GASTRIC EVACUATION AND DIGESTION STATE INDICES FOR GAG Mycteroperca microlepis CONSUMING FISH AND CRUSTACEAN PREY

By ELIZABETH JOANNE BERENS

A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2005

Copyright 2005 by Elizabeth Joanne Berens

To my parents, Charles and Joanne, For their support and encouragement which has allowed me to pursue my interests in the marine environment and the various organisms within.

Department of Fisheries and Aquatic Sciences. and Dr. I thank the American Fisheries Society. I thank Day Cherry and DJ White for their assistance with invertebrate collections and for giving me invaluable insight into a commercial fishing industry. Rick Kline. University of Florida. In particular. Karen Bjorndal. iv . Dr. Department of Zoology. Mark Luttenton for introducing me to fisheries science many years ago at Grand Valley State University.ACKNOWLEDGMENTS I would like to thank Dr. Debra Murie. Florida Chapter. Steve Larsen. and Doug Marcinek for assistance with laboratory set-up and specimen collection. University of Florida. William Lindberg. and Doug Colle and Larry Tolbert for advice and assistance with certain aspects of this study. This study would not have been possible without the help of many people and I especially thank Mark Butler. Eddie Leonard. Jackie Debicella. I thank my committee. Daryl Parkyn. and the University of Florida’s Fisheries and Aquatic Sciences Department for financial and logistical assistance during this study. Dr. for offering advice and assistance along the way. Florida Sea Grant. Dr.

..19 Gag Collections and Maintenance ....................................................................................................................................................................................37 Indices of Digestion for Baitfish Consumed by Gag.............................................................29 3 RESULTS ............................TABLE OF CONTENTS page ACKNOWLEDGMENTS ..............................................................34 Comparative Gastric Evacuation Models for Gag Consuming Baitfish versus Crab Prey ...............................32 Gastric Evacuation Models for Gag Consuming Baitfish Prey ......1 METHODS......................................... xi CHAPTER 1 2 INTRODUCTION .................................................................................................................25 Energetic Models ...........................................................69 Effects of Prey Type............................................................................................................................................................................................................................................................ iv LIST OF TABLES....................................................................................................................................................................................................................................................................................................... ix ABSTRACT.22 Gastric Evacuation Models............21 Caloric Analysis of Prey and Stomach Contents.............................................................................................................................................................................................................................................................28 Models of Average Digestion Codes..................................40 Indices of Digestion for Crab Prey Consumed by Gag .............................19 Experimental Feeding Trials for Gastric Evacuation Rates .............................32 Gag and Prey Collections ..................................................................36 Energy Values for Recovered Gag Stomach Contents .....36 Caloric Values of Baitfish and Crab Prey........................................................32 Gastric Evacuation Models for Gag Consuming Crab Prey.74 v ..................................................................................................................................................................................................................................................................................................... vii LIST OF FIGURES ........................................................................................41 4 DISCUSSION................69 Gastric Evacuation Models..........................................26 Indices of Digestive States.......................

......................................99 vi ......................79 Stomach Content Composition ........................................................................90 LIST OF REFERENCES.............................................92 BIOGRAPHICAL SKETCH .......................................Effects of Predator Size ...................................................................................................................................................................................................................................83 Indices of Digestion.....86 Consumption...........................................................................................................................................................................................................................77 Prey Composition ........................88 Conclusions...................................................................................................

................ and large gag (n=8) .....................55 Regression parameters of the gastric evacuation dry weight data of gag consuming baitfish prey fit to each model................................................31 Regression parameters of the gastric evacuation wet weight data of gag consuming baitfish prey fit to each model..61 Regression parameters for models describing the gross energy (kcal/g dry weight) of the stomach contents as a function of post-prandial time (PPT) by all gag consuming baitfish prey fit to the linear................ for small gag (n=8)...... and large gag (n = 11)............. . medium gag (n = 16)...............................................................58 Regression parameters of the gastric evacuation dry weight data of gag consuming crab prey fit to each model.............................E.......................... medium gag (n=10).......... medium gag (n = 16)....... medium gag (n=10)..........................................).. medium gag (n=15).......................... and large gag (n=8) ... for small gag (n = 13)....................... for small gag (n=8).........................................................................................................59 Regression parameters of the pooled gastric evacuation data (n=26) of gag consuming crab prey on a wet and dry weight basis fit to the expanded power exponential models with either gag weight or TL scaling exponents .. exponential...................................57 Regression parameters of the gastric evacuation wet weight data of gag consuming crab prey fit to each model. for small gag (n = 13)..........................................30 Indices of crab Portunus gibbesii digestion by gag over post-prandial time (hr).................. ...........LIST OF TABLES Table 1 2 3 page Indices of baitfish Harengula jaguana digestion by gag over post-prandial time (hr)..........60 Composition of representative baitfish Harengula jaguana and crab Portunus gibbesii prey types used in gastric evacuation trials of gag. for small gag (n=12).................................................. and square root models.......... and large gag (n = 11)....62 4 5 6 7 8 9 10 vii .56 Regression parameters of the pooled gastric evacuation data (n=40) of gag consuming baitfish prey on a wet and dry weight basis fit to the expanded power exponential models with either gag weight or TL scaling exponents ... values are means (±S................. and large gag (n=11) .................

... and large gag (n=7)........... and square root models..67 Regression parameters of the digestion code data of gag consuming crab prey fit to each model............. medium gag (n=16).............11 Regression parameters modeling the gross energy (kcal/g dry weight) present over PPT by all gag consuming crab prey fit to the linear........... exponential.. and large gag (n=11) . for small gag (n=11)............................. for small gag (n=13).............63 Regression parameters modeling the percent of gross energy digested over PPT by all gag consuming baitfish prey fit to each model........65 Mean (±S..................... medium gag (n=16)............................................E. small gag (n=7)................................ medium gag (n=11)..... medium gag (n=7).................66 Regression parameters of the average digestion code data of gag consuming baitfish prey fit to each model..........................................................) and range of post-prandial times (PPT) in relation to digestion codes and % digestion for gag consuming baitfish Harengula jaguana versus crab prey Portunus gibbesi...... small gag (n=7).............. medium gag (n=8).... and large gag (n=5)..... for small gag (n=8)..... and large gag (n=8) ...................................64 Regression parameters modeling the percent of gross energy digested over PPT by all gag consuming crab prey fit to each model. and large gag (n=8)....................68 12 13 14 15 16 viii ...

...... medium............ .........LIST OF FIGURES Figure 1 2 3 page Commonly used gastric evacuation models depicting the digestion processes of different fish species........................................... ............................................ .................................... ........................... medium..... and large gag consuming baitfish prey (scaled sardines): (a) wet weight basis and (b) dry weight basis........44 The power exponential model describing the gastric evacuation processes of small.............48 The expanded power exponential model describing the combined gastric evacuation wet weight data of all gag consuming both baitfish (scaled sardines) and crab (Portunus gibbesii) prey incorporated with: (a) weight (W) and (b) total length (TL) scalers... ......................................................49 Models describing the gross energy of recovered stomach contents from small............................... fit to a square-root model and (b) crab prey (Portunus gibbesii)..................51 4 5 6 7 8 9 ix ........ and large gag consuming: (a) baitfish prey (scaled sardines).... fit to a linear model...45 The power exponential model expanded to include weight (W) or TL as scalers describing the combined gastric evacuation data of all gag consuming baitfish prey (scaled sardines): (a) wet weight basis and (b) dry weight basis..................... medium....................17 The relationship of gag weight (W) as a function of total length (TL) for gag between 300 and 750 mm TL................47 The power exponential model expanded to include weight (W) or TL as scalers describing the combined gastric evacuation data of all gag consuming crab prey (Portunus gibbesii): (a) wet weight basis and (b) dry weight basis.......46 The power exponential model describing the gastric evacuation processes of small.............................. ................... and large gag consuming crab prey (Portunus gibbesii): (a) wet weight basis and (b) dry weight basis.......50 The power exponential model describing the percentage of stomach content energy digested over elapsed time for small................... medium. and large gag consuming: (a) baitfish prey (scaled sardines) and (b) crab prey (Portunus gibbesii).........................

....................... medium.............. medium................................. ...54 11 12 x ...10 The power exponential model: (a) expanded to include weight (W) or TL as scalers describing the combined stomach content energy digestion data for all gag consuming baitfish prey (scaled sardines) and (b) describing the combined stomach content energy digestion data for all gag consuming crab prey (Portunus gibbesii)..........................................52 The power exponential model describing the average digestion code values of gag consuming baitfish prey (scaled sardines) over elapsed time: (a) small....................................................................... and large gag and (b) all gag fit to the expanded model using weight (W) or TL as scalers...........................................................53 The power exponential model describing the average digestion code values of gag consuming crab prey (Portunus gibbesii) over elapsed time: (a) small... ....................................... and large gag and (b) all gag fit to the expanded model using weight (W) or TL as scalers........

and the average digestion codes over PPT. exponential. requires prey-specific evacuation models. regardless of prey type or xi . especially off the west coast of Florida.0008) and best fit the wet weight and dry weight gastric evacuation data. the proportion of the gag meals. that remained after a pre-determined post-prandial time (PPT) was fit to linear. which feed primarily on fish prey and secondarily on crustacean prey. logistic. square root. Murie Major Department: Fisheries and Aquatic Sciences Gag Mycteroperca microlepis comprise one of the most valuable fisheries in the Gulf of Mexico.Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science GASTRIC EVACUATION AND DIGESTION STATE INDICES FOR GAG Mycteroperca microlepis CONSUMING FISH AND CRUSTACEAN PREY By Elizabeth Joanne Berens May 2005 Chair: Debra J. depends on the total amount of surplus energy available to the fish for growth after losses due to metabolism and wastes. as measured through growth. Estimating consumption rates of gag. Production of juvenile and prereproductive female gag. The power exponential models were significant (p 0. the percentage of prey energy digested over PPT. consisting of either baitfish (scaled sardine Harengula jaguana) or crab (purple swimmer crab Portunus gibbesii). on a wet weight and dry weight basis. and power exponential models. To develop these models.

87) gastric evacuation models. 2 =88. 2 =88. p <0. p<0. power exponential models fit to the crab data for each size of gag were coincident. these data were pooled and fit to an unexpanded power exponential model. Therefore. df=3.40. After correcting for energy from chitin. which is unavailable to the gag.0 hrs PPT in crab digestion only. df=3. Therefore.0001.0001) and highly predictive (R2=0. p<0. respectively).91).0001). t=-31.0001. proved significant (p<0. with W or TL scalers.gag size (R2 0. The power exponential model. digestion of crab prey was associated with a 5-6 hr lag period.98) for gag consuming baitfish when fit to the percentage of baitfish energy digested over PPT. and digestion over a longer period of time relative to fish prey. however. n=48. with lag phases of up to 5.22 kcal/g dry wt) was significantly lower than that of the baitfish prey (2-tailed Satterthwaite t-test for unequal variance.0001) and predictive model (R2=0. xii .0001 for each model) and highly predictive (R2 0. which also yielded a significant (p<0.81). dry weight. resulting in significant (p<0.26. n=66. total crab energy density (2. low prey energy densities. for gag consuming a mixture of fish and crab prey it will be necessary to develop a multiplicative or additive consumption model that also incorporates mixed-prey gastric evacuation models. and average digestion code data across all gag sizes by prey type. The expanded power exponential models with W and TL exponential scalers fit to the wet weight baitfish and crab gastric evacuation data differed significantly (Maximum Likelihood Ratio: n=66.05. Gag weight (W) or total length (TL) exponential scalers were incorporated into the power exponential model and fit to the wet weight. In general.

1998). p. and in 2001. 57660). 2003). One important grouper species in the Gulf of Mexico. 63 No. 1971). but are absent from most Caribbean waters (Smith. including fish. 1992. Ross & Moser. In recent years. Such studies are important for understanding the growth and 1 . such as groupers. grouper stocks have been overfished. late maturing reef fishes that consume a broad range of prey.8 million lbs) of gag were landed in the state of Florida alone (Turner et al.538 metric tons (7. 1998. Commercial and recreational catches in the southeastern U. Florida Fish and Wildlife Conservation Commission [FFWCC].. in part due to their slow growth and late maturity. and skewing traditional methods used to estimate their abundance (Federal Register.. Gag range from New York to Brazil and through the Gulf of Mexico. and cephalopods (Polovina & Ralston. 1987). 2001. Most species of groupers are demersal predators that generally occupy a top niche in the food chain of tropical and subtropical marine waters and have been recognized as slow-growing. Reef fish. 208. Collins et al. crustaceans. thereby making the species vulnerable to overfishing. Within the past decade. the gag Mycteroperca microlepis supports valuable recreational and commercial fisheries. tend to aggregate over specific habitats characterized by a patchy distribution.S. have exceeded 2. 1995. over 3. Vol.268 metric tons (5 million lbs) annually. many studies have examined the life history of gag and the effect of fishing on its populations (Hood & Schlieder.CHAPTER 1 INTRODUCTION Groupers (Pisces: Serranidae) support major commercial and recreational fisheries throughout the world.

including diet and activity demands. A generalized energy budget can be modeled as (Winberg. growth. Ma = metabolic rate increase above the standard rate due to activity. equals the energy lost to metabolism (Mr + Ma + SDA). and for helping managers cope with a variety of issues. and Gr = growth rate due to gonad (reproductive) synthesis. 1990). wastes (F + U). 1990. This generalized energy budget is a balanced equation where all energy consumed by the animal (C). and analyzing exploitation issues. 1993). Warren & Davis. An energy budget relates all the energy an organism acquires through ingestion to its energy used in metabolic processes. such as setting catch limits.2 production of the gag fishery. and growth (Gs + Gr). lost as wastes through excretion and egestion. F + U = waste losses due to egestion (feces) and excretion (urine) rates. Jobling. energy for growth is only available after all metabolic and waste demands have been subtracted from the total amount of energy consumed: Gs Gr C Mr Ma SDA F U (2) Due to the equation being theoretically balanced. SDA = metabolic rate increase due to specific dynamic action. Energy budgets can be used to evaluate the importance of many different factors controlling individual growth. or synthesized into new somatic or reproductive tissue (Adams & Breck. is an integral part of a fish’s energy budget. size restrictions. 1956. One of these topics. Hence. 1967) C Mr Ma SDA F U Gs Gr (1) where C = rate of energy consumption. Mr = standard metabolic rate. subject to different environmental conditions (Adams and Breck. researchers have the advantage of estimating one component of the energy budget by subtracting it from the other measured . Gs = somatic growth rate due to protein synthesis [and lipid deposition].

which quantifies the rate at which food passes out of the stomach (Adams & Breck. and even physical parameters. or the rate of gastric evacuation. abundance and distribution of prey. others in the lab. that predator’s evacuation rate will tend to be lower than if that fish had consumed a small single meal of the same food (Swenson & Smith. the types of prey consumed. 1990. Some of these complex factors are best studied in the field. and the predator’s digestion rate. 1978). 1990). which may or may not be feasible depending on species. and growth all depend on the amount of energy consumed. and hence accurate estimates of consumption rates are imperative. even though they are one of the most direct “inter-links” between the trophic components in an ecosystem (Klekowski & Duncan. Consumption rates are typically difficult to estimate because of many complex factors that influence the amount of food that a fish consumes. but some require the capture of large numbers of fish. however. the size. these factors can affect the number of meals consumed per day. waste. 1994). such as water temperature (Windell. although this method tends to pool all error into that one component (Adams & Breck.3 components. most studies take into account only two or three of these interrelated factors (Windell. including: the fish’s size. Consumption rates can partially affect gastric evacuation rates by causing an increase or decrease in the amount of time food remains in the gut. 1978. the amount of food consumed in a single meal. if a fish consumes a large single meal of a certain prey type. 1975). the predator’s feeding history. Total metabolism. Adams & Breck. Bromley. For example. . Consumption rates. 1973). 1990). Considering the many different biotic and abiotic factors that influence consumption and the fact that consumption rates are difficult to quantify outside of the laboratory. In addition. are often the least developed part of the energy budget.

1970. or the time between two sampling periods: Ct ( St So e 1 e kt kt ) kt (4) where So = amount of food present at the beginning of the sampling interval. 1978. or more . 1990). the Continuous Feeding Model involves sampling a population over a certain amount of time. Grove et al. (1978) found that an increase in the rate of gastric evacuation occurred with the consumption of low-energy food and was correlated to a rapid return of appetite and a high frequency of consumption. which directly incorporates the rate of gastric evacuation and can be modeled as C Ct (3) where C = daily ration (% body weight/day). some models require estimates of the original prey weight at time of capture. and then incorporating the length of the sampling period and the instantaneous rate of gastric evacuation into the model (Adams & Breck. and Ct = the amount of food ingested in a block of time. or the sum all Ct values in each time block. 1978). Although consumption affects rates of gastric evacuation. and k = instantaneous rate of gastric evacuation.. For example. Grove et al. calculated as k loge So St t 1. Elliott & Persson. calculating the amount of food in the gut at the beginning and end of the sampling period. St = amount of food present at the end of the sampling interval.4 Most consumption models require the calculation of gastric evacuation rates and. Elliott and Persson (1978) developed a commonly used Continuous Feeding Model for modeling consumption rate in brown trout Salmo trutta. additionally. gastric evacuation rates can also affect the amount of food a fish consumes (Brett & Higgs. t = length of the sampling interval.

research using consumption models has depended more heavily on rates of gastric evacuation than research in other fields. Butler et al.. 2004. such as animal nutrition. Digestion is the act of mechanical and enzymatic breakdown in the fish’s stomach that converts food into soluble and diffusible products capable of being absorbed. by cells in the fish’s stomach and intestine (Knutsen & Salvanes. Sponheimer et al. 2004). In fisheries.e. such as mammals (Stevens & Hume.5 meals consumed. Conversely. estimating food consumption in many animals requires indirect methods.. Hence. 1978). 2002.. or assimilated. gastric evacuation rates impact consumption and therefore they must be included in consumption models in order to calculate accurate estimates of total energy intake. 1995) and food consumption estimates are usually determined through direct gut content analysis using lavage techniques or by sacrificing large numbers of animals (Adams & Breck. Clearly. The terms digestion rate and gastric evacuation rate are often used interchangeably to denote the rate at which food passes from the stomach into the intestine even though this is something of a misnomer (Windell. Henriques et al. 1999). 2003. 1990).. and upon defecation are often referred to as indigestible matter. Roxburgh & Pinshow. Materials that can’t be absorbed are simply passed through the body. 1967). food that has been broken down and is no longer present in the stomach (i. such as inferring feeding rates of wild animals from feeding rates of .. In general. the alimentary tracts of fish are much simplier than the alimentary tracts of other animals. 1997. food that has been evacuated from the stomach) does not necessarily represent food that has been assimilated completely in the pyloric cecae (possibly) or intestine and utilized by the fish (Pandian. which often focus more on the rate of transit through the entire gastrointestinal track (Dorcas et al.

which tend to consume only a few fairly large prey items over a feeding cycle. including using X-ray and radioisotopic procedures and through serial slaughter (Beamish. 1990). Therefore. The fraction of original prey weight digested is plotted against hours post-feeding.. Serial slaughter methods in situ are generally not feasible since it is very difficult to get large groups of wild fish to simultaneously consume a measured amount of food. The slope of the resulting regression is then used to calculate the gastric evacuation rate and determine the gastric evacuation model (Adams & Breck. 1979. 1973. Flowerdew & Grove. A linear model often describes the gastric evacuation processes of top carnivores. Flowerdew & Grove. 1979). and then are either input directly into. thereby causing relatively long digestion times compared to the length of their feeding . 1977.6 captive animals (Innes et al. this method is usually used on captive fish where specific amounts of food can be given to individual fish. 1979). 1978). and their stomach contents easily recovered by killing the fish or by pumping its stomach (Adams & Breck. because lavaging certain types of animals or sacrificing large numbers of those animals may be logistically difficult. Swenson & Smith. 1990). prohibited. or simply undesirable (Winship et al. Diana. or used to meet the assumptions of a consumption model (Figure 1). 1987) or through bioenergetic modeling. and at predetermined times collect their stomach contents (Windell.. the fish can be monitored.. and square root gastric evacuation models have commonly been used to quantify the gastric evacuation processes of fish species. hold them in pens. There are several ways to estimate gastric evacuation rates. 2002). or piscivores. 1972. X-ray and radioisotopic procedures have been used most often to monitor the movement of hard parts through the entire digestion process (Jobling et al. Linear. exponential.

low fragmentation rates. Problems with the exponential model occur when the lower portion of the curve levels off and reaches its lower asymptote because it leads to overestimates of the amount of food remaining in the stomach at later stages of evacuation (Brodeur & Pearcy. Adams & Breck. 1987. and omnivores that feed at lower trophic levels on diets composed of many small food items. 1987). detritivores. 1990). and lowering the animal’s total food consumption rate (Rindorf. high fragmentation rates. 2002). These food items would be expected to have high surface-to-volume ratios. Due to the fact that the exponential model levels off. This slowing of gastric evacuation means that the fish’s motivation to feed would return more slowly. 1990). These types of prey items tend to be evacuated from the stomach at a constant rate. 1990). Previous work with piscivores and linear digestion processes have included studies on black and yellow rockfish Sebastes chrysomelas (Hopkins & Larson. plaice Pleuronectes platessa (Jobling. such as zooplankton. the digestion rate increases exponentially with time until some point at which refractory materials slow the rate and basically level off the digestion process. 1973). Jobling (1987) attributes the linear model to large food particles with lower surface-to-volume ratios. In these fish. and high dietary energy densities.7 period (Adams & Breck. . and tend to be low in energy densities (Jobling. and walleye Stizostedion vitreum vitreum (Swenson & Smith. planktivores. 1990). thereby increasing the amount of time between feedings. 1980b). The exponential model generally illustrates the digestion process of herbivores. and eat more or less continuously throughout the day (Adams & Breck. Brodeur and Pearcy (1987) considered the active part of the curve to be between 0 and 90% evacuation.

Brodeur (1984) suggested that an alternate way to choose the most appropriate evacuation model may be whether or not it is interpretable in terms of the inherent biological processes that occur during digestion. 1979. 1981). Temming & Andersen. t = elapsed time after ingestion. R = estimated parameter(s). 1982. 1984. 1990) (Figure 1). temperature. and meal size as predicting variables (R parameter) to determine values of a shape parameter (B) as dS / dt RS B (5) where S = residual stomach contents (g).8 The square root model has been used to describe digestion in predators such as plaice. theoretically. Atlantic cod Gadus morhua. and B = shape parameter. 1981). and bluegill sunfish Lepomis macrochirus (Jobling & Davies. Jobling. 1992. Hopkins & Larson. Recently. Hopkins & Larson. This shape parameter describes the degree of . 1994) (Figure 1). therefore. The power exponential model has also been used to describe predators with and without lag phases because it allows the shape of the evacuation curve to vary from sigmoidal to concave.. This volume-based model assumes that the instantaneous rate of evacuation is dependent upon the amount of food in the stomach. European perch Perca fluviatili. although it can not account for linear rates of digestion (Elashoff et al. The logistic model fits this idea considering that it accounts for the lag phases often seen in the early stages of digestion of many different fish species (Brodeur. Temming and Andersen (1994) have developed a general gastric evacuation model that integrates time after ingestion. evacuation patterns of small meals correspond to the later stages of large meal evacuation patterns and. 1990. dos Santos & Jobling. results in regression lines for different meal sizes having the same slopes (Jobling. the weight of the predator.

when in fact studies have shown that at high temperatures the evacuation rate will decrease (Tyler. Considering that exact times of 100% evacuation can not be determined from empty stomachs. exponential. 1970.9 curvilinearity. In effect. 1986). For example. Temming and Herrmann (2001) determined that B=0.7 on a dry weight basis for horse mackerel Trachurus trachurus thereby indicating that the model was more exponential than linear. whether it is convex. thereby resulting in biased evacuation rate estimates with exaggerated curvilinearity (Olson & Mullen. Temming & Andersen. linear. 1994). the distribution includes both faster and slower digesting fish throughout most of the distribution but only the slower digesting fish at the later time periods. an increasing proportion of the fish with faster digestion are typically excluded from the sample distribution. which truncates the sample distribution and reduces bias . 1986). 1986). excluding all of the faster digesting fish from the gastric evacuation distribution may have led many studies to choose exponential gastric evacuation models rather than square root or linear models (Olson & Mullen. the evacuation model is exponential. In part. 2001). If B=1. or any curve in between (Temming & Herrmann. As post-prandial time (PPT) increases. gastric evacuation models often include times to 90% or 95% digestion only. 1994). but B=0 indicates a linear evacuation process (Temming & Andersen. Empty stomachs are normally dropped from the distribution because it cannot be determined when 100% digestion occurred (Olson & Mullen. This general model of gastric evacuation is only valid within a limited temperature range because it assumes that the evacuation constant (R) increases exponentially with temperature.

Many environmental. 1979. therefore. and meal sizes (Bromley. thereby exhibiting an increased digestion rate (Flowerdew & Grove. 1980b. 1995). 1993). Additionally. prey sizes. thereby having another very important influence on consumption and digestion rates. prey. High-energy fats tend to slow gastric emptying more than . can vary greatly with prey type and prey size. such as enzyme production and kinetics. and. Jobling. 1970. or energy density. Most fish are ectotherms. and predator characteristics influence both consumption and gastric evacuation rates by either speeding or slowing food digestion. Hopkins & Larson. Temperature is among the most important environmental variables that influence consumption and digestion rates in fish. 1994). Previous studies have reported that consumption rates and digestion rates increase with rising water temperatures (Brett & Higgs. Many studies on fish have found that meals high in energy result in an increase in time to 100% gastric evacuation (Flowerdew & Grove. Body temperatures and digestion rates can therefore vary considerably throughout the year. 1990). the surrounding water temperature determines their body temperature (Hazel. 1973. including temperature.10 from fitting models to data that may include empty stomachs (Swenson & Smith. 1980a). are therefore directly related to water temperatures (Diana. predator sizes. Hopkins & Larson. He & Wurtsbaugh. Jobling. diets with an added diluent. prey type. Metabolic rates of fish and their corresponding physical and chemical processes. Meal composition. if temperature fluctuates seasonally. 1990). a non-digestible marker that lowers a meal’s energy density. 1993). Studies dealing with these two different rates must take many different conditions into account. were evacuated more rapidly from fish stomachs than those with higher energy content. 1979.

Therefore. which may be high in protein. not only do mature individuals contain more lipid.11 proteins or carbohydrates (Jobling. 1982. but they also have a lower surface area to volume ratio which slows gastric evacuation rates even more. lipid-rich mature prey are physically larger than immature prey of the same species. commonly report the means or the ranges of prey energy densities on an ash-free basis because these values exclude all inorganic elements.. NAG or D-glucosamine. Therefore. or ash. have been shown to remain in the stomach for long periods of time compared to food with fewer indigestible hard parts. The least digestible. and therefore. Crustacean exoskeletons. which is not a source of energy for the predator (Brett & Higgs. a carbohydrate (polysaccharide) that most fish breakdown to N-acetyl-D-glucosamine (NAG) and D-glucosamine and pass out of the body (Jackson et al. 1935. 1973).. mature prey fish high in lipid content will slow digestion compared to juvenile prey fish. Hopkins & Larson. fish are not known to assimilate chitin. 1970. must be taken into account when determining the energy density available from . 1985. 1980a). However. unavailable energy locked up in the crustacean’s exoskeleton. including crustaceans. in particular. which slows gastric evacuation rates. MacDonald et al. Clearly. unlike baitfish prey. Medved. 1990). such as fish prey (Hopkins & Larson. chitin contains energy that is unavailable to the predator (Battle. as well as inorganic ash. Additionally. 1990). 1992). Lindsay & Gooday. 1979. Beamish. any increase in prey size decreases the surface area to volume ratio available for enzymatic digestion (Swenson & Smith. 1972). While. but low in lipids. crustacean exoskeletons contain chitin. hard skeletal elements of prey are generally low in energy and are often the last part of a meal to be emptied from the stomach (Flowerdew & Grove. 1985). Gastric evacuation studies using baitfish and invertebrate prey. generally.

For example. is a proxy for fitness (Krebs & Kacelnik. 1978. The effect of predator size on gastric evacuation rates must be considered when using standardized meals to prevent an underestimation of consumption for small fish and an overestimation of . On the other hand. Predator size is another factor that can affect consumption and digestion models. MacArthur & Pianka. This idea originates from Optimal Foraging Theory. Jobling. fish often appear to maintain a relatively constant energy intake for metabolic function and growth. which states that animals will maximize their food (energy) intake per unit of time or minimize the time required to meet their energy requirements (Emlen.12 crustacean prey. 1980b. 1980a). As the availability of a gag’s food resources decreases. Schoener. 1966. 1991). Absolute gastric evacuation rates (grams of food leaving the stomach per hour) tend to increase with increasing predator body size. Jobling. but relative rates (per unit body weight) either decrease with increasing predator size or stay the same (Flowerdew & Grove. a 500 g grouper fed a 15 g meal would have a slower absolute gastric evacuation rate than a 1000 g grouper fed an identical 15 g meal because the smaller grouper would be consuming a much larger meal relative to its body size. if food resources are lower in energy then predators will tend to eat more. Bromley. For example. 1994). 1966. Optimal Foraging Theory assumes that the rate of energy intake. a 500 g grouper fed a 10 g meal (meal = 2% of grouper’s body weight) would have a faster relative gastric evacuation rate than a 1000 g grouper fed a 20 g meal (meal = 2% of grouper’s body weight). but if prey are high in energy then they eat less and maintain similar energy consumption levels (Grove et al. the gag’s dietary niche breadth should expand to include lower energy prey and increase the chance of prey encounters.. 1979. 1971). or foraging success. In a form of self-regulation.

Swenson & Smith. 1994).13 consumption for large fish (Adams & Breck. although some have shown negative correlations or no correlations at all (Bromley.. Both predator weight and length has been used when quantifying the influences of body size on gastric evacuation rates. 2002): .. Like gastric evacuation estimates. One method involves calculating the original prey weights based on vertebral column length. Differing definitions of gastric evacuation rates. Ruggerone. Lindberg et al. 1977). Studies have determined relative gastric evacuation rates most often (Beamish. and maximum length regressions. expressed as Absolute Rate = Relative to Body Size = Weight of Food time (6) Weight of Food Fish Body Weight time 1 (7) have led to these conflicting conclusions and have made comparisons of different evacuation results problematic (Bromley.. dos Santos & Jobling.. MacDonald et al. 1972. 1994). standard length. it has been shown that an increase in meal size leads to an increase in the rate of gastric emptying and an increase in time to 100% evacuation (Jobling et al. depending on season and individual growth (Koed. 1982. length because it is easily measured and predator weight because it can vary much more than length. 2001). Many studies have shown positive correlations between gastric evacuation rates and meal size. 1992). In general. Adams et al.. and backcalculating the time of prey ingestion based on its stage of digestion (Minton et al. 1989. 1990). 1973. 1982. 1981. including absolute rates and rates relative to body size. calculating the original time of prey ingestion by a wild predator and incorporating this required variable into a consumption model has been difficult.

An index of digestion may be a faster method due to the fact that a general time of prey ingestion can be estimated based on the prey’s appearance without having to back-calculate its size through regression analysis. Weaver (1996)... Several factors affecting this assumption of linear digestion include the type of prey gag consume and the amount of energy within that prey. but it may not be as accurate a method since it can only provide a general.. 1995). rather than specific.. gastric evacuation rates have only been estimated from field collections of gag consuming baitfish from artificial reefs off the west coast of Florida during the warmer months of the year (Lindberg et al. time after feeding. Both methods provide inaccurate estimates of ingestion times at later periods of digestion due to the fact that the prey eventually become unrecognizable to species and tend to have broken or missing vertebral columns. time to 90% evacuation has been estimated at 15 hours and 100% evacuation at 16 hours for baitfish prey (Lindberg et al. gag off the west coast of Florida were observed to have a significant proportion of crabs in their diet (13-14%) during the warmer months of the year . 2002). Kao.1% index of relative importance for crabs in the stomachs of gag between 300 and 400 mm standard length (374-489 mm TL). with invertebrates accounting for no more than 5% of the total food volume in their stomachs (Naughton & Saloman. noted a 17. Through preliminary field estimates based on back-calculated original prey weights and an assumed linear model. In addition.Digested Weight of Prey Original Weight of Prey (8) Another method involves creating a quantitative and qualitative visual index of digestion stages using numerical codes and prey descriptions (MacDonald et al. Currently. 1982. 2002). however.14 Stage of Digestion = 1 . 2002). 2000. Lindberg et al. Gag are commonly considered to be highly piscivorous.

crab prey are less calorie dense than baitfish prey because they commonly contain less lipid but also because the chitin in their exoskeletons contain energy that is unavailable to most fish (Battle.. Lindberg et al. 1990).. thereby lowering the frequency with which gag would consume meals. Medved. 1985. These proportions contribute between 7% and 24% of the total diet on a gross energy basis. 2002). 1982.. and less energy than baitfish prey.000 kcal/g dry weight (Lindberg et al.15 (Lindberg et al. Hard materials. there have been no studies that have attempted to correct for unavailable energy contained within the chitin of crab prey exoskeletons. 2002). followed by a rapid increase in evacuation rate and a subsequent leveling off as meal remnants are retained in the stomach (Hopkins & Larson. have been shown to cause significant lag times in digestion. with whole crab caloric densities estimated as 1. changes in digestion rates can lead to a slower return of appetite. more ash. MacDonald et al. Generally. 1985). To maintain growth rates similar to gag consuming baitfish. gag consuming crab prey must compensate for their reduction in rates of gastric evacuation and feeding frequency by consuming more crabs per meal. 1935. Considering that crabs contain unabsorbable chitin in their exoskeletons. The hard chitinous material found in crab exoskeletons should have a strong influence on the gag’s digestion rate. Interestingly. scaled . juvenile tomtate Haemulon aurolineatum. As discussed earlier. To date. such as crab exoskeleton. Lindsay & Gooday. (2002) have shown that gag on artificial reefs (Suwannee Regional Reef System) off the west coast of Florida are consuming baitfish prey (round scad Decapterus punctatus. gag consuming higher percentages of crab in their diet should consume prey less frequently and may have less energy available for growth after their metabolic and waste removal energy needs have been met.

. The specific objectives were: (1) experimentally determine gastric evacuation rates for gag as a function of prey type. Spanish sardine Sardinella aurita) that are abundant but not necessarily high in energy (1. depending on species) compared to relatively lipid-rich adult baitfish or other fish species that often are 2-3 times more calorie dense. especially for portunid crab prey. (3) create qualitative and quantitative indices of prey digestive states in order to estimate consumption times of prey sampled from the stomach contents of wild gag. (2) compare the effect of prey type (baitfish versus crab) on the gastric evacuation rates of gag. either baitfish or crab prey. The overall goal of this study was to develop gastric evacuation models for gag consuming baitfish and crustacean prey. and (4) model evacuation rates of gag and compare the models for best fit by prey type and gag size. Lindberg et al. in relation to gag size. The organic composition of individual prey (in terms of energy content) and the varying amounts of different prey items in the diet of wild gag during the warmer months of the year may therefore influence digestion rates significantly. evacuation models specific to prey type and fish size are currently unavailable for gag.16 sardine Harengula jaguana. However. (2002) suggested that estimates of consumption rates of wild gag may be improved with additional knowledge of prey-specific evacuation models.03-1.14 kcal/g wet weight.

17 100 80 60 40 20 0 0 2 4 6 Linear Model Y Where. A Bt Y = % prey remaining in the stomach A = Y-intercept B = gastric evacuation rate t = elapsed time after ingestion 8 10 12 14 16 18 Elapsed Time Since Feeding 100 80 60 40 20 0 0 2 4 Exponential Model Y Where. A Bt Y = % prey remaining in the stomach A = Y-intercept B = gastric evacuation rate t = elapsed time after ingestion 4 6 8 10 12 14 16 18 Elapsed Time Since Feeding Figure 1. Ae Bt Y = % prey remaining in the stomach A = Y-intercept B = gastric evacuation rate t = elapsed time after ingestion 6 8 10 12 14 16 18 Elapsed Time Since Feeding 100 80 60 40 20 0 0 2 Square Root Model Y Where. Commonly used models depicting the gastric evacuation processes of different fish species. .

Y = % prey remaining in the stomach A = half life of decaying prey B = shape coefficient t = elapsed time after ingestion and dashed lines show the potential variability in the shape coefficient 12 14 16 18 Elapsed Time Since Feeding Figure 1. 100 A 1 e B t C Y = % prey remaining in the stomach A = estimated parameter B = scale parameter C = x-ordinate of the point of inflection of the curve t = elapsed time after ingestion 100 80 60 40 20 0 0 2 4 6 8 10 Power Exponential Model Y 2 t B A Where.18 Logistic Model 100 80 60 40 20 0 0 2 4 6 8 10 12 14 16 18 Elapsed Time Since Feeding Y Where. (continued) .

46 m high by 0. and chlorine neutralizer) were used as necessary to ease stress and bring the gag back in the best possible condition. Small (300-449 mm total length.5 hrs of darkness per day.71 W).9 m. The gag were lifted to the surface in the traps. a series of artificial reefs off the west coast of Florida (28°59. Light regimes averaged 13. and ChlorAm-X (an ammonia. chloramines. a 189.. at a depth of 13 m (40 feet) (Lindberg et al. and featured an opening approximately 0.10 W to 29°20.. 2002) between February 2003 and January 2004.0 m by 0.91 N. Grouper traps measured approximately 1.3 L bead filter. and large (600750 mm TL) gag were trapped by SCUBA divers or caught underwater on hook-and-line using baited hooks. were constructed out of plastic-coated wire. and a mesh bag filter covering the tanks’ drainage pipe within the lower sump tank.30 m wide. Gag were held in 378 L (87-cm diameter) to 473 L (147-cm diameter) fiberglass tanks in a recirculating saltwater system of approximately 4. 83°19. TL). While in transport to UF. 2002). and they were transported 101 km to the University of Florida’s (UF) Fisheries and Aquatic Science’s aquatic facility using aerated coolers with 100% diffused oxygen supplied through carbon stones. their air bladders were vented (Lindberg et al. 83°31.CHAPTER 2 METHODS Gag Collections and Maintenance Gag were collected from the Suwannee Regional Reef System. water changes. Filtration equipment included a 187 L sand filter.06 N.163 L. following the photoperiod of the eastern Gulf of 19 .5 hrs of light and 10. medium (450-599 mm TL).

. and salinity (30-35 ppt) parameters were kept within acceptable limits for marine fish by weekly tests. 2002). 2002).. 2000. Monthly water changes replaced between 10-15% of the system’s total water capacity. All gag were measured for maximum TL. 1990). Fish exhibiting any health problems were anesthetized with Tricaine-S (Methanesulfonate. Pieces of PVC cut 45-cm long with 20-cm diameters were placed in the tanks to provide shelter for the gag. Gag were maintained individually or in small groups of 2-3 individuals at a mean water temperature of 28.2 and 1.8% body weight (Lindberg et al. Small transitional lights mimicked dusk and dawn periods. along with supplemental water changes.0-0.20 Mexico during the warmer months of the year (~May to November).2 m (40 ft) depth where the gag were collected (Bledsoe & Phlips. ChlorAm-X was used as needed. nitrite (0.3 mg/L). Ammonia (0. while nitrate levels (030 mg/L) were tested monthly (Stickney & Kohler.3). Only fish feeding voluntarily on the maintenance diet were used in feeding experiments.0oC (+ 1.0% body weight every other day in the morning or evening. or MS-222).0-0. to neutralize any ammonia spikes. Phlips & Bledsoe. Gag that did not feed within 1 week of initial capture were anesthetized with Tricaine-S and tube-fed a slurry of ground fish to stimulate subsequent natural feeding. and weighed within 2 weeks of capture.5 mg/L). their bodies scraped and their gills and fins clipped for microscopic analysis and possible bacterial. Gag were fed on maintenance rations of thawed. or parasitic identification. fungal. turning on or off 1 hr before or after the main lights.0 oC). fork length (FL). whole baitfish or crab prey at 3.0-8. which corresponded to the average temperature during the warmer months of the year at 12. pH (8. based on current estimates of baitfish prey average daily consumption during the warmer months being between 1.

Crab prey were measured for carapace length (CL). FL (29°25. and frozen until use.. Gag were acclimated for a minimum of 2 weeks or until they voluntarily fed on baitfish or crab prey. vertebral column length (VCL: measured from the atlas/axis to the hypural plate). all skin and fins intact in the sardine prey and all legs attached in the crab prey). carapace width (CW). sardine prey were thawed and then measured for TL. and frozen. Stomach contents were . Crabs were immediately placed on ice for transport.10 N. were caught by underwater cast nets towed by SCUBA divers over reef areas occupied by gag during the Fall of 2002 and 2003. in the eastern Gulf of Mexico during the spring of 2003 and 2004.30 W to 29°30. and weighed.. They were immediately placed on ice for transport to UF. also a potential gag prey item (Lindberg et al. 1990).21 Experimental Feeding Trials for Gastric Evacuation Rates The model pelagic baitfish species and potential gag prey. Portunid crab prey Portunus gibbesii. were caught in standard shrimp trawls at night off Horseshoe Beach or Keaton Beach. medium. 83°25.5% of the grouper’s body weight on a wet weight basis were fed to individual small. Only completely whole scaled sardines or portunid crabs were used in the feeding trials (e. 1987. FL.g. the scaled sardine. and large gag. 2002). Prior to consumption by the gag. Hopkins & Larson. vacuum-sealed with a FoodSaver 550. Acclimation periods for fish in previous studies have varied from 2 to 6 weeks (Brodeur & Pearcy. with the help of commercial bait-shrimp fishermen. and weighed. vacuum-sealed while being covered with bubble wrap to prevent their spines from cutting the sealing bag. Food was withheld for 1-2 days before baitfish feeding trials and for 2 days before crab trials to insure that all maintenance rations had been evacuated from the gag’s stomach. Only trials where all of the prey were consumed were included in this study. 83°15. Meals weighing approximately 1.50 N.60 W).

CW.22 recovered via serial slaughter after various time intervals. A serial slaughter method was necessary since previous lavage techniques have only been between 65% and 77% complete.0001 g. opening up the stomach from the esophageal opening to the pylorus. without scraping so hard as to remove large amounts of mucus off the stomach lining. The stomach contents were recovered by removing the entire gastrointestinal tract from the esophagus to the anus. pieces of crab exoskeleton. Each individual prey was then chopped up. VCL. 2002). large whole fish. and weighed (g). After a predetermined time interval. Wet weight of the recovered stomach contents was taken immediately by placing the contents on a damp sponge covered with a damp Kimwipe (paper tissue) to remove excess water.. The % wet weight of the stomach contents remaining in the gag’s stomach was calculated as % Wet Weight = Wet Weight of Stomach Contents (g) Remaining Wet Weight of Items Consumed (g) x 100 (9) Caloric Analysis of Prey and Stomach Contents To determine the total caloric density of representative prey types. and gently scraping the contents out. samples of whole scaled sardines and portunid crabs were measured for TL.18 hrs (5 min) and 24 hrs. respectively. transferring the stomach contents from the Kimwipe to a preweighed weigh boat. leaving whole crabs. FL. .08 and 925. gag were sacrificed by applying a brain-spinal pith (American Fisheries Society [AFS]. or CL. Association of Official Analytical Chemists [AOAC] 1990). Freeze dried prey were then individually ground in a Braun coffee grinder until homogenized. and then weighing the contents to the nearest 0. 2004). and % moisture content determined (methods 926.09. and weighed (g). freeze-dried to a constant weight. and pieces of fish vertebrae and spines behind (Lindberg et al. ranging between 0.

For this correction.04-1.23 Additionally. some crab prey had to be chopped or cut by hand using small dissecting scissors to fully homogenize the remaining hard pieces of exoskeleton. The ash-free caloric densities of the initial whole baitfish and crab prey were calculated in kcal/g dry weight by dividing the total available dry weight caloric value by the ash-free dry weight: Kcal/g Ash-Free = Total Energy kcal/g Dry Weight Dry Weight Ash-Free Dry Weight (11) Due to the fact that portunid crabs contain chitin in their exoskeletons. Individual prey weighing <0.70 g dry weight were spiked using benzoic acid tablets to facilitate burning and gross heat determinations. Standardized corrections to gross energy were made for fuse wire burn (15.57 g dry weight sample) in a muffle furnace for 12 hours at 450oC to determine their % ash-free dry weight (g) (method 923. were measured for CL and CW and weighed (g). mean energy densities. To correct for inorganic materials within each individual. IL). Soft body tissue was removed by cutting the . and % ash content between baitfish and crab prey were determined using a 2-tailed Satterthwaite t-test for unequal variances after a determination of homogeneity of variance was made using Levene’s Test ( =0. Moline.0 mm) and acid production (10. the percentage of ash was determined by ashing a subsample of individual freeze-dried baitfish and crabs (0.70-0.0 ml). whole crab prey. throughout a size range.03. energy density determinations were used to correct for the unavailable energy contained within the chitin component.05).99 g sample) were then determined using an isoperibol bomb calorimeter (Parr 1261. Energy densities in kcal/g dry weight of each individual prey type (0. AOAC 1990): % Ash-Free = (Tissue Dry Weight – Tissue Ash Dry Weight) x 100 Dry Weight Dry Tissue Weight (10) Differences in % moisture content.

The regression estimates of energy in the chitin exoskeleton of the initial whole portunid crab were then subtracted from the crab’s total energy density.24 body and legs length-wise and placing the crab in a Pyrex beaker containing 15% KOH solution. the residual chitinous exoskeleton from each crab was placed against a light box (a lit background) to check for any remaining tissue. and freezedried to a constant weight. The exoskeleton was then ground in a coffee grinder and chopped or cut by hand using small dissecting scissors until homogenized. Initial whole crab wet weights (W). were regressed as a function of their exoskeleton’s total energy density (kcal/g dry weight) and the resulting linear regression was used to estimate the amount of unavailable energy in the chitin from the initial whole crab prey that had been used for % moisture and ash determinations. resulting in the total available energy density in kcal/g dry weight of each initial individual whole crab. the exoskeleton was weighed for damp wet weight (as before). If any crab tissue was remaining the crab was placed back in the 15% KOH solution for another 2-3 days or until it was free of all tissue. The KOH solution digested all protein completely through alkaline hydrolysis while excluding chitin.4-linked polysaccharide (Pandian. To determine the caloric densities of the individually recovered stomach contents. The energy density of each exoskeleton was determined in the same manner as the whole baitfish and crab prey. After approximately 1 week. 1967). a large beta-1. frozen (-80ºC). To determine the caloric density of the individual chitinous exoskeleton. after which the exoskeletons were rinsed with distilled water. the recovered stomach contents were freeze-dried to a constant weight. The stomach contents were then homogenized and a subsample was analyzed for caloric density (as . before the KOH treatments.

In the power exponential model. exponential. the total available crab energy density per meal in kcal/g dry weight was estimated by back-calculating and then subtracting the average exoskeletal energy density of the entire meal (using the previous regression of individual whole crab W plotted as a function of gross exoskeletal energy) from that meal’s total energy density. 1986).. or the first feeding trials resulting in 0% remaining at PPT. increasing the proportion of fish with faster digestion that are eliminated from the distribution (Olson & Mullen. Hopkins & Larson. 1990). fish or crab) (kcal/g dry weight) was multiplied by the estimated dry weight of the meal fed to determine the original caloric energy content of each meal fed to each gag. 1990). and (3) had lower asymptotes showing less than 5% prey remaining (Hopkins . and power exponential evacuation models were fit to the wet weight and dry weight gastric evacuation data (% remaining in the stomach) separately for each prey type and gag size in order to model the percentage of food remaining with PPT (Hopkins & Larson. as well as to prevent biases from including zeros past endpoints. were included in the data set for each gag size class in order to prevent biases associated with excluding all zeros..e. and thereby. For energetic regression analyses. all Y-intercepts had to be multiplied by 100 to be comparable with the outputs of other models (Elashoff et al. Gastric Evacuation Models Linear. (2) had yintercept values (estimates of the % prey remaining at time 0) between 95-105% prey remaining. Only initial zeroes.25 per previous samples). A model was considered adequate if it: (1) showed homoscedasticity of variances. therefore. the percentage of food remaining in the stomach was divided by 100 in order to fit the proportion of food remaining in the stomachs to the model and. For stomach contents containing crab prey. square-root. logistic. the average caloric content of each meal type (i. 1982.

1999) R2 1 RSS CSS Total (12) where RSS = residual sums of squares and CSSTotal = corrected sums of squares total. Energetic Models The linear. 2001). medium. and power exponential models were fit to the gross energy (kcal/g dry weight) of the stomach content data.05) (Kimura. Haddon. 8. for each size of gag consuming either baitfish or crab prey in order to model the gross energy of the stomach contents as a function of PPT. exponential. 1982. 1999). the data were pooled by prey type and modeled with either a gag weight (W) or TL scaling exponent to create prey-specific models of gastric evacuation that account for differences in gag size as Y Y function W C function TLC (13) (14) where Y = % prey remaining in the stomach. 1990.among gag size groups. differed significantly). SAS v. Differences among gastric evacuation models for small. The baitfish and crab prey data sets were truncated as all recoverable stomach content energy values remained constantly high and variation tended to increase as zero points.e. or empty ..1... For curves that were not coincident (i. function = evacuation model. All nonlinear R2 values were calculated as (Elashoff et al. and C = W or TL scaling exponent.26 & Larson. 1980. r2 or R2 values were compared to determine which model best explained the gastric evacuation data (Brodeur & Pearcy. square-root. Zar. logistic. as determined by bomb calorimetry. SAS Institute Inc. and large gag were compared using Kimura’s Likelihood Ratio test ( =0. 1987). For models that fit these initial selection criteria.

. 1987) and all nonlinear R2 values were calculated using Eqn (12). exponential. the model of best fit was determined to be the model that fit all selection criteria and had the highest r2 or R2 value (Brodeur & Pearcy. Next. Again. based on the average energy density of each prey type. the linear. square-root. Again. As per previous analyses. Haddon. 1999). 1980. 1982.05) (Kimura. 1990. each of the models fit to the gastric evacuation data (i. A model was considered adequate if it showed homogeneity of variances. and power exponential models) were fit to the percentage of energy (kcal/g dry weight) digested as a function of PPT for each prey type and gag size. . Again. Kimura’s Likelihood Ratio test was used to compare the energy digestion models among gag sizes ( =0. with non-linear R2 values calculated using Eqn (12). logistic. and (3) upper asymptotes greater than 85% energy digested (Hopkins & Larson. W and TL exponential scalers were given to models that differed with gag size to create prey-specific models of the percentage of energy digested over time [as per Eqns (13) and (14)]. Like the gastric evacuation data.e. 1987). only initial zeroes were included in the data set for each gag size class in order to prevent biases (Olson & Mullen. the percentage of energy digested in the stomach was divided by 100 and all Y-intercepts multiplied by 100 to fit the proportion of energy digested to the power exponential model and to allow comparisons between Y-intercept estimates (Elashoff et al. Adequate models showed: (1) homoscedasticity of variances.5 and 0. 1986). Hopkins & Larson.27 stomachs. began to show up.. the highest r2 or R2 values determined which model best explained the gross energy data (Brodeur & Pearcy. 2001). (2) Y-intercept values (estimates of the % energy digested at time 0) between -0. 1990). Zar.5% energy digested.

Numeric values. thereby creating both a quantitative (codes and percentages of prey remaining) and qualitative (prey descriptions) index of prey digestive states over time. 2002). Prey digestive states were evaluated for each prey item recovered in the digestion rate feeding trials. skulls. These average code values for each size of gag consuming either baitfish or crab prey were plotted as a function of PPT. muscle. Again. On the other hand. or codes. .. absence. The maximum digestion index of 6 indicated well-digested prey. and bones/exoskeleton. Digestion codes correlated to prey descriptions. and gut tracts) that were less than 5% digested (Lindberg et al. then the mean digestion code per gag was determined to create indices of scaled sardine and portunid crab digestive states after PPT. 2002). For example. an index value of 1 would indicate 5 to 10 % digestion and was correlated with stomach contents that had prey with pieces of skin and muscle removed by digestion (Lindberg et al. were given to each individual prey item based on the approximate percentage of the prey remaining (Tables 1 and 2) (modified from Lindberg et al.28 Indices of Digestive States Gag stomach contents recovered for evacuation rate models were immediately analyzed for the state of prey digestion using the presence. eyes.. skin. assigning a digestion index value of 0 to baitfish prey would correlate to stomach contents that had complete prey items (intact eyes.. and appearance of prey skin/carapace. the first feeding trials in each gag size class that resulted in 0% remaining at time were included in the model analyses but all other 0 codes were excluded because it could not be determined exactly when 100% evacuation occurred. or prey digested over 90%. 2002).

therefore. exponential..5. and (3) had upper asymptotes showing greater than 82. square-root. In addition.05) (Kimura. only initial codes of 6 were included in the data set for each gag size class. Haddon.29 Models of Average Digestion Codes Linear.. As with evacuation rates. 2001). the power exponential model used the proportions of average digestion codes and. the highest r2 or R2 value determined which model best fit the gastric evacuation data (Brodeur & Pearcy. . 1980.5. all Y-intercepts were multiplied by 6 (a maximum digestion code of 6 equated to prey being 90% digested at PPT) to facilitate comparison with the other models (Elashoff et al. 1990). or a digestion code=5. 1982. All nonlinear R2 values were calculated using Eqn (12). logistic. and large gag gastric evacuation rates were tested using the maximum likelihood ratio test ( =0.5 and 0. For models that fit the initial selection criteria. medium. As with the evacuation models. 1982). 1987).5% prey digested. differences among models for small. Hopkins & Larson. A model was considered adequate if it: (1) showed homoscedasticity of variances. and power exponential models were fit to the average digestion code data for each prey type by gag size in order to model the average digestion code of food remaining with PPT after feeding (MacDonald et al. (2) had y-intercept values (estimates of the average digestion code at time 0) between -0.

90 6 >90 . bits of or no skin. complete VCL. no skull. most bones present.25 Recognizable fish but maybe not complete. most skin. little meat. partial or no skull. skull and otoliths present. much chyme/digesta 4 50 . no otoliths. no chyme/digesta 1 5 -10 Mostly whole fish. most finrays. complete or partial skull and otoliths present. no guts present. no head. some bones present.50 Mostly recognizable fish.75 5 75 . complete VCL. complete or incomplete VCL. no skin. complete or partial head. most meat but more missing than code 1. otoliths absent. no skull. some meat. most guts present. some guts present. little chyme/digesta May or may not be a recognizable fish. no finrays. skull. some guts present. very little chyme/digesta 3 25 . complete VCL. most skin. most bones present. no finrays. more chyme/digesta than code 4 Not a recognizable fish. most skin but more missing than in code 1. Code Percent of Total Fish Digested Description 0 <5 Whole fish. partial head. otoliths present. some finrays maybe present. all bones present. incomplete VCL. most meat but maybe bits missing. little or no skin. otoliths present. all guts. head. some chyme/digesta Not a recognizable fish. no head. bones present. complete VCL. bones present. incomplete VCL. most or all finrays gone. some skin.30 Table 1. no finrays. skull. head. no head. otoliths present or absent. no guts. all meat. no finrays. no chyme/digesta 2 10 . Indices of digestion for baitfish Harengula jaguana by gag over post-prandial time (hr). bits of or no meat. all guts and all bones present. some meat.

no chyme/digesta Partial crab. no chyme/digesta 1 5 -10 2 10 . possibly recognizable to species. little chyme/digesta 5 75 . all meat. Code Percent of Total Crab Digested <5 Description 0 Whole crab recognizable to species.90 Partial crab. no spines. all meat. carapace anterior/posterior missing. spines soft if present. few guts. carapace folded in or top/bottom missing. some meat. most meat.75 Partial crab. all legs. no spines. some meat present but exposed. partial soft carapace. few or no legs. all guts. Indices of digestion for crab Portunus gibbesii by gag over post-prandial time (hr). more chyme/digesta than code 4 Mostly still recognizable as a crab based on shell parts and color. carapace folded in or top/bottom missing. partial soft carapace. more chyme/digesta than code 5 6 >90 . partial very soft carapace. no chyme/digesta Whole crab recognizable to species. some guts. spines getting soft. carapace usually folded in. all guts. no legs. all spines. few or no legs. some guts.50 4 50 . complete and hard carapace. little exposed meat present. no chyme/digesta Partial crab. no spines.25 3 25 . all meat. most legs. complete carapace but getting soft and folding. partial soft carapace. spines soft if present. all guts. no legs. no legs.31 Table 2. partial very soft carapace. possibly recognizable to species. carapace usually folded in. most guts.

E.E.) of their body weight. in g) as a function of gag maximum total length (TL. whereas meals fed to gag consuming meals of whole crabs averaged 1.06%) was attributed to gag being fed prey items that were completely whole.5% of the fish’s body weight. n=231) and weighed an average of 5.9-8. 21 small gag (300-449 mm TL.8865 R2 = 0. y-intercepts between 95-105% prey remaining.31 mm S. in mm) was given by (Figure 2): W = 2E-05 x TL2. range 64-93 mm. 2153-4702 g). and 19 large gag (600-750 mm TL.06 g S. range 2.E.. Gastric Evacuation Models for Gag Consuming Baitfish Prey Baitfish feeding trials were completed on 40 gag. and therefore individual prey items were not “pruned” in order to feed meals that were exactly 1. and lower asymptotes 32 . n=111) and had an average mass of 5.E. Crabs fed to gag averaged 19. 26 medium gag (450-599 mm TL.CHAPTER 3 RESULTS Gag and Prey Collections A total of 66 gag were collected for gastric evacuation experiments.0-23..) of their body weight..08 g (±0.16 g S.45% (±0.E.6 mm TL (±0. Individual gag consumed meals of whole baitfish averaging 1. The small difference in average size of fish and crab meals (0. Based on the model selection criteria (homoscedasticity of variances. 988-2295 g). and 11 large gag.5 mm..E.02% S.02% S. range 2.93 g (±0.98 n=71 (15) Baitfish fed to gag for use in the experimental feeding trials were on average 78.77-11. including 13 small gag. The relationship between gag weight (W. range 15.15 mm S.68 g). 16 medium gag. 370-940 g).4 mm CL (±0.51% (±0.5 g).

All power exponential models fit to the wet weight gastric evacuation data for gag consuming baitfish prey were significant (p<0. p=0. 2 2 =21.5. and large gag consuming baitfish prey (Table 4. Figure 3a). with 92. The active parts of the evacuation curves. or times to 5% remaining. On a dry weight basis. the power exponential models also best described the gastric evacuation processes of small.4 hrs PPT.0001). medium. 19. the three initial zero points were dropped to prevent model bias.5 hours PPT.18. Among the three different size classes of gag consuming baitfish.7. 16.5% remaining 1. Figure 3b). and large gag were calculated to be 14. one baitfish point was dropped as it was more than 2 standard deviations away from the mean. and 17. Additionally. There was no apparent lag phase in the digestion of baitfish by gag as 5-30 min trials showed between 99. the power exponential models best described the gastric evacuation processes of small.0001).33 showing less than 5% prey remaining). =11.0040. medium. and size-specific models were therefore retained. respectively. df=2. respectively. All power exponential models fit to the dry weight gastric evacuation data were also significant (p<0. medium.9-98.5 and 12. p<0. medium. on a wet weight basis for small. Scaling factors for gag W or TL incorporated into the wet weight and dry . and large gag consuming baitfish prey when using recovered stomach contents on a wet weight basis (Table 3.03. Again.5 hr after feeding. Initial zero points from fish digesting their meals faster were included in the wet weight analysis while all zero points (n=3) following this were dropped to prevent model bias.0001 and n=40. Times to 5% remaining for small. respectively).5% of the prey remaining.4. the power exponential evacuation models were not coincident on either a wet weight or dry weight basis (Maximum Likelihood Ratio [ML]: n=40. df=2. and large gag on a dry weight basis were calculated as 9.

0001). As with gag consuming baitfish. R2=0.78. respectively). initial zero points from the faster digesting fish were included in the model analyses.99). and 8 large gag.0001) but did not meet selection criteria at every gag size class using either the wet weight or dry weight data.88) when using the gastric evacuation data on a dry weight basis. the logistic models could only meet the selection criteria for small gag when modeling the wet weight data (R2=0. Figure 5a). as opposed to 90-96% of the wet weight variation explained by the power exponential model. The crab prey . Again. the power exponential models best described the gastric evacuation processes of small.98 and 0. the models were highly predictive (R2=0. including 8 small gag. Gastric Evacuation Models for Gag Consuming Crab Prey Crab feeding trials were completed on 26 gag. medium.0001). Figure 4). The square root and logistic models were significant (p<0.34 weight power exponential models were significant (p<0. However. The exponential models did not meet selection criteria for any gag size using the wet weight or dry weight data. the linear model met selection criteria for each size of gag. and explained between 93% and 95% of the variation in the data (Table 3). and large gag consuming crab prey on a wet weight basis (Table 6. however one zero point following these fish was dropped to prevent model bias.97).87) (Table 5. On a wet weight basis.00134).760. Only small gag met criteria when fit to the square root model using both the wet weight and dry weight data and was significant (p<0. Although highly predictive and significant (p<0. the linear model provided a much poorer fit (R2=0. Although gag W and TL scaling exponents were small ( 0.0001). was significant (p<0.87-0. All power exponential models fit to the crab wet weight gastric evacuation data were significant (p<0.0001 for both expanded models fit to the wet or dry weight data) and each met model selection criteria and had high R2 values ( 0. 10 medium gag.0001.

35 caused long lag phases in digestion with noticeable breakdown only starting approximately 6 hrs after ingestion. The active parts of the curves, or times to 5% prey remaining, for small, medium, and large gag were calculated as 21.0, 19.6, and 24.5 hours PPT, respectively. On a dry weight basis, the power exponential models were also the most adequate models to describe the gastric evacuation processes of small, medium, and large gag consuming crab prey, although lag phases were approximately 3.0 hrs PPT (Table 7, Figure 5b). As with the crab wet weight data, all power exponential models fit to the crab dry weight gastric evacuation data were significant (p 0.0007). Again, one initial zero point was dropped to prevent model bias. Times to 95% gastric evacuation were calculated as 20.1, 17.2, and 26.2 hrs PPT for small, medium, and large gag consuming crab prey on dry weight basis, respectively. Models of gastric evacuation for crab prey among the three different gag size classes differed significantly for both wet weight and dry weight crab data (ML: n=26,
2

=7.36, df=2, p=0.025 and n=26,

2

=8.48,

df=2, p=0.014, respectively). Because gag size had a significant effect on the gastric evacuation rate, predator size-specific models of crab evacuation rates were retained. Power exponential models with scaling factors for gag W or TL incorporated were significant (p<0.0001 for both expanded models fit to the wet weight or dry weight data) and met selection criteria using both the wet weight and dry weight data (Table 8, Figure 6). As with the baitfish gastric evacuation data, gag W and TL scaling exponents were small ( 0.00123) but the models were highly predictive (R2 0.94). The logistic models also met selection criteria, were significant (p 0.0003), and fit the crab prey wet weight data well (R2=0.96) (Table 6). However, on a dry weight basis, the logistic model could only meet selection criteria using the medium gag data (R2=0.96)

36 (Table 7). In addition, the linear model adequately met selection criteria and was significant (p 0.0001) when using the medium gag dry weight data (R2=0.90), although the model could not meet criteria when data were expressed in wet weight. The exponential and square root models did not meet the selection criteria for any gag size class. Comparative Gastric Evacuation Models for Gag Consuming Baitfish versus Crab Prey The expanded power exponential model with either W or TL exponential scalers fit to the baitfish or crab wet weight data, pooled across all gag size classes by prey type, differed significantly from one another (ML: n=66,
2 2

=88.26, df=3, p<0.0001; n=66,

=88.40, df=3, p<0.0001, respectively) (Figure 7). Gastric evacuation of fish prey in

gag occurred earlier than crab prey on both a wet weight and dry weight basis, with no lag period obvious with fish prey and at least a 5 hr lag period evident with crab prey. Caloric Values of Baitfish and Crab Prey Scaled sardine used in the prey composition analysis ranged from 67 to 111 mm TL and had an average mass of 6.09 g (Table 9). Crabs ranged from 11.9 to 32.4 mm maximum CL and had a mean mass of 7.03 g. At a mean of 73.47 % moisture, the baitfish prey had a significantly greater moisture content than the mean for crab prey at 69.59% moisture (Levene: p=0.001, 2-tailed Satterthwaite t-test for unequal variance; n=49, t=-4.77, p 0.0001). As expected, the ash content of crabs (49.6%) was significantly higher than the ash content of the baitfish prey (24.2%) (Levene: p<0.0001, 2-tailed Satterthwaite t-test for unequal variance; n=38, t=23.98, p<0.0001). The mean caloric energy density of the baitfish prey was significantly greater than that of the crab prey (Levene: p=0.009, 2-tailed Satterthwaite t-test for unequal variance; n=48, t=-31.05,

37 p 0.0001), being approximately double (4.24 kcal/g dry weight versus 2.22 kcal/g dry weight, respectively). Baitfish mean ash-free caloric energy density (5.60 kcal/g dry wt) was also greater than the mean ash-free caloric energy density of the crab prey (4.55 kcal/g dry wt). A representative size range of crabs (15.1–28.0 mm CL, mean mass=6.71 g) from both 2003 and 2004 were soaked in 15% potassium hydroxide (KOH) to dissolve away all soft tissues and proteins (Table 9). The remaining chitinous exoskeletons had an average mass of 2.57 g and were used to correct for energy unavailable to the gag for assimilation. The mean caloric energy density of the remaining chitinous crab exoskeletons was 0.76 kcal/g dry weight. When the mean energy density of the crab exoskeletons was subtracted from the mean total energy density of the initial whole crabs, the mean caloric energy density available to the gag for assimilation was 2.03 kcal/g dry weight. The mean available crab energy density was significantly lower than the mean energy density of the baitfish prey (Levene: p=0.006, 2-tailed Satterthwaite t-test for unequal variance; n=48, t=-40.50, p 0.0001). Energy Values for Recovered Gag Stomach Contents The square-root model best described the total gross energy (kcal/g dry weight) of the recovered baitfish stomach contents from each size of gag when plotted as a function of PPT (Table X, Figure 8a). The square-root and exponential models fit to the baitfish gross energy data were significant, for each size of gag (p<0.0001). However, the square root model’s R2 values were identical to the exponential model’s R2 values for each size of gag (R2=0.01-0.29) and both had very low R2 values, indicating that the square-root and exponential models fit to the baitfish data were descriptive rather than predictive. The baitfish data set was truncated at 16.5 hrs PPT because stomach content energy

Additionally. respectively) resulting in the exclusion of three baitfish data points.29. medium. regardless of gag size). 0. exponential. and 0.89-0. Y-intercept energy densities for the stomach contents from baitfish trials were 4.42-3.0 hrs PPT because it had a gross energy of zero.51. and 2.31. Model R2 values only differed by 0. 4. exponential.01-0. and square-root models were significant (p 0.5 hrs PPT. The linear model met the selection criterion of homogeneity of variances but was insignificant at each gag size (p=0. and large gag at time zero and remained fairly constant.58. 2. ending at 4. respectively). respectively. medium.92. and 0.53 kcal/g dry weight. The linear. and large gag. and 4.0 hrs PPT.24 kcal/g dry weight for small. and square-root models.0907 for small.0627. The logistic and power exponential models . respectively (Figure 8b). Conversely.41. Crab data sets for total gross energy recovered were truncated at 20.5 hrs PPT. and large gag at PPT=0 and increased to 3. one crab data point was dropped at 20. medium.90. Figure 8b).24 kcal/g dry weight).02 when small gag were fed crab (R2=0. and 3. 0.39 kcal/g dry weight at 20.38.7962.95 kcal/g dry weight at 16.92.53-5.38 densities remained constantly high (2. for each model.91 for the linear. respectively (Figure 8a). one baitfish data point was dropped because it had a gross energy of zero at 16. and 4. 3.0159) regardless of gag size and R2 values were indistinguishable (R2=0. the linear model best fit the total gross energy (kcal/g dry weight) of the crab prey stomach contents consumed by each size of gag when plotted as a function of PPT due to its simplicity (Table 11. Additionally. resulting in the exclusion six crab data points. Energy values for the stomach contents from crab trials using the linear model were 2. because again.71.0 hrs PPT.59 kcal/g dry weight for small.21. stomach content energy densities remained constantly high (3. 4.

While initial points representing 100% gross energy digested at PPT were included in the regression analyses. Both the power exponential and linear models met model criteria and were significant (p<0. the power exponential models best fit the evacuation processes of each size of gag consuming either baitfish or crab prey (Tables 12 and 13. Baitfish prey energy did not begin to digest until 2-3 hrs PPT. Exponential scaling factors incorporating gag W and TL were added to the original power exponential model to retain the size-specific models of baitfish energy digestion (Figure 10a).0003) when gag were consuming crab prey. while crab prey energy did not begin to digest until approximately 4. 2 =13.0001) at each size of gag. Figure 9).22. p=0. and these models were therefore excluded. df=2. medium. however r2 values for the linear model were lower (Table 13). and large gag consuming baitfish prey.39 could not converge on parameter estimates on either the baitfish or the crab gross energy data.5 hrs PPT. When the percentage of gross energy digested (energy passed out of stomach) was regressed against PPT. No other models met both the Y-intercept selection criteria of 5% energy digested and an upper asymptote of 85% energy digested for small. three gag fed baitfish and one gag fed crab were dropped to prevent model bias. The power exponential models describing the percentage of baitfish energy density digested over time for each gag size differed significantly (ML: n=35. Both the baitfish and crab prey energy digested data exhibited lag phases. Both expanded models with W and TL scaling factors met the selection criteria and were significant (p<0. regardless of prey type. All power exponential models were significant (p<0. The W and TL scaling exponents were . Six gag fed baitfish and three gag fed crab were excluded from the analysis due to analytical problems.0013).0001).

the data were pooled and fit to the original model (Figure 10b). The power exponential model fit to the combined percentage data of crab energy digestion fit the selection criteria.0180 and 0. therefore.23.0001). df=2. loose bones.91) (Table 15. incomplete baitfish vertebral columns began to appear. respectively). skulls.92. Indices of Digestion for Baitfish Consumed by Gag All gag consumed the baitfish and crab prey whole with very minimal. At 9 hrs PPT. respectively) but highly predictive (R2=0.91 and 0. all recovered stomach contents 90% digested were given a digestion code=6. loss of scales and skin of fish prey. At 12 hrs PPT the baitfish were unrecognizable with only incomplete vertebral columns and very small bits of meat present (code=6). however. p=0. and heads. and was highly predictive (R2=0.91. and fin rays were often absent (code>4) . was significant (p<0. and digesta (code=6).40 small (C=0.). 2 =2. guts. Figure 11a). Due to the fact that the baitfish became unrecognizable and indistinguishable from one another after being 90% digested. Initial points with digestion codes of 6 were included in the analyses. the baitfish consisted only of chyme. skin.33). Digestion was observed to be most rapid in prey located in the pylorus of the stomach. otoliths. three data points with a code of 6 were dropped to prevent model bias. The power exponential model best described the relationship between the average digestion code for each gag and PPT for small. and large gag consuming baitfish prey (R2 0. or the cracking of carapaces and limb removal of crab prey.0403. Between 16 and 18 hrs after consumption. if any. The power exponential models describing the percentage of crab energy digested over time for each gag size were coincident (ML: n=22. There were slight lag phases . medium. Baitfish prey digested continuously after their initial consumption (Table 14).0001). All power exponential models fit to the average digestion code data for gag consuming baitfish prey were significant (p<0.

Subsequently. the carapace began to soften and fold in on itself while the carapace spines began to erode.93). respectively). although the scaling exponents were small (C=0. While the logistic models met selection criteria. the power exponential model’s upper asymptote equaled or exceeded an average digestion code of 5.0202 and 0.5.91). df=2. the expanded models were highly predictive (both R2=0.5 for large gag. folded. Indices of Digestion for Crab Prey Consumed by Gag Crab prey stomach content analyses showed that several legs were commonly detached from the crab carapace located toward the pyloric end of the stomach after the lag phase.12 or less.5 hr trials resulted in average digestion codes of 1.0231) when fit to the power exponential model. approximately 6-8 hrs PPT. exponential. Digestion continued with all appendages detaching from the carapace(s) and with further softening. and.63). and square root models could not meet selection criteria for any gag size (Y-intercepts 0. For each size of gag. The carapace softened. This pattern continued until around 15 hrs . the gastric evacuation data for gag consuming baitfish prey were pooled and scaling factors for gag W or TL incorporated (Figure 11b). 2 =7.5% digestion. the exoskeleton was not yet softened nor had it been noticeably digested at 6-8 hrs PPT. which equates to approximately 82. p=0. However. The expanded models scaled for W or TL met selection criteria and were significant (p<0.54.41 detected as 2. The linear. the model’s upper asymptotes did not reach an average digestion code of 5.0001).0355.0001). The average digestion code data differed significantly between the three different gag size classes (ML: n=40. and eroded until holes formed and allowed digestive enzymes access to the body cavity around 12 hrs PPT. and had high R2 values for each size of gag (all R2 0. were significant (p<0. Because predator size had a significant effect on the gag’s average digestion code values over PPT.

or appendages.5601. Because gag size had a significant effect on the average digestion code values of crab prey over PPT. all power exponential models fit to the average digestion code data for small.5 at 24 hrs PPT. but slightly overestimated the average digestion code at the earliest time intervals (Y-intercepts = 0. void of any meat. Again.0008). From 18 to 24 hrs PPT. The .5% of the total prey digested at time.92) for each size of gag consuming crab prey.95) (Table 16. Average digestion codes for each size of gag consuming crab prey differed significantly (ML: n=26. 2 =10.0188) or TL (C=0. which equates to approximately 82. The resulting expanded power exponential models met selection criteria. df=2. The power exponential model best described the average digestion code values as a function of PPT for small.12 -0. initial points with a digestion code of 6 were included in the regression analyses but one point with a code of 6 was dropped to prevent model bias. As with the baitfish data. p=0. medium. size-specific models were retained and scaling factors for either W (C=0. the power exponential models’ average digestion code estimates equaled or exceeded code 5.0001). the crabs were often unrecognizable because only tiny bits of carapace were present in the digesta.42 PPT when the crab prey was recognizable only by scattered pigmented carapace piece(s) and small bits of partial carapace. and large gag consuming crab prey (R2 0. medium.0051).95). For all sizes of gag. Figure 12a). and large gag consuming crab were significant (p 0.0001) and had high R2 values (0.41 hrs PPT). The average digestion code data clearly showed a lag in crab digestion for each size of gag as codes at 6 hrs PPT were still <1.0258) were added (Figure 12b).860. were significant (p<0. The square root models were significant (p<0. and were highly predictive (R2=0. gut.0.

the logistic models could not adequately estimate the average digestion code at time of large gag consuming baitfish. In addition. . and 0. fit the small.01. 0. both of which are well below acceptable criteria. the logistic models were significant (p 0. estimating Y-intercepts at 0. respectively. 0.94. however this model projected the Y-intercept of the small gag size class at -2. and had R2 values at 0.43 linear model met selection criteria and was significant (p=0. Again.0008).15.06. medium. as the upper asymptote estimate could not reach an average code of 5. the exponential model could not meet selection criteria for any size of gag consuming crab prey.5 at 24 hrs. and 0.04 hrs and medium gag at -0.0002) when fit to the large gag data.80 hrs PPT. However.96. and large gag size classes well. respectively.99.

44 5000 4000 3000 2000 1000 0 300 y = 2E-05x 2. The relationship of gag weight (W) as a function of total length (TL) for gag between 300 and 750 mm TL.8865 R2 = 0.9878 400 500 600 700 800 Total Length (TL) Figure 2. .

44 1 .47 t 80 S ---- t 60 40 20 S L S S L S M L S M M S 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 3. .89 8. The power exponential model describing the gastric evacuation processes of small. and large gag consuming baitfish prey (scaled sardines) on a: (a) wet weight basis (see Table 3) and (b) dry weight basis (see Table 4).85 7.01 ____ t 80 ---L M M M S L M M L L M M L M L S S M t 60 40 S 20 S S S L M S 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) (b) 120 M S L S 100 L SM L L M M M S L M M M M S M L L M M S M L …. 52 7. ____ Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 3 .34 5. medium.16 1.45 (a) 120 ….68 6.64 6.54 2.26 1 . 100 SS ML S M L L L M M S M S M Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 1.

0044 ____ t 80 60 40 20 0 0 5 10 15 20 25 Time Time (PPT) (hr) Post-PrandialElapsed Figure 4.64 7.12 1.85 6.16 80 60 40 20 0 0 5 10 15 20 25 Time Elapsed Post-Prandial Time (PPT) (hr) (b) 120 ….46 (a) 120 ….65 W 0. The power exponential model expanded to include weight (W) or TL as scalers describing the combined gastric evacuation data of all gag consuming baitfish prey (scaled sardines) on a: (a) wet weight basis and (b) dry weight basis (see Table 5).0024 0. 100 Weight TL Y Y 2 2 t 1.0008 ____ t 7. .88 6.46 W TL 0 .0013 TL0.40 1. 100 Weight TL Y Y 2 2 t 1.

. ____ M S Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 1.07 12. medium. 85 12.91 100 ____ S L M S M M L t ---S M S L M L S t 80 60 40 20 S M M S M S L L M 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) (b) 120 M 100 M L …. L M Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 2.19 1.47 (a) 120 ….00 t 80 S L ---L S M L S L t 60 M 40 20 S M S L M S S L M L 0 0 5 10 15 M 20 25 Post-Prandial Time (PPT) (hr) Figure 5.32 9 .49 3. 23 8.28 12.17 2. 80 8. The power exponential model describing the gastric evacuation processes of small.92 2. and large gag consuming crab prey (Portunus gibbesii) on a: (a) wet weight basis (see Table 6) and (b) dry weight basis (see Table 7).

64 12. . The power exponential model expanded to include weight (W) or TL as scalers describing the combined gastric evacuation data of all gag consuming crab prey (Portunus gibbesii) on a: (a) wet weight basis and (b) dry weight basis (see Table 8).0012 t 100 80 60 40 20 0 0 5 10 15 20 25 Time Time (PPT) (hr) Post-Prandial Elapsed (b) 120 ….48 (a) 120 ….0008 TL0.88 9.47 2. ____ Weight TL Y Y 2 2 t 1.26 W TL 0 .65 12.86 9.0037 t 100 80 60 40 20 0 0 5 10 15 20 25 Time Elapsed Post-Prandial Time (PPT) (hr) Figure 6.46 W 0.30 1.0037 0. ____ Weight TL Y Y 2 2 t 2 .

16 2.65 7 .0013 W 0.46 TL0.0008 TL0.0008 100 ….65 W 0. 47 80 60 C C C F F F F F C C F F C C C C C C C 40 F 20 F F F 0 0 5 10 15 20 25 Time Time (PPT) (hr) Post-PrandialElapsed (b) 120 ____ F FF C FC F F F F F F F F F F F F F F F F F F Baitfish (F) Crab (C) Y Y 2 2 t 1 . C C CF C C C C C C C F F t 12.12 2. C C C C C C C F F t 80 60 C C C F F F F F C C F F C C C C C C C 40 F 20 F F F 0 0 5 10 15 20 25 Time Time (PPT) (hr) Post-Prandial Elapsed Figure 7.0012 100 C C CF ….64 12.49 (a) ____ 120 F FC FF C F F F F F F F F F F F F F F F F F F Baitfish (F) Crab (C) Y Y 2 2 t 1.64 7 . The expanded power exponential model describing the combined gastric evacuation wet weight data of all gag consuming both baitfish (scaled sardines) and crab (Portunus gibbesii) prey incorporated with: (a) weight (W) and (b) total length (TL) scalers (see Tables 5 and 8). .

50 (a) 6.4139 0. medium.0436t 2.0 S L S M M S L L M M M S L S M M L M S S L L L S M M M M S L M L S M S 4.0015t 4.5 L S S M L 2.2935 0.1 …. and large gag consuming: (a) baitfish prey (scaled sardines).5871 0.0051t 1.0645t ____ Y Y 2.0 ---0 5 10 0 15 20 25 Post-Prandial Time (PPT) (hr) (b) 4.7 ---- 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 8.0578t 0. ____ Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 4. fit to a square-root model (see Table 10) and (b) crab prey (Portunus gibbesii).8 L S M M L S S M M L S 2.2388 0.2 3.0 ….4 Small Gag (S) Medium Gag (M) Large Gag (L) Y 2.0 L 5.0 2. fit to a linear model (see Table 11). .2053 0. 1. Models describing the gross energy of recovered stomach contents from small.0 S 3.0073t 4.5073 0.

64 1.51 (a) 100 S S S M S L M S M MM L 80 M S S M L M L L 60 S M M S L M L M 40 M …. ____ Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 1. The power exponential model describing the percentage of stomach content energy digested over elapsed time for small. 98 5.33 1.46 6.89 7.98 10.07 2.64 1. ____ Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 1.23 t 20 ---S L M t 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 9.61 t 20 S S M M L ---- t 0 M ML L 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) (b) 100 M S M S M L L L S M 80 S 60 S L M S L 40 M L …. 78 9. .73 9. and large gag consuming: (a) baitfish prey (scaled sardines) (see Table 12) and (b) crab prey (Portunus gibbesii) (see Table 13). medium.

32 7.0403 t 20 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) (b) 100 80 60 40 __ ____ 20 All Gag Y 2 t 2.52 (a) 100 80 60 40 …. ____ Weight TL Y Y 2 2 t 1. The power exponential model: (a) expanded to include W or TL as scalers describing the combined stomach content energy digestion data for all gag consuming baitfish prey (scaled sardines) and (b) describing the combined stomach content energy digestion data for all gag consuming crab prey (Portunus gibbesii).51 10.0180 TL0.21 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 10.12 8 .69 W 0. .39 1.

medium. and large gag (see Table 15) and (b) all gag fit to the expanded model using W or TL as scalers.18 1. ____ Weight TL Y Y 2 2 t 1. The power exponential model describing the average digestion code values of gag consuming baitfish prey (scaled sardines) over elapsed time: (a) small.30 5.0355 t 1 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 11.33 W 0. .0202 TL0.66 4. 30 6. S Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 2 2 2 t 1.71 1. 97 5.56 1.53 (a) 6 M S S L M S L M S L 5 S LM S M L L M 4 S M S M L M 3 ….79 2 M ____ t 1 LML SS M S M L M L ---- t 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) (b) 6 5 4 3 2 ….84 4.

17 3. and large gag (see Table 16) and (b) all gag fit to the expanded model using W or TL as scalers.47 12.47 11.29 2. medium.97 2 . .0258 ____ t 0 0 5 10 15 20 25 Post-Prandial Time (PPT) (hr) Figure 12.72 t 1 M S L M S ---10 15 t 0 0 L M 5 20 Post-Prandial Time (PPT) (hr) (b) 6 5 4 3 2 ….54 (a) 6 S M M S S L M L 5 M S M L 4 L M S S 3 L 2 M ….80 11.38 9.59 11 .19 W 0. 1 Weight TL Y Y 2 2 t 2. The power exponential model describing the average digestion code values of gag consuming crab prey (Portunus gibbesii) over elapsed time: (a) small.0188 TL0. ____ Small Gag (S) Medium Gag (M) Large Gag (L) Y Y Y 25 2 2 2 t 4.

55 .

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the square root model (Jobling. increasing gastric evacuation rates as temperature increases (Bromley. 1982). 1970. 1974. Other studies have fit gastric evacuation data using the linear model (Swenson & Smith. 1970. 6 and 7. based on consuming meals at 1.CHAPTER 4 DISCUSSION Gastric Evacuation Models The power exponential model (Elashoff et al. Koed. Andersen.81-0. 1981). 1995). The gastric evacuation rates of several piscivorous fish species have previously been described using the power exponential model. 1990).. regardless of prey type or method of measuring stomach contents. 1973. 2001). 4. and meal size as predicting variables (Temming & Andersen.99). R2 values were very high (0. Tyler. MacDonald et al. In all cases. In the present study. 1979.. In particular. 1990. Olson & Boggs. gastric evacuation rates 69 . 1994. Nelson & Ross. the logistic model (Hopkins & Larson. and Atlantic cod consuming capelin Mallotus villosus and Atlantic herring Clupea harengus (dos Santos & Jobling. Figures 3 and 5). 1999. including black and yellow rockfish consuming whitebait smelt Allosmerus elongatusus and purple shore crab Hemigrapsus nudus (Hopkins & Larson. 1992). 1982) provided the best description of the relationship of evacuated digesta as a function of PPT for gag. 1994). temperature. temperature has one of the strongest influences on rates of gastric evacuation in fish. 1986). Persson. and with expanded power exponential models incorporating time after ingestion. explaining >81% of the variation in both the baitfish and crab digestion data. the exponential model (Brett & Higgs. predator weight.5% of the gag’s body weight (Tables 3. Jones.

1987. Rao. 1984). 1973.70 were quantified at a mean temperature found on artificial reefs in the eastern Gulf of Mexico during the warmer months of the year (mean=28°C from May-November) because baitfish are only present on these reefs during these warmer months. upper/lower asymptote criteria. 1974. 1998). and/or the coefficient of determination (R2) (Swenson & Smith. standard errors. with steeper regressions resulting in higher R2 values (Barrett. homogeneity of variance. 1982. Specifically. Using the R2 value. Rao. 1984. Adjusted R2 values comprise the proportion of variance accounted for in the data and takes into account both the number of predictor variables and the sample size. Brodeur & Pearcy. Ruggerone. 1989). Rao. Using R2 values to compare among different models can also be problematic if the models use different numbers of predictor variables. 1984. which represents the total variability in y that can be explained by the fitted regression (Zar. may not by itself reliably evaluate regression models (Draper. Goodness of fit evaluations among different gastric evacuation models has often involved using a combination of Y-intercept values. 1999). since R2 will increase with an increasing number of predictor variables (Healy. small R2 values can be statistically significant while large R2 values can be insignificant (Rao. Eleshoff. residual mean square values. and R2 values. 1998). In addition. 1998). including Y-intercepts. equal values of the residual sums of squares can result in different R2 values depending on the steepness of the regression. 1998). residual plots. While in the present study goodness of fit was determined by multiple critieria. 1984) 2 Radj 1 MS Residual MS Total (16) . 1984. and is calculated as (Healy. using adjusted R2 values to determine model fit may evaluate regression models more reliably (Healy. Healy.

1986. dos Santos & Jobling. In this study. although this likely reflects the curvilinear flexibility of the power exponential model. gastric evacuation rate estimates and the shape of the evacuation curve have often depended on whether the stomach contents were measured by wet weight. . enzymes. dry weight measures of prey remaining over PPT should have corrected for measurement error but it also introduced new error because the original baitfish and crab dry weights had to be back-calculated based on predictive regressions for wet weight versus dry weight of whole representative prey. they should better evaluate fit between models with differing numbers of variables. Because wet weight measurements of the amount of food remaining in the stomach are the most common measurement taken in the field (Bromley. Theoretically. dry weight. or volume (Brodeur. Because of this measurement error. 1990). 1984. or body fluids may have remained and contributed to an overestimation of the amount of prey remaining over PPT (Hopkins & Larson. 1990. 1994). such as the logistic and power exponential models in this study. 1992). Wet weight measurement error had the potential to affect this study due to the fact that prey items recovered at later time periods could not be easily blotted prior to weighing. model selection was only minimally influenced by the way in which the stomach contents were measured considering that the power exponential model best fit both baitfish and crab wet weight and dry weight data. Since adjusted R2 values are not influenced by the number of predictor variables within each model. Olson & Boggs.71 where MSResidual = residual mean squares and MSTotal = total mean squares. the wet weight gastric evacuation curves generated from gag feeding trials in this study may be the most useful. Hopkins & Larson. and so water.

Previous work has shown that prey (whole Cape anchovy Engraulis capensis.72 Both baitfish and crab prey used in all gastric evacuation feeding trials were vacuum-sealed fresh. pieces of hake Merluccius sp. as compared to thawed pellets. pieces of squid Loligo vulgaris. Results from the present . field and laboratory experiments with tagged. Conversely. MacDonald et al. In contrast. 1987). frozen immediately. (1982) also found that ocean pout had lower indices of digestion at 20 hrs PPT when fed pellets of thawed Y. respectively. incise at 5 and 20 hrs PPT. respectively. and stored frozen because live baitfish were only present on artificial reefs in the eastern Gulf of Mexico in the warmer months (capturing and maintaining scaled sardines was not feasible) and collections of crab prey were only successful during February and March. Additionally.. MacDonald et al. (1982) tested the effects of frozen versus live prey on digestion indices and found that Atlantic cod and ocean pout Macrozoarces americanus had significantly lower indices of digestion when consuming pellets of fresh bivalves Yoldia sapotilla or polychaete worms Nephtys incise at 5 hrs and 20 hrs PPT. Atlantic cod and ocean pout had lower digestion indices when fed thawed versus fresh polychaete worms N. and detached rock lobster tails Jasus lalandii) that had been frozen and then thawed before use in in vitro digestion rate experiments digested faster in pepsin than controls of fresh prey (Jackson et al. Using previously frozen prey in gastric evacuation feeding trials may therefore result in underestimating the amount of food remaining in the stomach at time and consequently lead to overestimating the gastric evacuation rate. free-swimming black and yellow rockfish indicated that the evacuation rates of thawed purple shore crab determined in the laboratory gave acceptable approximations of evacuation rates in situ using freshly fed purple shore crabs (Hopkins & Larson. sapotilla. 1990).

evidence suggests that the extrapolation of single-meal models can still yield reasonable estimates of total daily ration (Persson. Brodeur. 1989. dos Santos & Jobling. Hopkins & Larson. Further work is necessary to determine if the single-meal gastric evacuation rates determined in this study are biased when applied to sequential-meal. 1984. Andersen. 1990. 1995. with times to 90% gastric evacuation being approximately 15 hrs PPT (Lindberg et al. Temming & Andersen. Even so. and considering this may not be the case. He & Wurtsbaugh. Koed. 2001). Evacuation trials for meals of combined fish . These results correspond very closely with preliminary field estimates on the stomach contents of wild gag. Ruggerone. 1993. 1996. Nelson & Ross. and sequential-meal/mixed-meal situations.73 study indicate that thawed baitfish prey are 95% evacuated from the stomachs of gag (300-750 mm TL) between 14. 1992). Most gastric evacuation experiments have described predator digestion patterns based on single-meals of a single prey type. 1982. or meals of brown shrimp and lugworm Arenicola marina were not significantly different than the gastric evacuation rates of single-prey meals. mixed-meal. 1970.. 1999. models may tend to underestimate the evacuation rates of early meals and overestimate the rates of later meals (Persson. Applying single-meal models to sequential-meal experimental conditions assumes that stomach contents are homogenous. 2002). Singh-Renton & Bromley. MacDonald et al. dos Santos & Jobling. and whether or not their extrapolation will yield acceptable estimates of total daily ration.7–19. even though multiple prey types may be incorporated into a multivariate model (Tyler..5 hrs PPT. SinghRenton and Bromley (1996) determined that the gastric evacuation rates of Atlantic cod and whiting Merlangius merlangus fed mixed-prey meals of brown shrimp Crangon vulgaris and Atlantic herring. 1984. 1994. 1984. 1992).

and 80-90% prey remaining at 2-5 hrs PPT. and on Atlantic cod consuming capelin. The power exponential model in the present study accounted for the lack of a lag phase seen in the gastric evacuation of scaled sardine prey based on wet weight and with the slight lag phase observed in the dry weight data (0-3 hrs PPT) (Figure 3). or krill were initially faster that rates of gastric evacuation considered on a wet weight basis. The scaled sardine prey chosen for this study were . As with most fish predators. the power exponential model best fit the gastric evacuation data (Tables 3. 6. dos Santos and Jobling (1992) used the power exponential model and found that on a dry weight basis the gastric evacuation rates of Atlantic cod fed capelin. since some stomach contents recovered in the field contain mixed prey types (personal observation). prawn Pandalus borealis. along with mechanically macerating the prey. both measurement types showed lag phases in digestion. on both a wet weight and dry weight basis. 1992). 1995). gut. and 7. 1990). likely due to the lower temperature (14. Likewise. and krill Meganyctiphanes norvegica (dos Santos & Jobling. Figures 3 and 5). 4. 1990). and other tissues can be easily digested (Diana. herring. The power exponential model best fit black and yellow rockfish gastric evacuation data based on the wet weight and dry weight measurements of recovered whitebait smelt. digestive enzymes must work through the prey’s scales and skin or carapace before the muscle. gag capture and swallow their prey whole and therefore. Effects of Prey Type For all sizes of gag consuming baitfish or crab prey.5°C) (Hopkins & Larson. including those on black and yellow rockfish consuming whitebait smelt (Hopkins & Larson. Atlantic herring.74 and crab prey were not addressed in the present study but would be useful to develop for gag. prawn. Other studies have used the power exponential model.

relatively small and low in energy density compared to mature sardines.. Although low in energy compared to mature sardines. the sardine prey had fairly small scales and thin skin. 2002). the wet weight and volume gastric evacuation data from black and yellow rockfish fed purple shore crabs were modeled using a power exponential model. they were of a similar size to prey found in the stomach contents of gag during warmer months on reefs in the eastern Gulf of Mexico (Lindberg et al. these immature sardines are high-energy prey when compared to other prey types such as portunid crabs (Table 9). which all tend to possess thinner exoskeletons and therefore less chitin material than most crab exoskeletons (Tyler. although the dry weight gastric evacuation data was better fit using a linear model (Hopkins & Larson. Therefore. In fact. the linear model described the continuous gastric evacuation of baitfish prey on a wet weight basis for each gag size class relatively well (r2>0. . MacDonald et al. Because of their size. or polychaete prey. The gag’s digestive enzymes were likely better able to quickly and continuously break down these comparatively low-energy. However. amphipods.75 immature and therefore. therefore these sardines were considered highly friable compared to mature sardines of the same species.. immature sardines. The power exponential model also provided the best fit for both the wet weight and dry weight gastric evacuation data for all sizes of gag consuming crab prey (Tables 6 and 7. the fact that a linear model adequately describes the gastric evacuation process of baitfish prey is still consistent with the continuous digestion of high energy prey types over PPT as described by Jobling (1987). 1970. krill.93) (Table 3). Few studies have modeled the gastric evacuation of crab prey fed to piscivorous fish species. Most studies have concentrated on shrimp. Figure 5). Similarly. 1990).

when gag consumed crab prey on a wet weight basis. Digestion was then slowed again as residual hard parts persisted in the stomach. The power exponential model was able to capture the large digestive lag phases seen in the gastric evacuation patterns of the crab prey (Figure 5). 1992. 1994. Andersen. Hopkins and Larson (1990) found that modeling the gastric evacuation data from black and yellow rockfish consuming the purple shore crab on a dry weight basis reduced the digestive lag phase. and starvation. lag phases of approximately 0-6. followed by a rapid increase in digestion for approximately the next 12-15 hrs. thereby causing the large lag phases seen in the wet weight data (Figure 5). this fluid retention may vary greatly with crab molt stage. dos Santos & Jobling. as compared to baitfish prey. This trend was likely caused by measurement error because the crabs may have more fluid within their exoskeleton normally. Temming & Andersen. and in addition.5 hrs with little or no digestion occurred. 1966. At these later time intervals. 1999). force-feeding. digestion-resistant exoskeleton material and other residual hard parts are known to cause evacuation curves to level off and form the lower asymptotes of various gastric evacuation models (Battle.76 1982. 1992). Basing models on the dry weight data tended to decrease the lag phases (0-3 hrs). Windell. The crabs could not be easily damp-blotted for weighing and therefore water and body fluids may have remained within the crabs’ exoskeletons. dos Santos & Jobling. as compared to modeling the data on a wet weight basis. thereby appearing more similar to the dry weight digestion curves of gag consuming baitfish prey. Similarly. 1935. accounting for the model’s lower asymptote estimates. Lags in digestion after prey consumption have been attributed to impaired enzyme secretion. and have the potential to cause bias by . Generally.

and mechanical peristalsis necessary to breakup the chitinous material in the softened exoskeleton (Pandian. Atlantic cod. 1982. 1998). prawn. Atlantic herring. 1984. through the secretion of hydrochloric acid to decalcify the calcium carbonate material found in their exoskeletons. 1967.. the lag was due to the gastric softening and digestive break down of whole crustaceans. found that predator size had a significant effect on the rate of gastric evacuation. thereby playing a role in the retention of exoskeletal material and the formation of digestive lag phases. 1995).. In addition. and whether stomach contents were measured on a wet weight or dry weight basis. food was withheld from gag for <48 hr prior to testing and the gag voluntarily consumed thawed baitfish or crab prey. most gag either gained a very slight amount of weight or maintained their weight while in captivity. 1979). chitobiase. or krill (dos Santos & Jobling. the observed lag phase was not due to force-feeding or starvation. however. and whiting fed saithe Pollachius virens (Jones. MacDonald et al. In contrast. Effects of Predator Size Gag size had a significant effect on their rates of gastric evacuation. turbot Scophthalmus maximus fed processed pellets (Flowerdew & Grove. In the present study. dab Limanda limanda fed an artificial paste diet (Jobling et al. Hopkins & Larson. 1970). studies on haddock Melanogrannus aeglefinus. the pyloric sphincter may limit the size of items that pass into the intestine. Jones. Lindsay. other studies on Atlantic cod that were fed shrimp Pandalus montagui (Tyler. Lovell. plus chitinase. and plaice Pleuronectes platessa fed fish-paste . 1987). Therefore. whether they were consuming baitfish prey or crab prey. More likely. 1974. with smaller predators having faster rates of gastric evacuation. 1974). and Atlantic cod fed capelin. 1973. 1977).77 overestimating the percentages of prey remaining at PPT (Swenson & Smith.. Jackson et al. Similarly. In addition. 1990.

Therefore. Meal size was not included as a predicting factor in the gastric evacuation models because initial meal sizes cannot be estimated in wild gag caught in the field.0 created multiplicative scaling factors for the evacuation models that were very close to 1.g. while gag size significantly affected the gastric evacuation rates of both baitfish and crab prey. prey size. the present study shows that gag size significantly affects rates of gastric evacuation despite feeding meals of baitfish or crab relative to each gag’s body weight. based on the assumption that the effects of predator size on gastric evacuation times are multiplicative (Andersen. whereas their evacuation rates differed when they were fed equal-sized meals (Swenson & Smith. W=1). predator weight (W). 1980a) did not observe size-specific evacuation rates.78 (Jobling. and predator total length (TL) based on this assumption (dos Santos & Jobling. Koed. These small but significant scaling factors are reasonable considering that the gag were fed meals on a . Swenson and Smith (1973) found that large and small fish evacuate meals at approximately the same rate when meals where fed relative to their body weight. In the present study. prey energy density. Previous studies have incorporated scaling factors such as temperature.00134. meal size. 1994.. 1999) (Tables 5 and 8. Andersen. Scaling factors were incorporated into the power exponential models for both prey types and measures of stomach contents (wet weights and dry weights) to account for significant differences in either gag weight or total length. 1973). Previous work has shown that feeding predators meal sizes relative to their body weight accounts for the variability associated with gastric evacuation rates in different sized predators (Swenson & Smith. values for W and TL raised to a power approaching 0. Andersen.0 (e. Figures 4 and 6). 1973). 1999. 2001). Temming & Andersen. 2001. all W and TL coefficients were 0. 1992. In contrast.

the role of chitinase in the gastric processes of fish has been somewhat controversial. into an absorbable form and therefore can not utilize chitin as an energy source because the -linkages of glucose within the chitin molecule cannot be broken by amylase enzymes (which only breaks -linkages) (Battle.. However. because using the percentage of meal recovered has been known to take out much of the variation in the weight of meals recovered from different sized predators (Bromley. and 25% chitin. i. MacDonald et al. a percentage of gag body weight. Lindsay (1984) hypothesized that the primary function of chitinase in the gut of fish may be to chemically disrupt the outer chitinous material of prey.79 relative weight basis. especially marine fish. Prey Composition Many fish secrete chitinase but it has been commonly accepted that piscivorous fish. (1992) suggested that absorbing NAG and D- . Vollhardt & Schore.e. 1984). other work has shown that high levels of gastric N-acetyl-D-glucosamine (NAG) were caused by high levels of chitinolytic enzymes (chitinase and chitobiase) and resulted in significantly lower growth rates in fish fed diets containing 4. such as crabs. are not efficient at converting carbohydrates. and better predict the percentages of prey remaining in the stomachs of all sizes of gag.. 1985. 1984). such as chitin. Lindsay & Gooday. Lower levels of chitinase activity were found in fish that mechanically disrupted prey before ingestion compared to fish that swallowed prey whole and likely required more chitinase to breakup chitinous material (Lindsay. better determine the extent to which the W and TL exponential scalers account for gag size. Incorporating W or TL scaling factors into the power exponential model with data on gag <200 mm TL and >750 mm TL should further improve the R2 values of the power exponential model. as compared to diets containing starch (Lindsay et al. 1985. 10. 1994). Medved. 1987). Conversely. Jackson et al. 1982.. 1935.

However. The presence of calcium carbonate.80 glucosamine (Gln) (i. One advantage of chitinolysis seems to be the increase in gastric evacuation rates associated with the increase in mechanical breakdown of the exoskeleton. 1992).. 1992). which possibly inhibited individual growth because NAG and Gln could not be utilized as quickly as D-glucose for metabolic. (1992) hypothesized that... (1962) reported that mammals metabolize NAG and Gln more slowly than D-glucose. (2002) has shown. Kohn et al. Jackson et al.e. the mean energy densities of both baitfish and crab prey were determined. and potentially. 1967). waste removal. fish that consume chitin may have adapted to inefficiently absorb chitin end-products because of the possible costs associated with reduced growth. however. Compere et al. versus approximately 89-96% of the total energy when fish prey are consumed (Diana. estimations of exoskeletal energy densities may have been biased. thereby allowing easier access to more readily digestible tissues and allowing proteolytic enzymes better access to cuticle proteins (Jackson et al. to better estimate the mean energy density available for assimilation by individual gag consuming crab prey. and growth processes (Jackson et al. that many proteins can be extracted from the exoskeleton without decalcification and that many proteins in the exoskeleton have relatively high apparent molecular weights. It has been commonly accepted that crustaceans with their chitinous exoskeletons allow piscivorous predators to assimilate only between 70–80% of the total crustacean energy consumed. In the present study. although chitin may be available to assimilate. along with the mean energy density of the crab shells by themselves. . the products of chitin digestion) may actually be inhibiting growth. 1995). Potassium hydroxide treatments for tissue removal of the crab prey did not decalcify the exoskeleton or extract the cuticle’s pigments (Pandian.

and large gag . were within the size range (CL) and mass (g) of the whole crabs used for the prey analyses. would be expected to be lower in energy density and have slower gastric evacuation rates (Flowerdew & Grove. All representative baitfish and crab prey used in the prey analyses reflected size (TL and CL) ranges that mirrored those normally found in the stomach contents of wild gag (Murie. higher mean % moisture. fat and ash content. 1992). In fact. 1979. times to 95% evacuation at 28. Jobling. digestionresistant materials have been shown to slow rates of gastric evacuation due to the relatively large amounts of residual matter and ash that must be digested and passed from the body (Battle. Diets containing unabsorbable materials tend to lower a meal’s energy density and increase rates of gastric evacuation (Flowerdew & Grove. higher mean available caloric energy density. Therefore. unpublished data).0oC for small. Windell. 1966. The composition of food prey. 1978. Jobling. specifically unabsorbable chitin energy. On the other hand. such as crabs with exoskeletal coverings. The representative baitfish prey had lower mean mass. 1980a). Hopkins & Larson. 1979. Quantifying baitfish energy densities in terms of their ash-free dry weight corrects for the inorganic materials of their bones and other parts. remaining in the crab exoskeletons may have lowered the estimates of exoskeletal energy density. such as scales. lower mean % ash. and higher mean available ash-free caloric energy density estimates than the crab prey (Table 9). is known to affect rates of gastric emptying (Elliott & Persson. in the present study. prey with high amounts of digestion-resistant materials. 1990. 1980a). 1980a). The crab prey used in the regression analysis to correct for the known unavailable energy in crab fed to gag. 1935. in terms of energy density and specifically. dos Santos & Jobling.81 pigments. medium. Jobling.

2001). there was no lag phase in digestion for black and yellow rockfish fed whitebait smelt. Crab samples collected in the spring may have contained higher percentages of pre-molting .3 hrs longer than gag consuming baitfish (Figure 7). a 22. 1990). In the present study. Similarly. including immature scaled sardines.9% difference (Hopkins & Larson. however.5 hrs PPT. immature scaled sardines were collected during late October or November of 2002 and 2003. 2002) were of a similar size range. compared to a large lag phase seen when rockfish were fed the purple shore crab and 99. Spring samples of crab could have biased the rates of gastric evacuation and estimates of energy density determined in this study. Most digestion and consumption models do not account for fluctuating prey energy densities between species and seasons (Koed.8% of the meal was remaining at 10 hrs PPT. are only present on artificial reefs in the eastern Gulf of Mexico in the warmer months. While these earlier findings are corroborated by the present study. were collected in February and March of 2003 and 2004. 1990).7% of the meal was remaining 10 hrs PPT..82 consuming crab prey on a wet weight basis took 4. a 52. while purple shore crabs were not evacuated to 95% until 49. black and yellow rockfish evacuated whitebait smelt to 95% in 27 hrs. relative to the scaled sardine and portunid crab prey used in the present study. Due to the fact that all baitfish.5 hr difference (Hopkins & Larson. In a previous study. as 46. and all sardines collected or observed on artificial reefs in the eastern Gulf of Mexico (Lindberg et al. fluctuating baitfish prey energy densities between seasons did not need to be determined. it is important to recognize that the whitebait smelt used in the black and yellow rockfish feeding trials were thin-skinned and almost scaleless. while the purple shore crabs were smaller. Crab prey.

Considering that prey composition and energy densities greatly affect both gastric evacuation and consumption rates.=0. and their seasonal molting patterns. Determining the extent of fluctuating crab prey energy densities by season may be necessary depending on when they begin to show up in the gags’ diet. resulting in greatly reduced energy densities. although the measurements of mean mass for both prey types was slightly smaller (Tables 9 and 10). any biases caused by pre-molt or post-molt individuals were likely small and within error.83 and post-molted individuals that have no gonad development. The baitfish stomach contents had higher mean % moisture and mean available caloric energy density estimates but lower mean % ash and mean available ash-free caloric energy density estimates than the representative baitfish prey. Stomach Content Composition Both the scaled sardines and portunid crabs used in the gastric evacuation and average digestive code regressions were within the size range of the subsampled prey used in the whole prey regression analyses.06) and more than 81% of the variation in the gastric evacuation data was explained by the power exponential model. it is important to know the true energy densities of the prey consumed by season. or if they are continuously found in their diets. considering that all crabs used in the prey composition determinations were similar in energy density (mean=2.22. All values are very close to those of the representative baitfish prey and likely reflect the natural variation seen in the scaled sardine population residing on artificial reefs in the eastern Gulf of Mexico. mean available caloric energy density. S. and mean available . However. The crab prey stomach contents had higher mean % moisture. Comparing the energy of crabs sampled in the present study to crabs sampled from the stomachs of wild gag could determine the extent to which premolt and post-molt crabs potentially biased results from the present study.E.

the square-root model fit to the baitfish data was more descriptive than predictive. 1980a). Figure 8).84 ash-free caloric energy density estimates but a lower mean % ash estimate than the representative crab prey. The fact that mean % moisture differed significantly between whole baitfish and crab prey but did not differ between baitfish and crab stomach contents may reflect wet weight measurement error ( i.. non-nutritive bulk . that may preferentially remain in the stomach for longer periods. or the addition of secreted digestive juices (hydrochloric acid. Jobling (1980a) fed plaice meals of fish paste with kaolin (inert material) and regressed the square-root of the meal’s energy density by time with a linear model and found that energy decreased over PPT. respectively. pepsin. A linear model assumes that energy turnover is maintained at a constant level. should be instantaneously evacuated from the stomach (Jobling. the ability to damp-blot excess water off recovered prey). This likely resulted in a homogenous mixture of baitfish with a relatively consistent energy density in the gag’s stomach. Again. therefore. the crab prey may not comprise such a homogenous mixture because the crabs contain higher amounts of indigestible matter. and mucous). such as chitin. all values are very close to those of the representative crab prey. muscular activity and rates of . likely reflecting the friable nature of the baitfish prey that was easily broken down through mechanical and enzymatic action in the gag’s stomach. most likely reflecting the natural variation seen in the crab population. regressed as a function of PPT (Tables 11 and 12. The square-root and linear models provided adequate fits for the gross energy (kcal/g dry wt) of baitfish and crab prey stomach contents. In contrast. Jobling (1980a) hypothesized that as the energy density of meals increased.e. such as chitin. Few studies have looked at stomach content gross energy as a function of PPT. However.

but did not fit a quantitative function to the data. A greater weight change in the energy:weight ratio compared to the energy change due to low energy skeletal or exoskeletal material remaining in the gut may be causing the increase in the gross caloric energy densities of the recovered gag stomach contents over time. One study by Beamish (1972) on largemouth bass Micropterus salmoides fed emerald shiners Notropis atherinoides noted that the caloric energy density of the stomach contents increased over time when expressed as cal/g dry weight. except small gag consuming baitfish (Figure 8). Conversely. and that protein (i. that is.85 gastric peristalsis may restrain energy turnover. the function was not modeled. at the same time.96) (Tables 13 and 14. in the stomach contents of whiting declined over time and attributed the decline to changes in the ratio of readily digestible soft tissue to skeletal material but again. nitrogen) digested more quickly than lipid. Figure . The power exponential model best described the percentage of stomach content energy digested over PPT for both prey types and in all sizes of gag. large portions of digesta and heavy macromolecules are evacuated from the gut.e. the caloric energy density of the stomach contents for both prey types increased over PPT for each size of gag. a larger portion of low energy unabsorbable skeletal or exoskeletal material remains in the stomach longer compared to proteins or lipids but. Bromley (1988) found that the gross energy density (kcal/g dry weight) of sandeel Ammodytes sp. with predictive models explaining over 87% of the variation (R2 = 0. These results most likely reflect energy:weight ratio changes over PPT. Relatively heavy macromolecules.87-0. In the present study.. such as calcium carbonate and some proteins within the exoskeleton. may be extracted from the exoskeleton with the addition of digestive enzymes (including HCl) more quickly then lighter macromolecules.

1967).07 to 10. and B ranging from -1. Figures 5 and 9).23 to 2.32. Indices of Digestion Visual indices of prey digestive stages over PPT are one way that researchers can estimate times of prey consumption in the field based on the average digestion codes given to recovered stomach contents instead of quantitative measurements (i.73 to -2. However. with the lag in the percentage of digested crab prey energy being approximately 5 hrs PPT and the wet weight gastric evacuation lag phase being 6 hrs PPT. When each size class of gag consumed .89. respectively (Tables 7 and 14. In the present study. the power exponential model fit the percentage of digested crab energy from the gag stomach contents in a pattern most similar to the gastric evacuation of the dry weight crab prey data. Lag phases were more similar to the wet weight gastric evacuation crab data.19. such as that of Beamish (1972) and Bromley (1988). Previous work. backcalculation of original prey sizes and weights).. converted. the lag phases present in the baitfish energy digested data (2-3 hrs PPT) were more similar to the lag phases seen in the dry weight gastric evacuation data of small and large gag (2-2.78 and from 1.98 and from 1. Figures 3 and 9). with A ranging from 9.5 hrs PPT). the power exponential model fit the percentage of digested baitfish energy in a pattern most similar to the gastric evacuation of the wet weight data for baitfish (A ranging from 5.01. and B ranging from -1. respectively) (Tables 3 and 8. for the digested energy data and dry weight gastric evacuation data.61 and from 6. absorbed.23 and from 8. have determined gross energy densities over time.00 to 9.52 to 1.16 to 8. and residual energy densities per day as a function fish body weight for estimations of absorption efficiency (Pandian.86 9). Conversely.46 to -1. others have estimated the amount of energy consumed.e.64 to 7. for the digested energy data and wet weight gastric evacuation data.

Therefore. only average digestion codes of prey at time will need to be estimated. 2002). as the crab’s exoskeleton acts as a barrier to prevent quick enzymatic and mechanical digestion of the crab tissues (MacDonald et al. Evacuation studies on Atlantic cod. Again. 1982)..2-1.. Using the visual indices of prey digestion qualified in this study. sapotilla. based on preliminary average daily consumption estimates of between 1. incisa have also found that visual indices of prey digestion are correlated with quantitative differences in evacuation rates between prey types (MacDonald et al. 1990. 1992.. Lag phases of up to 6.. Figures 11 and 12). and American plaice Hippoglossoides platessoide fed bivalves Y. 1982. will allow researchers to approximate the wet weight and dry weight percentages of prey remaining in a gag’s stomach and estimate an approximate time of prey consumption for wild gag. little or no lag phase was seen in the average digestion code data when gag consumed baitfish prey due to the fact that the gag’s digestive enzymes and mechanical peristalsis of the stomach could easily breakup the individual prey items. Hopkins & Larson. dos Santos & Jobling.87 either baitfish or crab prey the power exponential model best described their average digestive codes over PPT (Tables 15 and 16. winter flounder Pseudopleuronectes americanus. Bromley. These visual indices of scaled sardine and portunid crab digestive stages clearly support gastric evacuation results from the present study and the contention that prey containing higher percentages of chitin will remain recognizable for longer periods in the gut (MacDonald et al. along with their corresponding models of average digestion codes at time. ocean pout.5 hrs in digestion of crab were seen in both the average digestion code data and the wet weight gastric evacuation data. amphipods Tmetonyx cicada. 1982). and polychaetes N.8% body weight per day (Lindberg et al. 1994). thereby saving measurement effort and processing .

i. such as baitfish. squid. both fish and crustacean prey. such as crab prey. If baitfish and crab energy densities were equal. capture. and gastric . 2004) or an additive model. Consumption Based on results from this study. these back-calculated times of prey consumption can be used to determine the frequency with which gag feed and their diurnal pattern of feeding. handling. will need to increase their consumption of that prey to grow at rates similar to predators consuming more energy-rich prey. such as baitfish.e. the rate of crab prey exploitation would likely better approximate a Type II Functional Response curve (Holling. gag will likely require a multiplicative consumption model. crab. Both feeding frequency and the diurnal feeding pattern (synchronous or asynchronous) of gag will affect which type of consumption model is most appropriate for wild gag (Adams & Breck. Water temperature influences on gag gastric evacuation rates must be determined on a species-specific basis. 1990). such as that of Atlantic cod feeding on Atlantic herring in the Barents Sea (Johansen et al. Such models can account for the differing gastric evacuation rates of various representative prey items. handling time.88 time for stomach contents collected in the field.. which includes the pursuit. and possibly larger pelagic prey. with a delay for gag consuming crab.. each species’ seasonal energy density fluctuations should be incorporated into a comprehensive gastric evacuation model for gag. Most importantly. 1959) compared to baitfish prey as gag became satiated. taking into account the seasonality of prey quality and quantity Predators consuming prey that contain less available energy and higher amounts of indigestible materials that slow rates of gastric evacuation. Specifically. Similarly. The multiplicative or additive model chosen must incorporate gastric evacuation models explicit to the prey types consumed by gag.

2003). the consumption of crab prey by gag will level off at a maximum rate that is lower than the maximum rate of baitfish consumption because crabs require more handling time in digesting food. Gill..5 hrs PPT. with times to 95% gastric evacuation of baitfish and crab prey between 14. resulting in the number of prey consumed per predator initially rising as prey density increases and then leveling off with further increases in prey density (Holling. 1959). respectively . even when gag are fed to satiation on crab.89 evacuation of prey. 2001). 1966. if the time and effort needed to capture baitfish and crab prey were similar. a gag should preferentially choose energy-rich baitfish prey over lower energy crab prey. 2002). MacArthur and Pianka. Optimal Foraging Theory predicts that prey are detected and consumed based on their energy density. Based on the Type II Functional Response curve.. i. Gag fed to satiation on friable baitfish prey should be limited less by prey handling times. therefore. 1966. density-dependent effects may reduce a gag’s successful foraging episodes due to increased competition with a reduction in prey .e. differences in consumption rates of baitfish and crab prey may be a function of their differing vulnerabilities (Sih & Christensen. However. in addition to the time. it is unclear why some gag may be consuming significant amounts of energy-poor crab prey (Lindberg et al.7-19. more gastric evacuation time) than baitfish prey. in terms of encounter and capture success rates. Therefore. Based on preliminary analyses of gag stomach contents.6-24. Results from the present study clearly show that crab prey require more handling time (at a minimum. is taken into account. effort and risk involved in capture of prey (Emlen.5 hrs and 19. resulting in more energy available for individual gag growth. Different prey have different vulnerabilities. energy available for gag growth will be limited to the time required for individual gag to handle the crab prey.

7-17. thereby causing an increase in the gag’s optimal diet breadth to include lower energy prey.4 hrs PPT determined from experimental feeding trials of captured gag in this study. previous density-dependant effects on foraging successes. 2002). Results from the present study independently verify preliminary gastric evacuation rate estimates of wild gag consuming baitfish prey on artificial reefs off the west coast of Florida (Lindberg et al. Interestingly. This feeding disparity may be evidence of differing prey recognition capabilities. This range in gag size reflects the stage-specific size range of gag normally encountered on coastal reefs in the eastern Gulf of Mexico (Lindberg et al. previous experience. 1966). most probably a result of the crab exoskeleton acting as a barrier to digestion. the power exponential model best fit both the wet weight and dry weight gastric evacuation data irrespective of gag size and prey type. differing levels of unabsorbable material. Gag size (300-750 mm TL) significantly affected both the scaled sardine and portunid crab gastric evacuation models when using either the wet weight or dry weight data. some gag seemed to naturally take to crab prey much more readily in the laboratory than others (personal observation). such as crabs (Emlen. 2002). Conclusions In conclusion. The flexibility of the power exponential model also best fit the percentage of stomach content energy digested over time and the average scaled . in terms of the curve’s shape and total prey retention times. Gag gastric evacuation rates as a function of PPT were significantly affected by baitfish and crab prey composition. individual food preferences in gag. Based on field estimates. compared to 95% gastric evacuation between 14. wild gag evacuated 90% of a baitfish meal in 15 hrs and 100% in 16 hrs. or differing portunid crab and baitfish abundances in the environment affecting predator-prey encounter rates. and energy density..90 numbers. 1966. MacArthur and Pianka.

sequential meals. with different relative meal sizes. and the average digestion codes over PPT of wild gag. Additional work on quantifying the gastric evacuation rates of gag < 300 mm TL and >750 mm TL. at temperatures <28°C and >28°C. The power exponential gastric evacuation models for gag determined in the present study have shown that prey type and gag size are not only significant parameters when determining rates of gastric evacuation but they. . are variables that must be included in any future consumption model for wild gag. the percentages of prey energy digested over PPT. along with back-calculated times of prey consumption by wild gag.91 sardine and portunid crab digestion code data for each size of gag. mixed meals. and differing prey types will improve the ability of the best-fit power exponential model to predict the percentages of prey remaining over PPT.

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and the various human influences affecting these relationships. While pursuing a Master of Science in biology at the University of Florida. she served as Treasurer for Students United in the Research of Fisheries (SURF). During September of 2004. she began work at Mote Marine Laboratory in Sarasota. Michigan. and resulted in a Bachelor of Science in biology with an aquatic emphasis and minors in Russian studies and studio art. predator-prey relationships. Her curiosity developed into an interest in fisheries science at Grand Valley State University in Allendale. 1975. After working as a Lake Management Assistant on Kiawah Island. 99 . She plans to continue work on commercially important marine species. the author decided to pursue a graduate-level education to further her interest in marine fisheries. species of special concern. as a staff biologist with the Sarasota Dolphin Research Program.BIOGRAPHICAL SKETCH Born on April 15. the author showed a great curiosity for various aquatic organisms and their environments at an early age. South Carolina. in Grand Rapids. Michigan. Florida.