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Reconstruction of Original Body Size and Estimation of Allometric 101 Reconstruction of original body size tie length in squid) in population and other types of surveys, information and estimation of allometric relationships on total length may not always be for the Iongtin inshore squid <Loligo pealeit1 readily available. Therefore, knowl­ edge of allometric relationships may and northern shortfin squid (///ex il/ecebrosus> be useful to. accurately assess trophic interactions and predator-prey rela­ Michelle D. Staudinger' tionships. For the majority of ceph­ alopod species, there are currently 1 Francis Juanes no predictive equations to estimate Suzanne Carlson2 total length from mantle length and Email address for contact author: [email protected] to account for variability in growth. 1 Department of Natural Resources Conservation To improve descriptions of the feed­ University of Massachusetts Amherst ing habits of teuthophagous preda­ Amherst, Massachusetts, 01003-9285 tors and to increase the number of 2 School of Natural Science evaluations of size-based predation Hampshire CoRege on cephalopod prey we present 1) pre­ Amherst, Massachusetts, 01002 dictive equations for reconstructing original prey size and 2) allometric relationships of mantle length to total body length for the two most common species of cephalopods in the Northwest Atlantic Ocean, L. pealeii Quantification of predator-prey body mammals (Gannon et al., 1997; Wil­ and I. illecebrosus. size relationships is essential to under­ liams, 1999). standing trophic dynamics in marine As with the bones and otoliths of ecosystems. Prey lengths recovered prey fish, cephalopod beaks are often Materials and methods from predator stomachs help deter­ recovered from predator stomachs mine the sizes of prey most influential and may be used for identification of Loligo pealeii were collected by otter in supporting predator growth and prey species and the reconstruction trawl from coastal waters off Mas­ to ascertain size-specific effects of of original prey body size (Clarke, sachusetts during the months of May natural mortality on prey populations 1986). Many predators (e.g., marine through September of 2006 and 2007. (Bax, 1998; Claessen et al., 2002). mammals) cannot digest the chitin­ Illex illecebrosus were collected from Estimating prey size from stomach ous beaks and thousands of beaks outer shelf waters from New Jersey content analyses is often hindered may accumulate in the stomachs to North Carolina during February because of the degradation of tissue until they are regurgitated (Clarke, 2007 on both the winter and spring and bone by digestion. Furthermore, 1980). Predictive equations for es­ bottom trawl surveys conducted by reconstruction of original prey size timating body size in the two most the National Marine Fisheries Ser­ from digested remains requires spe­ common species of cephalopods in the vice (Northeast Fisheries Science cies-specific reference materials and Northwest Atlantic Ocean (Bowman Center) (Azarovitz, 1981). All squid techniques. A number of diagnostic et al., 2000) are either based on few were preserved by freezing until guides for freshwater (Hansel et al., observations (n=25) as seen in the they were processed in the labora­ 1988) and marine <Watt et al., 1997; longfin inshore squid (Loligo pealeii) tory. Specimens were thawed to room Granadeiro and Silva, 2000) prey spe­ (Gannon et al., 1997), or are nonexis­ temperature and then measured for cies exist; however they are limited tent as is true for the northern short­ dorsal mantle length (DML), total to specific geographic regions (Smale fin squid (lllex illecebrosus). length (TL), and maximum length et al., 1995; Gosztonyi et al., 2007). Trophic niche breadth is the range (LMax) to the nearest 1.0 millimeter. Predictive equations for reconstruct­ of relative prey sizes consumed onto­ Dorsal mantle length was measured ing original prey size from diag­ genetically by a predator (Scharf et as the distance between the posterior nostic bones in marine fishes have al., 2000). In previous diet studies, been developed in several studies of trophic niche breadth has been used piscivorous fishes of the Northwest to predict shifts in foraging modes Atlantic Ocean (Scharf et al., 1998; and physical limitations on feeding Manuscript submitted 2 June 2008. Wood, 2005). Conversely, morphomet­ patterns (Bethea et al., 2004; Beau­ Manuscript accepted 28 August 2008. ric relationships for cephalopods in champ et al., 2007). Calculation of Fish. Bull. 107:101-105 (2009). this region are scarce despite their trophic niche breadth requires mea­ importance to a wide range of preda­ surements of the total lengths of The views and opinions expressed tors, such as finfish (Bowman et al., or implied in this article are those predators and prey. Depending on of the author and do not necessarily 2000; Staudinger, 2006), elasmo­ how a species is traditionally mea­ reflect the poeition of theNational branchs (Kohler, 1987), and marine sured (e.g., fork length in fish, man- Marine Fisheries Service, NOAA. 102 Fishery Bulletin 107(1) DML TL Figure 1 _ Morphological features used for measuring dorsal mantle length (DML), total length (TL), and maximum length (LMax> of squid. and anterior tips of the dorsal side of the mantle, total L. pealeii and I. illecebrosus. The r2 values for all body length was measured as the distance between the pos­ size relationships ranged from 0.88 to 0.98 and were terior tip of the mantle to the end of the longest arm, highly significant <P<O.OOOl) (Table 1). A total of 434 L. and maximum length was measured as the distance pealeii ranging in size from 1.9 to 28.0 em (DML) and between the posterior tip of the mantle to the end of 158 I. illecebrosus ranging in size from 4.4 to 28.4 ern the longest tentacle (Fig. 1). Beaks were extracted from (DML) were measured to develop allometric relationships the buccal mass and the lower rostrum length (LRL) of between DML, TL, and LMax· the lower beak was measured to the nearest 0.01 mil­ Equations for reconstructing original squid size limeter. Lower beaks were held so that the rostrum tip (DML) from lower rostrum lengths (LRL) were highly was facing the observer and then turned to a left-facing significant (P<0.0001) in both L. pealeii and I. illece­ orientation <Fig. 2); the lower beak was best viewed brosus (Table 1). The model developed for L. pealeii when held against a white background for contrast. The improved the only known equation for this species by LRL of L. pealeii was measured by placing the tip of the expanding the sample size from n;;::25 and the coeffi­ moving arm of the calipers inside the jaw angle of the cient of determination (r2) of 0.73 (Gannon et al., 1997) lower beak and extending it to the tip of the rostrum to n=144 and an r2 of 0.83 (Table 1). Lower rostrum (Clarke, 1986) (Fig. 2A). The LRL of I. illecebrosus lengths ( LRL) were measured from L. pealeii rang­ was measured from the tip of the rostrum to the jaw ing from 2.6 to 24.7 em (DML). The predictive model angle. In I. illecebrosus the shoulder forms a tooth which for estimating DML from LRL in I. illecebrosus was facilitates location of the jaw angle (Fig. 2B). Beaks from developed from 89 specimens ranging from 4.4 to 28.4 both species were measured either under a dissecting em (DML). The relationship between LRL and DML in microscope or a magnifying glass. I. illecebrosus was stronger and less variable (r2=0.94, Least squares regression analysis was used to esti­ coefficient of variation [CV]=8.15) in comparison to L. mate the relationship between mantle length and total pealeii (rZ=0.83, CV=15.94). Measurement of the lower length, mantle length and maximum length, and LRL rostral length in I. illecebrosus is greatly facilitated by and mantle length for both squid species. Using the the presence of a tooth located in the angle point. This PROC UNIVARIATE command in SAS, vers. 9.1 (SAS structure is absent in the lower beak of L. pealeii, and Institute Inc., Cary, NC), we found that all variables may make measuring beaks from this species more dif­ were in compliance with assumptions of normality and ficult and prone to error. no outliers were detected. Linear models were used to develop predictive equations for all pairings of morpho­ logical structures. All statistical analyses were per­ Discussion formed by using the PROC REG command in SAS. The results of the present study are intended to assist and encourage quantitative assessments of cephalo­ Results pod prey in the diets of a broad range of finfish, elas­ mobranch, marine mammal, and seabird predators. Total length (TL) and maximum length (LMax> were Although methods for identification and reconstruc­ strongly related to dorsal mantle length (DML) in both tion of original body size from cephalopod beaks have NOTE Staudinger et al.: Reconstruction or original body size and allometric relationships ror Loligo peofeii and fffex iffecebrosus 103 Rostrum tip A R o~; trum tip B I / .lawttngl.: Jaw angle ,' :' Shoulder ·~. ,/ Shoulder Figure 2 Orientation and key morphological structures of the lower beak of <Al the longfin Inshore (Loligo pealeii! and (8 ) the northern shortfin llllex illecebro~tus! squids. Lower rostrum length (LRLJ Ia measured from the tip of the rostrum to the jaw angle. Table 1 Least squares regression equations for describing the relationship of total length <TL) to dorsal mantle length (D.ML) and maxi­ mum length (ly..,.), and the relation ofDML to the lower rostral length of the lower beak (LRL) In Iongtin inshore <Loligo pealeii! and
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