#

 # #

#

800 m

Peloponissos

**Zakinthos**

Epirus

Peloponissos

Egina Isl. Salami na Isl

The highly diversified diet observed in *A. foliacea* is typical of bathyal penaeoideans in the Western Mediterranean (Cartes, 1995). The feeding activity of *A. foliacea* in the Eastern Ionian Sea was examined studing the stomach fullness according the equations (i) wet food weight (g) per 100 g shrimp wet weight [% body weight (BW) Wet = (SWWBW) x 100] and (ii) dry food weight (g) per 100 g wet weight [%BWDry = (SWDBW) \* 100] (Héroux & Magnan, 1996). The nutritional quality (food quality) of the preys has been estimated by two ways: (a) % dry weight (DW) = (SWDSWW) 100 and (b) % ash free dry weight (AFDW) = (AFDWSWD) x 100, where SWW=stomach wet weight, g, SWD= stomach dry weight, gr after 24 h of oven drying at 70o C), ash-free dry weight (AFDW; as loss on ignition at 450o C for 3 h) and BW is the body weight. All the weights were measured to an accuracy of 0.0001 g). The food quality indices are a measure of total organic matter and form a better estimation of food value that wet weight, which includes substantial amounts of inorganic material (Hiller-Adams & Childress, 1983). The stomach fullness of *A. foliacea* varied Seasonally in both sexes and both fullness indices (%BW Wet, %BW Dry) were significantly higher in females than in males for each Season. The maximum values of %BW Wet in both

Diet quality (%BW Dry and %AFDW) also differed significantly among Seasons for both sexes of *A. foliacea*. In general, few significant differences in food quality were detected between males and females for each Season. Males and females of *A. foliacea* presented the

highest values of both quality indices in spring and the minimum in winter.

Eastern Ionian are given below.

Fig. 6. Study area in the Eastern Greek Ionian Sea

sexes occurred in winter and the minimum in spring.

**4.2** *foliacea's* **feeding habits** 

**4.2.1 Feeding activity and food quality** 

The feeding habits of this decapods identified in the Ionian Sea are generally comparable to those reported in other regions of the Mediterranean, such as in the Catalan Sea (Cartes, 1995), Sicilian Channel (Gristina et al., 1992) and Aegean Sea (Chartosia et al., 2005). Any difference observed in the whole Mediterranean, such as food diversity, different food categories and mean number of prey could be due to bottom morphology (Cartes 1995) and to the oligotrophic conditions of the Eastern Mediterranean. This characteristic of the Eastern Mediterranean could also explain the increased number of pelagic prey consumed by *A. foliacea* compared to the western part of the basin (Cartes, 1995). The considerably higher water temperature of the Eastern Mediterranean (Politou et al., 2004) may also play a role, resulting in a higher metabolic rate of this species, in comparison with those from the western part of the basin.

Trophic diversity ((H', Shannon-Wiener index) varied slightly among Seasons in both sexes (Figure 7) and no statistically significant differences were established between sexes. The maximum diversity (3.00 and 3.04 for males and females, respectively) and mean number of prey items (2.9 and 3.1 for males and females, respectively) were found in summer for both sexes of *A. foliacea*.

The observed low number of empty stomachs [(number of empty stomachs per number of stomachs examined) \* 100] (Hyslop, 1980) in the present study, ranging from 4,5 to 18,1%, indicating either a high feeding rate or slow digestion rate, could be explained by their high metabolic rates. The lowest proportion of empty stomachs of *A. foliacea* was found in spring for both sexes, followed by summer. In contrast, the highest number of empty stomachs was found in autumn for females and summer for males.

Fig. 7. Diversity index (H', Shannon-Wiener index) values for *A. foliacea* per sex and season in the Ionian Sea.

In general, a decrease in diversity and mean prey items with increasing overlap was observed. In the Eastern Ionian Sea, the giant red shrimp fed on a greater proportion of pelagic resources and prey with a good swimming ability, such as the natantian decapods,

Feeding Habits of Both Deep-Water Red Shrimps, *Aristaeomorpha foliacea*

assemblages rather than a preference for specific items.

specimens, but in different proportions

**4.3 Aristeus antennatus' feeding habits 4.3.1 Feeding activity and food quality** 

(Cartes et al., 2008a).

and *Aristeus antennatus* (Decapoda, Aristeidae) in the Ionian Sea (E. Mediterranean) 123

end of winter and by spring almost all females are inseminated (Kapiris, 2004). The minimum value of the stomach fullness in spring, in combination to the highest food quality value and the lowest vacuity index in females in the same Season, suggests that egg maturation is connected to the feeding habits of *A. foliacea*. During winter, *A. foliacea* had the highest stomach fullness, but with decreased food quality. This increase of food consumption by the giant red shrimp of the Ionian Sea during the pre-reproductive period has also been observed in *A. antennatus* off the Balearic Islands. Increased feeding rates could be the main reason for its egg development and could allow earlier gonad maturity

Besides the Seasonal feeding adaptation to the biological requirements (reproductive process), food availability also plays an important role for these species in the Eastern Ionian Sea. The highest densities in the suprabenthic fauna (mysids, cumaceans, amphipods, isopods, tanaidaceans) have been observed during spring, but zooplankton (chiefly copepods, ostracods and chaetognaths) were more abundant in summer and autumn. Such fluctuations in food availability have also been shown in the diets of both sexes of *A. foliacea* in this study. Thus, the diet of the giant red shrimp probably reflects localized forage

The size-related changes in diet composition are an important factor in determining ecological relationships of marine organisms during their life span. Comparison of diet composition, dietary diversity, and feeding activity among small, medium and (only for females) large individuals reveals that this decapod undergoes slight changes in feeding habits with increasing body size, as well as gonad maturity, in the Eastern Ionian Sea. Small males and females (immature individuals) consumed fewer prey due to their smaller stomachs, with more frequent occurrence of epibenthic prey in their foreguts. Larger, mature individuals of both sexes are more efficient predators due to their greater swimming ability and larger mandibles. A positive trend of ingesting larger prey with increased size was observed only for females. This is the first time where this gradation, probably due to the population structure and to morphological variation among size classes and sexes, has been observed for *A. foliacea*. In general, somatic growth and gonad development induce a change in this species' feeding behavior as the body grows an increase in the mean weight of prey and a decrease in the mean number of prey items per stomach was obvious. However, almost the same prey occurred in the stomachs of small, medium and large

A differentiation has been presented in *A. antennatus* diet according to the depth in the western Mediterranean (Cartes, 1994), the feeding time (Cartes, 1993a) and the daily consumption of food (Maynou & Cartes, 1997, 1998; Cartes & Maynou, 1998). The diet of *A. antennatus* changed as a function of depth at around 1000 m depth in the Catalan Sea, as a function of Seasonality influences by planktonic prey in deeper zones and by possible nocturnal movements upward along the slope canyons (Cartes, 1993a, 1994; Cartes et al., 2010). The importance of spatial patterns in its diet and feeding habits and the main environmental variables controlling these trophic aspects has been studied by Cartes et

and to a lesser extent on benthic prey, indicating that this shrimp is an active and effective predator of the bathyal zone in the Eastern Mediterranean. The characteristic of its active predation could be also confirmed by the very low abundance of infaunal and epibentic prey (e.g. polychaetes, bivalves and gastropods) in the stomachs of this species. The increased abundance of fishes and cephalopods in their foreguts most probably reflects the great scavenging ability of this species. In any case, this does not exclude the possibility that this species feeds actively upon fishes and cephalopods.

### **4.2.2 Food habits in relation to sex, season and size**

The diets of both sexes of *A. foliacea* consisted of 60 different prey categories (most as species-level prey categories). The preys belonged chiefly to three major groups: (i) crustaceans – particularly decapods, reptantia (anomurans, brachyurans), amphipods, euphausiids, ostracods, copepods, mysids, tanaidaceans, cumaceans, (ii) cephalopods and (iii) fishes. These three prey categories constituted 72–82% of the relative abundance and total occurrence for males and 70–88% of the relative abundance and the total occurrence in females. The most dominant natantians found were the nektobenthic *Plesionika martia, Plesionika heterocarpus* and *Plesionika giglioli*, followed by *Pasiphaea sp., Sergestes sp.* and *Solenocera sp*. Some appendages from *Aristeus antennatus* were also found mainly in female *A. foliacea*. These findings could be accidental, as they were found in the sampling stations where both species coexisted and, thus, some body appendages could have been destroyed and mixed during the net tow (net feeding). It is also possible that the smaller individuals of each species were consumed by larger adults of the other, due to their voracious character, but further study of this hypothesis is required. Among cephalopods, the dominant species were *Abraliopsis pfefferi, Pyroteuthis margarifera* and *Abralia veranyi*. For fishes, specimens of Myctophidae and Macrouridae were the most abundant in the foreguts.

Only a partial differentiation in the feeding behaviour, in terms of both diet composition and feeding activity, was observed between sexes of *A. foliacea*. In general, both sexes fed upon natantian decapods, particularly *Plesionika spp., Sergestes sp., Pasiphaea sp.*, and fishes throughout the year, while 'other crustaceans' and polychaetes were ingested on a secondary basis. The consumption of the same prey items, but in different abundance and occurrence, may be attributed to sexual dimorphism and to size difference between the sexes.

In general, the existence of regular Seasonal rhythms in the feeding activity of deep water species is related mainly to Seasonal fluctuations in various factors including the abundance of their prey, depth, local geographical characteristics, submarine canyons, bottom type, Seabed features, Seasonal horizontal or diurnal vertical migrations, etc. (Cartes 1993, 1998). In the Eastern Ionian Sea the Seasonal feeding habits of the giant red shrimp seem to be related to reproduction, and perhaps to other biological processes, and food availability.

High observed values of trophic overlap between Seasons for both sexes indicated that Season is not the main factor affecting the diet of deep-water shrimps in the Eastern Ionian Sea. In spite of this, most feeding activity values (empty stomachs, quality indices, mean number of prey items found into the stomachs, diversity index) support the finding that feeding activity increased during spring–summer for both sexes. This increase could be attributed to the increased reproductive activity (gonad maturity, egg-laying) observed in this period (Papaconstantinou & Kapiris 2001, 2003). In addition, copulation begins at the

and to a lesser extent on benthic prey, indicating that this shrimp is an active and effective predator of the bathyal zone in the Eastern Mediterranean. The characteristic of its active predation could be also confirmed by the very low abundance of infaunal and epibentic prey (e.g. polychaetes, bivalves and gastropods) in the stomachs of this species. The increased abundance of fishes and cephalopods in their foreguts most probably reflects the great scavenging ability of this species. In any case, this does not exclude the possibility that

The diets of both sexes of *A. foliacea* consisted of 60 different prey categories (most as species-level prey categories). The preys belonged chiefly to three major groups: (i) crustaceans – particularly decapods, reptantia (anomurans, brachyurans), amphipods, euphausiids, ostracods, copepods, mysids, tanaidaceans, cumaceans, (ii) cephalopods and (iii) fishes. These three prey categories constituted 72–82% of the relative abundance and total occurrence for males and 70–88% of the relative abundance and the total occurrence in females. The most dominant natantians found were the nektobenthic *Plesionika martia, Plesionika heterocarpus* and *Plesionika giglioli*, followed by *Pasiphaea sp., Sergestes sp.* and *Solenocera sp*. Some appendages from *Aristeus antennatus* were also found mainly in female *A. foliacea*. These findings could be accidental, as they were found in the sampling stations where both species coexisted and, thus, some body appendages could have been destroyed and mixed during the net tow (net feeding). It is also possible that the smaller individuals of each species were consumed by larger adults of the other, due to their voracious character, but further study of this hypothesis is required. Among cephalopods, the dominant species were *Abraliopsis pfefferi, Pyroteuthis margarifera* and *Abralia veranyi*. For fishes, specimens of

Only a partial differentiation in the feeding behaviour, in terms of both diet composition and feeding activity, was observed between sexes of *A. foliacea*. In general, both sexes fed upon natantian decapods, particularly *Plesionika spp., Sergestes sp., Pasiphaea sp.*, and fishes throughout the year, while 'other crustaceans' and polychaetes were ingested on a secondary basis. The consumption of the same prey items, but in different abundance and occurrence,

In general, the existence of regular Seasonal rhythms in the feeding activity of deep water species is related mainly to Seasonal fluctuations in various factors including the abundance of their prey, depth, local geographical characteristics, submarine canyons, bottom type, Seabed features, Seasonal horizontal or diurnal vertical migrations, etc. (Cartes 1993, 1998). In the Eastern Ionian Sea the Seasonal feeding habits of the giant red shrimp seem to be related to reproduction, and perhaps to other biological processes, and food availability.

High observed values of trophic overlap between Seasons for both sexes indicated that Season is not the main factor affecting the diet of deep-water shrimps in the Eastern Ionian Sea. In spite of this, most feeding activity values (empty stomachs, quality indices, mean number of prey items found into the stomachs, diversity index) support the finding that feeding activity increased during spring–summer for both sexes. This increase could be attributed to the increased reproductive activity (gonad maturity, egg-laying) observed in this period (Papaconstantinou & Kapiris 2001, 2003). In addition, copulation begins at the

this species feeds actively upon fishes and cephalopods.

**4.2.2 Food habits in relation to sex, season and size** 

Myctophidae and Macrouridae were the most abundant in the foreguts.

may be attributed to sexual dimorphism and to size difference between the sexes.

end of winter and by spring almost all females are inseminated (Kapiris, 2004). The minimum value of the stomach fullness in spring, in combination to the highest food quality value and the lowest vacuity index in females in the same Season, suggests that egg maturation is connected to the feeding habits of *A. foliacea*. During winter, *A. foliacea* had the highest stomach fullness, but with decreased food quality. This increase of food consumption by the giant red shrimp of the Ionian Sea during the pre-reproductive period has also been observed in *A. antennatus* off the Balearic Islands. Increased feeding rates could be the main reason for its egg development and could allow earlier gonad maturity (Cartes et al., 2008a).

Besides the Seasonal feeding adaptation to the biological requirements (reproductive process), food availability also plays an important role for these species in the Eastern Ionian Sea. The highest densities in the suprabenthic fauna (mysids, cumaceans, amphipods, isopods, tanaidaceans) have been observed during spring, but zooplankton (chiefly copepods, ostracods and chaetognaths) were more abundant in summer and autumn. Such fluctuations in food availability have also been shown in the diets of both sexes of *A. foliacea* in this study. Thus, the diet of the giant red shrimp probably reflects localized forage assemblages rather than a preference for specific items.

The size-related changes in diet composition are an important factor in determining ecological relationships of marine organisms during their life span. Comparison of diet composition, dietary diversity, and feeding activity among small, medium and (only for females) large individuals reveals that this decapod undergoes slight changes in feeding habits with increasing body size, as well as gonad maturity, in the Eastern Ionian Sea. Small males and females (immature individuals) consumed fewer prey due to their smaller stomachs, with more frequent occurrence of epibenthic prey in their foreguts. Larger, mature individuals of both sexes are more efficient predators due to their greater swimming ability and larger mandibles. A positive trend of ingesting larger prey with increased size was observed only for females. This is the first time where this gradation, probably due to the population structure and to morphological variation among size classes and sexes, has been observed for *A. foliacea*. In general, somatic growth and gonad development induce a change in this species' feeding behavior as the body grows an increase in the mean weight of prey and a decrease in the mean number of prey items per stomach was obvious. However, almost the same prey occurred in the stomachs of small, medium and large specimens, but in different proportions

### **4.3 Aristeus antennatus' feeding habits**

### **4.3.1 Feeding activity and food quality**

A differentiation has been presented in *A. antennatus* diet according to the depth in the western Mediterranean (Cartes, 1994), the feeding time (Cartes, 1993a) and the daily consumption of food (Maynou & Cartes, 1997, 1998; Cartes & Maynou, 1998). The diet of *A. antennatus* changed as a function of depth at around 1000 m depth in the Catalan Sea, as a function of Seasonality influences by planktonic prey in deeper zones and by possible nocturnal movements upward along the slope canyons (Cartes, 1993a, 1994; Cartes et al., 2010). The importance of spatial patterns in its diet and feeding habits and the main environmental variables controlling these trophic aspects has been studied by Cartes et

Feeding Habits of Both Deep-Water Red Shrimps, *Aristaeomorpha foliacea*

probably reflects availability in the marine environment.

the adults of the other one. In any case, further study is necessary.

and to size difference between the sexes.

**4.3.2 Seasonal differences** 

and *Aristeus antennatus* (Decapoda, Aristeidae) in the Ionian Sea (E. Mediterranean) 125

The diet of *A. antennatus* both sexes consisted of 54 prey categories. These prey items belonged mainly to smaller crustaceans (e.g. natantian decapods, *Plesionika sp., Sergestes sp.*, euphausiids, tanaidaceans), molluscs primarily gastropods, bivalves, polychaetes (Eunicidae, Spionidae, and Nereididae), chaetognaths and, to a lesser extent, fishes. The above prey categories consisted of 71–82% of the relative abundance and total occurrence for males and 61–81% of the relative abundance and the total occurrence in females. Its diversified diet in the present study area consists of increased endobenthic and epibenthic invertebrates and includes organisms that are related with the Seabed, nekton and decapods. This species is among the few megabenthic predators whose diet is mainly based on benthos in the deep Mediterranean (Cartes & Carrassón, 2004). The increased abundance of gastropods, echinoderms, polychaetes — chiefly Eunicidae, sipunculans and chaetognaths in the stomachs, confirms that this species in the Greek Ionian Sea could be considered a "slow hunter", foraging mainly on organisms that live completely or partially buried in the substratum. The macrophyte consumption was rare in both sexes and

The data of the present study confirm that *A. antennatus* could be considered a less active and slower hunter than the other aristeid species (*A. foliacea*) found in the same area (Kapiris et al., 2010) and preys on detrivores or small predators occupying a lower position in the benthopelagic food chain (Maynou & Cartes, 1997). The feeding activity patterns of *A. antennatus* in the Greek Ionian Sea are, more or less, comparable to those reported in other geographical regions, such as the central (e.g. Relini & Orsi Relini, 1987; Follesa et al., 2009) or in the western Mediterranean (Cartes & Sardà, 1989; Maynou & Cartes, 1998). Apparent differences in the activity patterns should be attributed to the more oligotrophic character of the Ionian Sea (E. Mediterranean) in relation to the western one and to the bottom morphology (Cartes, 1995). The above mentioned oligotrophic character of the eastern Mediterranean could explain the presence of the increased number of some pelagic preys in its stomachs, in comparison to the western one (Cartes, 1995), but – as we said before – these preys constitute the minority comparing to the benthic ones. Some remains of the sympatric *A. foliacea* in the stomachs of *A. antennatus* and vice versa could be accidental, since they have been found in the sampling stations where both species coexisted and, thus, some body appendages were destroyed and mixed during the net tow (net feeding). It is possible that the smaller individuals of each species, due to their voracious character, can be fed by

Only a partial differentiation in the feeding behaviour between sexes, in terms of both diet composition and feeding activity, is observed. Males exhibit lower values of fullness, food quality indices and evenness than females. Both sexes consume the same prey items, but in different abundance and occurrence. From the above results, a slightly higher predatory ability of females is shown. These differences could also be attributed to sexual dimorphism

Taking into consideration the narrow depth sampling range, the estimated values of the trophic overlap indicate that, Season could not be considered as the main factor affecting the diet of blue–red shrimp in the Greek Ionian Sea, like in *A. foliacea*. Notwithstanding, some particular topics are analyzed below. The existence of regular Seasonal rhythms in the

al.(2008b) in Western Mediterranean. In the whole E. Mediterranean, the feeding habits of *A. antennatus* have been studied in detail in the Ionian (Kapiris & Thessalou-Legaki, 2011) and the Aegean Sea (Chartosia et al., 2005).

The observed low number of empty stomachs in the Greek Ionian (mean value of the empty stomachs in males was 6,53 and for females was 8,54) could be explained by their high metabolic rates (Company, 1995). Significant statistical differences amongst the Seasonal medians of both fullness indices were found [%BW Wet (for both sexes) and %BW Dry (only in females)]. The maximum values of %BW Wet were determined in winter in both sexes and the minimum in spring. Both fullness indices were statistically higher in females than those of males (Figure 8).

Significant statistical differences amongst the Seasonal medians of both indices of food quality (%DW, %AFDW) were established only for females. Females presented a lower value of %DW and higher of %AFDW than males, in spring, while their highest values of both quality indices were found in spring (Figure 8).

Fig. 8. Seasonal values of stomach fullness and food quality of both sexes of *A. antennatus* in the Greek Ionian Sea.

al.(2008b) in Western Mediterranean. In the whole E. Mediterranean, the feeding habits of *A. antennatus* have been studied in detail in the Ionian (Kapiris & Thessalou-Legaki, 2011) and

The observed low number of empty stomachs in the Greek Ionian (mean value of the empty stomachs in males was 6,53 and for females was 8,54) could be explained by their high metabolic rates (Company, 1995). Significant statistical differences amongst the Seasonal medians of both fullness indices were found [%BW Wet (for both sexes) and %BW Dry (only in females)]. The maximum values of %BW Wet were determined in winter in both sexes and the minimum in spring. Both fullness indices were statistically higher in females than

Significant statistical differences amongst the Seasonal medians of both indices of food quality (%DW, %AFDW) were established only for females. Females presented a lower value of %DW and higher of %AFDW than males, in spring, while their highest values of

**Females**

Winter Spring Summer Autumn **Season**

%DW % AFDW % BW Wet % BW Dry

**Males**

Winter Spring Summer Autumn **Season**

%DW % AFDW % BW Wet % BW Dry

Fig. 8. Seasonal values of stomach fullness and food quality of both sexes of *A. antennatus* in

0 0,05 0,1 0,15 0,2 0,25

0 0,02 0,04 0,06 0,08 0,1 0,12

**Fullness**

**Fullness**

the Aegean Sea (Chartosia et al., 2005).

both quality indices were found in spring (Figure 8).

**Food quality**

the Greek Ionian Sea.

**Food quality**

those of males (Figure 8).

The diet of *A. antennatus* both sexes consisted of 54 prey categories. These prey items belonged mainly to smaller crustaceans (e.g. natantian decapods, *Plesionika sp., Sergestes sp.*, euphausiids, tanaidaceans), molluscs primarily gastropods, bivalves, polychaetes (Eunicidae, Spionidae, and Nereididae), chaetognaths and, to a lesser extent, fishes. The above prey categories consisted of 71–82% of the relative abundance and total occurrence for males and 61–81% of the relative abundance and the total occurrence in females. Its diversified diet in the present study area consists of increased endobenthic and epibenthic invertebrates and includes organisms that are related with the Seabed, nekton and decapods. This species is among the few megabenthic predators whose diet is mainly based on benthos in the deep Mediterranean (Cartes & Carrassón, 2004). The increased abundance of gastropods, echinoderms, polychaetes — chiefly Eunicidae, sipunculans and chaetognaths in the stomachs, confirms that this species in the Greek Ionian Sea could be considered a "slow hunter", foraging mainly on organisms that live completely or partially buried in the substratum. The macrophyte consumption was rare in both sexes and probably reflects availability in the marine environment.

The data of the present study confirm that *A. antennatus* could be considered a less active and slower hunter than the other aristeid species (*A. foliacea*) found in the same area (Kapiris et al., 2010) and preys on detrivores or small predators occupying a lower position in the benthopelagic food chain (Maynou & Cartes, 1997). The feeding activity patterns of *A. antennatus* in the Greek Ionian Sea are, more or less, comparable to those reported in other geographical regions, such as the central (e.g. Relini & Orsi Relini, 1987; Follesa et al., 2009) or in the western Mediterranean (Cartes & Sardà, 1989; Maynou & Cartes, 1998). Apparent differences in the activity patterns should be attributed to the more oligotrophic character of the Ionian Sea (E. Mediterranean) in relation to the western one and to the bottom morphology (Cartes, 1995). The above mentioned oligotrophic character of the eastern Mediterranean could explain the presence of the increased number of some pelagic preys in its stomachs, in comparison to the western one (Cartes, 1995), but – as we said before – these preys constitute the minority comparing to the benthic ones. Some remains of the sympatric *A. foliacea* in the stomachs of *A. antennatus* and vice versa could be accidental, since they have been found in the sampling stations where both species coexisted and, thus, some body appendages were destroyed and mixed during the net tow (net feeding). It is possible that the smaller individuals of each species, due to their voracious character, can be fed by the adults of the other one. In any case, further study is necessary.

Only a partial differentiation in the feeding behaviour between sexes, in terms of both diet composition and feeding activity, is observed. Males exhibit lower values of fullness, food quality indices and evenness than females. Both sexes consume the same prey items, but in different abundance and occurrence. From the above results, a slightly higher predatory ability of females is shown. These differences could also be attributed to sexual dimorphism and to size difference between the sexes.

### **4.3.2 Seasonal differences**

Taking into consideration the narrow depth sampling range, the estimated values of the trophic overlap indicate that, Season could not be considered as the main factor affecting the diet of blue–red shrimp in the Greek Ionian Sea, like in *A. foliacea*. Notwithstanding, some particular topics are analyzed below. The existence of regular Seasonal rhythms in the

Feeding Habits of Both Deep-Water Red Shrimps, *Aristaeomorpha foliacea*

observations) and oxygen consumption rates (Company & Sardà 1998).

also observed in deep-water decapods (Fanelli & Cartes, 2008).

DG XIV, Contract nº MED92/005.

I.T.P.P. Special Publication.

2, No 45, pp. 1-6.

**5. Conclusions** 

**6. References** 

integrated management in the future.

and *Aristeus antennatus* (Decapoda, Aristeidae) in the Ionian Sea (E. Mediterranean) 127

Our results on the feeding ecology of both deep water shrimps could be considered as primary importance for the future management of deep water assemblages, since they play an important role. Since the deep waters in the E. Ionian Sea are almost unexploited, the present data could elucidate the relationships between species in this ecosystem improving, thus, the knowledge and the trophic relationships among the species helping in their

According all the studies carried out on both decapods feeding habits, *A. foliacea* exploits different resources from those used by *A. antennatus* and, despite both shrimps have similar morphologies and size ranges, the exploitation of different resources probably both species to coexist in the same areas (Cartes, 1995). In addition to this, since both deep-sea red shrimps belonging in the same family, have an almost similar depth distribution It is expected that they have similar energy values (in terms of wet mass), water body content (K. Kapiris unpublished

Concluding, the increased demand of the large energetic content and the food availability in the same period make us suggest that both facts could stimulate fecundity in the deep-sea blue–red shrimp in the E. Mediterranean. A similar trend has been shown for the same species in the western Mediterranean (Cartes et al., 2008a, b). Generally, energy reserves strongly affect fecundity and reproduction in fishes (e.g. Lloret et al., 2005) and have been

Aquastudio (1996). *Survey of red shrimp fishing in the Western Italian basins.* Final Report. CE

Atkinson, D.B. (1995). The biology and fishery of roundnose grenadier (*Coryphaenoides* 

Bianchi, N. & Morri, C. (2000). Marine biodiversity of the Mediterranean Sea, situation,

Bouchet, P.H. & Taviani, M. (1992). The Mediterranean deep sea fauna, pseudopopulations

Brian, A. (1931). La biologia del fondo a "scampi" nel Mare Ligure. V. *Aristaeomorpha,* 

Burukovsky, R.N., Romensky, L.L., Kozyaistva, R. & Okeanografii, I. (AtantNIRO) (1972).

of Atlantic species?. *Deep-Sea Res*., Vol. 39, No 2, pp. 169-184.

Sciences, Vol. 296, pp. 51-111, Kluwer Academic Publishers, Dorbrecht. Bianchini, M.L. & Ragonese, S. (1994). Life cycles and fisheries of the deepwater red shrimps

*rupestris* Gunnerus, 1765) in the north west Atlantic. In *Deep-water fisheries of the south Atlantic Oceanic Slope*, Hopper, A. G. (Ed.), NATO Asi. Series E., Applied

*A. foliacea* and *A. antennatus*. *Proceedings of the International workshop held in the Istituto di Tecnologia della Pesca e del Pescato*, pp. 1-87, Mazara del Vallo. N.T.R.-

problems and prospects for future research. *Mar. Poll. Bull*. Vol. 40, No 5, pp. 367-376.

*Aristeus* ed altri macruri natante. *Boll. Mus. Zool. Anat. Comp. R. Univ. Genova*, Vol.

On the variability of the rostrum in the *Aristeus varidens* (Decapoda, Penaeidae). *Trudy Atlanticheskii Nauchna-issledovatel'skii Inst.* Vol. 42, pp. 156-161 [In Russian].

feeding activity of deep-water species is mainly related to the Seasonal fluctuations of abundance of prey they consume, the depth, the local geographical characteristics, the submarine canyons, the type of bottom, the Seabed, the Seasonal horizontal and diurnal vertical migrations, etc. (Cartes, 1993a, 1998). In addition to this, the Seasonal changes in stomach fullness of blue–red shrimp could be possibly linked to the oceanographic processes and to the several water masses, at least in the W. Mediterranean (Cartes et al., 2008b; Maynou, 2008).

The above slight Seasonal changes in the feeding dynamics of this aristeid in the Greek Ionian Sea seem to be related mainly to their biological processes (e.g. mating and reproduction) and to the food availability. The increased values of food quality indices and diversity support the finding that feeding activity seemed to increase qualitatively – in the period spring–summer, mainly for females. In addition to this, the observed highest empty stomachs found in these Seasons, mainly for females, could be attributed to the increased volume of the gonads which press the stomach. This increase of the highly energetic diet could be attributed to the increased pre- and reproductive activity observed in this period (Kapiris and Thessalou-Legaki, 2006, 2009). As Cartes et al. (2008a) noted *A. antennatus* seemed to increase the energy intake in its diet from February to April-June in the western Mediterranean. During winter both sexes of *A. antennatus* in Greek Ionian Sea consume an increased number of prey items, having as a result the highest stomach fullness, but of decreased quality. This phenomenon could be related to the mating period which takes place in this Season (Kapiris & Thessalou-Legaki, 2006, 2009). Besides the Seasonal feeding adaptation to the biological requirements, the food availability also plays an important role for this species in the Greek Ionian Sea. Madurell & Cartes (2005) point out that, in the same study area, the suprabenthos fauna (mysids, cumaceans, amphipods, isopods, and tanaidaceans) showed the highest densities in spring, while the zooplankton fauna (chiefly copepods, ostracods and chaetognaths) was more abundant in autumn and summer. In agreement with the results of the present study, the above fluctuations of food availability are also shown in the diet of *A. antennatus*. Thus, the diet of the blue–red shrimp probably reflects localized forage assemblages rather than a preference for specific items. In addition to this, these results reinforce the opinion concerning the "accidental hunting" of *A. antennatus*.

### **4.3.3 Ontogenetic differences**

Comparison of the diet composition, dietary diversity and feeding activity between the small size, medium size and – only in females – large size individuals reveals that this species undergoes changes in feeding habits with increasing body size and gonad maturity in the Greek Ionian Sea. Small immature individuals consume less prey, mainly epibenthic, – but of increased quality – due to their smaller stomach. Larger mature specimens of both sexes are more efficient predators because of their greater swimming ability and their larger mandibles. The positive trend between increasing females' body size and consumption of larger prey is observed could be attributed to the population structure and to the morphological characteristics of the different size classes and sexes. In general, somatic growth and gonad development induce a change of *A. antennatus* feeding behaviour in the Greek Ionian Sea: as the body grows, an increasing mean weight of prey and mean number of prey items per stomach was obvious. However, almost the same prey occurred in the stomachs of small, medium and large specimens, but in different proportions.

### **5. Conclusions**

126 Food Quality

feeding activity of deep-water species is mainly related to the Seasonal fluctuations of abundance of prey they consume, the depth, the local geographical characteristics, the submarine canyons, the type of bottom, the Seabed, the Seasonal horizontal and diurnal vertical migrations, etc. (Cartes, 1993a, 1998). In addition to this, the Seasonal changes in stomach fullness of blue–red shrimp could be possibly linked to the oceanographic processes and to the several water masses, at least in the W. Mediterranean (Cartes et al.,

The above slight Seasonal changes in the feeding dynamics of this aristeid in the Greek Ionian Sea seem to be related mainly to their biological processes (e.g. mating and reproduction) and to the food availability. The increased values of food quality indices and diversity support the finding that feeding activity seemed to increase qualitatively – in the period spring–summer, mainly for females. In addition to this, the observed highest empty stomachs found in these Seasons, mainly for females, could be attributed to the increased volume of the gonads which press the stomach. This increase of the highly energetic diet could be attributed to the increased pre- and reproductive activity observed in this period (Kapiris and Thessalou-Legaki, 2006, 2009). As Cartes et al. (2008a) noted *A. antennatus* seemed to increase the energy intake in its diet from February to April-June in the western Mediterranean. During winter both sexes of *A. antennatus* in Greek Ionian Sea consume an increased number of prey items, having as a result the highest stomach fullness, but of decreased quality. This phenomenon could be related to the mating period which takes place in this Season (Kapiris & Thessalou-Legaki, 2006, 2009). Besides the Seasonal feeding adaptation to the biological requirements, the food availability also plays an important role for this species in the Greek Ionian Sea. Madurell & Cartes (2005) point out that, in the same study area, the suprabenthos fauna (mysids, cumaceans, amphipods, isopods, and tanaidaceans) showed the highest densities in spring, while the zooplankton fauna (chiefly copepods, ostracods and chaetognaths) was more abundant in autumn and summer. In agreement with the results of the present study, the above fluctuations of food availability are also shown in the diet of *A. antennatus*. Thus, the diet of the blue–red shrimp probably reflects localized forage assemblages rather than a preference for specific items. In addition to this, these results reinforce the opinion concerning

Comparison of the diet composition, dietary diversity and feeding activity between the small size, medium size and – only in females – large size individuals reveals that this species undergoes changes in feeding habits with increasing body size and gonad maturity in the Greek Ionian Sea. Small immature individuals consume less prey, mainly epibenthic, – but of increased quality – due to their smaller stomach. Larger mature specimens of both sexes are more efficient predators because of their greater swimming ability and their larger mandibles. The positive trend between increasing females' body size and consumption of larger prey is observed could be attributed to the population structure and to the morphological characteristics of the different size classes and sexes. In general, somatic growth and gonad development induce a change of *A. antennatus* feeding behaviour in the Greek Ionian Sea: as the body grows, an increasing mean weight of prey and mean number of prey items per stomach was obvious. However, almost the same prey occurred in the

stomachs of small, medium and large specimens, but in different proportions.

2008b; Maynou, 2008).

the "accidental hunting" of *A. antennatus*.

**4.3.3 Ontogenetic differences** 

Our results on the feeding ecology of both deep water shrimps could be considered as primary importance for the future management of deep water assemblages, since they play an important role. Since the deep waters in the E. Ionian Sea are almost unexploited, the present data could elucidate the relationships between species in this ecosystem improving, thus, the knowledge and the trophic relationships among the species helping in their integrated management in the future.

According all the studies carried out on both decapods feeding habits, *A. foliacea* exploits different resources from those used by *A. antennatus* and, despite both shrimps have similar morphologies and size ranges, the exploitation of different resources probably both species to coexist in the same areas (Cartes, 1995). In addition to this, since both deep-sea red shrimps belonging in the same family, have an almost similar depth distribution It is expected that they have similar energy values (in terms of wet mass), water body content (K. Kapiris unpublished observations) and oxygen consumption rates (Company & Sardà 1998).

Concluding, the increased demand of the large energetic content and the food availability in the same period make us suggest that both facts could stimulate fecundity in the deep-sea blue–red shrimp in the E. Mediterranean. A similar trend has been shown for the same species in the western Mediterranean (Cartes et al., 2008a, b). Generally, energy reserves strongly affect fecundity and reproduction in fishes (e.g. Lloret et al., 2005) and have been also observed in deep-water decapods (Fanelli & Cartes, 2008).

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### *Edited by Kostas Kapiris*

The book discusses the novel scientific approaches for the improvement of the food quality and offers food scientists valuable assistance for the future. The detailed methodologies and their practical applications could serve as a fundamental reference work for the industry and a requisite guide for the research worker, food scientist and food analyst. It will serve as a valuable tool for the analysts improving their knowledge with new scientific data for quality evaluation. Two case study chapters provide data on the improvement of food quality in marine and land organisms in the natural environment.

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Food Quality

Food Quality