Gulf of Mexico Science Volume 26 Article 3 Number 1 Number 1

2008 Dietary Analysis of Batfishes (Lophiiformes: Ogcocephalidae) in the Gulf of Mexico Bronson H. Nagareda Anuenue Fisheries Research Center

Jonathan M. Shenker Florida Institute of Technology

DOI: 10.18785/goms.2601.03 Follow this and additional works at: https://aquila.usm.edu/goms

Recommended Citation Nagareda, B. H. and J. M. Shenker. 2008. Dietary Analysis of Batfishes (Lophiiformes: Ogcocephalidae) in the Gulf of Mexico. Gulf of Mexico Science 26 (1). Retrieved from https://aquila.usm.edu/goms/vol26/iss1/3

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Dietary Analysis of Batfishes (Lophiifonnes: Ogcocephalidae) in the Gulf of Mexico

BRONSON H. NAGAREDA AND jONATHAN M, SHENKER

Stomach content analyses, perfomted on three species of batftshes, Halieutic!Jtl.tys ac-uleatus, Ogcocep!Jalus decllviroshis, and Ogcocep!Jalm pantosticus collected in tlte Gulf of Mexico in summer (Jlme-July) and fall (Oct.-Nov.) 2002 and 2003, revealed a variety of benthic invertebrates, particularly gastropods, wonns, and xanthid crabs. Schoener's dietary overlap indices (Sl) were calculated between the three species within the same seasons, and within each species between seasons. SI values indicated that each species consumed a different assemblage of prey and that two of the species exhibited temporal variation in diet.

he (order Lophiiformes) have attractant rather than acting as a visual stimulus T evolved some of the most unusual morpho· for prey. logical and ecological adaptations in the tremen· All lophiiform fishes stud!ed to date arc dously diverse and varied clade of bony fishes primarily piscivorous (Bertelsen, 1951; Randall, (class Osteichthyes) and are among the most 1967; Bigelow and Schroeder, 1953; Pietsch and specialized groups of fishes. The defining feature Grobecker, 1987), utilizing their lures to regu· of the angletiishes is the modified first dorsal-fin larly obtain food. However, the few existing spine (illicium) and terminal bait (esca) that studies on the diets of batfishes suggest a senres as an apparatus to attract prey. The different feeding strategy. Rather than the batfishes, family Ogcocephalidae, represent one obvious adaptations for piscivorous macrophagy of 18 families within the order, presently of nearly all other lophiiform taxa, exemplified containing 10 genera and 68 species, of which best by their large anterodorsally directed five genera and 15 species are known from the mouths, ogcocephalids possess small, ventrally western Atlantic Ocean (Bradbury, 2003). It is a directed mouths, small villiform teeth, and a poorly known group of small ( < 300 mm) short gut, all adaptations for the capture of small benthic fishes found worldwide in tropical and demersal prey (durophagy), such as gastropods, subtropical seas, from shallow inshore waters to small crustaceans, and polychaete worms (Woot· depths as great as 3,000 m. They generally reside ton, 1994; McEachran and Fechhelm, 1998; on continental shelves and slopes, on flat, Gibran and Castro, I999). In an effort to relatively open-bottom habitats of rubble, sand, demonstrate these trophic adaptations, as a and mud (Bradbmy, 1980, Richards and Brad­ unique approach to feeding among anglerfishes, bury, 1999). dietary analyses assessing numerical importance Previous studies of ogcocephalids have fo· and numerical similarities were conducted on cused primarily on and systematics three species of batfishes (H. acu/eatus, OgcocejJh­ (Hubbs, 1958; Bradbury, 1967, 1980, 1988, 1998, alus declivirostlis Bradbury, 1980; and I 999; Bradbury et al., I 999; En do and Shinohara, pantostictus Bradbury, 1980) commonly found on I999), although ogcocephalids have been briefly the continental shelf of the northern and discussed in a few studies of the trophic ecology western Gulf of Mexico. of a given region (e.g., Reid, 1954; Randall, 1967). Of the few studies on the natural history METHODS of batfishes, \Vinans (1975) examined the stomach contents of OgcocejJ!wlus rostellum Brad­ Specimens were obtained during the National bmy, 1980, and Ha/ieutichtii)'S aculealus (rvlitchell, Oceanic and Atmospheric Administration/Na­ 1818) off Cape Canaveral, Florida; and Gibran tional :Marine Fisheries Service (NOAA/N:MFS) and Castro (1999) examined the stomach biannual groundfish survey between Brownsville, contents of Ogcocephalus ve..spntilio (Linnaeus, TX, and Pascagoula, MS, conducted by the R/V 1758) at Sao Sebastiao Channel, in southeastern Oregon II in June-July and Oct.-Nov. 2002 and Brazil. A histological study of the esca of 2003. Trawls were made at randomly selected Ogcocephalus cubifrons (Richardson, 1836) was depth-stratified stations betv-leen 12 and 90 m, conducted by Combs (1973) who hypothesized using a 16.8·m net otter trawl (2.54-cm mesh), that the glandular esca secreted a chemical with gear towed at an approximate speed of

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c 100% liB teleosts ~ 80% ~bryozoans a.0"' E 60% LS pelecypods 0 0 40% o:t ~ D crustaceans cu 20% .!!!,- 11 gastropods 0% c -Q) '- :::: '- :::: '- :::: ~ -Ill -Ill ~ c..Q) i(/) i(/) i(/) Halieutichthys Ogcocephalus Ogcocephalus acu/eatus declivirostris pantostictus Batfish species

Fig. 1. Comparison of the dietary composition (%) of three species of batfishcs taken from NOAA/Ni\Jl

1.3 m/sec for no less than 10 min, but not more indicate significant dieta1y overlaps (Zaret and than 55 min. During the survey, batfishes were Rand, 1971; Mathnr, 1977). opporltmistically collected and frozen aboard ship for subsequent analysis. Once back at the REsuLTS laboratOI)', fishes were thawed, standard length and sex recorded, and stomach contents exam­ The stomachs of H. aculeatus [n = 230, 37- ined under a dissecting microscope. Prey items 72 mm standard length (SL)] (Table 1) con­ were counted, identified to the lowest possible tained primarily polychaetes (Nereidae), small taxon, and identified by H. Perry (Gulf Coast gastropods (a species of Nassmius thought to be Research Laboratory, University of Southern undescribed; H. Peny, pers. comm.), and small .ivlississippi) and D. Shelton (Alabama J\·Ialaco­ shrimps (Mysidae), although two additional logical Research Center). species of gastropods and four other crustaceans Prey consumption was summadzed in tenns of were taken in low abundance (Fig. 1, Tables 1 percentage of frequency of occurrence (%F; and 2). Nassmius sp. was consumed only in the proportion of stomachs containing a specific prey summer, whereas the other two dominant prey item) and percentage of the total numerical taxa occurred in both the summer and fall. This dietary composition (%N; proportion of the seasonal absence of gastropod prey is reflected in number of a specific prey item to the total number an SI value between the summer and fall samples of prey items in all stomachs examined). Scho­ of 0.568 (Table 3). Relatively few prey items were ener's (1970) dietaty overlap index (SI) was found in any one stomach, with a maximum of calculated to assess dietary overlap among the six gastropods and two polychaetes per stomach, three co-occurring species ·within the same seasons although one contained 20 mysids. The hard­ (summer and fall) and within each species shelled prey items recovered were all small, between seasons (:Mathur, 1977), as follows: approximately 1-'1 nun in maximum dimension. The stomachs of 0. derlivirostJis (n = 168, 25- n 1<16 mm SL) (Table 1) contained a higher species diversity than those of H. aculeatus, but a ~ I - o.s(z= IP,, - P,,l) the prey were dominated by small (1-5 mm) I~ I gastropods, predominately Cosmioconcha calli­ glypta (Dall and Simpson, 1901) and Nassmius where Pxi was the dietaty composition proportion sp., two taxa that comprised over 90% of the diet of food categ01y i in the diet of sample .\~ P>b the (Fig. 1, Table 4), with as many as 28 gastropods dietary composition proportion of food category i found in a single stomach. Nereid polychaetes, in the diet of sample); and n, the number of food pel}•cepods, crustaceans, and bryozoans were also categories. SI values of ~ 0.60 were taken to found. Two specimens contained the shrimp eel https://aquila.usm.edu/goms/vol26/iss1/3 2 DOI: 10.18785/goms.2601.03 Nagareda and Shenker: Dietary Analysis of Batfishes (Lophiiformes: Ogcocephalidae) in t 30 GULF OF MEXICO SCIENCE, 2008, VOL. 26(I)

TABLE 1. Sample sizes and standard lengths of three species of ogcocephalids from the Gulf of Mexico. Summer samples were collected in June and Julr, fall samples in Oct. and Nov., 2002 and 2003. n = Number of fishes collected; SL = standard length.

Species n 51. (mm) Ucan SL (mm) HalieutichthJS aculeatus Summer sample 125 49-67 58.2 Fall sample 105 37-72 53.5 Ogcocephalus declivirostris Summer sample 93 43-115 70.9 Fall sample 75 25-146 62.5 Ogcocephalus pantostictus Summer sample 26 47-275 190.1 Fall sample 11 39-268 145.6

Ophicthus gomesii (Castelnau, 1855). The diets of hivitlatum (Valenciennes, 1828), ·were found in a 0. declivirostlis remained consistent, ·with summer 224-mm specimen, and small shrimp eels ( 0. and fall collections having a SI value of 0.637 gomesit) were found in the stomachs of several 0. (Table 3). pantostictusspecimens (176-275 mm SL). The diet of 0. pantfJstictus (n = 37, 39-275 mm There was no significant dietary overlap (SI ~ SL) (Table 1) was dominated by xanthid crabs in 0.60) between the three species in either the summer samples, and the lesser blue crab summer or fall (Table 3). In summer, H. Callinecles simi/is (\Villiams, 1966) and the gastro­ aculeatus showed SI values of 0.383 and 0.153, pod C. caUigl)pta in fall samples (Fig. 2, Table 5). with 0. declivirosl1is and 0. panlostictus, respec­ Benveen seasons, 0. jJantostictus had anSI value of tively. Ogcocephalus declivirosttis and 0. pantostictus 0.376. Prey items "\vere generally larger (10- also showed very low dietary overlap vl'ith an SI 25 mm) than those of the other two species, value of 0.222 in the summer. Fall samples although most gastropod prey was similar in size showed low dietary overlap as well, with H (1-5 mm) than those found in the stomachs of 0. aculeatus having SI values of 0.206 and 0.241 declivimshis. Unlike the other species examined, 0. between 0. declivirost1is and 0. pantosticlus. The pantostictus occasionally consumed fish: the re­ overlap in diet between 0. declivirosl1is and 0. mains of a 51-mm dwarf sand perch, DifJleclntm pantostictus in the fall increased somewhat,

TABLE 2. Stomach contents of llalieutichthJs aculeatuscollcctcd in the Gulf of Mexico in june-July and Oct.-Nov., 2002 and 2003. n = Number of stomachs examined; %F = frequency of occunence; %N = numerical dieta1y composition; Max = maximum number of individuals of prey type in a gut.

June-July Oct.--i\'o\'. n 125 n 105 ~~ .1 Prey taxa %F Max %N %F Max %N ) Polychaeta Nereidac 22.40 2 15.42 22.86 27.91 Gastropoda Nassmius sp. 24.8 6 26.06 C)'lichnella bidentala 14.40 3 11.70 0.10 2 2.33 Cosmiocoudta mlligl;jJ!a 4.80 2 3.72 0.10 1.16 Pelecypoda Nuculana awta 5.60 2 4.26 Crustacea Xanthidae 9.60 3 9.04 8.57 1 10.57 Stomatopoda 4.00 2.66 2.85 3.49 Isopoda 4.80 3.19 Amphipoda Mysidacea 27.2 3 22.3 14.29 20 54.65 Pagunts spp. 1.60 2 1.60

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TAnLE 3. Schoener dietary overlap indices for three species ofbatfishes collected in the Gulf of Mexico. Values in bold indicate intraspecific comparisons between summer (June-July, 2002 and 2003) and fall (Oct.-Nov., 2002 and 2003) samples. Values above the bold print indicate interspecific comparisons from fall collections, Values below the bold print indicate interspecific comparisons from summer collections,

llafinaichthys awleaiiH Ogroaphalus pantMrictus

Halieutichth)'S awlmtus 0.568 0.206 0.241 Ogcocephalus dedivirost1is 0.383 0.637 0.459 Ogcocep!wlus Jmntoslictus 0.153 0.222 0.376

although not to any significant level (SI species and between seasons within a species OA59). (except 0. decliviros/Jis; see Table 3) suggesting there is relatively little overlap in diets among

DISCUSSION species in the Gulf of 1-lexico. This may be in part because of spatial habitat differences as Ogcoccphalids are an exception to the pri­ batfish species do not regularly co-occur in trawl marily macrophagus, piscivorous feeding habits collections. Most of the lowest dietat)' overlap of other lophiiform fishes. In support of the few values occurred benveen H. aculeatus and both existing studies of batfish feeding (e.g., 'Vinans, species of Ogcoceplwlus. Halieutichth)'S aculeatus 1975; Gibran and Castro, 1999), batfishes in this rarely occurred in trawls with 0. declivirostJis and study consumed almost exclusively small benthic never with 0. pantostictus. Furthermore, 0. invertebrates such as small snails, shrimps, crabs, declivimsl1is and 0. pantostictus also rarely oc­ and polychaetes. The three species of batfish in curred in the same collections. The SI values this study fed on somev.,hat different assemblages betv-:een these two species ·were low in the of prey. Stomach contents of fl. aculeatus were summer but increased (although not significant­ dominated by nereid polychaetes and mysids. ly) in the fall. This was reflected by the increase Ogcocep!wlus declivirost1is consumed negligible in diet of C. calliglypta by 0. pantostictus in the fall amounts of polychaetes and no mysids, instead (Tables 3 and 5). The summer SI value between feeding primarily on gastropods (Nassmius sp. H. aculeatus and 0. declivirostris was higher than and C. ca.Zligl)1Jla) and small xanthid crabs. most other values between species and this Although there was some dietary overlap, with resulted from the presence of Nassmius sp. in both species consuming Nassarius sp. in the the diet of H. a.culealus during the summer. summer, the majority of the diet was vet)' Although spatial habitat differences between different. Ogcocephalus Jmntostictus fed primarily species may explain the large differences in on crabs (Xanthidae and C. similis) and gastro­ dietary overlap values, in some collections pods (C. calligl)jila). Small fishes [e.g., shrimp batfi:shes co-occurred (II. aculeatus/ 0. dediv-iros­ eels ( 0. gomesil) and dwatf sand perch (D. t1is and 0. declivirosttis/ 0. pantostictus). Despite bivittatum)] were rarely found inside the stom~ this co-occurrence, the SI values between species achs of 0. declivirostds and 0. pantostictus, and remained low. This may reflect prey resource their overall contribution to the diet appears partitioning by the different batfishes where they limited. Larger individuals of 0. Jmntostictus co-occur. Seasonal changes in some of the (> 215 mm SL) and two 0. dedivirosllis (89 and available prey species of these fishes may also 1•16 mm SL) were the only specimens found to account for the differences in diet, although the contain fishes as prey items. 'While small fishes possibilily cannot be discounted that that these may not be a regular prey item, it is not differences simply reflect different microhabitats surprising to find them in the stomachs of large sampled by individual trawls. batfishes as an opportunistic prey item. In Only 0. dedivirosl1is showed a persistent and addition to co-occurring in the same habitat, significant dietary similarity benveen the summer small fishes may mistake batfishes for a foraging and fall samples; its primary prey of gastropods or shelter structure in the same manner pro­ remained consistent between seasons. The SI posed for frogfishes by Pietsch and Grobecker value of 0.568 for H. aculeatus bcD.veen seasons (1987). indicated some similarity as it approached the Although some prey taxa were commonly 0.60 threshold value (Zaret and Rand, 1971; consumed by all three species, the consumption lvfathur, 1977) to indicate significant dietary of other prey types resulted in low values ( < 0.60) overlap. Dominant prey items such as nereid of Schoener's dietmy overlap index among polychaetes and mysids were consistent in the https://aquila.usm.edu/goms/vol26/iss1/3 4 DOI: 10.18785/goms.2601.03 Nagareda and Shenker: Dietary Analysis of Batfishes (Lophiiformes: Ogcocephalidae) in t 32 GULF OF MEXICO SCIENCE, 2008, VOL. 26(1)

TAnLE 4. Stomach contents of Ogcncephalus d.eclivirosltis collected in the Gulf of Mexico in June-July and Oct.- Nov., 2002 and 2003. n = Number of stomachs examined; %F = frequency of occurrence; %N = numerical dietary composition; Max = maximum number of prey type in a gut.

Summer Fall

n 93 n 75

Prey taxa %F Max %N %F Max %N Polychacta Ncrcidac 4.3D D.9D 10.67 2 6.77 Gastropoda Nassmius sp. 26.88 18 23.67 16.DD 2D 24.81 C)·lidmella bidentata ID.75 9 6.18 6.67 2 5.26 Cosmioconclla calliglypta 35.48 21 31.56 14.67 5 14.29 Niso aeglees 2.15 I DAD 4.0D 2.26 Polyslira albida 15.05 17 ID.87 6.67 3.76 Kurlzie/la sp. 8.6 28 8.74 4.DD I 3.DI Terebra areas 12.9D 6 4.D5 1.33 I 0.75 Natica marochiensis 2.15 3 l.D7 1.33 0.75 Compsodti.llia sp. 12.DD 3 14.89 Pelecypoda Argopecten sp. 1.08 7 1.49 Nuculana acuta 12.9D 3 3.2D 5.33 2 3.76 Cntstacea Xanthidae 31.51 2 6.37 16.DD 4 12.78 Raninoides sp. 2.15 DAO 2.67 3 3.76 Tmch)jmweus similis 1.33 0.75 Pagurus spp. l.D8 D.2D Bryozoa l.D8 I D.2D 1.33 2 1.5D Teleostei Opldchthus gomesii 2.67 1.5D

diet of this species between seasons; however, also contained larger prey items than those of Nassarius sp. was a numerically important dietary the other two species studied. This may be a item only in the summer samples of H. acu./eatus, result of the larger size of specimens of 0. as it was absent in fall samples. This may reflect pantostictus collected in the study (summer prey availability differences; however, Nassmius mean length, 190.1 mm; fall mean length, sp. was consistent in the diet of 0. dedivirostris 145.6 nun). beaveen seasons suggesting it may be available Size, howe\'er, may not necessarily be a major throughout the year as a potential prey item and factor in the size and type of prey taken. that resource partitioning occurs between these Nagareda (2005) analyzed a sample of 0. two species. Alternatively, if spatial habitat declivimstds for relationships between prey size differences are normally an important factor in and batfish body and mouth size. The results batfish diets, the increase of Nassmius sp. in the showed only a slight increase in prey size with diet H. aculeatus may indicate increased sympatry increasing predator body size and mouth gape. with 0. declivirosl1is during the summer. Prey Although the gape parameters increased with consumption of 0. pantosliclus changed consid­ increasing SL, this increase was not reflected in erably between summer and fall. This species the consumption of significantly larger prey. may be taking advantage of differences in Except for polrchaetes and shrimp eels, all prey recruitment and availability of prey by taking taken were short in length, with a maximum xanthid crabs in the summer and then switching cross-sectional dimension of less than 5 mm. to C. similis and C. calligl;pta in the fall. A review Despite being longer prey items, polychaetes and of unpublished NOAA/NMFS collection data snake eels are soft and easily compressible, and suggests that small juvenile C. similis were more had cross-sectional measurements similar to abundant in fall trawl collections. A more those of the gastropods and crustaceans taken intensive and focused study on C. similis may by the batfish. Other constraints, including the help determine whether this increase is because availability of prey of larger sizes, or other of recruitment. The stomachs of 0. pantostictus morphological constraints such as pha1yngeal

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TARLE 5. Stomach contents of Ogcocephalus pantostictus collected in the Gulf of Mexico in June-Julr and Oct.­ Nov., 2002 and 2003. n = Number of stomachs examined; %F = frequency of occurrence; %N = numerical dietary composition; Max = maximum number of individuals of prey type in a gut.

Summer Fall

n 26 n II Prey ta_xa %F Max %N %F Ma_x %N Polychacta Ncreidae 3.85 I 2.44 9.09 3 4.17 Gastropoda Nassmius sp. 3.85 I 2.44 18.18 I 2.78 Cosmiocmuha callig!Jpla 3.85 4 9.76 45.45 8 30.56 Kwtz.iella sp. 27.27 3 9.72 Terebra areas 9.09 3 9.72 Nafica mamchiensis 3.85 2.44 Oliva Sa)'ana 7.69 4 12.20 Crustacea Xanthidae 19.23 4 24.39 Raninoides sp. 9.09 1.39 Cnllinectes similis 54.5

gape, may affect prey selection. The latter may be sumed the gastropod Nassmius consensus (Rave­ a significant constraint in batfish feeding be­ nel, 1861) and, in the cooler months, the scallop cause batfishes regularly take prey with hard Argopecten gibbus (Linnaeus, 1758). Winans (1975) exoskeletons or shells and do not crush them to suggested that the minimal overlap of prey items get to the soft tissues. Further investigation in between the sympatric batfishes was because of this aspect of batfish feeding biology may help resource partitioning. Although spatial habitat determine what morphological limitations may differences in this study appeared to be a more exist in batfish feeding. important factor in the different diets of the Gibran and Castro (1999) suggested that species studied, the limited co-occurrence and gastropod shells recovered from the stomachs low dietary overlap of these species also suggest of batfish reflected predation upon mobile that some resource partitioning occurs between hermit crabs. In contrast to their study, almost batfishes in the Gulf of :Mexico as well. Although all of the gastropod prey analyzed here included the summer samples for H. aculeatus and 0. easily recognizable soft parts (antennae, siphons, dedivirostlis in this study both showed similar opercula), demonstrating that the batfishes amounts of Nassmius sp. taken as prey, the preyed on the gastropods themselves and not primary types of prey taken by H. aculeatus on hermit crabs. Compared to the overall (polychaete worms, small crustaceans) and the number of gastropods, relatively few hermit two species of Ogcoceplwlus (gastropods) v.. 'erc crabs (Pagums sp.) were found in the stomachs similar in both studies. 'Winans (1 975) interpreted of the batfishes studied. They appear to be minor his observations in the diet shift of 0. rostellum components of the diets of I!. aculeatus and 0. from Nassmius to Argopecten as an adaptation to a dedivirostlis; however, Pagunts sp. was a moder­ locally abundant prey source because the shift ately numerically important prey item in the diet coincided with scallop recruitment. In this study, of summer-sampled 0. pantostictus. only 0. declivirostris showed a significantly consis~ 'Vinans (1975) investigated the sympatric spe­ tent diet between seasons. Although the SI value cies 0. rostellum and II. aculeatus off Cape was not significant, tvm of the three primary prey Canaveral, FL, and found that the diet of H. items for H aculeatus remained consistent be~ aculeatus there also consisted primmily of nereid tween seasons, although the lack of Nassmius sp. polychaetes, whereas 0. rostellum primarily con- in fall samples for this species may indicate a diet https://aquila.usm.edu/goms/vol26/iss1/3 6 DOI: 10.18785/goms.2601.03 Nagareda and Shenker: Dietary Analysis of Batfishes (Lophiiformes: Ogcocephalidae) in t 34 GULF OF MEXICO SCIENCE, 2008, VOL. 26(1)

shift for this species. Ogcoceplwlus pantostictus was ---. 1998. A new species of Maltlwpsis (Lophii­ the only batfish in this study that appeared to formes: Ogcocephalidae) from the westem Atlantic show a diet shift to possibly take advantage of the Ocean. Bull. MaL Sci. 63:207-211. changes in its available prey bet\veen seasons. ---. 1999. A review of the fish genus Dibranchus, with descriptions of new species and a new genus, Benthic sampling of the invertebrates and prey Snlncisquama (Lophiiformes: Ogcocephalidae). Proc. species in the same habitat as the batfishes may Calif. Acad. Sci. 51 (5):259-310. help clarify whether these fishes are taking prey ---. 2003. Family Ogcocephalidae Jordan 1895- because of availability or selection. Such studies batfishes. Calif. Acad. Sci. Annat. Checklists Fish. may compare the feeding biology of batfish 17:H7. species taken at different locations for similari­ ---,J. E. McCosKl:R, A.'.TI W. C. SCHROEDER. 1953. Fishes of the .A3441 no. 417. Gulf of Maine, U.S. Fish Wild!. Sen•. Fish. Bull. 53:1-577. ScHOENER, T. W. 1970. Nonsynchronous spatial overlap llRAOI\URY, M. G. 1967. The genera of batfishes (family of lizards in patchy habitats. Ecology 51:408-418. Ogcocephalidae). Copeia 1967:399-422. WINA.'\S, G. A. 1975. Food habits of two sympatric ---. 1980. A revision of the fish genus Ogcacephalus, batfishcs (Ogcoccphalidae) offshore Cape Canav­ with descriptions of new species from the western eral, Florida. Unpubl. M.S. thesis, Florida Institute of North Atlantic Ocean (Ogcocephalidae; Lophii­ Technology, Melbourne, FL. formes). Proc, Calif. Acad. Sci. 42(7):229-285. Woonox, R. J. 1999. Ecology of teleost fishes Kluwer ---. 1988. Rare fishes of the deep-sea genus Academic Publishers, Dordrecht, The Netherlands. Halieutopsis. a review with descriptions of four new ZARET, T. M., ANI> A. S. "&\Nil. 1971. Competition in species (Lophiifonnes: Ogcocephalidac). Fieldiana tropical stream fishes: support for the competitive Zoo!. (n.s.) 44:1-22. exclusion principle. Ecology 52:336--342.

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(BHN) DErARTIIL'\fTOF L-\_l'\'DAt"\'D NATURAL REsoURCF.S, OF BIOLOGICAL SciEI\'CES, fLORIDA JNSTO""UIE OF DIVISION oF AQUATIC REsouRcES, A"\'uENUE fiSHERIES TECHNOLOGY, 150 "'\VEST UNIVERSIIT BOULE."YARD, RESEARCH CENTER, 1039 SAND lSL\J"\'ll PARKWAY, ~vfELBOU&"\'E, FLORIDA 32901. Date accepted: Oc­ HONOLULU, I-L\WAII 96819;AND {JMS) DEPARTMEl\'T tober 27, 2008

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