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Effects of Food Availability on Survival, Growth, and Reproduction of the Grass Shrimp Palaemonetes Pugio: a Laboratory Study

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Effects of Food Availability on Survival, Growth, and Reproduction of the Grass Shrimp Palaemonetes Pugio: a Laboratory Study MARINE ECOLOGY PROGRESS SERIES Vol. 220: 231–239, 2001 Published September 27 Mar Ecol Prog Ser Effects of food availability on survival, growth, and reproduction of the grass shrimp Palaemonetes pugio: a laboratory study Kathleen A. Reinsel1,*, Patricia S. Glas2, James R. Rayburn3, M. Karen Pritchard4, William S. Fisher5 1Department of Biology, Wittenberg University, PO Box 720, Springfield, Ohio 45504, USA 2Department of Biology, The Citadel, 171 Moultrie Street, Charleston, South Carolina 29409, USA 3Department of Biology, Jacksonville State University, 700 Pelham Road. N., Jacksonville, Alabama 36265-1602, USA 4Biology Department, Pensacola Junior College, Pensacola, Florida 32504, USA 5Gulf Ecology Division, National Health and Environmental Effects Research Laboratory, US Environmental Protection Agency, Gulf Breeze, Florida 32561, USA ABSTRACT: Grass shrimp are abundant, ecologically important inhabitants of estuarine ecosystems that have been used extensively in laboratory experiments. However, optimal laboratory feeding conditions have not been determined. We used a laboratory experiment to determine the effects of food availability on survival, growth and reproduction in adult Palaemonetes pugio Holthuis and hatching success of their offspring. Fifteen groups of 50 shrimp were fed 1 of 5 rations of TetraMin® flake food, a standard laboratory diet, for 12 wk; supplementary amounts of newly hatched Artemia sp. nauplii were also given twice weekly. Adult survival, female reproductive condition, and embryo hatching success were determined every 2 wk. Growth was determined from measurements at the beginning and end of the study. Limited food availability caused severe mortality and reduced growth of female shrimp, whereas male shrimp were not affected. Reproduction, as measured by percent gravid females and individual clutch size, was not affected by food availability. However, estimated population-level embryo production was reduced indirectly for the lower rations through mortality and smaller size of females. Hatching success was low in all treatments. Further studies are needed to determine optimal laboratory feeding conditions for grass shrimp so they may be better utilized as predictors of responses in the field. KEY WORDS: Feeding · Reproduction · Grass shrimp · Palaemonetes · Food limitation · Nutrition · Hatching success Resale or republication not permitted without written consent of the publisher INTRODUCTION tidal creeks (Scott et al. 1992). As deposit-feeding omnivores, grass shrimp break down decaying marsh Grass shrimp are conspicuous inhabitants of estuar- grass and thereby influence the cycling of nutrients, ies throughout the United States. Three species that particulates and energy in estuarine systems (Welsh often co-occur along the Atlantic and Gulf Coasts, 1975). In addition, they are important prey for early Palaemonetes pugio Holthuis, P. vulgaris Say and P. and juvenile life stages of several commercially intermedius Holthuis, can comprise up to 56% of the important fish species, including red drum and spot- total abundance of pelagic macrofauna in estuarine ted sea trout (Gunter 1945, Moody 1950, Wenner 1995, Wenner & Archambault 1996). Because of their ecological significance, they have potential utility as *E-mail: [email protected] sentinels of ecosystem integrity. To develop this © Inter-Research 2001 232 Mar Ecol Prog Ser 220: 231–239, 2001 potential, it is important that we understand the bio- MATERIALS AND METHODS logical characteristics of grass shrimp and environ- mental factors that influence their survival, growth, Field collection. Approximately 1500 grass shrimp and reproduction. were collected with dip nets on 5 August 1997 from the Like many decapod crustaceans, development of Escambia River near Pensacola, Florida, USA. Shrimp grass shrimp embryos occurs externally on the pleo- were collected from a 30 m-long stretch of sandy pods of the female. The reproductive pattern of grass shoreline with Spartina alterniflora growing along the shrimp is similar to that described for Palaemonetes bank. On the day of collection, water temperature was varians (Antheunisse et al. 1968) and P. kadiakensis 30°C and salinity was 10 ppt. Shrimp were transported (White 1949). Soon after a female molts, 1 or more to the laboratory in coolers containing water from the males may deposit spermatophores externally on her collection site. Water in the coolers was aerated, and ventral thorax. Eggs are extruded within 2 to 3 h of ma- shrimp were maintained in the coolers overnight. In ting, fertilized as they pass over the spermatheca, and the laboratory, the salinity of the water in the coolers attached with ‘glaire‘ or cement to pleopodal setae that was raised in increments of 2 to 4 ppt every 4 to 6 h to form the incubation chamber (Williams 1984). During acclimate shrimp to 20 ppt used in experimental hold- the embryonic development period (roughly 2 wk dur- ing aquaria, which were fed by flowing, filtered sea- ing summer months, but dependent on temperature) water. the female may re-initiate oogenesis to provide new Laboratory conditions. On the day after field collec- ova for extrusion at the next molt (White 1949). Repro- tion, 30 female and 20 male grass shrimp identified as duction in Palaemonetes species in the field occurs Palaemonetes pugio Holthuis (Williams 1984) were from early spring through fall, depending upon the randomly assigned to each of 15 aquaria (40 l). The location. total length of each shrimp, from the tip of the rostrum Because of their ecological importance and their rel- to the end of the telson, was measured with calipers to ative hardiness in the laboratory, grass shrimp have the nearest 0.1 mm. In addition, the presence or ab- historically been used in a variety of laboratory exper- sence of egg masses on the pleopods and visible oocy- iments. These range from studies of their physiology tes in the ovary were recorded for all female shrimp. (e.g. Goddard & Forward 1991, Okazaki & Freeman To examine the effects of food availability on sur- 1993) and ecology (e.g. Bell & Coull 1978, Kneib 1987, vival, growth and reproduction, grass shrimp were Olmi & Lipcius 1991, Vernberg & Piyatiratitivorakul maintained on 5 different food rations in a 12 wk labo- 1998) to toxicology studies (e.g. Tatem et al. 1976, Wil- ratory experiment (6 August to 30 October 1997). son et al. 1985, Weis et al. 1992, Khan et al. 1993, Key Three replicate 40 l aquaria for each of the 5 food treat- et al. 1998). Due to the relatively short developmental ments were arranged in a randomized block design. period, embryos of Palaemonetes pugio have recently Aquaria were supplied with flow-through filtered, been used in an in vitro assay to examine developmen- salinity-controlled (20 ppt) seawater and were aerated tal effects of environmental toxicants (Fisher & Foss with airstones. The flow rate and aeration were similar 1993, Rayburn et al. 1996, Rayburn & Fisher 1997, in all aquaria and were monitored daily. The room was 1999, Shelton et al. 1999). maintained on a 16 h:8 h light:dark cycle and aquaria 1 Results of toxicological studies using grass shrimp were covered with /4 inch (6.4 mm) plastic mesh to adults and embryos have the potential to be extremely prevent escape of shrimp. useful in predicting the effects of environmental cont- Shrimp were fed daily with 1 of 5 different amounts aminants in field situations. However, for laboratory of TetraMin® Standard Mix flake food (TetraWerke, results to be interpreted accurately, it is imperative Melle, Germany). Previous experience with laboratory that laboratory holding conditions be appropriate for culture had shown that a daily ration of 0.025 g survival, growth and reproduction of adult shrimp. In shrimp–1 resulted in little excess food after 24 h (Foss & particular, an optimal diet for production of viable off- Rayburn 1997). This standard ration was selected as 1 spring is essential for studies involving shrimp em- of the 5 experimental treatments. The other treatments 1 1 bryos. Although the subject of food quality has been were /4, /2, 2 and 4 times the standard ration, result- extensively studied for shrimp species used in aqua- ing in daily food rations ranging from 0.0062 to 0.1 g culture (e.g. review by Harrison 1990), specific studies shrimp–1. The shrimp were fed at the same time each addressing nutritional requirements for adult survival, day. In order to maintain a constant amount of food per growth, and production of viable embryos have not shrimp, the total food amounts were adjusted every been performed for Palaemonetes species. In this study 2 wk to compensate for any mortalities that occurred in we performed a laboratory experiment examining the the aquaria. Equal numbers of newly hatched Artemia effects of food availability on survival, growth and sp. nauplii were added to each aquarium twice weekly reproductive output of P. pugio. as a supplemental food source, since many crustaceans Reinsel et al.: Effects of food availability on survival, growth, and reproduction 233 cannot synthesize cholesterol and some polyunsatu- interaction effects in repeated-measures designs were rated fatty acids (Whitney 1969, 1970, Kanazawa & followed by 1-way ANOVAs for each sampling time. Teshima 1971). Naupliar concentrations (100 l–1 in all Survival data were analyzed with separate 2-way aquaria) were significantly lower than the 5000 l–1 typ- (ration, time) repeated-measures ANOVAs for female ically used to rear crustacean larvae (Welch & Epifanio and male shrimp. Data were expressed as the propor- 1995) in order to prevent significant alteration of the tion of the initial number of shrimp that survived to food levels in the lower treatments. each census. Since 1 female shrimp was removed from Measurements. Aquaria were examined daily for the each aquarium for embryo survival tests at each cen- presence of gravid females, dead individuals and sus, proportions of female survivors were corrected for the amount of food remaining in each aquarium. The these removals. water quality in all aquaria was analyzed weekly. Tem- Growth data were analyzed separately for female perature and dissolved oxygen measurements were and male shrimp.
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