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Bibliography of the Maurice Lamontagne Institute

Bernard SAINTE-MARIE

C. GUAY, B. SAINTE-MARIE, J.-C. BRÊTHES, 2011. Strong maternal effects and extreme heterogeneity of progeny development in the caridean shrimp Sclerocrangon boreas (Crangonidae). Mar. Biol., 158(12): 2835-2845 .

Sclerocrangon boreas is uncommon among marine coastal carideans in having a non-dispersing, abbreviated (2-stage) larval phase. We investigated the implications of this strategy in terms of fecundity, oVspring provisioning and brood care in S. boreas from the St. Lawrence Gulf and Estuary in 2009–2010. Fecundity scaled positively to female body size but was low due to the production of large, lipid-rich eggs. Ovspring size at all stages of development was positively related to female size. Larval traits and lipid dynamics indicate obligatory lecithotrophic development from hatching to juvenile. The larva becomes a juvenile on the mother and remains associated with her for sometime after. The co-occurrence of early egg stages among many juveniles in some clutches raises the possibility that maternal care of juveniles includes the provisioning of trophic eggs or eggs reclaimed from other females.©2011 Springer

SAINTE-MARIE, B., S. DUBÉ, 2011. Processus consultatif scientifique régional sur l’évaluation du stock de flétan atlantique du golfe du Saint-Laurent (4RST), 16 février 2011, Institut Maurice-Lamontagne ; Regional Science Advisory Process on the Assessment of the Gulf of St.Lawrence (4RST) Atlantic Halibut Stock, February 16, 2011, Maurice Lamontagne Institute. MPO, Secrétariat canadien de consultation scientifique, Série des comptes rendus; DFO, Canadian Science Advisory Secretariat, Proceedings Series, 2011/008, 19 p .

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SAINTE-MARIE, B., 2010. The First 30 Years of the Journal of Crustacean Biology – A Bibliometric Study. J. Crust. Biol., 30(4): 541-549 .

This review examines some of the characteristics and highlights some notable articles of the Journal of Crustacean Biology (JCB) in its first 30 years of existence. A total of 2052 articles appeared in JCB from the first issue in February 1981 to the end of 2009. The number of articles by volume increased from 50 in 1981 to 93 in 2002 and then declined to around 70. From 1981 to 2009, article size varied around a mean of 11 pages (pre-2005 format) but mean number of authors and references by article increased by a factor of 1.87 and 2.20, respectively. JCB content is predominated numerically by taxonomy and systematics (36% of all articles), but other research areas (anatomy, physiology, development, growth-reproduction, life history, behavior, ecology, conservation) were also represented from the outset. JCB's 2-year impact factor increased significantly from 1991 to 2009. Longer-term impact of JCB is evident in the fact that almost half of all JCB articles were cited in 2009 and that the mean age of those cited articles was only slightly less than the mean age of all JCB articles (12.6 vs 13.3 years). However, citations to JCB differ widely across research areas, with articles in taxonomy cited on average at less than half the rate of articles in the areas of ecology or conservation. The most cited JCB articles by combination of research area and decade of publication deal primarily with higher crustaceans (malacostracans) and are reviews or original research articles with cross-disciplinary appeal

ÉMOND, K., B. SAINTE-MARIE, L. GENDRON, 2010. Relative growth, life-history phases, and sexual maturity of American lobster (Homarus americanus). Can. J. Zool., 88: 347-358 .

Previous studies of relative growth in crustaceans have focused primarily on body parts representing sexual characters for the purpose of determining size at onset of sexual maturity. We have revisited the relative growth of abdomen and crusher claw in American lobster (Homarus americanus H. Milne Edwards, 1837) in a general life-history perspective using a broad spectrum of lobster sizes (6–160 mm cephalothorax length (CL)). Growth phases were recognized by inflections in scatterplots of a body-part measurement against CL. The abdomen is characterized by three growth phases in the male and female, whereas the crusher claw has at least three growth phases in the male and at least two in the female. Additionally, we explored relative growth of gonopod and vas deferens for males of 35–150 mm CL. Both organs exhibit a synchronous change from strong to weak positive growth allometry. The growth phases can be associated with major life-history events including the transition from a cryptic to an overt lifestyle and the onset of physiological, functional, and morphometric maturity. The onset of morphometric maturation inferred from relative growth of abdomen and crusher claw precedes functional maturity in females and follows it in males©2010 National Council Research Canada

BURMEISTER, A., B. SAINTE-MARIE, 2010. Pattern and causes of a temperature-dependent gradient of size at terminal moult in Snow Crab (Chionoecetes opilio) along West Greenland. Polar Biol., 33(6): 775-788 .

Geographic variation in size of male and female snow crabs (Chionoecetes opilio) was investigated along the west coast of Greenland to test the hypothesis that size at terminal molt (=adulthood) is temperature dependent. A total of 81,490 snow crabs were collected in small-mesh traps in Disko Bay (68–69°N) and six sites near Sisimiut (66–67°N) in May and June from 2000 to 2005. Average bottom temperature over the study period ranged from -0.8 to 3.2 deg;C across the sampling sites. Mean carapace width was positively correlated with temperature in both sexes, a pattern often described as a converse Jame’s cline. We infer that temperature per se is the causative factor and discount season length, food availability or density as ultimate causes of the cline. Temperature effects on body size of crabs apparently result from a change in the number of instars before terminal molt. This interpretation is supported by size frequency analysis showing that in general crabs were larger at instar in a colder than in a warmer site. We briefly discuss the implications of our findings for population reproductive potential and the effectiveness of a fixed legal size limit in protecting some adult males from exploitation at different temperature regimes.©2010 Springer-Verlag

DUTIL, J.-D., R. LAROCQUE, S. VALOIS, E. MAYRAND, B. SAINTE-MARIE, 2009. Spatial, annual and seasonal patterns in the condition and muscle size of snow crab. Mar. Biol., 156(9): 1903-1916 .

The extent of spatial (depth and locality) and temporal (season and year) variabilities in condition and relative muscle size (a direct proxy of growth) were examined in male and female adult and non-adult snow crabs Chionoecetes opilio. Condition, determined from the relative size of the digestive gland and moisture content of the muscle and digestive gland, and muscle size, determined as the ratio of merus muscle mass over merus volume, separated as diVerent processes in a principal component analysis. Snow crabs showed a wide range of condition and muscle size values. Overall, the condition was better in non-adult than in adult crabs, with adult females being in worst condition, and muscle size was larger in males than in females. Condition variability was greater for seasonal compared to annual samples, probably reXecting annual molt cycles. In contrast, the muscle size variability was greater for annual compared to seasonal samples, possibly as a result of changing crab abundance and competition intensity during recruitment pulses. Condition and muscle size increased through summer in males and immature females, although to diVerent extents depending on instar, but did not change in adult females. Both condition and muscle size were highly variable at the investigated spatial scales. Condition and muscle size had a signiWcant eVect on gonad size, once the eVect of crab size was removed, suggesting a direct link between these two parameters and reproductive capability.©2009 Springer

LACOURSIÈRE-ROUSSEL, A., B. SAINTE-MARIE, 2009. Sexual system and female spawning frequency in the sculptured shrimp Sclerocrangon boreas (decapoda : Caridea : Crangonidae). J. Crust. Biol., 29(2): 192-200 .

The nature of the sexual system (protandry, gonochory) and the frequency of female spawning (semelparous, iteroparous annual, iteroparous biennial) in the crangonid Sclerocrangon boreas remain uncertain. We addressed these questions by examining population sex ratio, anatomy and histology, and gonad and oö cyte sizes. Gonochory is supported by several facts: sex ratio was balanced at the smallest shrimp sizes, no intermediate sex form was found, and there was no evidence of degeneration of male gonad at the sizes where this might have been expected to occur. A majority of larger females, if not all, had a structure linking the ovary to the base of the fifth pereiopod but the presence of this structure on small females suggests it is not a degenerate vas deferens. Iteroparity is supported by the presence of two generations of oöcytes in females, including those gravid, and female biennial spawning is supported by the small size of ovaries/oöcytes in gravid females.©2009 The Crustacean Society

TURCOTTE, C., B. SAINTE-MARIE, 2009. Biological synopsis of the Japanese Skeleton Shrimp (Caprella mutica). Can. Manuscr. Rep. Fish. Aquat. Sci., 2903, 33 p .

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Caprella mutica is an amphipod crustacean that was first described in 1935 from the Siberian shores of the Japan Sea. In its indigenous habitat, C. mutica is mainly found in stands of eelgrass or macroalgae in the infralittoral zone where temperature varies from –1.8 °C in winter to 25  °C in summer and salinity from 11 to 35. C. mutica has a high reproductive potential: there are two generations per year, females mature at a small size, are relatively fecund and reproduce more than once, and "direct" development of progeny ensures a good survival rate. C. mutica was introduced to North America first on the Pacific side in the early 1970s and on the Atlantic side in the late 1990s, as well as in Europe during the mid 1990s. The sites where C. mutica was introduced in the northern hemisphere are at temperate, boreal and subarctic latitudes between 35 and 70 °N. Transoceanic introductions probably occurred by way of transport of aquaculture organisms and ballast waters. Secondary dispersal of the species along coasts may be due to commercial and recreational navigation, drifting on floatsam, and at a smaller spatial scale by swimming and creeping. In its new habitat in the northern hemisphere, C. mutica is found mainly or only on artificial structures such as nets, ropes and buoys that are used in aquaculture. Good environmental tolerance, rapid growth and a high reproductive rate, combined with a flexible diet, an aggressive nature and a high degree of commensalism with human activities, seem to contribute to make C. mutica a good invader. C. mutica apparently has a negative impact on some aquaculture activities but its effect on natural ecosystems remains unknown.

TURCOTTE, C., B. SAINTE-MARIE, 2009. Synthèse de la biologie de la caprelle japonaise (Caprella mutica). Rapp. manus. can. sci. halieut. aquat., 2903, 35 p .

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Caprella mutica is an amphipod crustacean that was first described in 1935 from the Siberian shores of the Japan Sea. In its indigenous habitat, C. mutica is mainly found in stands of eelgrass or macroalgae in the infralittoral zone where temperature varies from –1.8 °C in winter to 25  °C in summer and salinity from 11 to 35. C. mutica has a high reproductive potential: there are two generations per year, females mature at a small size, are relatively fecund and reproduce more than once, and "direct" development of progeny ensures a good survival rate. C. mutica was introduced to North America first on the Pacific side in the early 1970s and on the Atlantic side in the late 1990s, as well as in Europe during the mid 1990s. The sites where C. mutica was introduced in the northern hemisphere are at temperate, boreal and subarctic latitudes between 35 and 70 °N. Transoceanic introductions probably occurred by way of transport of aquaculture organisms and ballast waters. Secondary dispersal of the species along coasts may be due to commercial and recreational navigation, drifting on floatsam, and at a smaller spatial scale by swimming and creeping. In its new habitat in the northern hemisphere, C. mutica is found mainly or only on artificial structures such as nets, ropes and buoys that are used in aquaculture. Good environmental tolerance, rapid growth and a high reproductive rate, combined with a flexible diet, an aggressive nature and a high degree of commensalism with human activities, seem to contribute to make C. mutica a good invader. C. mutica apparently has a negative impact on some aquaculture activities but its effect on natural ecosystems remains unknown.

RADULOVICI, A.E., B. SAINTE-MARIE, F. DUFRESNE, 2009. DNA barcoding of marine crustaceans from the Estuary and Gulf of St Lawrence : a regional-scale approach. Mol. Ecol. Resour., 9(Suppl.1): 181-187 .

Marine crustaceans are known as a group with a high level of morphological and ecological diversity but are difficult to identify by traditional approaches and usually require the help of highly trained taxonomists. A faster identification method, DNA barcoding, was found to be an effective tool for species identification in many metazoan groups including some crustaceans. Here we expand the DNA barcode database with a case study involving 80 malacostracan species from the Estuary and Gulf of St Lawrence. DNA sequences for 460 specimens grouped into clusters corresponding to known morphological species in 95 % of cases. Genetic distances between species were on average 25 times higher than within species. Intraspecific divergence was high (3.78–13.6 %) in specimens belonging to four morphological species, suggesting the occurrence of cryptic species. Moreover, we detected the presence of an invasive amphipod species in the St Lawrence Estuary. This study reconfirms the usefulness.©2009 Blackwell Publishing Ltd

BUBLITZ, R., B. SAINTE-MARIE, C. NEWCOMB-HODGETTS, N. FLETCHER, M. SMITH, J. HARDEGE, 2008. Interspecific activity of the sex pheromone of the European shore crab (Carcinus maenas). Behaviour, 145(10): 1465-1478 .

The recent identification of uridine diphosphate (UDP) as the female sex-pheromone in the European shore crab Carcinus maenas demonstrated not only the link between moult and pheromone production, but also how it may have evolved from a `simple' metabolic byproduct. Consequently, it is expected to be present in other moulting crustaceans, thus raising issues involving species specificity of the female pheromone. Bioassays were conducted using synthetic pheromone (UDP, 10-3-10-4 M) to examine if it induced sexual behaviour in other crustacean species that are neither closely related nor occur in the same ecosystem. The snow crab, Chionoecetes opilio, and the yellowline arrow crab, Stenorhynchus seticornis, both belonging to a different superfamily (Majoidea) and occurring in different habitats than C. maneas (Portunoidea), displayed significant sexual behaviour towards UDP treated objects (p < 0.005). These and other examples demonstrate that the female sex-pheromone UDP is not species-specific but is present and active in some other decapod crustaceans.©2008 Brill

PUEBLA, O., J.-M. SÉVIGNY, B. SAINTE-MARIE, J.-C. BRÊTHES, A. BURMEISTER, E.G. DAWE, M. MORIYASU, 2008. Population genetic structure of the snow crab (Chionoecetes opilio) at the Northwest Atlantic scale. Can. J. Fish. Aquat. Sci., 65: 425-436 .

Marine species with planktonic larval durations of several months (teleplanic larvae) can potentially maintain demographic connectivity across large geographical distances. This perspective has important fundamental and applied implications, notably for the understanding of evolutionary and ecological processes in the marine realm, the implementation of marine protected areas, and fisheries management. Here we present, at the scale of the Northwest Atlantic, a spatial analysis of snow crab (Chionoecetes opilio, Majoidea) population genetic structure, a species that has a planktonic larval phase of 3 to 5 months. Eight microsatellite markers analysed on 847 C. opilio samples from 13 locations revealed an absence of significant genetic structure along the west coast of Greenland and within Atlantic Canada from southern Labrador to Nova Scotia. These results are consistent with a scenario of extensive demographic connectivity among C. opilio populations and have implications for the management of this species, which supports one of the most important Canadian and Greenlandic fisheries in terms of economic value. A genetic break is nevertheless identified between Greenland and Atlantic Canada, showing that genetic structure can develop within seas (the Labrador Sea in this case) despite the occurrence of very long planktonic larval stages.©2008 NRC Canada

CHABOT, D., B. SAINTE-MARIE, K. BRIAND, J.M. HANSON, 2008. Atlantic cod and snow crab predator-prey size relationship in the Gulf of St. Lawrence, Canada. Mar. Ecol. Prog. Ser., 363: 227-240 .

Atlantic cod Gadus morhua stomach contents (n = 30 973, including 28 377 non-empty stomachs) and morphometric measurements on live snow crab Chionoecetes opilio and cod were examined to assess the predator-prey relationship between these 2 species. The most common snow crab instars found in cod stomachs were III and V (˜6 to 8 and ˜12 to 16 mm carapace width [CW], respectively) in the northern Gulf of St. Lawrence (GSL) and VI and VII (˜17 to 23 and ˜23 to 31 mm CW, respectively) in the southern GSL. A significant positive relationship was found between cod length and the largest and smallest CW of snow crab ingested by cod. Positive relationships were also found between gape width and body length in cod and between 3 measures of size (maximum span, width at rest, length at rest) and CW in snow crab. Snow crab length at rest was closely related to cod gape width, suggesting that the largest snow crab ingested by cod must be attacked from the side. There appears to be a plateau at 65.1 mm in the relationship between maximum snow crab CW and cod length, caused by the absence of large (adolescent and adult) male snow crab in cod stomachs. Other studies have found recently moulted, soft-shell snow crabs in cod stomachs, but this appears to be rare. Thus, snow crabs are susceptible to predation by cod mostly for the first 4 yr of postsettlement in the GSL. Any effect of cod predation on the snowcrab fishery would be felt 6 to 11 yr later, given growth models established for the GSL.©2008 Inter-Research

SAINTE-MARIE, B., T. GOSSELIN, T., J.-M. SÉVIGNY, N. URBANI, 2008. The snow crab mating system: opportunity for natural and unnatural selection in a changing environment. Bull. Mar. Sci., 83(1): 131-161 .

The impact of fishing as a driver of sexual selection is not well understood in crustaceans. Fishing must be viewed as acting in conjunction with, or in opposition to, natural factors, which also modify the context for sexual competition, mate choice, and sexual conflict. We review knowledge of the polygynandrous mating system of the snow crab and evaluate the likely interplay between natural and fishing forces in the process of sexual selection. The snow crab has determinate growth and two female reproductive stages (primiparous and multiparous) with discrete and disjunct mating seasons. Temperature shifts the spectrum of size at maturity in both sexes and determines female reproductive tempo, thereby altering sperm supply, egg production, and operational sex ratio. Population dynamics modulates the phenotype of receptive individuals and the direction and intensity of sexual competition over time. Fishing directed only at large males may attenuate or exacerbate some aspects of sexual conflict at primiparous mating, depending on the natural context, but otherwise it consistently promotes mating of less fecund males, reduces opportunity for female mate choice, and increases the likelihood of sperm limitation. These changes have mixed but still incompletely appreciated effects on female reproductive fitness. The long-term potential for selection against large size at maturity remains uncertain. ©2008 Rosenstiel School of Marine and Atmospheric Science of the University of Miami

SAINTE-MARIE, B., 2008. Évaluation de l’oursin vert de la Côte-Nord de l'estuaire du Saint-Laurent en 2008. MPO, Secrétariat canadien de consultation scientifique, Avis scientifique, 2008/048, 14 p .

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SAINTE-MARIE, B., 2008. Assessment of green sea urchin of the North Shore of the St. Lawrence Estuary in 2008. DFO, Canadian Science Advisory Secretariat, Science Advisory Report, 2008/048, 13 p .

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FONSECA, D.B., B. SAINTE-MARIE, F. HAZEL, 2008. Longevity and change in shell condition of adult male snow crab >i>Chionoecetes opilio inferred from dactyl wear and mark–recapture data. Trans. Am. Fish. Soc., 137(4): 1029-1043 .

Postmolt longevity and changes in the shell condition and body integrity of male snow crab Chionoecetes opilio after their terminal molt were assessed through a mark-recapture experiment and population censuses in a commercially unfished locality of eastern Canada. The experiment explored the value of dactyl wear as a quantitative measure and shell condition (SC; measured on a five-stage scale) as a relative index of shell age. Males were recaptured up to 6 years after release. Much of the extensive variation in observed dactyl wear was explained by time at liberty (Dt) and male size, and the extent of change in SC was positively correlated with Dt. The conservative wear-based estimate of male longevity was 7.7 years, a value 1-3 years greater than previously estimated. Dactyl wear and recapture data confirmed that SC is a relative, albeit rough, index of shell age. Shell hardness was positively correlated with male size and peaked in stage 3 about 3.5 years after the terminal molt. The number of missing pereopods increased with shell age and SC stage and overall was negatively correlated with male size. The commercial value of adult males may be highest at 1-4.5 years post-terminal molt and the reproductive value at 2-5.5 years.©2008 American Fisheries Society

CHABOT, D., A. RONDEAU, B. SAINTE-MARIE, L. SAVARD, T. SURETTE, P. ARCHAMBAULT, 2007. Distribution des invertébrés benthiques dans l'estuaire et le golfe du Saint-Laurent ; Distribution of benthic invertebrates in the Estuary and Gulf of St. Lawrence. MPO, Secrétariat canadien de consultation scientifique, Document de recherche ; DFO, Canadian Science Advisory Secretariat, Research Document, 2007/018, 108 p .

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This study had two objectives. The first was to gather all the available data from scientific surveys both at the Maurice-Lamontagne Institute (MLI) and the Gulf Fisheries Centre (GFC) in order to establish distribution patterns and relative abundance of benthic invertebrates in the lower estuary and the gulf of St. Lawrence (LEGSL). The second objective was to propose Ecologically and Biologically Significant Areas (EBSAs) for benthic invertebrates of the LEGSL, according to these distribution and relative abundance data. Surveys from the MLI are mainly conducted in the lower estuary and in the northern gulf of St. Lawrence (nGSL) but some cover Gaspesie, including Chaleurs bay, and the Magdalen Islands. The southern gulf of St. Lawrence (sGSL) is covered by the GFC surveys. The bulk of the information presented here comes from annual scientific surveys carried out by the two regions: the multi-species survey and the mobile gear Sentinel survey by the MLI and the fall survey, snow crab survey, and Northumberland survey by the GFC. Several other surveys by MLI provided useful data, some even with a long time series but with limited geographical coverage (surveys for snow crab, scallop and surf clam), while others were conducted less frequently and at a small geographic scale (surveys for clam and whelk). Cod and Greenland halibut stomach contents have also been used as a sampling device for the distribution on some benthic invertebrates. Despite the large number of surveys considered here, the coastal zone (less than 50 m deep in the Estuary and the nGSL and less than 30 m deep in the sGSL) was not adequately sampled, except for the Northumberland Strait. In the main section of this document, distributions of 44 taxa are presented and have guided the identification of EBSAs : 4 general groups (soft corals, anemones, sponges, ascidians), 5 echinoderms, 6 molluscs, 1 mysid, 22 shrimps, and 6 crabs. Zones of maximum relative abundance of each taxa, weighted inversely to their surface area of high abundance, were used to calculate an index of benthic invertebrate concentration for each 10 x 10 km square sampled in the study area. This index was the primary tool in the identification of potential EBSAs. As a result, 17 EBSAs for benthic invertebrates are proposed. However, it is important to keep in mind that only a small proportion (approximately 0.02 %) of the benthic invertebrate species known to be present in the study area was considered in the process. In particular, the lack of data for the coastal zone is a major gap. We present in appendix to this report the data on 6 coastal species that we were able to obtain.

KRUSE, G.H, A.V. TYLER, B. SAINTE-MARIE, D. PENGILLY, 2007. A workshop on mechanisms affecting year-class strength formation in snow crabs (Chionoecetes opilio) in the eastern Bering Sea. Alaska Fish. Res. Bull., 12(2): 277-290 .

A group of specialists on subarctic crab biology and fisheries participated in a workshop to discuss stage-specific recruitment processes of snow crab Chionoecetes opilio populations. The goal was to develop from experience with North Pacific and North Atlantic stocks a comprehensive set of hypotheses on the physical and biotic factors that may contribute to the variation in year-class success of the snow crab stock in the eastern Bering Sea. Participants identified 15 life history stages and associated survival and productivity processes. Some of the processes deemed to be most critical include match of hatching larvae with adequate densities of suitable prey, advection (or retention) of larvae to nursery areas suitable for settlement, predation by Pacific cod Gadus macrocephalus and other predators, and cannibalism of newly settled juveniles by older, larger snow crab juveniles. A comprehensive set of hypotheses is intended to serve as a vehicle by which to direct future field and laboratory research programs to better understand snow crab population and fishery dynamics.©2007 Alaska Department of Fish and Game

SAINTE-MARIE, B., 2007. Sperm demand and allocation in decapod crustaceans. Pages 191-210 in J.E. Duffy & M. Thiel (ed.). Evolutionary ecology of social and sexual systems : crustaceans as model organisms .

This chapter examines trends in male gametic strategies in relation to female sperm demand among gonochoristic decapod crustaceans. Female lifetime fecundity and number of eggs per spawn can vary much more among decapod species than in other taxa in which gametic strategies have been explored. Female anatomy and life history determine a gradient of complexity and potential duration for sperm storage, lasting from a few hours to several years. The spermathecae of some decapods have enormous capacity and filling may be antagonistic to ovary development. Sperm requirements associated with high female fecundity can be met by large sperm investments from individual males, promiscuity, and/or very effective use of sperm. As in many other taxa, male decapods may allocate sperm strategically as a function of female size (fecundity) and mated status, number of mating opportunities, and risk and intensity of sexual (sperm) competition.©2007 Oxford University Press, Inc.

GOSSELIN, T., B. SAINTE-MARIE, J.-M. SÉVIGNY, 2007. Individual identification of decapod crustaceans II : natural and genetic markers in snow crab (Chionoecetes opilio). J. Crust. Biol., 27(3) : 399-403 .

Methods for the identification of individual crustaceans are needed in many types of studies. Snow crab (Chionoecetes opilio) individuals have distinctive natural patterns of tubercles and spines on the carapace. The results of a double-marking experiment using these natural markers along with genetic (microsatellite) markers confirm that natural markings are a reliable means of recognizing individuals within groups of tens to hundreds of snow crabs. These natural markings are persistent through at least two molts. They have already demonstrated their usefulness in laboratory studies of molting and mating and could be applied to a wider spectrum of investigations. A cursory examination suggests that similar carapace features could be used to identify individuals in other crustacean species as well.©2007 The Crustacean Society

MACDIARMID, A.B., B. SAINTE-MARIE, 2006. Reproduction. Pages 45-77 in B.F. Phillips (ed.). Lobsters : biology, management, aquaculture and fisheries. Blackwell Publishing .

GENDRON, L., B. SAINTE-MARIE, 2006. Growth of juvenile lobster Homarus americanus off the Magdalen Islands (Quebec, Canada) and projection of instar and age at commercial size. Mar. Ecol. Prog. Ser., 326: 221-233 .

Juvenile American lobster Homarus americanus were quantitatively surveyed in the southeast Baie de Plaisance, Magdalen Islands, from 1995 to 2004. Lobster were collected by SCUBA, in summer and/or early fall after the annual settlement period. Growth for up to 3 yr following settlement was assessed by modal analysis of carapace length (CL)-frequency distributions and was confirmed for the first year by in situ rearing of Stage IV lobster collected from the plankton. In every sampling year, some modes (attributed to instars) were apparent in CL-frequency distributions up to 50 mm CL (estimated to be Instar XVI on average). The mean percent molt increment declined gradually from about 23% at Instar IV to about 15% at Instar XV. Strong year-classes could be tracked reasonably well for up to 2-3 yr after settlement. From this point, we projected growth to fishery recruitment and suggest that lobster reach commercial size at a higher instar and older age than previously believed. This is the first study to explain lobster juvenile growth in such detail, and it constitutes an essential step toward the understanding of potential settler-to-recruit relationships.© 2006 Inter-Research.

GASCON, D., B. SAINTE-MARIE, 2006. Examination of industry trawl survey of April 2006 in relation to the assessment of the snow crab in CFA 16. DFO, Canadian Science Advisory Secretariat, Science Advisory Report, 2006/026, 6 p .

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SAINTE-MARIE, B., I. BÉRUBÉ, S. BRILLON, F. HAZEL, 2006. Observations on the growth of the sculptured shrimp Sclerocrangon boreas (Decapoda : caridea). J. Crust. Biol., 26(1) : 55-62 .

Little is known of the growth of the sculptured shrimp Sclerocrangon boreas, a remarkable member of arctic and subarctic marine shelf communities. We determined the length-weight relationship, abdomen allometry, and size structure of shrimp in a population of the Gulf of Saint Lawrence, eastern Canada. We also reared shrimp for up to 3.5 years to measure their growth. The presence of very small immature females in the wild population indicates that the sculptured shrimp is not obligatorily a protandric hermaphrodite, if at all. Females grow faster, reach a greater size, and live longer than males. Males may be ≥4 years old at 17 mm cephalothorax length (CL) and females ≥6 years old at 29 mm CL. Ovigerous females have a broader abdomen with longer pleopod setae than similarly-sized immature females. After releasing their progeny, some females may molt and grow in length but revert to a condition of narrow abdomen and short pleopod setae, and then molt again to a condition of broad abdomen with long pleopod setae. This finding and demographic data suggest that some females are alternate-year spawners. Other females did not molt for ≥2 years and may spawn in successive years.©2007 The Crustacean Society

GASCON, D., B. SAINTE-MARIE, 2006. Examen du relevé au chalut de l'industrie d'avril 2006 dans le contexte de l'évaluation du crabe des neiges de la zone 16. MPO, Secrétariat canadien de consultation scientifique, Avis scientifique, 2006/026, 6 p .

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GOSSELIN, T., B. SAINTE-MARIE, L. BERNATCHEZ, 2005. Geographic variation of multiple paternity in the American lobster, Homarus americanus. Mol. Ecol., 14(5): 1517-1525 .

We studied the frequency of multiple paternity for American lobster (Homarus americanus) at three Canadian sites differing in exploitation rate and mean adult size. The probability of detecting multiple paternity using four microsatellite loci and 100 eggs per female was in excess of 99 % under various scenarios of paternal contribution. Overall, 13 % of the 108 examined females carried a clutch sired by two or three males. Multiple paternity was observed at the two most exploited sites (11 % at Magdalen Islands and 28 % at Grand Manan Island), whereas single paternity only was observed at the least exploited site (Anticosti Island). Within populations females with a clutch sired by more than one male tended to be smaller than females with a clutch sired by a single male. Based on these and other findings, we postulate a link between female promiscuity and sperm limitation in the American lobster.©2005 Blackwell Publishing Ltd

DULUC, C., B. SAINTE-MARIE, J.-C. BRÊTHES, 2005. Value of indicators for assessing recent mating in brachyuran crabs. J. Mar. Biol. Assoc. U.K., 85(1): 129-135 .

Methods for assessing the occurrence and recency of mating are important for the management and conservation of exploited brachyuran crabs. Using multiparous females of the snow crab Chionoecetes opilio, we evaluated by experiment the efficacy of three indicators of recent mating: a white deposit in the spermathecae, extended mate-guarding, and fresh grasping marks on the female pereiopods. This was done by contrasting sperm counts between the left and the right spermatheca of females that were exposed to males with the right gonopod ablated, at treatment sex ratios of 20♀:3♂ or 50♀:3♂. We expected that sperm reserves would be balanced between the two spermathecae of non-mated females and larger in the right than in the left spermatheca of mated females. Although no mating indicator was infallible, the presence of a white deposit was the most accurate because it maximized the median difference of sperm counts between the two spermathecae for the group of presumably mated females and minimized it for the group of presumably non-mated females. The use of grasping marks overestimated the mating frequency and resulted in the misclassification of a large proportion of females. Extended mate-guarding was a slightly better mating indicator than grasping marks, but it is not practical for field studies. Classification errors associated with each indicator can be explained mostly by female and male behaviours and may vary in magnitude with sociosexual context.©2005 Marine Biological Association of the United Kingdom

SAINTE-MARIE, B., R. DUFOUR, L. BOURASSA, D. CHABOT, M. DIONNE, D. GILBERT, A. RONDEAU, J.-M. SÉVIGNY, 2005. Critères et proposition pour une définition des unités de production du crabe des neiges (Chionoecetes opilio) dans l'estuaire et le nord du golfe du Saint-Laurent ; Criteria and proposition for the definition of snow crab (Chionoecetes opilio) production units in the estuary and northern Gulf of St. Lawrence. MPO, Secrétariat canadien de consultation scientifique, Document de recherche ; DFO, Canadian Science Advisory Secretariat, Research Document, 2005/059, 20 p .

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The current borders of snow crab management areas in the Department of Fisheries and Oceans (DFO) Québec region are not based on biological or oceanographical criteria and some times enclose territories with very different characteristics. As noted by the FRCC (2005), there is a need to define biological production units to better monitor the status and ensure conservation of populations. Toward that goal, we conducted a literature review and used trawl data from the Gulf of Saint Lawrence to identify the biological and physical factors that constrain the distribution and dispersal of snow crab. This information allows us to characterize snow crab habitat, which can be defined as a territory with bottoms of soft sediments, bathed by waters with a salinity > 26 ‰, part of which is at a temperature of 0-2 °C and the remainder at a temperature varying from about -1.5 to 4 °C. The surface waters above this territory must generally have a salinity > 26 ‰ and warm up to 5-15 °C for several weeks for larvae to survive and grow. Snow crab larvae can disperse themselves over long distances and this may explain the weak genetic differentiation of populations within the Gulf. However, the various benthic stages generally have a much smaller dispersal capability. We describe the criteria used to define biological production units and on that basis we propose to divide the territory under the responsibility of DFO-Québec into seven units.

SIDDEEK, M.S.M., B. SAINTE-MARIE, J. BOUTILLIER, G. BISHOP, 2004. Comparison of reference points estimated using a size-based method for two high-latitude crab species in the United States and Canada. Can. J. Fish. Aquat. Sci., 61: 1404-1430 .

We briefly reviewed the decision rules currently used for managing two major high-latitude crab stocks, snow crab (Chionoecetes opilio) and Dungeness crab (Cancer magister), in the United States and Canada and compared them with model-based reference points, harvest rate, and biomass proportion relative to virgin biomass, developed using species- and area-specific parameters. The model followed a size-based approach, which incorporated Beverton-Holt and Ricker stock-recruitment models and estimated mean and median reference points. The recruitment was also perturbed to generate distributions of reference points. The Beverton-Holt stock-recruitment model provided a lower harvest rate than that of the Ricker model. Harvest rates were lower for combined sexes spawning biomass than for female-only spawning biomass. Increasing the minimum size at first capture and decreasing the handling mortality resulted in increased harvest rates. Changes in fishery duration and timing of fishery open date did not change the harvest rate appreciably. The harvest rates for the Canadian snow and Dungeness crabs were mostly higher than those estimated for the Bering Sea and Southeast Alaska stocks. Reliable estimates of a number of life history parameters are lacking for both species, and hence, the results of this exercise need to be treated in a precautionary manner.©2004 NRC Cananda

SMITH, S.J., B. SAINTE-MARIE, 2004. Biological reference points for invertebrate fisheries: introduction. Can. J. Fish. Aquat. Sci., 61: 1303-1306 .

In the context of applying the precautionary approach to fisheries management, much of the current attention on developing reference points has been focussed on finfish species. However, at present the largest marine fisheries in Canada in terms of landings and value are those for invertebrate species. A workshop on reference points for invertebrate fisheries was held in Halifax, Nova Scotia, Canada, from 2 to 5 December 2002 to address the particular needs of fisheries on this diverse group of species. Seven of the papers presented at that workshop were published as a group in this issue of the Canadian Journal of Fisheries and Aquatic Science. Reproduction and recruitment were identified by all of these papers as key elements of population resilience and productivity that needed to be monitored in some way. The monitoring of reproductive capacity should be extended to aspects important in the mating systems, such as local densities, spatial patterns of age-size structure, and spatial and temporal patterns of breeding areas for broadcast spawners with poorly mobile or sessile adults. Similarly, such monitoring would also include female and male size and the number of female reproductive opportunities for crustacean species with copulating adults.©2004 NRC Canada

DUFOUR, R., B. SAINTE-MARIE, L. BOURASSA, 2003. Snow crab of the St. Lawrence estuary and the northern Gulf of St. Lawrence (Areas 13 to 17 and 12A, 12B and 12C) in 2002. Science, Stock Status Report, 2003/011, 24 p .

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DUFOUR, R., SAINTE-MARIE, B., BOURASSA, L., 2003. Crabe des neiges de l'estuaire et du nord du golfe Saint-Laurent (zones 13 a 17 et 12A, 12B et 12C) en 2002. Rapport sur l'état des stocks, 2003/011, 23 p .

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SAINTE-MARIE, B., C. TURCOTTE, 2003. Assessment of snow crab (Chionoecetes opilio) catchability by Japanese trap. Can. Tech. Rep. Fish. Aquat. Sci., 2508, 21 p .

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Experiments were performed in summer 1992 on the south shore of the Saint Lawrence Estuary to determine catchability of snow crab by the Japanese trap. Catchability, or trap effective fishing area, is the quotient of the mean number of individuals captured by a trap over the density of these individuals on the bottom. The units for expressing catchability are square metres. We evaluated crab density on the bottom with a beam trawl. Catchability was determined for males in various categories of maturity (adolescent versus adult), relative age (time elapsed since last moult) and size. Furthermore, we measured the effects on catch composition and yield of Japanese traps of prior occupancy by one or more live snow crabs, either hard-shell adolescent males, soft-shell (recently moulted) adult males or hard-shell adult males. Japanese traps were highly selective for adult males of legal size (≥95 mm carapace width, CW) with an intermediate shell condition (aged about 2-3 years since terminal moult). The catchability of these males was 1000 times greater (about 63,000 m2) than that of legal-size adult males moulted since a few weeks and 10 times greater than that of legal-size adult males with an old shell (aged about 5-6 years since terminal moult). Adolescent males of legal size were about 30 times less catchable than legal-size adult males with an intermediate shell. The relationship between catchability and size of adult males with an intermediate shell was dome-shaped, with males smaller than 80 mm CW and males larger than 135 mm CW having low catchability. Reasons for the differences of catchability among male categories are discussed. In general, the prior occupancy by hard-shell adult males resulted in a moderate reduction of the yield of Japanese traps, probably because crabs outside the traps were intimidated. Prior occupancy of Japanese traps by soft-shell adult males resulted in a large reduction of yield. However, this decrease can be attributed to the high mortality of soft-shell crabs and the snow crab’s aversion for dead conspecifics. The results of the present study could be used to adjust the yield of Japanese traps used in the commercial fishery or post-season trap surveys so that derived abundance indices better reflect the demography of the sampled populations.

SAINTE-MARIE, B., C. TURCOTTE, 2003. Évaluation de la capturabilité du crabe des neiges (chionoecetes opilio) par le casier japonais. Rapp. tech. can. sci. halieut. aquat., 2508, 23 p .

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Experiments were performed in summer 1992 on the south shore of the Saint Lawrence Estuary to determine catchability of snow crab by the Japanese trap. Catchability, or trap effective fishing area, is the quotient of the mean number of individuals captured by a trap over the density of these individuals on the bottom. The units for expressing catchability are square metres. We evaluated crab density on the bottom with a beam trawl. Catchability was determined for males in various categories of maturity (adolescent versus adult), relative age (time elapsed since last moult) and size. Furthermore, we measured the effects on catch composition and yield of Japanese traps of prior occupancy by one or more live snow crabs, either hard-shell adolescent males, soft-shell (recently moulted) adult males or hard-shell adult males. Japanese traps were highly selective for adult males of legal size (≥95 mm carapace width, CW) with an intermediate shell condition (aged about 2-3 years since terminal moult). The catchability of these males was 1000 times greater (about 63,000 m2) than that of legal-size adult males moulted since a few weeks and 10 times greater than that of legal-size adult males with an old shell (aged about 5-6 years since terminal moult). Adolescent males of legal size were about 30 times less catchable than legal-size adult males with an intermediate shell. The relationship between catchability and size of adult males with an intermediate shell was dome-shaped, with males smaller than 80 mm CW and males larger than 135 mm CW having low catchability. Reasons for the differences of catchability among male categories are discussed. In general, the prior occupancy by hard-shell adult males resulted in a moderate reduction of the yield of Japanese traps, probably because crabs outside the traps were intimidated. Prior occupancy of Japanese traps by soft-shell adult males resulted in a large reduction of yield. However, this decrease can be attributed to the high mortality of soft-shell crabs and the snow crab’s aversion for dead conspecifics. The results of the present study could be used to adjust the yield of Japanese traps used in the commercial fishery or post-season trap surveys so that derived abundance indices better reflect the demography of the sampled populations.

GOSSELIN, T., B. SAINTE-MARIE, L. BERNATCHEZ, 2003. Patterns of sexual cohabitation and female ejaculate storage in the American lobster (Homarus americanus). Behav. Ecol. Sociobiol., 55: 151-160 .

DIONNE, M., B. SAINTE-MARIE, E. BOURGET, D. GILBERT, 2003. Distribution and habitat selection of early benthic stages of snow crab Chionoecetes opilio. Mar. Ecol. Prog. Ser., 259: 117-128 .

SAINTE-MARIE, B., D. CHABOT, 2002. Ontogenetic shifts in natural diet during benthic stages of American lobster (Homarus americanus), off the Magdalen Islands. Fish. Bull., 100: 106-116 .

SAINTE-MARIE, B., 2002. DFO Quebec Region : data and analytical framework. Pages 7-11 in J.K. Rice & J. Moores (ed.). Zonal snow crab workshop ; atelier zonal sur le crabe des neiges (DFO, CSAS, Proceedings Series ; MPO, Secrétariat canadien de consultation scientifique, Série des comptes rendus, 2002/022) .

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PAILLE, N., B. SAINTE-MARIE, J.-C. BRÊTHES, 2002. Behavior, growth and survival of stage V lobsters (Homarus americanus) in relation to shelter availability and lobster density. Mar. Fresh. Behav. Physiol., 35(4): 203-219 .

PAUL, A.J., E.G. DAWE, R.W. ELNER, G.S. JAMIESON, G.H. KRUSE, R.S. OTTO, B. SAINTE-MARIE, T.C. SHIRLEY, D. WOODBY, D., 2002. Crabs in cold water regions : biology, management, and economics : proceedings of the symposium Crab2001, Crabs in Cold Water Regions : Biology, Management, and Economics, January 17-20, 2001, Anchorage, Alaska, USA. Alaska Sea Grant College Program (Alaska Sea Grant College Program report, AK-SG-02-01 ; Lowell Wakefield fisheries symposia series, 19), 866 p .

SAINTE-MARIE, B., J.-M. SÉVIGNY, M. CARPENTIER, 2002. Interannual variability of sperm reserves and fecundity of primiparous females of the snow crab (Chionoecetes opilio) in relation to sex ratio. Can. J. Fish. Aquat. Sci., 59: 1932-1940 .

Demographics of adults and reproductive condition of primiparous (first brood) females of the snow crab (Chionoecetes opilio) were monitored annually from 1994 to 2002 at a fished site to investigate the possibility that sperm supply limits embryo production. Abundance of primipara fluctuated 533-fold because of a recruitment pulse, and this caused a large oscillation in the sex ratio of adult males to primipara. Annual mean of stored ejaculate weight (SL) and potential fecundity index (PF, clutch weight x percent fertilized eggs) adjusted to constant primipara carapace width ranged from 31 to 130 mg by spermatheca and from 1.97 to 3.43 g by clutch, respectively. Annual mean of SL and number of stored sperm (range 3.81 x 106 to 35.00 x 106 sperm by spermatheca) decreased when sex ratio decreased, probably because of a combined reduction of sperm allocation and female promiscuity. Annual mean PF was negatively correlated with abundance of small males, which may reflect egg losses during postoviposition matings. Althoug sociosexual context has a large impact on reproductive condition of primipara, the possibility that sperm supply limits embryo production could not be confirmed or excluded because of the complexity of snow crab mating behavior.

SAINTE-MARIE, B., 2002. DFO Quebec Region : biological basis for the stock assessment. Pages 5-7 in J.K. Rice & J. Moores (ed.). Zonal snow crab workshop ; atelier zonal sur le crabe des neiges (DFO, CSAS, Proceedings Series ; MPO, Secrétariat canadien de consultation scientifique, Série des comptes rendus, 2002/022) .

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SAINTE-MARIE, B., M. LAFRANCE, 2002. Growth and survival of recently settled snow crab Chionoecetes opilio in relation to intra- and intercohort competition and cannibalism : a laboratory study. Mar. Ecol. Prog. Ser., 244: 191-203 .

TREMBLAY, M.J., B. SAINTE-MARIE, 2001. Canadian Lobster Atlantic Wide Studies (CLAWS) Symposium : abstracts and proceedings summary. Can. Tech. Rep. Fish. Aquat. Sci., 2328, 111 p .

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A symposium on lobster research conducted under the umbrella of Canadian Lobster Atlantic Wide Studies (CLAWS) was held in Moncton, N.B. from March 28-30, 2000. CLAWS is a multidisciplinary effort directed at understanding lobster production in the Canadian Atlantic. The objectives of the Moncton Symposium were to discuss recent and ongoing lobster research, to receive feedback from stakeholders, and to plan new research under CLAWS II. Participants at the Symposium included representatives from industry, universities and Fisheries and Oceans Science. The Symposium was structured around five Science sessions on days 1 and 2 : Larval drift, Juvenile lobsters, Catchability, Assessment Parameters, Fishermen and Science, and a general Poster session. On day 3 (Industry Day) overviews of each session were followed by general discussion. The final session of the Symposium discussed the research planned for CLAWS II.

TREMBLAY, M.J., B. SAINTE-MARIE, 2001. Symposium sur le Programme intégré sur le homard canadien et son environnement (PINHCE) : résumés et sommaire des travaux. Rapp. tech. can. sci. halieut. aquat., 2328, 130 p .

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A symposium on lobster research conducted under the umbrella of Canadian Lobster Atlantic Wide Studies (CLAWS) was held in Moncton, N.B. from March 28-30, 2000. CLAWS is a multidisciplinary effort directed at understanding lobster production in the Canadian Atlantic. The objectives of the Moncton Symposium were to discuss recent and ongoing lobster research, to receive feedback from stakeholders, and to plan new research under CLAWS II. Participants at the Symposium included representatives from industry, universities and Fisheries and Oceans Science. The Symposium was structured around five Science sessions on days 1 and 2 : Larval drift, Juvenile lobsters, Catchability, Assessment Parameters, Fishermen and Science, and a general Poster session. On day 3 (Industry Day) overviews of each session were followed by general discussion. The final session of the Symposium discussed the research planned for CLAWS II.

DUFOUR, R., B. SAINTE-MARIE, 2001. Snow crab of the estuary and northern Gulf of St. Lawrence (areas 13 to 17). Science, Stock Status Report, C4-01, 15 p .

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DUFOUR, R., B. SAINTE-MARIE, 2001. Crabe des neiges de l'estuaire et du nord du golfe du Saint-Laurent (zones 13 à 17). Rapport sur l'état des stocks, C4-01, 16 p .

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RONDEAU, A., B. SAINTE-MARIE, 2001. Variable mate-guarding time and sperm allocation by male snow crabs (Chionoecetes opilio) in response to sexual competition, and their impact on the mating success of females. Biol. Bull., 201: 204-217 .

DUFOUR, R., B. SAINTE-MARIE, 2000. Crabe des neiges de l'estuaire et du nord du golfe Saint-Laurent (zones 13 à 17). Rapport sur l'état des stocks, C4-01, 14 p .

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SAINTE-MARIE, G., B. SAINTE-MARIE, J.M. SEVIGNY, 2000. Ejaculate-storage patterns and the site of fertilization in female snow crabs (Chionoecetes opilio; Brachyura, Majidae). Can. J. Zool., 78: 1902-1917 .

A light-microscopy study of mated female snow crabs (Chionoecetes opilio) was conducted to investigate the site of fertilization and to resolve how multiple ejaculates are stored in the spermathecae. In its basic configuration, an ejaculate consisted of a layer or patch of spermatophores enclosing mature spermatids that was capped by a relatively large volume of amorphous matter, which in turn could include a patch of spermatophores containing immature spermatids. Up to 10-12 ejaculates were stored in the spermathecae with the largest loads. An ejaculate was initially deposited in the intermediate chamber and ventral part of the spermatheca, and was displaced toward the dorsal part of the spermatheca when a new ejaculate was inserted. Ejaculates were neatly stratified along the ventrodorsal axis of spermathecae with small to moderate loads, but they were disrupted and the storage pattern was disorderly in spermathecae with large loads. Ejaculate stratification favors last-male sperm precedence and single-male paternity. However, multiple-male paternity might occur in females with large spermathecal loads, in part because several ejaculates can co-occur close to the oviduct opening. Mixing of male and female gametes in preparation for oviposition, and probably also fertilization, occurs to some° in the ovaries©2000 National Research Council Canada

DUFOUR, R., B. SAINTE-MARIE, 2000. Snow crab of the estuary and northern Gulf of St. Lawrence (areas 13 to 17). Science, Stock Status Report, C4-01, 13 p .

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SAINTE-MARIE, B., N. URBANI, J.-M. SÉVIGNY, F. HAZEL, U. KUHNLEIN, 1999. Multiple choice criteria and the dynamics of assortative mating during the first breeding season of female snow crab Chionoecetes opilio (Brachyura, Majidae). Mar. Ecol. Prog. Ser., 181: 141-153 .

SAINTE-MARIE, G., B. SAINTE-MARIE, 1999. Reproductive products in the adult snow crab (Chionoecetes opilio). I, Observations on spermiogenesis and spermatophore formation in the vas deferens. Can. J. Zool., 77: 440-450 .

Some of the events infolding in the vas deferens of the adult snow crab (Chionoecetes opilio) were examined by means of light microscopy. Sperm celles entered the vas deferens as precursors of immature spermatids and developed into immature or mature spermatids within it. However, spermatozoa were not observed in the male reproductive tract. Two types of amorphous matter were added successively to sperm cells in the vas deferens. The first type was periodic acid - Schiff (PAS)-positive and apparently induced spermiogenesis when present in a sufficiently large amount. However, a smaller amount of this amorphous matter was sufficient to form the basal pellicle of spermatophores. The second type was PAS-negative and thickened the wall of spermatophores. Immature and mature spermatids were usually enclosed within distinct spermatophores. Enclosed mature spermatids were connected together by bridges formed by Feulgen-positive spikes coated and extended by PAS-positive amorphous matter. Once broken, the bridges appeared as arms that radiated from a spermatid. Peripheral mature spermatids were furthermore linked to the spermatophore wall by threads of PAS-positive amorphous matter. The bridges and threads may form a pathway for the diffusion of extraneous substances through the spermatophore wall to the innermost cells© 1999 National Research Council Canada

SAINTE-MARIE, G., B. SAINTE-MARIE, 1999. Reproductive products in the adult snow crab (Chionoecetes opilio). II, Multiple types of sperm cells and of spermatophores in the spermathecae of mated females. Can. J. Zool., 77: 451-462 .

Contents of the spermathecae of mated adult snow crabs (Chionoecetes opilio) were examined by light microscopy. The contents could consist of water and three basic types of amorphous matter and of spermatophores. Water was present in the form of large patches or smaller spheres. Of the two major types of amorphous matter, one reacted positively and one negatively to periodic acid - Schiff's reagent (PAS), and one was only, and one predominantly, of male origin. The minor type of amorphous matter was orange and of female origin and could include dark cellular debris. Spermatophores enclosed either immature spermatids, mature spermatids, hitherto unreported spermatozoa, or cell forms intermediate between these three types. Peripheral mature spermatids and spermatozoa had a polarized orientation and were attaches to the spermatophore wall. Overall, amorphous matter and spermatophores were topographically segregated by type within a spermatheca, and spermatophores enclosing immature spermatids occurred mostly in PAS-negative amorphous matter. Spermatid differentiation can unfold in the female reproductive tract as well as in the vas deferens, while the transformation of spermatids into spermatozoa was observed only in the female. The diversity of sperm cell types and the ordered placement of semen constituent within the spermatheca suggest that sperm are partitioned for short- or long-term use©1999 National Research Council Canada

DUFOUR, R., B. SAINTE-MARIE, 1999. Crabe des neiges de l'estuaire et du nord du golfe du Saint-Laurent (zones 13 à 17). Rapport sur l'état des stocks, C4-01, 14 p .

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DUFOUR, R., B. SAINTE-MARIE, 1999. Snow crab of the estuary and northern Gulf of St. Lawrence (Areas 13 to 17). Science, Stock Status Report, C4-01, 14 p .

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URBANI, N., B. SAINTE-MARIE, J.-M. SÉVIGNY, D. ZADWORNY, U. KUHNLEIN, 1998. Sperm competition and paternity assurance during the first breeding period of female snow crab (Chionoecetes opilio) (Brachyura : Majidae). Can. J. Fish. Aquat. Sci., 55: 1104-1113 .

Two highly polymorphic microsatellite DNA loci were used to determine paternity of primiparous snow crab Chionoecetes opilio females. In addition, female spermathecal contents were genotyped at the two loci and histological analysis of the organs was carried out to elucidate patterns of sperm competition. Spermathecal contents from wild-caught females were cut into several cross sections and each section genotyped individually. Both wild and laboratory females commonly mated with several males whose ejaculates were stored and stratified in the spermathecae. Genetic typing of the offspring of laboratory-reared females revealed single paternity and indicated that the last mate to inseminate a female before oviposition gained paternity of the clutch. The predominant mechanism ensuring single paternity appeared to be sperm stratification. In wild-caught females, the microsatellite typing of the offspring also revealed single paternity, but larvae appeared to be sired by males whose genotypes were found in the spermathecal cross sections towards the dorsal end (blind end) of the spermathecae. This suggested that they were the first males to mate with females that they guarded until oviposition, and females remated with other males thereafter.

DUFOUR, R., B. SAINTE-MARIE, 1998. Snow crab of the estuary and northern Gulf of St. Lawrence (areas 13 to 17). Science, Stock Status Report, C4-01, 13 p .

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ALUNNO-BRUSCIA, M., B. SAINTE-MARIE, 1998. Abdomen allometry, ovary development, and growth of female snow crab, Chionoecetes opilio (Brachyura, Majidae), in the northwestern Gulf of St. Lawrence. Can. J. Fish. Aquat. Sci., 55: 459-477 .

Growth and maturation of female snow crab (Chionoecetes opilio) in the northwestern Gulf of St. Lawrence were elucidated using carapace size-frequency distributions, indices of molting activity, ovarian mass, and laboratory and field measures of growth per molt. Females develop in three major stages : immature, with narrow abdomen and no detectable ovaries; prepubescent, with narrow abdomen and previtellogenic followed by vitellogenic ovaries; and adult, with broad abdomen and reproductive capability. Additionally, there is an ephemeral pubescent stage represented by females temporally close to the maturity molt. A reduction in the rate of carapace and abdomen growth occurs at the passage from immature to prepubescent, because energy is diverted into germinal growth, and adulthood is reached at a terminal molt to maturity. The pattern of abdomen growth relative to carapace is complex, consisting in successive phases of low, high, and again low positive allometry with increasing size of immature-prepubescent females. Over the period 1989-1996, a few females became adult at instar 8 at 4.5 yr postlarval age, but more commonly, maturity occurred at instars 9 and 10 at ages 5.5 and 6.5 yr, respectively. Average size at maturity may be temperature dependent, and within cohorts, larger females may mature earlier than smaller females.

OUELLET, P., B. SAINTE-MARIE, 1998. Distribution and abundance of lobster (Homarus americanus) larvae in the Magdalen Islands : implications for the availability of postlarvae and benthic settlement. Pages 101-107 in L. Gendron (ed.). Proceedings of a Workshop on Lobster Stock Enhancement held in the Magdalen Islands (Quebec) from October 29 to 31, 1997 . Dept. of Fisheries and Oceans (Can. Ind. Rep. Fish. Aquat. Sci., 244) .

SAINTE-MARIE, G., B. SAINTE-MARIE, 1998. Morphology of the spermatheca, oviduct, intermediate chamber, and vagina of the adult snow crab (Chionoecetes opilio). Can. J. Zool., 76: 1589-1604 .

The morphology of the reproductive tract of the adult female snow crab (Chionoecetes opilio) at different stages of reproductive life was examined within the framework of studies on sperm competition. Apart from an ovary and oviduct, the tract consists of an epithelium-lined spermatheca and a cuticle-lined intermediate chamber and vagina. The morphological traits of these components are markedly influenced by the° of spermathecal loading. Several amendments to previous descriptions are necessary : (i) pouches are not a permanent feature of the spermatheca; (ii) the outer part of the spermatheca wall is a true capsule that overlies a multilayered epithelium divided into anchoring, proliferative, and secretory strata; (iii) spermathecal secretions are of the merocrine type; and (iv) the epithelium of the spermatheca is not pigmented. Tridimensional analysis revealed that the vagina connects to the intermediate chamber via a parietal opening. The intermediate chamber is a flexible structure circled dorsally by 3-5 cuticular folds that project into its lumen. The oviduct crosses the thicker terminal segment of the spermathecal epithelium with a dorsoventral incidence, to a point opposite the parietal opening of the vagina, in such a way that it delivers its contents almost directly into the intermediate chamber. Novel functional implications of these findings are hypothesized©1998 National Research Council Canada

SAINTE-MARIE, B., D. GILBERT, 1998. Possible effects of changes in CIL temperature and thickness on population dynamics of snow crab, Chionoecetes opilio, in the Gulf of Saint Lawrence. DFO, Canadian Stock Assessment Secretariat, Research Document, 98/38, 19 p .

URBANI, N., B. SAINTE-MARIE, J.-M. SÉVIGNY, D. ZADWORNY, U. KUHNLEIN, 1998. Mating dynamics of the snow crab (Chionoecetes opilio, Brachyura : Majidae) : an analysis using DNA microsatellite markers. Pages 241-244 in Le Gal & Halvorson (ed.). New developments in marine biotechnology. Plenum Press, New York .

URBANI, N., J.-M. SÉVIGNY, B. SAINTE-MARIE, D. ZADWORNY, U. KUHNLEIN, 1998. Identification of microsatellite markers in the snow crab Chionoecetes opilio. Mol. Ecol., 7: 357-358 .

OUELLET, P., B. SAINTE-MARIE, 1998. Distribution et abondance des larves de homard (Homarus americanus) aux Îles-de-la-Madeleine : implications pour la disponibilité des postlarves et le succès de l'établissement benthique. Pages 109-115 in L. Gendron (éd.). Compte-rendu d'un atelier de travail sur l'ensemencement des stocks de homard, tenu aux Îles-de-la-Madeleine (Québec) du 29 au 31 octobre 1997. MPO (Rapp. Can. Ind. Sci. Halieut. Aquat., 244) .

DUFOUR, R., B. SAINTE-MARIE, 1998. Crabe des neiges de l'estuaire et du nord du golfe du Saint-Laurent (zones 13 à 17). Rapport sur l'état des stocks, C4-01, 15 p .

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SAINTE-MARIE, B., J.-M. SÉVIGNY, Y. GAUTHIER, 1997. Laboratory behavior of adolescent and adult males of the snow crab (Chionoecetes opilio) (Brachyura : Majidae) mated noncompetitively and competitively with primiparous females. Can. J. Fish. Aquat. Sci., 54: 239-248 .

In the laboratory, virgin females of snow crab (Chionoecetes opilio) were placed singly with one or two males to investigate mating behavior. Physiologically mature males, both adolescent and adult mated with females. Typically, matings comprised a precopulatory period, one or more copulations, and a postcopulatory period during which the female extruded eggs. Some females copulated after oviposition. In noncompetitive mating, adult males of 40-60 mm carapace width had repeated but brief copulations, while adult males of 120-140 mm carapace width had one long copulation. In pairs of competing adult males, large body size was an advantage both in taking over a female from a rival and in preventing female takeover by a rival. In competitions between an adult male and a larger adolescent male, the adult was more likely to dispossess his rival or revent takeover. The variance in mass of ejaculate stored by females was explained (98 %) by total copulatory time and number of copulations. Females with two mates obtained more copulations and larger ejaculate stores than females with one mate, suggesting that interannual variability in ejaculate stores reported for wild females is related to the level of male sexual competition.

SAINTE-MARIE, B., 1997. Comment - An improved link between industry, management and science : review of case history of the southwestern Gulf of St. Lawrence snow crab fishery. Can. J. Fish. Aquat. Sci., 54: 496-500 .

DUFOUR, R., B. SAINTE-MARIE, 1997. Snow crab of the estuary and northern Gulf of St. Lawrence (areas 13 to 17). Science, Stock Status Report, C4-01, 19 p .

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LOVRICH, G.A., B. SAINTE-MARIE, 1997. Cannibalism in the snow crab, Chionocetes opilio (O. Fabricius) (Brachyura : Majidae), and its potential importance to recruitment. J. Exp. Mar. Biol. Ecol., 211: 225-245 .

BOSSÉ, L., B. SAINTE-MARIE, J. FOURNIER, 1996. Les invertébrés des fonds meubles et la biogéographie du fjord du Saguenay. Rapp. tech. can. sci. halieut. aquat., 2132, 45 p .

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During the summers of 1990 and 1991, the bottom of the Saguenay Fjord at depths greater than 60 m was photographed and sampled with baited traps, a box corer, a grab and an epibenthic sledge. The photographic survey shows a progressive change in sediment and epibenthic communities along the Fjord. Over 320 species of benthic invertebrates were collected, among which 178 represent first records for the Fjord. Fourteen of the latter were hitherto unknown to the estuary or Gulf of St. Lawrence. Amphipods, polychaetes, gastropods and bivalves are the predominant taxa in number of species. Including those benthic invertebrates listed in previous studies, the Fjord should comprise at least 410 species. Among these species, 11 % are of arctic affiliation and 58 % are of boreal affiliation. Nearly 80 % of the Fjord species are also present in the Lower St. Lawrence estuary. Sixteen of the 78 species which are found in the Fjord but not in the Lower estuary are arctic affiliated. Populations of these 16 arctic species are probably relics of ancestral populations which have persisted in the Fjord due to an appropriate combination of hydrothermal and edaphic conditions. However, relic populations of arctic species also exist in other restricted parts of the Estuary and the Gulf of Saint Lawrence. In light of these results, the status of arctic enclave which is often ascribed to the Fjord seems questionable. This term leaves the impression that the Saguenay Fjord's fauna is a replica of Arctic fauna, when it is in fact mostly of boreal affiliation.

SÉVIGNY, J.-M., B. SAINTE-MARIE, 1996. Electrophoretic data support the last-male sperm precedence hypothesis in the snow crab, Chionoecetes opilio (Brachyura : Majidae). J. Shellfish Res., 15: 437-440 .

DUFOUR, R., B. SAINTE-MARIE, 1996. Snow crab of the estuary and northern Gulf of St. Lawrence. DFO, Atlantic Fisheries, Stock Status Report, 96/6, 13 p .

BOUCHARD, S., B. SAINTE-MARIE, J.N. McNEIL, 1996. Indirect evidence indicates female semiochemicals release male precopulatory behaviour in the snow crab, Chionoecetes opilio (Brachyura : Majidae). Chemoecology, 7: 39-44 .

SAINTE-MARIE, B., J.-M. SÉVIGNY, B.D. SMITH, G.A. LOVRICH, 1996. Recruitment variability in snow crab (Chionoecetes opilio) : pattern, possible causes, and implications for fishery management. Pages 451-478 In High latitude crabs : biology, management and economics : proceedings of the International Symposium. University of Alaska Fairbanks (Alaska Sea Grant report, 96-02) .

SAINTE-MARIE, B., S. RAYMOND, J.-C. BRÊTHES, 1995. Growth and maturation of the benthic stages of male snow crab, Chionoecetes opilio (Brachyura : Majidae). Can. J. Fish. Aquat. Sci., 52: 903-924 .

SAINTE-MARIE, B., C. CARRIÈRE, 1995. Fertilization of the second clutch of eggs of snow crab, Chionoecetes opilio, from females mated once or twice after their molt to maturity. Fish. Bull., 93: 759-764 .

LOVRICH, G.A., B. SAINTE-MARIE, B.D. SMITH, 1995. Depth distribution and seasonal movements of Chionoecetes opilio (Brachyura : Majidae) in Baie Sainte-Marguerite, Gulf of Saint Lawrence. Can. J. Zool., 73: 1712-1726 .

Bimonthly beam-trawling from April 1991 to May 1992 and diver observations were used to assess distribution and large-scale movement of snow crabs, Chionoecetes opilio, over depths of 4-140 m in a bay of the northern Gulf of Saint Lawrence. Megalopae settled during October 1991. Immature crabs of instars I-IV (i.e., 3.3-9.7 mm mean carapace width, CW) were found predominantly on bottoms similar to 50-80 m deep and were cryptic and sedentary. Most immature crabs of instars V-VIII (i.e., 14.1-34.5 mm mean CW) and adolescent males (i.e., producing sperm but not terminally moulted) migrated in winter to subtidal grounds, where they moulted. Movement to shallow waters was massive and resulted in a mean density of 860 crabs per 1000 m2 at similar to 15 m in December 1991. Adult males (i.e., producing sperm and terminally moulted) of <70 mm mean CW also moved to the shallow grounds from October to December 1991, where some mated with pubescent-primiparous females (i.e., adult, first brood) from January to April 1992. Adult males of >90 mm mean CW were mainly found at depths >80 m over most of the year, but from March to May 1992 an increase in mean CW of adult males at <80 m indicated some upslope movement, probably to mate with multiparous females (i.e., adult, second or ulterior brood). Adult females were more gregarious and sedentary than adult males. We expand on the hypothesis that interannual variability in recruitment to adulthood tends to reflect differences in year-class strength, and that year-class strength varies in accordance with megalopal supply and (or) survivorship of cryptic instars.©1995 National Research Council Canada

CYR, C., B. SAINTE-MARIE, 1995. Catch of Japanese crab traps in relation to bait quantity and shielding. Fish. Res., 24: 129-139 .

SAINTE-MARIE, B., R. DUFOUR, 1995. Snow crab in the estuary and northern Gulf of St. Lawrence (fishing areas 13, 14, 15, 16 and 17). Pages 2-26 in L. Savard (ed.). Status report on invertebrates in 1994 : crustaceans and molluscs on the Québec coast, northern shrimp and zooplankton in the estuary and Gulf of St. Lawrence. DFO (Can. Manuscr. Rep. Fish. Aquat. Sci., 2323) .

SAINTE-MARIE, B., R. DUFOUR, 1995. Crabe des neiges de l'estuaire et du nord du golfe du Saint-Laurent (zones de pêche 13, 14, 15, 16 et 17). Pages 2-27 in L. Savard (éd.). Rapport sur l'état des invertébrés en 1994 : crustacés et mollusques des côtes du Québec, crevette nordique et zooplancton de l'estuaire et du golfe du Saint-Laurent. MPO (Rapp. manus. can. sci. halieut. aquat., 2323) .

SAINTE-MARIE, B., G.A. LOVRICH, 1994. Delivery and storage of sperm at first mating of female Chionoecetes opilio (Brachyura : Majidae) in relation to size and morphometric maturity of male parent. J. Crust. Biol., 14(3): 508-521 .

SAINTE-MARIE, B., 1994. The nascent amphipod fishery in Québec : exploiting a low-trophic level, multispecies resource. Pages 97-102 in L. Gendron & S. Robinson (ed.). The development of underutilized invertebrate fisheries in eastern Canada. Dept. of Fisheries and Oceans (Can. Manus. Rep. Fish. Aquat. Sci., 2247) .

BOSSÉ, L., B. SAINTE-MARIE, J. FOURNIER, P. BRUNEL, 1994. Inventaire et biogéographie des invertébrés des fonds meubles du fjord du Saguenay. Pages 44-49 in J.-M. Sévigny & C.M. Couillard (éd.). Le fjord du Saguenay : un milieu exceptionnel de recherche. Ministère des pêches et des océans (Rapp. manus. can. sci. halieut. aquat., 2270) .

ANGERS, A., F. POTHIER, J.-M. SÉVIGNY, B. SAINTE-MARIE, 1994. Tissue specificity and ontogeny of lactate dehydrogenase in snow crab, Chionoecetes opilio (Brachyura, Majidae). Comp. Biochem. Physiol., 108B: 385-395 .

SAINTE-MARIE, B., R. DUFOUR, 1994. Crabe des neiges de l'estuaire et du nord du golfe du Saint-Laurent (zones de pêche 13, 14, 15, 16 et 17). Pages 2-23 in L. Savard (éd.). Rapport sur l'état des invertébrés en 1993 : crustacés et mollusques des côtes du Québec et crevette nordique de l'estuaire et du golfe du Saint-Laurent. MPO (Rapp. manus. can. sci. halieut. aquat., 2257) .

HODSON, P.V., E. PELLETIER, R. McLEOD, J. HELLOU, B. SAINTE-MARIE, C.M. COUILLARD, J.-M. SÉVIGNY, 1994. Chemical contamination of surface sediments and biota of the Saguenay Fjord. Pages 97-104 in J.-M. Sévigny & C.M. Couillard (éd.). Le fjord du Saguenay : un milieu exceptionnel de recherche. Ministère des pêches et des océans (Rapp. manus. can. sci. halieut. aquat., 2270) .

SAINTE-MARIE, B., R. DUFOUR, 1994. Snow crab in the estuary and northern Gulf of St. Lawrence (fishing areas 13, 14, 15, 16 and 17). Pages 2-20 in L. Savard (ed.). Status report on invertebrates 1993 : crustaceans and molluscs on the Québec coast and northern shrimp in the Estuary and Gulf of St. Lawrence. DFO (Can. Manuscr. Rep. Fish. Aquat. Sci., 2257) .

SAINTE-MARIE, B., 1993. Reproductive cycle and fecundity of primiparous and multiparous female snow crab, Chionoecetes opilio, in the northwest Gulf of Saint Lawrence. Can. J. Fish. Aquat. Sci., 50: 2147-2156 .

DUTIL, J.-D., J. MUNRO, R. DUFOUR, Y. GAUTHIER, F. HAZEL, B. SAINTE-MARIE, L. PROVENCHER, J. BOYER, D. HARDY, 1993. L'élevage du crabe des neiges : bilan des travaux de recherche. Ministère des pêches et des océans, Programme fédéral de développement des pêches du Québec, 12 p .

SAINTE-MARIE, B., V. LAPOINTE, D. ARCHAMBAULT, R. DUFOUR, 1992. Distribution spatiale et structure de taille du crabe des neiges, Chionoecetes opilio (O. Fabricius), dans le fjord du Saguenay. Rapp. manus. can. sci. halieut. aquat., 2156, 12 p .

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Exploratory fishing with baited traps was conducted in the Saguenay Fjord in July and August 1988 to determine whether or not Chionoecetes opilio occurred there and, if need be, to estimate the size of the population. The C. opilio in the Saguenay Fjord constitute a breeding population and there is local recruitment of juveniles. Mingling of individuals from C. opilio populations in the Fjord and in the Saint Lawrence Estuary may be limited by the 20-m deep external sill and by marked stratification of the water column. The population of C. opilio is apparently small and is in a quasi-virgin state.

SAINTE-MARIE, B., F. HAZEL, 1992. Moulting and mating of snow crab, Chionoecetes opilio (O. Fabricius), in shallow waters of the northwestern Gulf of Saint Lawrence. Can. J. Fish. Aquat. Sci., 49: 1282-1293 .

SAINTE-MARIE, B., 1992. Foraging of scavenging deep-sea lysianassoid amphipods. Pages 105-124 in G.T. Rowe & V. Pariente (ed.). Deep-sea food chains and the global carbon cycle. Kluwer Academic Publishers .

LAPOINTE, V., B. SAINTE-MARIE, 1992. Currents, predators, and the aggregation of the gastropod Buccinum undatum around bait. Mar. Ecol. Prog. Ser., 85: 245-257 .

LEGENDRE, L., K. JUNNIPER, D. BOOTH, A. CEMBELLA, Y. DE LAFONTAINE, J. GAGNÉ, W.G. HARRISON, S. ROY, B. SAINTE-MARIE, A. SINCLAIR, R. TRITES, A.F. VÉZINA, 1991. Report of the Workshop on Biological Oceanography. Pages 23-30 in J.-C. Therriault (ed.). The Gulf of St. Lawrence : small ocean or big estuary ? Dept. of Fisheries and Oceans (Can. Spec. Publ. Fish. Aquat. Sci., 113) .

LEGENDRE, L., K. JUNNIPER, D. BOOTH, A. CEMBELLA, Y. DE LAFONTAINE, J. GAGNÉ, W.G. HARRISON, S. ROY, B. SAINTE-MARIE, A. SINCLAIR, R. TRITES, A.F. VÉZINA, 1991. Rapport de l'atelier de travail sur l'océanographie biologique. Pages 23-30 in J.-C. Therriault (éd.). Le golfe du Saint-Laurent : petit océan ou grand estuaire? Ministère des pêches et des océans (Publ. spéc. can. sci. halieut. aquat., 113) .

SAINTE-MARIE, B., 1991. Whelk (Buccinum undatum) movement and its implications for the use of tag-recapture methods for the determination of baited trap fishing parameters. Can. J. Fish. Aquat. Sci., 48: 751-756 .

SAINTE-MARIE, B., 1991. A review of the reproductive bionomics of aquatic gammaridean amphipods : variation of life history traits with latitude, depth, salinity and superfamily. Hydrobiologia, 223: 189-227 .

SAINTE-MARIE, B., G. LAMARCHE, J.-M. GAGNON, 1990. Reproductive bionomics of some shallow-water lysianossoids in the Saint Lawrence Estuary, with a review on the fecundity of the Lysianassoidea (Crustacea, Amphipoda). Can. J. Zool., 68: 1639-1644 .

The life history and fecundity of five shallow-water lysianassoids from the Saint Lawrence Estuary were examined. Orchomenella minuta and Or. pinguis are annual and iteroparous (two broods), Psammonyx terranovae is iteroparous (two broods) and probably biennial, and Anonyx sarsi and Onisimus litoralis are biennial and semelparous. Females of A. sarsi and On. litoralis cease feeding on bait shortly before or after oviposition, whereas females of the iteroparous Or. pinguis stop feeding on bait only when broods are in the latest stages of embryo development. These ontogenetic changes may result from gut constriction caused by developing ovaries and broods, or may be due to behavioral changes. Data on the fecundity of the Lysianassoidea are reviewed, and it is concluded that deep-living species are probably much less fecund than shallow-living species. Anonyx nugax, Or. pinguis, and A. sarsi are more fecund than other lysianassoids, possibly because of their high-risk carrion-feeding and suprabenthic foraging activities©1990 National Research Council Canada

SAINTE-MARIE, B., J.A. PERCY, J.R. SHEA, 1989. A comparison of meal size and feeding rate of the lysianassid amphipods Anonyx nugax, Onisimus (=Pseudalibrotus) litoralis and Orchomenella pinguis. Mar. Biol., 102: 361-368 .

SAINTE-MARIE, B., R. DUFOUR, C. DESJARDINS, 1988. Beaching of snow crabs (Chionoecetes opilio) on the north shore of the Gulf of Saint Lawrence. Naturaliste can., 115 :105-109 .

SAINTE-MARIE, B., 1987. Meal size and feeding rate of the shallow-water lysianassid Anonyx sarsi (Crustacea : Amphipoda). Mar. Ecol. Prog. Ser., 40: 209-219 .

SAINTE-MARIE, B., B.T. HARGRAVE, 1987. Estimation of scavenger abundance and distance of attraction to bait. Mar. Biol., 94: 431-443 .