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Food and Prey Detection in Two Deep-Sea Cephalopods: Opisthoteuthis Agassizi and O

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Food and Prey Detection in Two Deep-Sea Cephalopods: Opisthoteuthis Agassizi and O BULLETINOFMARINESCIENCE,49(1-2): 288-299,1991 FOOD AND PREY DETECTION IN TWO DEEP-SEA CEPHALOPODS: OPISTHOTEUTHIS AGASSIZI AND O. VOSSI (OCTOPODA: CIRRATA) Roger Villanueva and Angel Guerra ABSTRACT The diet of Opisthoteuthis agassizi and O. vossi in the southeast Atlantic was studied from 171 and 121 individuals respectively. Small epibenthic and supra benthic crustaceans and polychaetes are the most frequent prey items in both species, suggesting that they feed on suprabenthic and epib<:nthic material. Diel analysis of feeding by O. agassizi at 490 m and O. vossi at 836 m depth demonstrated a pattern of continuous feeding. Relationships oftotal body length and beak measurements to total weight were also studied. The ultrastructure of sucker and cirri are described for both species and their relationship with prey detection mechanisms is discussed. The cirrate octopods represent the oldest evolutionary lineage of octopods, adapted to the deep-sea environment, where they have been recorded at depths of up to 7,229 m (Voss, 1988a; Aldred et al., 1983). The trophic position of these octo pods in the benthopelagic ecosystem is still unknown. The aim of the present study was to provide some insight onto the feeding patterns of the cirrates, which live in an area where light intensity and temperature are characterized by their low levels and lack of drastic changes. Furthermore, sucker and cirrus ultrastruc- ture was investigated with the aim of relating their morphology to prey detection and capture. Two Opisthoteuthis species with overlapping distributions in Na- mibian waters (SE Atlantic) were selected: O. agassizi and O. vossi. Opisthoteuthis agassizi Verrill, 1883 is distributed on both sides of the Atlantic: from the Caribbean Sea to north of Brazil (Voss, 1988b), from the south coast of Ireland to the north coast of Spain and the Mediterranean Sea and from the Gulf of Guinea to Namibian waters (Morales, 1959; Adam, 1962; Sanchez, 1988; Voss, 1988b). Depth of capture ranges between 452 m (present study) and 2,250 m (Verrill, 1885). Opisthoteuthis vossi Sanchez and Guerra, 1989 is a species that lives off Namibian waters at 900 m depth. MATERIALS AND METHODS Sample Collection. -A total of 186 individuals of O. agassizi and 124 of 0, vossi were collected in Namibian waters (Table I Fig. I). Specimens were found in 42 stations during the scientific surveys Benguela-XIII (July, 1988), Benguela-XIV (January, 1989), Benguela-XV (July, 1989), and Benguela- XVI (March, 1990) on board the research ship CHICHA- TouZA and during commercial surveys on board the Freezer Trawler HERMANOSTOUZA(March, 1989) and the Freezer Trawler ITXAS-LuR(April, 1989). Depth ranges varied from 452 to 823 m for O. agassizi and from 778 to 888 m for O. vossi. Samples were taken by means of bottom trawls. Abundances in individuals per square kilometer were obtained from knowledge of the trawling device area and trawling speeds. Two time periods were sampled to study feeding rhythms. O. agassizi was sampled at 25°43'S, 13°40'E, between 483 and 490 m depth, during a 32-h cycle. Seven samples were collected: from 0920-1250, 1355-1620, 1730-2030, 22 I5-{)120, 022Q-{)600, 0710-1055, and 1300-1700 hr., 28-29 March 1989. A total of79 specimens was collected. O. vossi was sampled at 23°52'S, 13°1'E, at depths between 829 and 836 m. Four samples were collected: from 1645-1845, 230Q-{)100, 050Q-{)700 and 0950-1150 hr., 2-3 March 1990. A total of 96 individuals was collected. Immediately after collection, dorsal mantle length (DML), total length (TL) and total weight (TW) were measured for all individuals and their sex recorded. Individuals were dissected and their stomachs stored in 70% ethyl alcohol for further analysis. Mandibles of selected individuals were also dissected and fixed in 70% ethyl alcohol for upper crest length (DC) and lower crest length (LC) measurement. 288 VILLANUEVA AND GUERRA: FEEDING ECOLOGY IN Op/STHOTEUTHIS 289 Table 1. Opistoteuthis agassizi and O. vossi. Material examined. DML and TL in mm, TW in g. (SD, standard deviation; n, number of specimens; DML, dorsal mantle length; TL, total length; TW, total weight) o. agassizi o. vossi Mean SO N Range Mean SO N Range DMLmales 67.8 18.4 118 13-108 85.3 22.6 75 40-140 DMLfemales 59.1 10.9 64 32-85 68 14.7 48 34-101 DML all specimens 64.4 17 184 13-108 78.5 21.6 123 34-140 TL males 238.6 88.8 108 38-482 288.4 83 71 125-465 TL females 195 45.5 59 104-342 222.4 44.2 46 126-355 TL all specimens 221.6 79.9 169 38-482 262.4 77.3 117 125-465 TW males 840.2 1,109.6 112 4-5,400 907.2 765.2 76 75-3,050 TW females 400.9 289.3 63 29-1,650 362.3 236.7 48 72-1,300 TW all specimens 674.5 925.9 177 4-5,400 696.3 671.6 124 72-3,050 Samples of suckers and cirri were fixed in 5% formalin water for optical microscope analysis. For scanning electron microscope analysis samples of suckers and cirri were fixed for I h in 2% glutar- aldehyde in buffered phosphate solution, washed in the same buffered solution and postfixed in 1% OsO •. Sample Processing.-In the laboratory stomachs were dissected and their contents identified to the lowest taxonomic level possible. The reference papers of McLaughlin (1980), Barnard et a!. (1962), and Kensley (1978) were used for identification purposes. A subjective index of stomach fullness (SF!) was applied, from 0 (denoting empty stomach) to 4 (denoting distended). Samples of cirri were dehydrated in a series of ethanol and embedded in paraffin. Sections (6-8 /Lm) were obtained and stained with haematoxylin and eosin for optic microscope analysis. Electron mi- croscope samples were critical point dried in CO2 and examined with a Hitachi S-5 70 scanning electron microscope at 15 Kv. Data Treatment. -Student's '-tests were applied to the slopes of the different curves estimates. Krus- kall-Wallis non-parametric analysis of variance was applied to the stomach-fullness results, and x2 to analyze the day-night abundances recorded. REsULTS Length-weight Regressions.-A total of 105 males, 58 females and 2 non-sexed specimens of O. agassizi and 71 males and 46 females of O. vossi were examined. DML and TL were analyzed and TL was chosen over DML due to its best fitting with TW. Total length (TL) versus Total Weight (TW) regressions are presented in Figures 2 and 3. No significant differences were observed between males and females within species or between species (t-test, P > 0.05). In the O. agassizi regression equation: TW = 2·IQ-5·TL3·1, with a correlation coefficient ofr = 0.96 (sexes mixed). In the O. vossi regression equation: TW = 19·IQ-5·TU·7, with r = 0.93 (sexes mixed). Mandibular Growth. - In O. agassizi the size of mandibles varied between 5.1 and 20.8 mm UC length and between 4.1 and 15.6 mm LC length. Regression equations are: logUC = 0.46 + 0.22 'logTW, with R = 0.94, on a total number of specimens, N = 58 (Fig. 4); and logLC = 0.36 + 0.22 'logTW, with r = 0.99 and N = 58 (always sexes mixed). In O. vossi mandibles varied between 6.9 and 20.5 mm UC length and between 6.1 and 16.2 mm LC length. Regression equations are: logUC = 0.43 + 0.24 'logTW, with R = 0.87 and N = 115 (Fig. 5); and logLC = 0.38 + 0.23 'logTW, with r = 0.86 and N = 115 (sexes mixed). No significant differences were observed between species (t-test, P > 0.05). 290 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991 • • I I • • I I I I • I I • , I '- 15- Zambezl River Limpopo River 35° 40· - 40·S I I I I Itt I I I t I I I I I I I I I I I I • I • • • • 15· 20· 25° 30· 35·E Figure 1. Sampled area. Food and Feeding. - The remains found in the stomachs included setae and man- dibles of polychaetes, appendages, mandibles and carapace pieces of crustaceans, pieces of microgastropods and microbivalves, sand and foraminifers. Exuviae of the calanoid copepod :species Undinopsis sp., Oothrix sp. and Ateidopsis sp. were recorded in four stomachs. The fact that Opisthotheuthis species do not have radulae permitted the identification of abundant whole prey items. Prey categories iden- tified for the two species are shown in Table 2. A total of 125 individuals of O. agassizi and 93 of 0. vossi had prey items in their stomachs. Benthic gammarid amphipods and polychaetes comprised the major prey categories in both species (Table 3). For both species a maximum of 6 prey categories was observed in a single stomach. An average of 1.9 and 2.0 prey categories per stomach were observed for O. agassizi and O. vossi respectively. Whole prey sizes per category ranged 1- 8 mm in Crustacea, 1--2.5 mm in Gastropoda and 1.5-2.3 mm in Bivalvia. The diet of males and females proved to be similar. No differences in the type offood prey were observed between daytime and nighttime samples. Although octopod weight was independent of prey categories; for both species, polychaetes were more abundant as prey in the stomachs of individuals larger than 500 g, while amphipods dominated in individuals below 500 g (Table 3). VILLANUEVA AND GUERRA: FEEDING ECOLOGY IN OPISTHOTEUTHIS 291 o 5 Q,~gassizi o -m ~ 4 o Males -.s:: ,--m CD 3 + Females ~ ('is 2 -0 •••• 1 + + 200 400 Total Length (mm) Figure 2.
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