (2005 – 2012).

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Interpretation

Four different indices were evaluated for their potential as moult predictors for the American lobster. The challenge was to identify a way to properly capture moult timing to allow for comparisons among locations and among years. Although the dataset was significant with more than 115,000 individual records from 2004 to 2012, collecting data at consistent intervals simultaneously for all locations was impractical and proved to be a limiting factor in the data analysis.

All data in this project were collected using commercial lobster traps. This was the only practical method of sampling lobsters on the scale needed. Trap sampling could be supplemented by other methods (SCUBA, trawl) but all methods have potential sampling bias. The trap sampling program used here may have selected lobsters that were not representative of the entire natural lobster population. If the likelihood for lobsters to enter a trap was not consistent across the entire population, then it was always possible that only certain types of lobsters would have been selected for the study. Perimoult lobsters could be adopting shelter- specific behaviours and therefore, the true seasonal pattern of moulting in the population might not be captured by the sampling program. The three main methods used to assess moult also had intrinsic limitations. Although the Brix refractometric method is objective, the protein levels can be affected by handling. The pleopod staging method is semi-objective, and some intra-animal variation can be observed (Lavallée, unpublished data). Finally, shell hardness was assessed by manual squeezing of the lobster carapace in three different areas. This method is subjective and can have repeatability and reproducibility issues associated with it.

Several options were considered when trying to identify a hemolymph protein levels cut-offs that could be used as a moult predictor. The cut-off value of 16 on the Brix refractometers was selected as the predictor; it was assumed that moulting would start shortly after the highest proportion of lobsters have reached a Brix value of 16 or higher. From the data collected in this project, it is clear that the timing of the moult varied by location and year, although the patterns for these variations are unclear. In a follow-up paper from a review on growth completed 20 years earlier, Hartnoll (2001) raised the issue that little is known in terms of what determines the wide intrinsic intermoult period both within and between crustacean species. From these data, lobsters from Yarmouth Outside and Inside appeared to be moulting later in the year compared to lobsters from Lobster Bay and Port Latour Inside. These findings are consistent with an earlier and preliminary analysis of some of these data (Retzlaff et al., 2007). There was also a general although not consistent tendency for lobsters to moult somewhat earlier each year. This yearly variation in moult timing is consistent with other studies. When looking at the lobsters in the southern Gulf of St. Lawrence, Comeau & Savoie (2001) reported that premoult lobsters were always observed in May and June; female premoult lobsters were not observed later than early September, while premoult males could still be found by the end of October in some years. The reasons for the tendency for earlier moulting over the years are not clear. Ocean water temperature may be a factor and more work is needed on obtaining the best temperature data for the sites and periods examined. Significantly, water temperatures in 2012 were the warmest in many years (DFO, 2013), yet this study indicated that moulting occurred later in that year. A potential hypothesis for the contradictory results for 2012 could

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be explained by warm water temperatures in the fall of 2011, which could have helped triggered a late fall 2011 moult in animals that normally would have only moulted in 2012.

The pleopod staging method was used in this project to assess where sampled lobsters were in their premoult cycle. Two variations of this method were used to define moult timing in lobsters; when the highest proportion of lobsters had reach the active premoult or stage group 3 (committed to moult), and when the highest proportion of lobsters had reached the late active premoult or stage 4+. When looking at moult timing based on those two variables, we observed that moulting patterns varied by location and year. However, these variations were unclear and inconsistent among the locations. As per the hemolymph protein levels, we noted a general tendency to have moult happening earlier in the year with the exception of 2012. Campbell (1983) suggested that lobsters with pleopod stages 3 to 3.5 would likely moult 3 to 6 weeks later while lobsters with pleopod stages 4 to 4.5 would likely moult 0.5- 3 weeks later, and this was reflected in this study with an earlier moult timing identified by the moult stage group of 3 normally being reached before the premoult stage 4+. Again, lobsters from Lobster Bay and Port Latour Inside appeared to be moulting before lobsters from the other locations, suggesting a variation shift in moult timing between inshore and more offshore locations.

Finally, shell hardness was also assessed as a potential indicator of moult timing. The value and role of this parameter alone as an indicator seems lower than those of other parameters; this could be due to the fact that shell hardness is a semi-subjective test. More objective methods to assess shell hardness should be investigated, and potentially incorporated into the sampling procedure. The durometer has been used in the past by industry and the scientific community, and could represent an option. However, the reproducibility and repeatability of this technique is less than optimal. The general tendency for lobsters to moult earlier each year was not observed when looking at the lowest proportion of lobsters being classified as having hard shell. The proportions of lobsters in the soft, and to a certain degree the medium shell categories were small, making the data analysis less rigid and more difficult to complete. Over the last 10 years or so, anecdotal reports have been made in regards of increased number of soft shelled lobsters at the start of the fishing season in SWNS. One of the hypotheses often put forward by industry stakeholders is that this increase in soft shelled lobsters could be due to a double moult. Comeau & Savoie, (2001) reported that there was virtually no evidence of double moulting in the southern Gulf of St. Lawrence. The increased proportion of soft shelled lobsters in late fall is likely due to another segment of the lobster population that normally would have only moulted the following year. However, due to favorable environmental conditions, these lobsters were able to successfully complete the ecdysis before the water temperatures started to decrease substantially. This is likely what is being observed in SWNS. However, this may be a phenomenon that doesn’t necessarily apply to all lobsters as Comeau & Savoie (2001) reported that there were likely very few lobsters in the 50 to 90 mm carapace length that would skip moulting one year in the southern Gulf of St. Lawrence.

The data collected in this project showed a significant amount of variation in all parameters measured, and assessment of the duration of the moult was not achieved. This was likely due to the non-normal distribution of several parameters (high number of outliers). Some evidence show that mature female lobsters can purposely stagger their moult at intervals throughout the

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summer so they could consequently mate with dominant males (Cowan & Atema, 1990), while immature female lobsters have been shown to exhibit moulting synchrony (Karnofsky et al., 1989). This would indicate that the timing of the moult for immature females should not be influenced by the availability of dominant mature males. Additionally, Waddy & Aiken (1999) were able to show in stage III larval American lobsters the existence of a population-based moulting rhythm which has an endogenous component and is also triggered by environmental factors such as photoperiod. This staggering is likely indicative of active control of moult timing in female lobsters. If this moult staggering was happening when lobsters were sampled, it is likely to be reflected in these outliers.

The analysis accounted for size, sex, and sex by size interaction, when generating the moult proportion values for the protein levels assessed via the Brix refractometer. Moult timing was slightly different among the size and sex groups, usually within a two week period. This difference could be real but could also partially reflect the sampling interval. Some of the data became very sparse when adjusting for size and sex simultaneously, especially when looking at premoult stages and shell hardness. One of the major goals of this project was to identify possible ways to predict moult timing. Breaking down the data further by size and sex categories may provide limited benefit as the moult timing prediction should be for the entire population, and not only commercial size (≥82.5mm). Because of some variation in moult- timing according to size and gender, these variables could be further investigated for fishery management protocols (gender and/or size specific fishery). Moult timing should vary with population demographics and this could contribute to the lack of consistency found in this study. However, other factors, such as: environmental conditions, genetic differences, geographic location of sampling, sampling timing and sampling interval, could likely have a greater impact on moult timing.

The lack of consistent agreement among moult predicting variables could be due to measurement error, variability in scoring among data collection technicians, and issues pertaining to the truncation of data records. Additionally, the chosen cut-points for the variables to indicate moult vs. non-moult status may not be biologically equivalent. Further evaluation through Bayesian analysis may reveal the diagnostic performance of the moult indication measures.

Gaussian curve fitting was attempted to deal with the issue of data truncation, arising from the lack of continuous data collection over the span of the study. This approach restricts the curves of the moult indicator proportions to a common shape based on evidence from the dataset. This allows for the identification/confirmation of peak moulting when dealing with truncated data where the peak moult period may fall outside of the data range. This methodology could provide a peak moulting time and a standard error for the estimate allowing statistical comparison within locations and across years. An attempt at using this approach was implemented using the open source statistical program R. However, the proportion data for the moult indicator variables contained numerous values close to zero requiring the transformation of the proportion variables to a better scale. This process of optimizing the transformation of the data to a better scale could prove to be extremely time consuming, but could warrant further investigation.

ALMQ data analysis Final Report

Summary  The extensive dataset on moult indicators collected from 2005-2012 was analyzed to assess differences in moult timing.  The analysis of data on temperature and other factors with the potential to affect moult timing was beyond the scope of the current project.  Moulting appears to occur at different times in different areas. Areas further from shore appeared to have later moulting periods.  There were important differences in moult timing between males and females, and lobsters of different sizes.  There was a tendency for the lobster moult to become progressively earlier from 2005 to 2011; this was not uniform across sites.  Moulting appears to have been later in 2012, even though this was an exceptionally warm year.

Recommendations & next steps  Data summaries as presented are of interest to create a baseline of tangible observations that reflect biological status of sampled lobsters and indicate that moulting appears to occur at different times in different areas. Important to separate gender and sizes in groupings that may help interpret highest moulting rates in smaller vs. larger lobsters. Some sites have more males sampled than females and it may skew results.  The goals of a revamped ALMQ program need to be clearly identified from the perspective of industry (i.e., how much effort should be directed at understanding the biology of moult timing vs. predicting the shell hardness of commercial size lobsters during the fishing season).  Sampling methodology for the ALMQ program should be reviewed after consideration of the program goals. Possible expansion of the sampling program to extend throughout the entire study period to ensure that peak moulting is captured among all locations. Expansion could include greater frequency of sampling over entire study periods (e.g., weekly sampling), using the current sampling procedure.  Cost benefit analysis of the current sampling strategy vs. alternatives should be evaluated with the program goals in mind.  Choice of moult timing variable for the determination of moult timing needs to be further evaluated with respect to choices in cut-points (protein levels), and the repeatability and reproducibility of the other more subjective moult scoring variables (i.e. shell hardness). Improved agreement among the moult timing variables may allow for reliance upon one moult indicator variables for determination of moult timing, thereby allowing for increased data collection efficiency. Additionally, evaluation of incorporating multiple scores to determine moult timing may be beneficial.

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 Further work with Gaussian curve fitting methodologies may provide alternatives to expanding or modifying the current sampling program and provide a greater accuracy in determination of moult timing.  Water temperature data and other environmental parameters should be incorporated in the sampling program and in further data analysis. This could allow for the establishment of correlations between these parameters and moult timing. The incorporation of ocean forecasting models should also be considered to generate long- term moult timing predictions. These novel prediction models could have fishery management applications with lower post-harvest product downgrading and greater economic return to the industry. Among other things, the following options could be considered: dynamic fishing season opening/closing dates, sub-area and smaller scale management, at-sea management procedures, and establishment of grading/quality standards with corresponding labeling on Canadian lobster harvests.

Acknowledgements

The authours wish to acknowledge the contributions of the various professional and technical staff that made the field sampling, data management and website possible; S. Allain, B. Bernard, E, Branton, B. Cabot, J. Dagley, A. Darling, C. Denton, L. Edwards, G. Jackson, C. MacDonald, N. MacDonald, T. Pearo, B. Petrie, A. Retzlaff and J. Silver. Their efforts and countless hours of dedication are greatly valued.

The assistance of the ALMQ steering committee is gratefully acknowledged. The intellectual contributions and dedicated support of the following individuals were invaluable to the success of this undertaking: J. Garland, L. Greening, S. Greenwood, P. King, C. MacKenzie, D. Morrow, R. Claytor, W. Smith, A. Spinney, M. Surette.

Finally, this project would not have been possible without the countless hours of hard work of our industry participants; M. Angleson, L. Blackland, P. Conrad, G. Cottreau, G. Cunninham, S. Denton, N. Farstad, E. Garron, A. Gray, R. Goodwin, E. MacKay, C. Nickerson, F. Nickerson, S. Lyonas, H. Saulnier, S. Scobey, J. Simpson, G. Smith, W. Smith, A. Spinney, J. Spinney. Their dedication and contribution allowed the technical staff to carry out the field sampling in a professional and safe environment. To all, thank you.

ALMQ data analysis Final Report

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Retzlaff, A., Claytor, R., Petrie, B., Frail, C., Tremblay. M.J., Pezzack, D., & Lavallée, J. (2007) Variation in Moult Timing and Market Quality in the American Lobster (Homarus americanus). Bedford Institute of Oceanography: 2006 in Review. Her Majesty the Queen in Right of Canada, Cat. No. Fs101-3/2006E, ISBN: 978-0-662-46177-7, ISSN: 1499-9951. Pp. 22-26

Spindler-Barth, M. (1976). Changes in the chemical composition of the common shore crab, carcinus maenas, during the molting cycle. Journal of Comparative Physiology B, 105, 197-205.

Stewart, J. E. & Li, M. F. (1969). A Study of Lobster (Homarus Americanus) Ecology Using Serum Protein Concentration as an Index. Canadian Journal of Zoology 47, 21-28.

Travis, D. F. (1955). The molting cycle of the spiny lobster, panulirus argus latreille. III. physiological changes which occur in the blood and urine during the normal molting cycle. The Biological Bulletin, 109, 484-503.

Waddy, S. L., & Aiken, D. E. (1990). Intermolt insemination, an alternative mating strategy for the American lobster (Homarus americanus). Canadian Journal Of Fisheries And Aquatic Sciences, 47(12), 2402-2406.

Waddy, S. L., & Aiken, D. E. (1992). Seasonal variation in spawning by preovigerous American lobster (Homarus americanus) in response to temperature and photoperiod manipulation. Canadian Journal Of Fisheries And Aquatic Sciences, 49(6), 1114-1117.

Waddy, S. L., & Aiken, D. E. (1999). Timing of the metamorphic molt of the American lobster (Homarus americanus) is governed by a population-based, photoperiodically entrained daily rhythm. Canadian Journal Of Fisheries And Aquatic Sciences, 56(12), 2324-2330.

Waddy, S. L., & Aiken, D. E. (2000). Problems with the terminology proposed by Lavalli & Lawton for certain life history phases of the American lobster Homarus americanus. Marine Ecology- Progress Series, 197309-310.

Waddy, S. L., Aiken, D. E., & De Kleijn, D. P. V. (1995). Control and growth of reproduction. In J. R. Factor (Ed.), Biology of the lobster homarus americanus (pp. 217-266). San Diego, CA: Academic Press, Inc.

ALMQ data analysis Final Report

Appendices

Appendix 1. Predicted mean lobster size for location, year, gender and interactions.

Predictive margins Number of obs = 76171

Expression : Linear prediction, predict()

------| Number of | Comparisons ------+------location | 5 year | 8 location#year | 40 sex | 2 location#sex | 10 year#sex | 16 location#year#sex | 80 ------

------| Delta-method Bonferroni Bonferroni | Margin Std. Err. z P>|z| [95% Conf. Interval] ------+------location | 1 | 86.30299 .0905047 953.58 0.000 86.06986 86.53611 2 | 87.56035 .0806986 1085.03 0.000 87.35249 87.76822 3 | 88.16356 .0947671 930.32 0.000 87.91946 88.40767 4 | 85.59752 .0827985 1033.81 0.000 85.38425 85.8108 5 | 88.97288 .0990553 898.21 0.000 88.71773 89.22803 | year | 2 | 88.49662 .1032418 857.18 0.000 88.21432 88.77892 3 | 87.59357 .0953906 918.26 0.000 87.33274 87.85441 4 | 86.61377 .099154 873.53 0.000 86.34265 86.8849 5 | 86.70415 .1223615 708.59 0.000 86.36957 87.03873 6 | 86.33793 .1238438 697.15 0.000 85.9993 86.67657 7 | 86.85784 .1176021 738.57 0.000 86.53628 87.17941 8 | 87.4054 .1165193 750.14 0.000 87.0868 87.72401 9 | 88.4826 .1266009 698.91 0.000 88.13642 88.82877 | location#year | 1 2 | 85.3672 .2365309 360.91 0.000 84.60386 86.13054 1 3 | 86.46032 .2242288 385.59 0.000 85.73668 87.18395 1 4 | 84.52187 .2344851 360.46 0.000 83.76514 85.27861 1 5 | 85.79596 .2659353 322.62 0.000 84.93773 86.65419 1 6 | 85.18538 .2764477 308.14 0.000 84.29322 86.07753 1 7 | 86.84419 .2757695 314.92 0.000 85.95422 87.73416 1 8 | 88.09302 .2760133 319.16 0.000 87.20227 88.98378 1 9 | 89.45229 .2759678 324.14 0.000 88.56168 90.3429 2 2 | 89.90754 .2223774 404.30 0.000 89.18988 90.6252 2 3 | 88.26455 .1972084 447.57 0.000 87.62811 88.90098 2 4 | 86.83636 .2171724 399.85 0.000 86.13549 87.53722 2 5 | 86.50721 .2345847 368.77 0.000 85.75016 87.26427 2 6 | 85.30156 .2532327 336.85 0.000 84.48432 86.1188 2 7 | 86.81771 .2210097 392.82 0.000 86.10446 87.53095 2 8 | 87.0191 .2279116 381.81 0.000 86.28358 87.75462 2 9 | 89.29135 .2646781 337.36 0.000 88.43718 90.14553 3 2 | 88.93044 .2497521 356.07 0.000 88.12444 89.73644 3 3 | 89.49082 .2467218 362.72 0.000 88.6946 90.28705 3 4 | 88.3457 .2810517 314.34 0.000 87.43868 89.25271 3 5 | 87.9809 .2494333 352.72 0.000 87.17592 88.78587 3 6 | 88.04907 .2855769 308.32 0.000 87.12745 88.97069 3 7 | 87.06487 .2488775 349.83 0.000 86.26168 87.86805 3 8 | 86.59933 .2778143 311.72 0.000 85.70276 87.49589 3 9 | 87.94903 .2755441 319.18 0.000 87.05979 88.83827 4 2 | 84.86809 .2133057 397.87 0.000 84.1797 85.55647 4 3 | 86.12106 .1958324 439.77 0.000 85.48906 86.75305 4 4 | 85.8888 .1876936 457.60 0.000 85.28307 86.49453 4 5 | 84.59156 .2327282 363.48 0.000 83.8405 85.34262 4 6 | 85.33848 .2765837 308.54 0.000 84.44589 86.23108 4 7 | 85.75028 .2479508 345.84 0.000 84.95009 86.55047 4 8 | 85.25311 .2771263 307.63 0.000 84.35876 86.14745 4 9 | 87.02595 .2950414 294.96 0.000 86.07379 87.97811 5 2 | 94.10027 .2506505 375.42 0.000 93.29136 94.90917 5 3 | 87.22393 .2191576 398.00 0.000 86.51666 87.9312 5 4 | 86.99493 .2028417 428.88 0.000 86.34032 87.64955

ALMQ data analysis Final Report

Appendix 1. Predicted mean lobster size for location, year, gender and interactions (cont.)

5 5 | 88.77836 .4188606 211.95 0.000 87.42661 90.13012 5 6 | 88.39031 .3066177 288.28 0.000 87.40078 89.37983 5 7 | 87.69518 .3371431 260.11 0.000 86.60714 88.78321 5 8 | 90.49941 .2456551 368.40 0.000 89.70663 91.29219 5 9 | 88.39866 .2901657 304.65 0.000 87.46223 89.33508 | sex | 1 | 89.28467 .0531267 1680.60 0.000 89.1656 89.40375 2 | 84.81299 .0575376 1474.04 0.000 84.68402 84.94195 | location#sex | 1 1 | 88.08565 .1291133 682.24 0.000 87.72323 88.44808 1 2 | 84.24579 .1255315 671.11 0.000 83.89341 84.59816 2 1 | 90.45123 .1004128 900.79 0.000 90.16936 90.73309 2 2 | 84.21886 .1297435 649.12 0.000 83.85467 84.58306 3 1 | 90.51151 .1229719 736.03 0.000 90.16632 90.85669 3 2 | 85.43823 .1470141 581.16 0.000 85.02556 85.8509 4 1 | 87.39136 .1162524 751.74 0.000 87.06503 87.71768 4 2 | 83.52506 .1175271 710.69 0.000 83.19516 83.85496 5 1 | 90.35054 .1442537 626.33 0.000 89.94562 90.75547 5 2 | 87.39118 .1325104 659.50 0.000 87.01922 87.76314 | year#sex | 2 1 | 91.14526 .1431814 636.57 0.000 90.72214 91.56839 2 2 | 85.42476 .1525746 559.89 0.000 84.97387 85.87564 3 1 | 89.27894 .1311627 680.67 0.000 88.89133 89.66654 3 2 | 85.39495 .1412932 604.38 0.000 84.9774 85.81249 4 1 | 88.0955 .1344072 655.44 0.000 87.69831 88.4927 4 2 | 84.61805 .1506025 561.86 0.000 84.173 85.06311 5 1 | 88.06783 .1740725 505.93 0.000 87.55342 88.58224 5 2 | 84.89954 .1790836 474.08 0.000 84.37032 85.42877 6 1 | 88.27317 .165944 531.95 0.000 87.78277 88.76356 6 2 | 84.02993 .1884599 445.88 0.000 83.473 84.58686 7 1 | 88.90967 .1597589 556.52 0.000 88.43756 89.38179 7 2 | 84.30458 .1747344 482.47 0.000 83.78821 84.82095 8 1 | 89.76054 .1607329 558.45 0.000 89.28555 90.23553 8 2 | 84.5653 .1705935 495.71 0.000 84.06117 85.06944 9 1 | 91.28183 .1756415 519.71 0.000 90.76278 91.80088 9 2 | 85.09605 .181224 469.56 0.000 84.56051 85.6316 | location#year#sex | 1 2 1 | 87.73717 .3452244 254.15 0.000 86.55632 88.91802 1 2 2 | 82.61042 .3170881 260.53 0.000 81.52582 83.69503 1 3 1 | 88.08901 .3215211 273.98 0.000 86.98924 89.18878 1 3 2 | 84.5658 .3088751 273.79 0.000 83.50928 85.62232 1 4 1 | 85.32004 .3368625 253.28 0.000 84.16779 86.47229 1 4 2 | 83.59344 .3221068 259.52 0.000 82.49166 84.69521 1 5 1 | 87.12941 .3857203 225.89 0.000 85.81005 88.44878 1 5 2 | 84.24487 .3600443 233.98 0.000 83.01333 85.47642 1 6 1 | 86.55989 .3913891 221.16 0.000 85.22114 87.89865 1 6 2 | 83.58653 .387753 215.57 0.000 82.26021 84.91285 1 7 1 | 88.7834 .3842165 231.08 0.000 87.46918 90.09762 1 7 2 | 84.58848 .3951294 214.08 0.000 83.23693 85.94003 1 8 1 | 90.55599 .3872418 233.85 0.000 89.23142 91.88057 1 8 2 | 85.22807 .391917 217.46 0.000 83.88751 86.56863 1 9 1 | 92.16426 .3867326 238.32 0.000 90.84143 93.48709 1 9 2 | 86.2977 .3924469 219.90 0.000 84.95532 87.64007 2 2 1 | 92.78975 .2734613 339.32 0.000 91.85437 93.72513 2 2 2 | 86.55492 .3608672 239.85 0.000 85.32056 87.78928 2 3 1 | 91.01348 .247703 367.43 0.000 90.1662 91.86075 2 3 2 | 85.06696 .3145968 270.40 0.000 83.99087 86.14304 2 4 1 | 89.89691 .2627198 342.18 0.000 88.99827 90.79555 2 4 2 | 83.27629 .3568078 233.39 0.000 82.05582 84.49676 2 5 1 | 89.04604 .2716821 327.76 0.000 88.11675 89.97534 2 5 2 | 83.55402 .3970403 210.44 0.000 82.19593 84.9121 2 6 1 | 87.20226 .3143235 277.43 0.000 86.1271 88.27741 2 6 2 | 83.09064 .4079201 203.69 0.000 81.69534 84.48594 2 7 1 | 90.00839 .2820222 319.15 0.000 89.04372 90.97305 2 7 2 | 83.10627 .347783 238.96 0.000 81.91667 84.29587 2 8 1 | 89.99856 .2864606 314.17 0.000 89.01871 90.9784 2 8 2 | 83.55336 .3633704 229.94 0.000 82.31045 84.79628 2 9 1 | 93.02938 .3586175 259.41 0.000 91.80272 94.25604 2 9 2 | 84.94324 .3921817 216.59 0.000 83.60178 86.28471 3 2 1 | 91.1313 .3290802 276.93 0.000 90.00568 92.25693 3 2 2 | 86.37037 .3812609 226.54 0.000 85.06626 87.67448 3 3 1 | 92.20235 .3206486 287.55 0.000 91.10556 93.29914 3 3 2 | 86.33675 .3817488 226.16 0.000 85.03097 87.64253 3 4 1 | 90.37259 .3590234 251.72 0.000 89.14454 91.60064 3 4 2 | 85.98799 .4418439 194.61 0.000 84.47665 87.49933

ALMQ data analysis Final Report

Appendix 1. Predicted mean lobster size for location, year, gender and interactions (cont.)

3 5 1 | 89.94791 .3117631 288.51 0.000 88.88151 91.0143 3 5 2 | 85.69285 .3995383 214.48 0.000 84.32622 87.05948 3 6 1 | 90.73632 .3528824 257.13 0.000 89.52928 91.94337 3 6 2 | 84.92322 .4616709 183.95 0.000 83.34406 86.50238 3 7 1 | 89.07404 .3148707 282.89 0.000 87.99702 90.15107 3 7 2 | 84.72777 .3945885 214.72 0.000 83.37807 86.07747 3 8 1 | 89.19652 .3762488 237.07 0.000 87.90955 90.48349 3 8 2 | 83.57824 .4118527 202.93 0.000 82.16949 84.98699 3 9 1 | 90.59898 .3800494 238.39 0.000 89.29902 91.89895 3 9 2 | 84.86657 .3998188 212.26 0.000 83.49898 86.23416 4 2 1 | 87.44576 .3001936 291.30 0.000 86.41894 88.47258 4 2 2 | 81.8697 .30163 271.42 0.000 80.83797 82.90144 4 3 1 | 87.41661 .2772207 315.33 0.000 86.46837 88.36485 4 3 2 | 84.61406 .2747277 307.99 0.000 83.67434 85.55377 4 4 1 | 87.13566 .2661308 327.42 0.000 86.22535 88.04596 4 4 2 | 84.43845 .2627198 321.40 0.000 83.53981 85.33709 4 5 1 | 85.54062 .3317744 257.83 0.000 84.40578 86.67546 4 5 2 | 83.4876 .3232877 258.25 0.000 82.38179 84.59342 4 6 1 | 87.42773 .3657113 239.06 0.000 86.17681 88.67866 4 6 2 | 82.90824 .4207244 197.06 0.000 81.46914 84.34734 4 7 1 | 87.70921 .3450438 254.20 0.000 86.52897 88.88944 4 7 2 | 83.47164 .3558142 234.59 0.000 82.25456 84.68871 4 8 1 | 87.07597 .3964915 219.62 0.000 85.71976 88.43218 4 8 2 | 83.13273 .3829767 217.07 0.000 81.82275 84.44271 4 9 1 | 89.83658 .4130858 217.48 0.000 88.42361 91.24955 4 9 2 | 83.75659 .4200716 199.39 0.000 82.31972 85.19346 5 2 1 | 97.23937 .3718804 261.48 0.000 95.96734 98.51139 5 2 2 | 90.44883 .3269101 276.68 0.000 89.33062 91.56703 5 3 1 | 87.60904 .3176498 275.80 0.000 86.52251 88.69557 5 3 2 | 86.77597 .2970355 292.14 0.000 85.75995 87.79199 5 4 1 | 87.50763 .2947593 296.88 0.000 86.4994 88.51587 5 4 2 | 86.39855 .2738213 315.53 0.000 85.46194 87.33516 5 5 1 | 89.06406 .6364291 139.94 0.000 86.88713 91.24098 5 5 2 | 88.44604 .5224384 169.29 0.000 86.65903 90.23306 5 6 1 | 90.16609 .4437509 203.19 0.000 88.64823 91.68395 5 6 2 | 86.3247 .4165347 207.24 0.000 84.89993 87.74946 5 7 1 | 88.90588 .4724089 188.20 0.000 87.29 90.52177 5 7 2 | 86.28687 .4795132 179.95 0.000 84.64668 87.92706 5 8 1 | 92.58723 .3479679 266.08 0.000 91.397 93.77747 5 8 2 | 88.07083 .3443242 255.78 0.000 86.89306 89.2486 5 9 1 | 90.48255 .4155856 217.72 0.000 89.06103 91.90407 5 9 2 | 85.97465 .4003815 214.73 0.000 84.60513 87.34416 ------

ALMQ data analysis Final Report

Appendix 2. Box Plots of lobster size (mm) by gender, for Jacquard’s Ridge, 2005 - 2012.

Jacquard’s Ridge

200

150

SIZE

100 50

Male Male Male Male Male Male Male Male Female Female Female Female Female Female Female Female 2005 2006 2007 2008 2009 2010 2011 2012 Graphs by location

Appendix 3. Box Plots of lobster size (mm) by gender, for Lobster Bay, 2005 - 2012.

Lobster Bay

200

150

100

SIZE 50

Male Male Male Male Male Male Male Male Female Female Female Female Female Female Female Female 2005 2006 2007 2008 2009 2010 2011 2012 Graphs by location

ALMQ data analysis Final Report

Appendix 4. Box Plots of lobster size (mm) by gender, for Port Latour Inside, 2005 - 2012.

Port Latour Inside

160

140

120

100

SIZE

80 60

Male Male Male Male Male Male Male Male Female Female Female Female Female Female Female Female 2005 2006 2007 2008 2009 2010 2011 2012 Graphs by location

Appendix 5. Box Plots of lobster size (mm) by gender, for Yarmouth Inside, 2005 - 2012.

Yarmouth Inside

200

150

SIZE

100 50

Male Male Male Male Male Male Male Male Female Female Female Female Female Female Female Female 2005 2006 2007 2008 2009 2010 2011 2012 Graphs by location

ALMQ data analysis Final Report

Appendix 6. Box Plots of lobster size (mm) by gender, for Yarmouth Outside, 2005 - 2012.

Yarmouth Outside

200

150

SIZE

100 50

Male Male Male Male Male Male Male Male Female Female Female Female Female Female Female Female 2005 2006 2007 2008 2009 2010 2011 2012 Graphs by location

Appendix 7. Jacquard’s Ridge overlaid plot of protein Brix (16+) proportion, 2005 - 2012.

Moult Timing - Jaquard's Ridge 2005 - 2012

Based on Protein Brix (16+)

prot_brix_molt_prop 0

25/Jul/2006 02/Jul/2008 20/Jul/2010 04/Jul/2011 03/Jul/2012 16/Aug/2005 29/Aug/2007 25/Jun/2009 date

ALMQ data analysis Final Report

Appendix 8. Lobster Bay overlaid plot of protein Brix (16+) proportion, 2005 - 2012.

Moult Timing - Lobster Bay 2005 - 2012

Based on Protein Brix (16+)

prot_brix_molt_prop

.1 0

04/Jul/2012 06/May/2005 09/May/2006 10/May/2007 19/Jun/200813/May/2009 25/May/2010 25/May/2011 date

Appendix 9 Port Latour Inside overlaid plot of protein Brix (16+) proportion, 2005 - 2012.

Moult Timing - Port Latour Inside 2005 - 2012

Based on Protein Brix (16+)

prot_brix_molt_prop 0

20/Jul/2005 07/Jul/2008 01/Jul/2009 14/Aug/200616/Jun/2007 18/May/2010 27/May/2011 21/Aug/2012 date

ALMQ data analysis Final Report

Appendix 10. Yarmouth Inside overlaid plot of protein Brix (16+) proportion, 2005 - 2012.

Moult Timing - Yarmouth Inside 2005 - 2012

Based on Protein Brix (16+)

prot_brix_molt_prop 0

10/Jul/2008 14/Jul/2009 25/Aug/2005 31/Aug/200608/Aug/2007 29/Jun/2010 27/Jun/2011 29/Jun/2012 date

Appendix 11. Yarmouth Outside overlaid plot of protein Brix (16+) proportion, 2005 - 2012.

Moult Timing - Yarmouth Outside 2005 - 2011

Based on Protein Brix (16+)

prot_brix_molt_prop

.1 0

25/Jul/2011 09/Sep/200503/Aug/2006 07/Aug/2007 21/Aug/200830/Jun/200902/Jun/2010 15/Jun/2012 date

ALMQ data analysis Final Report

Appendix 12. Jacquard’s Ridge overlaid plot of moult proportion by gender, for all years standardized by days since January 1st.

Moult Timing - Protein Brix Jacquard's Ridge 2005 - 2012

Male Female

prot_brix_molt_prop_sex 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by sex

Appendix 13. Lobster Bay overlaid plot of moult proportion by gender, for all years standardized by days since January 1st.

Moult Timing - Protein Brix Lobster Bay 2005 - 2012

Male Female

prot_brix_molt_prop_sex 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by sex

ALMQ data analysis Final Report

Appendix 14. Port Latour Inside overlaid plot of moult proportion by gender, for all years standardized by days since January 1st.

Moult Timing - Protein Brix Port Latour Inside 2005 - 2012

Male Female

prot_brix_molt_prop_sex 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by sex

Appendix 15. Yarmouth Inside overlaid plot of moult proportion by gender, for all years standardized by days since January 1st.

Moult Timing - Protein Brix Yarmouth Inside 2005 - 2012

Male Female

prot_brix_molt_prop_sex 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by sex

ALMQ data analysis Final Report

Appendix 16. Yarmouth Outside overlaid plot of moult proportion by gender, for all years standardized by days since January 1st.

Moult Timing - Protein Brix Yarmouth Outside 2005 - 2012

Male Female

prot_brix_molt_prop_sex 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by sex

Appendix 17. Jacquard’s Ridge overlaid plot of moult proportion by lobster size category (mm), st for all years, standardized by days since January 1 .

Moult Timing - Protein Brix Jacquard's Ridge 2005 - 2012

<82.5 >=82.5

prot_brix_molt_prop_size 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by size_ct

ALMQ data analysis Final Report

Appendix 18. Lobster Bay overlaid plot of moult proportion by lobster size category (mm), for st all years, standardized by days since January 1 .

Moult Timing - Protein Brix Lobster Bay 2005 - 2012

<82.5 >=82.5

prot_brix_molt_prop_size 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by size_ct

Appendix 19. Port Latour Inside overlaid plot of moult proportion by lobster size category st (mm), for all years, standardized by days since January 1 .

Moult Timing - Protein Brix Port Latour Inside 2005 - 2012

<82.5 >=82.5

prot_brix_molt_prop_size 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by size_ct

ALMQ data analysis Final Report

Appendix 20. Yarmouth Inside overlaid plot of moult proportion by lobster size category (mm), st for all years, standardized by days since January 1 .

Moult Timing - Protein Brix Yarmouth Inside 2005 - 2012

<82.5 >=82.5

prot_brix_molt_prop_size 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by size_ct

Appendix 21. Yarmouth Outside overlaid plot of moult proportion by lobster size category st (mm), for all years, standardized by days since January 1 .

Moult Timing - Protein Brix Yarmouth Outside 2005 - 2012

<82.5 >=82.5

prot_brix_molt_prop_size 0

0 100 200 300 400 0 100 200 300 400 day 2005 2006 2007 2008 2009 2010 2011 2012

Graphs by size_ct

ALMQ data analysis Final Report

Appendix 22. Yarmouth Outside overlaid plot of premoult stage (4+) proportion, 2005 – 2012.

Moult Timing - Jaquard's Ridge 2005 - 2012

Based on Moultstage (4+)

.25

.15

moltstage_molt_prop

.05 0

14/Jul/2007 02/Jul/2008 20/Jul/2010 20/Jul/2011 02/Aug/2005 06/Sep/2006 05/Aug/2009 01/Aug/2012 date

Appendix 23. Lobster Bay overlaid plot of premoult stage (4+) proportion, 2005 – 2012.

Moult Timing - Lobster Bay 2005 - 2012

Based on Moultstage (4+)

.15

.05

moltstage_molt_prop 0

29/Jul/2008 22/Jul/2009 17/Sep/200509/Aug/2006 28/Aug/2007 25/May/2010 03/Aug/2011 06/Sep/2012 date

ALMQ data analysis Final Report

Appendix 24. Port Latour Inside overlaid plot of premoult stage (4+) proportion, 2005 – 2012.

Moult Timing - Port Latour Inside 2005 - 2012

Based on Moultstage (4+)

.15

moltstage_molt_prop

.05 0

20/Jul/2005 10/Jul/2007 01/Jul/2009 29/Aug/2006 24/Jun/2008 30/Jun/201027/May/2011 07/Aug/2012 date

Appendix 25. Yarmouth Inside overlaid plot of premoult stage (4+) proportion, 2005 – 2012.

Moult Timing - Yarmouth Inside 2005 - 2012

Based on Moultstage (4+)

moltstage_molt_prop 0

19/Sep/2005 14/Sep/2006 06/Sep/200707/Aug/200819/Jun/2009 18/Jun/2010 01/Sep/2011 12/Oct/2012 date

ALMQ data analysis Final Report

Appendix 26. Yarmouth Outside overlaid plot of premoult stage (4+) proportion, 2005 – 2012.

Moult Timing - Yarmouth Outside 2005 - 2012

Based on Moultstage (4+)

moltstage_molt_prop 0

27/Jul/2009 25/Jul/2011 11/Jul/2012 09/Aug/2005 15/Sep/200605/Sep/200721/Aug/2008 30/Aug/2010 date

Appendix 27. Jacquard’s Ridge overlaid plot of premoult stage group (3) proportion, 2005 – 2012.

Moult Timing - Jaquard's Ridge 2005 - 2012

Based on Moultstage Group (3)

moltstage_group_molt_prop 0

25/Jul/2006 02/Jul/2008 03/Jul/2012 16/Sep/2005 14/Aug/2007 05/Aug/200922/Jun/2010 21/Jun/2011 date

ALMQ data analysis Final Report

Appendix 28. Lobster Bay overlaid plot of premoult stage group (3) proportion, 2005 – 2012.

Moult Timing - Lobster Bay 2005 - 2012

Based on Moultstage Group (3)

moltstage_group_molt_prop 0

06/Jul/2005 29/Jun/200622/May/2007 19/Jun/200820/Apr/2009 08/Apr/2010 10/May/2011 06/Sep/2012 date

Appendix 29. Port Latour Inside overlaid plot of premoult stage group (3) proportion, 2005 – 2012.

Moult Timing - Port Latour Inside 2005 - 2012

Based on Moultstage Group (3)

moltstage_group_molt_prop 0

08/Jul/2005 05/Jul/2006 25/Jul/2007 07/Jul/2008 14/May/2009 23/Apr/2010 27/May/2011 12/Jun/2012 date

ALMQ data analysis Final Report

Appendix 30. Yarmouth Inside overlaid plot of premoult stage group (3) proportion, 2005 – 2012.

Moult Timing - Yarmouth Inside 2005 - 2012

Based on Moultstage Group (3)

moltstage_group_molt_prop 0

12/Jul/2005 04/Jul/2006 08/Aug/2007 07/Aug/200829/Jun/200921/Apr/2010 04/May/2011 12/Oct/2012 date

Appendix 31. Yarmouth Outside overlaid plot of premoult stage group (3) proportion, 2005 – 2012.

Moult Timing - Yarmouth Outside 2005 - 2012

Based on Moultstage Group (3)

moltstage_group_molt_prop 0

11/Jul/2012 09/Sep/200503/Aug/2006 05/Sep/200721/Aug/200818/Jun/2009 02/Jun/2010 11/Aug/2011 date

ALMQ data analysis Final Report

Appendix 32. Jacquard’s Ridge overlaid plot of lowest hard-shell proportion, 2005 – 2012.

Moult Timing - Jaquard's Ridge 2005 - 2012

Based on Shell Hardness Proportion

molt_shell_group_prop 0

17/Jul/2012 16/Sep/200523/Aug/2006 21/Sep/200725/Aug/2008 16/Sep/2009 20/Oct/201023/Aug/2011 date

Appendix 33. Lobster Bay overlaid plot of lowest hard-shell proportion, 2005 – 2012.

Moult Timing - Lobster Bay 2005 - 2012

Based on Shell Hardness Proportion

molt_shell_group_prop 0

13/Jul/2007 29/Jul/2008 17/Aug/2005 07/Sep/2006 03/Sep/2009 28/Oct/201003/Aug/2011 01/Oct/2012 date

ALMQ data analysis Final Report

Appendix 34. Port Latour Inside overlaid plot of lowest hard-shell proportion, 2005 – 2012.

Moult Timing - Port Latour Inside 2005 - 2012

Based on Shell Hardness Proportion

molt_shell_group_prop 0

18/Jul/2006 14/Jul/2010 26/Jul/2011 02/Sep/2005 07/Aug/2007 05/Aug/2008 12/Aug/2009 21/Aug/2012 date

Appendix 35. Yarmouth Inside overlaid plot of lowest hard-shell proportion, 2005 – 2012.

Moult Timing - Yarmouth Inside 2005 - 2012

Based on Shell Hardness Proportion

molt_shell_group_prop 0

28/Jul/2009 27/Jul/2011 08/Sep/200517/Aug/2006 20/Sep/2007 02/Oct/2008 10/Aug/2010 28/Sep/2012 date

ALMQ data analysis Final Report

Appendix 36. Yarmouth Outside overlaid plot of lowest hard-shell proportion, 2005 – 2012.

Moult Timing - Yarmouth Outside 2005 - 2012

Based on Shell Hardness Proportion

molt_shell_group_prop 0

30/May/2005 27/Sep/2006 03/Oct/2007 28/Oct/200810/Aug/2009 30/Aug/2010 28/Sep/2011 27/Sep/2012 date